Medical Entomology by William B Herms SC.D.

MEDICAL ENTOMOLOGY
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THE MACMILLAN COMPANY
NEW YORK. BOSTON. CHICAGO
DA1.l.AS • ATLANTA. SAN FRANCISCO
MACMILLAN AND CO., LIMITED

LONDON. BOMBAY. CALCUTTA
MADRAS. MELBOURNE
THE MAC)ULLAN COMPANY

OF CANADA, LIMITED
~TORONTO
MEDICAL
E·NTOMOLOGY
With Special Reference to the Health and
Well-being of Man and Animals
BY vVILLIAM B. HERMS_, SC.D.

Late Professor of ParaSitology, Emeritus, VI . 't f
, ,mverSl yo
California; one-time Lecturer m TroplCal M d' ,
e lClne,
University of California, Medical School, San F '
. ranClSCO;
late Chairman, Division of Entomology and Pi 't logy
, 0 f C al1 'orma
University f' araSl 0 ,
FOURTH EDITION
Based on the book known as
MEDICAL AND VETERINARY
ENTOMOLOGY
THE MACMILLAN COMPANY: ~EW YORK

1953
Copyright, 1915, 1923, 1939, and 1950 by
THE MACMILLAN COMPANY
Copyright, 1943, by
WILLIAM B. HERMS
All rights reserved-no part of this book may be

reproduced in any form without permission in writing
from the publisher, except by a rez'iewer who wishes
to quote brief passages in connection with a review
written for inclusion in magazine or newspaper.
SECOND PRINTING, 1953

I gratefully and proudly dedicate the fourth edition
of Medical Entomology to my former graduate stu-
dents of the University of California, more than fifty of
whom served during World War II in many parts of
the world in various capacities in the mammoth strug-
gle to protect our fighting forces against the ravages of
arthropod-borne tropical diseases, particularly malaria.
PREFACE TO THE FOURTH EDITION
._.
THE ORIGINAL MANUSCRIPT for this book was prepared about forty
years ago, and the first edition, entitled Medical and Veterinary Ento-
mology, appeared in 1915. The book was most generously received from
the beginning and has been widely and increasingly used in each suc-
ceeding edition. Jt took considerable perseverance to complete another
revision for a fourth edition. With the encouragement of long-time users
of the work and the assistance of colleagues, graduate students, and
others this has now been accomplished.
The science of medical entomol~has been fully accepted as one
of the important fundamentals for ~plete knowledge and under-
standing of public health operations as well as for an understanding of
the health of domestic animals; it is essential to a complete understand-
ing of the epidemiology of disease. Medical entomology is now taught
in the entomological curricula of most colleges and universities, as well
as in curricula of public health, preventive medicine, and tropical medi-
cine.
The general plan of the book remains essentially unchanged; but ow-
ing to the rapid expansion of knowledge pertaining to arthropods as
vectors of infections of man and animals and because of changes in con-
trol technique due to the discovery of new insecticide, such as DDT, a
complete revision of nearly all chapters became necessary. Stress con-
tinues to be placed on biology (life history and ecology) as fundamen-
tal to rational control and as basic to sound epidemiological procedures.
While it is recognized that the advent of DDT did measurably change
arthropod-control concepts, the use of this and other new insecticides
precipitated new problems or new phases of old ones, particularly as
these newer insecticides affect the public .health either directly or in-
directly and to some degree also the "balance" in nature. Instead of
requiring less knowledge concerning the ecology of living things (so-
called "shot-gun" methods of application have too largely prevailed),
the wise and successful use of these insecticides requires much more
fundamental knowledge of the· ecology of the offending organisms as
well as of others living in the same niche. Hazards to the health of man
and his domestic animals must be well considered. Those in charge of
training military personnel for vector control operations ( to check
malaria, tsutsugamushi disease, etc.) under military conditions must
recognize, as do public health officials, that sole reliance on the tech-
vii
viii PREF ACE TO THE FOURTH EDITION
niques of chemical control (use of insecticides) and the neglect of knowl-

edge of vectors can lead only to failure in the long run.
Among the many persons to whom I am indebted for assistance I
wish to thank Professor E. O. Essig, my successor as head of the Divi-
sion of Entomology and Parasitology, University of California, for plac-
ing clerical and other assistance at my disposal. Thanks are due Professors
W. M. Hoskins, M. A. Stewart, R. Craig, and Dean Furman for technical
assistance, and to Dr. W. C. Reeves, H. F. Gray, Willis Doetschmann,
and Richard Coleman for critical reading of certain chapters. I wish to
thank Arve Dahl and Richard Peters of the Bureau of Vector Control of
the California State Department of Public Health for many courtesies
extended during the preparation of this edition. My thanks go partic-
ularly to Mrs. K. E. Frick for the task of typing the manuscript. To my
wife for long-continued encouragement lowe more than words can
express.
W. B. H.
Berkeley, California
April, 1949
PREFACE TO THE FIRST EDITION
._.
MUCH of the matter contained in the following pages was prepared

for the press more than six years ago, but owing to the rapid advances
made in the field of parasitology, particularly concerning insects, the
writer has withheld it until this time, when, after considerable revision
and addition, it has seemed expedient to publish the same. The manu-
script has been in almost constant use for a period of six years in teach-
ing classes in parasitology, both in the University of California and in
the San Francisco Veterinary College. It has been the aim to include
herewith a large part of the writer's original work, some of which has
until now remained unpublished, as well as the published observations
of many other investigators in this field, all of which has- gone to build
up the foundation of the new science of medical entomology.
This book is not intended to be a comprehensive treatise touching all
the investigations in the field of medical entomology, but rather an at-
tempt to systematize the subject and to assist in securing for it a place
among the applied biological sciences. However, a discussion is included
of all of the more important diseases and irritations of man and of the
domesticated animals in which insects and arachnids are concerned,
either as carriers or as causative organisms.
Owing to the immense literature on insects as relating to disease,
much of which is widely scattered., the student in this field must spend
considerable time in searching for the desired information and, what
is more important, the information is not readily accessible to the physi-
cian, the veterinaria'n, the health officer, and the sanitarian. It is there-
fore to be hoped that this book will not only prove useful as a text, but
also as a handbook for all individuals who are professionally interested
in the health and well-being of man and beast, as affected by insects
and arachnids.
In the second place detailed accounts of experiments are included
here and there, so that the investigator might employ the methods de-
scribed in either the repetition of the work or in carrying on further
investigations along the lines suggested.
Although many special papers have been consulted in the prepara-
tion of this work, a bibliography is not included herewith, inasmuch as
this information is obtainable in much more complete form in the bibli-
ographical works of other writers. Reference to special papers is usually
ix
x PREFACE TO THE FIRST EDITION
made in footnote form, but where certain facts have long been accepted
as common knowledge, reference is ordinarily omitted.
Sources from which assistance has been drawn are too numerous to
enumerate adequately, but to all who have contributed toward the
preparation of this work I wish to express my sincere appreciation and
thanks, but most particularly to my advanced students in parasitology,
who have contributed much valuable data, and to my colleagues, Pro-
fessor C. W. Woodworth, Dr. Edwin C. Van Dyke, Dr. W. A. Sawyer,
and Dr. S. B. Freeborn, and to my wife, Lillie M. Herms, for generous
cooperation and kindly criticism.
Unless otherwise credited the illustrations are from photographs and
drawings made by the author and various assistants. Thanks are due
particularly to Dr. William Colby Rucker for the use of flea drawings,
to Professor Herbert Osborn for permission to reproduce certain draw-
ings of biting and sucking lice, to Dr. Bruce Mayne for photographs of
Tabanas striatus, and stomachs of infected Anopheles mosquitoes, to
Professor J. S. Hine for photographs of certain other tabanids, and to
Mr. W. C. Matthews, scientific illustrator, for valuable assistance in the
preparation of many of the figures.
W. B. H.
Berkeley, California
CONTENTS
._.
PREFACE to the fourth edition vii
PREFACE to the first edition ix
Chapter I
INTRODUCTION 1
HISTORICAL
Chapter II
SCOPE AND METHOD 11
SCOPE; TRAINING; OBJECTIVES; METHOD; IMPORTANCE OF ECOLOGY;
CONTROL OF INSECT-BORNE DISEASES; VECTOR CONTROL; INSECTICIDAL
PRACTICES; VITAL L~ES DUE TO INSECTS; HOW MUCH TO SPEND ON
CONTROL
Chapter III
PARASITES AND PARASITISM 22
SYMBIOSIS AND PARASITISM; PARASITISM; CLASSES OF PARASITES;
EFFECTS OF PARASITISM; ORIGIN OF PARASITISM; SYSTEMATIC POSITION
OF ANIMAL PARASITES
Chapter IV
HOW ARTHROPODS CAUSE AND CARRY DISEASE 35
ARTHROPOD AND PATHOGEN; CHAIN OF INFECTION; PATHOLOGICAL CON-
DITIONS CAUSED BY ARTHROPODS; ENTOMOPHOBIA; ENVENOMIZATION;
DERMATOSIS; MYIASIS; ALLERGY CAUSED BY INSECTS; MECHANICAL
CARRIERS OF INFECTION; CYCLICO-PROPAGATIVE TRANSMISSION; CY-
CLICO-DEVELOPMENTAL TRANSMISSION; PROPAGATIVE TRANSMISSION;
FECAL CONTAMINATION; HEREDITARY TRANSMISSION; ARTHROPODS AS
INTERMEDIATE HOSTS OF HELMINTHS; RESERVOIR ANIMALS
Chapter V
STRUCTURE, DEVELOPMENT, AND CLASSIFICATION OF
INSECTS AND ARACHNIDS
THE INSECTA (HEXAPODA); EXTERNAL ANATOMY; WINGS; META-
MORPHOSIS; IMPORTANCE OF KNOWING INTERNAL ANATOMY; DIGES-
TIVE SYSTEM; INSECT LARVAE; THE ORDERS OF INSECTS; THE
ARACHNIDA; ARACHNID DEVELOPMENT; ARACHNID CLASSIFICATION.
KEY TO THE TERRESTRIAL ORDERS OF THE CLASS ARACHNIDA
xi
xii CONTENTS
Chapter VI
INSECT AND ARACHNID MOUTH PARTS 60
IMPORTANCE OF MOUTH PARTS; CLASSIFICATION OF MOUTH PARTS.
MORPHOLOGY OF MOUTH PARTS: ORTHOPTERON TYPE; THYSANOPTERON
TYPE; HEMIPTERON TYPE; ANOPLURON TYPE; DIPTERON TYPE;
SIPHONAPTERON TYPE; HYMENOPTERON TYPE; LEPIDOPTERON TYPE.
ORDERS OF INSECTS ARRANGED ACCORDING TO MOUTH PARTS. ARACHNID
MOUTH PARTS
Chapter VII
COCKROACHES AND BEETLES 78
COCKROACHES; FEEDING HABITS; LIFE HISTORY; COCKROACHES AS
VECTORS; COCKROACHES AS INTERMEDIATE HOSTS OF NEMATODE
PARASITES; COCKROACH CONTROL. BEETLES: THE ORDER COLEOPTERA;
CHARACTERISTICS; SCAVENGER BEETLES; BEETLES AS INTERMEDIATE
HOSTS OF HELMINTHS; CANTHARIASIS; VESICATING BEE~LES; ROSE
CHAFERS POISONOUS TO POULTRY; SUNDRY ANNOYING BEETLES; BEETLES
AS PARASITES. KEY TO FAMILIES OF COLEOPTERA
Chapter VIII
BUGS 94
ORDER HEMIPTERA. BEDBUGS; FAMILY CIMICIDAE; THE COMMON BED-
BUG; METHODS OF DISTRIBUTION; BEDBUG BITES; BEDBUG CONTROL.
KEY TO NORTH AMERICAN CIMICIDAE. THE CONENOSES: F AMIL Y REDU-
VIIDAE; LIFE HISTORY; THE ALIMENTARY CANAL OF Triatoma
protracta (UHL.); CONENOSE BITES; CHAGAS' DISEASE; TRANSMISSION
OF INFECTION; CONTROL. KEY TO SOME PREDACEOUS REDUVIIDAE
LIKELY TO BE OF MEDICAL IMPORTANCE. SOME OTHER BUGS. KEY TO
PRINCIPAL FAMILIES OF HEMIPTERA-HETEROPTERA OF NORTH AMERICA
WHICH CONTAIN PREDACEOUS SPECI,ES
Chapter IX
THE LICE 113
THE SUCKING LICE: GENERAL CHARACTERISTICS; CLASSIFICATION OF
ANOPLURA; THE LICE OF MAN AND OTHER PRIMATES. SUCKING LICE OF
MAN: THE HUMAN HEAD LOUSE; THE PUBIC LOUSE: THE BODY LOUSE;
DISSEMINATION OF BODY LICE: PEDICULOSIS: FAVUS AND IMPETIGO;
EPIDEMIC RELAPSING FEVER; TYPHUS FEVER; TRANSMISSION BY LICE;
MURINE TYPHUS FEVER; TRENCH FEVER; DELOUSING; LOUSICIDES. SUCK-
ING LICE OF MAMMALS: ANOPLURA AFFECTING DOMESTICATED
MAMMALS. THE BITING LICE: CLASSIFICATION; INJURY BY BITING LICE;
LIFE HISTORY; LICE INFESTING DOMESTIC FOWLS; CONTROL OF POULTRY
LICE; BITING LICE OF DOMESTICATED MAMMALS; CONTROL OF BITING
LICE ON MAMMALS; LICE AND TAENIASIS
Chapter X
GNATS (EXCLUSIVE OF MOSQUITOES) 143
ORDER DIPTERA; CLASSIFICATION OF THE DIPTERA; SOME FAMILIES OF
THE ORDER DIPTERA. FAMILY SIMULIDAE: CHARACTERISTICS; LIFE
CONTENTS xiii
lUSTORY; CLASSIFICATION; COMMON SPECIES; THE BITE; RELATION TO

DISEASE; BOVINE ONCHOCERCIASIS; LEUCOCYTOZOON INFECTION OF
POULTRY; BLACK-GNAT CONTROL. FAMILY PSYCHODIDAE: PSYCHODA
FLIES; PHLEBOTOMUS FLIES; LIFE HISTOt\y OF SAND FLIES; CARRION'S
DISEASE; SAND FLY FEVER; LEISHMANIASIS; SPECIES OF PHLEBOTOMUS.
FAMILY CERATOPOGONIDAE: CHARACTERISTICS. FAMILY CHIRONOMIDAE.
FAMILY DIXIDAE. FAMILY CHAOBORIDAE. FAMILY CHLOROPIDAE:
Hippelates FLIES; RELATION TO CONJUNCTIVITIS; RELATION TO YAWS;
BOVINE MASTITIS; LIFE HISTORY OF Hippelates pusio LOEW; CLASSIFI-
CATION; CONTROL OF HIPPELATES FLIES
Chapter XI
MOSQUITOES: CLASSIFICATION AND BIOLOGY 175
IMPORTANCE; MALE TERMINALIA; LIFE HISTORY; FLIGHT HABITS;
LONGEVITY; INTERNAL ANATOMY. TRIBE MEGARHININI: CHARACTERIS-
TICS. TRIBE CULICINI. TRIBE AEDINI. SALT-MARSH MOSQUITOES; FLOOD-
WATER Aedes; TREE-HOLE MOSQUITOES; BOREAL Aedes OR SNOW
MOSQUITOES; Aedes aegypti; OTHER GENERA. TRIBE ANOPHELINI:
CHARACTERISTICS; MATING AND OVIPOSITION; EGG CHARACTERS; BREED-
ING HABITS; BITING HABITS; SOME NORTH AMERICAN ANOPHELINES.
KEY TO CULICID TRIBES AND GENERA OF THE UNITED STATES
Chapter XII
MOSQUITOES AS VECTORS OF DISEASE 213
THE MALARIAS: HUMAN MALARIA; HISTORICAL; THE PLASMODIA; LIFE
CYCLE OF THE PLASMODIA; WHAT MAKES A MOSQUITO A GOOD NATURAL
VECTOR?; "ANOPHELISM WITHOUT MALARIA"; NUMBER OF PERSONS IN-
,FECTED BY ONE MOSQUITO; EFFECT OF TEMPERATURE ON PLASMODIA
IN THE MOSQUITO; HIBERNATING ANOPHELINES NOT CARRIERS; VECTORS
OF HUMAN MALARIA; MALARIA SURVEYS. ANIMAL MALARIAS: AVIAN
MALARIA; SIMIAN MALARIA; SAURIAN AND AMPHIBIAN MALARIA. YEL-
LOW FEVER: WALTER REED AND YELLOW FEVER; MECHANISM OF
TRANSMISSION; SUMMARY OF RECENT DISCOVERIES; JUNGLE YELLOW
FEVER. DENGUE FEVER: MOSQUITO TRANSMISSION. ARTHROPOD-BORNE
VIRUS ENCEPHALITIDES: EQUINE ENCEPHALOMYELITIS; JAPANESE "B"
VIRUS. FILARIASIS: MOSQUITO SPECIES INVOLVED; HEARTWORM OF DOGS;
OTHER FILARIAL WORMS. RABBIT MYXOMATOSIS. FOWLPOX
Chapter XIII
MOSQUITO ABATEMENT 256
HISTORICAL; ORGANIZATION FOR ABATEMENT WORK; PERSONNEL;
TRAINING OF PERSONNEL AND EDUCATION OF THE PUBLIC; MOSQUITO
SURVEYS; ESSENTIALS OF MOSQUITO ABATEMENT; WATER MANAGE-
MENT; IMPOUNDED WATER; CONTROLLED REFLOODING; FLUSHING;
SALINIFICATION; CREEKS AND SMALL STREAMS; DRAINAGE; DITCHING
WITH DYNAMITE; MAINTENANCE OF DITCHES; SALT-MARSH DRAINAGE;
FILLING AND PUMPING; MOSQUITO BREEDING IN RICE FIELDS; ANOPHE-
xiv CONTENTS
LINE CONTROL IN BROMELIADS; SEWER INLETS AND CATCH BASINS;

PUBLIC UTILITIES STREET VAULTS; CESSPOOLS, PRIVIES, LIQUID MANURE
PITS; TREE HOLES; CEMETERY URNS; SUNDRY NUISANCES; ROUTINE
INSPECTIONS FOR MOSQUITO BREEDING; LARVICIDES; RESIDUAL TREAT-
MENT; APPLICATION OF LARVICIDES; AIRCRAFT IN MOSQUITO CONTROL;
REPELLENTS; SCREENS TO EXCLUDE MOSQUITOES; DUCK CLUBS; MOS-
QUITO CONTROL AND WILD LIFE CONSERVATION; NATURAL ENEMIES
OF MOSQUITOES; TRANSPORTATION OF EXOTIC MOSQUITOES BY AIR-
PLANE
Chapter XIV
HORSEFLIES, DEER FLIES, AND SNIPE FLIES 297
HORSEFLIES: TABANIDAE; BREEDING HABITS AND LIFE HISTORY; BITES;
RELATION TO ANTHRAX; SURRA; TULAREMIA; LOIASIS; EL DEBAB; CON-
TROL; THE SPECIES OF TABANIDAE. KEY TO THE TABANID GENERA OF
NEARCTIC AMERICA. SNIPE FLIES
Chapter XV
HOUSEFLIES 316
ORDER DIPTERA, SUPERFAMILY MUSCOIDEA: HOUSE-INVADING FLIES;
FAMILY MUSCIDAE; THE TRUE HOUSEFLY; RANGE OF FLIGHT; LONGE-
VITY; OTHER HOUSE-INVADING FLIES; BLOWFLIES AND FLESH FLIES;
TUBERCULOSIS; EGGS OF PARASITIC WORMS; INTESTINAL PROTOZOA;
MURRINA; BOVINE MASTITIS; CUTANEOUS HABRONEMIASIS; FOWL
TAENIASIS. FLY CONTROL: RURAL; MANURE WASTAGE; MANURE DIS-
POSAL; COMPOSTING PITS; CLOSE PACKING; CHEMICAL TREATMENT OF
MANURE; DDT IN RURAL FLY CONTROL; COMMUNITY FLY CONTROL;
DDT-RESISTANT HOUSEFLIES; USE OF MANURE ON LAWNS; LAWN
CLIPPINGS; FLIES FROM SEPTIC TANKS; SEWAGE TREATMENT PLANTS;
PRIVIES; FLYTRAPS
Chapter XVI
BLOODSUCKING MUSCOID FLlES-Tsetse flies, Stomoxys flies,
Horn flies 352
TSETSE FLIES: GENUS Glossina; GENERAL CHARACTERISTICS; LIFE HIS-
TORY; TRYPANOSOMIASIS; Glossina SPECIES; TSETSE-FLY CONTROL.
STOMOXYS FLIES: GENERAL CHARACTERISTICS; HABITS; BREEDING HABITS
AND LIFE HISTORY; LONGEVITY; POLIOMYELITIS; CONTROL OF STABLE
FLIES. HORN FLY: CHARACTERISTICS; LIFE HISTORY; DAMAGE DONE;
CONTROL; OTHER SPECIES OF BLOODSUCKING MUSCOID FLIES
Chapter XVII
MYIASIS 372
ACCIDENTAL MYIASIS: IDENTIFICATION OF FLY MAGGOTS; INTESTINAL
MYIASIS; GASTRIC MYIASIS; URINARY MYIASIS; TRAUMATIC DERMAL
MYIASIS. SEMI-OBLIGATE MYIASIS. SUNDRY MAGGOT INFESTATIONS:
TUMBU FLY AND CONGO FLOOR MAGGOT; WOOL MAGGOTS; CONTROL OF
WOOL-MAGGOT FLIES; TOXIC EFFECT OF INGESTED FLY LARVAE; BLOOD-
SUCKING MAGGOTS OF BIRDS. OBLIGATE MYIASIS: OESTRID FLIES;
CONTENTS XV
HORSE BOTFLIES; CATTLE GRUBS; LIFE HISTORY AND HABITS; HEAD

MAGGOTS, GRUB IN THE HEAD, OR SHEEP BOTS; HEAD MAGGOT OF
HORSES, DEER; DERMAL CREEPING MYIASIS; OPHTHALMOMYIASIS;
RODENT BOTS; SURGICAL MAGGOTS
Chapter XVIII
LOUSE FLIES 410
HIPPOBOSCIDAE: CHARACTERISTICS; THE SHEEP "TICK" OR KED; LIFE
HISTORY; DAMAGE DONE; CONTROL; LOUSE FLIES OF DEER. GENUS Hip-
pobosca. LOUSE FLIES OF BIRDS; BAT FLIES
Chapter XIX
FLEAS 416
ORDER SIPHONAPTERA: CHARACTERISTICS; DIGESTIVE TRACT; LIFE HIS-
TORY; LONGEVITY OF FLEAS; HOSTS AND OCCURRENCE OF SPECIES;
PLAGUE; THE DISEASE IN MAN; FLEAS AS VECTORS; ROLE OF FLEA IN
PLAGUE TRANSMISSION; PLAGUE IN FIELD RODENTS; WILD RODENT
FLEAS; INFECTED AND INFECTIVE FLEAS; MURINE (ENDEMIC) TYPHUS
FEVER. THE COMMONER SPECIES OF FLEAS. KEY TO THE FAMILIES OF
SIPHONAPTERA; FAMILY PULICIDAE; FAMILY DOLICHOPSYLLIDAE;
FAMILY HECTOPSYLLIDAE; FAMILY HYSTRICHOPSYLLIDAE; THE CHIGOE
FLEA; WESTERN HEN FLEA; FLEAS IN THE HOUSEHOLD; FLEAS AS IN-
TERMEDIATE HOSTS OF CESTODES; FLEA REPELLENTS; RODENT CON-
TROL; METHODS OF RAT CONTROL; FIELD RODENTS
Chapter XX
TICKS AND TICK-BORNE DISEASES 464
HIGH VECTOR POTENTIAL OF TICKS; MAN NOT A NATURAL HOST FOR
TICKS; HISTORICAL; CHARACTERISTICS; LIFE HISTORY; LONGEVITY;
CLASSIFICATION. THE FAMILY IXODIDAE (HARD-BODIED TICKS): GENUS
Ixodes;GENUS Dermacentor; KEY TO ADULTS OF GENUS Dermacentor;
GENUS Haemaphysalis; GENUS Rhipicentor; GENUS Rhipicephalus;
GENUS Margaropus; GENUS Hyalomma; GENUS Amblyomma; GENUS
Boophilus; THE TEXAS CATTLE FEVER TICK; LIFE HISTORY. DISEASES
CARRIED BY IXODINE TICKS: TEXAS CATTLE FEVER; ROCKY MOUNTAIN
SPOTTED FEVER; TICK TRANSMISSION OF SPOTTED FEVER; INFECTION
IN NATURE; MECHANISM OF INFECTION; CONTROL AND PREVENTION;
COLORADO TICK FEVER; TICK TRANSMISSION OF TULAREMIA; TICK
PARALYSIS; BULLIS FEVER; Q FEVER; LYMPHOCYTIC CHORIOMENINGITIS;
EAST COAST FEVER; EQUINE PIROPLASMOSIS; CANINE BABESIOSIS
(PIROPLASMOSIS); HEARTWATER; BOVINE ANAPLASMOSIS. COMBATING
TICKS: TICKS ON LIVE STOCK; TO FREE DOGS OF TICKS; CONTROL OF
TICKS ON VEGETATION. FAMILY ARGASIDAE (SOFT-BODIED TICKS):
GENUS Ornithodoros; TICK-BORNE RELAPSING FEVER-IN THE UNITED
STATES; SPIROCHETES OF ENDEMIC RELAPSING FEVER; ERADICATION
OF INFECTION FROM SUMMER CABINS; GENUS Otobius; THE SPINOSE
EAR TICK; GENUS Argas; Argas persicus (OKEN); OTHER SPECIES OF
Argas; GENUS Antricola
xvi CONTENTS
Chapter XXI
MITES 525
CHARACTERISTICS. MANGE OR ITCH MITES: CHARACTERISTICS; SAR-
COPTIC MITES; HUMAN SCABIES; SWINE MANGE; EQUINE MANGE; TREAT-
MENT FOR EQUINE MANGE; BOVINE MANGE; CANINE MANGE; NOTOED-
RIC MANGE; OTHER MANGE MITES; SCALY LEG MITE ON POULTRY;
DEPLUMING MITE. FOOT AND TAIL MANGE: CHORIOPTIC AND SYM-
BIOTIC SCABIES. SCAB MITES; CHARACTERISTICS; OVINE SCABIES;
PSORPOTIC MANGE OF HORSES; BOVINE SCABIES. AURICULAR MITES.
FOLLICLE MITES-FOLLICULAR MANGE. MACRONYSSIDAE; TROPICAL RAT
MITES; TROPICAL FOWL MITE; NORTHERN FOWL MITE. THE FAMILY
Dermanyssidae. CHIGGER MITES: CHIGGER DERMATITIS. LOUSE-LIKE
MITES. FLOUR AND MEAL MITES, GROCER'S ITCH. RED SPIDERS. QUILL
MITES. AIR-SAC MITES. TONGUE WORMS; PENTASTOMIASIS
Chapter XXII
VENOMOUS AND URTICARIAL ARTHROPODS 563
INSECT VENOMS; HOW THE VENOM IS INTRODUCED; STINGING INSECTS;
MORPHOLOGY OF BEE STING; OPERATION OF THE STING; REACTION TO
BEE STINGS; TREATMENT FOR BEE STING; YELLOW JACKETS OR
HORNETS; MUD DAUBERS; ANTS; MUTILLID WASPS. BITING INSECTS:
CONENOSES OR KISSING BUGS; BITING WATER BUGS; BLOODSUCKING
PHYTOPHAGOUS BUGS; THRIPS BITING MAN; BLISTER BEETLES. SPIDERS:
TARANTULAS; DANGEROUS SPIDERS; THE BLACK WIDOW SPIDER; ARACH-
NIDISM-SPIDER BITE; NATURE OF VENOM; TREATMENT; CONTROL.
SCORPIONS; SCORPION STING. WHIP SCORPIONS. SUN SPIDERS. VENOMOUS
TICKS. CENTIPEDES AND MILLIPEDES
INDEX 611
MEDICAL ENTOMOLOGY
CHAPTER I
._.
INTRODUCTION
Historical. In the King James version of the Old Testament (Exodus

8: 24) we read, " ... and there came a grievous swarm of flies into the
house of Pharoh, and into his servants' houses, and into all the land of
Egypt: the land was corrupted by reason of the swarm of flies." The
Douay Version reads, "and the land was corrupted by this kind of flies."
Whether the term "flies" as used in this passage is coextensive with the
modern use of the word may be questioned, but it is interesting to con-
template the implications. As early as 1577 Mercurialis 1 '" expressed the
belief that flies carried the virus of plague from those ill or dead of the
disease to the food of the well. Although we now know that this is not the
usual mode of plague transmission, the principal role that flies playas
vectors of disease was correctly interpreted; i.e., they are food contam-
inators.
In 1587 Gabriel Soares de Souza 2 stated that flies suck poisons from
sores (Framboesia tropica) and leave them in skin abrasions on healthy
individuals, thus infecting many persons. In 1769 Edward Bancrofp ad-
vanced a similar theory, but it was not until 1907 that Castellani4 demon-
strated experimentally that flies do transmit Treponema pertenue Cas-
tellani, the causal agent of yaws. The role of the housefly as a vector of
disease was, however, not fully appreciated until 1898 when, as the result
of investigations made during the Spanish-American War, Veeder5 wrote:
I have mad~ cultures of bacteria from fly tracks and from the excrement
of flies and there seems to be not the slightest difficulty in so doing. Indeed
the evidence of every sort is so clear that I have reached the conclusion that
the conveyance of infection in the manner indicated is the chief factor in
decimating the army. Certainly so far as is known to the writer, nothing
adequate has been said about it in current discussions.
Although popular beliefs in many parts of the world had for some
time connected mosquitoes with various tropical fevers, no well-formu-
lated ideas were advanced until 1848, when Josiah Nott 6 of New Orleans
published his belief that mosquitoes gave rise to both malaria and yellow
~ Superior figures in the text refer to Bibliography at the end of each chapter.
1
2 MEDICAL ENTOMOLOGY
fever. In 1854 Beauperthuy/ a French physician in the West Indies,

formulated an excellent theory that mosquitoes were responsible for the
transmission of yellow fever, believing, however, that the unknown
disease factor was carried by the insect from certain decomposing matter
and in turn introduced by it into the human body.
While considerable was known by early naturalists and physicians
concerning the larger intestinal parasites, such as roundworms and tape-
worms, little information relating to microorganisms was available until
after the development of the microscope by Anton van Leeuwenhoek
( 1695),8 who found that his "material contained many tiny animals
which moved about in a most amusing fashion; the largest of these
showed the liveliest and most active motion, moving through the water
or saliva as a fish of prey darts through the sea." This discovery led to
the study of hitherto invisible organisms and eventually to the formula-
tion of the "germ theory" by Pasteur 9 in 1877.
Although according to Howard1o no standard medical treatise men-
tioned any specific disease as insect-borne prior to 1871, Raimbert1 '
showed in 1869 by experiment (inoculation of proboscides, wings, etc.,
of nonbiting muscids into guinea pigs) that anthrax (Bacillus anthracis
Cohn) could be disseminated by Bies, which as early as 1776 was be-
lieved by Montfils 12 to be the case. The first discovery of primary im-
portance in the field of medical entomology, however, was made in 1878
by Patrick Manson 13 who, working in China, observed the development of
Wuchereria (= Filaria) bancrofti (Cobbold) in the body of a mosquito,
Culex quinquefasciatus Say (= Culex fatigans Wied.), and eventually,
together with Bancroft, Low, and others, proved the mosquito to be the
intermediary host and vector of the causal organism of filariasis.
The discovery by Laveran 14 in 1880 of the causal organism of malaria
(Plasmodium malariae) living parasitically in the red blood cells of man,
marks an epoch in protozoology. Taking rank with Laveran's discovery of
the malaria parasite is the discovery by Theobald Smith in 1889 (Smith
and Kilbourne, 1893'5 of the causal protozoon organism (Babesia bi-
gemina) of Texas cattle fever, also living parasitically within the red
blood corpuscles of the host. Associated with Smith in the investigation
of the disease was F. L. Kilbourne, and together in 1893 they made the
second great fundamental discovery in the field of medical entomology,
namely that the cattle tick, Boophilus annulatus (Say) = (B. bovis
Riley), is the necessary intermediate host of the causal agent of the
disease. This knowledge, combined with Manson's discovery concerning
mosquitoes and filariasis, established a new basis for the control and
prevention of disease in both man and domestic animals.
In quick succession there followed a series of famous discoveries. In
1895 Bruce16 investigated nagana, the fatal tsetse-By disease of Africa
INTRODUCTION 3
(Zululand) and established the fact that the infection is conveyed from
animal to animal through the agency of Glossina morsitans Westwood.
In 1897 Ronald Ross"7 announced that he had found the zygotes of
the malaria parasite in two "dapple-wingled mosquitoes" (anophelines)
which had been bred from the larva and fed on a patient whose blood
contained crescents. In the discovery that mosquitoes carry malaria there
are linked the names of Ross; Manson;"8 MacCallum;"9 Bastianelli, Big-
nami, and Grassi;20 Koch;21 and Sambon and Low,22 the last two having
( 1900) demonstrated beyond a doubt the fact of transmission.
One of the world's outstanding achievements in the field of experi-
mental medicine is that of the United States Army Yellow Fever Com-
mission, consisting of Reed,23 Carrol, Lazear, and Agramonte, which in
1900 on the island of Cuba proved conclusively that yellow fever is car-
ried by a mosquito, Aedes aegypti (Linn.) [then known as Culex fasci-
atus Fabr. and later as Stegomyia fasciata (Fabr.)]. Carlos Finlay,24 a
Cuban physician, had as early as 1880 propounded the theory and con-
ducted experiments in an attempt to prove it; hence he, too, amply
deserves recognition and great praise.
These two discoveries concerning malaria and yellow fever gave great
impetus to the subject of mosquito control, although L. O. Howard had
already demonstrated the value of kerosene in his experiments in the
Catskill Mountains in 1892. Howard's pioneer book entitled Mosquitoes:
How They Live; How They Carry Disease; How They Are Classified;
How They May Be Destroyed, appeared in 1901.
During almost a third of a century following these fundamental dis-
coveries little advance was made in knowledge concerning the transmis-
sion of malaria and yellow fever, and the complete solution of the prob-
lem of control of both diseases seemed to be within reach-that is, simply
mosquito control. However, in 1937 malaria was again referred to as a
mysterious disease by Hackett 25 in his treatise Malaria in Europe vis.:
" . . . under close examination malaria became only more intricate and
impenetrable, more protean in its character, more diverse in its local
manifestations." The expression "anophelism without malaria" came into
use, and malariologists became more interested, as Hackett points out, in
the anophelines which did not transmit malaria than in those that did.
The discovery by Falleronj26 in 1926 that Anopheles maculipennis
Meigen, an important vector of malaria, was in reality separable into
races on differences in the' egg pattern, led Hackett, Martini, and Missi-
roli 27 (1932) to the discovery that the races of this species differ markedly
in their vector relationship to malaria, thus opening new avenues of re-
search. Now what appeared to be a clear-cut situation in 1898 became
once more a malaria "puzzle."
Furthermore, the apparently well-solved problem of yellow fever
control through the control of Aedes aegypti (Linn.) was again com-
pletely thrown open for further investigation by the discovery of Stokes,
Bauer, and Hudson 28 in 1927 that experimental animals (monkeys) can
be infected with yellow fever. Now, because of the availability of experi-
mental animals, more than a dozen species of mosquito instead of just
one species are known to have the ability to transmit the infection from
monkey to monkey by the bite.
In 1932 there was first observed in the Valle de Chanaan, Espirito
Santo, Brazil, a type of yellow fever designated as jungle yellow fever
( Soper, 1936) ,29 differing from the known type, transmitted usually by
Aedes aegypti (Linn.), only in that it occurs under conditions suggesting
that infection takes place away from urban habitations, that man may not
be an essential factor in the continuity of infection, indeed "man may be
but an accident in the course of an epizootic in the lower animals, 'or it
may even be due to the persistence of the virus in invertebrate vectors
for long periods of time."
In 1898 Simond 30 succeeded in transmitting plague from a sick rat to
a healthy rat through the agen'cy of infected fleas. This discovery was at
first discredited, but the experiments were successfully repeated by
Verjbitskpl in 1903 and Liston 32 in 1904,
The designation sylva tic (selva tic) plague has come into use par-
ticularly since 1928 (Ricardo Jorge 33 ) to specify plague of wild rodents
in which fleas play an important role as invertebrate reservoirs as well
as vectors.
At this juncture of our historical review of the subject it is appro-
priate to call attention to the first comprehensive treatise dealing with
arthropods as carriers of disease, namely the work of the late Professor
George H. F. Nutta1l 34 entitled, On the Role of Insects, Arachnids and
Myriapods as Carriers in the Spread of Bacterial and Parasitic Diseases
of Man and Animals. A Critical and Historical Study. Every student of
medical entomology should be familiar with this publication. The follow-
ing quotation from that work is significant:
Whilst hygienists have given much attention to the study of pathogenic
organisms in air, water, soil and food, their behavior under different chemical
and physical conditions, as also to the possibility of their direct or indirect
transmission from diseased to healthy individuals; relatively little attention
has been paid to one of the means by which infectious diseases are spread, to
the role played especially by insects, which may serve either as carriers or
intermediary hosts of disease-agents. The most thorough work in this direc-
tion has been done by parasitologists. Very few of the works on hygiene even
mention the role of insects as carriers of infection, and those that do, gener-
ally speak vaguely on the subject.
Nuttall deserves to be called the father of medical entomology.
INTRODUCTION 5
In 1901 Forde 35 observed certain parasites in the blood of persons
suffering from Gambian sleeping sickness, which Dutton 36 ,37 recognized
as trypanosomes and named Trypanosoma gambiense; and in 1903 Bruce
and Nabarro 38 showed that Glossina palpalis (Robineau-Desvoidy) was
the carrier, thus adding another tsetse-fly disease to the list. Stephens and
Fantham39 in 1910 described Trypanosoma rhodesiense as the causal
organism of Rhodesian sleeping sickness, and Kinghorn and Yorke 40 in
1912 proved Glossina morsitans Westwood to be the responsible vector.
Graham,41 while working in Syria in 1902, found that dengue, or
breakbone fever, a Widely distributed disease particularly of warm
climates, though frequently occurring elsewhere, is mosquito-borne. He,
and later Ashburn and Craig,<2 reported that possibly several species of
mosquitoes, notably Aedes aegypti (Linn.), are able to transmit the
infection. A closely related disease is pappataci fever, also known as
"three-day fever" and "sand fly fever," transmitted by Phlebotomus
papatasii Scopoli, as proved by Doerr, Franz, and Taussig43 in 1909.
In 1903 Marchoux and Salimbeni44 proved that fowl spirochetosis
caused by Spirochaeta gallinarum Blanchard is tick-borne and that Argas
persicus (Oken), the common fowl tick, is a vector. Another tick-borne
disease came to light when Dutton and Todd 45 and Ross and Milne 46
in 1904 discovered that African relapsing fever is carried by the tick
Ornithodoros moubata (Murray), the causal organism being Borrelia
(= Spirochaeta) recurrentis Lebert = (Spirochaeta duttoni N ovy and
Knapp). Furthermore, in 1906 Ricketts,<7 working in Montana (U.S.A.),
proved conclusively that ailOther tick, which he believed to be Derma-
center occidentalis Neum., but now known to be Dermacentor andersoni
Stiles [Dermacentor venustus (Banks)], is the principal vector of Rocky
Mountain spotted fever. Wolbach' s (1919) named the causal organism
Dermacentroxenus rickettsi.
Although lice have for centuries been associated with filth and disease,
apparently little thought was given these insects as possible carriers of
infection, even though Melnikoff49 had shown in 1869 that the biting
dog louse, Trichodectes canis DeGeer, was an intermediate host of the
double-pored dog tapeworm, Dipylidium caninum (Linn.), which also
occasionally occurs in humans. Aubert50 (1879), according to Nuttall,
considered that pediculi were spreaders of impetigo, and the cause of
prurigo, pityriasis, etc. Also, in experiments conducted by Dewevre 51 in
1892, lice were shown to carry the specific microorganisms mechanically
on their front legs, and infection was thus accidentally transmitted to
healthy persons. Furthermore, Fliigge 52 in 1891 and Tictin 53 in 1897
both supposed that disease might be carried by vermin and conducted
experiments with bedbugs. In 1907 Mackie 54 working in India found that
relapsing fever was transmitted by the body louse, Pediculus humanus
Linn., in whose body the causal organism, Borrelia (= Spirochaeta)
recurrentis Lebert (S. carteri Manson), multiplies.
Nicolle, Comte, and Conseil,55 working in Tunis in 1909, and Ricketts
and Wilder 56 working independently in Mexico in 1910, proved experi-
mentally that the body louse (Pediculus humanus Linn.) is a carrier of
typhus fever, the causal organism of which, Rickettsia prowazeki, was
described and named by Da Rocha-Lima 51 in 1916.
Members of the insect family Reduviidae (conenose bugs or kissing
bugs) have been long known for their fierce bites and bloodthirstiness,
but it was apparently not until 1909 that insects of this group were ex-
perimentally proved to be disease carriers. In that year Chagas,58 who
had already described the causal organism [Schizotrypanum (= Try-
panosoma) cruzi,] of Chagas' disease, also known as "Brazilian trypano-
somiasis," demonstrated that this disease is carried by the conenose bug
Mestor megistus (Burm.) [Triatoma megista (Burm.) = Panstrongylus
megistus (Burm.)]. In 1933 Kofoid and Donat59 showed that the try-
panosome of the conenose bug Triatoma protracta (Uhler) in California
is identical with that found in Mestor megistus (Burm.).
Flies of the family Tabanidae (horseflies, gadflies, earflies, deer flies,
etc.) were looked upon with suspiCion as early as 1776, but apparently
no satisfactory evidence against them was forthcoming until 1913, when
Mitzmain60 (Mayne), working in the Philippine Islands, demonstrated
transmission of surra of the carabao by Tabanus striatus Fabr., which he
regarded as the principal carrier. Strong evidence against tabanid flies
of the genus Chrysops as intermediary hosts of Loa (= Filaria) loa
(Cobbold) was advanced by Leiper,61 also in 1913.
Bloodsucking gnats belonging to the dipteron family Simuliidae are a
terrible scourge to both man and beast in many parts of the world and
have long been under suspicion as vectors of disease. In 1926 Blacklock62
reported Simulium damnosum Theob. as the vector of the filarial worm
Onchocerca volvulus (Leuckart), the causal agent of onchocercosis. In
1934 O'Roke 63 reported Simulium venustum Say to be the vector of a
disease of ducks caused by Leucocytozoon anatis Wickware.
Tularemia, also known as Pahvant Valley plague (Utah, U.S.A.) or
deer fly fever, was shown by Francis and Mayne 64 in 1921 to be carried
from rodent to rodent by the tabanid fly Chrysops discalis Williston, and
presumably from rodent to man in the same manner. The causal organism
of this disease, Pasteurella (= Bacterium) tularensis, was described in
1911 by McCoy and Chapin as the cause of a plague-like disease of
California ground squirrels. Though transmitted in nature by the deer
fly and several other species of arthropods, particularly the tick Derma-
centor andersoni Stiles, which is involved hereditarily, the infection is
most commonly contracted by handling infected rabbits.
INTRODUCTION 7
In 1933 Kelser 65 announced that he had succeeded in transmitting the
virus of equine encephalomyelitis from inoculated guinea pigs to a horse
by the bite of the mosquito Aedes aegypti (Linn.).
The discovery of the insecticidal value of the compound dichloro-
diphenyl-trichloroethane (DDT) represents the beginning of. a new era.
in the prevention of insect-borne diseases as pertains to both plants and
animals, particularly of such a devastating disease of man as malaria.
This compound was first synthesized by Othmar Zeidler 66 in 1874 at
Strasbourg, Germany. Zeidler apparently was ignorant of its insecticidal
properties. It was not until about 1939 that Paul Miiller, (Lauger et al. 67 )
then a member of the scientific staff of Geigy Company in Basle, Switzer-
land, discovered its remarkable insecticidal value. For this discovery
Miiller was awarded the Nobel Prize in Medicine for 1948.
Both historically and for future investigation the relation which the
protozoon subfamily Herpetomoninae of Castellani and Chalmers bears
to insects and their relation to animal and plant diseases, is one of interest
to parasitologists. Numerous insects are kn~n to harbor Leptomonas
(inclusive of Herpetomonas), Crithidia, Leishmania, and other genera,
SO,me of doubtful classification, but the problem of segregating those
which are zol:i- and phytopathogenic from those which are merely en-
tomoparasitic in exceedingly difficult and fraught with snares and pitfalls.
Apparently the earliest discovery in this connection concerning plants
was made by Lafont 68 in 1910 when he demonstrated that Leptomonas
davidi Lafont, the cause of "flagellosis" in three species of Euphorbiaceae,
required as its intermediary host the bug, Nysius euphorbiae Horvath.
BIBLIOGRAPHY
1. Mercurialis (Hieronymus), 1577 (?). De pestes in universum, praeser-
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2. de Souza, Gabriel Soares, 1587. Tratado descriptivo do Brazil em
1587, obra de Gabriel de Souza. Rio de Janeiro: Typographia universal de
Laemmert, 120 pp. (Cited by Fran<;a, C., in Tr. Roy. Soc. Trop. Med. & Hyg.,
15:58-60.)
3. Bancroft, Edward, 1769. An Essay on the Natural History of Guiana
in So. America. London: T. Becker and P. A. De Houdt, 402 pp.
4. Castellani, Aldo, 1907. "Experimental investigation on Framboesia
tropica (Yaws)," J. Hyg., 7:558-59.
5. Veeder, M. A., 1898. "Files as spreaders of sickness in camps," Med.
Rec., 54:429-30.
6. Nott, Josiah C., 1848. "On the origin of yellow fever," New Orleans
M. & S. J., 4:563-601.
7. Beauperthuy, L. D., 1854. "Transmission of yellow fever and other
diseases by mosquito," Gazeta aficial de Cumana, Ano. 4, no. 57, May 23.
(Cited by Howard, Dyer, and Knab.)
8. Leeuwenhoek, Anton van, 1695. Arcana naturae detecta ope micro-
scopiorum. Kronevelt: Delphis Batavorum, 568 pp. (Cited by Howard, Dyer,
and Knab.) See also The Select Works of Antony van Leeuwenhoek, translated
by Samuel Hoole, London, 1798.
9. Pasteur et Joubert, 1877. "Chimie Physiologique: Etude sur la maladie
charbonneuse," Compt. rend. Acad. d. sc., 84:900-906.
10. Howard, L. 0., 1921. "Sketch History of Medical Entomology," in
A Half Century of Public Health, M. P. Ravenel. New York: Amer. Pub.
Health Assn., pp. 412-38.
11. Raimbert, A., 1869. "Recherches experimentales sur Ie transmission du
charbon par les mouches," Compt. rend. Acad. d. sc., 69:805-12. (Cited by
Nuttall.)
12. Montfils, A. J., 1776. "D'une maladie fn3quente connue en Bourgogne
sous Ie nom de Puce maligne," J. de med., 45:500. (Cited by Nuttall.)
13. Manson, Patrick, 1878. "On the development of Filaria senguinus
hominis, and on the mosquito considered as a nurse," J. Linn. Soc., Zool.,
London, 14:304-11.
14. Laveran, A., 1880. "Note sur un nouveau parasite trouve dans de sang de
plusieurs malades atteints de'1ievre palustre," Bull. Acad. de med., Paris, 9:1235.
15. Smith, Theobald, and Kilbourne, F. L., 1893. Investigations into the
Nature, Causation, and Prevention of Texas or Southern Cattle Fever. Wash-
ington, D. C.: U. S. Dept. Agric. in Bur. Animal Indust. Bull., no. 1. 301 pp.
16. Bruce, David, 1895. Tsetse-Fly Disease or Nagana in Zululand:
Preliminary report. Durban: Bennett and Davis. (Cited by Nuttall.)
17. Ross, R, 1897. "On some peculiar pigmented cells found in two
mosquitoes fed on malarial blood," Brit. M.]., 2:1786-88.
18. Manson, P., 1898. "Surgeon-Major Ronald Rose: Recent investigations
on mosquito-malaria theory," Brit. M.]., 1:1575-77.
19. MacCallum, W. C., 1898. "On the haematozoan infections of birds,"
J. Exper. Med., 3:117-36.
20. Bastianelli, G.; Bignami, A. E.; Grassi, B.; 1898. "Coltivazione delle
semilune malariche dell' uomo nell' Anopheles claviger Fabr: Note pre-
liminare." Atti della Reale Accad. del Lincei, Nov. 28, p. 313. (Cited by Nuttall.)
21. Koch, R, 1899. "Ueber die Entwicklung der Malariaparasiten," Ztschr.
f. Hyg. u. Infektionskr., 32:1-24.
22. Sambon, L. W., and Low, G., 1900. "The malaria experiments in the
Campagna," Brit. M. J., 2:1679-82.
23. Reed, Walter, 1900. "The etiology of yellow fever," Philadelphia Med.
J., 6:790-96.
24. Finlay, Carlos J., 1881 et seq. Traba;os selectos. Havana: Republica de
Cuba, Secretaria de Sanidad y Beneficencia, 1912. xxxiv + 657 pp.
25. Hackett, L. W., 1937. Malaria in Europe: An ecological study. Oxford
University Press, London: Humphrey Milford. xvi + 336 pp.
26. Falleroni, D., 1926. "Fauna anofelica italiana e suo habitat' (paludi,
risaie, canali). Metodi di lotta contro la malaria," Riv. di Malariol., 5:553-93.
27. Hackett, L. W.; Martini, E.; and Missiroli, A.; 1932. "The races of
A. maculipennis," Am. ]. Hyg., 16:137-62.
INTRODUCTION 9
28. Stokes, Adrian; Bauer, J. H.; and Hudson, N. Paul; 1928. "The trans-
mission of yellow fever to Macacus rhesus: Preliminary note," /.A.M.A., 90:
253-54.
29. Soper, Fred L., 1936. "Jungle yellow fever: A new epidemiological
entity in South America," Rev. de hyg. e saude pub., 10:107-44.
30. Simond, P. L., 1898. "La propagation de la peste," Ann. Inst. Pasteur,
12:625.
31. Verjbitski, D. T., 1908. "The part played by insects in the epidemiol-
ogy of plague," J. Hyg., 8:162-208.
32. Liston, W. G., 1905. "Plague rats and fleas," J. Bombay Nat. Hist.
Soc., 16:253-73.
33. Jorge, Ricardo, 1928. Les faunes regionales des rongeurs et des puces
dans leurs rapports avec la peste. Paris: Masson et Cie. 306 pp.
34. Nuttall, G. H. F., 1899. "On the role of insects, arachnids, and myria-
pods as carriers in the spread of bacterial and parasitic disease of man and ani-
mals. A critical and historical study." Johns Hopkins Hospital Reports, 8:1-154.
35. Forde, R. M., 1902. "Some clinical notes on a European patient in
whose blood a Typanosoma was observed," J. Trop. Med., 5:261.
36. Dutton, J. E., 1902. "Trypanosoma in man," Brit. M. ]., 1:42.
37. ---, 1902. "Note on a Trypanosoma occurring in the blood of man,"
Brit. M. J., 2:881-884.
38. Bruce, D., and Nabarro, D., 1903. Progress Report on Sleeping Sick-
ness in Uganda. "Rept. Sleeping Sickness Comm., Roy. Soc. London," no. 1.
39. Stephens, J. W. W., and Fantham, H. B., 1910. "On the peculiar
morphology of a trypanosome from a case of sleeping sickness and the possi-
bility of its being a new species (T. rhodesiense)," Proc. Roy. Soc. London,
ser. B, 83:28-33.
40. Kinghorn, A., and Yorke, W., 1912. "On the transmission of human
trypanosomes by Glossina morsitans Westw., and on the occurrence of human
trypanosomes in game," Ann. Trop. Med., 6:1-23.
41. Graham, H., 1902. "Dengue: A study of its mode of propagation and
pathology," Med. Rec., 61:204-7.
42. Ashburn, P. M., and Craig, C. F., 1907. "Experimental investigations
regarding the etiology of dengue fever," J. Infect. Dis., 4:440-75.
43. Doerr, R.; Franz, K.; and Taussig, S.; 1909. Das Pappatacifieber.
Leipzig u. Wien.: Franz Deutiche. 166 pp.
44. Marchoux, E., and Salimbeni, A., 1903. "La spirillose des poules,"
Ann. Inst. Pasteur, 17:569-80.
45. Dutton, J. E., and Todd, J. L., 1905. The Nature of Human Tick
Fever in the Eastern Part of the Congo Free State, with Notes on the Distribu-
tion and Bionomics of the Tick. Liverpool School Trop. Med. in Memoir no.
17. 18 pp.
46. Ross, P. H., and Milne, A. D., 1904. "Tick fever," Brit. M. ]., 2:1453-
54.
47. Ricketts, H. T., 1906. "The transmission of Rocky Mountain spotted
fever by the bite of the wood tick (Dermacentor occidentalis)," J.A.M.A.,
57:358.
48. Wolbach, S. B., 1919. "Studies on Rocky Mountain spotted fever."
J. Med. Research, 41:1-193.
49. Melnikoff, H., 1869. "Ueber die Jugendzustiinde der Taenia cucme-
rina," Arch. f. Naturgesch., 25:62-69.
50. Aubert, 1879. "Les pous et les ecoles: Un point d'hygi(me scolaire,"
Reviewed in Ann. de Dermat. et Syph., 1880, 2 ser., 1:292-93.
51. Dewevre, 1892. "Note sur Ie role des pediculi dans la propagation de
l'impetigo," Compt. rend. Soc. de biol., 4:232-34.
52. Fltigge, C., 1891. Grundriss der Hygiene. Leipsig: veit & Co. x +
560 pp. (Cited by Nuttall.)
53. Tictin, J., 1897. "Zur Lehre vom Rtickfalltyphus," Centralbl. f. Bakt.,
1 Abt., 21:179-86.
54. Mackie, F. P., 1907. "The part played by Pediculus corporis in the
transmission of relapsing fever," Brit. M. /., 2:1706-9.
55. Nicolle, Charles; Comte, C.; et Conseil, E.; 1909. "Transmission ex-
perimentals du typhus exanthimatique par Ie pou du corps," Compt. rend.
Acad. d. sc., 149:486-89.
56. Ricketts, H. T., and Wilder, R. M., 1910. "The transmission of the
typhus fever of Mexico (tarbardilIo) by means of the louse (Pediculus vesti-
menU)," I.AM.A, 54:1304-7.
57. Da Rocha-Lima, H., 1916. "Untersuchungen tiber Fleckfieber," Miin-
chen. Med. Wchnschr., 63:1381-84.
58. Chagas, C., 1909. "Ueber eine neue Trypanosomiasis des Menschen,"
Mem. lnst. Oswaldo Cruz, 1:159-218.
59. Kofoid, Charles A., and Donat, F., 1933. "Experimenntal infection
with Tryanosoma cruzi from intestine of conenose bug Triatoma protracta,"
Proc. Soc. Exper. Biol. & Med., 30:489-91.
60. Mitzmain, M. B., 1913. "The mechanical transmission of surra by
Tabanus striatus Fabr," Philippine J. Sc., 8 (ser. B) :223-29.
61. Leiper, Robert R., 1913. "Metamorphosis of Filaria loa," Lancet, 1:5l.
62. Blacklock, D. B., 1926. "Development of Onchocerca volvulus in
Simulium dammosum," Ann. Trop. Med., 20:1-48 and 203-18.
63. O'Roke, Earl C., 1934. A Malaria-like Disease of Ducks Caused by
Leucocytozoon anatis Wickware. Ann Arbor: University of Michigan Press.
64. Francis, Edward, and Mayne, Bruce, 1921. "Experimental transmis-
sion of tularaemia by flies of the species Chrysops discalis," U. S. Public
Health Service, Pub. Health Rep., 36:1738-46.
65. KeIser, R. A., 1933. "Mosquitoes as vectors of the virus of equine en-
cephalomyelitis," I. Am. Vet. M. A, 82, n.s. 35:767-7l.
66. Zeidler, Othmar, 1874. "Verbindungen von Chloral mit Bram und
Chlorbenzol," Ber. d. Deutsch. Chem. Gesellsch., 7:1180.
67. Uiuger, P.; Martin, H.; and Muller, P.; 1944. "Dber Konstitution und
toxische Wirkung von nattirlichen und neuen synthetischen insektentotenden
Stoffen," Helvet. chimica acta, 27:892-928.
68. Lafont, A., 1910. "Sur la presence d'un Leptomonas parasite de la
classe des Flagelles dans Ie latex de trois Euphorbiacees," Ann. lnst. Pasteur
24:205-9.
CHAPTER II
SCOPE AND METHOD
Scope. The preceding brief outline of the historical development of

knowledge pertaining to the relationship of arthropods to diseases, para-
sitic infestations, and various disorders of man and his domesticated ani-
mals will in itself give the student some understanding of the scope of
medical entomology. The name, medical entomology, as a designation
for this field of knowledge apparently did not come into use until about
1909. 1 Even at that late date there were those in high places who ridi-
culed the importance of arthropods as vectors of disease in spite of im-
portant entomological discoveries relating to the transmission of plague,
malaria, and yellow fever. Today medical entomology is not only recog-
nized as a science in its own right, but also takes equal rank with some
of the older sciences contributing to the fields of public health, tropical
medicine, preventive medicine, and veterinary medicine.
Medical entomology may be defined as the science which deals with
the relation of insects, arachnids, and other arthropods to parasitic infes-
tations and diseases of man and beast: they may be causal agents, as in
scabies; or vectors, as in the malarias. The science is directly concerned
with the biology and control of the offending arthropods. It contributes
to the conservation of the public health and the health and well-being
of animals.
Training. The medical entomologist must be thoroughly trained in
general zoology; he must be particularly well trained in entomology and
arachnology; and his training must include protozoology, helminthology,
and bacteriology. He must be a paraSitologist in the truest and widest
sense. He must be well versed in field ecology. Many insect-borne diseases
of man and his domesticated animals are maintained in nature in wild
mammalian and other reservoir animals; hence the medical entomologist
must have a thorough understanding of vertebrate zoology and also a
wider knowledge of invertebrates (particularly fresh-water forms) than
is offered in the usual courses in zoology and entomology. Familiarity
with aquatic organisms is also helpful in determining public health im-
plications when these organisms are found in domestic water supplies.
Familiarity with insects attacking stored and dried foods, stored grain,
11
and foods in general is valuable. A knowledge of house-invading animals
is essential. Entomological training must emphasize immature stages
in the developmental cycle. Although mycology is not usually considered
essential, the author has derived much benefit from a course in that field
and recommends it to students of medical entomology. Much direct
benefit will be derived from many of the courses offered in medical,
veterinary, and public health curricula, such as general anatomy, general
physiology, epidemiology, pathology, histology, toxicology, hematology,
and coprology. If available, training in public health administration
should be acquired. A substantial knowledge of chemistry is essential.
In order to solve certain field problems involving arthropods, for example,
mosquitoes and flies, knowledge of such phases of engineering as pertain
to drainage and sewage disposal is useful. Familiarity with these last-
named subjects will enable the medical entomologist to cooperate in-
telligently with other medical investigators and with engineers or, on
occasion, to make intelligent use of the professional services of phy-
sicians, veterinarians, engineers, epidemiologists, and other experts and
specialists in the solution of complicated problems. For a more extended
consideration of the preparation required for a career as a medical
entomologist the student is referred to an article by the author in the
Journal of Economic Entomology for February, 1943. 2
Objectives. The aim of medical entomology is the pursuit of knowl-
edge concerning disease vectors and particularly concerning the control
and prevention of arthropod-borne diseases. There are many notable
examples of the service rendered by workers in this fertile field, such as
the control of malaria and yellow fever in Cuba and the Panama Canal
Zone,3 and the campaign against rats and fleas in San Francisco in 1907. 4
The latter resulted in the eradication of plague from that city, thus
averting the spread of this terrible scourge to the surrounding area.
The eradication during 1939-1940 of Anopheles gambiae Giles in Brazil
after its devastating introduction from tropical Africa is an excellent
recent example. 5 The benefit derived by animal industry is well illustrated
in the control of the tick vector [Boophilus annulatus (Say)] of Texas
cattle fever in the southern United States.
Method. In common with researchers in other related fields, the medi-
cal entomologist must employ the experimental method to advance his
science. 6 The experimental method is described by Thomas Hunt Morgan
( 1907) 1 in Experimental Zoology (by permission of The Macmillan
Company, publishers) as "the most important tool of research that
scientists employ. . . . The essence of the experimental method consists
in requiring that every suggestion (or hypothesis) be put to the test 6f
experiment before it is admitted to a scientific status. . . . It is the
method of attacking problems that is the chief characteristic of experi-
SCOPE AND METHOD 13
mental work. . . . We demand in the case of a problem in experimental
science that the conditions under which an event takes place be dis-
covered, and that, if possible, we reproduce artificially the result by con-
trolling the conditions. In fact the control of natural phenomena is the
goal of experimental work." Ivy (1948) 8 states, "It has been through the
experimental method of controlled or conditioned observation, and only
through this method, that scientists have discovered and will continue
to discover the most intimate secrets of Nature."
Since discovery of disease vectors and development of methods for
the control of arthropod vectors and reservoir animals are aims of medical
entomology, it is highly important that the experimental method be em-
ployed. An examination of the literature pertaining to arthropods and
disease shows that much of the work in this field is still largely in the
descriptive stage. Knowledge of taxonomy, morphology, anatomy, his-
tology, and physiology of arthropods is fundamental to sound work in
the field of medical entomology, but such knowledge alone, even of in-
sects of public health importance, does not per se qualify one as a medi-
cal entomologist. Referring to the importance of fundamental knowledge,
Morgan (loc. cit.) states: "The carrying out of an experiment implies
the formulation of a working hypothesis, and this usually presupposes
some knowledge of the possible conditions that control the phenomena.
The experimental work becomes more explicit and accurate the more
we know beforehand of the possible conditions that may enter into the
result. . . . "
In the experimental study of insect vectors one would ordinarily first
determine by experiment whether or not laboratory animals are sus-
ceptible to a certain disease. If they are, further experimentation is
greatly simplified. After long years of slow progress, rapid strides were
made in our knowledge of yellow fever when it was discovered that
laboratory animals such as monkeys and white mice could be used in
experimentation. 9 ,lD
Well does the author remember the experiment completed December
16, 1909, which proved DiamanU8 montanu8 (Baker) = (Ceratophyllus
acutu8 Baker), the ground-squirrel flea, to be a transmitter of plague
from ground squirrel to ground squirrel. The cage used by the United
States Public Health Service in San Francisco for the experimental ani-
mals was an ordinary galvanized iron garbage can 33 inches high by 19
inches in diameter, suitably screened and smeared at the rim with "tangle-
foot" to prevent the escape of the fleas. Every possible precaution was
taken to obviate danger. McCoy11 describes the experiment as follows:
A ground squirrel was inoculated subcutaneously with a broth culture

of Bacillus pestis derived from a human case of plague. This anima] died on
the fifth day. Three days before its death, one hundred fleas (Ceratophyllus
acutus) were put in the cage with it. While yet warm the dead rodent was re-
moved from the cage, and twenty-seven live fleas were taken from its body.
Two of these were crushed, and staining of the resulting smears showed an
abundance of pest-like bacilli in each. The twenty-five fleas remaining were
put in a clean cage with a healthy squirrel. This animal died of subacute
plague ten days later . . . . This experiment is conclusive in showing that
C. acutus may convey plague from a sick to a healthy squirrel. It should be
stated that all of the squirrels were kept in quarantine for at least a month
prior to their being used for the experiment. In fact, all of these squirrels
were obtained in a region in which no plague squirrels have ever been found.
Although human malaria was not known to be transmissible to labo-

ratory animals, knowledge of the vector was hastened because a related
malaria occurs in birds. Discoveries by MacCallum (lac. cit.) on bird
malaria furnished the key to the famous discoveries of Ross, Grassi,
Bignami, and others. Naturally induced (mosquito-transmitted) as well as
blood-induced malaria in the routine therapy of neurosyphilitics has
enabled investigators to make progress, heretofore impossible, in the
understanding of the behavior of Plasmodium vivax and other human
malaria plasmodia (Young, 1944) _12 Also the discovery of a malaria in-
fection in monkeys caused by Plasmodium knowlesi opened the way for
much experimental work applicable to human malariaY
Faust14 has aptly stated: "In most experimental work with human
parasitic infections laboratory animals can be utilized for all practical
experimental tests, thus obviating the need for any potential risk by
human volunteers. . .. Yet in certain crucial types of experimentation
it has been found highly desirable to know if human host-parasite rela-
tionships are directly parallel to those of susceptible animals." Less
dangerous diseases such as tick-borne relapSing fever have been given
human tests (see Chapter XX). Ivy (1948), lac. cit.) states: "In the
medical sciences, the only method which can clearly reveal and establish
the cause, prevention, and treatment of disease is the method of con-
trolled experimentation on animals and volunteer human subjects."
In experiments in the field of medical entomology an unusual amount
of care is required to prevent the escape of infected insects. Also much
skill and patience are needed to rear an adequate supply of living insects
in the laboratory for experimental use and to secure normal feeding
responses on the part of imprisoned insects.
Importance of ecology. The importance of ecological knowledge in
the investigation of insect-borne diseases has been long stressed by the
author. As early as 1909 (lac. cit.) he pointed out that "It is essential
that the student become familiar with the habits and habitat of the insect
in the field, its life history under normal and unusual conditions." Ecology
SCOPE AND METHOD 15
is variously defined, but few of the later proposed definitions define it
as well as did Haeckel (1869) when he described it as the "relation of
the animal to its organic as well as its inorganic environment." Chap-
man l5 said, . . . "he [Haeckel] considered oekologie to include the
general economy of the household of nature." In many instances man
will need to learn how to live in the same world with these now threaten-
ing members of that "household." Pearse 16 in a paper on the ecology of
parasites points out that "Man has succeeded by changing the environ-
ment or by changing his own characteristics as a habitat, in ridding him-
self of many of his parasites."
In dealing with the importance of ecology in relation to malaria,
Richard P. Strong17 has stated
In addition to these effects of the immediate environment upon the human
host, ecological studies must often consider its effects upon the intermediate
hosts in instances where they exist. Here, also, climate plays an important
role, not only in the character of the vertebrate fauna which the region har-
bors, but especially of the invertebrate fauna. Also, at temperatures below
a certain degree, the parasites in the insects which transmit them may be
unable to multiply or the insects satisfactorily to breed or even exist, as, for
example, the parasites and insects concerned in the transmission of sleeping
sickness and of malaria.
The epidemic of malaria with its high mortality which has recently been
raging in Ceylon, India, is a striking example of the effect that climatic con-
ditions and environment may exert upon a disease. This epidemic has occurred
in what has been hitherto regarded as the most healthy and prosperous por-
tion of the island, the southwestern part, in which there has usually been
a high annual rainfall and where there has been evidence that the percent-
age of the population infected with malarial parasites has been but small,
and hence the population relatively non-immune to the disease. This year
the prevailing rains which are brought so regularly by the southwest monsoon
failed to supply the usual amount of water, resulting in a prolonged drought.
Then came a few heavy rains and drought again. Thus conditions arose greatly
favoring the breeding of the mosquito. Anopheles culicifacies, which trans-
mits the disease in this region, as many shallow pools were formed along the
river beds and streams. Through these innumerable temporary breeding places,
more perfect conditions for the production of mosquitoes could probably
not have been devised. The outbreak of malaria was followed by failure of
the crops, also due particularly to the lack of rain. Thus the people became
further impoverished and the general state of their health reduced, and with-
in five months there were 113,811 deaths, of which 66,704 were estimated to
be due to malaria.
The complexity of ecological factors as pertains to medical ento-
mology is well shown in malaria, in which three animal species are in-
volved, namely, man the victim, the vector mosquito, and the causal
plasmodium-each species having characteristic ecological requirements.
The complexity becomes even greater when a reservoir animal enters
the epidemiological picture, as in endemic (tick-borne) relapsing fever
in California. In the cycle of this disease man is again involved, then
there is a tick vector, a spirochete as the pathogen, and a fourth species,
a chipmunk, as a reservoir. Meyer I8 has clearly portrayed the complexity
of an ecological approach to the study of plague in his De Lamar lecture,
entitled "The Ecology of Plague."
The student of medical entomology will do well to sturdy Uvarov'sl9
Insects and Climate, Martini's20 Wege der Seuchen, and Buxton's21
"The Effect of Climatic Conditions upon Populations of Insects." The
latter author (p. 326) remarks, " . . . the geographical spread of human
diseases and the seasonal occurrence of certain epidemics appear to be
directly due to alteration in the numbers of insects which are the essen-
tial vectors of these diseases. Our ultimate objective is to know the num-
bers of particular sorts of insects which are capable of infecting us with
the organisms which they carry."
Control of insect-borne diseases. The control of insect-borne diseases
involves not only the control of the responsible insect vectors, often very
difficult or even at times impossible, but depends also upon the control
of the fomites from which the arthropod receives its infection.
In the simplest form of insect transmission, i.e., by mechanical con-
tamination of food and drink, the source of infection may be found in
human excreta or other dangerous animal wastes, in which case the
possibility of spread by insects may be largely overcome by correcting
the defect in sanitation. Properly constructed fly-tight privies and septic
tanks would largely prevent the spread of typhoid fever and related filth
diseases by flies in rural areas. Rodent control is intended to destroy the
natural reservoirs of plague, as the flea is usually only an agent of trans-
mission. The control of plague through flea control alone does not satisfy
all requirements.
The handling of persons with infectious diseases is a matter of great
importance, as for example the screening of yellow fever patients against
the mosquito vector during the critical infectious stage of the disease.
Furthermore the proper screening of malaria patients against mosquitoes
is a factor in malaria control. Searching out and adequately treating.
carriers, while beset with many difficulties, should not be overlooked in
a program for control of insect-borne disease. There are situations where
vector control is not economically possible; for example, there are highly
malarial areas where mosquito control cannot be economically practiced,
yet certain agricultural procedures or construction work must go on. In
such cases suppressive (prophylactic) treatment is indicated.
SCOPE AND METHOD 17
In the face of universal rapid transportation by airplane and other
means, including automobile, and in the light of wartime experience,
health authorities of states and nations should be thoroughly familiar
with the vector potentialities of arthropods within their respective geo-
graphical boundaries (Herms, 1947).22 Vigilance at the point of depar-
ture and proper quarantine measures at the destination should be main-
tained to avoid the spread of arthropod-borne diseases. 23
Vector control. In the control of disease-transmitting arthropods the
most vulnerable point in the life history is usually sought and the most
effective control means are then employed. This involves an intimate
knowledge of biology. The more familiar one is with the life history,
habits, and ecology of the vector, the better equipped one will be to
cope with the problems of control.
Control measures may be only of a temporary nature for purposes of
immediate relief and hence must be repeated; or they may be intended
to have permanent effect. Temporary control measures involve holding
a nuisance in check for a short time, a few hours or a few days, and
require constant repetition: for example, the use of aerosols to kill adult
mosquitoes and flies, the use of mosquito repellents, or even the applica-
tion of oil to mosquito-breeding pools. Permanent control, on the other
hand, involves correction of breeding places by mechanical or other
means in order to prevent vector breeding: for example, draining or fill-
ing unnecessary ponds and pools of standing water in which mosquitoes
may breed; the correction of irrigation defects, particularly errors in
drainage; the disposal of manures and organic wastes in such a manner
as to prevent breeding of houseflies, e.g., close packing or immediate
spreading of manure. Permanent control measures are usually more eco-
nomical in the long run.
Species sanitation implies that control measures aimed at disease
vectors must fit the particular offending species. The importance of this
procedure is particularly applicable in vector-control operations where
closely related species or subspecies occur in the same general area but
only one is of public health importance. By concentrating appropriate
efforts on the proven vector species or subspecies, for example, in malaria
control operations in the presence of two or more anopheline species,
good results may be economically obtained.
Naturalistic control implies a planned change in the natural habitat
of an offending species, such as a disease vector, so as to make it im-
possible for the species to continue breeding in effective numbers. For
example, Anopheles albimanus Wied., a potent vector of malaria in the
Caribbean area, breeds abundantly in certain coastal lagoons having a
salinity between 15 and 25 per cent. By simply connecting these lagoons
with the sea so as to facilitate tidal action and thus increase the salinity
2. ---, 1943. "Preparation for a career as a medical entomologist,"

]. Econ. Entomol., 36:18-22.
3. Le Prince, Joseph A., and Orenstein, A. J. 1916. Mosquito Control
in Panama: The eradication of malaria and yellow fever in Cuba and Panama.
New York: G. P. Putnam's Sons. 355 pp.
4. Todd, F. M., 1909. Eradicating Plague from San Francisco: Report
of Citizens Health Committee. San Francisco: C. A. Murdock Co. Press. 313
pp.
5. Soper, Fred L., and Wilson, D. Bruce, 1943. Anopheles gambiae in
Brazil 1930 to 1949. New York: The Rockefeller Foundation. xviii + 262 pp.
6. Herms, William B., 1929. "The experimental method as applied to
entomological investigations," ]. Econ. Entomol., 22:45-59.
7. Morgan, Thomas Hunt, 1907. Experimental Zoology. New York: The
Macmillan Company. xiii + 454 pp.
8. Ivy, A. C., 1948. "The history and ethics of the use of human subjects
in medical experiments," Science, 108:1-5.
9. Stokes, Adrian; Bauer, J. H.; and Hudson, N. Paul; 1928. "The
tnm'Smlssion o~ ye\\ow \e"VeI to MucacuS' 'rhesus. Yle\imin....ry note," J.A.M .A.,
90:253-54.
10. Theiler, Max, 1930. "Studies on the action of yellow fever virus in
mice," Ann. Trop. Med., 24:249-72.
11. McCoy, G. W., 1911. "Studies upon plague in ground squirrels. Part
iv: Insect transmission in relation to plague among ground squirrels." Wash-
ington, D. C.; Govt. Print. Office, in Pub. Health Bull., no. 43, pp. 41-51.
12. Young, M. D., 1944. "Studies on the periodicity of induced Plasmo-
dium vivax," ]. Nat. Malaria Soc., 3:237-40.
13. Knowles, R., and Das Gupta, B. M., 1932. "A study of monkey
malaria and its experimental transmission to man," Indian Med. Gaz. 67:301-
20.
14. Faust, Ernest C., 1933. "The use of the experimental method in the
study of human parasitic infections," Scient. Monthly, 37:139-46.
15. Chapman, R. N., 1931. Animal Ecology, New York: McGraw-Hill
Book Co., Inc. 464 pp.
16. Pearse, A. S., 1926. "The ecology of parasites," Ecology, 7:113-19.
17. Strong, Richard P., 1935. "The importance of ecology in relation to
disease," Science, 82:307-17.
18. Meyer, K. F., 1942. "The ecology of plague," Medicine, 21:143-74.
19. Uvarov, B. P., 1931. "Insects and climate." Tr. Entomolog. Soc.
London, vol. 79, part 1, 247 pp.
20. Martini, E., 1936. Wege der Seuchen, Lebensgemeinschaft, Kultur,
Boden und Klima als Grundlagen von Epidemien. Stuttgart: Ferdinand Enke.
109 pp.
21. Buxton, P. A., 1933. "The effect of climatic conditions upon popula c
tions of insects," Tr. Roy. Soc. Trop. Med. & Hyg., 26:325-64.
22. Herms, William B., 1947. "Vector potentialities with respect to the
spread C?f insect-borne diseases of man in California," California Med., 67:95-
99.
SCOPE AND METHOD 21
23. Herms, W. M., 1946. "Wartime aviation quarantine: Pests and their
control," J. Pest Control lndust., 14:24-25.
24. Hunter, W. D., 1913. "American interelit in medical entomology
,. Econ. Entomol., 6, 27-39.
25. Fernald, H. T., 1926. Applied Entomolo§y. New York: McGraw-HilI
Book Co., Inc., xiv + 395 pp.
26. Carter, H. R., 1919. "The malaria problew of the South," U. S. Public
27. Herms, William B., 1919. "What shall ,'Ie do with our information
concerning malaria in California?" Calif. State Bd. of Health, Month. Bull.,
15:181-89.
CHAPTER III
. _-
PARASITES AND PARASITISM
Symbiosis and parasitism. Biologists are not agreed as to the definition

of symbiosis, but for our purpose it may be regarded as a condition of
conjoint life existing between different organisms, and in its most perfect
form the associated organisms or symbionts "are completely adapted to
a life in common," while on the other hand a poor adaptation to a sym-
biotic existence may lead to serious pathological reactions and even "to
the death of the organism that is invaded." This interpretation of sym-
biosis by Nuttall! differs from the usual definition, which denotes a con-
dition of conjoint life that is more or less beneficial to the associated
organisms.
Nuttall (lac. cit.) well said" ... it is difficult to imagine that sym-
biosis originated otherwise than through a preliminary stage of parasitism
on the part of one or the other of the associated organisms, the conflict
between them in the course of time ending in mutual adaptation." When
the symbiotic relationship is of benefit to both organisms (i.e., recipro-
cal) it may be termed mutualism. For example, the tiny staphylinid bee-
tles Xenodusa cava (Lec.) and Xenodusa montana (Csy.) secrete a fluid
which ants, Formica rufa Linn. and other species, suck from glandular
hairs; and in return for this favor the ants feed the beetles, which are
said to be unable to feed themselves and hence perish if unassisted by
the ants. Also the corn root aphid, Aphis maidiradicis Forbes, is cared
for in a most solicitous fashion, from egg to adult, by several species of
ants, Lasius sp., which feed on the so-called honeydew secreted by the
aphids.
When only one of the two organisms is benefited by the symbiosis, .
the relationship is known as commensalism. Thus a minute species of
cockroach, Attaphila fungicola Wheeler, is said by Wheeler2 (p. 397)
(reprinted by permission of Columbia University Press) to lick the sur-
faces, feeding on oily secretions of Atta ants, which tolerate the little
roaches, "without the slightest signs of hostility." No harm, of course,
results to the ants, and evidently no benefit is derived by them.
Parasitism. The definition of parasitism is well within the meaning of
the term symbiosis; however, the parasitologist instead of using the term
22
symbionts as applied to the associated species employs the terms host,
the physically larger of these, and parasite, the smaller in size, assuming
that the latter lives at the expense of the former. This conception of the
relationship of the associated species has led to a commonly used defini-
tion, viz.: parasitism is the state of one organism, the parasite, gaining
nourishment and other advantage, such as shelter, at the expense of
another living organism, the host. However, the latter (the host) must
not be destroyed as the result of this association before at least the
growth period of the former is completed, otherwise the result would
be disastrous to the parasite as well as the host. The life of a true parasite
is closely tuned to the life cycle and habits of the host. Essentially "a
paraSite is an organism which lives at the expense of its host, giving
nothing of value in return" (Stunkard, 1929 3 ). In spite of the fact that
the life of a parasite is commonly regarded as an easy one, it is rigor-
ously circumscribed and full of dangers. It is sometimes referred to as
a form of "hopeless speCialization," since it leads eventually to extinction,
unless conceivably the parasite were able to work back gradually from
parasitism to a free-living condition. On the other hand if parasitism is
but the beginning of an ill-adapted symbiotic life, then conceivably it
might gradually end through progression into a condition of mutual
adaptation. Indeed Nuttall (loc. cit.) has gone so far as to point out that
" . . . it is difficult to imagine that symbiosis [in the restricted sense,
that is, mutual adaptation] originated otherwise than through a prelim-
inary stage of parasitism on the part of one or other of the associated
organisms."
Swellengrebel4 describes a heavy malaria parasite incidence in the
blood of isolated bush Negroes in the interior of Dutch Guiana, yet with
no malarial fever (disease) resulting from this infection. He reports that
this high and constant malarial infection (mostly falciparum) leaves not
only the adult natives unaffected but does little harm even to the
youngest children. Swellengrebel himself suffered an infection while
among these bushmen after stopping "quinine prophylaxis," and in due
time (11 days) became ill with fa1ciparum malaria. The parasites, there-
fore, had lost none of their potential virulence when introduced in the
usual manner by mosquitoes into the body of a nonbushman.
Swellengrebel argues that this phenomenon in the case of these
isolated bush Negroes is the outcome of an interaction, a struggle, be-
tween host and parasite in which "both have had to sacrifice something,"
"a nicely adjusted equilibrium" having been achieved. "It is an adaptation
of the host to a greatly subdued parasite" which does no harm to its host
and consequently none to itself; it behaves as an "efficient parasite."
Swellengrebel points out that the title of his paper might perhaps better
have been "The Efficient Host."
There are many species ~f insects which are parasitic on other insects,
and some of these are very useful in holding certain agricultural pests in
check. Such "parasitoid" insects, as they are better termed, are often
reared in insectaries in enormous numbers to be liberated at a proper
time for control purposes. This practice is known as "biological ~ontrol."
While only of indirect concern to the medical entomologist, it is never-
theless a subject which will interest him in insect-control operations. The
"parasitism" of "parasitoids" results in the slow death of the parasitized
host insects. Predator insects kill their prey quickly, e.g., ladybird beetles
of the family Coccinellidae; also the Chrysopidae, commonly known
as aphid lions. These latter predators capture their prey and literally
eat it; while still others, such as the kissing bugs of the family Reduviidae,
suck the vital juices; but in all cases \he prey is killed quickly.
An interesting form of social parasitism occurs in social insects such
as ants and hornets, where one species lives in the colony of another
species which is "deluded" into feeding both the adults and progeny of
the invaders. It has been -suggested that this form of parasitism exists
among human societies as well. The extreme case, as suggested by Root 5
would appear to exist among certain wasps that construct no cells of
their own and do not hunt, but provide for the future of their progeny
by placing their eggs in the cells which have been constructed and
stocked with food by some other species of wasp, and in some cases the
egg or larva which originally inhabited the cell is bundled out to starve,
or is devoured.
Parasitism in its various forms is widespread in nature among many
. groups of animals from the Protozoa to the Chordata, but as Stunkard
(lac. cit.) has so well stated, " . . . few clearly apprehend that it com-
prises one of the most distinctive categories of animal existence, and
constitutes a discrete and characteristic phenomenon. Careful and critical
study of large numbers of parasite~ belonging to various classes of ani-
mals, has demonstrated that parasitism has certain basic and fundamental
. characteristics, no matter in what group of animals the parasitic habit
may have been developed. The same general tendencies are manifest,
similar attributes and relationships appear, and similar consequences
inevitably follow the adoption of the parasitic habit."
Classes of parasites. Animal parasites are known as zooparasites, while
parasitic organisms belonging to the plant kingdom are known as
phytoparasitt3s. Parasites which live either temporarily or permanently on
the outside of the body are termed ectoparasites, and animals of this
group are known as ectozoa, e.g., the lice. Parasites which live within
the body or its cavities are called endoparasites, the animals being called
entozoa, e.g., tongue worms. Obligatory parasites are forced to remain
during their entire life in or on the body of the host, e.g., both biting
and suckling lice. Facultative parasites are able to exist as free-living or-
ganisms, and they can also live as parasites, e.g., the larvae of various
species of blowflies and flesh flies. Adult female mosquitoes are example
of intermittent parasites which prey upon the host at times only and are
free-living during intervals between meals.
Many other forms of parasitism are recognized, such as transitory
parasitism, which is said to refer to cases in which the organism is
parasitic during only a part of its life history, e.g., botflies. When two
or more hosts are required for the development of an obligatory parasite,
that host in which the parasite reaches sexual maturity is termed the
primary or definitive host, while the other, or others if more than one
species is required, should be referred to as the secondary host or hosts.
This is illustrated in the life history of the malaria Plasmodium, which
reaches sexual maturity in the body of certain anopheline mosquitoes,
the primary or definitive host, while the human being is the secondary
host. The lung fluke, Paragonimus westermani (Kerbert), requires both
a melaniid snail (Melania libertina, etc.) and a fresh-water crab or cray-
fish (Astacus ;aponicus W. de Haan, etc.) as secondary hosts, while man
(and certain other mammals) is the definitive or primary host.
Effects of parasitism on host and parasite. Parasites are adapted to
their mode of life in two general respects: physiologically, in that they
require a certain host species or group of species as food; and mor-
phologically, in that their structure is modified to this end. These adap-
tations become the more intense and exclusive as this mode of life prog-
resses, i.e., the further back one goes in the history of a parasitic species,
the more nearly like its free-living relations it becomes. This accommoda-
tion is obvious in those species in which the young are free-living and
resemble the young of related wholly free-living species. The extreme
case of the cirriped "louse," Sacculina carcini (Rathke), is illustrative of
this recapitulation. The parasitic Cirripedes, fish "lice," show striking
structural adaptations to a parasitic mode of living: thus the first pair of
antennae may become well-formed hooks for holding fast and the first
pair of maxillipedes become suckers; the mandibles are transformed
into piercing organs, and the mouth becomes a sucking tube.
With the introduction of easily assimilable substances into the diges-
tive tract of the parasite, the digestive organs gradually become modi-
fied; and with the entire body in direct contact with "predigested food,"
as is the case with tapeworms and other intestinal parasites also, to
a certain extent, with fleshfly larvae, the outer covering of the parasite
becomes modified to allow direct absorption, and thus the digestive tract
becomes wholly unnecessary and may be entirely lost. However, modifi-
cation of the digestive tract is not restricted to parasites by any means.
The modification or loss of wings in wholly parasitic insects, such
as the Mallophaga, Anoplura, and certain Hippoboscidae, is commonly
said to be the result of the parasitic mode of life; however, this phenom-
enon is again not characteristic of parasitic forms alone, as all will
readily concede. There are numerous examples of wing reduction and
extreme modification of appendages in nonparasitic forms.
Keilin and Nuttall 6 report that while Pediculus humanus capitis
DeGeer, the head louse of man, and Pediculus humanus corporis
DeGeer, the body louse, can be distinguished on morphological grounds,
"typical capitis lose all their distinctive morphological characters when
raised experimentally on man under conditions which are favourable for
the propagation of corporis, and they acquire all the morphological
characters of corporis after four or more generations" (NuttaIF). Adap-
tations of each of these races to the host are apparently quickly acquired.
Typical capitis is more active at a lower temperature than is corporis
owing, no doubt, to the exposed environment on the scalp of the host,
while the latter is protected among the clothing. Capitis is smaller in size
and has stouter legs, hence can climb hairs more rapidly than can cor-
poris. Under a normal environment corporis imbibes larger meals at
longer intervals in conformance with the resting habits of the host, while
capitis with continuous opportunity for feeding takes a small amount of
food at short intervals. Furthermore, Nuttall (loc. cit.) remarks that,
"The effect of darkness no doubt is responSible for corporis possessing
longer and slimmer antennae and legs than capitis. The latter is more
exposed to light upon the head than is corporis beneath the clothing in
most instances. It is, of course, well known that arthropods inhabiting
dark places have longer antennae and legs than those living exposed to
light." Adaptation to the skin color of the host appears to be common.
Lice collected from the heads of the brown-skinned Gilbertese during
the allthor's investigations among these people on Fanning IslandS were
strikingly sooty in color. Rearing lice in pill boxes of different colors in-
side, as carried on by Nuttall,9 shows that change of color is rather
easily accomplished.
Nuttall's conclusions based on the observations cited above are very
interesting, namely, "There is little doubt in my mind that capitis is being
converted into corporis today in nature, and that the latter, wben man
has become hairless, will constitute a species whose birth we are wit-
nessing."
The wider adaptability of parasites, i.e., adaptation to different host
species as compared with different parts of the body of the same host, as
in Pediculus humanus Linn., is shown in the different races of Sarcoptes
scabiei (Linn.), the itch or mange mite of humans, swine, horses, and
other animals. While there appear to be specific differences in pigmen-
tation, cuticular markings, and chaetotaxy, they are not constant, though
some insist on the validity of these characters. Transfer from host to host
of different species can be more or less readily accomplished. Other
examples could be cited to illustrate this adaptability, such as certain
polymorphic mammalian tryanosomes, namely, T. brucei Plimmer and
Bradford, T. gambiense, and T. rhodesiense, as cited by Duke/o who
refers to these as phYSiological variants of a single species.
The relation, both as to behavior and structure, between the parasite
and the host becomes more perfect as the symbiosis grows more intimate.
The true parasite and its host represent a type of machine with all its
parts functioning coordinately, hence it is difficult to discuss the behavior
of the former without also dwelling on the behavior of the latter. The
stage in this relationship when the former cannot exist without the latter
is certainly reached in many instances, but one may well wonder whether
the reverse condition is ever actually achieved through the agency of
parasitism.
In the case of insects which suck the blood of human beings, one is
impressed with the large degree of tolerance that is manifested toward
those species which are wholly or largely dependent on man; no doubt
immunity is an important factor, and in turn one wonders just how far
the parasite has gone in making its blood lust less offensive. Thus several
extreme cases will illustrate what is meant. Gilbert Islanders with whom
the author spent some time during the summer of 1924 will reluctantly
give a few head lice on request, but prefer keeping them for festive
reasons. To an old timer the body louse is not so offensive as it is to a
tenderfoot.
The bite of Anopheles maculipennis Meigen is generally benign, while
the bite of Aedes dorsalis (Meigen), a common salt-marsh species, is
almost always viciously irritating. The former species has become closely
associated with man and is a potent vector of malaria, the latter is a
"wild" species and not a disease vector in nature. The bite of Triatoma
protracta (Uhler) is very painful to most persons, and one is inclined to
suggest that this species must moderate its bite before it can become a
successful disease vector. Besides adaptations of mouth parts for piercing
and bloodsucking, and moderation of venoms to lessen pain in the host,
there is also the factor which prevents blood coagulation. This factor is
particularly well developed in the bloodsucking h~minths such as leeches
and hookworms. Much needs to be done to explain the specific symptoms
caused by arthropod salivary fluids following insect and tick bites, such
as urticaria following the bite of Triatoma protracta (Uhler) and paraly-
sis following the bite of Dermacentor andersoni Stiles.
The study of parasitism has contributed much during the past few
years to the field of pathology Itnd clinical medicine. Disturbances re-
sulting from parasitism are usually of a specific nature, and hitherto un-
known causes of certain symptoms are now often readily attributable to
specifl.c parasite infestations. The student of medical entomology will
certainly consult Parasitism and Disease by Theobald Smithl l (Princeton
University Press).
Origin of parasitism. Parasitism is one of the ways in which organisms
acquire food and is only one of the avenues, though an important one,
that bring the arthropod into relation with man and other animals as
potential pathogenic agents. Scavenger insects with indiscriminate feed-
ing habits, for example, scavenger Hies and cockroaches, which may feed
on excrement, often become food contaminators and consequently may
be important factors in the dissemination of filth diseases such as typhoid
fever. Also predaceous arthropods, such as the black widow spider and
the conenose bugs, may attack man and inject venoms. While the female
black widow spider probably bites human beings only in self-defense or
in defense of her cocoon, certain reduviid bugs, such as Triatoma,
are said actually to suck the blood of sleeping persons, though many
other warm-blooded animals may also serve as hosts. It is but a short
step from sucking blood by tapping the body of a bedbug which has
fed on a person and tapping the body of the person directly. This pro-
cedure is followed at least in some instances by Triatoma protracta
( Uhler). Various species of bloodsucking arthropods secure blood meals
by tapping the abdomens of their blood-engorged associates.
Parasites may be restricted more or less completely to particular
species of host animals (host specifl.c); hence, the parasite must have
developed its parasitic habit after the host came into existence, i.e., para-
sitism must be a recently acquired habit on the part of a one-time free-
living organism. This becomes more apparent by a study of the life
history of the parasite; the earlier stages commonly point to a primitively
free-living existence. Perhaps the ancestors of a given group of present-
day parasites were attracted to waste food, offal, and exudations of cer-
tain animals. When the search for food became simplified, they began
living as messmates, or commensalists, or as scavengers; thus the asso-
ciation between the two species became closer and eventually the para-
sitic habit was established. This is borne out by a study of the nearest
allies of a given parasite, in which the gradation from the free-living
animal to the parasite may be traced. The very close structural similarity
between the free-living, wingless book louse, Troctes divinatoria (Mull.)
(a member of the order Psocoptera, family Psocidae), and a common
hen louse, Menopon gallinae Linn. (= M. pallidum Nitzsch) (a member
of the order Mallophaga), leads us to believe that the parasitic Mallo-
phaga have been derived from the primitive Psocidae. Knowing the
habits of the book louse, we can easily imagine how the practice of
parasitism might eventually have become established, i.e., from the eat-
ing of feathers, scales, and excretions off the animal to the eating of the
same on the animal as a host.
Degrees of parasitism may also be illustrated by examples from the
biting lice, Mallophaga, in which there are species having the power to
run freely and live for a considerable length of time off the host, e.g.,
Menopon gallinae Linn. (= M. pallidum N itzsch), the common hen louse,
while other related species have become entirely sessile, as in the extreme
case of the worm-like louse, Menopon titan Piaget, inhabiting the gular
pouch of the pelican. Among the fleas there are also good examples of
gradation in habit and structure: the human flea, Pulex irritans Linn.,
has developed remarkable springing power and is comparatively free to
move from place to place, while the mature female sticktight flea, Tunga
penetrans (Linn.), is quite sessile, holding fast to one spot much like a
tick.
The medical entomologist is continually pressed with question con-
cerning the origin of the present intimate relationship existing between
insects and causal organisms of diseases of higher animals. It is interest-
ing to know that Galli-Valerio 12 found that Herpetomonas pyrrhocoris
Zotta et Galli-Valerio lives part of the time in flagellate form in the
rotting blossoms of the meadow saffron, Colchicum autumnale, where it
is picked up apparently by the bug in the spring and is deposited in
nonflagellate form again in the blossom during the autumn by means of
the insect's excreta. This is believed to be a very primitive degree of the
adaptation of a saprophytic protozoon to an internal parasitic life in an
insect. The relationship existing between Leptomonas davidi, already
referred to, living in the latex of Euphorbia and carried by the bug
Nysius euphorbiae Horv., represents a step in advance.
Systematic position of animal parasites. Though parasitic animals are
found in other phyla, those of economic importance affecting man and
beast are included almost exclusively in the following groups:
a. Protozoa: unicellular animals; e.g., Endamoeba histolytica (Schau-

dinn), causing amoebic dysentery; Plasmodium vivax ( Grassi and
Feletti), causing malaria; Trypanosoma gambiense Dutton, causing
African sleeping sickness.
b. Nemathelminthes: bilateral, unsegmented worms of cylindrical
form; e.g., Trichinella spira lis (Owen), causing trichinosis; Ascaris lum-
bricoides Linn., roundworm of man; Ancylostoma duodenale (Dubini),
a hookworm of man. Development is usually direct (Fig. 1).
c. Platyhelminthes: bilateral worms; flattened dorsoventrally; no anal
opening. Usually requiring an intermediate host.
1. Cestoda: scolex with separable segments called proglottids;
e.g., Taenia solium Linn., the pork tapeworm of man; Taenia
FIG. 1. Examples of para-
sitic roundworms (Phylum
N em a thelmin thes, Class
Nematoda). (A) roundworm
of swine (Ascaris lumbri-
coides); (B) Trichinella
spiralis, greatly enlarged.
IA IB
FIG. 2. Examples of parasitic flat-

worms (Phylum Platyhelminthes, Class
Cestoda). (Left) a poultry tapeworm
(Choanotaenia infundibulum , X 1) , and
(right) a common tapeworm of sheep
(Moniezia expansa, greatly reduced).
2
3 4 5
FIG. 3. Example of parasitic flatworms (Phylum Platyhelminthes, Class
Trematoda). A liver fluke of sheep (Fasciola hepatica). X 1.
FIG. 4. Example of segmented cylindrical worms (Phylum Annelida, Class
Chaetopoda) . Earthworm (Lumbricus sp., X .5) nonparasitic, but may serve
as an intermediary host for certain poultry tapeworms.
FIG. 5. Example of segmented cylindrical worms (Phylum Annelida, Class
Hirudinea). Leech (Hirudo medicinalis). X .5.
30
saginata Goeze, the beef tapeworm of man; Dipylidium caninum
(Linn. ), a common tapeworm of the dog (Fig. 2).
2. Trematoda: alimentary canal branched; mouth in a sucker;
e.g., Fasciola hepatica Linn., the liver fluke of sheep (Fig. 3).
d. Annelida: bilaterally symmetrical, segmented or annulated worms.
1. Chaetopoda: locomoter chaetae; segmentation extending to in-
ternal organs; e.g., Lumbricus terrestris Linn., a common earth-
worm (nonparasitic) (Fig. 4).
2. Hirudinea: flattened; sucker at each end of body; arrangement
of internal organs does not correspond to external segmenta-
tion; e.g., Hirudo medicinalis Linn.,. the medicinal leech (Fig.
5 ).
e. Arthropoda: segmented body with paired jOinted appendages;
chitinous exoskeleton; bilaterally symmetrical; heart dorsal; ventral nerve
cord.
1. Crustacea: head and thorax often united to form a cephalo-
thorax; numerous paired, biramous appendages; two pairs of
antennae; respiration usually branchial; habitat usually aquatic;
e.g., shrimp, crayfish, and the sow bug (the latter terrestrial).
These examples are nonparasitic (Fig. 6).
2. Onychophora (Protracheata): vermiform and externally unseg-
mented; numerous paired, imperfectly segmented legs; one pair
of antennae; tracheal respiration; habitat terrestrial; e.g.,
Peripatus (nonparasitic) (Fig. 7).
3. Chilopoda: body flattened dorsoventrally; one pair of jointed
legs on each segment except first two and last one; one pair of
antennae; eyes simple; genital pore posteriorly located as in
insects; habitat terrestrial; e.g., centipedes (nonparasitic) (Fig.
8a).
4. Diplopoda: cylindrical in form; two pairs of jointed legs on
each segment except first two and last one; simple eyes; one
pair of antennae; genital pore anteriorly located; habitat terres-
trial; e.g., millipedes (nonparasitic) (Fig. 8b).
5. Symphyla: very small; body flattened dorsoventrally; one pair
of jointed legs per segment and not more than 12 pairs in all;
spinnerets present at posterior end; eyeless; one pair of anten-
nae; genital pore anterior; e.g., garden centipedes (non-
parasitic) .
6. Pauropoda: minute; body flattened; nine, seldom ten, pairs of
jointed legs; antennae branched; eyes generally absent; e.g.,
pauropods (nonparasitic). •
7. Insecta or Hexapoda: body divided into three pairs (head,
thorax, and abdomen); three pairs of walking appendages on
FIG. 6. Examples of the

Ph YI u m Arthropoda, Class
Crustacea. (A) shrimp, X 1.2;
(B) crayfish, x.6; (C) sow-
bug, X 2. (All three examples
are nonparasitic.)
thorax; two pairs of wings on thorax (may be reduced or

absent); one pair of antennae; compound eyes; usually three
simple eyes; trachea ted respiratory system; e.g., Triatoma pro-
tracta (Uhler) (conenose bug); Cimex lectularius Linn. (bed-
bug); Anopheles maculipennis Meig. (malaria mosquito) (Fig.
9).
8. Arachnida: head and thorax fused to form cephalothorax; four
pairs of walking appendages on cephalothorax (larvae may be
7 8
FIG. 7. Example of Phylum Arthropoda, Class Onychopora. Peripatus.
x.5. (After Folsom.)
FIG. 8. Examples of the Phylum Arthropoda: (left) Class Chilopoda, a
centipede, X .5; (right) Class Diplopoda, a millipede, X .7.
PAHASITES AND PAHASlTISM 33
hexapod); wingless; no antennae; eyes simple, when present;
e.g., Latrodectus mactans (Fabr.), a poisonous spider; Hadrurus
hirsutus (Wood), scorpion; Boophilus annulatus (Say) , the
9 10
FIG. 9. Examples of the Class Insecta. (A) a reduviid (conenose) , Xl;

(B) a mosquito (Anopheles), x2; and (C) bedbug (Cimex), x2.S.
FIG. 10. Examples of the Phylum Arthropoda, Class Arachnida. (A) a
spider, x.5; (B) a tick, x1.3; and (C) a mite, x30.
Texas fever tick; Dermanyssus gallinae (DeGeer) , the poultry

mite; Psoroptes communis val'. ovis (Hering), the sheep scab
mite (Fig. 10).
BIBLIOGRAPHY
1. Nuttall, George H. F., 1923. Symbiosis in Animals and Plants . Section
1. Physiology. Liverpool: British Association. 18 pp.
2. Wheeler, W. M., 1926. Ants: Their Structure, Development and Be-
havior. New York : Columbia Univ. Press. xxv + 663 pp.
3. Stunkard, Horace W., 1929. "Parasitism as a biological phenomenon,"
Scient. Monthly, 28 :349-62.
4. Swellengrebel, N. R., 1940. "The efficient parasite," Science, 92:465-
69.
5. Root, Francis M., 1924. "Parasitism among the insects," Scient .
Monthly, 9:479-95.
6. Kellin, D ., and Nuttall, G. R. F., 1919. "Hermaphroditism and other
abnormalities in Pediculus humanus," Parasitology, 11 :279-328 (7 plates) .
7. Nuttall, G. H. F., 1919. "The systematic position, synonymy, and
iconography of Pediculus humanus and Phthirus pubis," Parasitology, 11 :329-
46.
8. Herms, W. B., 1925. "Entomological observations on Fanning and
Washington Islands, together with general biological notes," Pan-Pacific
Entomologist, 2:49-54.
9. Nuttall, G. H. F., 1919. "The biology of Pediculus humanus,"
Parasitology, 11:201-20 (1 plate).
10. Duke, H. L., 1923. "Further inquiries into the zoological status of
the polymorphic mammalian trypanosomes of Africa," Parasitology, 15:358-95.
11. Smith, Theobald, 1934. Parasitism and Disease. Princeton Univ. Press.
xiii + 196 pp.
12. Galli-Valerio, B., 1920. "Le cycle evolutif probable de' I'Herpeto-
monas pyrrhocoris, Zotta & Galli-Valerio," Schweiz. med. Wchnschr., 1:401-
2. (Cited in Rev. Applied Entomol. Med. & Vet., 1922, 10 (ser. B, part 1) :8.
CHAPTER IV
.- .
HOW ARTHROPODS CAUSE AND
CARRY DISEASE
Arthropod and pathogen. When one considers the fact that man and
his domesticated animals are daily closely associated with scores of
species of insects'" and their kin, the wonder is that there are not more
arthropodan infestations and insect-borne diseases. Many of the one-
time free-living arthropods have in time become parasitic; some now
burrow into the skin, as do certain mites (ac'ariasis); others have invaded
the alimentary tract, as do larvae of botflies (myiasis); still others have
become bloodsuckers, as are bedbugs, sucking lice, horseflies, mosqui-
toes, etc. Bloodsuckers, by virtue of their bloodsucking habit, may readily
become vectors of pathogenic blood-inhabiting microorganisms. It has
been well said that no bloodsucking arthropod can be trusted; eventually
many more species will prove to be vectors of disease.
The medical entomologist must acquaint himself with the detailed
biology of the disease-producing microorganisms as well as with that of
the vectors. In order to know where and how the arthropod acquires
the pathogenic agent and how it, the vector, becomes infectious, the be-
havior of such organisms must be studied; their habitat in the body of
the diseased host or reservoir must be known; the gateways of escape
must be ascertained; their longevity and virulence when removed from
- the host as well as many other pertinent factors must be determined.
After having acquired the pathogenic organism, its course within the
body of the arthropod must be studied in order to know how escape is
effected and how it reaches the body of the next host. A knowledge of
the feeding habits and mouth parts of arthropods is essential. (See
mechanism of infection, p. 36.)
Bubonic plague, for example, is a bacillary disease caused by Pas-
teurella pestis, of which the rat, among other animals, is an important
host. Man readily succumbs to the infection. While the pathogenic or-
ganisms may pass from host to host in several ways, it has been found
"Z. P. Metcalf estimates the number of species of insects alone, described for
the whole world between 1758 (Linnaeus) and 1940, at 1,500,000. (Entomological
News, 1940, 51:219-22.)
35
that rat fleas are the most important vectors. The bacilli are found in
great abundance in the buboes which are situated largely in the axillary
and inguinal regions of the infected rat, and it has been observed that
these regions are favored by fleas, which, because of their bloodsucking
habits, imbibe the highly infectious fluids. If the rat dies of the plague,
the fleas leave the body and seek another host; this interval in the change
of hosts raises the question of environmental resistance and longevity of
the bacilli. Do all species of rat fleas lend themselves equally well as
hosts? Can the bacilli resist the digestive fluids of the flea, and if so,
how long do they remain infective in the body of the flea? In the mean-
time what course do these organisms take within the body of the insect,
and how do they make their escape? How does the flea find another host,
and what animals will it attack? How long can the flea live without food?
How are the plague bacilli introduced into the body of the next victim?
Will the flea remain infective throughout the rest of its normal life ? What
is the length of life of a flea, and does this vary with the species?
Malaria of man, another example, is caused by certain species of
plasmodia which live under far more restricted conditions than do the
plague bacilli. In the human being these invade the red corpuscles part
of the time and so far as is known do not live in the blood of other warm~
blooded animals. They require certain mosquitoes belonging to the genus
Anopheles as definitive hosts. Although many other bloodsucking in-
sects imbibe parasitized blood, these do not offer the necessary environ-
ment within the alimentary canal for the completion of the life cycle of
the parasite.
Chain of infection. For each arthropod-borne disease there is a chain
of events which leads to the successful transfer of an infection by an
arthropod vector from one animal to another. This chain consists of
several links, each more or less vulnerable and hence subject to rupture,
which could prevent successful transfer of the infection. The first link in
this chain is the developmental cycle of the pathogen within the body
of an appropriate host. This link may be broken by medicinal treatment,
as in malaria. The second link in the infection chain involves the vector,
its access to infection, its feeding habits, its manner of ingesting food, the
structure of its mouth parts, etc.; the third link represents the progress of
development of the pathogen (also more or less complicated) within
the body, stomach, or other organs of the vector; the fourth link repre-
sents the manner in which the now infectious microorganism leaves the
body of the now infectious vector and thereupon enters the body of the
next susceptible animal, thus resulting in infection-specifically it is the
mechanism of infection, e.g., (1) simple contamination by means of
germ-laden houseflies or cockroaches coming in contact with food, milk,
etc.; (2) simple mechanical inoculation of anthrax bacilli by m"'eans of
ARTHROPODS AND DI~EASE 37
contaminated horsefly mouth parts, or inoc\ilation of sexually mature
malaria plasmodia by an Anopheles mosq\iito; (3) regurgitation of
plague bacilli upon the skin by fleas while btting after a proventricular
blockage; (4) fecal contamination where trypa_nosomes of Chagas' disease
are deposited on the skin when the conenos e bug bites; (5) skin con-
tamination with crushed tissues of infectioul) arthropods, e.g., crushed
infected ticks may result in spotted fever inf~ction.
The terminal link in the infection chain i~ the successful completion
of a cycle similar to that of the first link but tn. the body of another host
animal after the intervention of an arthropo<l vector. Other links in the
chain of infection as well as other variations in the mechanism of infec-
tion will occur to the student as he progresses in the field of medical
entomology.
Pathological conditions caused by arthrollods. The ways in which
arthropods relate to the public health and t4e health and well-being of
animals are almost as numerous as the species of responsible arthropods
('iRnnsmves; Eat-'lf :>pcR-'lt%; lruwevm- :>'limikt- rtf I('S next of KriI, lIas certain
characteristics which affect the host differently.. It is nevertheless possible
to classify the relationships of arthropods to health, putting them into
two divisions, each of which is subject to fUHher segregation. The two
major relationships may be designated as. (,,\) pathological conditions
caused directly by arthropods, and (B) artlu'opods as vectors of patho-
genic organisms. The first. major division (A) includes such categories
as (1) entomophobia, (2) accidental injury tG sense organs. (3) enveno-
mization, (4) dermatosis, (5) myiasis, (6) allergy. The second major
division (B) may be subdivided into (1) a<xidental carriers, and (2)
obligatory vectors involving some degree of development (cyclical or
propagative) within the arthropod.
As our subject develops in the remaining chapters of this book, the
student will classify the way or ways in whi(~h the particular groups of
arthropods fit into the scheme presented in tltis chapter.
Entomophobia. Insects and spiders, even though they may be wholly
innocuous, frequently cause man acute annoYance and worry which may
eventually lead to a nervous disorder with s~nsory hallucinations. 1 This
class of mental disturbances may perhaps de~erve a specific deSignation
such as entomophobia.
The author has observed two or three cas,es of this nature each year
for many years. The cases are usually referreq to him by physiCians, pri-
marily because the patient is assumed to h qve confidence in one who
has knowledge of the control of tormenting ~nsects. The patient usually
imagines that insects of very minute size persist in threatening him and
biting him, usually at particular sites of the l,ody, and that they achieve
this contact in the most cunning ways in spite of every precaution.
S8 MEDICAL EN1'OMOLOGY
Accidental injury to sense organs. Various species of insects, like other
minute objects, may accidentally enter the eye; this is most likely to be
true of small flying insects. Some of these, notably several species of
rove beetles, Staphylinidae, cause extreme pain because of an irritating
secretion. Many species of insects discharge odoriferous fluid or vapor,
and in some instances the fluid is sO forcibly ejected that it may be
thrown some distance from the insect. Stewart (1937) 2 records the case
of a phasmid walking stick, Anisomorpha buprestoides (Stoll), squirting
fluid a distance of two feet and striking a person in the eye. "The pain
in the left eye was immediately excruciating; being reported to be as
severe as if it had been caused by molten lead. Quick, thorough drench-
ing with cool water allayed the burning agony to a dull, aching pain.
The pain eased considerably within the course of a few hours. Upon
awaking the next morning the entire cornea was almost a brilliant scarlet
in color and the eye was so sensitive to light and pressure for the next
forty-eight hours that the patient was incapacitated for work. Vision was
impaired for about five days. The corIlea gradually cleared of congestion
and vision improved so that at the end of this time the eye was perfectly
normal again. No subsequent ill effects were experienced." Stewart re-
marks that, "In Texas there is a rather common belief that devil's horses
will spit in human eyes."
Injury to the human eye is often caused by the spiny larva of the
sheep botfly or head maggot fly, Oestrus avis Linn. This fly deposits liv-
ing young in the nostrils of sheep, and persons working in the field with
sheep are occasionally "struck" in the eye by the fly and one or more
larvae may be deposited. This indicates that the fly may "strike" sheep
Similarly at times.
Insects commonly enter the ears of sleeping persons and may cause
much pain before they are removed. Thus in the author's experience a
carpet beetle larva (Dermestes sp.) was taken from the ear of an elderly
man, and a hyaline grass bug, Corizus hyalinus (Fabr.), family Coreidae,
from the ear of a child, having caused much pain in each case. The
spinose ear tick, Otobius megnini (Dtlges) also invades the ears of man
and many animals.
Envenomization. Venoms of insects cause reactions which are remark-
ably characteristics. The effects may be classified under the following
headings: (1) hemolytic, (2) hemorrhagic, (3) neurotoxic, (4) vesi-
cating. The venoms are introduced in the following ways: (1) by the
bite, as of conenose bugs and black widow spiders; (2) by the sting, as
of bees, wasps, scorpions; (3) by urticating hairs, as with the brown-
tail moth; and (4) by contact, as with vesicating fluids from blister
beetles. (See Chapter XXII. )
Dermatosis. Various skin irritations are caused by arthropods, either
ARTHROPODS AND DISEASE 39
by bites or by skin invasions. Some of these irritations could be appro-

priately classified as envenomizations. Skin irritations commonly result
from the bites of such insects as mosquitoes, fleas, lice, and bedbugs.
Various species of burrowing mites cause skin irritations known as
acariasis. Among the latter are the itch mites, Sarcoptes scabiei (Linn.);
the scab mites, Psoroptes communis Hering; the follicle mites, Demodex
folliculorum Simon; the chigger mites, Eutrombicula alfreddugesi (Oude-
mans), and other trombiculids.
Myiasis. An invasion of organs and tissues of man and beast by mag-
gots, the larvae of Diptera, is termed myiasis. The invading maggots may
be specific myiasis-producing forms, i.e., obligatory sarcobionts, which
invade cutaneous tissues, as does Dermatobia hominis (Linn.) in man
and Hypoderma bovis (DeGeer), the warble fly of cattle; invading the
gastric and intestinal tract are the botflies of horses, Gasterophilus intes-
tinalis (DeGeer); and invading the nasal and frontal sinuses are the
head maggots of sheep, Oestrus ovis Linn. The invading maggots may be
necrobionts or facultative sarcobionts, in which case traumatic dermal
myiasis may result, as with infestations of screwworms, Callitroga (=
Cochliomyia) americana C. & P. Accidental myiasis may be the result of
fly larvae in food, or the result of flies attracted by anal, vaginal, or nasal
discharges. The larvae of blowflies, bluebottle flies, and greenbottle flies
commonly occur in accidental intestinal myiasis as well as in traumatic
myiasis.
Allergy caused by insects. The condition of being specifically hyper-
sensitive to certain insect proteins is a fairly common and widespread
phenomenon among persons working habitually with bees or collections
.of dead insects, or exposed for longer periods to pulverized insect parts,
scales of butterflies, moths, and caddis flies. Residents on the shores of
lakes where cast skins of mayflies abound, are subject to attacks of
"asthma" as the result of allergens (Figley, 1929).3
Mechanical (simple) carriers of infection. Many species of insects may
accidentally contribute to the transmission of various filth diseases; how-
ever, when insects habitually and alternately feed and/or breed in excre-
ment and then feed on human food prepared for the table, they may
be classified as food contaminators and a real menace to the public
health, e.g., flies, particularly the common housefly, and cockroaches.
The causal organisms of filth diseases, such as typhoid fever, cholera, and
amoebiasis, may adhere to the mouth parts and feet of these insects and
may be deposited on human food, with resulting infection to the con-
sumer. The mechanical transmission of yaws (a spirochete infection)
and of certain eye infections (so-called pinkeye) is similarly effected
by flies, particularly Hippelates. The eggs of helminth parasites, notably
pinworms [Enterobius vermicularis (Linn.)] may also be so dissemi-
nated. Not only do pathogenic bacteria, protozoa, and helminthic ova
cling to the mouth parts, feet, wings, and other parts of the insect body,
but they may also be swallowed by the insect and pass uninjured
through its alimentary canal and be deposited on food with the insects'
feces or be regurgitated with similar effect.
It has been amply proved that coprophagous fly larvae (maggots),
which feed on and develop in human excrement, may transfer bacteria
taken up in this stage through the pupal stage to the mature flies. In this
manner the infection of anthrax (Bacillus anthracis) may be disseminated
by flesh flies, bred in carcasses of animals dead of this disease. This is a
strong argument in favor of the incineration of dead animals.
Another purely mechanical method of disease transmission is by
means of contaminated piercing mouth parts, in which these organs in
the act of feeding become contaminated with blood-inhabiting patho-
genic organisms, and simple inoculation may result from the next feeding.
Here again, as in the aforementioned cases, the pathogenic organisms
undergo no interim developmental change. Insects that belong to this
class of simple carriers generally have strong, piercing mouth parts, cap-
able of drawing considerable blood and are intermittent feeders, flying
readily and quickly from one host to another, e.g., horseflies (Tabani-
dae), which may be vectors of anthrax in this manner.
Cyclico-propagative transmission. Of the several ways in which so-
called biological transmission by arthropods is effected, the cyclico-pro-
pagative type is the one most likely to be used to illustrate insect trans-
mission of disease; however, Huff' has pointed out that there are actually
at least two other ways in which this occurs, namely, cyclico-developmen-
tal and propagative. In the cyclico-propagative type of transmission the
causal organisms "undergo cyclical change and multiply" in the body of
the arthropod, as in the transmission of malaria plasmodia by anopheline
mosquitoes and in the transmission of Babesia bigemina of Texas cattle
fever by the Texas fever tick, Boophilus annulatus.
Cyclico-developmentaI transmission. When the causal organisms "un-
dergo cyclical change but do not multiply" in the body of the arthropod,
transmission may be classified as cyclico-developmental, as in the mos-
quito transmission of the worm Wuchereria bancrofti, causal organism of
filariasis.
Propagative transmission. When "the organisms undergo no cyclical
change, but multiply" in the body of the vector, transmission is said to be
propagative only. The transmission of bubonic plague is probably prop-
agative, since it is known that the causal organism, Pasteurella pestis,
can multiply in the fore-gut of the flea. It is also very probable that the
transmission of relapsing fever by ticks falls into this category.
Fecal contamination. The feces of several species of disease-trans-
mitting arthropods are known to be infectious after the insect has fed on
infectious animals, for example, the fece~ of fleas fed on plague-infected
rodents, also the feces of conenose bugs, Triatoma, after feeding on ani-
mals infected with Brazilian trypanosomiasis, Chagas' disease. The feces
of infective yellow fever mosquitoes are said to be highly dangerous.
The Rockefeller Foundation (Annual Report, 1930) states that a dilution
of between one ten-millionth and one-billionth of one cubic centimeter of
yellow fever virus has frequently proved fatal to monkeys and that
infection through the unbroken skin is readily accomplished.
Hereditary transmission. Transmission of infection through the ar-
thropodan egg was discovered by Smith and Kilbourne in 1893 in their
investigation of Texas cattle fever. When the cattle tick, Boophilus an-
nulatus (Say), reaches maturity on the body of the animal, it drops to
the ground, where the female deposits her eggs and dies. The young
ticks on emerging from the eggs are infective and transmit the causal
protozoon, Babesia big emina, to susceptible cattle. The term "hereditary
transmission" is not correctly applied to the transmission of this disease,
since no hereditary or genetic mechanism is involved; however, it is now
widely used and may be permissible under the circumstances. Substitutes
such as congenital contamination, ovarian contamination, or transovarian
transmission have been suggested.
Other infections transmitted transovarianly are Rocky Mountain
spotted fever, caused by !lickettsia rickettsi (Wolbach) and transmitted
by ticks, e.g., Dermacentor andersoni Stiles; tularemia caused by Pas-
teurella (= Bacterium) tularensis and transmitted by the tick Derma-
centor andersoni Stiles; relapsing fever, caused by Borrelia (Spirochaeta)
recurrentis and transmitted by several species of ticks belonging to the
. genus Ornithodoros. Ticks play an important role in the hereditary
transmission of infections involving protozoa, rickettsiae, bacteria, and
spirochetes. Hereditary transmission of virus diseases is not so clear,
although sand fly fever, or Pappataci fever, has been produced by the
bites of Phlebotomus papatasii Scopoli bred from infected parents
(Whittingham 1922, according to Hinman). 5
Arthropods as intermediate hosts of helminths. Numerous arthropods
serve as intermediate hosts of many species of parasitic worms. There
are about 150 known species of worms parasitic in vertebrates which
have arthropods as intermediate hosts and for which the primary hosts
are known. About 275 species of arthropods are known to serve in this
capacity; of these nearly 200 are insects, about 75 are crustaceans, and the
remainder are arachnids and myriapods. No doubt hundreds of such life
histories are still to be ascertained. Hall 6 states that only 1 per cent of the
life histories of the known tapeworms have been worked out.
The reasons for the frequent host-parasite relationships between
arthropods and helminths are no doubt mainly the following: (1) the
arthropods are an exceedingly large group of animals; (2) they constitute
an important food supply for fish, amphibians, birds, and some mam-
mals; (3) herbivorous animals are sure to ingest arthropods which are
habitually present on plants; (4) many species of insects feed on and
breed in manures, and are thus exposed to infection from eggs and
larvae of worms parasitic in hosts responsible for the manure; (5) many
arthropods are transient or permanent ectoparasites feeding on blood in
which there may be parasitic worms.
Monoxenous worms such as Ascaris and Enterobius during the course
of their life histories transfer from one host animal to a similar host
animal without the intervention of an intermediate host. "The heterox-
enous worms have life histories in which in most cases the worms pass
from mature stages in one host animal to larval stages in a host animal
of a different sort, the intermediate host, and then return to a host animal
of the first sort or a more or less closely related species and develop in
this animal to maturity. In some instances two intermediate hosts are
utilized in sequence for larval stages." (Hall, loco cit. )
As already pointed out, knowledge concerning arthropods serving as
intermediate hosts for the Cestoda (tapeworms) is quite fragmentary.
Among the better-known instances are the double-pored dog tapeworm,
Dipylidium caninum (Linn.), which requires either the biting dog louse,
Trichodectes canis DeGeer, or a flea, Ctenocephalides canis (Curtis), as
intermediate host. Choanotaenia infundibulum (Bloch), a fowl tapeworm,
utilizes the housefly, Musca domestica Linn. (See appropriate chapters.)
Stunkard7 has shown that Moniezia expansa (Rudolphi), an important
tapeworm of sheep and other herbivores, uses a species of oribatid mite,
Galumna sp., as intermediate host. Eggs of the tapeworm were fed to
these mites and in time the onchospheres were recovered in large num-
bers from the body cavity of the mites. More recently Stunkard8 (1944)
reported a similar developmental pattern in oribatid mites for anoplocep-
haline cestodes of rabbits and for the monkey-human species.
Numerous trematodes (flukes) of fish, amphibians, reptiles, and
other insectivorous animals use arthropods as intermediate hosts. Since
many of the vertebrate hosts are aquatic or semiaquatic, so most of the
arthropods are also aquatic, such as dragonflies (Odonata), caseworms
(Trichoptera ), mayflies (Ephemerida ), and stoneflies (Plecoptera ).
Among these flukes are the poultry fluke, Prosthogonimus pellucidus
(v. Linstow), particularly of ducks, which use the larvae of the dragon-
fly, Libellula quadrimaculata Linn., as intermediate host. The important
lung fluke of man, Paragonimus westermani (Kerbert), requires as its
second intermediate host (the first is a melaniid snail) a crustacean,
Astacus spp., crayfish.
Among the Nematoda (threadworms) are numerous species that
use arthropods as intermediate hosts; These include the Gongylonema
worms (Spiruridae), such as G. pulchrum Molin, which causes an infec-
tion of humans (also of the pig, sheep, ox, etc.) known as gongylone-
miasis. These worms occur as larvae in such insects as cockroaches
(Blattidae), meal worms (Tenebrionidae), and a few other forms. The
mature worms, extremely slender (0.5 mm diameter), reach a length of
145 mm in the female. In the vertebrate hosts the worms are found in
burrows of the mucosa and submucosa of the mouth, tongue, and esopha-
gus. The eggs are evacuated with the fecal material of the host and do
not develop until ingested by an insect. The eggs hatch in the digestive
tract of the insect and soon penetrate the intestinal wall, coming to rest
as encapsulated larvae in the body cavity. There they remain until the
insect is ingested intact or in fragments by an appropriate vertebrate
host. In this host the .larvae are freed and migrate along the digestive
tract to the oral cavity, where they mature. Sambon 9 ,lo based his deduc-
tions concerning cancer on a study of Gongylonema worms. (See Chapter
VII.)
Other nematodes which require arthropods as intermediate hosts are
certain species belonging to the family Filariidae, such as Wuchereria
bancrofti and Onchocerca volvulus, the former requiring mosquitoes and
the latter black gnats (see later chapters).
The thornheaded worms (Acanthocephala) use beetles (Scara-
baeidae) mainly as intermediate hosts, e.g., the thornheaded worm of
swine, Macracanthorhynchus hirudinaceus (Pallas).
The famous guinea worm of the Nile Valley and equatorial Africa,
Dracunculus medinensis (Linn.), a worm which as an adult female
may measure from 70 to 120 cm in length, requires Crustacea belonging
to the genus Cyclops as intermediate hosts.
Reservoir animals. Reservoir animals play an important role in the
natural distribution of insect-borne diseases. Since true reservoir animals
suffer little or no ill effect from certain microorganisms pathogenic to
man their presence may go unnoticed, as is the case with rabbit reservoirs
of Rocky Mountain spotted fever; however, rat epizootics are commonly
the forerunners of human plague epidemics. The human being may him-
self be a reservoir of certain insect-borne infections, even plague. Since
there are numerous vertebrates which serve as disease reservoirs, it be-
hooves the medical entomologist to acquaint himself thoroughly with
the broad field of vertebrate zoology, particularly the ecological aspects,
and with the parasitic ectozoa of wild mammals. Greater attention must
be given to the problems of forest sanitation in recreation areas.
There are at least seventy species of mammals which may serve as
possible reservoirs of plague, according to StallybrassY The rickettsial
infections are notable for their wild-animal reservoirs: Rocky Mountain

spotted fever with its rabbits, badgers, woodchucks, field mice, and
others; typhus fever with rats; Japanese flood fever with voles. The
spirochete infections are equally. noteworthy in that the relapsing-fever
reservoirs are evidently fairly numerous; thus young porcupines, as well
as armadillos and opossums, have been listed, and in California certain
squirrels and chipmunks. Trypanosome infections are well represented
by sleeping sickness with its numerous big-game reservoirs, and Chagas'
disease with its armadillos, opossums, dogs, and cats.
Hudson12 calls attention to the fact that the animal reservoir host
plays a prime role in tropical medicine. In tropical regions of the earth
as well as in the less highly organized countries of the temperate zone
there is a striking increase in the number of diseases drawn from animal
reservoirs. Here huge numbers of human victims are "encompassed by
an intricate web of animal and insect life, in a physical environment of
extraordinary complexity and variability."
Recently a remarkable suggestion has been made by Maldonado 13
that certain plants may act as reservoirs of the causal organism of verruga,
a disease ef man, and that the species of Phlebotomus flies that transmit
the disease may feed on the latex. It is also suggested that this would
explain why these sand fl.ies are so abundant during the rains (January
to April), when this particular plant growth (Jatropa basiacantha and
Orthopterygium huancui) is most luxuriant.
nIBLIOGRAPHY
1. Smith, Roger C., 1934. "Hallucinations of insect infestation causing
annoyance to man," Bull. Brooklyn Entomolog. Soc., 29:208-12.
2. Stewart, Morris A., 1937. "Phasmid injury to the human eye," Ganad.
Entomolgist, 69:84-86.
3. Figley, K. D., 1929. "Asthma due to the may-fly," Am. f. M. Se., 178:
338-45.
4. Huff, Clay G., 1931. "A proposed classification of disease transmission
by arthopods," Science, 74:456-57.
5. Hinman, E. Harold, 1933. "Hereditary transmission of infections
through arthropods," Am. f. Trop. Med., 13:415-23.
6. Hall, M. C., 1929. Arthropods As Intermediate Hosts of Helminths,
"Smithsonian Misc. Collect., vol. 81, no. 15, Washington D. c.: Smithsonian
Inst. (Publ. no. 3024).77 pp.
7. Stunkard, H. W., 1937. "The life cycle of Moniezia expansa," Science,
86: 312.
8. ---, 1944. "How do tapeworms of herbivorous animals complete
their life cycles?" Tr. New York Aead. Se., ser. 2, 6:108-21.
9. Sambon, L. W., 1925. "Researches on the epidemiology of cancer
made in Iceland and Italy, July-Oct., 1924." J. Trop. Med. 28:39-71.
10. ---, 1926. "O,vations and researches on the epidemiology of
cancer made in Holland and Italy, May-Sept., 1925," J. Trap. Med., 29:233-
81.
11. Stallybrass, C. 0., 1931. The Principles of Epidemiology and the Proc-
esS of Infection. London: C. Routledge & Son. 696 pp. (1 plate, I chart).
12. Hudson, E. H., 1944. "The role of the reservoir host in tropical dis-
ease," Am. J. Trap. Med., 24:125-30.
13. Maldonado, A., 1931. "Role probable de quelques plantes caracter-
istique de la region verruqueuse sur l'etiologie de la verruga du Perou," Bull.
Soc. path. exot., 24:27-28.
CHAPTER V
STRUCTURE, DEVELOPMENT, AND

CLASSIFICATION OF INSECTS
AND ARACHNIDS
The Insecta (Hexapodaj constitute the largest class in numbers of

species (see footnote, p. 35) in the Phylum Arthropoda, which in turn
comprises a greater number of species thin all of the animal kingdom
combined. As members of the Phylum Arthropoda, insects share the fol-
lowing arthropodan characteristics: segmented body with paired, jOinted
appendages; bilateral symmetry; dorsal heart; ventral nerve cord; and
chitinous exoskeleton. Insects have the body divided into three more or
less distinct parts, the head, the thorax, and the abdomen. There were
primitively 20 or 21 segments, but owing to the specialization of the
head and posterior terminal segments, there are now only about ~2
recognizable ones. The head of the adult insect bears a pair of antennae,
the mouth parts, and the eyes; the thorax bears the locomotor appendages,
usually two pairs of wi\lgs, (these may be reduced or absent), and three
pairs of jointed legs; hIe abdomen bears no appendages except the
terminalia. Respiration is effected by means of a complex system of
microscopic tracheal tubules opening through the body wall and carrying
air directly to all parts of the body of the insect. Immature insects be-
longing to many of the orders differ markedly from the mature forms,
e.g., maggots of flies, but all possess tracheal tubules.
External anatomy. In order to familiarize himself with the external
anatomy of insects, especially with the parts upon which classification is
mainly based, the student should study carefully some large hard-bodied
insect, such as the horsefly (Fig. 11) or the cockroach. He should give
sufficient time to this exercise in the laboratory to become thoroughly
informed.
Wings. The earliest systems of insect classification were based on
wing characters, which together with the mouth parts and metamorphosis
afford a basis for modern classification. The venation of insect wings is so
markedly characteristic for most species that even a part of a wing is
often all that is necessary for determination. The winged insects are
46
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w
::2:
o
o
(l)
<l:
~
w "-
E
X
<l:
0::
o
I
I-
o
<l:
w
I
47
usually referred to as the Pterygota, and may be either Exopterygota
(wings developing externally), e.g., cockroaches; or Endopterygota
(wings developing internally), e.g., beetles (see under orders). There
are typically two pairs of wings present, situated on the mesothorax and
metathorax, though in many parasitic insects, such as the bedbugs, lice,
fleas, certain louse flies, etc., the wings are absent. The wingless insects
just mentioned should, of course, not be included with the Apterygota,
a term designating a group of primitively wingless (apterous) forms.
The above-mentioned parasitic, wingless insects belong to several dif-
ferent orders, as will be seen later.
In form the wing presents a more or less triangular appearance. The
three sides are called margins: the costal margin is cephalic or anterior,
the anal margin is posterior or caudad, and the outer margin (apical) is
between these. The three angles connecting the margins are humeral (at
the base), apical (apex of wing) and anal (between the apical and anal
margins). Generally the fore and hind wings differ considerably in size;
the fore wing in some groups, such as the mayflies, many butterflies and
moths, and the bees and wasps, is larger than the hind wing; while in the
grasshoppers, cockroaches, beetles, etc., the fore wing is narrow and
serves largely as a cover (tegmina) to the hind wing, which folds fan-like.
In the dragonflies, termites, and ant lions, the fore and hind wings are
nearly equal. In the flies, the hind pair of wings is replaced by club-
shaped organs known as halteres, leaving consequently only one pair of
wings, hence the name Diptera (two-winged). In the calypterate Diptera
(muscoid flies) there are present large lobes (squamae, also called
"alulae") at the junction of the wings and the thorax.
There are some differences in the structure of the wings within the
order, though for each order a certain general pattern prevails; e.g., the
Neuroptera have thin membranous wings, often quite filmy; however,
the wings of Diptera and many Hemiptera have the same texture through-
out but possess fewer and differently arranged veins. The Diptera can, of
course, be readily distinguished, as stated above, by the presence of but
a single pair of wings. The winged Hemiptera-Heteroptera have the
front wings thickened at the base, while the apical portion is membranous;
The winged Homoptera have two pairs of wings of even texture.
The venation of the insect wing is an important element in classi-
fication, on account of the great variety of arrangement and the reliability
of this character for the identification of the family and genus. The veins
are hollow, rib-like structures which give strength to the wing. The
areas of membrane between the veins are called cells; they are said to
be open if the membranous area extends to the wing margin. By a care-
ful study of the evidence, a fundamental type of wing venation has been
constructed by Comstock and Needham and revised by Tillyard. The
INSECT AND ARACHNID STRUCTURE 49
accompanying figure (Fig. 12A) illustrating this type will be useful in
determining the identity of the principal veins. Fig. 12B illustrates Till-
yard's revision of the R-C-N (Redtenbacher, Comstock, and Needham)
system of nomenclature, which is used in this book.
Metamorphosis. In order to achieve the size and development of the
parent, the young insect undergoes greater or less change in size, form,
and structure. This series of changes is termed metamorphosis. The least
A PRIMITIVE WING TILLYARD B TABANUS
FIG. 12. (A) diagram of wing venation after the R-C-N nomenclature
(Tillyard revision) with associated sclerites. A, anal veins = V, vllnnal veins
except the first; Ax, axillary sclerites (1, 2, 3, 4); AxC, axillary cord; C, costa;
=
Cu, cubitus; h, humeral cross-vein hm; HP, humeral plate; i-m, inter-medial
cross-vein; ir, interradial cross-vein (often spoken of as a spur on the third
vein); ], jugal veins; it, jugal fold; M, media; mt, medial form; m, m', median
plates; m-m, median cross-vein = im, intermedial cross-vein; m-cu, medio-
subital cross-vein; Feu, postcubitus (first anal); R, radius; r, radial cross-vein;
rt, radial fork; rf', 1st radial fork; r-m, radio-medial cross-vein; Rs, radial sector;
S, sectorial cross-vein; Sc, subcosta; tg, rudiment of tegula; V, vannal veins;
vf, vannal fold.
(B) wing of Tabanus labeled (in parenthesis) according to terminology of
the Dipterists. Veins: C (costal); Sc (auxilIary); R, (1st longitudinal); R.+,
(2nd longitudinal); R" R5 (3rd longitudinal); M" M, (4th longitudinal); Ms,
M.Cu (5th longitudinal); A (6th longitudinal). Cells: C (costal); Se (sub-
costal); R, (marginal); R3 (1st submarginal); R, (2nd submarginal); R5 (1st
posterior); M, (2nd posterior); 2ci M, (3rd posterior); M3 (4th posterior); M,
(5th posterior); 1st M, (discal cell); R (1st basal); 2dM (2nd basal); eu,
(anal cell); A, (axillary).
change is found in the Apterygota (silverfish and springtails), which

are primitively wingless insects, and hence the newly emerged young
individual is externally unlike the parent only in size: this type of
metamorphosis is termed primitive (Fig. 13).
A greater qegree of metamorphosis occurs in the grasshopper. There
is not only a great difference in size, but the absence of wings in the
young is at once apparent. In order to reach the winged condition, the
young individual casts its skin at intervals, and with each ecdysis achieves
longer wings until, after a certain number of molts, the fully developed
wings appear. The following stages may be recognized: (l) egg, (2)
nymph, (3) imago or sexually mature adult. This type of metamorphosis

is called simple or incomplete, and the orders comprising these are known
as the H eterometabola (Fig. 14).
The greatest difference between the newly hatched young and the
parents occurs in such forms as the housefly (Fig. 15) and the butterfly.
In these forms the newly hatched insect has no r~emblance whatsoever
\\
L'
c'
\\
"u"
"
"
'\
FIG . 13. Illustrating primitive meta-

morphosis . (a) young of a Thysanuran
insect (Campodea), (b) adult of the
same. (After Kellogg.)
to the adult, but looks more like a segmented worm. However, the in-
ternal anatomy and certain other features are distinctly insectan. The
fact that the young are mandibulate and the adults haustellate in Diptera
and Lepidoptera offers much interesting ground for ecological discussion.
In order to attain the winged condition of the adult, the wingless, worm-
A B c
14. Illustrating simple metamorphosis. (A) young wingless grass-
FIG.
hopper; (B) grasshopper showing wing pads after the first molt; (C) adult of
the same. (Redrawn after Packard.)
like form must undergo many profound changes., and a new stage is
interjected, the pupa, or resting stage, in which this transformation is
accomplished. The newly hatched young insect emerging from the egg
is called the larva, hence the following stageg: (1) egg, (2) larva, (3)
pupa, and (4) imago. This type is termed complex or complete meta-
morphosis, and the orders comprising these are known as the Holo-
metabola.
Importance of knowing internal anatomy. It is important that the
student familiarize himself with the internal anatomy of insects,! giving
special attention to the digestive system and its accessory structures, such
as the salivary glands. Two cases will illustrate reasons for this:
1. The simplest condition in which the internal organs of insects are
involved in disease transmission is in the case of the common housefly,
in which pathogenic organisms may be sucked up with infectious dejecta
from those ill with cholera or typhoid fever and passed out with the
feces of the fly, which may be deposited on human food , either in their
original virulent condition or more or less attenuated. Regurgitation on
the part of the insect may be equally effective.
I
FIC. 15. Illustrating complex metamorphosis in the life cycle of the common
housefly: egg, larva, pupa, adult .
2. A more complicated situation exists in the case of the Anopheles

mosquito, which sucks up pathogenic organisms (plasmodia) with its
meal of human blood, whereupon these parasites undergo vital sexual
changes within the body of the insect, eventually finding lodgment in the
salivary glands before final introduction by the "bite" into the next
human victim. The insect in this case is the essential natural vector.
Digestive system. There are three distinct regions to the insect intestine
(Fig. 16); namely, (1) the fore-gut, consisting of the mouth, pharynx,
esophagus, crop, and proventriculus; (2) the mid-gut, comprising the
stomach; and (3) the hind-gut, consisting of the ileum, colon, rectum ,
and anus. The crop presents merely a widened portion of the esophagus
in the more generalized forms and serves as a food receptacle. In the
more speciali"?ed groups, such as the Diptera and Lepidoptera, the crop
is expanded into a capacious pocket or pouch. In such forms as the
cockroach and grasshopper the proventriculus consists of a highly muscu-
lar dilation provided internally with chitinous teeth for grinding or
straining food. The stomach is a simple sac into which open the gastric
Pro
Labm.
FIG. 16. Showing digestive h'act of cockroach, Order Orthoptera (top);

of cone nose bug, Order Hemiptera (middle); and of anopheline mosquito,
Order Diptera (bottom). Explanation of abbreviations: Ac.G., accessory glands;
An., anus; Ao1'., aorta; AS.G., accessory salivary gland; Br., brain; Ca., cardia;
Co ., colon; C1'., crop; EjD., ejaculatory duct; Eso., esophagus; Eso.D., esopha-
geal diverticula; Cas.C ., gastric caeca; He., heart; Hyp., hypopharynx; Il.,
ilium; Labm., labium; Labr., labrum; Max.S ., maxillary stylet; Md.S.; mandi-
hular stylet; M.G., mid-gut; M.T., Malpighian tubules; N.C., nerve cord; Pe.,
penis; Ph., pharynx; Pr., proventriculus; Pre.C., preoral cavity; P.S.G., principal
salivary gland; R., rectum; R.O., reproductive organs; R.P., rectal papilla;
SaLD., salivary duct; S.C., salivary gland; Sal.P., salivary pump; S.R., salivary
reservoir; St ., stomach; T. , testis; VD., vas deferens. (Top figure adapted after
\Hall; middle figure adapted after Elson; and lower figure, Herms.)
52
caeca, generally few in number, which give rise to certain digestive
fluids. At both ends of the stomach are located valves which control the
flow of the food. There is much variation in the length and the degree
of convolution of the hind intestine, but usually the three regions men-
tioned, namely, ileum, colon, and rectum, can be located. Emptying into
the ileum are the excretory or Malpighian tubules, varying in number
and length in the several groups of insects. •
The salivary system consists of a pair of salivary glands which may
be lobed; they are situated within the head, often extending into the
thorax. Usually each gland empties into a salivary duct, the two ducts
. discharging into a common duct which opens into the mouth at the base
of the labium. In many species of insects there is present a pair of sali-
vary reservoirs; these may be located near the opening of the common
duct and then present a compound condition, or may be situated on
either side of the esophagus at the end of a long slender duct.
Insect larvae. When insect larvae, parasitic or accidental, are en-
countered in the body of man or beast, confusion may arise because of
the worm-like appearance of the invaders, which may be incorrectly
classified as worms, for example, muscoid fly larvae. These larvae are
short and plump, and have 11 or 12 well-marked segments. Microscopic
examination of fragments will reveal tracheal tubules" which are not
present in worms. The student of medical entomology must be thoroughly
familiar with the immature stages of arthropods.
Although the larvae of insects belonging to the Order Diptera (flies
and mosquitoes) are characteristically legless and usually have an un-
chitinized, eyeless head, the variations within the order are considerable.
Maggots of muscoid flies, for example, are smooth, with the body taper-
ing to the inconspicuous head, which bears hook-like mouth parts; while
the larvae (wrigglers) of mosquitoes have a chitinous, freely moving,
conspicuous head with faceted eyes, and both the head and body bear
many hairs, spines, and bristles. The larvae of fleas (Order Siphonaptera)
are also legless; the chitinous head is well developed; and each of the
thoracic and abdominal segments is well armed with a band of bristles.
The larvae of beetles (Order Coleoptera), commonly called grubs, have
three pairs of legs on the thorax only; the head is well developed; the
body may be hairy, spiny, or naked. The larvae of moths and butterflies
(Order Lepidoptera) have three pairs of thoracic legs and two to five
abdominal prolegs; the head is prominent, and the mouth parts are
usually well developed and mandibulate; they are commonly called
caterpillars and are hairy, spiny, or naked. The larvae of ants, bees, and
wasps (Order Hymenoptera) are usually called grubs; they are usual1~7
without legs (apodous); the head is more or less well developed; t':e
body is usually fairly smooth and .gourd-shaped.
Insect classification. The medical entomologist must be equipped with

a good knowledge of the basic principles of classification so that he may
be able to place the insect at hand correctly in at least its proper Order
and family; in the case of insects of importance in medical entomology
he should be able to run the specimen down to the species with the aid
of a key. To determine the order to which an insect belongs one need
usually know only the venation and structure of the wings, if present, and
the type of mouth parts. This knowledge will enable the student to place
at least 90 per cent of the commoner insects in their proper orders. Un-
fortunately, the parasitic forms have undergone many modmcations,
such as reduction or loss of wings and great alteration in form, but
generally the mouth parts will serve as a ready means of crude identm-
cation. Before passing on to a list of the orders of insects, the usual basis
for classification may be pointed out, viz.:
1. Wings: (a) presence or absence, (b) form, (c) structure.
2. Mouth parts: (a) biting (mandibulate), (b) sucking (haustellate).
3. Metamorphosis: (a) primitive, (b) simple (incomplete), (c) complex
( complete) .
The orders of insects. The following list of orders of the class Insecta
includes those generally recognized by present-day entomologists. The
Comstocks in 1901 listed 19 orders of insects. The present number, 33
(Essig, 1942 2 ) has been primarily achieved by splitting up the original
Orthoptera and Neuroptera.
Subclass I. Apterygota (Ap'ter-y-go'ta), apterous (primitively wingless) in-
sects.
1. Order Protura (Pro-tu'ra), (protos, first; oura, tail), proturans; (be-
cause of certain affinities with the Chilopoda and Symphyla, these in-
sects are sometimes placed in a separate class, Myrientomata Berlese);
biting mouth parts; primitive metamorphosis.
2. Order Thysanura (Thy'sa-nu'ra), (thysanos, a tassel; oura, tail); silver-
fish, bristletails; biting mouth parts; primitive metamorphosis.
--3. Order Aptera (Ap'ter-a), (6-, without; ptera, wings); campodeids;
biting mouth parts; primitive metamorphosis.
4. Order Collembola (Col-Iem'bo-la), (colla, glue; embolos, wedge or
peg); springtails, snowfleas; biting mouth parts; primitive metamorpho-
sis.
Subclass II. Pterygota (Pter'y-go'ta), winged insects.
Division I. Exopterygota (Exo-pter'y-go'ta).
(Heterometabola: Insects with simple metamorphosis.)
5. Order Orthoptera (Or-thop'ter-a), (orthos, straight; ptera, wings);
grasshoppers, locusts, katydids, crickets, mole crickets; biting mouth
parts; simple metamorphosis.
. 6. Order Blattaria (Blat-tar'ia), (blatta, insect that shuns the light);
cockroaches; biting mouth parts; simple metamorphosis.
7. Order Dermaptera (Der-map'ter-a), (derma, skin; ptera, wings); ear-
wigs; biting mouth parts; simple metamorphosis.
8. Order Grylloblattodea (Gryl'lo-blat-to-de' a), (gryllus, a cricket; blatta,
a cockroach); grylloblattids; biting mouth parts; simple metamorphosis.
9. Order Phasmida (Phas'mi-da), (phasma, an apparition); walking sticks,
leaf insects; biting mouth parts; simple metamorphosis.
10. Order Mantodea (Man-to' de-a), (mantis, a prophet; eidos, form, like);
preying mantids, praying mantids, soothsayers; biting mouth parts;
simple metamorphosis.
11. Order Diploglossata (Dip'lo-glos'sa-ta), (diplos, double; glosso,
tongue); hemimerids; biting mouth parts; simple metamorphosis.
12. Order Plecoptera (Ple-cop'ter-a), (plektos, to weave; ptera, wings);
stoneflies, salmonflies, perlids; biting mouth parts; simple metamorpho-
sis.
13. Order 1soptera (1-sop'ter-a), (isos, equal; ptera, wings); termites, white
ants; biting mouth parts; simple metamorphosis.
14. Order Zoraptera (Zor-ap'te-ra), (zoros, pure; ptera, wings); zorap-
terans; biting mouth parts; simple metamorphosis.
15. Order Embioptera (Em'bi-op'-ter-a), (embios, lively; ptera, wings);
embiids, webspinners; biting mouth parts; simple metamorphosis.
16. Order Corrodentia (Cor'ro-den'ti-a), (corrodens, gnawing); book lice,
bark lice, dust lice, psocids; biting mouth parts; simple metamorphosis.
17. Order Mallophaga (Mal-Ioph' a-ga), (mallos, a lock of wool; phagein,
to eat); bird lice, biting bird lice; biting mouth parts; simple metamor-
phosis.
18. Order Ephemerida (Eph' e-mer' i-da), (ephemeros, living but a day);
mayflies, dayflies, ephemerids; vestigial mouth parts; simple metamor-
phosis.
19. Order Odonata...fO-don'a-ta), (odontos, a tooth); damsel flies, dragon-
flies, mosquito hawks, devil's darning needles, snakedoctors; biting
mouth parts; simple metamorphosis.
20. Order Thysanoptera (Thy'sa-nop'ter-a), (thysanos, fringe; ptera,
wings); thrips; rasping-sucking mouth parts; simple metamorphosis.
21. Order Anoplura (An'o-plu'ra), (anoplos, unarmed, oura, tail) true lice,
sucking lice; piercing-sucking mouth parts; simple metamorphosis.
22. Order Hemiptera (He-mip'ter-a), (hemi, half; ptera, wings); bugs,
cicadas, treehoppers, leafhoppers, aphids, scale insects, etc.; piercing-
sucking mouth parts; simple metamorphosis.
Division II. Endopterygota (En-do-pter'y-go'ta).
(Holometabola-1nsects with complex metamorphosis) .
. 23. Order. Megaloptera (Meg' a-Iop'ter-a), (megalos, great; ptera, wings);
dobsonflies, alderflies, sialids; biting mouth parts; complex metamor-
phosis.
24. Order Neuroptera (Neu.rop'ter-a), (neura, nerves; ptera, wings); lace-
wings , nerve-wiugecl insects; biliug mouth parls; complex metam(Jr -

phosis.
25. Order Raphidiodea (Ra-phid'i-o'de-a), (raphis, needle; rderring [0
ovipositor of female) ; snakeflies, serpent fli es; biting mouth parts ; com
plex metamorphosis.
26. Order Mecoptera (Me-cop'ter-a) , ( mecus, length, long; ptera, wings):
scorpion£lies; biting mouth parts; complex metamorphosis.
27. Order Trichoptera (Tri-chop'ter-a) , (ti1rix, hair ; ptera, wings); caddis
flies, caseflies, water moths; biting mouth parts, complex metamorphosis.
28. Order Coleoptera (Col'e-op'ter-a), (c oleos, sheath; ptera, wings) ,
beetles, weevils, biting mouth parts; complex metamorphosis.
29. Order Strepsiptera (Strep-sip 'ter-a) , (st l'epsi, twist; ptera, wings);
stylops, twisted wings; biting mouth parts; complex metamorphosis.
30. Order Diptera (Dip'ter-a), (dis, two ; ptera, wings); flies, gnats, mos-
quitoes; sucking mouth parts; complex metamorphosis.
31. Order Siphonaptera (Si'pho-nap'ter-a) , (Siphon, a tube ; aptera, wing-
less); fleas, chigoes; piercing-sucking mouth parts ; complex metamor-
phosis. (See Snodgra.sS'. ' )
32. Order Hymenoptera (Hy'men-op 'ter-a), (hymen, a membrane; ptera,
wings); bees, wasps, ants, sawflies, horntails, etc. ; lapping-sucking
mouth parts; complex metamorphosis.
33. Order Lepidoptera (Lep'i-clop'ter-a), (lepidos , scale; ptera, wing) ;
moths, butterflies; sucking mouth parts ; complex metamorphosis.
The Arachnida. The class Arachnida includes the ticks, mites, spiders,
scorpions, and related forms. Am9ng the species of arachnids are some
of the most important parasites and vectors of diseases of man and beast,
such as the ticks which carry spotted fever and relapsing fever of man,
FIG. 17. Showing external morphology-of a spider. C, coxa; Cit., chelicera:

E, eyes; Fill., femur; Op., opisthosoma; Pat., patella; Pd. , peclipalp; Pr. ,
prosoma ; Ptar., pretarsus; Sp. , spinnerets; Tar ., tarsus; Th., tibia; Tr., troch-
anter.
and Texas cattle fever and bovine anaplasmosis. Parasitic mites cause
acariasis, often serious, such as mange, scabies, and various forms of itch,
and may, like the ticks, serve as vectors of disease, particularly Tsutsuga-
mushi disease.
The more important arachnids lack distinct segmentation of the body,
e.g., ticks, mites, and spiders; while scorpions, pseudoscorpions, and a
few others are clearly segmented. The body is divided into two parts
(Fig. 17): first the cephalothorax (prosoma) composed of combined
head and thorax, and second, the abdomen (opisthosoma). In the ticks
and mites there is a strong fusion of the cephalothorax and the abdomen
so that the body becomes sac-like in form.
Adult arachnids with few exceptions have four pairs of legs, though
the larvae of ticks and nearly all mites have but three pairs. In spiders
there is a pair of pedipalpi, which in scorpions, whip scorpions, and
pseudoscorpions are strongly chelate. All arachnids are devoid of wings
and antennae. Eyes, when present, are simple; compound eyes are want-
ing. The mouth parts usually consist of a pair of piercing chelicerae,
peclipalpi, and in the Acarina a hypostome. The respiratory system of
many arachnids, particularly ticks and mites, is tracheal as in insects,
FIG. 18. Showing intemal anatomy of a spider. A, anus; B.L ., book lung;
Chc., chelicera; D.M., dorsal muscle; D.M.G., diverticula of mid-gut; E., eye;
I-I., heart; M.G., mid-gut; lH.T ., Malpighian tubule; P., pericardium; Pd.,
pedipalp; Sp., spinneret; S.St., sucking stomach. (Redrawn and adapted after
various authors.)
except that there is usually but one pair of spiracles. In spiders the
respiratory organ is a combination of lung books and tracheae. There is
frequently a strong sexual dimorphism in the arachnids; the males are
. commonly smaller than the females . (See later chapters dealing with
ticks, mites, and spiders.)
In general the arachnids are predatory and perhaps most of them
are nocturnal, although there are some species of spiders and scorpions
which are largely diurnal.
Arachnid development. Arachnida deposit eggs in all the orders
except the scorpions and some mites (Pediculoides), which are vivipa-
rous. Eggs are usually numerous, particularly in the ticks, which may
deposit as many as 18,000 per female. The newly hatched individuals
have the general form of the adults, although the number of legs may
vary, e.g., newly hatched ticks and mites usually have three pairs of legs.
Metamorphosis is simple, as in cockroaches and grasshoppers. Molting
takes place as in insects, the various stages being termed instars as in
the Insecta. The longevity of many arachnids is remarkable: ticks have
been known to live for as many as six to seven years, and some species
are able to endure starvation for several years.
Internal anatomy. The digestive tract of arachnids (Fig. 18) is charac-
terized by various types of diverticulae and branched tubules. The
diverticulae, which diverge from the tract between the sucking organ
of the pharynx and the mesenteron, range, according to Savory' from
two short simple sacs directed forward in the cephalothorax to a condi-
tion of five pairs, four of which extend laterally, reaching the bases of
the legs, and enter the coxae for a short distance; also there is a very
complex type which branch and divide and become very large. Leading
from the mesenteron is a complex system of branched tubules which
occupy most of the abdomen and function partly as a digestive gland
and partly as a reservoir. The Arachnida are thus enabled to store large
quantities of food and to undergo long periods of fasting, an adaptation
which particularly favors the parasitic forms.
The excretory organs of the arachnids are Malpighian tubules, which
empty into the gut, and coxal glands which empty excretory products
into tubules and discharge to the exterior from openings which vary in
relations to the coxae with the several orders.
Arachnid classification. o Excluding the marine arachnid order Xipho-
sura, king-crabs or horseshoe crabs, the following orders, all terrestrial,
may be considered in a classification for the purposes of this work,
namely, (1) Scorpionida, scorpions; (2) Araneida (Araneae), spiders;
(3) Pedipalpida, whip scorpions; (4) Pseudoscorpionida (Chelonethida),
false scorpions; (5) Solpugida (Solifugae), sun spiders or wind scorpions;
(6) Phalangida (Opiliones), harvestmep pr harvest spiders; (7) Acarina
(Acari), mites and ticks.
Among the more important gross characters used to separate the
terrestrial arachnids into orders are the following: segmentation of the
body; presence or absence of pedicle; presence or absence of telson;
chelicerae, large or small; pedipalpi, chelate or unchelate; location and
form of spiracles. '.
o Some authors include among the Arachnida the Pycnogonida ( Pantopoda )

sea-spiders, the Tardigrada (water-bears), and the Pentastomida (Linguatulida)
tongue worms.
,
Key to the Terrestrial Orders of the Class Arachnidaf

1. Abdomen distinctly segmented ......... ............................... 2
Abdomen not segmented ...................................................................... 7
2. Abdomen with tail-like prolongation ............................................... 3
Abdomen without tail-like prolongation .............................. ........ 4
3. Tail stoutly armed with a sting at end ...... Scorpionida (Scorpions)
Tail slender, without sting . ..... . ............ Pedipalpida (Whip scorpions)
4. Palpi chelate .. ...... Pseudoscorpionida (False or Pseudoscorpions)
Palpi not chelate....... .. ... ............ ............. ..................................... 5
5. Abdomen constricted at base and narrowly jOined to the cephalo-
thorax ................... .. Pedipalpida
Abdomen not constricted at base and broadly joined to cephalothorax 6
6. Legs very long and slender, body hairless, whole body fused together
.................... .... ...................... . ......... . ... Phalangida (Harvest spiders)
Legs moderate; body hairy, appearing 3-segmented .. . ........... ..
.............. . . . . . . .............. Solpugida (Solpugids, Vinegarroons)
7. Abdomen constricted at base and jOined to cephalothorax by a narrow
stalk .... . .. Araneida (True spiders)
Abdomen fused with cephalothorax .. . .Acarina (Ticks and Mites)
BIBLIOGRAPHY
1. Snodgrass, R. E., 1935. Principles of Insect Morphology. New York:
McGraw-Hill Book Co., Inc. 667 pp.
2. Essig, E. 0., 1942. College Entomology. New York: The Macmillan
Co., vii + 900 pp. (308 figs) .
3. Snodgrass, R. E., 1946. The Skeletal Anatomy of Fleas. "Smithsonian
Misc. Collect.," vol. 104, no. 18, Washington, D. C.: Smithsonian Inst. (Publ.
no. 3815). 89 pp. + 21 plates.
4. Savory, Theodore H., 1935. The Arachnida. London: Edward Arnold
& Co. 218 pp.
t Adapted after various authors.

._ ..
CHAPTER VI
INSECT AND ARACHNID MOUTH PARTS
Importance of mouth parts. No doubt all insects possessing mouth

parts capable of piercing the skin of man and beast may be regarded
with some suspicion by the medical entomologist as being potential vec-
tors of blood-inhabiting microparasites, even though the bloodsucking
habit may not normally prevail. The nonpiercing insects manifestly can-
not be responsible for infections introduced directly into the circulation,
except through previously injured surfaces. Thus the housefly although
possessing nonpiercing mouth parts, is recognized as one of the vectors
of Trypanosoma hippicum Darling, a blood-inhabiting parasite and the
causal organism of murrina of horses and mules. It carries the parasites
on its proboscis, which becomes contaminated while it is feeding from
bleeding wounds caused by stable flies (Stomoxys) or other means, and
conveys the organisms thence to open wounds on healthy animals.
The student of arthropods in relation to disease must have a thorough
knowledge of their mouth parts and feeding habits for at least two rea-
sons: first, that he may know how the pathogenic organisms are acquired
as well as transmitted and, secondly, that he may know, if occasion re-
quires, what control measures may be best employed. The medical en-
tomologist will soon find that a knowledge of the feeding habits of the
adult insect alone is inadequate and that he must be familiar with the
feeding habits of the insect in all stages of its life cycle. The larvae of
many of the sucking insects have mandibulate (biting-chewing) mouth
parts. The biting mouth parts of the worm-like flea larva enable it to
ingest particles of excrement or other matter in which eggs of the double-
pored dog tapeworm occur, and it may thus become an intermediate
host of this worm, passing the infection on to the adult flea which, if
ingested by a suitable host, becomes the agent of infection. Insects with
complete metamorphosis commonly change from chewing mouth parts in
the larval stage to the sucking type in the adult. Insects with simple
metamorphosis, like the cockroaches, have biting mouth parts in all active
stages of development. Snodgrass 1 describes the feeding apparatus of
biting and sucking insects affecting man and animals in great detail.
Classification of mouth parts. All insect mouth parts, however highly
60
specialized, may be traced to the simple primitive type still existing with
some modification in the cockroach. Insect mouth parts are commonly
divided into two general classes: (1) mandibulate, i.e., biting and chew-
ing, as in the cockroaches, grasshoppers, and beetles; and (2) haustellate,
i.e., sucking, as in the bugs, flies, bees, butterflies, and moths. This
classification is far from satisfactory for use in the field of medical en-
tomology. For example, the common housefly, Musca domestica Linn.,
and the stable fly, Stomoxys calcitrans (Linn.), both possess haustellate
mouth parts and belong to the same family of insects, Muscidae, hence
are systematically closely related, yet are quite unrelated in their man-
ner of disease transmission. By virtue of its efficient piercing stylets the
stable fly has the power to pierce the skin and suck blood, thus enabling
it to become a direct infector; whereas the housefly, because of the struc-
ture of its proboscis and its inability to suck blood by piercing the skin,
is only indirectly responsible for infection, i.e., it is more particularly a
food contaminator.
Obviously insects could be grouped on the basis of mouth parts and
feeding habits, into two simple divisions: (1) piercing as in mosquitoes,
and (2) nonpiercing, as in cockroaches. This, however, leaves much to
be desired in the light of medical entomology, hence the following
grouping is suggested.
1. Orthopteron type: generalized mouth parts consisting of opposable jaws

used in biting and chewing; upper and lower lips easily recognized, as
in the cockroach and grasshopper.
2. Thysanopteron type: mouth parts representing a transitional type, minute
in size; approaching the biting form, more particularly rasping, but func-
tioning as suctorial organs; as in the thrips; the right mandible is greatly
reduced or possibly even absent, causing a peculiar asymmetry shown in
the drawing (Fig. 20).
3. Hemipteron type: mouth parts consisting of piercing-sucking organs, com-
prising four stylets closely ensheathed within the elongated labium, form-
ing a proboscis, as in the conenose bug and bedbug.
4. Anopluran type: mouth parts piercing-sucking, in a sac concealed within
the head but evertible when functioning. Three stylets, i.e., one consist-
ing of the pair of united maxillae, the other two consisting of the hypo-
pharynx and labium; mandibles vestigial, e.g., sucking lice.
5. Dipteran type: suctorial organs, piercing in some, nonpiercing in others;
no single representative is available to illustrate the entire group of Dip-
tera, hence the following subtypes may be recognized.
a. First subtype: mosquito; mouth parts consisting of six stylets, loosely
ensheathed within the labium.
h. Second subtype: horsefly; mouth parts consisting of six short blade-like
structures used for piercing and cutting, all loosely ensheathed within
the labium.
c. Third subtype: stable fly; piercing stylets reduced to two in number,
closely ensheathed within the labium.
d. Fourth subtype: housefly; mouth parts consisting of a muscular pro-
boscis, not suited for piercing; stylets rudimentat:y.
e. Fifth subtype: the louse fly; mouth parts closely related to those of
the third subtype.
6. Siphonapteron type: piercing-sucking, fully exposed mouth parts consist-
ing of a pair of broad maxillary lobes bearing long palpi; two (paired)
stylets, a pair of maxillary lacinae (lacinial blades), chief cutting organs
commonly regarded as "mandibles"; a slender labium with parallel palpi,
and a median unpaired stylet, the "epipharynx," the labrum is difficult to
interpret (Snodgrass), e.g., fleas. _
7. Hymenopteron type: mouth parts consisting of suctorial, lapping organs,
mandibles specialized for portage and combat, as in the bee, wasp, and
ant.
S. Lepidopteron type: mouth parts consisting of a suctorial coiled tube, as
in the butterfly.
MORPHOLOGY OF MOUTH PARTS

Orthopteron type. To illustrate the orthopteron type of mouth
structUre either the grasshopper or the cockroach may be used. This
type, the mandibulate or biting-chewing, is the generalized or primitive
form and will serve as a basis for later comparisons and derivations. It is
not directly of importance in medical entomology except as it furnishes
a basis for a better understanding of the haustellate or sucking types.
If the head of a cockroach (Fig. 19) is viewed from the side and
again from the front, the relative position of the separate parts will be
better understood. Separating the individual parts, the following struc-
tures will be observed. In front, low down on the head, hangs the
labrum or lip, easily lifted as one would raise a hinged lid, the hinge
line being at the lower edge of the sclerite or plate, known as the clypeus.
The labrum functions as does the upper lip in higher animals, i.e., it
draws the food toward the mandibles. In this the labrum is greatly
aided by a rough structure called the epipharynx, which forms the inner
lining of the labrum and clypeus. Because of the close association of
these two structures they are often referred to as a double organ, the
labrum-epipharynx. Removing the labrum, a pair of heavy, opposable
jaws, the mandibles, is exposed. These are biting structures par excel-
lence. They are toothed and movable laterally. Dislodging the mandibles
brings into view the pair of maxillae, or accessory jaws. These organs are
known as first maxillae. They are composite structures separable into
cardo, stipes, lacinia, galea, and palpus, which should be carefully ob-
served, inasmuch as they undergo great modification in the remaining
types of mouth parts. The two supporting sclerites of the maxillae are
'1
~IDE VIEW FRONT VIEW

MOUTH' PARTS
HEAD
LABRUM
MANDIBLE MANDIBLE
. ~
/ CARDO 'm"":"
,
" h
'.
HYPOPHARYNX
'I
MAXILLA MAXILLA
lABIUM '
FIG. 19. Head and mouth parts of a cockroach. Orthopteron (mandibulate)

type of mouth parts.
63
carda (basal) and stipes (the second), while the distal lobes are (1)
the maxillary palpus (a jointed structure), (2) the galea (median and
fleshy), (3) the lacinia (inner and toothed), capa ble of aiding in com-
minuting food .
Underneath the maxillae and forming the floor of the mouth lies the
lower lip or labium, a double structure frequently called the second
maxilla. On the same plan as the maxillae, the labium consists of a basal
sclerite, the submentum, followed by the mentum, upon which rest the
labial palpi (a pair of outer, jointed structures to the right and left),
and the ligula (a pair of strap-like plates which together correspond to
the upper lip.) The labium is also subject to much modification in insects.
The fleshy organ still remaining in the mouth cavity after the parts
just described have been removed is the tongue or hypopharynx, an organ
of taste, comparable in a measure to the tongue of higher animals.
H /y
+----ANTENNA
COMPOUND EYE
FIG. 20. Head and mouth parts

of a thrips. Thysanopteron type.
(Hedrawn after Borden. )
The mandibles are most useful landmarks, since they are almost
universally present in insects, though in various degrees of development,
from the strong mandibles of certain beetles (Lucanidae) to the vestigial
structures of fleas (Siphonaptera). In the Hymenoptera, even though the
order is haustellate, the mandibles are nevertheless present and important
structures, serving, however, in the honeybee as wax implements and
organs of defense, and in ants as organs of portage and combat. In
Hemiptera and many Diptera the mandibles are converted into piercing
organs while the maxillae are also greatly changed in form.
Thysanopteron type. This type (Fig. 20) is interesting phylogeneti-
cally as a connecting link between the biting-chewing and piercing-
sucking mouth parts. It is in the very minute thrips, order Thysanoptera,
that we find this transitional type of mouth parts, biting in general struc-
ture but sucking in function. According to various authors the right man-
dible is reduced, and by others it is said to be entirely wanting, making
the head and mouth parts asvmmetrical; the left mandibl e, maxillae, and
INSECT AND AHACllNlD MOUTH PA ins 65
hypopharynx are elongate, suggesting the stylet of the piercing type,
and adapted to move in and out through a circular opening at the apex
HEMIPTEHA
MOUTH PARTS
MANDIBLE
RIGHT MANDIBLE.._--------~
RIGHT MAXILLA
FOOD CHANNEL-~-~
,,
\
DUCT- - I
LEFT MAXILLA ,,I
LEn MANDIBLE
I
I
I
I
,I
I
I
I
iS1YLE1S MAXILLA
I
: LABIUM
I
LEFT
3 4 5 -- 6 - -
FIG. 21. Hemipteron type of mouth parts. (Adapted after various authors.)
of the head. No food channel is formed , but the sap from plants is lap-
ped up as it exudes from the abraded surface.
Hemipteron type. A very djfferent sort of organ from those above
desclibed is found in the order Hemiptera (Fig. 21). Here the cylindri-
cal labium forms a prominent beak-like proboscis which is usually three-

or four- (rarely one- or two-) jointed and telescopic. It is devoid of
palpi. The proboscis encloses a pair of mandibles, often with terminal
Head
PRESTOMAL
TEETH __
------
ROSTRUM
/OORSAL STYLET \ VENTRAL STYLET

DORSAL STYLET
\
\
VENTRAL~BUCCAL FUNNEL DORSAL VIEW
STYLET - PHARYNGEAL
_ \ TUBE --EYE
PROTRACTOR
'SAC TUBE RETRACTOR
DILATOR MUSCLES MUSCLES
ROSTRUM
SAC TUBE--
PHARYNX//
RIGHT HALF OF SUCKING TUBE
RIGHT HALF OF SAC TUBE
FIG . 22. Head and mouth parts of a sucking louse. Anopluron type of mouth
parts. (Redrawn and adapted after various authors.)
barbs, and a pair of maxillae, all four are stylet-like and efficient piercing
organs, the maxillae operating as a unit and the mandibles functioning
separately. The maxillae are closely apposed, forming the food and
INSECT AND AHACHNlD MOUTH PAHTS 69
are narrower and provided ,with conspicuous palpi. The hypopharynx
and labrwn-epipharynx are both lancet-like, In the male these pierci ng
parts are very weakly developed.
HEAD
SIDE VIEW LABIUM REMOVED
ANTENNA COMPOUND EYE
MAXILLARY PALP
MAXILLARY PAL?
MANDIBLE
HYPOPHARYNX LABRUM-EPIPHARYNX
SAliVARY
SAL IVARY PUMP PUMP
PALPUS
LABRUM LEFT MANDIBLE LEFT MAX ILLA LABELLA
FIG. 24. Head and mouth parts of a horsefly. (In part redrawn after
Snodgrass. )
(c) Third subtype, the stable fly. This subtype (Fig. 25) is repre-
sented by a group of flies in which the mouth parts are distinctly
specialized for piercing, and show, together with the next subtype, to
what extent these structures may become modified within the same
family of insects.
The proboscis at rest is carried at the position of a bayonet at charge,
and is therefore provided with a prominent muscular elbow or knee.
This conspicuous organ (the proboscis) is the labium terminating in the
labella, which are provided with a complex series of rasping denticles.
Within the folds of the labium and easily removable through the upper
groove lie two stylets: the labrum, the uppermost and heavier, and the
hypopharynx, a lower and weaker one, the two forming a sucking tube
SIDE VIEW FRONT VIEW
HEAD
__ - COMPOUND EYE- __
ANTENNA
MAXILLARY
LABRUM-EPIPHARYNX
FOOD CHANNEL
DUCT
~ ____ PROBOSCIS
~'-'
FE EDING POSIT ION
OOR~toi'l~TRAl ~tR~~ HYP. LAB.EP.
FIG. 25. Head and mouth parts of the stablefly. (Redrawn and adapted after
various authors.)
supported within the folds of the labium. The maxillary palpi are located
at the proximal end of the proboscis.
(d) Fourth subtype, the housefly. Here (Fig. 26) the prominent
fleshy proboscis consists mainly of the labium, which terminates in a pair
of corrugated rasping organs, the labella, and is attached in elbow-like
form to the elongated head. The entire structure is highly muscular and
may be either protruded in feeding or partially withdravffi while at rest.
Lying on top of the grooved labium is the inconspicuous spade-like
SIDE VIEW FRONT VIEW
INTERBIFID
SPACES~~~~(i=-~
MEMBRANOUS
LINING
SALIVARY SUPERFICIAL SLIT
DUCT LEADING INTO
PSEUDOTRACHEAf-~~-~
CH ITINOUS RINGS
OF PSEUDOTRACHEA
LABELLA EPIDERMIS
FIG . 26. Head and mouth parts of the common housefly. Lower right-hand
figure shows detailed cross section of a pseudotrachea in the labella. (Redrawn
and adapted after various authors.)
labrum, which forms, with the hypopharynx, a sucking tube, supported
by the labium, which latter also encloses the salivary canal. By an ex-
amination of the labrum it will be seen that this forms a sort of convex
covering to the concaved hypopharynx, thus giving rise to a food tube.

The maxillae have evidently become fused with the fleshy elbow of the
proboscis, and only the prominent maxillary palpi remain.
SIDE VIEW VENTRAL VIEW
CONCEALED
ANTE A
EPIPHARYNX
TRACHEA~
P.
EPIPHARYNX
1D.
00
'00,'
•
7
MAXILLARY
LACINIA
SALIVARY
GROOVE
FOOD CHANNEL
~~L ABIAL PALP
MAX IL L A R y r
PALP
~ MAX ILLARY
LOBE I
LABIAL MAXILLARY
MAXILLARY PALP PALP
CROSS SECTION I_ACINIA
EPIPHARYNX
FIG . 27. Head and mouth parts of a flea. See text for explanation of parts.
(Redrawn and adapted after vadous authors .)
(e) Fifth subtype, the louse fly. The It:mse flies, members of the
family Hippobos!Idae, have mouth parts closely related to those of the
third subtype, the stable fly; the characteristic tubular or cylindrical
hallstellum is adapted for penetration into the skin of the host. The
labrum- epipharynx is stylet-shaped; its proximal portion is strongly
chitinized and rigid, whereas the distal end is membranous and very
flexible. " The hypopharynx in the two common species, Pseudolynchia
canariensis (Macq.) = [Lynchia mama (Bigot)] and Melophagus
FIG. 28. Head and mouth parts of the honeybee (Apis mellifica). Both
types of mouth parts well developed, but the mandibles are used chiefly for
portage and modeling. (Hymenopteron type.) (A) front view of the head
showing (1) antennae, (2) compound eyes, (3) simple eye, (4) labrum,
(5) mandibles, (6) maxillae (galea), (7) labium (palpi only), (8) labium
(glossa), (9) bouton, (10) maxillary palpus. (B) mouth parts removed to
show (5) mandibles, (6) maxillae (lacinia), (7) labium (palpi only), (8)
labium (glossa), (9) bouton, (10) maxillary palpus, (11) prementum, (12)
mentum, (13) cardo, (14) stipes.
ovinus Linn., is nearly as long as the combined haustellum and labellum

and is a very slender and hyaline mouth part.
Siphonapteron type. The mouth parts of the Siphonaptera (fleas)
(Fig. 27), though typically of a piercing-sucking type, are peculiar to
this order of insects. The broad maxillary lobes bearing long palpi are
conspicuous landmarks (Fig. 27); the other organ (slender) bearing
long parallel palpi is the labium. The principal blade-like piercing organs
are a pair of independently moveable structures commonly referred to as
mandibles but are said to be maxillary laciniae by Snodgrass (lac. cit.)
and others. The mandibles are believed to be rudimentary in fleas. The
median stylet (unpaired) is said to be the epipharynx, not the labrum
of many authors, the labrum being difficult to demonstrate. The epi-
pharynx is said to be closely embraced by the lacinial blades. The three
stylets (a pair of maxillary laciniae and the epipharynx) are held in the
channel of the labium. The labrum is rudimentary, and the existence ot
a hypopharynx is not demonstrable. The wound is made by the protrac-
tion and retraction of the maxillary laciniae. As soon as the blood begins
to flow, it is drawn up into the pharynx by the action of both the cibarial
and the pharyngeal pumps.
Hymenopteron type. In this type the two general classes of mouth
structures, the mandibulate and haustellate, find full development in the
same species, though the mandibles are not involved in the feeding proc-
ess. The honeybee (Fig. 28) serves as a representative species. The
labrum is narrow and quite simple; the mandibles are eaSily distinguish-
FIG. 29. Head and mouth

parts of a butterfly (Vanessa
sp.). (A) side view: suctorial,
coiled tube, Lepidopteron type.
( 1) antennae; (2) compound
eye; (3) proboscis, consisting
only of galeae; (4) labial pal-
pus. (The labrum is not visible
in side view.) (B) section of
proboscis showing double
nature.
able and are useful wax implements. In ' ants the mandibles are highly
efficient organs of portage and weapon~ of defense. The maxillae form
the lateral conspicuous wings of the suctorial parts; the lacinia and galea
are fused, and the maxillary palpi are minute. The labium is represented
by the long structures to the right and left of the middle tube, which
is probably the hypopharynx. The hypopharynx terminates in the spoon-
like labellum or bouton which completes the lapping character of the
subtype.
Lepidopteron type. This type, represented by the commoner butter-
flies and moths, is typically a coiled, sucking tube capable of great
elongation. Taking the cabbage butterfly, Pieris rapae (Linn.), as an
example (Fig. 29) the labrum is seen to be greatly reduced, and the
mandibles are absent. (These may be weakly present in the lower
INSECT AND ARACHNID MGpTH PARTS 75
Lepidoptera.) The maxillae are apparently represented only by the
galeae, which by close approximation of their inner groovoo surfaces
form the long, coiled proboscis. The double structure of the proboscis
can be easily demonstrated by manipulation. The labium is represented
by the labial palpi.
Orders of Insects Arranged According to Type of Mouth Parts

I. Orthopteron type: <> Biting and chewing (mandibulate) mouth parts.
Orders Orthoptera, Blattaria, Dermaptera, Grylloblattodea, Phasmida,
Mantodea, Coleoptera, Isoptera, Odonata, Diploglossata, Plecoptera,
Zoraptera, Embioptera, Corrodentia, Mallophaga, Ephemerida (vesti-
gial mouth parts), Megaloptera, Neuroptera, Raphidiodea, Mecop-
tera, Trichoptera, Strepsiptera, Protura, Thysan';lra, Collembola, Ap-
tera.
II. Thysanopteron type: Rasping-sucking; biting in structure, but sucking
in function; right mandible reduced or wanting; represents a transi-
tional form between biting and sucking mouth parts.
Order Thysanoptera.
III. Hemipteron type: Elongated (beak-like), 3- or 4-segmented proboscis,
snugly enclosing stylet-like mandibles and maxillae; piercing and
suctorial.
Order Hemiptera (including Homoptera).
IV. Anopluran type: Piercing-sucking mouth parts lie in sac concealed in
head when not in use; mandibles vestigial; maxillae, hypopharynx,
and labium form functional stylets; circlet of evertible oral teeth
provides means of attachment when feeding.
Order Anoplura.
V. Dipteron types: Unsegmented proboscis, which mayor may not contain
piercing stylets (see previous description) .
Order Diptera.
VI. Siphonapteron type: Piercing-sucking mouth parts; principal blade-like,
piercing organs consisting of a pair of independently moveable
maxillary laciniae, which together with stylet-like epipharynx are
held within the channel of the labium.
Order Siphonaptera.
VII. Hymenopteron type: For feeding purposes the mouth parts are of a non-
piercing, lapping type; mandibles present, but used for combat and
portage.
Order Hymenoptera.
VIII. Lepidopteron type: Greatly elongated coiled sucking tube; nonpiercing.
Order Lepidoptera.
Arachnid mouth parts. In his recent excellent study of the feeding

organs of Arachnida, Snodgrass 3 points out that arachnids come from
Orthopteron is used here merely to indicate a biting-chewing type which varks
conSiderably within this grouping of the orders.
an ancestral line that never acquired organs for mastication and even
today have no true jaws, hence are forced to subsist on liquids . A liquid
DORSAL VIEW
CHELICERAL
SHEATH ~----~~-r'-~
SEGMENTS
OF THE
PEDIPALP
BASIS CAPITULI
CHEUCERAL SHEATH "'\11 /TEETH

VENTRAL V I / _
HYPOSTOME
BAS IS CAPITULI
c-n t ; 1'31 COXA
" .. / ."..../
FIG. 30. Tick, capitulum; showing mouth parts of a tick (Arachnida, Acarina).
diet requires an ingestion pump, and with all arachnids a highly
developed sucking apparatus constitutes the essential part of the feeding
mechanism. The chelicerae (a pair) are the first postoral appendages of
INSECT AND AHACHNlD MOUTH PAHTS 77
the arachnid (see Chapter XX), and while functioning more or less as
"jaws" they are not homologous with the mandibles of insects; they are
used for grasping, holding, tearing, crushing, or piercing. The leg-like
pedipalps are the second postoral appendages of the Arachnida and are
the homologues of the mandibles of mandibulate arthropods (Snod-
grass). These organs are modified in various ways, functioning as organs
of prehension, protection, and, in male spiders, as sperm-carrying organs.
In the scorpions the pedipalps are chelate and serve for catching, hold-
ing, and crushing the prey.
According to Snodgrass (loc. cit.) the only features of the acarine
(ticks and mites) mouth parts that cannot be homologized with struc-
tures generally present in other Arachnida are the variously developed
appendicular lobes or processes often associated with the distal part of
the hypostome (Fig. 30). Also among the trombidiform mites the
chelicerae become progressively adapted for piercing by a transforma-
tion of the moveable digits into hooks or stylets.
BIBLIOGRAPHY
I. Snodgrass, H. E., 1944. The Feeding' Apparatus of Biting and Sucking
Insects Affecting Man and Animals. "Smithsonian Misc. Collect.," vol. 104,
no. 7, (Publ. no. 3773). Washington, D. C.: Smithsonian lnst. 1I3 pp.
2. Jobling, B., 1926. "A comparative study of the structure of the head
and mouth parts in the Hippoboscidae (Diptera Pupipara) ," Parasitology,
18:319-49. .
3. Snodgrass, H. E., 1948. The Feeding Organs of Arachnida, Including
Mites and Ticks. "Smithsonian Misc. Collect.," vol. 1l0, no. 10. Washington,
D. C.: Smithsonian lnst. (Publ. no. 3944).93 pp. (29 figs).
CHAPTER VII
._.
COCKROACHES AND BEETLES
A. COCKROACHES
Order BlaUaria
Cockroaches of today are direct descendants of the huge ancient
cockroaches (Paleoblatta), some reaching about six inches in length,
which together with enoromous dragonHies (Protodonata) "had con-
quered the world"l some 250,000,000 years ago. Though much smaller
in size but still cosmopolitan, modern cockroaches have cbanged but
little in general structure. They are usually Hattened dorsoventrally with
smooth (sometimes pilose) tough integument, varying in color from
chestnut-brown to black in the more pestiferous house-invading species,
but commonly green, or orange and other colors, in tropical species. The
head is decidedly Hexed backward and downward when at rest. The
prominent antennae are filiform and many segmented. There are two
pairs of wings in most species; in some species the wings are vestigial, in
others the wings are well developed in the males and short in the females,
e.g., Blatta orientalis Linn. The outer pair of narrow wings (tegmina) is
thick and leathery, while the inner pair is membranous and folds fan-
like. While most cockroaches possess the power of Hight, they are typi-
cally runners (cursorial) and can move swiftly by means of their long
well-developed legs. They are highly gregarious and primarily nocturnal.
but most cockroaches may be seen occasionally singly during the day.
Metamorphosis is simple.
Feeding habits. The mouth parts of cockroaches are of the genera1ized
biting-chewing type (Orthopteron). They feed on a great variety of
foods (omnivorous), with preference for starchy and sugary materials.
They will sip milk, nibble at cheese, pastry, meats, grain products, sugar,
sweet chocolate-in fact, practically no edible material available for
human consumption is exempt from attack by these vile insects which
just as freely feed on book bindings, the sized inner lining of shoes, dead
insects, their own cast-off skins and dead and crippled kin, fresh and
dried blood, excrement, and sputum. They feed principally at night,
hence we live in ignorance as to their disgusting and dangerous feeding
habits.
78
Furthermore, roaches habitually disgorge portions of their partly
digested food at intervals and drop their feces wherever they go. They
also discharge a nauseous secretion from both the mouth and from glands
opening on the body, imparting a persistent and typical "cockroach" odor
to food and dishes with which they come in contact.
Species of sanitary importance. There are about 2,250 species of cock-
roaches belonging to some 250 genera heretofore placed in the Family
Blattidae of the Order Orthoptera, now generally put into the Order
Blattaria. Cockroaches occur throughout the world but are chiefly
tropical. World-wide distribution of certain species has been effected by
maritime trading; holds of vessels, the galleys, and crew's sleeping quar-
\I
FIG. 31. The German cockroach, Blatella gennanica, in various stages of

development. The adult female is shown with egg case or ootheca in normal
position protruding from the terminal abdominal segment. X 2.
ters are often overrun with cockroaches. Among the better known species
in~ading the household, restaurants, hotel kitchens, etc., particularly in
neotropical regions, are described below:
Blatella germanica (Linn.) , known 2.S the German cockroach, water
bug, or croton bug, belongs to the Family Phyllodromiidae. It is the best
known and probably the most widely distributed species. It is a small
species, native to Europe, measuring from 14 to 16 mm in length, and is
pale yellowish-brown in color with two dark brown longitudinal stripes
on the pronotum. Both sexes are fully winged. The female carries the egg
capsule partly protruding from the tip of the abdomen (Fig. 31) until
hatching time.
Blatta orientalis (Linn.), known as the Oriental cockroach and also
as the "black beetle," is very much darker than the German roach. It is
dark brown to black in color, whence its common name "black beetle"
(Fig. 32). It is about 25 mm in length; the wings of the female are rudi-
mentary, and those of the male do not quite reach the tip of the abdo-
men. Enormous colonies of the Oriental cockroach are often found in
damp basements where food is available. This species belongs to tllP
Family Blattidae.
Periplaneta americana (Linll.), commonly called the American cock-
roach (Fig. 33), is a very large (30-40 mm in length) chestnut-brown
species, native to "'lexica and Central America, but now widely distrib-
uted over the earth. Both sexes have long wings, which are frequently
llsed in flying short distances. This species also belongs to the Familv
Blattidae.
~ D
FIG. 32. The Oriental roaGh (female), Blatta orientalis. X 1.3.
FIG. 33. The American cockroach, Periplaneta americana. X 1.3.
Periplaneta australasiae (Fabr.), while commonly called the Austra-

lian roach, though probably not indigenous to Australia, is, like ameri-
cana, a cosmopolitan species. It is reddish-brown in color resembling
americana, but has a strong straw-colored streak extending about one-
third of the way down the outer margins of the wing covers (tegmina)
as well as a yellow area around the margin of the pronotum forming a
double dark area on the dorsum. The wings are well developed in both
sexes. This species, too, belongs to the Family Blattidae.
Supella supellectilium Serv., known as the brown-banded or tropical
cockroach, resembles the German cockroach in appearance but is smaller
(10-12 mm) and has tvvo yellow cross bands, one at the base of the
wings and the other about Vi \3 of an inch farther back. The wings do not
quite reach the tip of the abdomen; adults fly readily when disturbed.
The species is a decidedly gregarious one. Unlike the German cockroach,
which confines its activities to the kitchen or around water and heat
pipes, the tropical cockroach hides in cupboards and pantries, invades
all rooms of the house, and seems to prefer "high locations, such as
shelves in closets, behind pictures and picture moldings, and the like."
This species, like the German . cockroach, belongs to the Family
Phyllodromiidae. It is rapidly becoming cosmopolitan.
Leucophaea surinamensis (Linn.), the dusty-tail roach, or Surinam
cockroach, is a dark brown to blackish tropical species measuring from
14 to 20 mm in length. It is a burrowing species, burrowing under piles
of debris, leaves, etc. It, too, belongs to the Family Phyllodromiidae.
FIG. 34. Egg cases (oothecae) of

cockroaches: (left ) Oriental roach,
(right) croton bug. X 3. .
Panchlora cubensis Sauss. is the green Cuban roach which is fre-

quently brought to the ports of many lands in bunches of bananas from
Central America.
Tokuichi ShirakF lists 68 species of cockroaches for Japan, many of
these newly described.
Life history. The eggs of cockroaches are assembled in leathery bean-
shaped capsules, ootheca (Fig. 34), within the body of the female and
are thus carried for some time, often weeks , usually partly protruding
from her abdomen until just before hatching, when they are dropped.
There are several tropical species belonging to the Family Panch10riclae
which are ovoviviparous, i.e., give birth to living young. The number of
eggs within each capsule varies with the species. For Periplaneta ameri-
cana the number is usually 16, and occasionally 18 to 24 (Gould and
Deay)o; the normal number for Blatta orientalis is also 16, while the
number per capsule for Blattella germanica varies between 30 and 40
with a maximum of 48; and for Supella supellectili1l11l the average is 18.
82 MEDICAL ENTOMOLOCY
The latter species drops the egg capsule after a day or two and attaches
it to some bit of debris. The Oriental roach may do this too. Many egg
capsules are produced during the lifetime of the female cockroach, e.g.,
as many as 90 by the American cockroach, 18 by the Oriental, and but
4 to 6 by the German (Gould and Deay, loco cit.).
The length of the incubation period varies with temperature and
humidity. At a constant temperature of about 86° F and a relative
humidity of about 70 per cent, Gould .and Deay (loc. cit.) found the in-
cubation period for the American cockroach to be 31.8 days, at room
temperature it averaged 52.9 days. For the Oriental cockroach the same
authors found the incubation period at room temperature to vary from
42 to 81 days; for the German cockroach under similar conditions (room
temperature averaging 76° F) it was 28.4 days. The eggs of this latter
species hatched in 16 days at a temperature of 88° F or higher. The
brown-banded roach required 90 days to hatch at room temperature
of 73° F and 49 days at 82° F.
On hatching, the young cockroaches are almost white and quite wing-
less; the skin is cast on emergence, with a second molt usually in three or
four weeks, followed by other molts at intervals of several weeks until
maturity is reached. The American roach may have as many as 13 molts,
with wing pads appearing in the third or fourth molt; maturity being
attained in from 285 to 642 days; however, Gould and Deay report as
high as 971 days under similar conditions for one specimen. Rau 4 reports
the developmental period for the Oriental roach as one year. For the
smaller species, such as the German cockroach, the developmental period
is much shorter; an average of about two months (50 to 60 days accord-
ing to Seamans and Woodruff5; and 90 to 95 days at room temperature
according to Gould and Deay). There are normally six molts, but there
may be seven under adverse conditions. This permits the development of
two or more generations a year for the German cockroach.
The longevity of the American cockroach is reported by Gould and
Deay to range from 102 to 588 days under room conditions, and the
complete life span of three females of this species is reported to have
been 783, 793, and 913 days respectively. The mean length of life of
female German cockroaches is reported by these authors to be 200 days,
with a maximum of 303 days.
Cockroaches as vectors. Herms and Nelson 6 by means of a simple
bacteriological experiment showed that cockroaches (Blatella germanica)
can acquire specific bacteria by crawling over cultures and then deposit
the bacteria on food, e.g., by crawling over sugar. They also showed that
these roaches carried a minimum of 13,370 bacteria. There were more
bacteria on the hind pair of legs than on the other two pairs of legs,
probably due to the fact that the insect's hind legs are more constantly
in contact with surfaces and that the forelegs are very frequently brushed
clean by means of the antennae. The number of bacteria carried by a fly
ranges from 550 to 6,600,000 according to Esten and Mason,7 hence,
potentially at least, the fly is a more effective carrier of microorganisms.
Longfellow8 has shown that Escherichia coli (Migula), Proteus vul-
garis Hauser, Staphylococcus aureus Rosenbach, S. citreus (Migula),
and a bacillus of the subtilis type are carried on the legs of roaches and
that the same organisms are found in the feces, passing uninjured
through the alimentary tract. However, the most important results have
been obtained by Barber,9 experimenting with the roach Periplaneta
americana (Linn.). He fed dejections of cholera patients to roaches and
found that a single insect would frequently ingest as much as 0.2 cc. In
eight cases cholera vibrios were recovered from the insect's feces and in
one case as long as 79 hours after feeding. After the vibrios were dis-
charged by the roach on such moist materials as fresh beef, lettuce, etc.,
they remained mobile for at least 16 hours, and no loss of virulence was
observed when the vibrios remained in the insect's intestine as long as
29 hours. Barber also found that the roaches disgorged portions of their
meal at intervals after feeding and that such disgorged material contained
cholera vibrios.
Cockroaches as intermediate hosts of nematode parasites. It was very
early known that cockroaches may become infected with Spirura rytip-
leurites (Deslgch.) of the rat by feeding on rat feces, and that other rats
may become infected in turn by feeding on infected roaches. Caleb 10
( 1878) reports that he discovered numerous parasites in the adipose
tissue of the roach, Blatta orientalis Linn., which were identical with
nematodes found in the rat, Rattus r. norvegicus. He also observed hair
of the rat in the alimentary canal of the roach. On fe~ding rats (Rattus
r. rattus) on infected roaches and examining them after the expiration
of eight days, he found the parasites in the folds of the mucous mem-
brane of the rats' stomachs. Several nematodes (three females and one
male) had already developed sexual organs.
According to Fibiger12 much evidence is at hand to prove that certain
nematode parasites (Spiroptera) of the rat produce true malignant
tumors. Eggs of these nematodes were found in the rat feces but no em-
bryos, and he reports that none developed for six months and also that
rats could not be infected with the eggs, the roach being a necessary
intermediate host. Two sp,ecies of roaches were used, namely, Blatta
orientalis Linn. and Periplaneta americana (Linn.). Ransom and Hall l l
found that although Aphodius beetles are the normal carriers, the croton
bug, Blatella germanica (Linn.), might also serve as an intermediate host
for Gongylonema pulchrum Molin, a nematode parasite of sheep and
cattle. Referring to the discovery of Fibiger relative to sarcoma in rats
1,1EDlCAL ENTOMOLOGY
cited above and based on bis (Sambon' s) epidemiological studies on

human cancer Sam bon (1925, loco cit., Chapter IV) concluded:
Similar to the role of the filaria in elephantiasis is, no doubt, that of

gongylonema in cancer. Like the filaria, so the gongylonema is not the direct
essential cause of the disease it gives rise to, but it probably is, in certain
localities, an etiological factor of considerable importance. The gongylonema,
known to be a cancer-producing agent in the rat, seems an equally likely
factor in the incidence of cancer among human beings in certain regions,
such as the Romagna and the Trentino. Here the disease affects principally
the upper portion of the alimentary tract and more especially the buccal
cavity, the tongue, the oesophagus, the larynx, and the cardia (situations
invaded by these forms) and here cockroaches, meal beetles, and cellar beetles
(certain beetles such as Tenebrio and Blaps of the family Tenebrionidae are
suitable intermediary hosts of these worms) are an ever present vermin in the
old houses in which such cancer cases occur. These observations so clearly
designative of a relationship between worm and neoplasm, are strengthened
by the fact that gongylonema is present and common in the majority of the
domestic animals of these regions.
Although Sam bon dwells particularly on Gongylonema he points out

that other helminths bear a similar relationship, each according to the
organ which is usually invaded and in which irritation is set up, thus
inviting the cancer factor. It is pointed out that cockroaches, either in-
tact or in fragments, are only too frequently found in bread and other
food which may be insufficiently baked to kill the encysted Gongylonema.
Cockroaches appear to lend themselves to many species of parasitic
nematode worms (see Chitwood 1S ).
Cockroach control. Cockroach infestations are generally an indication
of a breakdown in good housekeeping practices, whether in the home, a
restaurant, a hotel, or a grocery store. There are many ways in which
these insects (in any stage of development) may be brought in to start
an infestation, e.g., in boxes of groceries, in sacks of produce from ware-
houses, in freight shipments. They may come in under their own power
at night from neighboring foci; they may follow water pipes, sewers, etc.
Many homes and nearly all restaurants, grocery stores, etc., have a few
cockroaches now and then. Kill the individual intruders at once in any
convenient way; do not let them gain a foothold by providing them with
food, because of poor garbage disposal or other poor housekeeping
practices. See to it that there are no convenient hiding places in cracks
in walls, etc.; use plastic wood to fill up such cracks. Dark, moist, clut-
tered-up situations favor cockroaches; open, clean and neatly kept spaces,
permitting plenty of daylight are not inviting to these darkness-loving
pests.
The abatement of infestations generally calls for the application of
an appropriate insecticide. The use of sodium fluoride is one of the most
effective methods for the control of roaches, of all species. It is, however,
toxic to man and other animals and hence must be used with caution.
As an insecticide it is usually given a bluish color so that it will not be
confused with flour, soda, etc. Do not store this insecticide in the kitchen
or pantry, and do not dust sodium fluoride in cupboards or where food
can be contaminated with it. The container should be labeled "Poison."
For use against cockroaches a mixture is made of sodium fluoride (25
per cent by volume) and tested pyrethrum powder (75 per cent). This
mixture is applied by means of a dust gun to all cracks and crevices and
places where cockroaches may be in hiding or where they congregate.
Applying the powder in the evening is advised, and the application
should be repeated in a week or two. The insects come in contact with
the dust when running over the treated surfaces, and afterward they
clean themselves and swallow the particles adhering to their feet and
bodies; thus the sodium fluoride acts as a stomach poison. Various other
materials may be used as substitutes for this insecticide, such as finely
powdered borax and boric acid, which may be dusted safely in cupboards
and kitchens. This powder is effective only as long as it is dry; it kills
very slowly.
DDT is useful in the control of all the common house-invading
cockroaches, but Blatella germanica, the croton bug, requires somewhat
special treatment. Using the same methods of application as for sodium
fluoride, force a 10 per cent DDT powder (in 90 per cent powdered
pyrophyllite) into cracks and crevices and over surfaces where cock-
roaches crawl. Against the German roach precede the dusting with a 5
per cent DDT-xylene spray. Apply a 5 per cent oil solution of DDT
spray to the undersides of shelves, sinks, drawers, and tables. Do not
allow DDT to fall on dishes, food, or cooking utensils. Outside areas,
dooryards, the vicinity of garbage cans, and other places where cock-
roaches lurk should be dusted with DDT.
Chlordane (empirical formula, C 1o H 6 ClR ) ranks high as a cockroach
poison when used as a 2 to 5 per cent dust. Its "knock-down" powers, in
common with some other newer insecticides is low, hence formulations
should contain pyrethrum or one of the thiocyanates. In oil formulations
used as sprays, 2 per cent chlordane is recommended. Precautionary
measures with reference to food, dishes, cooking utensils, etc., must be
exercised as for DDT.
B. BEETLES
Order Coleoptera
The Order Coleoptera consists of over 250,000 described species of
beetles, comprising 40 per cent of all known members of the Class In-
secta. Very few of the families of beetles concern the medical entomolo-
gist directly, but because of their abundance and successful invasion of
all sorts of environments it is certain that contact will be made with mem-
bers of this Order of insects sooner or later. Familiarity with the Order
is important.
Characteristics. Beetles are readily distinguishable from all other in-
sects. Their integument is leathery; their mouth parts are strongly man-
dibulate, i.e., biting-chewing. Although wings may be absent in some
species, there is usually a pair of wings present; the forward pair, elytra,
not used in flight are horny, and when at rest meet in a straight line down
the dorsum; the hind wings are membranous and functional, often fold-
ing both horizontally and vertically. Metamorphosis is complex (egg,
larva, pupa, imago), with hypermetamorphosis in certain species. The
larvae are commonly known as "grubs" and may usually be recognized
by the presence of three pairs of fairly well-developed legs; however,
the larvae of weevils are apodous.
Scavenger beetles. All scavenger beetles, of which there are several
families, are potentially of some public health importance since their
habit of feeding as larvae or' adults on dead animals, hides, or other
animal matter may accidentally bring them in contact with pathogenic
organisms. They may carry infeGtion in at least two ways, either me-
chanically on their legs, mouth parts, or body, or in their excreta after
feeding on infectious material. Destruction of carcasses by incineration
is urged.
Nuttall (lac. cit., p. 4) reports that, as early as 1894, "Proust in ex-
amining goatskins taken from anthracic animals, found quantities of
living Dermestes vulpinus Fabr. upon them. He found virulent anthrax
bacilli in their excrements, as also in the eggs and in the larvae. It is
evident from this that these insects which feed on the skins permit the
anthrax spores to pass uninjured through their alimentary tract. Heim
( 1894) also had occasion to examine some skins which were suspected
of having caused anthrax in three persons engaged in handling the
leather. He found larvae of Attagenus pellio Linn., Anthrenus museorum
Linn. (both Dermestidae), and Ptinus; also funy developed insects of the
latter species on the skins. All these insects had virulent anthrax bacilli
( spores) on their surface and in their excreta, from which Heim con-
cludes they might spread disease. He says the excreta are very light and
easily scattered by the slightest current of air. Heim does not believe the
_bacilli multiply in the bodies of these insects, but that the latter may be
dangerous through their scattering the spores about."
Among the families of scavenger beetles are the following:
1. Family Staphylinidae, commonly known as rove beetles, includes many
species that feed on carrion, dung, and decaying animal matter; they are
characterized by the abbreviated condition of the wing covers (elytra), which
leaves much of the abdomen exposed and gives these beetles a larval or worm-
like appearance; the abdomen is flexible and is often curled VP and thrown
forward dorsally; the long functional wings are folded up and concealed be-
neath the short wing covers when not in use. The scavenger species are com-
monly seen on turning over carrion, hides, heaps of bones, etc. Among these
are members of two representative genera, Creophilus and Staphylinus (Fig.
35).
2. Family Silphidae, commonly known as carrion beetles, burying beetles,
or sexton beetles, are as a rule attracted to dead animals which they undermine
and, in the case of small ones, may even bury. They deposit their eggs Oil
the dead animals , and the larvae feed on the decomposing flesh . Two genera,
Silpha and Necrophorus (Fig. 36 ) illustrate the familv.
35A 35B 36A 36B
FIG. 35. Rove beetles (Staphylinidae) ; (A) Creoplzilus, ( B) Staphy-

linus. X 1.5.
FrG. ,'36. Sexton beetles (Silphidae); (A) Silpha americana, (B) Necrop-
horus sp. X 1.5.
3. Family Dermestidae, hide beetles, larder beetles, museum pests, der-

mestids, are small oval or elongate beetles, often mottled, grayish, or brownish
in color. The hairy larvae as well as the adults feed on dead animals, museum
specimens, wool, cured meats, cheese, and many other animal as well as
vegetable products (cereals). Dermestes lardarius Linn. is known as the larder
beetle; Dermestes vulpinus Fabr. is commonly used to clean dried flesh from
bones for museum use; Anthrenus scrophulariae (Linn.) is known as the
carpet beetle, a serious household pest. Larvae of the carpet beetle may enter
the ears of persons and cause much discomfort (Fig. 37).
Beetles as intermediate hosts of helminths. Many species of beetles

serve as intermediate hosts of helminthic parasites of man and of wild
and domesticated animals. This common relationship is no doubt due
to the variety of feeding habits of beetles which enables them to ingest
fecal matter in which eggs of intestinal parasites of animals commonly
occur; thus many cereal and omnivorous feeders, as well as coprophagous
beetles may readily lend themselves as intermediate hosts. As suggested
previously (see cockroaches) the infectiv~ stage of nematode worms of
the genus Gongylonema belonging to the family Spiruridae commonly
88 MEDICAL ENT0110LOGY
occur in dung beetles belonging to the beetle family Scarabaeidae, such

as members of the genus Aphodius; also in meal worms, belonging to the
family Tenebrionidae such as Tenebrio molitor Linn. Gongylonema
/lulchrllln J\lolin parasitizes many species of mammals, such as goats,
swine, and occasionally man. It invades tissues of the oral cavity and
esophagus, causing gongylonemiasis. Adult females measure about 145
mm in length and are. very slender. Eggs are evacuated in feces and lie
dormant until ingested by an appropriate insect in whose gut the larva
hatches and comes to lie encapsulated in the body cavity until the insect
is ingested by an appropriate vertebrate host.
May beetles or cockchafers (family Scarabaeidae) are known to be
intermediate hosts both in the larval and adult stages of the thorn-headed
worm, Macracanthorhynchus hirudinaceus (Pallas) [= GigantorhYI1-
FIG. 37. The vm-ied carpet beetle ,

Anthrenus verbasci; showing larva
and adult.
I
chus gigas (Bloch)], a parasite of swine. also said to occur in man in

rare instances. This nematode worm in its adult stage measures from 20
to 30 cm in length and about 3 to 5 mm in thickness, and inhabits the
small intestine of its host. The eggs are deposited in the intestine and
pass out with the feces , which may be swallowed by larvae of cock-
chafers. These are often extremely abundant among the rootlets of grass
in heavily sodded pastures, and swine with free range are fond of these
grubs, in search of which they diligently root up the soil with their
snouts. Thus every opportunity is given for the grubs to become infected
and in turn the swine.
After the ova have been ingested the larvae hatch in a few days
within the intestine of the insect and proceed to burrow through the in-
testinal wall and into the muscles, where they are said to encyst them-
selves. In Europe the intermediate insect host is commonly M elolontha
melolontha (Linn.) or Cetonia aurata (Linn.). May beetles of the genus
Lachnosterna (according to Stiles, Lachnosterna arcuata Smith, and
others) are probably all more or less concerned. The life of nearly all
May beetles is quite long, the larval stage alone often requiring nearly
three years. In districts infested with the thorn-headed worm a systematic
crusade against cockchafers would be the logical means of control, to-
gether with the treatment of swine.
Numerous species of beetles have been proved to be intermediate
hosts of the fowl tapeworm, Raillietina cesticillus (Molin). The species
listed by Reid, Ackert, and Case l l belong to the following families,
Scarabaeidae, two species; Tenebrionidae, one species; Carabidae, sub-
family Harpalinae, 26 species, to which they added 12 not previously
reported, giving a total of 38 species in this family. The beetles belonging
to the genus Amara proved to be particularly favorable hosts although
the largest number of cysticercoids were produced by a species of
Pterostichus, a total of 626 by one beetle which had been fed on four
proglottids.
The adult tapeworms, which measure from 10 to 12 cm in length,
are generally attached to the lumen of the upper third of the chicken
intestine. The terminal gravid proglottids \vhich break off pass from the
body of the host with the feces and continue to be motile for a short
time after evacuation. If the proglottids are eaten by appropriate species
of beetles the hexacanth embryos develop, penetrate the intestinal wall
of the host, and become mature cysticercoids in the body cavity of the
beetle in from 14 to 21 days or longer, usually in 14 to 16 days during
the hot part of the summer. If the beetles are now eaten by chickens
the mature ovoid yellowish-brown cysticercoids, which measure from
363p. to 521p. in length by 199fJ. to 398fJ. in width, according to the above-
mentioned investigators, are freed in the duodenum of the birds and in
about two weeks adult tapeworms are produced. The onchosphere stage
of R. cesticillus can be distinguished from onchospheres of other species
of fowl tapeworms by two funnel-like structures in the membranes which
surround the hexacanth embryo, (Reid, Ackert, and Case).
Canthariasis is a term used to designate the rare accidental beetle
(larva or adult) infestations of organs of the body, e.g., infestations of
the alimentary canal by larvae of the churchyard beetle, Blaps mortisaga
Linn., as the result of superstitiously drinking foul graveyard water in
which these beetles may occur. The ingestion of meal- and flour-infest-
ing insects such as the meal worm, Tenebrio molitor Linn., has also been
known to occur.
Vesicating beetles. Many beetles possess a vesicating substance in
their body tissues which produces blisters when coming in contact with
the skin. The family Meloidae, commonly referred to as the "blister
beetles," contains the species which are the important source of can-
tharidin used medicinally. Among these species are the Spanish fly.
Lytta vesicatoria (Linn.) the telini fly, M ylabris cichorii (Linn.). and
Epicauta hirticornis (Haag-Rlftenberg) of India. At least two species of
the family Oedemeridae, Sessinia collaris (Sharp) and S. decolor Fairm.,
known as coconut beetles by the Gilbertese, cause severe· blistering if
the skin is contacted. (See Chapter XXII.)
Rose chafers poisonous to poultry. The rose chafer, Macrodactylus
subspinosus (Fab.), family Scarabaeidae, was found by Lamson15 to be
poisonous to chicken, ducklings, goslings, and young turkeys. He reported
that chickens fed in quantity on rose chafers showed the following symp-
toms after from four to five hours: "Drowsy appearance, with wings
dropping. the eyes closed, and a slight shaking of the body. This drowsi-
ness increased, leg weakness developed, until the chickens no longer
stood, but slept resting on their feet and legs, and later died. Death oc-
curred usually from five to twenty-four hours after they had fed upon the
rose chafers. Convulsions occurred in about five per cent of the deaths.
Chickens that survived the twenty-four-hour period after eating rose
chafers seldom died from the poison, though they did not seem alto-
gether normal for several days afterward."
Sundry annoying beetles. Aside from any economic or other injury for
which beetles may be responsible, the sheer abundance of certain species
at times causes annoyance. For example, while trying to do microscopic
work or write reports by lamp light the author, stripped to the waist
during warm evenings in the tropics, was greatly disturbed by the pres-
ence of hordes of the so-called copra "bug" (red-legged ham beetle),
Necrobia rufipes Fabr., family Corynetidae. 16 These small greenish-blue
beetles, 3-5 mm long, originated in vast numbers in copra stored in
neighboring sheds. The larvae of this species may also enter the ears of
persons.
Tiny (3 mm long) saw-toothed grain beetles, Oryzoephilus surinam-
ensis (Linn.), belonging to the family Cucu;idae, may invade bed cham-
bers in great numbers and crawl over the bodies of occupants and nibble
the skin. An infestation of this kind was traced to the bathroom and
thence out of the house through the yard and into an old barn where,
under the stalls, grain from the manger had collected, affording a breed-
ing ground for the beetles. Extreme dryness had apparently driven the
insects to the bathroom for moisture and the attack on occupants of the
adjoining bedchamber was merely incidental.
Minute species of Staphylinidae (such as Atheta occidentalis) Bern-
haeur, a blackish species 3 mm long, are often encountered on the wing
in the late autumn and may accidentally enter the eyes, causing a severe
burning sensation, and temporarily blinding the victim. S:uch a mishap
to a person driving a motor car might lead to a serious accident. These
minute species breed in cow dung and decomposing plant refuse.
The so-called tule beetle, or stink beetle [Platynus maculicollis
(Dej.)], of California is nQrmally a beneficial predator, but when its
natural habitat in the marshes becomes dry in summer it commonly
leaves in search of moisture and may invade homes in the neighborhood.
Heavy early winter rains and cold weather may likewise cause invasions.
The nauseous odor of this beetle is almost int<1lerable. It belongs to the
family Carabidae which contains many other species possessing vile
odors.
Beetles as parasites. Belonging to the family Platypsyllidae, indeed
the only representative of this family, is the coleopterous parasite of the
beaver, Platypsyllus castoris Ritsema (1869). This is a permanent, oblig-
ate parasite in all its stages. The eggs are deposited on the skin of the
beaver among dense hairs. It occurs on these animals in both Europe
and North America.
Another family of Coleoptera, the Leptinidae, includes beetles which
are parasitic on beavers, and certain other rodents. The three known
species are Leptinus testaceus Muller, parasitic on mice and shrews in
Europe and North America; Leptinillus validus (Horn) found on North
American beavers; and Leptinillus aplodontioe Ferris, taken on Aplodon-
tia, a genus of rodents known as mountain beavers, peculiar to the Pacific
coast (Ferris) Y .
Key to the Families of Coleoptera Referred to in this Chapter O
1. Antennae variable in structure but never lamellate 2

Antennae with a club composed of from three to seven leaf-like seg-
ments (lamellae); legs usually fossorial; tarsi five-segmented. (May
beetles, .June beetles, Cockchafers) .Scarabaeidae
2. Anterior and intermediate tarsi five-segmented; posterior tarsi four-seg-
mented ............. ............ ... 3
All of the tarsi either three-, four-, or five-segmented ..... ......... ..... 4
3. Head as wide as prothorax; anterior coxal cavities open behind; body
not heavily chitinized (Blister beetles) . . Meloidae
Head narrower than prothorax; anterior coxal cavities closed behind;
body usually heavily chitinized (Darkling ground beetles)
..................... .................... . ... Tenebrionidae
L Elytra long, covering most of abdomen; abdominal tergites mostly
membranous ..... ................ 5
Elytra short, exposing most of abdomen; abdominal tergites mostly
chitinous . .. .. .... .................... .. ........ ........ . ... . ..... .... .. ... ... .. .... .... . . ... 8
.5. Fourth segment of tarsus distinct, free; antennae clavate, capitate or
moniliform . . ... . .... ... .. ... ... .... .. ..... ............... 6
Fourth segment of tarsus very small, fused with fifth segment; antennae
usually filiform; body often brightly colored (Leaf beetles) .......... .
........................ ............... Chrysomelidae
Prepared by Dr. E. Gorton Linsley.

MEDICAL ENTOMOLOGY
6. Abdomen with five visible sternites. 7
Abdomen with six visible sternites; tibial spurs large; antennae gradually
thickened or clavate (Carrion beetles, Burying beetles) .... Silphidae
7. Anterior coxae conical,. prominent; antennae moniliform; fiat, nonscaly
beetles (Flat beetles, Grain beetles) . Cucuiidae
Anterior coxae globular, not prominent; antennae capitate or clavate;
convex, scaly beetles (Leather beetles, skin beetles). Dermestidae
8. Wingless; eyes wanting or abortive.. ............. ... .. 9
Winged; the wings folding under the short elytra; eyes usually well
developed; tarsi from three- to five-segmented .......... Staphylinidae
9. Anterior coxae globular (Beaver beetles) ...................... . Platypsyllidae
Anterior coxae flat (Rodent beetles). Leptinidae
BIBLIOGRAPHY
1. Cockerell, T. D. A., 1927. "The carboniferous insects of Maryland,"
Ann. & Magazine of Nat. History, 19:385.
2. Shiraki, Tokuichi, 1931. "Orthoptera of the Japanese Empire. Part
II Blattidae, Insects Matsumurana," Taihoku Imperial Univ. Entomolog. Lab.
Contrib. no. 29, 5 (Part 4):171-209.
3. Gould, George E., and Deay, H. 0., 1940. "The biology of six species
of cockroaches which inhabit buildings." Lafayette: Purdue Univ., in Agric.
Exper. Sta., Bull. no. 451, 31 pp.
4. Rau, P., 1924. "Biology of the roach," Tr. Acad. Sc. St. Louis, 25:57-
79.
5. Seamans, Lois, and Woodruff, Laurence C., 1939. "Some factors in-
fluencing the number of molts of the German roach," J. Kansas Entomolog.
Soc., 12:73-76.
6. Herms, W. B., and Nelson, Y., 1913. "The croton bug (Ectobia
germanica) as a factor in bacterial dissemination," Am. ]. Pub. Health, 3:929-
34.
7. Esten, W. N., and Mason, C. J., 1908. "Sources of bacteria in milk,"
Storrs Agric. Exper. Sta., Bull. no. 51, pp. 94-98.
8. Longfellow, R. C., 1913. The common house roach as a carrier of
disease. Am. ]. Pub. Health, 3:58-61.
9. Barber, M. A., 1914. "Cockroaches and ants as carriers of the vibrios
of Asiatic cholera," Philippine]. Sc., sec. B, 9:1-4.
10. Galeb, Osman, 1878. "Observations et experiences sur les migrations
du Filaria rytipleurites, parasite des blattes et des rats," Compt. rend. Acad.
d. sc., 87:75-77.
11. Ransom, B. H., and Hall, M. C., 1915. "The life history of Gongylon-
ema scutatum," J. Parasitol., 2:80-86.
12. Fibiger, Johannes, 1913. "Ueber eine durch Nematoden (Spiroptera
sp. n.) hervorgerufene papillomatOse und carcinomatOse Geschwulstbildung
im magen der Ratte," Berl. klin. Wchnschr., 50:289-98.
13. Chitwood, B. G., 1932. "A synposis of the nematodes parasitic in in-
sects of the family Blattidae," Ztschr. fpkarasitenkunde, 5:14-50.
14. Reid, W. M.; Ackert, J. E.; and Case, A. A; 1938. "Studies on the
life history and biology of the fowl tapeworm, Raillietina cesticillus (Molin) ,"
Tr. Am. Micr. Soc., 57:65-76.
15. Lamson, G. H., Jr., 1922. "The rose chafer as a cause of death of
chickens." Storrs Agric. Exper. Sta. Bull. no. llO, pp. 117-34.
16. Herms, Williaril B., 1926. "Diocalandra taitensis (Guerin) and other
coconut pests of Fanning and Washington Islands," Philippine J. Sc., 30:243-
71.
17. Ferris, G. F., 1918. "An apparently new species of Leptinillus," Ganad.
Entomologist, 50:125-:28.
. _.
CHAPTER VIII
THE BUGS
BEDBUGS, CONENOSES, AND OTHER BUGS

Order Hemiptera
Order Hemiptera. The Hemiptera comprise all the insects, about
48,000 species, included in the Heteroptera and the Homoptera. The
order Hemiptera is divided into two suborders: ( 1) H eteroptera, in
which the fore pair of wings (hemelytra) is usually thickened at the base
and the distal overlapping portion is membranous, and (2) Homoptera,
in which the fore pair of wings is usually of about the same thickness
(membranous) throughout.
The suborder Homoptera includes such important phytophagous
families as the Aphididae (plant lice), Cicadidae (cicadas or harvest
flies), Cicadellidae (leafhoppers, sharpshooters), Membracidae (tree-
hoppers) and many others of great agricultural importance, particularly
many important vectors of plant diseases. These families include plant-
feeding insects with piercing mouth parts, such as leafhoppers and tree-
hoppers, many of which have been reported as biting and sucking blood
from human beings. Usingerl attributes this rather rare phenomenon of
bloodsucking in the normally phytophagous groups of the Hemiptera to
three influences, namely, the "stimulus of artificial light or other unusual
conditions of the environment, the attractive qualities of exposed liquids,
mainly perspiration, and hunger." He remarks further that this change
"from plant feeding to bloodsucking, is not such a profound one as would
at first be supposed. This is evidenced by a comparison of the composi-
tion of plant juices and blood and by the various plant-feeding groups,
some members of which have adapted themselves to a predaceous habit
or have shown their ability occasionally to suck the blood of mammals."
The Hemiptera-Heteroptera are the true bugs and are characterized
by a jointed suctorial proboscis attached anteriorly, which, when not in
use, is commonly flexed under the head. The winged members of this
order normally have the wing covers, hemelytra (fore pair of wings),
thickened at the base and the distal overlapping portion more or less
membranous. The true bugs are separated into two divisions, (1) the
94
THE BUGS 95
Gymnocerata, in which the antennae are conspicuous and capable of
being moved freely in front of the head, e.g., Cimex lectularius Linn.,
the common bedbug; Anasa tristis ( DeGeer ), the squash bug; and
Triatoma protracta (Uhler), a conenose; and (2) the Cryptocerata, in
which the antennae are concealed in small concavities (foveae) and are
closely pressed to the under side of the head, e.g., Lethocerus americanus
(Leidy), the giant water bug. Metamorphosis is simple.
A. THE BEDBUGS
Family Cimicidae
Family Cimicidae. The family Cimicidae (Acanthiidae) iy.cludes the
bedbugs, swallow bugs, and the poultry bug, characterized by a very
short, broad head, broadly attached to the prothorax, an oval body, well-
developed compound eyes, absence of ocelli, four-jointed conspicuous
antennae, three jointed proboscis lying in a groove beneath the head
and thorax, very short pad-like hemelytra. The bodies are broad and flat,
enabling the bugs to creep into narrow crevices. A nasty pungent odor
is attached to the group as a whole with few exceptions. They are night-
prowling and bloodsucking in habit, some feeding on birds and bats
and others on human beings. Peculiar to these bugs is the organ of
Berlese located in the IV or V abdominal segments. The presence or
absence of this organ and its particular location when present provides
a character useful in identification of species.
The three principal genera of the family Cimicidae 2 are: (1) Cimex,
e.g., C. lectularius L., the cosmopolitan bedbug; C. hemipterus (Fabr.)
[C. rotundatus (Sign.)], tropical and subtropical, also known as the
Indian bedbug; C. boueti Joyeux,3 the tropical bedbug of Africa and
South America, all of which resemble each other very closely as do the
following: C. columbarius Jenyns, parasitic on pigeons in Europe, C.
pipistrelli Jenyns, parasitic on bats in Europe, and C. pilosellus (Horv.)
parasitic on bats in North America; (2) CEciacus, in which the body is
clothed with long silky hairs, the filiform third and fourth joints of the
antennae only a little thinner than the first and second, e.g., CEciacus
hirundinus (Jenyns), the European barn-swallow bug and O. vicarius
Horv., the corresponding American species; (3) H aematosiphon, in
which the rostrum is long, reaching to the posterior coxae, of which
Haematosiphon inodora (Duges) is the only known species. This species
infests poultry in the southwestern United States and in Mexico and is
often a serious household pest. Its native hosts are the California condor
and great horned owl (U singer4 ). Although there are less than 40 de-
scribed species in the family, a more intensive study will no doubt result
in the discovery of additional species and changes in genera, as sug-
gested by the work of LisU
The common bedbug. The adult of the common bedbug, Cimex

lectularius Linn. (Fig. 38), measures from 4 to 5 mm in length and
:3 mm in breadth, it is obovate in form and much flattened . The adult is
reddish brown in color, whereas the young are yellowish white. Among
the local names applied to bedbugs are "chinches," "chintzes," "red
coats," "mahogany flats," "wall louse," "bedbugs," or simply "bugs."
Bedbugs, like lice, have been the constant companions of man for
centuries; the earliest writings on natural history (Pliny and Aristotle)
mention them. Bedbugs occasionally gain a foothold among laboratory
animals such as white rats and guinea pigs, upon which they feed readily.
They are nocturnal in their feeding habits, hiding in crevices during the
FlG. 38. The com-

mon bedbug, Cim ex lec-
tularius. Eggs shown at
right.
day. At night they are very active, crawling out of their hiding places, .
often traveling considerable distances to attack their victims. This is
especially true 'where iron bedsteads are used, as these do not provide
convenient hiding places for the bugs. Ordinarily where the old-fashioned
wooden bedsteads are used the bugs stay closer to their point of attack.
Mattresses in any case may afford harborage. Bedbugs are gregarious,
hence often great assemblages may be found in some convenient crevice
or beneath some nearby loose wallpaper, where the eggs are deposited
and the tarry black excrement collects.
The females deposit eggs in batches of from 10 to 50, totaling from
200 to 500, spread out in a yellowish patch. The eggs are large and yel-
lowish white in color. Oviposition occurs at intervals during a period of
from two to three months, apparently limited to the spring and summer
months, notvvithstanding' the fact that the insects are commonly favored
by warm rooms during the winter. The young hatch in from 7 to 30 clays
(usually about ten days), according to temperature, which also affects
their later growth. The time required for development from egg to
maturity is given as from 45 days to 11 months , although the time is
greatly influenced by temperature; there may be three or more genera-
tions a year under average condition. Ordinarily they require six to
THE BUGS '.)7
eight weeks to reach maturity. Bedbugs are evidently sensitive to high

temperatures; even a temperature of 100° F with fairly high humidity
will kill many of them. Activity ceases at about 60° F. At 60° F to
65') F bedbugs have lived for 136 days without food. Normally fed indi-
viduals may survive from 54 to 316 days under ordinary room-temperature
conditions. The presence or absence of food influences longevity; without
food adults may survive from 17 to 42 days (Marlatt"). Bedbugs molt
five times, and the minute wing pads characteristic of the adult insect
make their appearance with the last molt. Ordinarily but one meal is
taken between each molt and one before egg deposition; an average
period of eight days is required between moltings. Metamorphosis is
simple.
Methods of distribution. Bedbugs, lice, or other organisms cannot
originate spontaneously in filth as is believed by many; infestations are
traceable to introduced eggs, young, or adults. Thus the introduction of
one impregnated female might furnish the nucleus for a well-developed
colony in a few months. Hence the best-regulated household is not
exemp't from invasion, though cleanliness is the best preventive against
the multiplication of any household pest.
Public conveyances are commonly means for the dissemination of
bedbugs. Furthermore, migration from house to house by way of water
pipes, walls, and the like is not at all unlikely when infested houses are
vacated and the food supply is cut off. The insects are also easily carried
in clothing, traveling bags, suit cases, etc.
Bedbug bites. Persons "bitten" by bedbugs respond differently; in
some the bite produces marked swellings and considerable irritation,
while in others not the slightest inconvenience is caused. (This is true
also in the case of £lea bites and mosquito bites.) The bite, so-called, of
the bedbug is produced by piercing organs of the hemipteron type al-
ready described. It is probable that puncture by these stylets, unattended
by contamination or specific poisons, would produce little pain. The
welts and local inflammation are unquestionably caused by a specific
poison secreted by the salivary glands of the insect and introduced in
the act of feeding. The bedbug is able to engorge itself completely with
blood in from three to five minutes. Although persons are usually bitten
at night while in bed because of the normal nocturnal habits of the bed-
bugs, the insects will bite freely in subdued light by day.
The fact that bedbugs are obliged to feed at least five times either
upon the same or a different host in order to reach maturity, has placed
these insects under grave suspicion as potential vectors of disease.
Disease transmission. In consequence of statements made by various
authors that bedbugs are capable of transmitting plague and other
septicemic infections, Nuttall (1899, lac. cit., p. 4) carried on a series
of experiments with these insects. Mice were used as the experimental
animals because they are very susceptible to the infections in question.
He allowed the bugs to bite mice which had just died or were dying of
anthrax, plague, and mouse septicemia and then transferred them to
healthy mice. His experiments with anthrax are particularly instructive.
Mice inoculated with anthrax died in from 18 to 24 hours, after which
they were placed in glass-covered dishes and hungry bugs were intro-
duced. As soon as the bugs had sucked a little blood they were removed
to test tubes by means of a small camel's hair brush and transferred to
a shaved spot on healthy mice by inverting the tubes. Eight mice bitten
by 124 infected bugs all remained healthy. Variations of this experiment
gave similar results. It was found that the anthrax bacilli died in the
stomach of the insect in 48 to 96 hours at 13° to 17° C, and in 24 to 28
hours at 37°, and that the feces from the bugs contained living bacilli
during the first 24 hours after feeding. In view of these experiments it
may be concluded that infection through the bite of a bedbug either
does not occur or is exceptional. Infection might occur if recently in-
fected bugs were crushed while feeding and the punctured parts were
thereupon scratched. The feces of such bugs may also be infectious.
NuttalF showed that spirochetes survive in the bodies of these bugs for a
period of six days at a temperature of 12° C and for a much shorter
period (six hours) at 20° to 24° C. He did succeed in transmitting the in-
fection to a mouse, in one instance, by transferring 35 bugs from an in-
fected mouse to an un infected mouse. Patton in 19078 reported experi-
ments with Cimex hemipterus (Fabr.) [= C. rotundatus (Sign.)] in
which he was able to trace the development of the Leishman-Donovan
body of kala-azar through all its intermediate stages up to the formation
of the mature flagellates. Francis and Lake 9 reported transmission of
tularemia to mice by means of Cimex lectularius; however, Davis 10 points
out that during the hour-long interrupted feeding the tails of the mice
became soiled with bug feces and that in his tests bedbugs failed to
transmit P. tularense when a "clean" method of feeding was used, i.e.,
when fecal contamination of the host was eliminated.
The bedbug would appear to be relatively unimportant as a disease
vector, as indicated by the experiments cited above as well as many
others performed by various investigators. In spite of the fact that bed-
bugs can experimentally transmit the pathogenic agents of plague, re-
lapsing fever, leprosy, kala-azar, and tularemia (?), there is no convincing
evidence that the bedbug is a vector of these or any other diseases at
present known. It is of course possible, as shown in laboratory tests, that
occasional transmissions may be effected.
Bedbug control. With the advent of "DDT" the older methods of
bedbug control have been largely discarded; h~wever, the use of hydro-
THE BUGS 99
cyanic acid gas for house fumigation in some types of pest-control
operations will continue to be a sound practice in the hands of pro-
fessional operators. 11 Where live steam under pressure is available this
may be employed with good results in bunkhouses and the like.
DDT in bedbug control. A bedbug infestation in dwellings, barracks,
etc., can be successfully eradicated by the thorough application of a 5
per cent DDT spray applied to beds, mattresses, wall crevices, and other
infested situations. A single correct treatment will "bug proof" for six
months or longer. A 5 per cent DDT spray (xylene-Triton X-lOO) is
recommended; it is applied' as a spray until misty-wet to all parts of
mattresses (standing on edge), bedsprings, and bedsteads. In heavy
infestations the walls around the beds, particularly cracks along base-
boards, surfaces back of loose wall paper, and all other harborages must
also be sprayed. Chicken coops infested with the chicken bedbug can
also be freed of these pests by using DDT spray. Not only should food
be protected against the spray, but spraying must not be done near an
open flame.
Key to North American Cimicidaeo
1. Head longer than broad, clypeus narrowed anteriorly. Labrum much
longer than broad. Legs very long, the femora extending beyond sides of
body for a distance greater than width of pronotum. Organ of Berlese
absent. Primicimex cavernis Barber
Head broader than long, clypeus broadened anteriorly. Labrum short,
about as broad as long. Legs relatively short; the femora not much
longer than width of pronotum and only brieRy projecting beyond sides
of body. Organ of Berlese present. 2
2. Middle and hind coxae subcontiguous, the metasternum compressed.
Large bristles of body dentate only at their tips. Organ of Berlese
dorsal .............................. ... ..... ..... 3
Middle and hind coxae rather widely separated, the metasternum broad
and plate-like, widening posteriorly. Large bristles of body dentate
on convex side. Organ of Berlese ventral 6
3. Rostrum long, reaching hind margin of mesosternum. Posterior ab-
dominal tergites in female strongly sinuate, organ of Berlese located at
middle of fifth segment. New Mexico, Texas, Oklahoma, California,
Mexico. Hosts: California condor, Great horned owl, domestic chicken.
Haematosiphon inodora Duges.
Rostrum shorter, not reaching mesosternum. Posterior abdominal sutures
only feebly sinuate... ... .... ... ...... 4
4. Pronotum strongly narrowed posteriorly. Posteroventral portion of head
and adjoining prosternum strongly convex. California, Nebraska. Host:
White-throated swift. . . ...... Synxenoderus comosus List
Pronotum not strongly narrowed posteriorly. Head beneath and proster-
num not strongly convex... ................................... 5
o Prepared by Dr. Robert L. USinger.
5. Body clothed with long bristles, sides of abdomen with some bristles
twice as long as width of an eye. Colorado, Mexico. Hosts: Graybreasted
martin and other swallows . H esperocimex coloradensis List
Body with very short, inconspicuous bristles, those of sides of ab-
dominal segments much shorter than width of an eye. Eastern and
central United States. Host: Chimney swift. Cimexopsis nyctalis List
6. Pubescence long and sericeous. Last two antennal segments subequal in
length.. .Oeciactls vicarius Horvath
Pubescence short, stiff. Third antennal segment longer than fourth.
Genus Cimex Linn. 7
7. Sides of pronotum not widely dilated and not reflexed, fringed with
sparse, nearly straight hairs. Hemelytra with apical margins distinctly
rounded. Tropicopolitan. Host: Man. Cimex lzemipterus (Fabr.)
Sides of pronotum widely dilated, broader than width of an eye, and
densely fringed with backward-curved hairs. Apical margins of heme-
lytra nearly straight, rounded towards inner anngles 8
8. Length of contiguous portions of hemelytra shorter than scutellum.
Second antennal segment slightly shorter than third. Fringing hairs of
pronotal margin shorter than width of an eye. Widely distributed in
temperate regions. Host: Man .. Cimex lectularius Linn.
Length of contiguous portions of hemelytra longer than scutellum.
Second and third antennal segments equal in length. Fringing hairs
of pronotum longer than width of an eye 9
9. Marginal hairs relatively long. Tergum strigate. Eastern North America.
Host: Bats.. Cimex aditmucfus Barber
Marginal hairs relatively short. Tergum not transversely strigate. West-
ern North America. Host: Bats.. ..Cimex pilosellus Horvath
B. THE CONENOSES
Family Reduviidae
Family Reduviidae. The Reduviidae are typical examples of the heter-
opterous Hemiptera. They are commonly known as conenoses, kissing
bugs, and assassin bugs. There are over 2,500 species divided among 15
subfamilies, of which the Harpactocorinae is the largest, containing
more than a third of all the species, and next in size is the subfamily
Reduviinae. A very large percentage of the reduviids are predaceous and
feed on insects, many of which are harmful; hence the family is in the
main beneficial. A number of the species when handled carelessly defend
themselves by biting, and a very few have developed a definite habit
of sucking blood. The subfamily Triatominae comprises those members
of the family which feed exclusively on blood of vertebrates. This sub-
family is predominantly American.
. In addition to the characteristic wings already described, the head
of these insects is more or less elongated or cone-shaped, giving rise to
THE BUGS 101
the name conenose; the head has remarkably free movement; the eyes
are conspicuous; the ocelli, if present: are located behind the compound
eyes; the sturdy, three-jOinted proboscis can be thrust forward, but in
repose lies beneath the head; the piercing stylets can be extended far
beyond the tip of the proboscis; the long, slender, four- or five-segmented
antennae are situated in front of the eyes or on the border of the head;
the prothorax is strongly developed; most of the species are able to fly
well.
Life history. The rather large, usually smooth, more or less barrel-
shaped eggs of reduviids (often with stellate or fringed caps) are gen-
erally deposited in situations where the adults occur, i.e., the ground-
inhabiting forms deposit their eggs on the ground; arboreal forms lay
their eggs on leaves and stems; and house-inhabiting forms in dusty
FIG. 39 . Egg (left) and larva

(right) of a cone nose bug, Triatoma
firotracta. X 4.4
corners. The eggs of many of the species are illustrated by Readio,'2

who has also published an excellent account of the biology of the family. '"
The eggs are commonly deposited Singly, but sometimes in small
clusters, the total number per fema le varying considerably from a few
dozen to upward of 600. The incubation period varies from 8 to 10 days
to nearly a month, depending upon the species and temperature. The
newly hatched nymphs are wingless (Fig. 39). The usual number of
nymphal instars is five, although Readio states that Melanolestes picipes
(Herrich-Schaeffer) passed through only four. Some species overwinter
in the egg stage, others as adults, and still others as nymphs. In most
cases there appears to be but one generation a year. The length of the
life cycle of Triatoma rubrofasciata (DeGeer) was found by Neiva to
cover 210 days and . for Mestor megistus (Burm.) 260 days. Usinger14
reports a two-year life cycle for Triatoma longipes Barber.
The alimentary canal of Triatoma protracta (Uhl.). The alimentary
canal of Triatoma protracta (UhI.) according to Elson 15 is, as in other
Hemiptera, divided into three regions, the fore-, the mid-, and the hind-
intestine (Fig. 16). The fore-intestine comprises the pharyngeal duct,
the pharynx, and the esophagus, and measures 4.3 mm in length. The
pharyngeal duct consists of a delicate tube which connects the pharynx
with the suction canal of the maxillae. It is a very short duct and difficult
to distinguish from the pharynx. The pharynx may be considered as a
highly specialized portion of the alimentary canal. It is a boat-shaped
organ, which measures on an average about 2 mm in length and .3 mm
in diameter in the adult. The esophagus is a delicate duct, measuring
about 2 mm in length and .1 mm in diameter. Before reaching the pro-
ventriculus the esophagus expands and forms the proventricular vestibule.
The mid-intestine comprises the main portion of the alimentary tract and
consists of the proventriculus, the ventriculus or stomach proper, and an
elongated section. The stomach is a large sac, which when filled with
blood, occupies the greater portion of the body cavity and crowds the
other organs to the sides and behind. When empty it is wrinkled, but
when replete with blood it is smooth and pear-shaped. The mid-intes-
tine-3 is a much-coiled and narrowed portion of the mid-intestine; mid-
intestine-4 is a short dilated portion. The hind-intestine is the shortest
portion of the alimentary canal and comprises the rectum. This is a large
muscular sac, capable of considerable distension and usually containing
fecal material. It is pear-shaped with the broadest part 2.5 mm in diame-
ter and the length 3 mm. At the termination of the mid-intestine there
appears a circular whitish zone, the rectal gland. The ampullae, or en-
larged bases of the Malpighian tubules (four in number and of equal
size), form a rosette at this point. The Malpighian tubules are about
twice the length of the insect, and it is difficult to unravel them.
The salivary apparatus consists of two pairs of glands, the principal
and the accessory. The principal glands measure from 1 to 1.5 mm in
length and .8 mm in diameter and are situated in the mesothorax, one
on either side of the proventriculus. The spherical accessory glands,
about .5 mm in diameter, are located in the meta thorax directly posterior
to the principal glands, with which they are connected by a duct.
Conenose bites. Many of the species of conenoses inflict a painful bite
when handled carelessly. Most notorious perhaps of all is the "kissing
bug," Reduvius personatus (Linn.), a widely distributed species (belong-
ing to the subfamily Reduviinae) particularly active in the middle west-
ern and eastern United States. As early as 1899 Howard,16 quoting
Le Conte, writes as follows:
This species is remarkable for the intense pain caused by its bite. I do
not know whether it ever willingly plunges its rostrum into any person, but
when caught or unskillfully handled it always stings (pierces). In this case
the pain is almost equal to that of the bite of a snake, and the swelling and
THE BUGS 103
irritation which result from it will sometimes last for a week. In very weak
i\I1d irritable constitutions it may even prove fatal.
The wheel bug, Arilus cristatus (Linn.), belonging to the subfamily

Harpactorinae, also has a bad reputation as a biter. Reporting on the
bite of this species, Hall' 7 states:
The finger became reddened and felt hot to the touch. In the course of
a few days, growths resembling papillomas developed at the sites of the punc-
tures, the largest of these projecting as a small hornlike structure. Both of
these growths persisted for months, the largest slowly disappearing between
six and nine months after the infliction of the bite. The injured finger remained
warmer than the other fingers during this period, and, according to the
patient's statement, still feels warmer than the other fingers, a year later. The
development of pronounced cutaneous growths after a bite appears indicative
of the action of some toxin as a stimulant irritant.
A B
FIG. 40. Members of the family Reduviidae. (A) Triatoma protracta, (8)
Rasahus thoracicus.
The symptoms produced by the bites of Triatoma protracta (Uhler)

(Fig. 40A), a widely distributed Pacific coast species commonly known
as the China bedbug or cross bug and normally occuring in the nests
of wood rats, Neotoma spp., are described as follows by reporting phy-
sicians: "In a few minutes after a bite the patient develops nausea,
flushed face, palpitation of the heart, rapid breathing, rapid pulse, fol-
lowed by profuse urticaria all over the body. The symptoms vary with
individuals in their intensity." A blood examination made on a man
about four hours after he was bitten on the left arm showed a leucocyte
count of 11,000 (polymorphonuclears, 66 per cent; lymphocytes, 28 per
cent; monocytes, 3 per cent; eosinophils, 3 per cent). This man's symp-
toms a few minutes after the bite were large hive-like swellings in the
regions of the glands; intense itching and redness, especially in the hands;
pulse rate, normally 60, increased to 140.
Inquiries concerning these insects in California are most frequent
during May and June. They often attack sleeping individuals during the
night. Tests made in the laboratory indicate that some persons experience
no ill effects from the bite of this species (Wood18) .
The bloodsucking conenose or "Mexican bedbug," Triatoma san-
guisuga (Lec.), inflicts a very severe bite, which, because of the uniform
character of the resulting symptoms, supports the view that a specific
venom is injected with the bite. The bite is said to result in "a burning
pain, intense itching and much swelling . . . with red blotches and
welts all over the body and limbs." The effects of the bite may last for
months; however, they usually disappear within a few days.
The "two spotted corsairs," Rasahus biguttatus (Say) and Rasahus
thoracicus Stal (Fig. 40B), belonging to the subfamily Piratinae, the
former common in the southern United States, Cuba, and South America,
and giving way to the latter in the Northwest and California, are the
cause of many complaints. Howard (1899, loco cit.) cites the following
paper concerning "so-called spider bites," published by Dr. A. Davidson
in the Therapeutic Gazette of February 15, 1897, in which that author
arrives at the conclusion that almost all of the so-called "spider bites" met
with in southern California are produced by no spider at all but by
Rasahus thoracicus Stal. The symptoms which he describes are as follows:
Next day the injured part shows a local cellulitis with a dark central spot;
around this spot there frequently appears a bulbous vesicle about the size
of a ten-cent piece and filled with a dark grumous fluid; a smaller ulcer forms
underneath the vesicle, the necrotic area being generally limited to the central
part, while the surrounding tissues are more or less swollen and somewhat
painful. In a few days with rest and proper care the swelling subsides, and
in a week all traces of the cellulitis are, usually gone. In some of the cases
no vesicle forms at the point of injury, the formation probably depending-on
the constitutional vitality of the individual or the amount of poison introduced.
The use of warm compresses of magnesium sulphate is recommended

for the bites of conenoses.
Chagas' disease (American trypanosomiasis also known as Brazilian
trypanosomiasis) was first described in 1909 by Chagas (loc. cit., p. 6)
from Brazil. He named the causal organism Schizotrypanum cruzi (now
more generally known as Trypanosoma cruzi) , a spindle-shaped try-
panosome, with a single flagellum and a characteristic undulating mem-
THE BUGS 105
brane. The trypanosome occurs sparsely in human blood, hence diagnosis

of the infection by recovery of the organism from the blood is very dif-
ficult. The diagnostic method of Brumpt'9 to which he gave the name
"Ie xenodiagnostic" is now widely used; he described the method as fol-
lows: "J e donne Ie nom de xenodiagnostico (zenos, hote) au diagnostic
pouvant etre fait au moyen de l'hote agent vecteur habituel de la maladie
parasitaire, ou it l'aide d'un hote vicariant pouvant assurer la culture et
l'evoluation du parasite. C'est en somme une culture naturelle du parasite
chez des hOtes favorables." Essentially this refers to the use of appropri-
ate noninfected (clean) Triatoma bugs, which are allowed to feed upon
a person suspected of having the disease, and after incubation in the
body of the insect the trypanosomes, if present, may be recovered very
easily from the digestive tract of the bug either by dissection or by
microscopic sampling of feces taken from the rectum by means of a
slender pipette.
The most apparent symptom of Chagas' disease in most cases is the
unilateral swelling of the eyelid and face known as the sign of Romana.
The disease is most prevalent in children in whom it causes a high, long-
continuous fever, facial edema, adenitis, and anemia. Infection may last
many years. Symptoms of the chronic stage are commonly cardiac, be-
cause of the cardiotropism of the causal organisms; death is frequently
due to chronic myocarditis and is commonly sudden. The Leishmania
form of the trypanosome invades and destroys endothelial and other cells
notably cardiac and striated muscle. In a recent contribution, Romana 20
( 1947) re-emphasizes the nervous forms of Chagas' disease, based on a
study of cases of chronic encephalopathy; the patients presented "a
syndrome of psychic states and of spastic paralysis" which agrees with
the classical descriptions of Chagas and Villela. The disease occurs in
man throughout most of South and Central America from Argentina to
Mexico. In reservoir animals the infection occurs also in the southwestern
United States.
Transmission of the infection. Chagas (loc. cit., Chapter I) reported
successful transmission of the infection through the agency of Triatoma
( = C onorhinus) megistus (Burm.), but believed it was effected by
means of the bite of the insect. Brumpt21 (1912), using Rhodnius prolixus
Still, disproved the salivary theory of transmission by demonstrating that
the infectious stage of the life cycle of the trypanosome is completed in
the hind-gut of the insect and that the infection reaches the victim in the
feces of the bug, which almost invariably defecates on the skin of its
victim while in the act of sucking blood. From the soiled skin trypanos-
omes are readily transferred by fingers or otherwise to the highly recep-
tive conjunctiva of the eyes or the mucosa of the mouth or nose, and
infection takes place. Inoculation may also be effected by rubbing in
the organisms through the excoriated skin, e.g., by scratching. The

incubation period in man is said to be from 10 to 12 days.
Although bugs may receive their infection from man, it is no doubt
usually received from reservoir animals. On ingestion, the metacyclic
trypanosomes lose the flagella, change to oval "Leishmania forms," and
multiply, giving rise to "crithidia I forms" and the final infectious meta-
cyclic stage in the hind-gut of the insect. This life cycle requires from
6 to 15 days. The percentage of Triatoma found infected in nature is
startling; thus studies of this nature by various workers in widely sep-
arated areas cited by Usinger (lac. cit. ) showed that 43 per cent of
4,181 bugs were infected.
The following species of conenose bugs, among others, have been
found naturally infected: Rhodnius prolixus Stftl in l\1exico and Vene-
zuela; Panstrongylus geniculatus (Latr.) in Brazil and Panama; Triatoma
A B c D
FIG. 41. Examples of Reduviidae: (A) Triatoma protracta, (B) Triatoma

sanguisuga, (C) Panstrongylus geniculatus, and (D) Rhodnius pallescens.
dimidiata (Latr.) in Mexico, Panama,22 Guatemala, and San Salvador;

Triatoma rubrofasciata (DeGeer) in Brazil; Triatoma rubida (Uh1.),
T. hegneri Mazzotti, T. barberi Usinger, and T. phyllosoma (Burm. ), all
in Mexico; Triatoma sordida (Still) and T . delpontei Romana y Abalos,
in Argentina; Triatoma flavida (Neiva) in Cuba; Triatoma sanguisuga
(Le Conte), T. ambigua Neiva, T. gerstaeckeri (Stal), T. heidemanni
Neiva, and T. protracta woodi Usinger in Texas; T. longipes Barber, T.
protracta (UhI. ) and T. rubida uhleri Neiva, in Arizona; in Calif~nia ,
S. F. Wood 2 3 found a high natural infection in Triatoma protracta (Uh1.) ,
namely 25 per cent of 816 bugs examined (Fig. 41).
The infection occurs in natural reservoir animals such as armadillos,
opossums, house mice, rats ( N orwa y ), bats (several species), cats,
dogs, squirrels (Sciurus) , and wood rats.
THE BUGS 107
Kofoid and Donat (1933 24 ) reported on the occurrence of the infec-
tion in the wood rat, N eotoma fuscipes macrotis Thomas, in San Diego
County, California, with Triatoma protracta (Uhler) the vector. F. D.
Wood,z5 reporting more fully later (1934), states that the bloodsucking
bug, Triatoma protracta (Uhler), and the wood rat, N eotoma fuscipes
macrotis Thomas, are natural carriers of Trypanosoma cruzi in south-
ern California. She was able experimentally to infect the following ani-
mals with the trypanosome: albino rats, albino mice, rhesus monkeys, a
puppy, an opossum, the dusky-footed wood rat, and five species of white-
footed mice. S. F. Wood 26 (1943) reports naturally infected conenose
bugs from Arizona. Packchanian 27 reports successful experimental infec-
tion of a human being in Texas, using Triatoma heidemanni Neiva as
the vector.
It is of interest to note that Morishita (1935) 28 has conducted experi-
ments with Trypanosoma conorhini Donovan, which is regarded as non-
pathogenic; it resembles T. cruzi in its insectan phase, occurring com-
monly in the gut of Triatoma rubrofasciata (DeGeer) in Formosa. The
natural vertebrate host has not yet been discovered.
Several species of bedbugs are capable of transmitting the infection
experimentally, as well as many species of ticks, among these Amblyomma
ca;ennense (Fabr. ), Rhipicephalus sanguineus (Latr. ), Ornithodoros
moubata (Murray), and Ornithodoros savignyi (Audouin). Ornithodoros
turicattJ (Duges) has been proved to be an experimental vector of the
Brazilian strain by Wheeler.29
Control. The medical and social importance of Chagas' disease is
being widely recognized with the growing knowledge of the disease and
particularly with the advance in the knowledge of its vectors. There
appears to be no effective etiologic treatment, and there seems to be no
spontaneous cure. Measures to prevent the spread of the infection should
be aimed at the suppression of the insect vectors and limitation of reser-
voir animals. The widespread distribution and common occurrence of
both vectors and reservoirs add tremendously to the difficulty of the
problem. Educational measures that will reach the general public in all
infested areas are of paramount importance.
It is generally agreed that one of the major approaches to the prob-
lem is improvement in housing conditions in endemic areas, so as to
reduce and exclude reservoir animals (both wild and domestic) and to
prevent entrance and multiplication of the responsible insects.
Disinfestation of sleeping quarters, dwellings, and outhouses by the
use of one or more of the newer insecticides, such as benzene hexa-
chloride, chlordane, and special formulations of DDT has been recom-
mended. Romafia and Abalos 30 consider the first named as the preferred
insecticide, to be used on roofs, walls, furniture, henhouses, corrals, etc.

at the rate of 500 mg "Gammexane" per square meter, giving six months'
protection.
Key to Some Predaceous Reduviidae Likely to Be of

Medical Importance"
1. Hemelytra with a quadrangular or discoidal cell at the base of the
membrane. Subfamily Harpactocorinae .. 2
Hemelytra without a quadrangular or discoidal cell at the base of the
membrane .................... . 4
2. Sides of mesosternum without a tubercle or fold in front. Basal seg-
ment of beak about as long as front portion of head. Tribe
Zelini . Zelus socius Uhler
Zelus exsanguis (Stal)
Sides of mesosternum with a tubercle or fold in front of the hind angles
of the prosternum. First segment of the beak longer than front por-
tion of head. Tribe Harpactocorini 3
3. Front femora but little, if at all, thickened, unarmed, a little granulated.
Pronotum produced posteriorly over scutellum and with a high, median,
tuberculate ridge. Form robust. Arilus cristatus (Linn)
Front femora thickened, spinose, densely granulated. Pronotum not
produced or elevated as above. Form slender .. Sinea diadema (Fabr.)
4. Pronotum constructed behind the middle. Front coxae with outer sides
flat or concave. Subfamily Piratinae. 5
Pronotum constricted at or before the middle. Front coxae not Battened,
their outer sides convex. Subfamily Reduviinae .. 6,
5. Apical portion of anterior tibia angulately dilated beneath, the spongy
fossa being preceded by a small prominence. Hemelytra entirely
black . Melanolestes picipes (H. S.)
Tibia not dilated, the spongy fossa elongate. Corium and membrane
of hemelytra each marked with a yellow spot Basahus thoracicus (Stal
Rasahus biguttatus (Say)
6. Antennae inserted on top of head between margins close to eyes. An-
tenniferous tubercles not projecting from sides of head. Beak stout,
distinctly curved. Front of head turned downward. Tribe Reduviini
Reduvius personatus (Linn.)
Antennae inserted in lateral or dorsolateral margins of head. Anten-
niferous tubercles projecting slightly from sides of head. Beak slender
and relatively straight. Head strongly produced anteriorly. Tribe
Triatomini ...... ri
7. Head very long and slender with antennae inserted near the apex
......... Bhodnius prolixus Stal
Head moderately long with the antennae inserted at or behind ante-
ocular region .. .Triatoma sanguisuga (Lee.)
Triatoma protracta (UhI.)
Prepared by Dr. Robert L. USinger for the purposes of this book.
THE BUGS 109
C. SOME OTHER BUGS

Family Polyctenidae
This family includes the relatively little known bat bugs, which have
led a very uncertain systematic existence since 1874, when Westwood
founded the family Polyctenidae to receive these insects as aberrant
Anoplura. Later they were placed with the Hippoboscidae, or louse Hies
(Waterhouse, 1879). The number of known species is quite small (less
than a score); all are bloodsucking ectoparasites of bats. They have a
four-jointed rostrum (three-jointed in a single species from Africa),
tarsi three-jointed and antennae four-jointed, eyes wanting, hemelytra
short; the body commonly bears ctenidia (combs). They are viviparous,
the young being born at an advanced stage of development.
The two species known from the United States have recently been
described by Ferris and Usinger. One of these, Hesperoctenes hermsi,
was taken on the free-tailed bat, Tadarida macrotis (Gray), in the Chisos
~10untains, Texas, at an elevation of 6,200 feet, by Mr. A. E. Borell. The
other, H esperoctenes eumops, has been taken at several localities in
southern California on another free-tailed bat, Eumops perotis californicus
(Merriam).
Key to the Principal Families of H emiptera-H eteroptera of

North America which contain predaceous species(>
1. Antennae at least as long as the head; either free or, in the Phymatidae,
fitting in a groove beneath the lateral margins of the pronotum. Sub-
order Gymnocerata . ....................... . 6
Antennae shorter than the head and nearly or quite concealed in a
cavity beneath the eyes. Suborder Cryptocerata 2
2. Ocelli present. Shore-frequenting insects. (Toad bugs)
.. Family Gelastocoridae
Ocelli absent. Aquatic forms 3
3. Hind tarsi with indistinct, setiform claws (except in the minute, 3
mm or less, Pleine). Swim with ventral side upward (Back swim-
mers). . . Family Notonectidae
Hind tarsi with distinct claws 4
4. Membrane of the hemelytra without veins. Abdomen without caudal
appendages (Water creepers) . Family Naucoridae
Membrane of the hemelytra with distinct veins. Abdomen with caudal
appendages . ... . .......... ........ ............................. 5
5. Caudal appendages of abdomen long and slender. Tarsi one-segmented
(Water scorpions). .. Family Nepidae
Caudal appendages of abdomen short, flat, and retractile. Tarsi two-
segmented (Giant water bugs) Family Belostomatidae
• Prepared by Dr. Robert L. USinger for the purposes of this book.
6. Apex of last tarsal segment more or less split. Claws of at least the
front tarsi inserted before the apex. Superfamily Gerroidea . 7
Last segment of tarsi entire, with claws of all the legs inserted
at apex '" ........ ..... 8
7. Hind femora much surpassing apex of abdomen. Middle and hind
coxae approximate, distant from front ones (Water striders) ..
........................ . .. Family Gerridae
Hind femora scarcely surpassing tip of abdomen. Middle coxae (ex-
cept in Rhagovelia) equally distant from front and hind ones. (Broad-
shouldered water striders) ........ .................. Family Veliidae
8. Head very long, at least as long as the three thoracic segments com-
bined. Body linear. Legs and antennae very long and slender (Marsh-
treaders) .......................... Family Hydrometridae
Head shorter than pronotum and scutellum together.. 9
9. Antennae five-segmented. (Only members of the subfamily Asopinae
are predaceous. These have nonraptorial front legs, three-segmented
tarsi, and the first segment of the beak short, thick, and free) (Stink-
bugs) ............. ..... Family Pentatomidae
Antennae four-segmented (except in one genus of the Hebridae and
the genus Pagasa of the Nabidae, where they are five-segmented, and
in the Ectrichodine genus Rhiginia of the Reduviidae, where they are
eight-segmented. These exceptions do not have the combination of
characters listed under Asopinae).... 10
10. Beak three-segmented... .................. 11
Beak four-segmented.. ................ .............. 16
11. Front legs more or less raptoriaL Head cylindrical. Prosternum with
a median longitudinal stridulatory groove which receives the tip of
the beak. ............... .............. .... ................................ 12
Front legs not raptorial. Head rarely cylindrical. Prosternum without
a median longitudinal stridulatory groove between the front coxae... 13
12. Front legs with greatly thickened femora (Ambush bugs) ..
................................... ............... ............................... Family Phymatidae
Front femora somewhat thickened but much less than half as wide
as long (Assassin bugs). .. Family Reduviidae
13. Hemelytra with a cuneus (e.g. apical triangular region of corium
separated by a fracture) (Flower bugs) . Family Anthocoridae
Hemelytra without a cuneus.. .. ...... 14
14. Ocelli absent. Hemelytra reduced to small pads, without any trace of
a membrane. Body oval in form (Bedbugs, bat bugs, etc.)
............... ................................. . ............................ Family Cimicidae
Ocelli present..................................... .. .. .. .. .. .. .. .. .. .. 15
15. Membrane, when present, without veins; Clavus of same texture as
membrane. Body linear (Water treaders) .......... Family Mesoveliidae
Membrane with distinct, looped veins. Clavus and corium alike in
texture. Body suboval (Shore bugs) .......................... Family Saldidae
16. Hemelytra with a cuneus (Leaf bugs) ...................... Family Miridae
Hemelytra without a cuneus ..................................................................... 17
THE BUGS III
17. Body densely clothed with a very short, velvety pile. Membrane with-
out veins (Velvet water bugs). Family Hebridae
Body not clothed with a velvet pile ............ .... .... ........... 18
18. Hemelytra reduced to small pads. Ctenidia present at least on under
side of head and often on antennae, head, pronotum, and elsewhere
(Bat bugs) ...... ...... ..... .... ... ..................... Family Polyctenidae
Hemelytra usually well developed. Without ctenidia.. 19
19. Front legs raptorial. Beak with basal segment stout and very short; 20
Front legs not raptorial. Beak long and slender, the first segment
long, held between the bucculae at resL. . ...................... 21
20. Hemelytra entirely membranous (Gnat bugs) .. Family Enicocephalidae
Hemelytra with the basal portions thickened to form a corium distinct
from the apical membrane (Damsel bugs) ... Family Nabidae
21. Membrane with numerous veins which more or less run together
(Squash bug, etc.).. Family Goreidae
Membrane with only four or five simple veins usually arising from
the base (Chinch bug, etc.). Family Lygaeidae
BIBLIOGRAPHY
1. Usinger, Robert L., 1934. "Bloodsucking among phytophagous Hemi-
ptera," Ganad. Entomologist, 66:97-100.
2. Horvath, G., 1912. "Revision of the American Cimicidae," Ann. Musei
Nationalis Hungarici, 10:257-62.
3. Joyeux, Charles, 1913. "Biologie de Cimex boueti," Arch. de. parasit.,
16:140-46.
4. Usinger, Robert L. 1947. "Native hosts of the Mexican chicken bug,
Haematosiphon inodora (Duges)," Pan-PaCific Entomologist, 23:140.
5. List, George M., 1925. "Three new genera and three new species of
Cirnicidae from North America," Proc. Biol. Soc. Washington, 38:103-10.
6. Marlatt, G. L., 1946. "Bedbugs and their control," Pest Control and
Sanitation, 1:8-11.
7. Nuttall, C. H. F., 1913. "The Herter Lectures. I. Spirochaetosis."
Parasitology, 5:262-74.
8. Patton, W. 5., 1907. The development of the Leishman-Donovan
parasite in Cimex rotundatus. Covt. of India: "Scientific Mem." (by Officers
of the Med. and San. Depts.), n.s. no. 31. 25 pp.
9. Francis, Edward, and Lake, C. C., 1922. "Transmission of tularaemia
by the bedbug, Cimex lectulariu8," U. S. Public Health Service, Pub. Health
Rep., 37:83-95.
10. Davis, Gordon E., 1943. "Further attempts to trasmit Pasteurella
tularensis by the bedbug (Cimex lectularills)," ]. Parasitol., 29:395-96.
11. Wakeland, Claude, 1935. Fumigation for the Control of Household
Insects. Moscow: Univ. Idaho, College Agric., Extension Circular no. 50.
12. Readio, P. A., 1926. "Studies on the eggs of some Reduviidae (Heter-
optera)," Lawrence: Univ. Kansas, in Sc. Bull., vol. 16, pp. 157-79.
13. ---, 1927. "Studies on the biology of the Reduviidae of America
north of Mexic<?" Lawrence: Univ. Kansas, in Sc. Bull., vol. 17. 289 pp.
14. Usinger, Robert L., 1944. The Triatominae of North and Central
America and the West Indies and Their Public Health Significance. Washing-
ton. D. C.: Govt. Print. Office, in Public Health Bull., no. 288. iv + 83 pp.
15. Elson, J. A., 1933. A Morphological and Histological Study of the
Mechanism of Suction, Salivary Apparatus and Alimentary Canal of Hemiptera,
with Special Reference to Triatoma protracta (Uhler). Doctorate thesis, Univ.
Calif.
16. Howard, L. 0., 1899. The Insects to Which the Name "Kissing Bugs"
Became Applied During the Summer of 1899. Washington, D. C.: Dept.
Agric., in Div. Entomol. Bull., no. 22.
17. Hall, Maurice C., 1924. "Lesions due to the bite of the wheel-bug,
Arilus cristatus (Hemiptera; Reduviidae)," Arch. Int. Med., 33:513-15.
18. Wood, Sherwin W., 1942. "Reactions of man to the feeding of
reduviid bugs," J. Parasitol., 28:43-49.
19. Brumpt, E., 1914. "Le xenodiagnostic application au diagnostic de
quelques infectious parasitaires et en particulier a la Trypanosome de Chagas,"
Rull. Soc. path. exot., 7:706-10.
20. Romafia, C., 1947. "Miocarditis croruca equizo trepanosica," An. d.
Inst. med. regional, Univ. Nacional de Tucuman, Argentina, 2:1-39.
21. Brumpt, E., 1912. "La Trypanosoma cruzi evolue chez Conorhinus
megistus, Cimex lectularius, Cimex boueti et Ornithodoros moubata, cycle
evolutiv de ce parasite," Bull. Soc. path. exot., 5:360.
22. Rozeboom, L. E., 1936. "Triatoma dimidiata Latr., found naturally
infected with Trypanosoma cruzi Chagas in Panama," Am. J. Trop. Med.,
16:481-84.
23. Wood, S. F., 1942. "Observations on vectors of Chagas' disease in the
United States. I. California," Bull. Calif. Acad. Sc., 41 (pt. 2) :61-69.
24. Kofoid, Charles A., and Donat, Fae, 1933. "South American trypanoso-
miasis of the human type-occurrence in mammals in the United States,"
Calif. &. West. Med., 38:245-49.
25. Wood, Fae Donat, 1934. "Natural and experimental infection of
Triatoma protracta (Uhler) and mammals in California with American
human trypanosomiasis," Am. ]. Trop. Med., 14:497-517.
26. Wood, S. E., 1943. "Observations on vectors of Chagas' disease in the
United States. II. Arizona," Am. J. Trop. Med., 23:315-20.
27. Packchanian, A., 1943. "Infectivity of the Texas strain of Trypanosoma
cruzi to man," Am. J. Trop. Med., 23:309-14.
28. Morishita, Kaoru, 1935. "An experimental study on the life history
and biology of Trypanosoma conorhini (Donovan), occurring in the alimentary
tract of Triatoma rubrofasciata (de Geer) in Formosa," Japanese ]. Zool.,
G:459-546.
29. Wheeler, Charles M., 1938. "Experimental infection of Ornithodoros
turicata (Duges) with a Brazilian strain of Trypanosoma cruzi Chagas," Proc.
Soc. Exper. Bioi. & Med., 38:191-93.
30. Romafia, C., and Abalos, J. W., 1948. "Accion del "Gammexane"
sobre los Triatomideos. "Control" domiciliario," An. d. Inst. med. regional,
Universidad NacionaI de Tucuman, Argentina, 2:95-106. .
CHAPTER IX
._.
THE LICE
A. THE SUCKING LICE

Order Anoplura
General characteristics. The sucking lice comprise the order Anoplura.
Some believe them so closely related to the biting lice that both groups
are placed in the same order, viz., Anoplura, with the Mallophaga (biting
lice) reduced to a suborder and the sucking lice placed in the suborder
Siphunculata. There is little to support plaCing these parasites in a sub-
order (Parasita) of the Hemiptera. For the purposes of this work the
lice are classified as members of two orders, Anoplura (sucking) and
Mallophaga (biting), resembling each other in many respects, but differ-
ing radically in their feeding habits, the former being bloodsuckers while
the latter feed on scales, secretions, and detritus of the skin. Wings are
absent in both groups, and metamorphosis is simple. The body is flat-
tened; the legs are in part adapted for clinging to hairs and feathers.
The Anoplura have a protrusible proboscis at the tip of the head; the
Mallophaga have a pair of tiny mandibles situated on the under side of
the head. The biting lice are, as a rule, much more active than the suck-
ing lice.
The Anoplura parasitize mammals, while the Mallophaga are parasites
of both mammals and birds and in all cases are normally host specific,
permanent ectoparasites, very rarely except accidentally or experimen-
tally transferable to a different species. The United States Bureau of Ento-
mology is successfully rearing the human body louse on selected rabbits.
The entire life cycle, egg to egg, is normally spent on one host.
Classification of Anoplura. Students technically concerned with the
sucking lice will need to consult the classical works of Ferris, particularly
his monograph appearing in eight parts, 1919 to 1935, in Publications of
Stanford University Press. 1 The order comprises some 30 genera and
500 species (Essig); these are arranged according to various authors in
four families: ( 1) Echinophthiriidae, with body thickly covered with
short stout spines and scales, antennae four- or five-jointed, spiracles
small; occurring exclusively on marine mammals, e.g., Antarctophthirus
113
trichechi (Bah. ) on the Pacific walrus, and Echinophthirus phocae
(Lucas) on the seal; (2) Haematopinidae, body spines or hairs in rows,
never with scales, tibiae with thumb-like process opposing the claw,
eyes lacking, antennae five-segmented, e.g., Haematopinus suis (Linn.)
on swine, and Linognathus vituli (Linn.) on cattle; (3) Haematopi-
noididae, lice of gophers in North America; (4) Pediculidae, the most im-
portant family of all from the public health standpoint, since it includes
body lice, head lice, and pubic lice of man and other primates. Haema-
tomyzidae, heretofore included as a family of the Anoplura, has been
placed by Ferris 2 in the Mallophagan suborder, Rhynchophthirina; it
includes but the single species Haematomyzus elephantis Pia get of the
elephant.
The lice of man and other primates. The family Pediculidae the only
I)
eye-possessing lice, includes the three genera, Pedicinus, Pediculus, and

Phthirus. The genus Pedicinus belongs exclusively, according to Ferris,
to the Cynomorpha monkeys, and includes eight known species,
PediCinus eurygaster (Burm.) on Macacus; Pedicinus longiceps Pia get
on Pithecus; Pedicinus albidus (Rudow) on Macaca (Barbary ape);
Pedicinus hamadryas Mjoberg on Hamadryas; Pedicinus patas (Fahren-
holtz) on Cercopithecus patas; Pedicinus ancoratus Ferris on Presby tis;
Pedicinus pictus Ferris on Colobus caudatus; Pedicinus obtusus (Rudow)
on Semnopithicus maurus.
The genus Pediculus, according to Ferris, includes only three species:
(1) Pediculus humanus Linn., the head louse and body louse of man,
commonly referred to in medical literature as two distinct species,
namely, Pediculus capitus, head louse, and Pediculus vestimenti, body
louse; (2) Pediculus mjobergi Ferris of Ateles apes; and (3) Pediculus
schaffi Fahrenholz, of the chimpanzee.
The genus Phthirus (also spelled Phthirius) includes the so-called
crab lice, Phthirus pubis (Linn.) of man; and Phthirus gorillae Ewing,
of the gorilla.
SUCKING LICE OF MEN
The human head louse, Pediculus humanus capitis DeGeer (Fig. 42),
is gray in color, but is said to vary according to the color of the hair and
color of the host' (Murray 3). The male averages 2 mm in length and
the female 3 mm. This species occurs on the head, about the ears and
occiput, but from reliable observations made by a number of observers it
may in heavy infestations establish itself on other hairy parts of the body.
In severe infestations the hair may become literally matted with eggs
.. For extensive list of references (961) see "Bibliography on Lice and Man,
with particular reference to wartime conditions," by Mary E. Grinnell and Ina
L. Hawes, U. S. Dept. Agric. Washington, D. C. Bibliographical Bull. no. 1, July,
1943.
THE LICE 115
(nits), parasites, and exudate from the pustules which originate from the
louse bite. The term plica palonica is applied (Stiles) to the fetid mass .
forming a sort of carapace (trichoma), in which fungus may develop ,
and beneath which myriads of lice may be found .
. The number of eggs deposited by the female ranges from 50 to 150.
These are glued to the hair and hatch in from 5 to 10 days, an average
of 7 days. Development is very rapid (incomplete metamorphosis with
three molts), three weeks usually covering the entire cycle from egg to
egg. Lice are easily disseminated by physical contact and otherwise,
hence slight infestations may occur under the best of sanitary conditions,
particularly among school children. However, the continued presence
of lice on head or body is inexcusable since eradication is simple (see
r, A B
FIG. 42. Life cycle of the head

louse, Pediculus humanus cap-itis.
(A) egg attached to hair, (B)
nymph, (C) male, and (D) female. c D
lousicides, p. 124) . The mere use of soap and water in washing the head
and hair is ineffective in destroying lice present in the hair.
The pubic louse, Phthirus pubis (Linn.) (= Phthirus inguina lis Redi),
also called crab louse (Fig. 43), is easily recognized by its crab-like ap-
pearance. It measures from 1.5 to 2 mm in length, is nearly as broad as
long, and is grayish white in color. It infests the pubic regions particu-
larly but also the armpits and more rarely other parts of the body, such
as the mustache, beard, eyelashes, and eyebrows. The writer has seen
soldiers infested with this species of louse from the ankles to the eye-
brows. These lice ,are remarkably stationary in their habits, often re-
maining attached for days at one point with mouth parts inserted into
the skin. The pruritus caused by the bites of these parasites is very in-
tense, and a discoloration of the skin usually results if infestation con-
tinues over a longer period of time. The term phthiriasis may be em-
llG MEDICAL ENTO~IOLOGY
played to designate infestations of pubic lice, although the term pubic

pediculosis is also used.
The female louse deposits its eggs on the coarser hairs of the body
where the parasites occur. The number of eggs deposited per female is ap-
parently quite small, although Nuttall' states that he would not be sur-
prised to learn that 50 or more eggs may be layed. The incubation period
seems to be from six to eight days. After three molts the adult stage was
reached in Nuttall's experiments in from 15 to 17 days, and the egg-to-
egg period was 22 to 27 days. Pubic louse infestations are very common
among adults. Spread is usually by physical contact with infested in-
dividuals or use of infested toilet seats, blankets, etc.
FIG. 43. The pubic louse, Phthirus

pubis . Egg attached to hair.
The body louse, Pediculus humanus corporis DeGeer (= Pediculus

vestimenti Nitzch) (Fig. 44), is the common clothing louse which dur-
ing World War I became known as the "cootie," also called the "gray-
back." During World War II it became known as "mechanized dandruff"
and "seam squirrel." Parasitologists now quite generally agree that there
is no specific difference between it and the head louse; both are regarded
as racial forms of Pediculus humanus Linn. Evidence concerning this
matter is presented by Nuttall,5 whose admirable treatises on lice and
their relation to disease and control should be read by all students in-
terested in the subject. Patrick A. Buxton's The Louse" is an important
contribution.
Body lice infest the clothing where it comes in close contact with
the body rather continuously, e.g., the fork of the trousers, the armpits,
the waist line, neck, and shoulders. In his inspection of troops at de-
lousing stations during World War I, the author usually found lice on
the underclothing, but the nits were generally found in the seams of the
breeches at the crotch if present at all. After all clothing was removed
lice were occasionally found on the body. Eggs are undoubtedly b~f
preference deposited on fibers in the seams of clothing as already men-
THE LICE 117
tioned. Nuttall and others have satisfactorily proved that the body louse
may at times attach its eggs to the coarser hairs of the body.
Nuttall states that a female body louse may lay from 275 to 300 eggs,
the average number laid per day being about 10 for 20 to 30 days. The
shortest incubation period varies from 5 to 7 days when eggs are kept
near the body at between 35 ° and 37° C. HatQhing, according to Leeson 1
(1941), does not occur when the temperatUte reaches 23° C or below,
nor at 38 ° C or higher. At 24° C the incubation period is 17 to 21 days;
at 29° C it is from 9 to 11 days; at 35° C it is 5 to 7 days; at 38° C the
FIG . 44. Human body louse, Pe(li-

culus lwmanu.s corporis. X 15.
eggs will not hatch. The effective zone for the egg stage is apparently
from about 23° to 38° C.
The young lice begin sucking blood at once on hatching from the
eggs and throughout their development feed frequently both day and
night, particularly when the host is quiet. :Mahlrity is reached in 16 to
18 days from the time the eggs are deposi~ed. There are three molts.
Females begin laying eggs a day or two after reaching maturity. The
egg-to-egg cycle averages about three weeks. Unfed lice soon die; prob-
ably 10 days would cover the longest period of survival without food.
However, if fed, lice may live from 30 to 40 days. Moist fecal matter in
masses or spiral threads is extruded as the louse feeds; the feces drys
quickly in the air. Death may lurk in the eX'::rement of a louse.
The following summary of temperature influences on lice given in a
lecture on the "Life History of the Human tome," by Pierce,S June 17,
ll8 MEDICAL ENTOMOLOGY
1918, while varying somewhat from Leeson's (loc. cit.) results, is worth
repeating:
In the absence of definite humidity data we may roughly describe the
zones of climatic influence on the lice as follows. The zone of minimum fatal
temperatures for eggs in below 20° C. (68° F.) and for adults lies below zero
centigrade (32° F.). The zone of dormancy in adults extends from about _10°
to 5° C. (14° to 41° F.). The zone of sluggish movement without reproduc-
tive activity and with practically no digestive processes extends from 5° to
20° C. (41° to 68° F.). Digestion ceases at 12° C. The zone of optimum
activity lies between 20° to 40° C. (68° to 104° F.) with the optimum about
30° C. Practically all egg hatching occurs within this zone, as does all oviposi-
tion, practically all assimilation of food, and all normal activity. From 40° to
44 ° C. the lice are wildly active; this zone represents one of exhaustion in
which death of eggs occurs. Above 44° C. (1l2° F.) lies the zone of maximum
fatal temperatures.
Dissemination of body lice. Lice live normally on the surface of the.
the body or in clothing being worn. Lice do not voluntarily leave unless
the body grows cold in death or becomes hot with high fever. Even then
they cannot travel far but are easily dislodged and may fall to the
ground; they will quickly invade a new host if there is one close enough.
Proof that lice are uncommon in blankets used by heavily infested troops
was obtained by Peacock9 who reports 0.8 per blanket and 31.4 per man
including clothing. Louse infestation is mainly the result of contact with
lousy persons or their infested clothing. As many as 10,428 lice and
10,253 nits have been reported on one shirt; in heavily infested popula-
tions one may readily collect 400 to 500 lice from one person.
Pediculosis. The presence of lice on any part of the body may be re-
ferred to as pediculosis. That louse bites may produce certain systemic·
disturbances seems to be indicated in a report made by Moore: 10
I started feeding about 700 to 800 twice a day. Almost immediately a
general tired feeling was noticed in the calf of the legs and along the shin
hones, while on the sales of the feet and underneath the toes this tired feel-
ing was so intense as often to prevent sleep until late in the night. An irritable
and pessimistic state of mind developed. An illness resulted with symptoms
very similar to grip and a rash similar to German measles was present, par-
ticularly over the shoulders ard abdomen.
The skin of persons who continuously harbor lice becomes hardened
and deeply pigmented, a condition designated as vagabond's disease or
morbus errorum. To relieve itching due to pediculosis a lotion made of
1 per cent spirits of thymol may be used, or a mixture of 5 per cent ben-
zocaine, 2 per cent methyl salicylate, and 0.5 per cent salicylic acid made
up with 70 per cent alcohol.
Favus and impetigo. Experiments conducted by Dewevre ( 1892,
THE LICE 119
loco cit., p. 5) and others prove that lice may carry the causal organ-
isms of both favus (Achorion schoenleini Lebert) and tropical impetigo
(Diplococcus pemphigi contagiosi Wherry). Nuttall (1899, loco cit., p.
4) states the Dewevre "removed ten pediculi from a child suffering
from impetigo and placed them on a healthy infant, which a few days
later developed impetigo. The experiment Was repeated several times
with the same results. In a second series of experiments, he took scrap-
ings from under the nails of children that had impetigo and placing
them on artificially scratched places, reproduced the disease. Lastly he
took pediculi from a child that was not affected with impetigo and
placed them on a child that had the disease; removing them after twenty
minutes, he replaced them on a healthy child. The latter acquired the
disease, as did 50 per cent of the children so experimented with. He
claims the specific microorganism adheres to the front legs especially.
also to the hairs of the insect, and the latter carries them as bees do
pollen. In the last set of experiments, he only allowed the pediculi to
remain half an hour on the healthy head, but this was sufficient to pro-
duce infection."
Epidemic relapsing fever has occurred in many parts of the world and
may be regarded as at least having been cosmopolitan in distribution.
Widespread epidemics no longer occur. Great epidemics did occur, how-
ever, during World War I, notably in Rumania and later (1921) in
Africa. There were frequent epidemics of this disease in Europe during
the 18th and 19th centuries, and it was in the 1868 epidemic in Berlin
that Obermeier observed "myriads of living and actively motile spirilla
in the blood of relapsing fever patients during the febrile attack." In
1873 during another epidemic he applied Ehrenberg's nomenclature and
called these organisms "Spirochaeta." To honor the discoverer of the or-
ganism in the blood of relapsing fever patients Cohn gave the name
Spirochaeta obermeieri. The designation Spirochaeta recurrentis (Lebert)
has gained wide acceptance; however, because of priority one should
now use Borrelia recurrentis (Lebert).
Although lice had long previously been under suspicion it was not
until 1907 that MackieH in India secured evidence that relapsing fever
is a louse-borne disease. He noted that the spirochetes multiplied within
the gut of the lice and that they could be found in the ovary, testis, and
the Malpighian tubules of the insects. Mackie believed that infection
might result from the insect's regurgitating the contents of its gut into
the wound in the act of feeding. Later (1912) Nicolle, Blaizot, and
Conseil12 failed to transmit the spirochetes through the bites of infected
lice and found that the only reliable successful experiments involved the
injection or subcutaneous inoculation of an extract of infected lice. On
the basis of experiments in which men and monkeys were exposed to
hundreds of bites, they came to the conclusion that infection results from
spirochetes which collect under the fingernails and on the finger tips of
an individual after he has crushed lice on his skin, and which, in scratch-
ing, he inoculates into the excoriated skin.
Nicolle and his colleagues also found that the spirochetes disappear
from and later reappear in the louse. Only a few remain in the insecfs
intestine up to 5 or 6 hours after infection, and none after 24 hours; but
they reappear in the insect in from 8 to 12 days and are then present
in the general body cavity, none being found in the alimentary canal.
The authors believed that the spirochetes were transmitted to the off-
spring of infected lice. Chung and Feng 13 (1936) state that transovarian
transmission of Borrelia recurrentis does not occur in lice. They also state
that the salivary glands and Malpighian tubules of infected lice do not
contain the spirochetes, and that the feces of infected lice are not in-
fectious. The gastric juice of lice is detrimental to the Borrelia; only
about 1 to 5 per cent or less of the ingested spirochetes gain access to
the tissue and coelomic cavity where multiplication takes place, multi-
plication being by transverse division.
Man is the reservoir of the louse-borne infection (man-to-Iouse-to-
man). There must be a human carrier in a lousy population. In tick-borne
or endemic relapsing fever, endemicity is maintained in reservoir animals
(rodents, etc.) Nuttall has reported that a single infective louse crushed
upon the excoriated skin has produced relapsing fever. The incubation·
period in the human is from 6 to 10 days. The onset of the disease is
sudden, with headache, chills and fever, and generalized pains. The
fever remains high for four to six days and subsides abruptly, with an
afebrile period of four to eight days followed by a relapse. There may be
several relapses. Mortality is usually low but may vary from 2 to 50 per
cent, or even higher in crowded poverty-stricken and louse-infested
populations. Microscopic (darkfield) examination of blood smears taken
during febrile periods will readily reveal Borrelia. Mouse inoculation with
the patient's blood will produce the organisms in the blood of the mouse
in from 24 to 48 hours. (For tick-borne or endemic relapsing fever consult
Chapter Xx. )
Typhus fever. Classical or old world typhus fever, known also as
tabardillo (Mexico), Brill's disease (United States), jail fever, or war
fever, is a disease of ancient origin and wide distribution, chiefly in
Europe, north Africa, Asia, and higher altitudes of Mexico and Central
and South America. The causal organism is Rickettsia prowazeki Da
Rocha-Lima. Wherever human beings are concentrated in close quarters,
especially in times of war and famine, this disease may become rampant.
It is chiefly a disease of winter and spring, with mortality from 15 to 75
per cent. The disease is characterized by a high fever continuing about
THE LICE 121
two weeks, backache, intense headache, bronchial disturbances, mental
confusion, stupor, a congested face (designated also as a "besotted
expression"), and on the fifth or sixth day by a brick-red macular erup-
tion on chest and abdomen, which later spreads to other parts of the
body, even to hands, feet, and face. This mottling led to the belief that
tabardillo of Mexico was identical with spotted fever of Montana, a
fact that was proved to be erroneous by Ricketts and Wilder.14 Ricketts
contracted typhus fever during the course of this investigation and died
of the disease. Laboratory tests for typhus consist of complement fixation
with typhus antigen, becoming positive in the second week; and Weil-
Felix reaction (agglutination of B. proteus strain OX-19).
Transmission by lice. That the louse (Pediculus humanus Linn.) is
probably the sole agent in the transmission of typhus fever from man to
man was shown by Nicolle, et al./ 5 (1909, working in Tunis) and Ricketts
and Wilder 16 (1910, working in Mexico). The latter found that Pithecus
(Macacus) rhesus (Desmarest) can be infected with tabardillo (Mexican
typhus) invariably by the injection of virulent blood from man taken
on the eighth to tenth day of fever, that the monkey may pass through
an attack of typhus so mild that it cannot be recognized clinically, and
that immunity results. Typhus was transmitted to the monkey by the bite
of the louse in two experiments, the lice in one instance deriving their
infection from man and in another from the monkey. Another monkey
was infected through the introduction of the feces and abdominal con-
tents of infected lice into small incisions. There is a tendency to disregard
the head louse (P. humanus capitis) as a vector of typhus; however, it
must be remembered that Goldberger and Anderson 17 (1912) did succeed
in transmitting typhus to a monkey by cutaneous injection of a saline
suspension of crushed head lice.
Nuttall 18 points out that if lice (Pediculus humanus corporis) are
crushed 9 or 10 days after an infective feed, or if their feces are collected
three to six days after they have fed on infective blood, their contents
and feces respectively are capable of producing infection if placed upon
excoriated skin. It is now believed that the usual channel of infection is
through fecal contamination. The Rickettsiae may remain alive and
virulent in louse feces kept dry at room temperature for more than 60
days, thus infection may be acquired through the respiratory passages
by inhalation of minute particles of louse excrement. It is generally be-
lieved that the normal channel is through scratches or abrasions and
through conjunctival contact by fingers contaminated with louse feces.
The typhus fever patient is infectious for the louse, generally speak-
ing, from early in the disease to about the tenth day, though later infec-
tions are possible. The Rickettsiae multiply enormously in the mid-gut
of the louse; the epithelial cells become so distended after a few days
that they rupture and enormous numbers of Rickettsiae appear in the
insect's feces. It is believed that infected lice may die in a week or 10
days because of the enormous multiplication of these organisms. It is
believed that few lice become infected while feeding on a typhus fever
patient, probably less than half. If a louse survives the infection it remains
infective for the rest of its life.
Murine typhus fever (flea-borne) is maintained in nature in rats,
and transmission to man is effected through the agency of rat fleas. The
causal organism is Rickettsia mooseri Montiero (sometimes designated
as Rickettsia prowazeki mooseri). This disease is clinically similar to
louse-borne typhus except that its course is milder and the rash is less
profuse. The mortality is low, perhaps about 3 per cent. It is widely
distributed in many parts of the world, particularly in warmer climates,
in contrast to epidemic typhus which occurs in cooler climates favoring
lousiness. Both may occur in the same areas. The use of the terms "en-
demic" and "epidemic" to distinguish the "flea-borne" from the '10use-
borne" strain of typhus is open to criticism, since both must have an
endemicity to survive and both may be epidemic at times. Furthermore
it is now generally believed that "flea-borne" murine typhus may become
"louse-borne" and epidemic, with high mortality. (The relation of fleas
to murine typhus is discussed in Chapter XIX.)
Trench fever, also known as five-day fever, Volhynia fever, shank
fever, His-Wernerische Krankheit, and Febris quintana, is a nonfatal
disease characterized by sudden onset of fever, headache, dizziness,
pains in the muscles and bones, particularly in the legs, with especial
tenderness of the shins, and lasting 24 to 48 hours or longer, followed at
intervals of about five days by other attacks of fever of less and less
severity. While this disease was of considerable importance during World
War I, it did not make its appearance during World War II.
The causal organism of trench fever is believed to be Rickettsia
quintana, so named by Da Rocha-Lima 19 in 1916, confirming the earlier
work of Toepfer. 20 This organism is found in the stomach lumen of the
louse, while Rickettsia prowazeki of typhus invades the epithelial cells.
Two commissions, one British consisting of Byam, Carroll, et al., and
the other American, consisting of Strong, Swift, et al., carried out very
thorough investigations, during World War I; the summary, in part, of
the findings of the British commission follows:
1. The whole blood from febrile trench fever cases, up to the 51st day
of disease, when injected intravenously, is capable of reproducing the dis-
ease. The incubation period in such infectious varies greatly-from 5 to 20 .
days.
2. The virus as contained in the circulating blood is destroyed by the
addition of distilled water in large quantities.
THE LICE 123
3. The bites alone of infective lice do not produce trench fever.

4. The excreta of infective lice when applied to a broken surface of
skin do readily reproduce trench fever. The incubation period of such infec-
tions is remarkably constant and averages 8 days.
5. The excreta passed by lice fed on trench lever patients are not in-
fective till the expiration of not less than 5 days £rum the commencement of
the feeding on trench fever blood, thus indicating a developmental cycle in
the louse or a period during which the organism multiplies.
6. Once lice are infective they remain so till at least the 23rd day from
date of their infection.
7. The virus of trench fever, as contained in infected louse excreta, is
capable of withstanding drying at room temperature, exposure to sunlight,
keeping for not less than 16 days, and heating to 5{l° C. for 20 minutes.
8. 80° C. for 10 minutes destroyed the virus, which is therefore not a
spore-bearing organism.
9. The bodies of infected lice when crushed llpon the broken skin are
capable of producing trench fever. When lice becorfle so infective remains to
~~ <i~\.~"'ffi\.\."",~<i.
9a. Active trench fever blood equivalent to the content of 11 lice does
not produce trench fever when rubbed into the broken skin.
10. Infection probably does not take place by the mouth or by inhala-
tion.
11. The excreta of lice are not normally capable of producing trench fever.
[Note: Some reports indicate that feces may be highly infectious for as long
as 4 months.]
12. Trench fever infected lice do not transmit the disease to their off-
spring.
13. There is a possibility of some attacks of t(ench fever being afebrile
throughout.
14. The percentage of individuals naturally iIflmune to trench fever is
exceedingly small.
15. Old age is no bar to infection.
16. Such immunity as results from an attack of trench fever is not perma-
nent, and may only persist for so long as the individual shows evidence of
the disease.
17. Even as late as the 79th day of disease a patient's blood may remain
infective and be capable of infecting lice fed on such a patient while febrile.
Delousing methods practiced over a long period of years prior to and
including the early part of World War II were cumbersome and usually
expensive. The results were as a rule not wholly satisfactory in that the
recently deloused individual was immediately subject to reinfestation if
exposed to lousy persons. Among the methoos available during the
period of World War I were the following:,
Fumigation of clothing, persons to be properly bllthed while this was being
done; required specially constructed sealed chambers, stationary or portable;
fumigants, even when vacuum was used did not as a rule kill louse eggs;
among gases used were hydrocyanic acid gas, carbon tetrachloride," and
chlorpicrin.
0
Laundry methods, washing of rough cotton goods" at 180 F. for 30
minutes destroyed lice and their eggs; woolen goods" required a different
treatment to avoid shrinkage.
Dry heat, temperature of 60° C. (140° F.) for 20 minutes killed both
lice and eggs; required special types of heat chambers in order to circulate
heat to insure equal distribution among the loosely hung garments.
Ironing with a hot iron.
Steam sterilization, insured disinfestation as well as disinfection; disad-
vantageous in that shrinkage and wrinkling resulted; required special expen-
sive equipment such as steam sterilizers, delousing stations, etc; vacuum of
from 12 to 15 inches for about 5 minutes, followed by 15 to 20 pounds steam
pressure for 15 minutes, then allowed steam to escape and create 15 inches of
vacuum, broke vacuum and unloaded contents when zero was reached. Serbian
barrels were also used to generate steam.
Storage of garments, to kill lice by starvation; it was assumed that dry
storage for about three weeks would prove effective.
Louse powder, N.C.!., consisting of commercial naphthalene 96 per cent,
creosote 2 per cent, and iodoform 2 per cent, dusted in underclothing once
a week.
FollOWing World War I and up to 1941 little attention was paid to

the development of new methods of louse control. Control methods fell
back into the old groove of temporary relief, in spite of the fact that
insect toxicology had advanced enormously as applied to agricultural
pests. There was added, however, a new fumigant for delousing, namely
methyl bromide, a very dangerous gas which destroys all stages of the
louse. It was employed in two ways for delousing clothing, (1) fumiga-
tion of quantities of clothing in large gas-tight vaults; 30 minutes' ex-
posure in a 330 cubic-foot vault with three pounds of the gas; lower
temperatures required either more of the gas or a longer exposure; (2)
fumigation of single outfits of clothing in individual gas-tight bags-
each bag, 56 by 30 inches, was equipped inside in a pocket with an
ampule containing about 20 cc of methyl bromide; after clothing was
placed inside the bag and the bag closed, the ampule was crushed,
liberating the gas; after 45 minutes the bag was opened and the clothing
shaken out and aired. The use of this gas (methyl bromide) and practi-
cally all other aforementioned methods of delousing finally gave way
to lousicidal powders in wartime practice.
Lousicides. Great credit is due to the group of workers stationed at
the Orlando, Florida, laboratory of the United States Bureau of Ento-
mology and Plant Quarantine who developed the "MYL" insecticide
powder, which according to Miscellaneous Publication No. 606, United
THE LICE 125
States Department of Agriculture, August 1946, is composed of the fol-
lowing ingredients: 0.2 per cent of pyrethrins, 2 per cent of N-isobuty-
lundecylenamine, 2 per cent of 2,4-dinitroanisole, and 0.25 per cent of
Phenol S (isopropyl cresols, by-product of thymole manufacture) in
pyrophyllite (all per cents by weight). It is effective against all species
of human lice and not only repels them but kills them and their eggs.
The powder is dusted lightly into the seams of the underclothing or on
the infested parts of the body at weekly intervals.
Then came the Swiss louse powder, Neocid, consisting of 5 per cent
mixture of DDT (dichloro-diphenyl-trichloroethane) in talc. With the
successful field tests made with the three lousicide powders-MYL, DDT,
and the British AL-63 (containing rotenone) -there was opened a new
era in the control of louse-borne diseases. The Germans, according to
Bushland24 ( 1948 ), developed chloromethyl p-chlorophenyl sulfone
("Lauseto neu"), which was highly toxic to lice and an excellent ovicide,
but not nearly so persistent in residual effect as DDT. The arresting of
small typhus epidemics in native villages of Egypt, Algeria, and Mexico
proved that the louse powders are practical. The enormity of this dis-
covery can only be measured in the light of the great loss of life due to
typhus during 1917-1921 (World War I) when, according to the Statis-
tical Bulletin of the Metropolitan Life Insurance Company (November,
1941), Russia alone is said to have lost 2,500,000 to 3,000,000 of her
people; millions more died as the epidemic swept through the Balkans,
Poland, and Germany.
Typhus fever was again threatening in the early years of World War
II. During 1942 there were some 3,000 cases in Egypt and about 80,000
cases in the rest of North Africa. When Allied forces landed in Italy
in September, 1943, a typhus epidemic was threatening Naples, a city with
a population of nearly a million persons, where there was congestion,
insanitary living conditions; food scarcity, undernourishment, and con-
fusion. The United States of America Typhus Commission was charged
with responsibility for control of the disease on January 3, 1944
(Wheeler25 ); however, actual dusting operations were instituted on De-
cember 15, 1943, by a dusting team previously trained by a member of
the Health Commission of the Rockefeller Foundation. Wheeler, a mem-
ber of the United Stat~s Typhus Commission describes the measures
used in the epoch-making campaign to control typhus in Italy. Although
both MYL and DDT powders were used in the beginning of the cam-
paign, it was essentially a DDT program, a mixture of 10 per cent DDT
in talc being used.
Procedure. The procedure used in applying louse powders is de-
scribed by Wheeler (lac. cit.) as follows:
It consisted essentially of forcefully blowing powder, by hand dusters
or power dusters between the layers of clothing worn by the individual and
between the innermost layer of clothing and the skin of the body. This was
accomplished by a uniform technique, inserting the nozzle of the duster up
the sleeves, down the neck (both front and back), around the waistline and
into the crotch area of clothing. Hair and any cap or hat were dusted
thoroughly. An infested person properly dusted is no longer a menace to
others and will remain so for a period of at least two weeks, at the end of
which time he should be redusted. Approximately 1 to Ph oz. of powder
per person is sufficient to insure the thorough dusting of all clothing worn.
The total number of dustings were reported as 3,265,786. By February

20, 1944, the typhus fever epidemic in Naples appeared to have been
definitely broken, and upon direction of Brigadier General L. A. Fox, the
majority of the United States Typhus Commission members were called
back to headquarters in Cairo.
In the use of louse powders under military conditions, dusting with
power dust guns should follow a certain routine to avoid missing some
parts of the clothing being worn. Also in spite of the fact that few lice
occur in blankets, bedding, extra clothing, and mattresses, these must
be dusted to prevent reinfestation. If the treatment is done by the in-
dividual himself with a "sifter can" it is necessary to remove all the
clothing from the body. From 1 Y2 to 2 oz of the powder is needed on an
average per person; however, under field conditions as much as 4 oz
may be needed. Lice are usually killed in 24 hours, but the eggs are not
killed; however, because of the lasting qualities of DDT all the young
lice will be killed eventually on hatching. Where troops or others are
working in intimate contact with infested populations, semimonthly
dusting is recommended to prevent reinfestation.
Classical typhus has been the scourge of armies throughout recorded
history. During World War I military hospitals in the Balkan countries
were crowded with typhus patients, thousands being admitted daily
during 1915. In striking contrast there were only 64 cases of louse-borne
typhus in the American Army during all of World War II (Philip) .
Treatment for head lice. Among the many and various methods em-
ployed in the eradication of head lice, from the primitive use of fingers,
still a practice among natives as many of us have observed, and the use
of a fine-toothed comb dipped in kerosene, which some of us have ex-
perienced, to the most modern use of DDT louse powder, only a few
need be mentioned here: .
1. A towel wet with 10 per cent solution tincture of larkspur wound turban-
like around the head for several hours, repeat in about a week.
2. Washing the hair and scalp with 2 per cent solution of creolin; repeated in
about a week.
THE LICE 127
3. A wash made of a mixture of 5 level tablespoonfuls of derris or cube
powder, 2 level tablespoonfuls of neutral soap, and one quart of warm
water, applied thoroughly to the hair, mopped up with a towel without
drying the hair, and allowed to remain for at least eight hours; repeat in
about ten days if necessary.
4. An organic thiocyanate lotion recommended by Busvine and Buxton"
(1942), Lethane 284 Special: 12.5 per cent N-butyl carbitol thiocyanate,
37.5 per cent beta-thiocyanoethyl laurate; 50 per cent refined paraffin
(Busvine and Buxton diluted the paraffin to 25 per cent); apply a little of
the fluid to several parts of the scalp, exposed by parting the hair, then
massage with the fingers.
5. Dusting the scalp with derris or cube powder containing 3 per cent
rotenone; part the hair little by little and bring the powder in contact with
the scalp with the fingers; leave the powder in the hair for 48 hours before
washing.
6. Dusting the hair and scalp with DDT or MYL louse powder previously
described, rubbing the powder in with the hands; an additional treatment
is suggested in a week to 10 days later; the head should not be washed
for at least 24 hours after each treatment.
7. NBIN formula, recommended by the United States Bureau of Entomology
and Plant Quarantine:" benzyl benzoate 68 parts by weight, DDT 6 parts,
benzocaine (or ethyl p-amino-benzoate) 12 parts, and Tween 80 (sorbitan
monooleate, polyoxyalkylene derivative) 14 parts; dilute in 5 volumes of
water and apply to scalp and hair; one treatment will kill all lice and nits.
Treatment for crab lice. Usually the treatments effective against head
lice can also be used against crab lice, Phthirus pubis. The liquid or
powder must be applied to the pubic and anal regions of the body,
under the arms and wherever the body is hairy; in particularly hairy
persons the lousicide must be applied from neck to foot, perhaps also to
the eyebrows and beard. Whatever material is used it must be well
distributed and must reach the skin. DDT, MYL, and NBIN (already
described) as well as 5 per cent rotenone in 10 parts of petrolatum, are
all effective if used as directed.
SUCKING LICE OF MAMMALS
Anoplura affecting domesticated mammals. The important sucking'

lice of domesticated ungulate mammals belong to the family Haematopi-
nidae and are arranged mainly under three genera, Haematopinus.
Linognathus, and Solenopotes. Swine have one species of louse, H aemato-
pinus suis (Linn.) = (H. urius N itzsch ) (Fig. 45). This is the largest
species of the entire group, measuring as much as 5 to 6 mm in length;
it is cosmopolitan in distribution. The members of this species occur
chiefly in the folds of the skin on the neck, at the base of and inside the
ears, along the belly, and on the inner sides of the legs. According to
" Publication 606 on DDT.
128 . MEDICAL ENTOMOLOGY
Florence"7 hog lice feed readily on man but will not feed on guinea pigs.
The same author summarizes, as follows, the life history of these lice at
35° C, when they were kept continuously next to the body in vials: in-
cubation period of eggs (Fig. 46), 18 to 15 days, first molt after five to
six days, second molt after four days, third and last molt after four to
five days, sexual maturity after three days, a total of 29 to 88 days from
egg to egg.
Cattle lice belonging to the Anoplura number five species, namely,
(1) Linognathus vituli (Linn.), commonly known as the long-nosed ox
louse or "blue louse," measuring about 2 mm in length and distinguished
45 46
FIG. 45. Hog louse, Haematopinus suis. X 7.
FIG. 46. Nits (eggs) of the hog louse attached to the hair of the host. One
of the eggs has hatched. X 10.
from the next species by its long nose and slender body; cosmopolitan in
distribution; (2) Haematopinus ellrysterntlS (Nitzsch) , the cosmopolitan
short-nosed ox louse, somewhat larger (8.5 to 4.75 mm in length) than the
former and much broader in proportion; (8) Haematopinus tuberculatus
(Burm.), the buffalo. louse, infests cattle in parts of Australia; common
on cattle and carabao in India; absent in North America; measures
from 8.5 to 5.5 mm; resembles H. eurysternus closely but differs in that
the number of setae at the margin of the abdominal segme'nts caudad of
the paratergal plates is usually eight or more (may be five or six); (4)
Haematopinus quadripertusus Fahr., the tail louse; (5) Solenopotes
capillatus Enderl . has been redescribed by Bishopp28 and shown to have
a wide distribution in the United States as well as in many other parts
of the world. It is known as the little blue cattle louse. It measures from
1.2 to 1.5 mm in lE'ngth and in general appE'arance resembles the short-
nosed ox louse.
All species exce pt the tail louse show a tenden(;y to attach on the
head, neck, and shoulders of the host, where the eggs are glued to hairs.
In heavy infestations manv parts of the animal are infested . The eggs
THE LICE 129
of the short-nosed ox louse are usually whitish in color, while those of the
long-nosed louse are dark blue, and those of tIle Solenopotes pale yel-
lowish. The incubation period averages about 12 days with a range of
from 8 to 15 days. The life cycle from egg to egg requires from 20 to 30
days. There are three molts at about four-day intervals. Lamson 29 reports
that the short-nosed ox louse lays from 35 to 50 eggs over a period of
from 10 to 15 days. Anyone concerned particularly with cattle lice will
certainly consult Bulletin 832, Cornell University Agricultural Experiment
Station (Matthysse O ).
Horses, mules, and asses are frequently infested with one species of
sucking louse, Haematopinus asmt (Linn.) = [H. macrocephalus
(Burm. )], which measures from 2.5 to 3.5 mm in length. It resembles
the hog louse except that the head is relatively longer and more robust.
The lice usually congregate at the base of the mane and forelock and the
root of the tail. Hall Oi (loc. cit.) reports that 22 out of 38 horses examined
were infested with this species. The same author reports the incubation
period for the egg to range from 10 to 19 days (eggs kept in Petri dishes
at from 21 0 to 31 0 C). It is quite probable that this species requires
about the same time as the hog louse for the completion of its life cycle,
i.e., from 29 to 33 days from egg to egg.
Sheep in some parts of the United States are affected by the so-called
foot louse, Linognathus pedalis (Osborn). The author has observed this
species on sheep from California and Nevada, ~nd Osborn has reported
it from Iowa. It occurs on the legs, espeCially jn the region of the dew
claws, but in heavy infestations it may actually invade the wool above
the knee. Deaths have been reported as due to this louse. The species
measures about 2 mm in length; and as it is the only sucking louse likely
to attack sheep in the manner described above, its identity can be easily
established. Goats often suffer heavy infestations of Linognathus stenopsis
(Burm.).
Sucking lice on dogs and rabbits. Dogs are commonly heavily infested
with Linognathus piliferus (Burm.), and rabbits harbor H aemodipsus
ventricosus (Denny).
Control of sucking lice of mammals. Poorly fed animals, crowded pens,
aJld dark, insanitary quarters are factors in the multiplication of lice, but
the parasites may gain a foothold in spite of plenty of food, clean quar-
ters, and adequate space; hence the stock farmer living in climates per-
mitting this should not neglect to install a good dipping vat as a part
of his equipment. Although lice will not breed away from their host,
they may drop off with hair and may remain aHve for probably not over
five days; and the same animals after dipping or other animals of the
same species, introduced into quarters before the dropped lice have died,
may become reinfested. Furthermore, the great majority of the ordinary
dips do not destroy the eggs present at the time of treatment; hence a
second dipping is usually necessary after the young lice have hatched.
This second dipping should be properly timed and in most cases should
be done 16 to 18 days after the first. Dipping procedures are described
by Babcock and Laake,32 who recommended for cattle-louse control two
dippings in a sulfur-rotenone dip, containing one pound of cube (5 per
cent rotenone) and 10 pounds of wettable sulfur in 100 gallons of water.
Application of dusts containing 10 per cent derris or cube (0.5 per cent
or more of actual rotenone) is highly recommended by Matthysse (loc.
cit.) for the control of all species of cattle lice. Experiments with DDT
indicate that it is not as effective as rotenone; however, a 0.3 per cent
DDT emulsion dip has been reported effective with one treatment (Fur-
man).
Furman 33 (1947) reports complete eradication of Haematopinus
eurysternus and Linognathus vituli with a single dip in a 0.3 per cent
suspension of benzene hexachloride (approximately equivalent to 0.036
per cent gamma isomer). This material, which is now being used widely,
should not be applied to lactating dairy animals. Furman and Hoskins 34
( 1948) have shown that benzene hexachloride appears in the milk for
several days following application. It is also known that this material may
occur in the body fat of treated animals as long as 30 days after treat-
ment where a concentration approximately double that used for louse
control.
B. THE BITING LICE

Order Mallophaga
Classification. The biting lice ( Mallophaga), of which there were
about 1,400 described species 35 in 1916, and about 2,500 in 1937 accord-
ing to Muesebeck, may be divided into two suborders: (1) Amblycera
with short, clavate or capitate, four-segmented antennae, concealed in
shallow cavities on the under side of the head; four-segmented maxillary
palpi, mandibles horizontal; and ( 2 ) Ischnocera, with short, slender
three- or five-segmented, exposed antennae, no palpi, mandibles vertical.
The Amblycera are divided into six families among them ( 1 )
Gyropidae, having tarsi with one claw, infesting mammals only, e.g.,
Gyropus ovalis N itzsch and Gliricola porcelli (Linn.), both of the guinea
pig; (2) Menoponidae, having tarsi with two claws, infesting birds
mainly, e.g., Menopon gallinae Linn. (= Menopon pallidum Nitzsch),
the common hen louse, and Trinoton luridum Nitzsch of ducks; (3)
Boopiidae, a small family of biting lice on kangaroos and wallabies, con-
fined to Australia, e.g., Heterodoxus longitarsus (= H. armiferus Paine),
may also occur on dogs. The Ischnocera are divided into four families,
THE LICE 131
among them (1) Trichodectidae, with antennae three-segmented, tarsi
with one claw, infesting mammals only, e.g., Bovicola bovis (Linn.)
[= Trichodectes bovis (Linn.)] [( = Trichodectes scalaris Nitzsch)] of
cattle; and (2) Philopteridae, with antennae five-segmented, tarsi with
two claws, infesting birds mainly, e.g., Goniodes stylifer Nitzsch, the
large turkey louse, and Lipeurus caponis (Linn.), the wing louse of
chickens.
Injury done by biting lice. The injury done by the biting lice is largely
restricted to poultry, although some trouble may result when mammals
are badly infested. The injury is largely due to irritation or to itching
caused by the creeping insects and their incessant gnawing at the skin.
This irritation causes the host to become exceedingly restless, thereby
affecting its feeding habits and proper digestion; egg production in fowls
FIG. 47. Eggs of biting lice (Mal-

lophaga) on feathers of a bird. J
is greatly reduced and development retarded. A lousy flock of poultry
is not a good investment. 'When lice are abundant uncleanliness and
overcrowded conditions usually exist.
The life history of a biting louse. One of the most complete life history
studies of a biting louse of birds was made by Martin 36 on the pigeon
louse, C olumbicola columbae ( Linn. ) (= Lipeurus baculus N itzsch ) ,
family Philopteridae. She found that as many as sixty white, opaque
eggs of this louse are attached to a single feather (Fig. 47) . The in-
cubation period at 37° C was normally from three to five days; at 33° C
it ran from 9 to 14 days. At the latter temperature the nymphs always
died in from one to six days. There are three instars, each requiring on
an average slightly less than seven days, making a total of between 24
and 25 days for the complete life cycle. At 37° C the adults live usually
from 30 to 40 days, !he longest time being 51 days . Temperature is the
chief factor in determining the length of the life cycle as well as the
survival of the young.
The life cycle of Bovicola bovis (Linn. ), the red louse of cattle (family
Trichodectidae) is described by Matthysse (loc. cit.) as follows: at
95° F and 75 per cent humidity, the egg stage averages 5 to 9 days, first
ins tar 6 to 10 days, second instar 4 to 10 days, third instar 4 to 9 days,
preoviposition period 3 to 4 days, cycle egg to egg 22 to 42 days; maxi-
mum adult longevity was 42 days. The egg-to-egg cycle under normal
conditions would probably average 29 to 30 days .
Lice infesting domestic fowls. More than forty species of lice are
said to occur on domestic fowls of which seven species are commonly
found on chickens. Losses due to poultry lice are most evident among
the young birds, but heavy infestations on older fowls result in loss of
48 49 50
FIG. 48. The common shaft louse of poultry, Menopon gallinae.
FIG. 49. The large hen louse, Goniodes gigas. X 10.
FIG. 50. A turkey louse, Goniodes stylifel'. X 14.
weight, lowered egg production, and lowered vitality. Although other

maladies may present similar symptoms, infested fowLs are droopy,
lower the wings, present an unkempt and ruffled appearance, and suffer
from diarrhea. The commoner lice of chickens are (1) the "body louse,"
Eomenacanthus stramineus (Nitzsch) (= Menopon biseriatum Piaget) ,
a rapidly running species occurring on all parts of the fowl; it is light
yellow in color and about 2 mm in length; it lays its eggs in large clusters,
particularly on the small feathers below the vent; the egg stage requires
about a week and maturity is reached in about two weeks thereafter;
(2) the "shaft louse," Menopon gallinae Linn. (= Menopon pallidum
Nitzsch) (Fig. 48) , which resembles the body louse very closely but is
smaller in size, and occurs mainly on the shafts of the feathers; it is said
to gain its nourishment from barbs and scales of the feathers and is
THE LICE 133
therefore not so irritating as the body louse; according to Bishopp and
Wood 37 it does not occur on young chickens and deposits its eggs singly,
at the base of the feathers between the main shaft and the after shaft;
the life history appears to require a somewhat longer time than that of
the body louse; (3) the "head louse" Cuclutogaster (= Lipeurus) heter-
ographus Nitzsch, a dark grayish species about 2 mm in length, infesting
the head and neck of young chickens to which it is most injurious; it
deposits its eggs singly on the down or small feathers about the head
and requires about the same time for complete development as the first
species mentioned; (4) the large hen louse or "blue bug," Goniodes gigas
Tasch. (Fig. 49), about 3 mm in length; broad with rounded head, and
smoky gray in color; it is generally distributed over the body and easily
recognized; Eomenacanthus stramineus Nitzsch (= Menopon stramineum
Nitzsch) is also referred to as the large poultry louse, infesting turkeys
and other fowl as well as chickens; (5) the "wing louse," Lipeurus caponis
(Linn.) (Lipeurus variabilis Nitzsch), also known as the "variable louse,"
a long slender species about 2 mm in length; the margins of the head
are black, the head is large and rounded, and the general appearance is
sufficiently distinct to separate it from all the other species; (6) the
"fluff louse" Goniocotes hologaster Nitzsch, a very small and broad species
about 1 mm in length, pale in color and seldom abundant; see Ancona 38
for morphology of this species; (7) the "brown louse" Goniodes dissimilis
Nitzsch, reported for the southern United States by Bishopp and Wood;
it is described by them as somewhat smaller than the large hen louse,
reddish brown in color, and found on the feathers of the body.
Turkeys are commonly infested with the large (3 mm long) Goniodes
stylifer N itzsch (Fig. 50), which has the posterior angles of the head
extended backward into long projections terminating in stylets or bristles.
Another louse found on turkeys is Lipeurus polytrapezius Nitzsch, a long
slender species measuring from 3 to 3.5 mm in length. Eomenacanthus
stramineus Nitzsch (2.0 to 2.5 mII,l) occurs on turkeys also. Ducks and
geese harbor a rather small-sized species, Docophorus icterodes Nitzsch,
measuring about 1 mm in length; the head of this species is curiously
expanded and rounded in front and is a darkish red; the thorax is also
red with darker bands. Another species infesting ducks and geese is
Lipeurus squalidus N itzsch (Fig. 51), which is a bout 4 mm in length,
with the head longer than broad; it is very slender and is light yellowish
in color. Another long species infesting ducks (4 mm) is Trinoton luridum
Nitzsch, dark grayish in color with triangular head about as long as broad.
None of these species appears to become abundant enough to be of any
great consequence. The common lice of the swan are Docophorus cygni
Denny, about 1 mm in length; "in color the head, thorax and legs are
bright reddish brown while the abdomen is white in the center and
dark brown at the sides, the brown occupying hard-like portions at the
side of each segment"; and the extremely large and common Ornithobius
bucephalus Pia get (4 mm long). The latter is conspicuous because of its
size; the body is white and quite transparent.
Pigeons are often abundantly infested with Columbicola columbae
(Linn.) (= Lipeurus baculus Nitzsch), a very slender species measuring
about 2 mm in length; Goniodes damicornis Nitzsch, a broad brownish
species about 2 mm long; and Goniocotes bidentatus Scopoli, about 1 mm
in length, whitish in color wIth a head rounded in front. Guinea fowls
are said to harbor Goniodes numidia nus Denny, Lipeurus numidae
FIG. 51. A duck louse, Lipeurus squalidus. X 19. (Re-

drawn after Osborn.)
Denny, and Menopon numidae Gieb.; pea fowls, Goniocotes rectangu-

latus Nitzsch, Goniodes falcicornis Nitzsch, and Menopon phaeostomum
Nitzsch; pheasants, Goniocotes chrysocephalus Gieb., Goniodes colchicus
Denny, Lipeurus heterographus Nitzsch, and Menopon fulvomaculatum
Denny.
Control of poultry lice. No remedy has given such uniformly satis-
factory results in the control of the lice of all kinds on domesticated
birds as has sodium fluoride (NaF), apparently first used against these
parasites by Bishopp and Wood in 1917. Sodium fluoride can be ob-
tained in two forms, a white powder or commercial form (90 to 98 per
cent pure) and in fine crystals or chemically pure form . For louse control
the former more finely powdered form is preferable. It retains its ef-
ficiency almost indefinitely if kept in a dry place in stoppered bottles or
THE LICE 135
cans. One application generally will destroy all lice present. It may be
applied in three ways, viz., the pinch method, dusting, and dipping.
The pinch method consists of placing on the skin of each fowl ap-
proximately ten "pinches" (amount held between thumb and forefinger)
of the commercial sodium fluoride, distributed on the breast, each thigh,
below the vent, on each side of the back, on the neck, on the head, and
finally one sprinkled on the underside of each outspread wing. The
birds, when treated, should be held over a shallow pan or newspaper in
order that the excess of the chemical may be saved.
For dusting, the powdered sodium fluoride is sometimes mixed with
three or four times its bulk of flour or talc. It is applied with a large
shaker, and the feathers of the bird are ruffled as this is being done.
This procedure is not as economical of material or as efficient as the
pinch method, and the excess of chemical in the air is irritating to birds
and operators.
Dipping in sodium fluoride solution is rapidly becoming a standard
method of treatment among a large group of producers that have over-
come the poultrymen's prejudice against wetting their birds. In warmer
climates, such as that of California, birds may be dipped safely in almost
every month of the year by choosing a warm day with little wind and
completing the operations an hour or two before sundown in order that
the fowls may dry thoroughly before roosting for the night. The solution
should be prepared in a wooden container, avoiding contact with galva-
nized iron. The ordinary wooden washtub is excellent for this purpose.
One ounce of the commercial sodium fluoride or two-thirds of an ounce
of the chemically pure crystals should be dissolved in each gallon of
tepid water. The best method for dipping the birds is to hold them with
the left hand by both wings. They are then placed feet foremost in the
warm dip and submerged until only the head remains above the surface.
They should be held in this position from 20 to 25 seconds while the
feathers are being ruffled to permit penetration of the liquid. Just before
removal the head should be ducked under the surface. The birds should
be held above the dip for two or three seconds to allow them to drip
before releasing them. One hundred birds will use up approximately 5
gallons of dip; material should be available at the start of operations to
keep the dip replenished on this basis.
The dipping method kills all lice immediately, but where the chemical
is applied as a powder three or four days pass before elimination is com-
plete. If the birds are caught and handed to the operator from 100 to 125
birds an hour can be treated by dipping or dusting and approximately
60 to 75 per hour by the "pinch" method.
Other methods. The very fact that poultry wallow in dust whenever
it is available indicates a means of partly controlling the bird lice. Special
box wallows, conveniently placed, broad and deep enough so that there
will be room for several birds at a time, should be partly filled wih;' fin e
road dust or ashes with the addition of a quantity of tobacco dust in the
proportion of about six parts of the form er to one of the latter. It is quite
desirable to add a few handfuls of sulphur. The finer the dust the better,
since it is believed by some that the dust particles enter and clog up the
breathing pores of the lice. However, it is more probable that the agita-
tion caused by the birds wallowing in the dust dislodges many of the
lice and they are thus lost in the shuffle.
Powdered derris root applied as a dust, or as a dip, one-fourth ounce
to a gallon of water, is said to be effective. This method is described by
Wells, Bishopp, and Laake.39
The application of 40 per cent nicotine (Black Leaf 40) to the roosts
in a thin layer with a brush or swab before the birds go to roost gives ex-
cellent results. The treatmen t should be repeated at an interval of 10 days
52 53 54
FIG. 52. The biting ox louse, Bovicola hovis. X 26.
FIG. 53. Biting louse of the Angora goat, Trichodectes hennsi. X 22.
FIG. 54. The biting dog louse, Trichodectes canis. X 35.
to kill the lice which have hatched from eggs on the birds. It is recom-
mended that the application be made when the weather is calm. Appar-
ently this treatment acts as a fumigant. Telford' o (1945) tested the
effects of 37 louse powders on several species of chicken lice and found
that practically complete control was given by DDT, 0.5 to 4.0 per cent;
derris (with 5 per cent rotenone) 10 per cent; sodium fluoride, 33 per
cent, cryolite, 30 per cent, and micronized wettable sulfur.
Biting lice of domesticated mammals. The biting lice of domesticated
mammals are for the most part rather easily identified by their presence
on a given host, as commonly not more than one species of Mallophaga
TIlE LICE 137
is found on each species of mammalian host. Cattle are often heavily
infested on the withers, root of tail, neck, and shoulders with Bovicola
bovis (Linn.) (= Trichodectes scalaris N itzsch ) (Fig. 52), a little red
louse about 1 Y2 nun in length, definitely marked with transverse bars
(ladder-like) on the abdominal segments. The biting lice are most
numerous on the animals during dry, cold weather when the hair is long.
Although they do not irritate the cattle as much as suckling lice do, the
following noteworthy observation is made by Imes: 41
When present in large numbers, however, they often form colonies or

groups around the base of the tail, over the withers, and on other parts of
the animal, and produce lesions resembling those of scab. These lesions vary
in size from that of a 25-cent piece to 4 or 5 inches in diameter. The skin
over these areas appears to be raised, and ringworm may be suspected; but
when the lesion is manipulated the scarf skin falls off, exposing the lice
grouped on the raw tissues beneath. Under such conditions the irritation is
very great and the damage to the animal may be fully equal to that caused
~~ili. .
Horses, mules, and asses, but horses more particularly, when poorly
or irregularly groomed may suffer from two species of biting lice,
Trichodectes equi Linn. (= Trichodectes parumpilosus Piaget) and
Trichodectes pilosus Giebel; the latter according to Kellogg and Ferris42
has not been reported from North America, although it is reported by
Hall (loc. cit.) on horses in Michigan. Trichodectes equi Linn. is
described thus by Osborn: 43
. The head is decidely rounded in front, the antenn~e inserted well back,
so that the head forms a full semicircle in front of the base of the antennae.
The abdomen is more slender and tapering than in bovis (scalaris) . . . . The
color is much as in the allied species, the head, thorax and legs being a bright
reddish brown, or chestnut, and the abdomen of a dusky yellowish color,
with about eight transverse dusky bands occupying the central or anterior
portions of the segments and extending from the middle line a little more
than halfway to the margin. They are hardly as conspicuous as in bovis
(scalaris).
T. pilasus Gieb. is a smaller species and the antennae are inserted well
forward, almost on a line with the anterior border. Hall states that these
"lice give rise to itching, and the results from this are often surprisingly
unpleasant. A barn full of horses may become a pandemonium as a re-
sult of lice. The itching animals attempt to relieve the itching by rubbing
and biting, other animals start to kick, presently the kicking becomes
general and there is a resultant clamor and din, with a substantial ele-
ment of danger to the horses and attendants." The life history is about
the same as for the biting cattle lice.
Sheep may at times show severe infestations of Trichodectes avis
Linn. (= T. sphaerocephalu8 Nitzsch). This species is about 1.5 mm in

length, the head being somewhat rounded, as long as broad, and red-
dish in color; the abdomen is whitish. Because of extensive dipping
operations against scab this louse has seldom had an opportunity to
thrive, but we have observed that when scab disappeared and con-
sequently no dipping was done, the biting lice appeared in troublesome
numbers.
Goats are very commonly enormously infested with biting lice .
. Several species from goats have been described, about which there is still
some confusion, but the common species is Bovicola caprae (Gurlt)
(= T. climax Nitzsch). The author has found Angora goats to be heavily
infested with Trichodectes hel'msi Kellogg and Nakayama (Fig. 53),
FIG . 55. A biting louse of deer, Trichodectes tibialis. (Left) male, (right)
female. X 31.
which resembles and is probably a synonym of T. penicillatus Piaget,

which in turn is said to be a synonym of T. limbatus Gerv., of T . crassipes
Rudow, and of T. maior Piaget. The irritation produced by the lice, par-
ticularly on Angora goats, causes the animals to rub or bite themselves
so much that the mohair is matted and pulled out, resulting in consider-
able loss. The individual hairs are weakened by the gnawing of the lice.
Sulfur dips (10 pounds wettable sulfur to 100 gallons of water) are
highly recommended. Two dips, with an ll-day interval between, are
recommended.
Dogs, particularly puppies, may suffer much irritation from a small
biting louse, Trichodectes canis De Geer (= T. latus Nitzsch) (Fig. 54).
It is a broad, short species, measuring about 1 mm in length. The author
THE LICE 139
took biting lice from a dog in Berkeley, California, many years ago,
which were described as H eterodoxus armiferus by Paine. 44 As this genus
is said to be restricted to the kangaroo the incident was interesting. The
later species is now considered to be a synonym of Heterodoxus longi-
tarsus Pia get, which today is frequently found on dogs in California.
Cats may become heavily infested with Felicola subrostratus
(Nitzsch) (Trichodectes subrostratus Nitzsch). Guinea pigs commonly
harbor two species, Gyropus ovalis N itzsch and Gliricola porcelli (Linne)
(= Gyropus graCilis Nitzsch). The llama harbors Trichodectes breviceps
Rudow. Trichodectes tibialis Pia get (Fig. 55) is very abundant on
California deer.
Control of biting lice on mammals. As a general rule lousicides effec-
tive against sucking lice are also effective agllinst biting lice, although
the reverse does not always hold. Sodium fluoride has proved effective
in the destruction of the biting lice on cattle, horses, goats, sheep, dogs
(dangerous to cats), and guinea pigs. Bishopp states that "a high degree
of effectiveness (90 to 100 per cent destruction) may be obtained by
applying the sodium fluoride with a dust gun to the flock in a pen as the
goats are driven through a chute. It does not seem to be necessary to
drive the dust into the mohair especially and only a small amount-
about one-third ounce per head-is necessary." Because of the toxicity
of sodium fluoride its use on dairy cattle is not recommended not only
because of possible danger to the animals but also because of milk con-
tamination. Furman (1947, loco cit.) found that Bovicola bovis infesta-
tions could be eliminated by a single dip in 0.15 per cent benzene
hexachloride. A persistent odor is a factor to be considered if this in-
secticide is used.
The following suggestion by Lamson (loc. cit.) for the treatment of
cattle lice in general is noteworthy, though perhaps of little importance:
Of the many different measures for the control of lice on dairy cows and
young stock, raw linseed oil gave the best results from the standpoint of econ-
omy of material and labor of application, killing all the lice but not injuring
the skin and at the same time not making it necessary to thoroughly drench
the cow. It has no poisonous properties. At the same time it is a logical remedy,
as the lack of oiliness in the skin of the cow is a fundamental reason for her
being lousy. Linseed oil can be put on at the time taken for grooming or
cleaning the cows, thus doing two things in one application. . . . From four
to five cows can be treated with a pint . . . treatment should be repeated
at least five times. Raw linseed oil, can be best applied with a brush having
bristles of unequal length. Do not rub the skin too vigorously when applying
the oil. Do not allow the animals that have been treated to go out in the
strong sunlight until at least twelve hours after applying the oil. Do not ex-
ercise the animals after the treatment. Do not use the boiled or refined lin-
seed oil.
14Q MEDICAL ENTOMOLOGY
Lice and taeniasis. Dipylidium caninum (Linn.), the double-pored
dog tapeworm, is a common parasite of the dog and is occasionally
found in humans, especially children. It measures from 10 to 14 inches
in length, has long seed-like proglottids and an armored scolex; as its
larval host it has the biting dog louse, Trichodectes canis DeGeer, the
dog flea, Ctenocephalides canis (Curtis), or the human flea, Pulex irritans
Linn. The cysticercoid stage has been experimentally produced in the
louse by placing ripe crushed proglottids of the tapeworm on the skin of
a dog infested with lice.
As has already been explained, the biting lice subsist on epidermal
scales, skin exudations, and other matter on the skin of the animal. This
habit makes it comparatively easy for the louse to become infected by
swallowing "eggs." The dog, on the other hand, readily infects itself by
devouring the lice (or fleas) which irritate its skin.
Persons, particularly children, while fondling louse-infested (or flea-
infested) dogs may easily become infected by accidentally swallowing
lice (or fleas) which contain bladder worms (larval tapeworms). This
is more readily accomplished if the person is eating while fondling dogs.
BIBLIOGRAPHY
1. Ferris, G. F., 1919-35. Contributions toward a Monograph of the
Sucking Lice, parts 1-8. Univ. ser. "Leland Stanford Junior Univ. PubI.," no.
41. and Univ. ser. "Stanford Univ. Publ. BioI. Sc.," vol. 2. nos. 2-8.
2. Ferris, G. F., 1934. "A summary of the sucking lice (Anoplura) ,"
Entomological News, 45:70-74, and 85-88.
3. Murray, Andrew, 1860. "On the pediculi infesting the different races
of mall," Tr. Roy. Soc. Edinburgh, 22:567. (Cited by Osborn.)
4. Nuttall, G. H. F., 1918. "The biology of Phthirus pubis," Parasitology,
10:383-405.
5. ---, 1917. "The biology of Pediculus humanus," Parasitology, 10:80-
185.
6. Buxton, P. A., 1939. The Louse. London: Edward Arnold & Co. viii
+ 164 pp.
7. Leeson, H. S., 1941. "The effect of temperature upon the hatching
of the eggs of Pediculus humanus corporis DeGeer," Parasitology, 33:243.
8. Pierce, W. Dwight, 1918. The Life History of the Human Louse.
Washington, D. C.: U. S. Dept. Agric., Bur. Entomol., in Proceedings of the
Class Formed to Study the Entomology of Disease, Hygiene, and Sanitation.
(Mimeograph copy received from author.)
9. Peacock, A. D., 1916. "Structure of the mouth parts and mechanism
of feeding in Pediculus humanus," Parasitology, 11:98-117 (1 plate).
10. Moore, W., 1918. "An interesting reaction to louse bites," 1.A.M.A.,
71:1481-82.
11. Mackie, F. P., 1907. "The part played by Pediculus corporis in the
transmission ?f relapsing fever," Brit. M. /., 2: 1706-09.
THE LICE 141
12. Nicolle, C. N.; Blaizot, L.; and Conseil, E.; 1913. "Etiologie de la
Fievre nkurrente. Son mode de transmission par les poux," Ann. Inst.
Pasteur, 27:204-25.
13. Chung, Huei-Lan, and Feng, Lan-Chou, 1936. "Studies on the de-
velopment of Spirochaeta recurrentis in body louse," Chinese M./., 50:1181-
84.
14. Ricketts, H. T., and Wilder, R. M., 1910. "Further investigations re-
garding the etiology of tabardillo, Mexican typhus fever," /.A.M.A., 55:309-11.
15. Nicolle, Charles; Comte, C.; et Conseil, E.; 1909. "Transmission ex-
perimentale du typhus exanthematique par Ie pou du corps," Compt. rend.
Acad. d. sc., 149:486-89.
16. Ricketts, H. T., and Wilder, R. M., 1910. "The transmission of the
typhus fever of Mexico (tabardillo) by means of the louse (Pediculus vesti-
menti)," J,A.M.A., 54:1304-7.
17. Goldberger, J., and Anderson, J. F., 1912. "The transmission of typhus
fever, with special reference to transmission by the head louse (Pediculus
capitis)," U. S. Public Health Service, Pub. Health Rep., 27:297-307.
18. Nuttall, G. H. F., 1917. "Lice and disease," Parasitology, 10:43-79.
19. Da Rocha-Lima, H., 1916. "Untersuchungen tiber Fleckfieber," Mun-
chen. med. Wchnschr., 63:1381-83.
20. Toepfer, H., 1916. "Zur Ursache and Uebertragung des Wolhynischn
Fiebers," Munchen. med. Wchnschr., 63:1495-96.
21. Foster, M. H., 1918. "Preliminary report on carbon tetrachloride vapor
as a delousing agent," U. S. Public Health Service, Pub. Health Rep., 33:1823-
27.
22. Moore, W., and Hirschfelder, A. D., 1919. An Investigation of the
Louse Problem. Minneapolis: Univ. Minnesota Press, in Research Publ., vol.
8. 86 pp.
23. Moore, William, 1918. "The effect of laundering upon lice (Pediculus
corporis) and their eggs," J. Parasitol., 5:61-68.
24. Bushland, R. C., 1948. "Insecticides for the control of lice attacking
man and animals." Proc. 4th Internat. Congo Trop Med. & Malaria, Abstracts.
Washington, D. C.
25. Wheeler, Charles M., 1946. "Control of typhus in Italy 1943-1944
by use of DDT," Am.].Pub.Health, 36:119-29.
26. Busvine, J. R., and Buxton, P. A., 1942. "A new method of con-
trolling the head louse," Brit.M.]., 1:464-66.
27. Florence, Laura, 1921. The Hog Louse, Haematopinus mis Linne: Its
biology, anatomy, and histology. Ithaca: Cornell Univ. in Agric. Exper. Sta.
Memoir, No. 51, pp. 641-743.
28. Bishopp, F. C., 1921. "Solenopotes capillatus, a sucking louse of cattle
not heretofore known in the United States," f. Agric. Research, 21: 797-
80l.
29. Lamson, G. H., Jr., 1918., 1918. "Cattle lice and their control," Storrs
Agric. Exp. Sta., Bull., No. 97, pp. 397-414.
30. Matthysse, John G., 1946. Cattle Lice, Their Biology and Control.
Ithaca: Cornell Univ., in Agric. Exper. Sta. Bull., no. 832, 67 pp.
31. Hall, M. C., 1917. "Notes in regard to horse lice, Trichodectes and
Haematopinus," J. Am. Vet. M. A., 51, n. s. 4:494-504.
32. Babcock, O. G., and Laake, E. W., 1942 The Control of Cattle Lice.
Washington, D. C.: Dept. Agric., in Bur. Entomol. & Plant Quar. Circular,
E 447: 1-2 revised (Mimeograph ser.)
33. Furman, Deane P., 1947. "Benzene hexachloride to control cattle
lice," J. Econ. Entomol., 40:672-75.
34. ---, and Hoskins, W. M., 1948. "Benzene hexachloride in cream from
cows' milk," ]. Econ. Entomal., 41:106.
35. Harrison, Launcelot, 1916. "The genera and species of Mallophaga,"
Parasitology, 9:1-156.
36. Martin, Margaret, 1934. "Life history and habits of the pigeon louse,
Columbicola columbae (Linn.) ," Canad. Entomologist, 66:6-16.
37. Bishopp, F. C., and Wood, H. P., 1917. Mites and Lice of Poultry.
Washington, D. C.: Dept. of Agric., in Farmers' Bull., no. 801, 27 pp.
38. Ancona H. L., 1935. "Contribucion al conocimiento de los piojos de
los animales de Mexico, III Goniocates hologaster Nitzsch," An.Inst.Biol.,
Mexico, 6:119-28.
39. Wells, R. W.: Bishopp, F. C., and Laake, E. W.; 1922. "Derris as a
promising insecticide," f. Econ. Entomol. 15:90-5.
40. Telford, Horace S., 1945. "New insecticide for chicken lice control,"
f.Econ. Entomol., 38:573-76.
41. Imes, Marion, 1925. Cattle Lice and How to Eradicate Them. Wash-
ington, D. C.: Dept. Agric., in Farmers' Bull., no. 909. 24 pp. (Revised).
42. Kellogg, V. L., and Ferris, G. F., 1915. The Anoplura and Mallophaga
of North America. Univ. ser. "Leland Stanford Junior Univ. Publ." 74 pp.
+ 8 plates.
43. Osborn, Herbert, 1896. Insects Affecting Domestic Animals. Washing-
ton, D. C.: Dept. Agric., in Div. Entomo!. Bull., no. 5 (n. s.) 302 pp.
44. Paine, J. H., 1912. "The Mallophagan genus Heterodoxus Le Souef
and Bullen," Entomological News, 23:359-62.
CHAPTER X
._.
GNATS (EXCLUSIVE OF MOSQUITOES)

Simuliid Gnats, Phlebotomus Flies, Hippelates Flies
( FAMILIES SIMULIIDAE, PSYCHODIDAE, CHLOROPIDAE)

Order Piptera
Order Diptera. The several families of gnats discussed in this chap-
ter are members of the Order Diptera, "the flies," which comprises some
75,000 described species belonging to some 140 families (see Curran,
1934).1 Many species of insects belonging to this order are involved in
the transmission of important diseases of man and animals; hence the
medical entomologist must be thoroughly familiar with the Diptera.
As the name implies, all of the winged members of the Order have
only one pair of wings; the posterior pair is represented in nearly all
species by a pair of short (often minute) knobbed organs known as
balancers or halteres. In certain families, e.g., muscoid flies, there is pres-
ent a pair of minute wing-like structures at the juncture of the wings
with the body; these are known as squamae or allulae. Conspicuous com-
pound eyes are present, and most species possess three simple eyes
( ocelli). The metamorphosis is typically complete, consisting of the four
stages-egg, larva, pupa, and imago (adult). Some species are vivip-
arous, notably the tsetse flies. The mouth parts, as previously described,
are subject to great variation although all are suctorial; many species are
provided with very effective piercing stylets which enables them to
"bite" fiercely and to suck blood.
Much attention must be given the larval stages because the larvae
frequently invade the tissues and organs of the body of man and animals,
causing myiasis (see Chapter XVII); also a wider knowledge of aquatic
larvae (of which there are many species) is important in pursuing work
with mosquitoes and other gnats, as well as in the study of the biology
of water supplies (see Johannsen 2 ).
The Diptera have a wide range of breeding habits. There are very
few habitats suitable for animal life which have not been invaded by the
flies. The petroleum fly, Psilopa petrolei Coq., as its name implies,
actually passes its larval life in crude oil.
143
Classification of the Diptera. In the classification of Diptera, knowl-

edge of wing venation is important (Fig. 56). The great diversity of
antennal structure provides a useful series of characters, as does the ar-
rangement of spines (chaetotaxy) on the body of certain species, such
as the blowflies ( see Walton 3 ). The terminalia are important taxonomic
structures in many Diptera.
The Diptera are usually separated into two major groups (1)
Orthorrhapha, referring to the species in which the winged insect escapes
1HEOBII.LO\1I. ANOPHELES
SIMULIUM MUSCA
STOMOXYS GLOSSINA
FIG. 56. Wings of Diptera. For explanation of venation, see Figure 12.
from the puparium through aT-shaped anterodorsal split, as horseflies,

buffalo gnats, and mosquitoes; and (2) Cyclorrhapha, in which the in-
sect escapes from the puparium through a circular opening; in fact it
pushes off the anterior cap by means of pressure exerted by the bladder-
like ptilinum on the front of the head of the insect, e.g., houseflies and
blowflies.
For the purposes of this book the Diptera are divided into three sub-
orders. (1) Nematocera, in which the antennae are filiform and many-
jointed, as in mosquitoes; (2) Brachycera, in which the antennae are
short, not filamentous, generally three-segmented, variously formed, as
in horseflies; (3) Cyclorrhapha as described above, and with antennae,
GNATS 145
brachycerous, generally three-segmented, and frequently bearing an
arista on the terminal joint; ptilinum usually present, as in houseflies and
blowflies.
SUBORDER I. NEMATOCERA
Larvae have well-developed, exserted head and horizontally biting

mandibles; pupa free. Antennae of imago are many-jointed, longer than
the head and thorax, and the majority of the joints are usually alike;
arista is lacking. Palpi are usually four- or five-jointed, pendulous. Discal
cell is generally absent; cubital cell when present, widely open.
SUBORDER II. BRACHYCERA
Larvae have incomplete head, usually retractile, and with vertically

biting mandibles; pupa free. Antennae of adult are shorter than thorax,
very variable, generally three-segmented with last elongate; arista or
style, when present, is terminal. Palpi are porrect, one- or two-segmented.
Discal call is almost always present, cubital cell is contracted before
wing margin or closed.
SUBORDER III. CYCLORRHAPHA
Larvae have vestigial head; pupa coarctate; antennae of adult are

three-segmented with arista usually dorsal in position. Palpi are one-
segmented. Discal cell is almost always present. Cubital cell'is contracted
or closed. Head with frontal lunule and usually with ptilinum.
Some Families of the Order DipteraO

Suborder N ematocera
A. Mesonotum with an entire V-shaped suture.. (Crane Flies) Tipulidae
AA. Mesonotal suture transverse, not V-shaped.
B. Costa continued around the margin of the wings, though weaker be-
hind the apex.
C. Wings short and broad, folded roof-like over the body when
at rest, usually pointed
(Moth Flies, Sand Flies, Owl Midges) Psychodidae
CC. Wings long, or if broad, the apex is very broadly rounded,
always lying Hat over the back when at rest.
D. Apical veins strongly arched. (Dixa Midges) Dixidae
DD. Veins straight or nearly so.
E. Proboscis not elongate, extending but little beyond the
clypeus; wings with scales (when present) confined
mostly to the fringe .. Chaoboridae
EE. Proboscis elongate, extending far beyond the clypeus;
wings with the veins and margins with scales
(Mosquitoes) Culicidae
• Classification adapted after Curran (lac. cit.).
BB. Costa ending at or near the apex of the wing.
C. Wings very broad, the posterior veins weak and poorly de-
veloped .. (Black Flies, Buffalo Gnats) Simuliidae
CC. Wings narrow and long, the posterior veins stronger.
D. Wings lying flat over the back when at rest; metanotum
short and without a longitudinal groove; femora some-
times swollen.. (Biting Midges) Ceratopogonidae
DD. Wings lying roof-like over the back when at rest; meta-
notum long and with a median longitudinal groove;
legs long and slender. (Midges) Chironomidae
Suborder Brachycera
A. Third antennal joint annulated; arista absent
(Horseflies, Deer Flies) Tabanidae
AA. Third antennal joint not annulated; but if annulated, with extremely
long flagellum with terminal arista; squamae vestigial; normally shaped
flies (Snipe Flies) Rhagionidae (Leptidae)
Suborder Cyclorrhapha
A. Anal cell closed very close to the wing margin; a spurious vein run-
ning obliquely between the third and fourth longitudinal veins
(Flower Flies) Syrphidae
AA. Anal cell usually shorter; no spurious vein.
B. Second antennal segment with a longitudinal seam along the upper
outer edge extending almost the whole length; posterior calli
definitely formed by a depression extending from behind the
base of the wings to above the base of the scutellum
(Calypteratae: Muscoidea)
C. Metascutellum developed, appearing as a strong convexity
below the scutellum; hypopleura (meron of Snodgrass) with
strong bristles (Tachina Flies) Tachinidae
CC. Metascutellum weak or absent, or if developed there is only
hair on the hypopleura.
D. Oral opening and mouth parts very small; hypopleura
with abundant long hair.
E. Scutellum extending far beyond the base of the
metanotum; metascutellum never developed
(Robust Botflies) Cuterebridae (Oestridae)
EE. Scutellum very short; metascutellum usually strongly
developed; palpi usually large
(Botflies) Oestridae
DD. Oral opening normal; hypopleura with a row of bristles
or only short sparse hair.
E. Hypopleura with a row of bristles.
F. Apical cell strongly narrowed apically
(Blowflies) Metopiidae·
• Includes the Sarcophagidae, part of the Tachinidae and Muscidae of Willis-
ton's Manual. (Curran.)
GNATS 147
FF. Apical cell not at all narrowed apically
Muscidae·
EE. Hypopleura with fine, short hair or bare . Muscidae
BB. Second antennal segment rarely with a well-developed dorsal seam;
the posterior calli not differentiated (except in Gasterophilus):
squamae small (Acalyptratae).
C. Mouth parts vestigial, sunken in a very small oral pit
(Horse Botflies) Gasterophilidae (Oestridae)
CC. Mouth parts well developed; the oral opening large. Ocellar
triangle large; fifth vein with a distinct curvature near the
middle of the discal cell
(Frit Flies) Chloropidae (Oscinidae)
FAMILY SIMULIIDAE
(Buffalo Gnats-Black Flies)
Characteristics. The family Simuliidae, consisting of over 600
species:,5 includes insects commonly known as buffalo gnats, black flies,
and turkey gnats. They are small (1 to 5 mm long) bloodsucking flies,
FIG. 57. A buffalo gnat, Eusimu-

lium pectlarum. (Redrawn after (Gar-
man.)
with mouth parts blade-like and piercing in the female but more or less
rudimentary in the male. They are stout bodied and usually almost black
in color. The thorax presents a strong development of the scutum and
reduction of prescutum resulting in a prominent hump Fig. 57). The
antennae are ten- to eleven-jointed; the eyes of the female are distinctly
separated (in the male they are usually close together and prominent,
o Includes the Scatophagidae (Cordyluridae), Anthomyidae, and those Muscidae
(of the Williston Manual) lacking hypopleural bristles. (Curran.)
i.e., holoptic); ocelli absent; palpi four-jointed; wings broad and irides-
cent, with distinct alulae, the venation is characterized by a strong
development of the costal veins (Fig. 56). Buffalo gnats often occur in
enormous swarms during late spring and early summer in hilly sections
where swiftly flowing streams provide well-aerated water for larval
development. They are particularly abundant in the north temperate
and subarctic zones.
Although running water, such as that of shallow mountain creeks is
favored as a breeding place, the gnats may also breed in roadside
ditches of more slowly moving water. The gnats may be found in abun-
dance a mile or two from water, probably in search of food.
Life history. Eggs to the number of 350 to 450 per female are de-
posited in masses at the water surface of aquatic plants, logs, and water-
splashed rocks. Comstock says he has often watched the gnats hovering
over the brink of a fall where there was a thin sheet of swiftly flowing
water, and has seen them dart into the water, and out again. At such
times he has always found the surface of the rock more or less thickly
coated with eggs, and has no doubt that an egg is fastened to the rock
each time a fly darts into the water. The shiny eggs are at first creamy
white, changing to almost black.
The time required for hatching is from 5 to 30 days, depending on
temperature and motion of water. In running water at a temperature
of 20° to 22° C the incubation period is four to five days.6 The newly
emerged larva attaches itself by means of a caudal sucker to any sub-
merged object, such as a stone or log, and is kept from being washed
away by a silken thread. Movement from place to place is achieved by
shifting anchorage. In some favorable location, such as the rimes on the
downstream side of an old log partially damming a little stream, there
may be thousands of these tiny black spindle-shaped larvae ..The larvae
as well as the pupae are provided with gill filaments and usually remain
submerged or partly so. The larval period of some species is said to
require but three to five weeks. The food of the larvae consists of small
crustacea, protozoa, and algae. The larval period for Simulium ornatum
Meigen is given by SmarF as 7 to 10 weeks when temperatures in the
stream ranged between go to 15.5° C. At the end of the larval period
the organism spins a rough reddish-brown basket-like cocoon in which
pupation takes place. These cocoons are firmly attached to shallowly
covered objects such as rocks.
The pupal period is quite short in some species, requiring not over
five or six days; while in other species it lasts three to four weeks. Tem-
perature influences this stage, Le., cooler weather retards the emergence
of adults. Smart gives the pupal period for S. ornatum Meig. as 3.75
days at a constant temperature of 21 0 C. In some species there is con-
GNATS 149
tinual breeding from early spring to late autumn with overlapping gen-
erations; in others there is evidently one sudden brood coming fairly
early in the spring with stragglers following. They overwinter in either
the larval or egg stage. The life history, egg to adult, ranges from 60 days
to 15 weeks and over, depending on species and climatic conditions.
Larvae. The light brown to black larvae are cylindrical, twelve-seg-
mented, slightly thinner in the mid-region, and when 'fully grown are
from 10 to 15 mm in length (Fig. 58A). The posterior end of the body
is provided with a toothed disk-like sucker, composed of two modified
parapodia. The anterior proleg is also modified into a prehensile toothed
disk. By means of these organs the larvae move from place to place with
a looping motion. They attach themselves to rocks or other supports in
FIG. 58. (A) larva and (B) pupa of

Simulium; latter removed from cone-shaped
cocoon. (Redrawn after Lugger from
Washburn.)
A B
the water by means of the posterior sucker, the hooks of which they
insert into the network of silken threads produced by secretions from
the salivary glands with which they they covered the substratum. The
larvae may hang from threads produced in similar fashion or travel along
their length.
Although the larvae are provided with a well-developed tracheal
system, and nine pairs of spiracles may be observed, these are not open,
and respiration is carried on by means of gills, recognized as branched
retractile structures located dorsally on the last abdominal segment. The
fan-shaped filamentous structures located on the head are for the pur-
pose of creating a current, by means of which food is drawn into the
mouth.
Pupae. When the larvae are ready to pupate, each spins a crude
pock ~t-like cocoon open at the upper end. The pupae are provided with
respiratory filaments attached anteriorly to the dorsal portion of the
thorax. The filaments are often quite numerous and because of their
constancy in number in a given species may be of diagnostic value (Fig.
58B).
Classification. The family Simuliidae is divided into four genera
according to Dyar and Shannon,s viz.:-the three genera with the radius
setose on its entire length: (1) Parasimulium, with the radius joining the
costa at the middle of the costal vein, the radial sector forked, antennae
ten-jointed; (2) Prosimulium, with the radius joining the costal vein far
beyond its middle, the radial sector with a long fork, the second hind
tarsus without dorsal incision, the front usually broad; (3) Eusimulium,
with the radius also joining the costal vein far beyond its middle, but
with the radial sector simple, the hind basitarsus produced or not pro-
duced apically, tbe second hind tarsus with or without dorsal incision
and less than twice the width of the basitarus, and the front narrowed;
( 4) Simulium, the radius bare between the stem vein and the base of the
radial sector, the radial sector simple, the hind basitarsus produced
apically, the second hind tarsus with dorsal incision and less than twice
the width of the basitarsus. Smart (1945, loc. cit.) recognizes six genera,
viz., Parasimulium, Prosimulium, Cnephia, Austrosimulium, Gigantodax,
and Simulium.
Common species. Only one species is given under the genus Para-
simulium by Dyar and Shannon, namely P. furcatum Malloch from Hum-
boldt County, California. Prosimulium fulvum Coq. is a widely distrib-
uted species in the mountainous regions of the west and along the
Pacific coast from Alaska to California; P. hirtipes (Fries) is said to be
confined to the region east of the Mississippi and north of the Carolinas.
It rarely attacks man, according to Dyar and Shannon, though it is
known on occasion to bite rather severely; it is not considered an im-
portant pest of livestock.
Eusimulium pecuarum (Riley) [= Cnephia pecuarum (Riley) ac-
cording to Smart] is known as the southern buffalo gnat and is a great
scourge of livestock as well as of man in the Mississippi Valley. During
the height of the gnat season in the early spring, work on plantations is
often greatly handicapped because of the annoyance to work animals.
E. minus D. and S. [= Cnephia minus (D. and S.)] is a widely distrib-
uted western and Pacific coast species with the type locality indicated
as Yosemite. It resembles the buffalo gnat but is smaller and darker.
Simulium pictipes Hagen occurs in the eastern United States. It is
said to be an inoffensive species. Simulium vittata Zetterstedt is wide-
spread throughout North America and is a common species in Europe.
It attacks man and livestock freely. Simulium occidentale Townsend and
Simulium meridionale Riley, known as "turkey gnats," are also common
GNATS 151
and widespread species in North America but particularly in the south-
ern states, where the gnats appear in late spring following the buffalo
gnat. They attack poultry, biting the combs and wattles, and are said
to cause symptoms similar to "cholera," hence the name "cholera gnat."
Simulium venustum Say is the black fly. It is one of the most annoying
and widespread species. It torments fishermen and campers in New
England and Canada. The gnats occur in the greatest numbers during
June and July.
The bite. There is perhaps no other insect of equal size that can in-
flict so painful a bite as can the buffalo gnat. It is a day biter and is
rarely found indoors. The mouth parts are of the dipteron type (similar
to those of the horsefly), consisting of six blade-like lancets.
Human beings as well as domesticated animals are viciously attacked.
The eyes, ears, nostrils, wrists, and all exposed parts of the body of man
are subject to attack. The extreme pain, intense itching, and the result-
ant local swelling, and occasional severe complications, indicate the
presence of an active venom. Losses due to the bite of this fly are con-
siderable and are estimated variously by stockmen. Myriads of these
gnats appear after the spring floods of the Mississippi River and its tribu-
taries. Horses, mules, and cattle are often killed in a few hours by the
venomous bites and loss of blood. This sudden appearance of the gnats
is explained by the large accumulation of eggs that have been washed
into the area during floods. These eggs do not hatch until the next flood
causes movement of the water, for it is in flowing and well-aerated water
that the larvae hatch and develop rapidly.
Simulium columbaschensis Fabr. is the famous Columbacz gnat of
middle and southern Europe. Patton and Evans 9 (p. 193), citing Ciurea
and Dinulescu, report that in 1923 two immense swarms of this fly
invaded southwest Roumania in May, June, and July, causing the death
of 16,474 domestic animals, including cattle, horses, pigs, sheep, and
goats. Large numbers of deer, foxes, and hares, as well as other wild
animals, were killed at the same time, according to these authors.
Relation to disease. Owing to the vicious intermittent bloodsucking
habits of the buffalo gnats, it has long been suspected that they might
playa role in the transmission of disease.
With the startling report of Dr. Louis W. Sambon'o in 1910 ascrib-
ing the transmission of pellagra to a buffalo gnat, the study of the
Simuliidae gained new impetus. The gnats, however, have no relation to
_ this disease. Goldbergerl l states (1914) that pellagra is neither infectious
nor contagious, that it is essentially of dietary origin, and that the disease
does not develop in those who consume a mixed, well-balanced, and
varied diet. The most important disease transmitted by Simuliid flies is
onchocerciasis (onchocercosis), a disease of natives of certain parts of
Africa, Mexico, and South and Central America. It is caused by Rlarial

worms, Onchocerca volvulus (Leuckart ) (= O. caecutiens Brumpt),
which require black gnats as intermediate hosts. The female worm meas-
ures from 35 to 50 cm in length, the male 20 to 40 mm. These worms oc-
cur in conspicuous subcutaneous nodular tumors located primarily on the
trunk, shoulders, and head of infected persons. Several adult worms and
numerous larvae (produced viviparously) usually occur in each tumor.
Serious involvements of the eye occur, often resulting in complete blind-
ness evidently due to migration of the larvae. Strong 12 points out that
from a clinical standpoint the association of ocular disturbances (such as
photophobia) with the disease is emphasized by the high percentage of
failing vision and blindness in a locality where at least 95 per cent of the
populations are infected with the parasite and have demonstrable nod-
ules. The student should particularly consult a treatise on ocular oncho-
cerciasis by Hisette.13
Blacklock,14 working in Sierra Leone, has shown that when the micro-
filariae are taken up with the bite of Simulium flies, they migrate from
the fly's stomach, finding lodgment in the thoracic muscles, where further
development takes place, and then travel to the head and finally to the
labial structures of the fly, where escape is made when the fly bites, and
infection of the human being is accomplished. Development in the gnat
requires about 10 to 14 days. The time required for development in the
human from inoculation by the bite to maturity of the worm in the skin
nodule is said to be about four months. The species of fly observed in
these experiments was Simulium damno sum Theob., a widely distributed
black gnat of tropical Africa. BequaerP5 points out that of 57 species of
Simuliidae described from the Ethiopian region, five only are definitely
reported as biting people, but all must be looked upon with suspicion as
possible carriers of onchocerciasis, although S. damnosum Theob. and
S. neavei Roubaud are the only ones positively incriminated.
Strong'6 investigated the disease in Guatemala, where he reports it is
characterized by the formation of nodular tumors (containing adult
filariae) situated on the scalp, although they may occur on the trunk and
in the pelviC region. Vargas states that three species of black gnats are
reported to be vectors of the infection in Guatemala and Mexico (in the
states of Oaxaca and Chiapas, Mexico, according to Dampf17 and Maz-
zottPS). Simulium ochraceum Walker is said by Vargas (verbal com-
munication) to be the principal vector in these areas; others of less im-
portance being Simulium callidum Dyar and Shannon (= Simulium
mooseri Dampf) and Simulium metallicum Bellardi. Vargas has recently
reported the presence of onchocerciasis in Venezuela, the vector being
unknown.
Bovine onchocerciasis. It has been pOinted out by Steward19 that
GNATS 153
bovine onchocerciasis is of considerable economic importance in Austral-
ia, that the "worm nodules" due to Onchocerca gibsoni Cleland and
Johnston cause losses to the state of Queensland estimated at £500,000
per annum. The work done by Steward in England with Onchocerca
gutturosa Neumann proved that this latter parasite is transmitted by
Simulium ornatum Meigen. He showed that the filariae are conveyed from
the skin of the cow with the bite of the fly, thence to the mid-gut of the
insect, where development commences. Ten days after ingestion they
reach the "sausage" stage in the thoracic muscles, and by the nineteenth
to twenty-second day they migrate forward to the head ready for emer-
gence from the probosciS when the gnat feeds again.
Leucocytozoon infections of poultry. The name Leucocytozoon was
given to certain Sporozoa found in the blood of birds by Danilewsky in
1890, and in 1895 Theobald Smith discovered a Leucocytozoon in the
blood of turkeys; this parasite was named Leucocytozoon smithi by
Volkmar. In 1932 Skidmore,2o working in Nebraska, reported the success-
ful transmission of this parasite by Simulium occidentale Town. In 1938
Johnson et al. 21 reported transmission through the agency of Simulium
nigroparvum (Twinn). Johnson and his associates state that when the
organism is taken into the stomach of the fly gametes are formed, macro-
gametes being clearly observable as well as the zygote.
An important infection of both domestic and wild ducks, caused by
the protozoon parasite, Leucocytozoon simondi Mathis and Leger, 1910
(= L. anatis Wickware 1915), occurs in Michigan according to O'Roke 22
who proved that the disease is transmitted by the black fly, Simulium
venustum Say. Other vectors are S. occidentale Town and S. nigroparvum
Twinn. The development of the organism within the body of the gnat is
cyclico-propagative, resembling closely the life cycle of the plasmodium
of malaria in anopheline mosquitoes. O'Roke states that the asexual cycle
in the duck requires 10 days and the sexual cycle in the gnat not to ex-
ceed five days, with field evidence that it may be as short as two days
or less.
Black-gnat control. Knowing the breeding habits of black gnats, it
will be appreCiated that larval control is a difficult task. The writer has
repeatedly recommended that, in the vicinity of communities, streams in
which these insects are breeding should be kept as free as possible from
debris, including submerged roots and dipping branches of overhanging
trees. Although this can be done, prevailing winds may nevertheless bring
swarms of gnats from a distance; but the removal of debris from streams
lessens the opportunity for them to disposit their eggs. Old logs lying
crosswise of a small stream are a particular menace because shallow
waterfalls are thus usually produced and afford ideal breeding places for
the gnats. The fact that the larvae tend to congregate in masses in cer-
tain parts of streams in which they are breeding makes it possible to
remove them in quantities when they are once located.
The newer repellents such as indalone, Rutgers 612, and dimethyl-
phthalate are useful but require frequent application according to
Vargas. 23 Experimental work in larval control now being carried out by
several investigators indicates that a good measure of success can be
achieved by the use of either DDT or TDE as applied in tests for the
control of chaoborid gnats. Romafia and Abalos report successful control
of larvae in streams by the use of gammexane in wettable mixtures at the
rate of 700-1,000 mg per l/sec of water. Ann. d Inst. med. Regional,
Univ. Nacional de Tucuman, Argentina, Dec., 1948.
R. D. Glasgow, New York State entomologist, reports economical
and successful control of black flies (adults) by applying a DDT aerosol
fog by means of a helicopter. About one quart of a 15 per cent solution
of DDT in a petroleum solvent was used per acre.
FAMILY PSYCHODIDAE
(Moth Flies-Phlebotomus Flies)
Family Psychodidae. The family consists of several hundred species
and includes the tiny gnats known as owl midges, moth flies, and
Phlebotomus flies (sand flies). The ovate, usually pOinted wings and the
body are densely covered with hairs, whence the name moth flies. In the
Psychoda flies the wings when at rest lie roof-like over the abdomen. Be-
cause of the faint transverse venation the wings appear to have only
longitudinal veins. The antennae are usually fairly long and from twelve-
to sixteen-segmented.
The family may be divided into two subfamilies, (1) Psychodinae,
known as moth flies or owl midges, which are not bloodsuckers and
whose wings are held roof-like over the body; and (2) Phlebotominae,
the sand flies, of which the females are bloodsuckers; they are never
aquatic.
Psychoda Hies. Several species of Psychoda are commonly found in
great numbers about sewage disposal plants, cesspools, and washbasins
in bathrooms where the larvae may develop in sink drains in spite of hot
water and soap. A common Pacific coast species is Psychoda paCifica
Kincaid, brown in color, measuring from 2 to 2.3 mm in length, and
moth-like. Although the flies of this genus are not bloodsuckers, they
may breed in such numbers in the filter beds of sewage disposal plants,
as the author has personally observed, as to constitute a real annoyance
to neighboring households. The corresponding eastern species, Psychoda
alternata Say is known as the "sprinkling sewage filter fly." These gnats
may become annoying in the house, where they originate in the surface
GNATS 155
of the gelatinous material in sink and bathroom drain traps. Their life
cycle is quite short, ranging from 8 to 24 days depending on temperature,
the average is about two weeks (see Mallis and Pence 24 ).
The adult gnats may be controlled by spraying their resting places
with a 5 per cent DDT emulsion or suspension at the rate of 1 qt to
250 sq ft of area. Caution must be exercised in the use of DDT emulsions
for the control of the larvae because of damage of filter growth by the
solvent. Simpson25 states that suspensions with a wetting agent eliminates
adverse effects. He recommends addition of a quantity of DDT at one
time equal to 1 p.p.m., based on daily flow, and retarding filter operation
Y2 to 1 hour for contact time and repeating the operation every two
weeks.
Phlebotomus ( = Flebotomus) flies. The genus Phlebotomus ( =
Flebotomus) of the family Psychodidae comprises many species of small-
sized hairy gnats or midges measuring from 1.5 to about 4 mm in length.
They are commonly known as sand flies. They differ from the Psychoda
flies, their nearest relatives, in that the wings are held upward and out-
ward so that the costal margins form angles of about 60 degrees with
each other and with the body. The second longitudinal vein branches
at a considerable distance from the base of the wing. The females alone
have piercing mouth parts and are bloodsuckers; some species are said
to feed on cold-blooded animals such as lizards and snakes;2G,27 others
feed on field mice; no doubt most of them feed on a variety of warm-
blooded animals including man. The males suck moisture from any
available source and are even said to suck up sweat from humans. These
gnats are active only at night when there is little or no wind, seeking
protection by day in shelters both out of doors in crevices, caves, and
among vegetation, and within buildings where they hide in dark corners.
Their noiseless flight, which is commonly described as weak, is usually
in short so-called "hops" when they are disturbed; however, in longer
flights their progress is slow and steady and can be followed with the
eye. Townsend 28 points out that "deep canyons, free from wind and dimly
lighted, are especially adapted to them."
Life history of sand flies. Hertig,27 who has had wide experience with
Phlebotomus flies, points out that "the breeding-places of sand flies are
notoriously hard to find and have been demonstrated in the case of only
a ' few species. They are typically under stones, in masonry cracks, in
stables, poultry houses, etc., in situations combining darkness, humidity,
and a supply of organic matter which serves as food for the larvae. In
no case is the breeding-place aquatic." To situations mentioned above
may be added hollow trees and animal burrows. The eggs are deposited
in small batches. The incubation period is said to be from 9 to 12 days
(Whittingham and Rook).26 The minute whitish larvae have long anal
156 MEDICAL ENTO!VIOLOCY
spines; the mouth parts are strongly mandibulate; they feed on organic
debris, excrement of lizards, etc.; larval stage with four instars requires
from four to six weeks; the pupa, which is naked, requires about 10 days
for development. The females usually lay eggs in five to seven days
after a blood meal. The egg-to-egg cycle requires from 7 to 10 weeks;
however, Hertig (loc. cit.) points out that where there are cold winters
Phlebotomus Hies are subject to a diapause in the fourth larval instar
which may last from several weeks to nearly a year. This. Hertig states,
was not the case in Peruvian sand flies (Fig. 59).
",\ '
3
FIG. 59. Life cycle of Phlebotomus verrllCarUIn. (1) eggs, (2) larva, (3) pupa
(P. noguchii), (4) adult female, (5) adult male. (Adapted after Hertig.)
Carrion's disease (bartonellosis) is known also under the names of its
two chief clinical forms, namely Oroya fever, a severe anemia, and
usually fatal, and verruga peruana, benign, and recognized by a charac-
teristic cutaneous nodular eruption. "The disease is endemic in a narrow
strip along the Pacific slope and in certain other parts of the Peruvian
Andes between latitudes 6 and 13 degrees south and at altitudes usually
between 800 and 3,000 meters ," affecting parts of Peru, Colombia
(Patiiio-Camarg0 29 ), Bolivia, Chile, and Ecuador. The etiological agent
is Bartonella bacilliformis Noguchi, a minute rod-like or coccoid organism,
"which occurs in or on the red cells and intracellularly in a number of
organs, notably in the endothelial cells of lymph glands."
The transmission of verruga was attributed to Phlebotomus Hies by
Townsend 3o ,31 in 1913. He described the vector as Phlebotomus verru-
r
GNATS 157
carum and attributed a human infection to this species. The author was
one of those who examined a British sailor presumably successfully in- .
fected experimentally (see Hertig, loco cit., p. 155). Although there was
some evidence of old cutaneous lesions no definite diagnosis of verruga
was made. Although the "Verruga Expedition Report"32 does not con-
sider Townsend's evidence as conclusive, the latter again presented his
arguments in favor of the Phlebotomus fly theory 33 (1915). Based on
careful field studies in the "verruga zone" Noguchi and Shannon, et. al.,34
substantially confirmed Townsend's contention that Phlebotomus verru-
carum is a vector of verruga and added another species Phlebotomus
noguchii Shannon as a probable vector. Hertig (loc. cit.) indicates some
of the taxonomic difficulties encountered in these investigations and cities
the experimental work of Battistini 1927-1931, whose publications were
not available to the author, in which positive transmission to a monkey
(positive in 18 days) was made through the agency (by bites) of wild
sand flies of a mixed lot of P. verruca rum and P. noguchii. In 1939 Hertig
in a personal communication to the author stated that he had discovered
a clue to the mechanism of infection in that the verruga organism was
recovered in pure culture from the extreme tip of the proboscis, the
piercing stylets themselves being thoroughly infected. In the commu-
nication, Hertig stated that this infection was apparently not acquired
from a blood meal. Transmission of verruga to monkeys under these cir-
cumstances was reported. The source of the proboscis infections is un-
known; however, the fact that the proboscides of male sand flies, which
are not bloodsuckers, are also thus infected, points to liquid sources other
than the usual blood meal of the females.
Sand fIy fever, also known as pappataci fever, three-day fever, Medi-
terranean dengue, or summer influenza, is a seasonal (hot, dry periods)
febrile virus disease of short duration occurring in the Phlebotomus in-
fested regions of the Mediterranean, South China, parts of India, Ceylon,
and parts of South America. It is a nonfatal infection. Commonly a large
percentage of a ship's company on shore leave, even for one night only
during the proper season, may be stricken with sand fly fever within two
or three days after leaving port. The incubation period in the insect is
from 7 to 10 days. There is some evidence that the infection may be
transmitted transovarianly, hence the insect may be itself a reservoir of
the infection. Phlebotomus papatasii Scopoli in the Mediterranean region
becomes infective six to eight days after an infecting blood meal; the
virus is present in man's blood for 24 hours prior to onset and for the
first 24 hours of the disease, hence the infective period for the sand fly
is limited to this length of time. After a person has been bitten by an
infected fly the incubation period is usually from three to six days; thus
in the Canal Zone the disease is known as Panama six-day fever; in the
J
158 / MEDICAL ENTOMOLOGY
Dutch East Indies it is known as van der Scheer five-day fever, and in
some parts of India as seven-day fever. Phlebotomus minutus Rond. P.
perniciosus Newst., P. papatasii Scopoli, and no doubt other species are
vectors of the virus.
Leishmaniasis is caused by various species of parasitic protozoa be-
longing to the genus Leishmania, round or oval intracellular bodies,
which develop flagellate herpetomonad stages in the gut of insects.
Leishmania infections are variously known. Kala-azar, dumdum fever,
or tropical splenomegaly is a visceral leishmaniasis, traceable to Leish-
mania donovani (Laveran et Mesnil) which localizes in the reticulo-
endothelial cells. It is a widespread disease, occurring in all countries on
the shores of the Mediterranean, south Russia, India, China, Manchuria,
northern (Mediterranean) and equatorial Africa, eastern Brazil and
other parts of South America. There is progressive enlargement of the
spleen and later of the liver. As the disease progresses the skin becomes
grayish in color whence the name "black disease." It is regarded as fre-
quently fatal, death resulting within a few weeks in acute infections
and in from two to three years in chronic cases. A case of exogenous
origin at the Marine Hospital, Baltimore, Md., is reported by Lewis
and Spicknall. 35 Various species of bloodsucking arthropods have been
suspected as being vectors. The low susceptibility of laboratory animals
made progress difficult, but with the discovery that hamsters were highly
susceptible to the infection rapid progress was made. Patton and
Hindle 36 (1927), as well as Young and Hertig 37 and Napier, Knowles,
and Smith before them (1925), observed that the Leishmania. bodies
underwent development in the intestinal track of Phlebotomus flies after
being ingested. Flagellation is said to take place in the mid-intestine.
The cycle in the gnat is completed in from four to five days. Evidence
of infection by the insect is very meager, though Shortt and Smith,
et al. 38 (1931), report successful transmission by the bite of Phlebotomus
argentipes Annandale and Brunetti, believed to be the Indian vector.
Other species, notably Phlebotomus chinensis Patton and Hindle in
China, and P. sergenti Parrot in northern Africa, are believed to be
vectors.
Transmission by the bite of the fly is doubted by Southwell and
Kirshner.39 These investigators point out that the bite of the sand fly
causes an irritation and the person bitten scratches the bite, thus crush-
ing and killing the infected insect. The leptomonads, the infective forms,
do not invade the mouth parts in either P. argentipes A. and B., or P.
chinensis P. and H. Thus infection by the bite is believed to be unlikely,
though infection as the result of crushing infected flies on the skin ap-
pears to be possible. Recently Gupta,40 1948, reporting on researches in
India states that sand flies kept alive on raisins after an infective meal,
GNATS 159
showed the phenomenon of "blocking," and kala-azar could be trans-
mitted almost invariably to hamsters and human volunteers by the "bite"
of the blocked sand fly.
Oriental sore. Oriental sore, also known as Bagdad or Delhi boil, is
a cutaneous leishmaniasis caused by Leishmania tropica (Wright); it has
a wide distribution in Mediterranean areas, Palestine, Arabia, Asia Minor,
Iraq, India, French Congo, and other parts of the world. It is not neces-
sarily coextensive with kala-azar. In oriental sore the leishmanias inhabit
the skin and do not invade the viscera. In a series of papers by Adler
and Theodor,41 evidence is advanced to incriminate Phlebotomus
papatasii Scopoli. These workers found a cyclical development of the
Leishmania in the fly requiring from 8 to 21 days~ Infection of a human
was accomplished by rubbing the infected mid-gut of the insect into the
scarified skin, and flies were reinfected successfully from the sore thus
produced. Infection by the bite did not occur. Adler 42 (1948) reports
that transmission by the bite of P. papatasii has been achieved. He re-
ports that P. sergenti Parrott is the most effective vector: The infection
is apparently perpetuated by sand flies inhabiting the burrows of various
species of gerbils, and this explains outbreaks among groups of human
beings passing through uninhabited regions.
American mucocutaneous leishmaniasis ( naso-oral), also known as
espundia or uta, is widely distributed in South and Central America.
The causal agent is Leishmania brasiliensis Vianna. The horrible disfigur-
ing effects of this infection are shown by Goldman,'3 who points out that
it is the mucosal involvement which is so characteristic of American
cutaneous leishmaniasis. Once the infection gets into the mucocutaneous
junction, it destroys all types of tissues including cartilage and bone.
Goldman states that "prevention of cutaneous leishmaniasis in the
endemic areas appears to be a well nigh hopeless affair. It is likely the
incidence could be reduced considerably by definite knowledge of vec·
tors, animal reservoir, and transmission. If the foreigner in the endemic
area wishes some simple practical advice, the use of modern repellents,
insecticides, adequate fine-mesh screening, and the other measures for
the maintenance of careful hygiene of the skin in the tropics, and early
attention to indolent lesions on exposed parts of the body may be sug-
gested." Various species of Phlebotomus flies are looked upon with sus-
picion based on experimental evidence, among these are P. intermedius
Lutz and Neiva, P. pessoai Coutinho, and P. longipalpus Lutz and Neiva.
Species of Phlebotomus. According to Addis" (1945) there are six
species of sand flies in the United States, none which have received
much attention; namely (1) Phlebotomus vexatar Coq. and (2) P. limai
Fonseca, both widely distributed in the southern states; (3) P. diabolicus
Hall from Texas, reported to feed on humans and experimentally on
Syrian hamsters; (4) P. texanus Dampf, also from Texas; (5) P. stewarti
Mangabeira and Galindo, from California, collected in wild rodent bur-
rows in arid fields; (6) P. anthophorus Addis, another Texas species, but
found also in Mexico. Addis reports that this species feeds only in the
morning, has never been observed after sundown, has a definite pref-
erence for rabbit blood, but will feed on hamsters in the laboratory; it
is apparently not annoying to man.
Investigators concerned with the numerous species of South Ameri-
can Phlebotomus Hies will need to consult the many excellent contribu-
tions of Mangabeira, particularly his 1942 contribution,'5 which c6ntains
a valuable bibliography.
Phlebotomus control. Marett46 suggests, as means of controlling Phle-
botomus, the facing of rock walls, removal of heaps of stones, blocking
of holes affording shelter for the gnats, encouragement of gardening
(cultivation of ground), -planting of embankments with native aromatic
plants, etc. As sand Hies are highly susceptible to pyrethrum sprays,
painting screens lightly each evening with a mixture of one part pyre-
thrum extract concentrate (20 to 1) and 20 parts of light lubricating oil
has been recommended. Because of the minute size of these gnats ordi-
nary mesh screens do not exclude them; therefore, a 0.0334-inch mesh is
recommended. Repellents such as "612" and dimethylphthalate applied
to skin and clothing give quite a bit of protection. Hertig47 states that
the habits and life history of sand Hies render them peculiarly vulnerable
to DDT. They make short Hights with relatively long pauses on entering
or leaving any shelter, breeding place, or structure-a procedure repeated
nearly every night of their adult life. Any surface capable of retaining
residual DDT, and on which sand Hies must alight, may become a lethal
barrier. House spraying gives virtually complete protection indoors.
Hertig states that, in Peru, house spraying, combined with treatment of
outdoor shelters such as stone walls, reduced the local sand Hy population
to negligible numbers, an effect which persisted up to a year and a half.
The short Hight range limits infiltration, and the long life cycle delays
recovery of a depleted sand Hy population.
FAMILY CERATOPOGONIDAE
(Biting Midges-Punkies)
Characteristics. The Ceratopogonidae are very small (0.6-5.0 mm

in length), slender, bloodsucking gnats (males do not bite) resembling
the nonbiting midges belonging to the family Chironomidae, to which
family they are commonly ascribed. In their biting habits they resemble
the black Hies (Simuliidae) and are .frequently mistaken for them. Among
GNATS 1ft!
the 20 or more genera comprising the family, three will serve the purpose
of this section, namely Culicoides (Fig. 60), Ceratopogon, and Lepto-
conops, commonly known as "punkies," "no-see-ums," "sand Hies," or
"Glack gnats." The wings, which are narrow, with few veins and no scales,
may be clear or hairy and are folded Hat over the abdomen when at rest;
alulae slender. The larvae are aquatic or semiaquatic, and are found in
fresh, brackish, or salt water; they may live in moist earth or in tree holes.
A key to the North American Culicoides numbering thirty species is given
by Root and Hoffman. 4 8
C1.Ilicoides canithorax Hoffm., C. melleus Coq., and C. dovei Hall
constitute a serious economic problem in the summer resort areas of the
Atlantic coast, particularly about fresh-water inlets and tide-water pools
FIG. 60. Female Culicoides sp. (After

Dampf.)
where these midges are most numerous. Dove, Hall, and Hu1l 49 report
that the larvae are found in decaying humus of the densely shaded
areas at the edges of the grass marshes of the upper Atlantic coast. The
period required for development appears to last from 6 to 12 months
according to these authors. The larvae and pupae of Culicoides gut-
tipennis Coq. have been taken from tree holes of the live oak in Missis-
sippi (Hinman). 50 Culicoides diabolicus Hoffman is a fiercely biting
species in Mexico.
Ceratopogon stellifer Coq. is reported to be a severe biter in Arizona
and New Mexico. Leptoconops torrens Townsend and Holocopops
kertesZ'i Kieff. constitue a serious pest in California. The former is known
as the "valley black gnat" and the latter as the "Bodega black gnat."
Both species bite ViCiously, feeding on man, domestic animals, and birds.
The bite usually produces a transient swelling, which may become
vesicular, rupture, and produce a open lesion which may exude moisture
for weeks. Itching resulting from the bite is intense. The following ac-
count of the two species is from a recent publication (1948) by Smith
and Lowe. 51 The valley black gnat passes the larval stage in clay or clay-
adobe soils situated along the western side of the Sacramento Valley and
in isolated deposits in the San Joaquin Valley. The Bodega black gnat
passes the larval stage in damp sand with some organic matter, at or
just above high-tide level in the mouths of fresh water streams which
enter Bodega and Tomales bays (California). A study of the morphology
of the larvae showed that the "Leptoconops group" is widely divergent
from typical ceratopogonids and warrants the acceptance of the sub-
family name Leptoconopinae (Enderlein 1936).
Adults of the Bodega black gnat occur from mid-April until early
October, in a continuous flow from the soil. Females may feed as many
as four times. Males form large swarms, dancing in the lee of windbreaks.
Eggs are laid on the surface of damp sand where the salt concentration
is about 640 p.p.m. The larval stage lasts for 8 to 10 months. Pupation
occurs in the sand; pupae wriggle to the surface and stand vertically be-
fore the adult emerges. The pupal period is 8 days. Adult females cap-
tured in the field lived a maximum of 11 days, with blood meals. Males
do not feed; they lived a maximum of 4 days after capture in the field.
Adults of the valley black gnat occur for 4 to 6 weeks, beginning
usually in the middle of May. Females feed only once; males do not feed.
Unfed gnats lived only 6 hours in captivity; with a blood meal, females
lived a maximum of 5 days. The larvae occur in clay-adobe soils at a
depth of 15 to 30 inches. Egress and entrance is dependent upon the
drying and cracking of the soil. The larval period is at least two years
in length. Larvae spend the summers in immobile estivation. If the soil
does not crack on schedule, the mature larvae enter a diapause. Some
evidence is given to indicate that larvae may diapuase for at least three
years. Larvae are found in summer in soil with a moisture content of 17
to 20 per cent, a salt concentration of 400 p.p.m., a pH of 9.6, and a
temperature of 65° to 68° F. Methods of identifying breeding grounds
consist of trapping adults as they emerge from the soil, and washing
larvae from the soil.
Culicoides austeni Carter, Ingram, and Macfie has been reported by
Sharp 52 as an intermediate host of Acanthocheilonema perstans (Manson)
[Dipetalonema perstans ( Manson) ]. The embryos of this worm are
found in the peripheral circulation both by day and by night. Sharp has
observed that diurnal periodicity is the more common. In the vast ma-
jority of cases it is said to be nonpathogenic. It is primarily equatorial and
African in distribution, though it occurs also in British Guiana and in
New Guinea. Sharp has shown that the microfilariae undergo meta-
morphosis in the body of Culicoides austeni Carter, Ingram, and Macfle,
GNATS 163
increasing to three times their original length before they appear in the
proboscis of the insect. The cycle in the By requires seven to nine days.
Sharp states that it is probable that Culicoides grahami Austen will also
prove to be a natural carrier. Buckley 53 found that Culicoides furens
Poey transmits the filarial worm, Mansonella ozzardi (Manson), in the
Antilles, and Dampf5' points out that this species of gnat is widely dis-
tributed along the coast of the Gulf of Mexico and the Caribbean Sea.
Control. Because of the variety of breeding places involved, it be-
comes necessary first of all to determine these for the species giving
trouble. Some species breed in salt marshes, in which case dikes, tide
gates and other salt-marsh control devices may be employed. Other
species breed in mud and plant debris along the margins of fresh-water
streams or ponds, and for the control of these removal of vegetation and
FIG. 61. A male midge (Chirono-

midae) , commonly mistaken for a
mosquito. X 12. (After Osborn.)
channelization are suggested, or filling in low ground may be helpful.
Still other species breed in holes in shade trees, in which case holes should
be treated with DDT or creosote, or otherwise made unfavorable for
breeding.
Punkies may be excluded from the house, according to the United
States Bureau of Entomology and Plant Quarantine, by applying a mix-
ture of 1 part pyrethrum extract concentrate (20 to 1) and 20 parts
lubricating oil (S.A.E. 5) to window screens by means of a brush or rag.
This mixture, it is reported, will exclude the gnats from the house for 24
to 48 hours. Some of the newer insecticides would, no doubt, prove

useful.
FAMILY CHIRONOMIDAE
(Midges)
Family Chironornidae. Although the midges are commonly mistaken
for mosquitoes they bear little resemblance to them on closer examina-
tion. In the midges the proboscis is short and not adapted for piercing,
the palpi are three- or four-jointed , the wings are bare or haired. The
antennae are plumose in the males and sparsely haired in the female
(Fig. 61) . Midges are widely distributed and may often be extremely
abundant in the vicinity of standing water, since the larvae are aquatic.
Occasionally great swarms of these insects hover in the air toward eve-
ning and produce 11 distinct humming sound. They are attracted to light
in great numbers. 1'he family is a very large one, comprising nearly 2,000
species.
Many of the larvae are red in color, hence the name "blood worm."
The larvae are worm-like (Fig. 62) and move by creeping or looping;
FIG. 62. Larva (left) and pupa (right) of a chirono-

mid gnat (midge). (Larva redrawn after Needham and
pupa redrawn after Grunberg.)
they have a closed (apneustic) respiratory system, hence need not come
to the surface for air as do mosquito larvae. Most species are bottom
feeders and scavengers in habit. While occurring most abundantly in
shallow shore water with vegetation such as reeds and tule, they have
been taken at great depths from the bottom of lakes; some species breed
in swiftly floWing water.
Burrill 55 in a very interesting paper on the swarming of midges states
that under the conditions observed they swarm an hour or two in the
early morning sunlight, then mostly stop flying and rest on such objects
as grass, the underside of tree leaves, tree trunks, and porch screens.
They may fly throughout a cloud~' day. He also observed a late summer
GNATS 165
swarm of Chironomus plumosus Burrill fly until after midnight. (See

also Gibson,5G 1945.)
FAMILY DIXIDAE
(Dixa Midges)
Dixa midges are usually placed in the family Culicidae (mosquitoes)
and are designated as a subfamily, Dixinae; however, for the purposes
of this work they are separated from the mosquitoes. They resemble
mosquitoes in wing venation but are almost devoid of hairs and scales;
and the proboscis, although somewhat projecting, is not fitted for piercing.
This family is mentioned here particularly because the larvae are fre-
quently mistaken for those of Anopheles, being commonly found in simi-
lar situations and also because the adults resemble and are related to the
true mosquitoes. Dixa larvae are usually seen at the surface of water
among vegetation and debris, moving in a horizontal U-shaped position.
The student is referred to O. A. Johannsen's'" "North American Dixidae,"
and H. G. Dyar and R. C. Shannon's "Some New Species of American
Dixa, Meigen."
FAMILY CHAOBORIDAE
(Chaoborid G11ats)
Family Chaoboridae. This family is usually regarded as a subfamily
(Chaoborinae) of the Culicidae, mosquitoes, but for the purposes of
FIG. 63. A chaoborid gnat, Chao-

horus astictopus, shoWing life cycle.
(Top left) female gnat. (top right)
eggs, (lower left) pupa, (lower right)
male gnat, (middle figure) larva.
this book the less commonly recognized family rank is used. The members
of this family are not bloodsuckers, and the gill-breathing larvae, which
live in deep water, are almost transparent and are seen with some dif-
ficulty, except when in motion, even in fairly clear water, hence their
name "phantom larvae." The tiny lead-colored cigar-shaped eggs are
deposited in great numbers on the surface of still water, such as ponds,
lagoons, lakes, etc. The eggs soon sink to the bottom. The incubation
period is less than 24 hours. The larvae grow slowly during the sum-
mer, reaching approximately full growth by winter, remaining thus
through the winter, and pupating in the early spring. The pupal stage
requires about two weeks. The pupae quickly come to the surface, where
the gmits literally "pop" out of the pupal skins, balance on the water
momentarily, and then fly shoreward. Chaoborus astictopus D. and S.
(= C. lacustris Freeborn) (Fig. 63) on which the foregoing life history
is based is a distinct nuisance along the shores of Clear Lake, California. 59
Lindquist 60 (1948) in an informal statement repqrted tests made with
TDE (dichloro-diphenyl-dichloroethane) which is less toxic to fish than
DDT, an important matter in the control of aquatic insects in large
bodies of water. He sprayed a small lake with an emulsion of TDE, and
some 85 per cent of the chaoborid larvae were killed. The dosage was
about one part of active ingredient to 45 million parts of water. The
material disseminated through the water to a depth of about 20 feet.
FAMILY CHLOROPIDAE
(Hippelates Flies-Eye Gnats)
Hippelates Hies. These flies are members of the family Chloropidae
(Oscinidae), commonly known as frit flies. Unlike the gnats discussed
above in this chapter, all Nematocera, the Chloropidae have short aristate
antennae. The members of the genus Hippelates are as a rule very small
flies (1.5 to 2.5 mm in length); they are frequently called "eye gnats"
or "eye flies" because they have a liking for lachrymal secretions, also
sebaceous secretions, pus, and blood. They may be distinguished by the
presence of a distinct curved, shining black apical or subapical spur on
the hind tibiae. They are extraordinarily persistent and if brushed away
will quickly return to continue engorging themselves. They are nonbiting;
however, the labellum is provided with spines which apparently act as
cutting instruments capable of producing minute multiple incisions, likely
to assist pathogenic organisms carried by the insects in gaining a foot-
hold (Graham-Smith61 ). The flies are easily mistaken for the pomace
fly, Drosophila. The larvae of most of the Chloropidae live in grass and
other plants (stem maggots); however, those of the genus Hippelates
develop in a wide variety of material such as decaying vegetable and
animal matter.
GNATS 167
Relation to conjUl").ctivitis. Siphunculina funicola de Meyere is known
as the "eye-fly" of India, Ceylon, and Java and is believed to be responsi-
ble for the spreading of conjunctivitis in these countries. Roy 192862 gives
a chart which shows that the seasonal prevalence of this fly in Assam
coincides closely with epidemic conjunctivitis.
Hippelates flies h:.tve long been looked upon with suspicion in certain
parts 01 the southern \)nlteu 'i)t'ates 'as posslb\e "\leeton di 'a IOTm oI con-
junctivitis commonly known as "sore eye," "pinkeye," etc. At a meeting
of the Entomological Society of Washington, held October 11, 1894,
E. A. Schwarz63 presented notes on Hippelates pusio Loew in the south-
ern states. Be stated that it was particularly abundant in Florida and
annoying to man and animals, and that it is attracted to eyes and to the
natural openings of the body as well as to infected wounds. In an article
in Insect Life (1895), Schwarz 64 throws much suspicion on Hippelates
flies as vectors of "sore eye."
For a number of years, at least since 1912, there have been numerous
cases of catarrhal conjunctivitis apparently of the follicular type in the
Coachella Valley of California where a veritable pest of Hippelates Hies
flourishes in season. Nowhere else in California are there such enormous
numbers of these flies and nowhere else in the state do as many cases of
so-called pinkeye exist. Several papers dealing with this subject were
published following investigations in the Coachella Valley.65-67
At the time of the Coachella investigations the flies were identified
as Hippelates flavipes Loew. In a letter dated May 19, 1927, J. M. Aldrich
writes, "The species which you sent from Coachella Valley, California, is
pusio Loew. It is the same species which was identified as flavipes Loew
by Malloch, Proc. U.S.N.M., 46, 1913, p. 245. His variety pusio on page
246 is a different form. The true flavipes of Loew, is the one described by
Malloch on page 243, as nitidifrons new species, as I have ascertained by
examining the types in the Museum of Comparative Zoology at Cam-
bridge, Mass. The earlier literature is somewhat uncertain since the
time of the original description, but probably your species is the same
one that has been referred to as flavipes when mentioning its annoying
habits." In the same letter Aldrich also writes, "In the Proceedings of the
California Academy of Sciences, Vol. 4, 619, Townsend described Oscinis
collusor from Lower California which he said was reported to cause
irritation of eyes of travelers and the 'mal de ojo' of natives. I examined
his types in the Academy shortly before their destruction by fire in the
spring of 1906 and found that they belonged to the genus Hippelates,
and it is quite prob[1ble that the species is pusio." The writer has traced
the Coachella Valley species through to the Mexican border at Mexicali.
The literature dealing with the Oscinidae as vectors of conjunctivitis
has been reviewed with great care by Graham-Smith. 68 This review indi-
cates a paucity of experimental evidence but a large amount of cir-
cumstantial evidence involving flies as spreaders of conjunctivitis in
Egypt, the West Indies, India, Ceylon, Java, and the United States.
Relation to yaws. As pointed out in the first chapter, flies have' for
many years been suspected as vectors of yaws (framboesia tropica),
and some experimental evidence has been advanced from time to time;
however, the evidence collected by Kumm 69 (1935) in Jamaica with
Hippelates pallipes Loew is most convincing. Kumm, as well as others,
has shown that it is relatively easy to demonstrate motile Treponema
,Pertenue Castellani in the "vomit drops" of eye gnats after they have
fed on infectious lesions of yaws. He found, however, that the spirochetes
were presumably digested in the mid-gut and hind-gut of the gnats very
soon after they were ingested, none being seen after an interval of two
days. There was no evidence of cyclical development.
The gnats receive the infection most readily by feeding on available
primary lesions which exude fresh infected serum with large numbers of
spirochetes. Inoculation is effected mechanically, i.e., the unchanged
spirochetes are deposited in "vomit drops"7o when infected gnats feed
on exuding serum from wounds, excoriated areas, or susceptible surfaces.
The manner in which the gnats receive the infection and their general
feeding habits are well described by Kumm, Turner, and Peat.'!
Bovine mastitis. Sanders 72 reports investigations at the Florida Agri-
cultural Experiment Station which incriminate Hippelates flies (species
not given) as well as the housefly, Musca domestica, as vectors of bovine
mastitis. Hippelates flies were seen to hover around the natural openings
of calves, yearlings, pregnant heifers, and lactating cows. They fed on
lachrymal fluid, fatty body secretions, milk droplets accidentally spilled,
and on secretion at the tip of the teats of animals in herds where mastitis
has prevailed. Exposure tests were made with flies feeding alternately
on infected material and the teat orifice; also the teat orifice was exposed
to flies taken directly from premises where mastitis prevailed. "Mastitis
developed in each of the experimental animals by the exposure technique
employed."
Life history of Hippelates pusio Loew. This species has a wide dis-
tribution in the southern United States where the winters are mild. The
adult flies are present throughout the year in the Coachella Valley (Cali-
fornia) and are particularly annoying during two periods, i.e., March,
April, May, and August, September, October. During June, July, and
early August the gnats are not abundant on account of extreme heat,
when the daily temperatures range well above 100 0 F. During the peak
months the adults are noticeable early in the morning and late in the
~fternoon and then in deep shade, such as densely planted shrubbery, in
GNATS 169
date gardens, and in the shade of the house. The fluted, distinctly curved
eggs are about .5 111m in length (Fig. 64). They are deposited on decay-
ing organic matter of wide range. The incubation period is about three
days. The larvae feed on a great variety of decaying organic matter in-
cluding excrement, provided the material is rather loose and well aerated.
According to Burgess (verbal communication) the larvae will not de-
velop naturally in closely compacted soil or putrid material, neither will
they breed naturally in excrement unless it is mixed with loose earth.
The larval stage under optimum conditions requires about 11 days. The
larvae may remain in this stage during the winter. Pupation takes place
FIG. 64. Hippelates }JlIsio. Egg; larva, showing cephalo-pharyngeal skeleton

and anterior spiracular process; pupa; adult fly. (After Herms and Burgess,
except adult fly which is redrawn after D. C . Hall.)
close to the surface of the material in which the larvae develop. The
pupal stage requires about six days, giving a total of about 21 days from
egg to adult fly. Except for overwintering adults, the first flies emerge
from the pupae of the overwintering larvae during late February and
early March, when the first great wave of the insects appears, as noted
above. Experiments performed by Hall' '' show that the larval stage
averaged about 1l.4 days on human excrement, 8.7 days on dog manure,
and about 17 days on decaying oranges. Burgess 7 .' points out that the
majority of Hippelates gnats are bred in soil that is (1) light and friable
(well drained), (2) freshly plowed (i.e., plowed not over three weeks
before), and (3) contains abundant humus or vegetable matter (cover
crops, manure).
Classification. The Hippelates flies possess a stout and distinct tibial
spur, which, with the following characte~s , will assist in the identification
of this genus: cepha liC bristles short, weak, and not conspicuolls; ace liars
very short, erect, and convergent or cruciate; fronto-orbital hairs minute,

slightly reclinate; no interhumeral bristles present (Sabrosky75). The
color of the body in the pusio group is black with a yellowish trace at the
base of the wings; the thorax is shiny black. Sabrosky places the follow-
ing species in the pusio group: H. pusio Loew, H. flavipes Loew, H.
pallipes (Loew), H. robensoni Sabr., H. bishoppi Sabr., H. bicolol'
Coq., and H. collusor Tns. The plebe;us group, in which the thorax is
more or less pollinose, with at least pre alar and prescutellar areas of
pollen, comprises the following species: H. plebe;us Loew, H. dorsalis
Loew, H. nobilis Loew, and H. proboscideus Will. The dissidens group,
according to Sabrosky, comprises smaller dark pollinose species, with
legs chiefly black, proboscis short and fleshy, hind tibial spur small. The
following species belong to this group: H. dissidens (Tucker), H. hermsi
Sabr., H. microcentrus Coq., H. montanus Sabr., H. convexus Loew, and
H. particeps Becker.
Control of Hippelates flies is difficult and involves a combination of
measures, such as trapping, with the use of finely chopped liver as bait;
sanitation, i.e., removal of garbage, manure piles, refuse heaps, and
decaying vegetable matter; also cultural methods such as light disking.
Tests made with DDT ground treatment in the form of water emul-
sion sprays and dusts proved unsatisfactory, and thermal aerosols gave
only temporary relief. Other insecticides, such as chlordane, C1oHsCls ,
are being tested.
Much more information concerning the breeding habits and behavior
of Hippelates flies is needed to provide a sound basis for permanent
control.
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Loew with spirochaetes of yaws," Tr.Roy.Soc. Trop. Med. & Hyg., 29:265-
72.
70. ---, and Turner, T. B., 1936. "The transmission of yaws from man
to rabbits by an insect vector, Hippelates pallipes Loew," Am.].Trop.Med.,
16:245-62 (5 plates).
71. ---; Turner, T. B.; and Peat, A. A.; 1935. "The duration of motility
of the spirochaetes of yaws in a small West Indian fly, Hippelates pallipes
Loew," Am. J. Trop. Med., 15:209-23.
72. Sanders, D. A., 1940. "Musca domestica and Hippelates flies, vectors
of bovine mastitis," Science, 92:286.
73. Hall, David G., 1932. "Some studies on the breeding media, develop-
ment, and stages of the eye gnat, Hippelates pusio Loew," Am.].Hyg., 16:
854-64.
74. Burgess, R. W., 1935. The eye gnat in the Coachella Valley, Califor-
nia, Washington, D. C.: Dept. Agric., in Bur. Entomo!' & Plant Quar. Circular
E-335 (Mimeograph ser.).
75. Sabrosky, Curtis W., 1941. "The Hippelates flies or eye gnats: Pre-
liminary notes," Canad. Entomologist, 73:23-27.
_.
CHAPTER XI
.
MOSQUITOES
ORDER DIPTERA, FAMILY CULICIDAE

Classification and Biology
Importance. Among the numerous species of bloodsucking arthropods
that annoy man and other warm-blooded animals, both wild and do·
mestic, mosquitoes stand out most prominently-their number is legion,
and they are an almost constant annoyance; there are some that bite
by night and others that bite by day. Great swarms may be produced
even in small quantities of water. They breed in practically all sorts of
water, fresh and salt, foul or potable; water in tin cans, broken gourds,
hoof prints, tree holes, reservoirs of impounded water; and vast stretches
of salt marsh. Great areas of seacoast are at times made uninhabitable by
salt-marsh mosquitoes, and agriculture may be retarded. Real estate
values suffer where the mosquito pest prevails, and losses resulting from
lowered industrial efficiency are frequently considerable because of mos-
quito annoyance. Economic losses due to mosquitoes would alone, no
doubt, amply justify the great sums now spent on mosquito abatement,
yet these losses are minor compared with the prodigious damage done to
the public health by mosquitoes as vectors of disease.
Mosquitoes are the sole vectors of the malarias, yellow fever, and
dengue, and they participate very importantly in the transmission of
filariasis and the encephalitides. It is reported that in an ordinary year
in India alone at least a hundred million persons suffer from, and a
million succumb to, the direct ravages of malaria. The indirect effect
in lowered vitality and susceptibility to other diseases accounts for
another million deaths. Considerable progress in spite of severe economic
handicaps is being made to alleviate this serious situation.
Family Culicidae. Although some authors, Edwards! and others, in-
clude the dixa midges and the chaoborid gnats (see Chapter X) as well
as the true mosquitoes in the Family Culicidae, only the latter are so
classified for the purposes of this book. Excepting the huge sturdily
built American gallinippers, Psorophora ciliata (Fab.) (body length
9 mm and wing spread 13 mm), and a few other species, mosquitoes
175
are small and fragile, ranging in body rength from 3 to 6 lllm. Abumlaut
scales are present on the wing veins and fringing the margins; the body,
head, and legs are scaly.
Mosquitoes are distinguished by their characteristic wing venatiull
(Fig. 65), as follows: subcosta (Sc) long, reaching the costa; radius
four-branched; R 2 + 3 forked; RH5 simple; no cross-vein connection of
Rl and Rz; media (M) two-branched; cross-veins r-m and m-cu both
present; Ax absent or very faint; cubitus (Cu) forked; An, long and
reaching wing-margin; characteristic scales clothing the wings, and more
or less abundant on the head and body (often scant or wanting); char-
acteristics of the thorax, such as the absence of a definite suture between
the pre scutum and scutum, completely divided pronotum (Edwards).
The true mosquitoes have elongate mouth parts formed for piercing, and
for bloodsucking in the females, though not all mosquitoes are blood-
suckers, and males are normally not bloodsuckers. The antennae are
long and filamentous with 14 or 15 segments, hairs in whorls, plumose il~
the males of most species; males of Opifex and Deinocerites do not
have plumose antennae.
There are about 1,600 described species of mosquitoes in the world;
about 120 in North America. 2 These ubiquitous insects occur at eleva-
tions of 14,000 feet in Kashmir and as low as 3,760 feet below sea level
in gold mines of south India (Russell et al. 3 ). For practical purposes
mosquitoes may be divided into four tribes: (1) Megarhinini, basal half
of proboscis rigid and distal portion flexible, the adults are flower-feeding
and the larvae are predaceous, e.g., Megarhinus inornatus Walker; (2)
Culicini, in which the palpi of the female are less than half as long as the
proboscis, scutellum trilobed, pulvilli present, eggs laid in rafts, e.g.,
Culex pipiens Linn.; (3) Aedini, generally included with the Culicini,
in which the palpi and scutellum are as in the Culicini, but the abdomen
of the female is pointed, postspiracular bristles are present, pulvilli absent
or hair-like, eggs are laid singly, e.g., Aedes aegypti (Linn.); (4) Anoph-
elini, in which the palpi of both sexes are as long or nearly as long as the
proboscis, scutellum rounded, without lobes, eggs laid singly, e.g., Anoph-
eles maculipennis Meigen (Fig. 65).
The larvae of mosquitoes are without exception aquatic and are dis-
tinguished from all other dipterous larvae by the possession of a complete
head capsule and the presence of only one pair of functional spiracles,
situated dorsally on the eighth abdominal segment. The medical ento-
mologist must be particularly familiar with larval characters in order that
he may make accurate identifications in the conduct of surveys and
inspections and in answering complaints (Fig. 65).
Male terminalia. The most critical differences used in the classification
of adult mosquitoes are to be found in the male terminalia 4 (Fig. 66).
CULEX PIPIEN S ANOPHELES MACUL\PE.NNIS AEDES AEGYPTI
f -- -- ---I ~--l
- \
I ~ !
-
FIG. 65. Morphological details and life histories of three genera of mos-
quitoes: Culex , Anopheles, and Aedes. Explanation of abbreviations: An. Pi.,
anal plate; An. pron. , anterior pronotal setae; Anl. Sdl., anal saddle (dorsal
plate); Ant., antenna; A.T., antennal tuft ; Cmb . sc., comb scale; D. Br., dor-
sal brush; D.H.H. , dorsal head hairs; Hal ., haltere; Hr. tft., siphon hair tuft;
I .C .H ., inner clypeal hair; L.A.T., lateral abdominal tuft; Mesemp ., mesepi-
meral setae; !lIn., mesonotum (tergum,) ; O.C.H., outer clypeal hair; Pal. Hrs .,
palmate or float hairs (tuft); Pec. Sc. , pecten scale; Pn., postnotum; Po. pron.,
posterior proTlotal setae', Pl'ea ., prealer &etae ; Proe-ps ., proepisternal setae;
P. Spr., postspiracular se tae; Resp . Trump. , respiratory tmmpet; Scl. , scutel-
lum; Sp. , spiracle ; St . pl. , sternopleural setae; V. Br., ventral brush. (Adapted
after various authors.)
177
it is of importance to know that the terminal portion of the abdomen

of the adult male begins to rotate on its axis within a few hours after
emergence from the pupa, and a rotation of 180 degrees is completed in
LT9
FIG . 66. Male termillalia of (A) Aedes squamiger, (8) Anopheles gam-
biae, (C) Anopheles freeborni, (D) Aedes aegypti, and (E) Culex quinque-
fasciatus. AL, apical lobe; ALC, apical lobe of cIaspette; Ap. D, appendage
of dististyle; Bs, basistyle; BL, basal lobe; BLC, basal lobe of claspette; BP,
basal plate; CL, claspette; CA, c1aspette appendage; Ds, dististyle; La, lacuna;
LT9 (=P-IXT), lobe of 9th tergite (=process of 9th tergite); P, para-
mere; Ph, phallosome; Ppt, paraproct; Pr, proctiger; SBL, spine of basal lobe;
VPP, ventral arm of paraproct, IXT = 9th tergite. (8 to E above after Ross
and Roberts.)
from 12 to 24 homs; this portion of the abdomen remains upside down
for the rest of the mosquito's life. The terminalia may be prepared for
microscopic study by first clipping off the tip of the male abdomen with
MOSQUITOES 179
fine scissors and then dropping it into 70 per cent alcohol for a few
moments, after which it is transferred to 20 per cent KOH for twenty
minutes, thence to a glass slide where the excess KOH is removed by
blotting the edge of the liquid; next a drop of glacial acetic acid is put
on the specimen, removing the excess by blotting; finally a drop of
chloral gum is placed on it and it is then covered with a cover-slip. After
drying, the specimen is ready to be sealed. Microscopic identification is
made by comparison with drawings of the terminalia of various species
in such publications as the Mosquito Atlas by Ross and Roberts. 5 A
description of the technique for dissecting male terminalia of mosquitoes
is given by Komp. 6
Life history of mosquitoes. All mosquitoes pass through the several
stages of a complex metamorphosis-egg, larva, pupa, and adult (Fig. 65).
Descriptions of the developmental stages of mosquitoes were published
as early as 1665 by Robert Hooke (Russell et al., lac. cit.). The larvae
are commonly known as wrigglers, and the pupae as tumblers. Water in
which to pass the larval and pupal stages is essential. Mosquitoes of
some species may deposit their eggs on moist earth or on the dry edges
of water-holding tree holes, and the larvae of some species may exist for
several hours under relatively dry conditions. Howard 7 states that "in no
case, however, were we able to revive larvae in mud from which water
had been drawn off for more than 48 hours, and after 24 hours only a
small proportion of the larvae revived." Eggs, on the contrary, in some
species may survive long periods of desiccation, notably those of the
yellow fever mosquito, which will hatch after being dry for a period of
six months. According to Dyar,S the eggs of Psorophora, with their spinose
protecting coat, are able to withstand desiccation on the dry ground for
months or years, hatching with the advent of water. Similarly the eggs
of various salt-marsh Aedes species may survive desiccation for several
years.
Mosquito eggs are deposited either singly or in rafts (Fig. 65) on
the surface of quiet pools of water and by some species along the mar-
gins, and even in fairly dry situations where pools may be formed later
by rains or tidal action. The incubation period varies greatly with the
species and temperature, from 16 to 24 hours in many of the Culicini;
in the case of the snow mosquitoes, boreal Aedini, eggs laid during the
summer and autumn remain buried under the snow through the winter
and hatch with the melting snow in spring; thus one brood is produced
annually.
The larvae of the various many-brooded species, most commonly
observed in rain barrels, watering troughs, and similar situations, hang
suspended diagonally from the surface by means of a prominent breath-
ing siphon with head downward as in the Culicini. Aedes larvae behave
similarly. The larvae of the tribe Anophelini lie horizontally just beneath
the surface of the water, suspended particularly by means of palmate
hairs (Fig. 65). The mandibulate larvae of practically all species secure
their food by browsing on microorganisms, both plants and animals. It is
not difficult to observe the feeding habits of the larvae as they squirm
about while breathing at the surface or wriggle down to the bottom or
along tfte sides niDbling food. Anopneline larvae are adapted for feeding
at the surface, as is indicated by the palmate hairs by means of which
they maintain a horizontal position and by their ability to rotate the head
through 180 0 while feeding against the surface film, which is laden with
bacteria and other microorganisms (Christophers and Puri). 9 The larvae
grow rapidly durin.g the warm summer, molting four times, the last molt
resulting in the pupa. An average of seven days is required for the larval
stage in several of our commoner Culicini under optimum conditions.
The larval stage of the Anophelini requires a somewhat longer time.
With the fourth molt the pupa or "tumbler" appears. In this non-
feeding stage ther~ is a pair of breathing "trumpets" situated dorsally on
the cephalothorax. The pupa is remarkably active and sensitive to dis-
turbances of the water, letting go suddenly and darting with a tumbling
motion to deeper \Vater and after a few moments rising with little motion
to the surface, where the breathing trumpets break the surface film and
contact with the air is re-established. The pupal stage is quite short,
usually from two to three days.
In a series of ~xperiments to determine the effect of various quanti-
ties of a given larval food on the development of a common species of
fresh-water mosqUito in California, namely Culiseta incidens (Thorn.),
the largest percentage of emergence was obtained when 2.5 grams ot
yeast were supplied per liter of distilled water. The egg rafts were first
placed in the water, each raft in a separate container, hatching in about
2Yz days. The ne\Vly hatched larvae were thoroughly mixed and trans,
ferred in lots of 100 to battery jars containing one liter of water with a.
measured quantity of Fleischmann's yeast; pH readings were taken.
before and after the yeast was added, and in all cases the pH reading~
were 6.6; daily pH readings were taken thereafter until all the mosquitoe~
had emerged, the pH remaining the same until pupation; the averag~
minimum room temperature was 19.20 C and the maximum 24.4 0 C. By
far the largest percentage of adults, 88 per cent, was produced in the jaJ:
with 2.5 grams of yeast, with a sex ratio of 5.1 males to 4.9 females. Th~
first molt took pla~e on the fourth day, the second molt on the sixth day,
the third on the ninth day, and the fourth on the twelfth day, when th~
first pupa appear~d; the first adult mosquito, a male, emerged on th~
eighteenth day, giving a total of about 21 days for the complete lif(:l
history, including the incubation period of the eggs. The complete record
MOSQUITOES 181
10
of the experiment is published elsewhere. When the tempe'rature was
maintained at 24° ± 1°C the life history was shortened to 12Y:z days-
egg stage 24 hours, larval stage 10 days, and pupal stage 36 hours.
Food habits of adult mosquitoes. The mouth parts of male mosquitoes
are not suited to piercing; hence males are not bloodsuckers. Their
nourishment is normally derived from nectar and plant juices and other
liquids. With the exception of a few species, such as the plant-feeding
Megarhinini and the Harpagomyia, which feed on regurgitated stomach
contents offered by ants (Cremastogaster) , all female mosquitoes are
able to pierce the skin of many kinds of animals and feed on blood. No
doubt vast numbers never have the opportunity to feed on blood. The
great majority of species are zoophilous, i.e., feed in nature on animals
other than man; some feed only on the blood of reptiles and amphibians,
e.g., Culex apicalis Adams. Species which feed on man by preference
are said to be anthropophilous. The feeding habits of mosquitoes may be
determined by applying the precipitin test to wild-caught engorged
females.
Flight habits of mosquitoes. Although most of the domestic species
remain fairly close to their point of origin, i.e., within a distance of a few
city blocks or half a mile, there are many species, particularly among
the Aedini, which may travel many miles. In searching for breeding
places of the common Culicini under urban conditions the point of origin
will usually be found not far from the points of complaint.
Salt-marsh mosquitoes are often a source of great annoyance far from
their breeding places, and a knowledge of their migratory habits is im-
portant in mosq uito control. Aedes sollicitans (Walker), an important
salt-marsh species of the Atlantic seaboard, is known to migrate at least
40 miles. The migrations of Aedes squamiger (Coquillett) have been
traced on the Pacific coast for a distance of 50 to 75 milesY Males are
seldom found far from their point of origin, although the males of Man-
sonia perturbans (Walker) are said to accompany the migrating females.
The flight range of Aedes vexans (Meig.) and A. aldrichi D. & K.
has been studied by Stage, Gjullin, and Yates 12 by using a stain, 1.5 per
cent aqueous solution of methylene blue or eosine, applied with a hand
compressed-air sprayer to newly emerged adult mosquitoes resting on
vegetation near their breeding places. Mosquitoes were collected in this
area at regular intervals until no more could be found. The collected
specimens were killed and each tested with one or two drops of solvent
made of 3 parts glycerin, 3 parts 95 per cent alcohol, and 1 part chloro-
form. The following results are recorded: (1) Both species and both
sexes were dispersed in all directions, with and against general wind
currents, for a distance of about two miles. (2) Males moved away from
the breeding areas more slowly than females. (3) Females of both species
traveled one-half mile across part of the Columbia River (Oregon)
within 24 hours after being stained. (4) One Aedes vexans (Meig.)
female was recovered 46 days after being stained, three miles distant
and across the Columbia River. (5) One Aedes sp. male was taken at
a point 5 miles distant 24 days after being stained. This was the greatest
distance for any positive flight record obtained. (6) The pests were
abundant to a distance of 15 miles from the breeding grounds and then
diminished rapidly until at about 30 miles only one female was taken
during a 10-minute search.
In conducting control operations for many years involving Anopheles
quadrimaculatus Say it has been assumed that the usual maximum flight
of this species is about one mile, and this procedure is in the main
sound; however, Eyles, Sabrosky, and Russell 13 recaptured marked in-
dividuals 2.0 to 2.7 miles from the point of release and a single in-
dividual at 3.63 miles. It is pointed out, however, that there was no
domestic blood supply sources between the flooded swamp and the
stations of recapture. The flight range of Anopheles freeborni Aitken
during dispersal and hibernation flight is, of course, much greater, being
as much as 25 miles.
Using the fluorescent dye, Rhodamine B, to mark the mosquitoes,
Reeves 14 and associates released over 20,000 Culex tarsalis Coq.; the
maximum range of recovery was 2.5 miles for females and 1.1 miles for
males. The majority of recoveries were made within one mile. On the
basis of these studies it is recommended that control measures for Culex
tarsalis be carried out in a zone not less than one and one-half miles be-
yond the human or animal hosts to be protected.
Longevity of mosquitoes. Male mosquitoes usually remain alive fm
but six or seven days, although Anopheles pseudopunctipennis francis-
canus Theob. males have been kept alive in our laboratories for over a
month, and Mayne15 was able to keep an A. punctipennis Say male alive
for 89 days and a female of the same species for 231 days; females with
ample food may live for four or five months, particularly under hibernat-
ing conditions. During their period of greatest activity it is likely that
the average lifetime of the females is not far from 30 days.
The staining experiment by Stage, et. al. (loc. cit.), produced im-
portant data relative to longevity. Thus six Aedes aldrichi D. & K. females
were taken 52 days after staining, one female of the same species 85 days
after staining, one Aedes vexans (Meig.) female after 55 days; also,
under especially favorable conditions, one 94-day-old A. aldrichi D. & K.
male was taken, and females of both Aedes aldrichi D. & K. and A.
vexans (Meig.) were collected from 104 to 113 days after staining. The
latter species is said to have the greater maximum longevity by approxi-
mately 15 to 20 days.
MOSQUITOES 183
Freeborn has found that increased humidity has a protective in-
16
fluence on the longevity of Anopheles freeborni Aitken kept at constant

temperature; but at a constant of 80° F no amount of relative humidity
can protect them for the full life span of a month. A relative humidity
of 55 per cent ensures the normal life span at 70° F. Freeborn points
out that 55 per cent humidity involves a saturation deficiency of 3.6
grains per cubic foot, by which can be expressed the drying power
of the air in the absence of wind currents. A deficiency of 3.8 grains at
76° F was tolerated for only three weeks instead of more than four.
It is pointed out that the lethal effect may be caused by either a fatal
600
110 500
400
o-cNumbers of adults (X'esent 300
<>-oNoonday temperature ( ~.F) 200
....-<>Atmorneter read1n25(WB,incc) 100 !!
.... Relative humidity 75 ~
'0
50 5
.8
z
25
20
20 May June July

FIG. 67. Weekly catches of adult Anopheles maculipennis with the existing
noonday temperatures in degrees Fahrenheit, the noonday relative humidity
in percentage, and the weekly average of 24-hour records of the white ball
atmometer in cubic centimeters. (After Freeborn.)
temperature or by desiccation of the insect's body. With an increase

of temperature or a decrease in relative humidity the saturation defiCiency
increases and the demand on the insect's moisture content becomes
greater (Fig. 67). The ability of a particular species to retain adsorbed
water in the presence"tpf existing saturation deficiencies undoubtedly
explains the vl\riability of resistance of the different species to desiccation,
according to Freeborn. The length of a life of A. freeborni as well as
other vectors of malaria has important bearing on their ability to transmit
the infection, i.e., if the mosquito dies before the malaria parasite is fully
developed, the latter must also perish.
Internal anatomy of mosquitoes. To be prepared to study the relation
of mosquitoes to such diseases as malaria and filariasis, the student must
be familiar with their internal anatomy. The attention of the student
is called particularly to the excellent treatise on the "Structure and
Biology of Anophelines" by Nuttall and ShipletT (1903).
The alimentary canal of the mosquito is separable into three regions,
the fore-, mid-, and hind-gut, each of which may be arbitrarily sub-
divided into more or less distinct divisions (Fig. 16, p. 52). Thus the
fore-gut consists of the sucking tube of the proboscis, the pharynx, in-
cluding the pumping organ, and the esophagus with its diverticula (three
in number and generally known as food reservoirs). The mid-gut con-
sists of a narrower anterior portion (false proventriculus) and a wider
posterior portion ( stomach) occupying the thorax and much of the
abdomen, and limited posteriorly by the origin of the five Malpighian
tubules, which indicate the beginning of the hind-gut. The hind-gut is
bent on itself several times and consists of the narrow, longer ileum, the
colon, and what is arbitrarily termed the rectum, indicated by the pres-
ence of rectal papillae.
The salivary system consists of two sets of salivary glands (right and
left), three glands to each set. These organs are situated ventrally in the
thorax near the neck. Each set of glands empties into a duct which com-
bines with the opposite one to form the common salivary duct. This
common duct empties its contents into the pharynx through the salivary
receptacle close to the base of the probosciS.
The reproductive system of the female mosquito occupies the pos-
terior portion of the abdomen and comprises a pair of ovaries joined by
a pair of oviducts terminating in the vagina, which opens ventrally in a
depression of the ninth sternite; spermathecae are present (one to three,
depending on the species). The spermathecae of an impregnated female
contain myriads of spermatozoa, and the ovaries when mature occupy
the larger part of the abdomen.
TRIBE MEGARHININI
Characteristics. The members of the tribe Megarhinini occur in tropi-
cal as well as temperate climates and are usually highly colored; they are
day Hiers; both sexes are Hower-feeders and do not suck blood. The
basal half of the proboscis is stout and rigid, while the distal portion is
Hexible, which accounts for the curious hook-like position of the proboscis
when at rest. The palpi vary in length from one-fourth the length of the
proboscis to nearly the same length. The huge larvae are predaceous and
cannibalistic. The mouth parts are particularly adapted for capturing
prey.
The eggs are deposited singly, and Edwards states that most species
MOSQUITOES 185
breed in small confined collections of water, such as may occur in bam-
boo stems, tree holes, pitcher plants, and the like.
Edwards lists 52 species, of which two species are said to occur in
North America, Megarhinus rutilus Coq. and M. septentrionalis Dyar and
Knab. The giant species Megarhinus inornatus Walker was introduced
from New Britain into the Hawaiian Islands 18 for purposes of mosquito
control, without practical result.
TRIBE CULICINI
Characters. All members of the tribe Culicini have the scutellum
trilobed with each lobe bearing bristles, but with areas between lobes
without bristles (Edwards). The abdomen is blunt and completely
clothed with broad scales, which nearly always lie Hat; the pulvilli are
broad and distinct; postspiracular bristles are absent; the larvae have
a prominent Siphon with well-developed pecten (Fig. 65), and usually
numerous hair tufts on the siphon. The eggs are usually deposited in
tight raft-like masses on the surface of the water.
The Tribe Culicini, exclusive of the Aedini, which are separated from
the Culicini for the purposes of this book, includes some 700 species dis-
tributed among more than 20 genera, of which the genus Culex alone
contains nearly 400 known species.
Culex pipiens Linnaeus. Two subspecies are generally recognized,
viz., C. pipiens pall ens Coq. and C. pipiens molestus ForskiU. This
is the common house mosquito or rain-barrel mosquito of many tem-
perate parts of the world. The thorax, abdomen, and proboscis are
brown; the latter is darker toward the tip. The basal white bands on the
abdomen join lateral basal triangular patches. This mosquito, a domestic
species, lays its eggs in rafts on water, in rain barrels, tanks, cisterns,
catch basins, and other small collections of water. Where breeding
places are favorable it may occur in enormous numbers. It invades houses
freely. Because of its vicious bites and high-pitched, tantalizing hum
continued late into the night, it may be a terrific pest. Although greatly
influenced by temperature the life history requires but about 10 days
under warm summer conditions, egg stage 18 to 24 hours, larva about 7
days, and the pupa about 2 days.
Woke 19 fed 38 Culex pipiens Linn. on man, and these mosquitoes de-
posited 29 egg masses, totaling 2,118 eggs, or an average of 73.0 eggs
per mass. At the same time 39 females fed on a canary deposited 22 egg
masses, totaling 4,473 eggs, or an average of 203.3 eggs per mass. Over
twice as many eggs per mass or per milligram of blood ingested were
produced by mosquitoes fed on canary blood as were produced by mos-
quitoes fed on the blood of man.
Culex quinquefasciatus Say [= Culex fatigans (Wied.)] is one of the
most widespread household pest mosquitoes of warmer climates. In

North America it is said to extend to 39° N. latitude. It is an important
vector of Wuchereria bancrofti. It is a severe nocturnal biter. It breeds
in all sorts of artificial water containers, but also in ground pools. It shows
a preference for polluted water. The adult females are medium- to
small-sized, resembling Culex pipiel1s very closely.
Culex tarsalis Coq. (Fig. 68) is an abundant and widespread species
of the semiarid regions of North America; 'however, it occurs as far east
as Illinois and southerly to western Florida. It has been taken at eleva-
tions of 9,000 feet. It is an important vector of the virus of 'western
FIG. 68. Culex tarsalis, an import-

ant vector of the virus of western
equine and St. Louis encephalitis.
(Photograph by R. Craig.)
equine and St. Louis encephalitis. It is a fairly large and robust species;
generally dark brown to black in color; the black abdomen has broad,
segmental, basal bands of yellowish-white scales; each segment of the
ventor with V-shaped marking of black scales, the apex of the V ante-
riorly; femora are black with a dotted white line along both sides, knees
white; tibiae black, also with white line, bases and apices white; tarsi
black; hind tarsi with apical and basal white bands on all segments, last
tarsal segment white; proboscis black, with a sharply outlined white
ring just before the middle. It breeds in all sorts of ground pools, road-
side ditches, pools in and around corrals, artificial containers, etc.
Culiseta incidens (Thomson), like other members of this genus, par-
takes of characters which would place it in either of the tribes Culicini
or Aedini, but for practical purposes, based largely on breeding habits,
the members of this genus are placed in the tribe Culicini. In this genus,
(Culiseta) , which includes seven North American species, the post-
spiracular bristles (Fig. 65) are absent and in the females of at least
MOSQUITOES 187
Pacific coast species the anterior and posterior cross-veins tend to lie in
one line, except in C. morsitans Theo., in which they are well separated,
i.e., by more than half their length. In Culiseta incidens (Thorn.), a
western species (west of the Rocky Mountains), the wings are spotted.
It breeds throughout the year, where temperature permits, in all sorts
of permanent pools and is a common domestic species. It lends itself
particularly well to laboratory experimentation. The life history of this
species is described earlier in this chapter.
Culiseta inornata (Williston) is found throughout the United States
and southern Canada. Its wings are broad and clear, the cross-veins are
scaled, and the very short black palpi have white scales at the tip. It
breeds throughout the year under favorable temperature conditions
wherever pools are present in wooded areas. .
TRIBE AEDINI
Characteristics. Ordinarily the Aedes mosquitoes are included in the

tribe Culicini, but because of their remarkable breeding habits and other
striking characteristics the author has taken the liberty to set apart the
Aedes, of which there are about 600 species, as a separate tribe, the
Aedini. Nearly half of all the species of mosquitoes in North America
belong to the genus Aedes.
Like the Culicini the Aedini have a trilobed scutellum. In most of
the species, as Edwards points out, the claws are toothed in the female,
postspiracular bristles are present, the pulvilli are absent or hair-like, and
the female abdomen tends to be more pointed and the cerci longer than
in other groups. The larvae have short siphons bearing one pair of pos-
teroventral hair tufts, and nearly always a distinct pecten (Fig. 65). The
eggs are deposited singly on the surface of the water, on mud, or even
in situations where there may be little moisture but where submergence
may follow. The females of all species bite, many of them viciously.
Many species are diurnal in biting habits, most of them biting toward
evening.
Salt-marsh mosquitoes. Aedes dorsalis (Meigen), a fierce day biter,
is widely distributed throughout the northern half of the United States,
Canada, Europe, and Asia. In general the body is straw-colored (tan),
varying from almost white to dark brown; the thorax has three longitu-
dinal bright brown stripes; the hind tarsi have white bands at bases and
apices of all segments, the last one wholly white. Although the species
breeds freely and abundantly in fresh water, such as flood water, rice
fields, and drainage from irrigation, it is nevertheless the commonest salt-
marsh mosquito of the Pacific coast north of Monterey. It is here a dis-
tinctly brackish-water breeder, generally breeding in pools reached only
by the monthly "rip" tides. There are thus several monthly broods, the
first appearing as early as March. The eggs are deposited singly, most
of them in the mud along the edge of receding pools; they may remain
unhatched for many months in situations from which water is excluded.
Development after hatching is rapid, and emergence of the adult mos-
quitoes may be within eight or nine days.
Aedes squamiger (Coquillett) is restricted to the Californian sea-
coast from San Francisco Bay to San Diego, breeding in salt-marsh pools
that are fed by fresh water from the winter rains. Like A. dorsalis
(Meig.) it is a fierce day biter, rather worse toward dusk, and travels
great distances. It is known as the gray salt-marsh mosquito. The vesti-
ture is characteristically composed of large scales. The proboscis is uni-
formly black scaled. Its egg-laying habits are similar to those of A. dor-
salis (Meig.); there is, however, but a single brood of A. squamiger in
the San Francisco Bay region coming as early as February.
Aedes taeniorhynchus (Wiedemann) is a typical salt-marsh breeder
distributed along the coastal area of the United States northward to
Connecticut on the east coast and to Santa Barbara on the west coast. It
is the brown salt-marsh mosquito; its proboscis is distinctly white-banded.
It is a fierce day biter, and its egg-laying habits are similar to those of
A. dorsalis (Meig.) and A. squamiger (Coq.). There are monthly broods
throughout the summer. Development is exceedingly rapid, the larval
stage may require but four days, the adults emerging in from 8 to 10
days.
Aedes sollicitans (Walker), the pestiferous salt-marsh mosquito of
the Atlantic coast, breeds from Maine to Florida and thence west along
the Gulf of Mexico to Texas. There are many broods, and in its southern
range breeding may be continuous.
The numbers of larvae appearing in certain pools is almost unbe-
lievable. Smith20 states:
I have found pools so crowded that an estimate of 100 wrigglers in an

area of one square inch was scarcely equal to the fact. Half that number is
a common occurrence. This means over 7,000 in an area of one square foot,
and it needs an area of less than 150 square feet-a pool roughly ten by fifteen
feet-to produce 1,000,000 mosquitoes at one hatching. If these figures seem
incredible they can be easily verified anywhere along shore by anyone who
will put on a pair of gum boots and will hunt out a few breeding places.
Flood-water Aedes. Aedes vexans (Meigen) is a typical Hood-water

mosquito having practically a world-wide holarctic distribution. It is. one
of the fiercest day biters and exceedingly abundant; it is truly a vexa-
tious mosquito. It breeds in greatest numbers along the edges of rivers
subject to overflow, and like other Aedes species lays its eggs along the
MOSQUITOES 189
muddy edges of receding pools, where they may hatch the same season
when water due to intermittent flooding or freshets reaches them, or
they carryover. There may thus be several broods where flooding occurs
as a result of melting mountain snows or thunderstorms, or there may
be only one brood where there is a single spring flood. The species is a
rapid breeder and migrates many miles. It varies in color from brown
to gray; the tarsi are basally narrowly banded; the wings are uniformly
brown.
Aedes dorsalis (Meigen), already referred to as a salt-marsh breeder,
is also a prolific breeder in open flood-water pools, particularly in irri-
gated pastures after flooding and in drainage pools due to excess water.
Where early flood waters occur along wooded river courses the single-
brooded Aedes vexans (Meig.), dominates, but as the season advances
Aedes dorsalis (Meig.), a many-brooded species, supersedes it in these
areas, but breeds particularly in open pastures where the temperature of
the water is high. In water of a temperature of 99.5° F we found enor-
mous numbers of larvae completing their cycle in about six days.
Tree-hole mosquitoes. Although the habit of breeding in water-holding
tree holes occurs in various species belonging to genera other than Aedes,
e.g., Anopheles barberi Coquillett, there are a number of typical tree-
hole breeders in the Aedini, notably Aedes varipalpus (Coq.) a Pacific
coast species; Aedes triseriatus (Say) of the eastern United States; Aedes
luteocephalus Newstead, Ethiopian; Aedes simpsoni Theobald, Ethiopian,
Aedes seoulensis Yamada, Chinese; and others.
Aedes varipalpus (Coquillett) has bright white markings on the legs
at both bases and apices of the tarsal segments and many white or silvery
scales distributed over the body so as to give the vestiture a silver mottled
appearance. It is one of our smallest mosquitoes but a fierce biter. This
Pacific coast species deposits its eggs on the sides of tree holes, notably
holes in live oaks (Quercus agrifolia), also California laurel (Umbellu-
laria californica) and valley oaks (Quercus lobata). Freeborn 21 states that
the eggs "hatch whenever they are wet by the rising waters. There is
some evidence that the eggs may drop off after a period of desiccation
or, as an alternative method, the larvae may hatch without the interven-
tion of actual wetting and fall into the water below. The straw-colored
larvae with bright brown heads and enormously developed gills swim
about their secluded medium with snake-like movements, but spend
most of their time with their heads and thoraces buried in the silty
deposit at the bottom of the tree holes. . . . The developmental period
is extremely long, lasting from one to seven months. Although there is
an intermingling of the broods, there are two pronounced peaks, one in
the early summer and another in the fall. The fall adults deposit eggs
which produce the larvae that overwinter."
Feng,n in his report on the tree-hole mosquitoes of Peiping, China,
states that the pH of water in which all species bred varied from 7.8 to
8.4. He also states that there was no specific relationship between the
species of the tree and the species of the mosquitoes, several species
being frequently found breeding in the hole of the same tree.
Boreal Aedes or snow mosquitoes. An interesting group of Aedes con-
sists of the so-called snow mosquitoes which appear in the early spring
in the high mountains and northern ranges of distribution, breeding in
the pools left by the melting snow (Fig. 69) . These Aedes have but one
generation and appear in enormous swarms in the higher elevations and
FIG. 69. Typical breeding place for snow mosquitoes on floor of Yosemite
Valley. (Photograph by H. F. Gray.)
northern ranges much to the dismay of the huntsman and alpine traveler.
These mosquitoes have been collected in many localities in the Sierra
Nevada by the writer. The following quotations are taken from an inter-
esting account of the mosquitoes by Dyar. 23
At an altitude of 6,000 feet, pupae were abundant May 25 and by the

first week in June the breeding was complete; even the pools that still con-
tained water or had only just thawed out were empty. Adults appeared by
the first of June, and by the 15th the woods were filled with them in all
directions.
Speaking especially of the Fallen Leaf Lake region (vicinity of Lake
Tahoe) a region in the heart of the Sierras to the north of the high peaks
and on the eastern side of the divide, A. (communis) tahoensis Dyar is the
commonest and earliest species, found everywhere, both in the hills and the
pines in level country. It breeds in the earliest pools of clear water held in
MOSQUITOES 191
rocky land, its home being in the mountains, but it soon spreads everywhere.
It was common in the early pools at the head of Fallen Leaf Lake, being the
only species present. It also bred in wave-pools at the lower end of the lake.
Dispersal of the adults was in general downward, they being abundant in
the pines at Tallac on June 17, though no breeding places were near. A. cata-
phylla Dyar is less abundant and less widely dispersed at Fallen Leaf. It was
commonest at the foot of the trail to Angora Lakes at the head of the lake,
rare at the outlet of the lake and absent at Tallac on Lake Tahoe. A. hexodon-
tus Dyar breeds in early pools, but especially those of a marshy character,
larvae being taken from hoofprints of cattle in the edge of a marsh. The adults
were well distributed and toward the end of June replaced A. (communis)
tahoensis Dyar as the dominant species. A. ventrovittis Dyar is a rare species,
taken only at one place near the outlet of Fallen Leaf Lake and then in small
numbers. It is presumably a marsh breeder, though the larvae were not found.
A. palustris breeds in open grassy marshes, not in large numbers. Dispersal
was general, adults being taken everywhere, although seldom commonly. A.
increpitus Dyar is the slowest breeder of any of the early species, the larvae
lingering after all the others are gone, frequently in the same pools. They
were abundant at the outlet of Fallen Leaf Lake with a downward dispersal,
the adults being common at Tallac, about 2 miles from the breeding places,
while only found a quarter of a mile up the lake and many days later.
The seasonal appearance of these mosquitoes varies with the altitude in
the ratio of about a month in time to 1,000 feet of elevation. At Yosemite,
at about 5,000 feet, all the species were about a month earlier than at Lake
Tahoe, at 6,000 feet, while at Summit, at 7,000 feet, they were still another
month later, larvae and pupae of tahoensis and hexodontus being taken there
on July 2, 1916, about the same stage that they were taken at Fallen Leaf on
June 1, 1916.
At an elevation of about 10,000 feet the author encountered a veri-
table plague of Aedes ventrovittis Dyar and A. communis tahoensis Dyar
along the shores of Young Lakes in the Sierra Nevada 1fountains July
22 to 27, 1936. Larvae and pupae were still present in small pools of
snow water along the shores of the lakes. Apparently only certain pools
were infested.
In northern Alaska the spring thaw is followed by a sudden emer-
gence of swarms of mosquitoes. Like other boreal mosquitoes they over-
winter in the egg stage and hatch as soon as the ice thaws from around
the eggs in the spring. There is but one generation a year. Jachowski
and Schultz,24,25 who studied Aedes punctor (Kirby), A. communis (De
Geer ), and A. nearcticus Dyar, classify the sites from which they col-
lected larvae as grassy sloughs, mossy pools ("tundra pools"), frost'
ditches, and willow-alder pools. The eggs hatch almost immediately after
the spring thaw, and if a freeze occurs after this, the larvae will freeze
into the ice and thaw out again when it warms up, The larval period was
observed to be about 28 days and the pupal period from 3 to 5 days.
During chilly night hours mosquitoes were observed to gather in large
numbers about the open doors of heated quarters or hovered in the warm
draft from chimneys. Blood meals, if necessary, are readily available
from herds of caribou, large populations of rodents, and nesting ducks,
geese, and other birds.
Of 18 species of mosquitoes found in central Alaska according to
Jenkins 26 the following species are the most abundant: Aedes punctor
(Kirby), A. communis (DeGeer), A. pionips Dyar, Culiseta alaskaensis
FIG. 70. Aedes aegypti, an important

vector of the viruses of yellow fever and
dengue. (Adapted after Soper, Wilson,
et al.)
~
AEDES (STEGOMYIA) AEGYPTI
( Lud. ), and C. impatiens (Walk.); A. fiavescens (M uell. ), locally a bun-

dant in the coastal marshes.
Aedes aegypti (Linn.) (Fig. 70), the yellow fever mosquito, will long
be known under the name of Stegomyia fasciata (Fabr.) although it
was known for several years under the names of Aedes calopus (Meig.)
and Aedes argenteus (Poiret). It is not only the most important vector of
yellow fever but likewise of dengue fever. This species is widely distrib-
uted within the limits of 40° Nand 40° S latitude, but it is highly
susceptible to temperature variations. According to Hindle 27 it soon dies
in the open air at a temperature of 7° to 8° C, succumbing in a few
seconds to an exposure of 0° C, and 37° C is rapidly fatal. Furthermore,
it does Bot thrive in dry hot climates. The adult insect (Fig. 70) is
beautifully marked with silvery-white or yellowish-white bands and
stripes on a nearly black background, whence the name "tiger mosquito."
MOSQUITOES 193
It has a "lyre-like" pattern dorsally on its thorax, i.e., two outer curved
yellowish-white lines and two median parallel lines. The legs are con-
spicuously banded, and the last joint of the hind leg is entirely white.
The head is covered with broad Hat scales with only a single row of
upright forked scales.
The yellow fever mosquito is a typical domestic species seldom found
far from man's habitations. Many observers believe it to be a day-Hying
and day-feeding species, but this habit apparently is restricted to the
younger individuals up to six or seven days after emergence, or rather
until a meal of blood is secured, when the insect becomes nocturnal.
Howard observes that "it prefers the blood of white races to that of dark
races, and attacks young, vigorous persons of fine skin and good color
in preference to anemic or aged people."
The eggs of the yellow fever mosquito are deposited singly on the
surface of the water, usually in containers at or near the waterline; they
are dark in color, and each egg is surrounded by air cells (Fig. 65).
Comparatively few eggs are deposited at one laying, and while there
may be two layings, pOSSibly more, the total averages about 140 (144
according to Woke) when fed on man. Woke 28 found that this species
produced greater numbers of eggs when fed on the frog or turtle.
Unlike the eggs of most species these can withstand desiccation to
a very marked degree, some authors declaring that this is possible for
several months. Ordinarily the eggs hatch in about 48 hours.
The larvae are quite robust, the breathing siphon is comparatively
short and heavy and black (Fig. 65), and thell: position in the water is
almost vertical, considerably more so than that of other culicine species.
The larval stage is ordinarily passed in about 9 or 10 days under average
conditions and in from 5 to 7 days under wanner conditions.
The pupae have broadly triangular breathing trumpets. Only about
36 hours is spent in the pupal stage.
According to Howard the shortest period of development from egg
to imago observed by Reed and Carroll in Cuba was nine and one-half
days, i.e., egg stage, two days; larval stage, six days; pupal stage, 36
hours. From this very short period the time ranges from 11 to 18 days
according to the same author.
The yellow fever mosquito breeds by preference in artificial con-
tainers of rain water. (It is known, however, at times to breed naturally
in brackish water.) Rain-water barrels, tanks, cisterns, tin cans, urns,
etc., provide suitable breeding places (wooden containers seem to be
preferred); water collected among the leaves of certain members of the
Agave family, also water collected in banana palms may produce many
mosquitoes.
Woke has shown that Aedes aegypti (Linn.) fed on frog blood and
turtle blood produced viable eggs, and that the larvae developed nor-
mally and produced normal adults.
Although Aedes aegypti (Linn.) is called the yellow fever mosquito
and is undoubtedly the most important vector of this disease under
natural conditions because of its domestic breeding habits, there are
nevertheless a dozen other species which are able to transmit the disease
(see p. 239).
Other genera. The genus Psorophora, which includes only nine North
American species, is distinguished by the presence of both prespiracular
and postspiracular bristles and by the second marginal cell of the wing,
which is more than half as long as its petiole. The larvae of some species
are predaceous, feeding on other mosquito larvae and other small aquatic
animals in temporary ground pools.
Psorophora columbiae (Dyar and Knab) is Widely distributed in the
eastern United States but ranges from parts of South America through
Cuba and Mexico to Canada. It is' strikingly speckled in appearance
and is a fierce biter. In 1932 this species is reported to have caused great
loss to livestock in the Everglades section of Florida. The United States
Insect Pest Survey Bulletin (Vol. 12, No. 10, p. 428) describes the
plague: " ... by evening of that day the buzzing was as loud as that
of a swarm of bees. During the night livestock could be heard running
and thrashing in the underbrush, and on the morning of September 6,
dead animals were found throughout the section. The recorded mortality
was 80 head of cattle, 3 horses, 1 mule, 67 hogs, 20 chickens, and 2 dogs.
Post-mortem examinations showed no mosquitoes in the respiratory ap-
paratus, indicating that the animals died either from loss of blood, nerv-
ous exhaustion, or the effects of some toxin." The milk supply was also
greatly reduced during the four days of the infestation.
The genus Mansonia is characterized in large measure by the scales
of the wings, which are very broad by comparison with those of other
species of mosquitoes. The larvae have the air tube sharply pointed, en-
abling them to pierce the stems or roots of aquatic plants from which
they obtain air and to which they remain attached (submerged) through-
out larval development.
Mansonia perturbans Walker is a small species (5 mm) which is
widely distributed throughout North America. It has severe biting habits
and evidently travels some distance from its breeding place. The biology
of this and other species of the genus is fascinating indeed.
The genus Orthopodomyia includes only two species from North
America. The adults are described by Matheson as "rather gaily orna-
mented and easily recognized by their coloring." The larvae are found
in tree holes, also in broken bamboo and leafaxils of certain plants.
Orthopodomyia signifera (Coquillett) extends along the eastern sea-
MOSQUITOES 195
board of the United States and westerly from Florida into southern
California, thence northerly to near SacramentO.
TRIBE ANOPHELINI
Characteristics. The following characters are generally employed to
characterize the tribe Anophelini: palpi of both sexes are usually about
ANTENNA
EYE---f---"j..
\""'... .
THORAX-i
i
SCUTE LLUM --t-:-;&.;;~~"':
POST, :, '."""1!1iJmn""",,""';;;;_ 11I'\'1" '"'
SCUTELLUM
HALTER
ABDOMENl
.- ____ I,
ANOPHELES MACULIPENNIS
MEIGEN
~
FIG. 71. Dorsal view of Anopheles mosquito (female) showing certain

structural details useful in classification. (Modified after Essig in College
Entomology, The Macmillan Company, New York, 1942.)
as long as the proboscis; the scutellum (Figs. 65 and 71) is evenly
rounded (except in Chagasia where a slightly trilobed condition occurs);
mandibles and maxillae of the females are well developed and toothed;
legs are very long and slender, there are no distinct tibial bristles and
no pulvilli; the abdomen is without scales, or at least with the sternites

largely bare (Edwards); the wings usually have distinct markings.
The tribe Anophelini has been divided into numerous genera such
as Myzorhynchus, Arribalzagia, Argyritarsis, Neomyzomyia, Myzomyia,
and more than thirty others. Edwards, as well as other culicidologists,
reduced the number of genera to three: Chagasia, with the scutellum
slightly trilobed; Bironella, with scutellum evenly rounded, wing with
stem of median (M) fork wavy; and Anopheles, with scutellum evenly
rounded, wing with stem of median (M) fork straight. There are three
species of Chagasia, all of tropical America. The genus Bironella in-
cludes two species, both of New Guinea. The genus Anopheles includes
FIG. 72. Anopheles freeborni in the act of sucking blood from the author's halld.
over 300 species and subspecies, (Russell et al., 1943, loc. cit) , of which
about 90 occur in the Americas, 14 in North America.
The common species rest with the proboscis, head, and abdomen
nearly in a straight line and when resting have the appearance of a splin-
ter lifted at an angle from a given surface (Fig. 72). In exceptional
cases, as in Anopheles culicifacies Giles of India, the resting position is
Culex-like. Hoffman 29 states that A. grabhamii Theobald rests with its
body almost at a right angle to the vertical surface. A. hyrcanus sinensis
Wied. takes a similar position . The hum of anophelines is distinctly Iow-
pitched and almost inaudible unless they are close to the ear or in a
bottle. Most of our common anophelines are not strong fliers and usually
take to cover even in a moderate breeze. In California, Anopheles free-
bomi Aitken previously recorded here as Anoplleles maculipennis Meigen ,
MOSQUITOES 197
engages in an annual dispersal Hight of overwintering females about
mid-February, during which time the mosquitoes may invade much ter-
ritory, traveling more than 25 miles. During this Hight the first eggs are
deposited. In the main this Hight favors the spread of the species. While
on the Hight the mosquitoes bite by day even in broad sunlight. Free-
borni also engages in a prehibernation Hight during late October and
November. The males having died soon after mating, the overwintering
forms are females only.
Although some individuals of a few species may overwinter in the
larval stage buried in mud and debris at the bottom of certain pools,
the usual method of overwintering is in the adult (females only) stage.
Mating and oviposition. Fertilization of the females takes place almost
immediately upon emergence. The males emerge first and may be seen
dancing over or near the breeding places in small swarms apparently
awaiting the appearance of the females; when these dart into the danc-
ing swarm mating occurs. This type of mating requiring wide spaces is
known as eurygamous, while those forms such as Anopheles sacharovi
Favr. (= elutus Edwards) which mate in confinement in a small space
are known as stenogamous. Overwintering females are fertilized by the
last brood of males during the autumn, and the eggs are deposited soon
after the spring dispersal Hight. In certain localities at least, there is a
period when the species exists only in the larval stage, all the adults
having died after egg deposition. There is probably only a single laying
at this time.
Under laboratory conditions the great majority of eggs are deposited
between sunset and eleven o'clock in the evening (vicinity of Chico,
California, May to August), although our records show layings later at
night and a few during late afternoons of highly overcast humid days.
The average number of eggs deposited by Anopheles freeborni Aitken is
slightly in excess of 200, with 385 as the maximum for one laying. 30 The
same average and somewhat smaller total number for one laying was
observed for Anopheles punctipennis (Say), a maximum of 321, with an
average of 203.31 During 1937 the largest number in one batch for this
species was 352, observed by Aitken. One A. punctipennis (Say) female
deposited 500 eggs in our laboratory in four layings from Mar. 2 to 22,
1938. For Anopheles pseudopunctipennis franciscanus McCracken, for-
merly recorded in California as Anopheles pseudopunctipennis Theobold,
a maximum of 283 was observed, with an average of 151. At least three
batches of eggs may be laid during the lifetime of a female. It is of in-
terest to note that in one of our observations a female A. freeborni de-
posited 174 eggs in 19 minutes, an egg every six to seven seconds with
intervening periods of rest. During the entire operation the female rest-
ing on the surface of the water remained motionless except for the monot-
onous jerking of the abdomen when the egg was released. The eggs fell
in a heap beneath the insect, pearly white in color, toppling over and
forming geometrical patterns and becomin~ deep brownish black in
about 45 minutes.
Egg characters. The characters of anopheline eggs used in classifica-
tion are: presence or absence of floats, position and length of the float,
presence or absence of frill, and pattern. (Fig. 73) Christophers and
Barraud 32 classify anopheline eggs as of four types:
1. Eggs probably of primitive type with full ..float surrounding egg,
2. Eggs with terminal frill (pseudopunctipem1is franciscanus of Herms and
Freeborn) ,
3. Whale-back eggs with floats separated frorrl dorsal surface,
4. Various types of boat-like eggs with floats touching margin of dorsal
surface. '
The egg of the Californian A. freeborni Aitken (Fig. 72) is fusiform,
sllgntly rounaea at eacn ena ana tapeimg 'to 'tIn:: exteIn "&.rcl't UIft '6
slightly broader than the other (Herms and ,Frost). 31 The upper surface
is flattened with a slight longitudinal concavity, while the lower surface
is broadly convex, the convexity becoming more pronounced at the broad
end of the egg. The upper surface is granular, bordered by a laterally
striated frill 161l in width, except at the flO;,1-ts, while the lower surface
shows, under proper light, a silvery reticuilltion. Medianly placed are
two roughly oval lateral floats, each divided in a majority of cases into
12 scalloped compartments. The larger part of the area covered by these
floats is on the lateral faces of the egg, but tVey project dorsally over the
margins, which are described as "gunwales" rather aptly by one author
who likens the egg to a boat. The eggs rallge in length from 5961l to
656p.. The floats vary in length from 122p. to 2;:;.4p..
Falleroni, Martini, Hackett, and Missiroli (see Hackett33 and Hackett
and Lewis34) have shown that Anopheles m{lculipennis Meigen actually
comprises seven races in Europe, which call be distinguished only by
egg patterns, namely var. melanoon Hackett, var. messeae Falleroni, var.
typicus Meigen, var. atroparvus van Thiel, vaf· labranchiae Falleroni, and
var. subalpinus Hackett and Lewis; another, var. elutus (= A. sacharovi
Favr.) has solid leaden-colored eggs (Fig. 7:»·
The eggs of A. punctipennis (Say) resemble those of the Californian
A. freeborni Aitken with these exceptions: ill the punctipennis form the
"frill" extends along the margins of the egg without interruption at the
site of the floats which are located on the upper portion of the ventral
surface, and extending farther along the sides of the egg are a greater
number of compartments, ranging from 16 to 22, which do not converge
in fan-wise fashion as in freeborni.
Anopheles punctipennis (Say)
A. pseudopunci'ipennis Theob. (usual form California)
A. pseudopunctipennis Theol:>. (rOTe form California)
A. maculipennis Meig. (California)
A. maculipennis var. maculipennis Meig. vor. labranchioe Follerani

(typicus)
var. messeae Falleroni vor. atroparvu5 van Thiel
vor. melanoon Hackett A . elutus Edwards
FIG. 73. Eggs of certain anopheline mosquitoes, including racial forms.

California A. maculipennis Meig. should read A. freeborni Aitken, and A.
fJseudopunctipennis should read A. pseudopunctipennis franciscanus Mc-
Cracken. (European races of A. maculipennis redrawn after Hackett.)
199
The eggs of Anopheles pseudopunctipennis franciscanus McCracken

occur in at least two different forms in California. Herms and Freeborn 30
described the eggs as having the upper surface nearly Hat, showing little
concavity longitudinally although the lower surface shows a marked
convexity. Both ends of the egg are rounded, one being considerably
broader than the other. The Hoats are represented by a fusiform closely
appressed area, approximately 270p. long, lying on the dorsal side of the
blunt end. This area is divided medianly by a line which is assumed to
be the line of contact of the two Hoats that have been forced up from
FIG. 74. Typical breeding place of California Anopheles freeborni in sunlit

weed-grown ditch with slowly moving water.
the sides. Lateral lines mark off each longitudinal half of the area into 12
sections representing the original compartments of the lateral Hoats. This
area is so appressed that its position is not distinguishable from a lateral
view. Near the narrow end of the egg the membranous covering Hares out
from the body of the egg to form a translucent, striated collar which
eompletely encircles the end, with the exception of a triangular incision
down the dorsal median line in a manner which reminds one of an over-
sized dress collar (Fig. 72). The egg hangs at an angle in the water,
supported by surface tension on the "collar." The eggs ranged in length
t-.fOSQUITOES 20]
from 512J.L to 528J.L. Rozeboom"" states that the eggs of A. pseucloJJlIllcti-
pennis Theobald in Panama resemble the eggs of the Califomia species
as described by Herms and Freeborn , except that ill the Panamaniall
form the floats are large and have mallY Roat ridges; the l:ollar-Iike frill
being identical. The Panamanian eggs ranged in length from 480 IJ. to
573p., the mean being 520p..
A second form of A. pseudopul1ctipel1nis egg, probably var. boyd;
Vargas was described by Herms and Frost (lac. cit.) as follows: floats are
not only present but completely encircle the egg, which lies flat upon
FIC. 75. Breeding place for Californian Anopheles treebomi in overflow

from river. Larvae particularly along margins and in hoofprints at edge.
(Photograph by L. L. Williams, Jr.)
the surface of the water with the floats extended in nearly every instance
(Fig. 73). The floats average from 45 to 47 in number, and from 15p. to
25J.L in width, being wider along the sides of the egg and narrowed at
the ends. The length of the eggs over all ranged from 510J.L to 544J.L.
Breeding habits. The breeding habits of anophelines differ consider-
ably for even very closely related species, e.g., the American Anopheles
freeborni Aitken and A. quadrimaculatus Say, both four-spotted anophe-
lines separable with accuracy as adults only on differences in male ter-
minalia, have widely different breeding requirements, the former, at
least in California, breeding largely in open sunlit shallow seepage
water (Figs. 74 and 75) and the latter in impounded water with floating
debris and aquatic vegetation. The European races of A. maculipennis
Meigen already referred to emphasize the need of accurate knowledge

as to breeding habits.
The following example illustrates the very great importance of know-
ing the breeding habits in the conduct of malaria control operations.
Williams 3G points out that in the Federated Malay States Anopheles
umbrosus Theobald is the vector of malaria in the coastal plain, breed-
_._. } I.d all*,~ -~-'-

i-,
-.,
I I
FIG.76. Showing life history of a
mosquito, Anoph eles maclIlipellllis.
ing in practically stagnant water densely shaded by mangrove. Its pro-

duction is controlled, as Williams points out, by clearing the swamps
and letting in the brilliant sunshine, or by cutting ditches and confining
the water to definite channels. The same type of work when practiced on
high inland plateaus increases the malaria rate, because here the vector
is Anopheles maculatus Theobald, which prefers the quiet edges of
MOSQUITOES 203
trickling streams in the open sunshine. Anopheles minimus Theobald,

the principal vector of malaria in the Philippines, breeds in small flowing
streams in the foothills (Russell 37 ). Several species of Anopheles (though
unimportant as vectors) are tree-hole breeders, viz.: A. plumbeus
Stephens (European) and A. barberi Coq. (American). A. bellator D.
& K., an important Caribbean vector, breeds in collections of water
among bromeliads.
An unusual situation is reported for Anopheles sergenti (Theobald),
a north African and Palestinian species, the larvae of which inhabit small
pools and springs among stones at the edge of the lake (Tiberias). The
larvae are often under the stones and not easily found (Buxton 3S ).
Life history of anophelines. Although there is much variation in the
life histories of the species of Anopheles mosquitoes (Fig. 76) as well
as considerable variation within the species due to temperature and
other factors, the length of time required for development from egg to
adult is generally longer than in other genera, except in genera where
the egg stage may be greatly prolonged, as in Aedes.
Incubation period. In a series of tests in which about 20,000 eggs of
Anopheles freeborni Aitken 39 were used the incubation period at room
temperature of 70° F + 3° was about 72 hours. Under field conditions
the incubation period ranged from two to four days, with an average of
2.5 days, under which conditions the incubation period of Anopheles
punctipennis (Say) ranged from two to six days with an average of 3.2
days.
The eggs of A. freeborni removed from the water and dried at tem-
peratures of 74° F and 65° F remained viable after a period of desic-
cation not over 72 hours. No hatching was obtained from eggs of A.
punctipennis (Say) after 24 hours' desiccation.
Larval period. Hatching generally took place during the evening and
night in the experiments cited. With yeast as food in distilled water at
pH of 6.6 to 7.6 the larvae of A. freeborni reached the pupal stage in 15
to 16 days.
Pupal stage. The pupal stage requires about three days. Thus the
entire life history from egg to adult in A. freeborni under experimental
conditions requires about 21 to 22 days; the same is true for A. puncti-
pennis (Say) and A. punctipennis franciscanus McC. Under field condi-
tions this period may be considerably prolonged. Adult mosquitoes
reared in the laboratory did not begin oviposition until 13 to 15 days
after receiving a blood meal.
The life cycle of Anopheles albimanus Wiedemann, the important
vector of malaria of the Panama Canal Zone, has been carefully studied
by Rozeboom. 40 With room temperature between 27° and 32° C and
water temperature for larvae from 21° to 27° C, and eggs and pupae
at 27° to 30° C, the entire cycle (egg to adult) required from 18 to 24
days, an average of three weeks. A period of seven days, or a little over,
was necessary for the development of the ovaries, an average of 435
eggs being deposited; the incubation period was 40 to 48 hours; the
larval stage required from 6 to 22 days, usually 8 to 13 days in hay in-
fusion water; the pupal stage took 30 to 33 hours; the longest observed
adult life of a female was 31 days, and of a male, 27 days.
Anopheles quadrimaculatus Say occurs in the United States from
Mexico to Canada throughout the Mississippi Valley and east to the
Atlantic. It is the chief vector of malaria in the eastern, central, and
southern United States. The wings have four distinct spots. According
to Williams (loc. cit.):
It breeds almost wholly in still water that is relatively clean. It requires

some sunshine, never being found in dense shade. However, it requires some
darkness, for it is never found in waters which are wholly unshaded, unless
they have a type of flotage which casts narrow strips of shade where the larvae
may lie during a portion of the daylight hours. . . . An ideal breeding place
for A. quadrimaculatus Say is in freshly impounded water which floods a basin
containing underbrush and which is sparsely covered with trees. Such a body
of water quickly gathers flotage of dead and dying land vegetation, twigs and
leaves, among which algae soan appear. Such flotage not only offers the req-
uisite amount of shade. but an abundant food supply. Such an impounding
will not acquire a large quantity of natural enemies, such as top minnows and
aquatic insects, for a number of years and seldom acquires enough entirely to
prevent production of the mosqUito.
The normal detritus passing down a narrow stream will clog the interstices
of a fallen tree or branch and create a dam. These natural impounded waters
are excellent breeding places for quadrimaculatus. Swamps covered by a
growth of virgin timber, on the other hand, are not good breeding places.
Such swamps are almost invariably covered with such a dense timber growth
that sunlight can reach the surface of the water only in those small areas
where an opening has been made by the fall of a dead tree. Swamps of this
description have a small seeding of quadrimaculatus, but not enough to
propagate malaria. When the lumberman enters, cutting out the large trees,
leaving the small ones, the branches and the tree tops, he changes a safe
water surface into one almost ideal for quadrimaculatus production. He has
let in the sunshine without removing all of the shade, and he has left behind
waste which not only creates flne flotage, but large portions of which tend to
clog the channel which traverses the average swamp, thus making a series of
ponds.
The brood peaks of this species in southwestern Georgia according

to Boyd41 are from 20 to 30 days apart, and there are from 8 to 10
annual broods, the first appearing from 20 to 30 days after the last
frost, and the last brood, the tenth, if there is favorable weather, in
MOSQUITOES 205
December. January and February, he states, are the only months when
no broods emerge. An excellent description of the techniques used in
large-scale rearing of Anopheles quadrimaculatus Say is given by Green-
wald, Cochran, and Scharff. 42
Biting habits. In determining the relative importance of various
species of mosquitoes as vectors of diseases, a knowledge of their biting
habits is essential. They may be preponderantly zoophilous, feeding on
animals other than man, or preponderantly anthropophilous i.e., feeding
on man; they are seldom exclusively either. That a species is a strong
vector of any given infection is probably not always due to an innate ca-
pacity but more probably due to a larger degree of domesticity. A close
and constant contact between the vectors of a disease and the human
victims is essential to extensive transmission of the disease. There are
some species, such as Anopheles minimus flavirostris ( Ludlow), an
important vector of malaria in the Philippine Islands, and A. albimanus
Wied., a vector in the Caribbean area, which do not remain indoors
during the day after feeding indoors at night, an important factor in
making malaria surveys.
It is often pointed out that the fact that certain species of mosquitoes
predominate indoors or are the sale entrants must be weighed against
the production .and proximity of these and other species under com"
parison. Carter, Le Prince, and Griffitts 43 have observed that Anopheles
quadrimaculatus Say is much more often found in residences than A.
punctipennis (Say). Mayne (Mitzmain)~4 in a table comparing the col-
lections of A. punctipennis (Say) and A. quadrimaculatus Say, totaling
1,377 specimens, compares the numbers of each species taken inside
dwellings, under dwellings, in privies, vacated buildings, cattle sheds,
horse and mule sheds, fowl roosts, and wagon and tool sheds. In com-
menting on his results he states:
It is indicated that in the three sources of direct human influence, namely,
inside dwellings, under dwellings, and in privies, the last produced the greatest
number of specimens of A. punctipennis. This species comprised less than
one-third (30 per cent) of the catch in houses, while under dwellings 62 per
cent of the mosquitoes collected proved to be A. punctipennis. . . . It has
been observed by service officers, conducting malarial surveys, that although
A. punctipenis rarely bites while inside a dwelling, it is found to attack persons
seated on the porch or gallery of the house, after which these mosquitoes
seek rest, presumably under the house in preference to the interior.
During the summer of 1920, from May 12 to July 13, daily collections
of mosquitoes were made by the author and his associates at Vina
(Northern Sacramento Valley, California), one series being indoors and
the other outdoors. The indoor collections were made regularly in the
same buildings-a cowshed, washhouse, showerbath, storehouse, and
dwelling, whereas the outdoor collections were made under a short
wooden bridge within 10 to 12 feet of an aggregation of shacks occupied
by Chinese and Negroes. The indoor series taken in an area where control
measures were in progress does not represent a large number of mos-
quitoes, but the fact remains that of 77 anophelines collected, 50 were
Anopheles freeborni Aitken (including only one male) and 27 were
A. punctipennis (Say) (including two males), or practically twice as
many A. freeborni. On the other hand, the outdoor series represented
a total of 343 anophelines of which 102 were A. freeborni (42 males
and 60 females) and 241 were A. punctipennis (130 males and III fe-
males) or something over twice as many A. punctipennis.
These collections bear out very well the general observations that
A B c
FIG. 77. Californian Anopheles: (A) A. freeborni, (B) A. punctipennis, and
(C) A. pseudopunctipennis franciscanus.
A. freeborni Aitken like A. quadrimaculatus Say readily invades houses
and is consequently of greater importance as a malaria vector, while
A. punctipennis (Say) is chiefly an outdoor biter, porch biter, etc., and
consequently probably of less importance as a vector. During the entire
period of two months the well-screened cottage which was occupied by
our party was not invaded a single time by A. punctipennis (Say),
while A. freeborni Aitken was a common visitor. Anopheles pseudo-
punctipennis franciscanus McC. is considered to be a field mosquito in
California and is apparently of no consequence as a vector. Methods used
to study the feeding habits of mosquitoes are well described by Reeves
and Hammon.' 5 In this study they used the precipitin test in which
mosquital blood smears were tested with the antisera of domestic rabbits
against the sera of the horse, cow, sheep, dog, man, and chicken. In
Mexico, Central America, and southerly, Anopheles pseudopunctipennis
pseudopunctipennis Theo. invades human habitations and is an impor-
tant malaria vector.
Some North American anophelines. Anopheles freeborni Aitken (Fig.
77A), Anopheles occidentalis (Dyar and Knab), and Anopheles aztecus
MOSQUITOES 207
Hoffman constitute for the present American maculipennis group ac-
cording to Aitken. 46 Freeborni is referred to as the brown, unicolorous
form of the irrigated western regions, west of the Rocky Mountains,
extending northerly into British Columbia and southerly into parts of
Arizona and New Mexico, the western tip of Texas, and Baja California
(Mexico). Occidentalis is referred to as the "silver" or "bronze-tipped"
melanic form of the cool western seacoasts, into Alaska, and across the
continent through southern Canada and the northern areas of the United
States to Connecticut; Vargas contends that the above-indicated easterly
range of occidentalis is taken over by earli. Aztecus is the large, long-
winged, linear scaled species of the Mexican plateau, at elevations of
5,000 to 7,000 feet and over.
Anopheles quadrimaculatus Say is the common malaria vector of the
eastern United States, extending from southern New Hampshire and
southern Ontario westerly to Minnesota and south to Vera Cruz, Mexico.
It is particularly abundant in the southeastern United States. The wing
scales are all dark, some forming four blackish spots; palpi and hind
tarsi are black; femora have small white spots at tip, "knee spots."
Anopheles punctipennis (Say) (Fig 77B) is said to be the most
widespread anopheline in North America, ranging from southern Can-
ada to the Mexican plateau. The wings have black and yellow scales,
the latter forming two spots on the costal margin, one of which is long
and situated beyond the middle, the second smaller and near the apex,
giving a mottled appearance to the wings. The proboscis is black, and
the palpi are unbanded. The general appearance of the body is dark
brown. This species breeds in clear, cool, shaded pools. The females
seldom enter dwellings but invade unscreened porches and bite in the
open.
Anopheles pseudopunctipennis Theobald is a widespread plainS and
highland species, extending from Argentina along the Andean region of
South America, through Central America, Mexico, and the south central
United States. It resembles A. punctipennis (Say) somewhat, but is
easily separated by the white banding of the palpi. The tip of each
palpus is white. It breeds primarily in sunlit pools along the courses of
receding streams, the larvae being exceedingly abundant in the mats
of green algae.
Anopheles pseudopunctipennis franciscanus McCracken (Fig. 77C),
together with the var. boydi Vargas, is very abundant in many parts of
California where it was previously recorded as Anopheles pseudopuncti-
pennis Theobald; it also occurs in Nevada, Utah, Arizona, New Mexico,
western Texas, and Baja California. Its breeding habits resemble those
of pseudopunctipennis. It differs from the latter in that the tips of the
palpi are black. It rarely enters human habitations.
208 MEDICAL ENTOI-.I0LOGY
Anopheles crucians Wiedemann, Anopheles bradleyi King, and

Anopheles georgianus King resemble each other very closely and con-
stitute the "crucians" complex. The palpi of all three species are banded
with white, and the last segment is entirely white. Separation of the
species can be made only in the larval stage. Crucians has been found
from Massachusetts southwesterly to Kansas, Oklahoma, and Texas. It
breeds in ponds, lake margins, swamps, and pools of both intermittent
and permanent character; it breeds extensively in acid waters as in the
cypress swamps of Florida and Georgia (Matheson). The females bite
freely during the day. Bradleyi occurs along the Atlantic and Gulf coasts
from Maryland to Vera Cruz, Mexico. It normally breeds in saline waters.
Georgianus occurs widely in fresh water in the Gulf states. It can be
distinguished from the other two members of the complex with dif-
ficulty.
Anopheles barberi Coquillett occurs in the eastern United States
from New York southward. It breeds commonly in water collected in
tree holes, but occasionally in artificial containers.
Anopheles walkeri Theobald is a widely distributed species in the
eastern United States to the Gulf of Mexico and north to Minnesota.
The presence of white "knee" spots and the golden color of the halters
distinguish it from A. atropos D. and K.
Anopheles atropos Dyar and Knab is strictly a salt-water mosquito
confined to the shores of the Gulf of Mexico from Florida to Louisiana.
It is small and very dark with unspotted wings. It lacks white knee spots.
Within its range it is said to outnumber A. crucians Wied. three to one
and breeds by preference in permanent salt pools or in shallow water
on muck or alluvial marshes. It is reported to be a tormenting pest, biting
in direct sunlight as well as by night.
Anopheles albimanus Wiedemann, is considered to be the most im-
portant anopheline species in the Caribbean region. The hind tarsi are
conspicuously white-banded; the apical segment of the palpi is entirely
white. It develops freely in brackish or salt water. This species has been
reported from several points in the extreme southern parts of the United
States, e.g., Key West, Florida, and the Rio Grande Valley (Browns-
ville), Texas.
Key to Culicid Tribes and Genera of the United States

(Prepared by T. H. C. Aitken after various authors)
1. Abdomen without scales, or at least with the sternites largely bare.

Scutellum never trilobed; crescent-shaped with the marginal setae
evenly distributed. Palpi (males and females) as long or almost as
long as the proboscis. Tribe Anophelini, Genus Anopheles Meigen 1818
Abdomen with both tegites and sternites completely clothed with
MOSQUITOES 209
scales. Scutellum trilobed (except Megarhinini-spurious vein extends

toward base of wing in upper basal cell from angle of R, + 5). Palpi
(females) much shorter than the proboscis 2
2. Proboscis rigid, outer half more slender and bent backward ..
. Tribe Megarhinini, Genus Megarhinus Robineau-Desvoidy 1827
Proboscis more flexible, of uniform thickness (unless swollen at tip),
outer half not bent backward ............ Tribe Culicini 3
3. Postnotum with setae.. . ............ Genus Wyeomyia Theobald 1901
Postnotum without setae... 4
4. Wing membrane without microtrichia. Cell R, shorter than its stem;
anal vein ends about opposite base of cubital fork. .
.Genus Uranotaenia Lynch-Arribalzaga 1891
Wing membrane with microtrichia. Cell R, longer than its petiole.
Anal vein extending well beyond fork of cubitus ............. . 5
5. Postspiracular bristles present ............................................................... . 6
Postspiracular bristles absent. .......................................... . 8
6. Prespiracular bristles present (even if few)
... Genus Psorophora Robineau-Desvoidy 1827
Prespiracular bristles absent. 7
7. Wing scales mostly narrow, or when broad, setae are present on upper
side of base of Rl (1st vein) Genus Aedes Meigen 1818
Wing scales broad; setae absent on uppper side of Rl
Genus Mansonia Blanchard 1901 (Subgenera Mansonia, Mansonioides,
Rhynchotaenia )
8. Prespiracular bristles present; lower side of base of Rl (1st vein)
distinctly pilose . .......... Genus Culiseta Felt 1904
Prespiracular bristles absent; lower side of base of Rl scaly or
bare ............ . 9
9. Pulvilli present .................................................................................... . 10
Pulvilli absent.. . .... ....... ...... .... ............... . 11
10. Second joint of antenna (first flagellar) very long in both sexes; an-
tenna of male not plumose ..... Genus Deinocerites Theobald 1901
Second joint of antenna normal; antenna of male nearly always
plumose ...... Genus Culex Linnaeus 1758
11. Fourth joint of front tarsus very short (both sexes); first segment of
front tarsi longer than the last four together ...
............. Genus Orthopodomyia Theobald 1904
Fourth joint of front tarsus not shortened in female; first segment of
front tarsi not longer than last four together ..
Genus Mansonia Blanchard 1901 (Subgenus Coquillettidia)
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alhimanus Wiedemann," Ann. Entomolog. Soc. Amer., 29:480-89.
41. Boyd, Mark F., 1929. "Studies on the bionomics of North American
2]2 1'.!EDTCAL ENTOMOLOGY
anophelines. I. The number of annual broods of A. quadrimaculatus," Am. ].
Hyg., 7:264-75.
42. Greenwald, Margaret; Cochran, J. H.; and Scharff, Donald K.; 1948.
"Large scale rearing of Anopheles quadrimaculatus Say at Orlando, Florida,"
Mosq. News, 8:50-56.
43. Carter, H. R.; Le Prince, J. A. A., and Griffitts, T. H. D.; 1916. Im-
pounded water. Washington, D. C.: Govt. Print. Office, in Pub. Health Bull.,
no. 79. 34 pp.
44. Mitzmain, M. B., 1917. "Anopheline mosquitoes, their distribution
and infection under field conditions," U. S. Public Health Service, Pub. Health
Rep. 32:536-40.
45. Reeves, W. C., and Hammon, W. McD., 1943. "Feeding habits of the
proven and possible mosquito vectors of western equine and St. Louis
encephalitis in the Yakima Valley, Washington," Am. ]. Trap. Med., 24:131-34.
46. Aitken, Thomas H. G., 1945. "Studies on the anopheline complex of
Western America," Univ. Calif., Publ. in Entomol. 7:273-364.
CHAPTER XII
._.
MOSQUITOES AS VECTORS OF DISEASE
A. THE MALARIAS
Human Malaria. Malaria is a widely distributed disease, prevalent
to a greater or less degree on every continent and on many islands of
the seas. It is considered to be the most important disease of man.
Whether or not malaria existed in North America prior to the discovery
of the continent has not been definitely established; however, this disease
was evidently recognized as a factor in colonization on the Massachusetts
coast and the Georgia-Carolina coast as early as the middle of the 17th
century.l In the United States, where malaria is on a rapid decline, there
were an estimated 900,000 cases as late as 1935; approximately 3,900
deaths were reported for 1934 in 13 southern states. 2 The number of
deaths in 1944 due to malaria was reported to have been 584, of which
540 occurred in the 14 southern states. 3
The presence of endemic malaria is dependent upon a complex of
environmental factors favorable to the development of large numbers
of vector (anthropophilous) mosquitoes, as well as to the Plasmodia
causing the disease. Temperature, particularly as it affects the develop-
ment of the plasmodium in the mosquito, and temperature combined
with humidity as it affects the life of the vector, are critical factors; a
mean summer isotherm of 15° to 16° C in general limits its geographical
distribution fairly well. The distribution of malaria is dependent upon
the availability of water for mosquito breeding, not necessarily heavy
rainfall; naturally arid regions may be seriously affected because of im-
perfections in irrigation if this is practiced. Although lowlands are more
likely to be affected, this does not hold as a general rule, because if one
or more important factors are lacking in a lowland region the area is
nonmalarious. The disease may occur at high elevations (9,000 feet in
Quito) under favorable circumstances. The writer found endemic malaria
in California at an elevation of about 5,500 feet; it is present in Mexico
at an elevation of near 7,500 feet.
Few diseases have so large a list of synonyms; among these are ague,
chills and fever, jungle fever, paludism, marsh fever, remittent fever,
intermittent fever, Wechselfieber, Kaltesfieber, etc. The symptoms are
213
2]2 MEDICA[. ENTOMOLOGY
anophelines. 1. The number of annual broods of A. quadrimaculatus," Am. J.

Hyg., 7:264-75.
42. Greenwald, Margaret; Cochran, J. H.; and Scharff, Donald K.; 1948.
"Large scale rearing of Anopheles quadrimaculatus Say at Orlando, Florida,"
Mosq. News, 8:50-56.
43. Carter, H. R.; Le Prince, J. A. A., and Griffitts, T. H. D.; 1916. Im-
pounded water. Washington, D. C.: Govt. Print. Office, in Pub. Health Bull.,
no. 79. 34 pp.
44. Mitzmain, M. B., 1917. "Anopheline mosquitoes, their distribution
and infection under field conditions," U. S. Public Health Service, Pub. Health
Rep. 32:536-40.
45. Reeves, W. C., and Hammon, W. McD., 1943. "Feeding habits of the
proven and possible mosquito vectors of western equine and St. Louis
encephalitis in the Yakima Valley, Washington," Am. /. Trop. Med., 24:131-34.
46. Aitken, Thomas H. G., 1945. "Studies on the anopheline complex of
Western America," Univ. Calif., Publ. in Entomol. 7:273-364.
CHAPTER XII
._.
MOSQUITOES AS VECTORS OF DISEASE
A. THE MALARIAS
Human Malaria. Malaria is a widely distributed disease, prevalent
to a greater or less degree on every continent and on many islands of
the seas. It is considered to be the most important disease of man.
Whether or not malaria existed in North America prior to the discovery
of the continent has not been definitely established; however, this disease
was evidently recognized as a factor in colonization on the Massachusetts
coast and the Georgia-Carolina coast as early as the middle of the 17th
century.1 In the United States, where malaria is on a rapid decline, there
were an estimated 900,000 cases as late as 1935; approximately 3,900
deaths were reported for 1934 in 13 southern states. 2 The number of
deaths in 1944 due to malaria was reported to have been 584, of which
540 occurred in the 14 southern states. 3
The presence of endemic malaria is dependent upon a complex of
environmental factors favorable to the development of large numbers
of vector (anthropophilous) mosquitoes, as well as to the Plasmodia
causing the disease. Temperature, particularly as it affects the develop-
ment of the plasmodium in the mosquito, and temperature combined
with humidity as it affects the life of the vector, are critical factors; a
mean summer isotherm of 15° to 16° C in general limits its geographical
distribution fairly well. The distribution of malaria is dependent upon
the availability of water for mosquito breeding, not necessarily heavy
rainfall; naturally arid regions may be seriously affected because of im-
perfections in irrigation if this is practiced. Although lowlands are more
likely to be affected, this does not hold as a general rule, because if one
or more important factors are lacking in a lowland region the area is
nonmalarious. The disease may occur at high elevations (9,000 feet in
Quito) under favorable circumstances. The writer found endemic malaria
in California at an elevation of about 5,500 feet; it is present in Mexico
at an elevation of near 7,500 feet.
Few diseases have so large a list of synonyms; among these are ague,
chills and fever, jungle fever, paludism, marsh fever, remittent fever,
intermittent fever, Wechselfieber, Kaltesfieber, etc. The symptoms are
213
commonly characterized by more or less regularly occurring febrile
paroxysms. In most cases there are three fairly well-defined stages, viz.:
the cold stage (the chill) in which the skin becomes pale and has the
appearance of "gooseflesh," the patient's teeth may chatter, and he may
shiver more or less violently; the next stage is the hot stage, or fever, the
temperature rising during the chill, the skin is hot and flushed; the third
stage is marked by the appearance of a general perspiration, the fever
falls, and the temperature approaches normal. The entire paroxysm may
last but a few hours. In many cases the stages are not well marked,
neither do the paroxysms recur at the same intervals. The intervals
depend largely on the type of infection. When the paroxysm occurs at
intervals of 24 hours, as is often true in the early stages of infection or
in multiple infections, it is quotidian; when the interval is 48 hours or
every third day it is tertian; and when the interval is 72 hours or every
fourth day, it is quartan.
The disease is caused by an infection with one or more of several
species of blood-inhabiting protozoa (Class Sporozoa) belonging to the
genus Plasmodium. Under the clincal term "malaria," there are really
combined four or more diseases, which, while caused by closely related
parasites and haVing a common means of transmission, nevertheless
possess individual characteristics, namely: vivax malaria, tertian; falci-
parum malaria, estivoautumnal; quartan malaria, quartan fever; ovale
malaria, ovale tertian.
The plasmodial parasites attack the red blood corpuscles, destroying
them while undergoing asexual reproduction; this asexual reproduction
or sporulation occurs at more or less regular intervals, i.e., 48 or 72
hours, depending upon the species of Plasmodium. The infection, ac-
cording to Reed,4 results in (1) changes in organs, such as enlargement
of the spleen and liver, and heart involvements; in fatal cases of sub-
tertian malaria capillaries in the brain and pia are found congested or
blocked by schizonts and sporulating forms of plasmodia, with pun~i
form hemorrhages in the white matter of the cerebral cortex; (2) leuco-
penia with increase of mononuclears and varying degrees of anemia as
the result of direct destruction of red cells by plasmodia and indirect de-
generation of others; (3) malarial pigment (melanin) in macrophages
in the splenic sinuses is characteristic, as are also the heavy pigment
deposits in the cells in the splenic vein and liver; it is the same pigment
as that produced in the red cells by the plasmodia and released with
the rupture of infected red cells; (4) changes in physiology such as
periodic febrile paroxysms; these are quite regular in benign tertian
malaria, but because of irregular maturing of plasmodia the periodicity
of the paroxysms is often concealed in falciparum (subtertian) malaria;
focal symptomatology due to localization in subtertian [Plasmodium
fa1ciparum (Welch)] arising from the "sticky tendency" of parasitized
red cells which causes agglutination and blockage; (5) malaria cachexia,
a chronic condition following repeated malarial attacks. Persons con-
cerned technically with malaria should consult Russell, West, and Man-
well's Practical Malariology" and Symposium on Human Malaria."1I>
Historical. Although the name "malaria" was not employed until the
middle of the eighteenth century, the disease was known for many
centuries, Hippocrates having divided periodic fevers into quotidian
( daily), tertian (every third day) and quartan (every fourth day). The
fable of Hercules and the Hydra is believed to refer to malaria. In 1753
Torti named the disease "malaria," believing it to be air-borne and
emanating from the bad air (mal aria) rising from swamps and marshes.
Credit for the discovery of the causal agent (1880) belongs to C. L. A.
Laveran, a French army surgeon who was then stationed in Algeria.
Although the mosquito transmission theory is said to have been held for
many years by Italian and Tyrolese peasants and the natives of what
was formerly German East Africa, the first well-formulated mosquito-
malaria theory was advanced by Dr. A. F. A. King in 1883. 7 In 1885 and
1886 GolgiB discovered that the course of the fevers corresponded to the
development of the parasites in the blood corpuscles and particularly to
their periodic sporulation. He demonstrated this for both the quartan and
terian parasites.
Manson expressed a strong belief in the malaria-mosquito theory as
early as 1884, and it was his sustained guidance and encouragement that
carried Ronald Ross9 on to those brilliant discoveries in India in 1897-
1898 that definitely incriminated mosquitoes as vectors of malaria, and
twice won for him the Nobel prize. Although Ross made important dis-
coveries in the field of human malaria and its anopheline vector, his
chief discovery was the complete life cycle of the causal organism of
bird malaria in a culicine mosquito; he established the bird-to-mosquito-
to-bird cycle.
Credit is due MacCallum 10 for his discovery in 1897 that the flagel-
lated bodies which Ross had found in the intestines of mosquitoes were
actually male parasites and that these fertilize the female cells, thus
giving the clue to the nature of the pigmented cells in the stomach wall
of the mosquitoes. MacCallum actually observed the process of fertiliza-
tion in blood taken from a patient suffering from aestivo-autumnal ma-
laria. After describing the process of flagellation in the bird infection
and entrance of the "flagellum" into the granular round form, MacCallum
remarks (loc. cit.): "Have we not here, without much doubt, a sexual
process in the organisms the result of which is the motile vermiculus.
. . . We can thus consider the two forms of adult organism found in
the fresh blood as male and female, the granular form containing more
II> Also see Boyd, Mark F., 1950. Malariology. Philadelphia: W. B. Saunders Co.
2 vols. 1643 pp.

chromatophilic substance than the hyaline male form, which we may
now say gives up its life in the production of four or more spermatozoa
or flagella."
In an addendum MacCallum states that he had
examined the blood of a woman suffering from an infection with the aestivo-
autumnal type of organism, in which a great number of crescents were to be
seen. These in a freshly made slide of blood, with very few exceptions, re-
tained their crescentic shape for only a few minutes. They soon drew them-
selves up, thus straightened out the curve of the crescent while shortening
themselves into the well-known ovoid form. After the lapse of 10 to 12 minutes
most of them were quite round and extra-corpuscular, the "bib" lying beside
them as a delicate circle or "shadow" of the red corpuscle.
After 20 to 25 minutes certain ones of these spherical forms became
flagellated; others, and especially those in which the pigment formed a definite
ring and was not diffused throughout the organism, remained quiet and did
not become flagellated. In a field where an example of each form could be
watched, the flagella broke from the flagellated form and struggled among
the corpuscles, finally approaching the quiet spherical form; one of them
entered, agitating the pigment greatly, sometimes spinning the ring around.
The rest were refused admission, but swarmed about, beating their heads
against the wall of the organism. This occurred after 35 to 45 minutes.
After the entrance of the flagellum the organism again became quiet and
rather swollen, but although in the two instances in which this process was
traced, the fertilized form was watched for a long time, no form analogous to
the "verrniculus" was seen.
During this same period of discovery in the field of malaria Grassi"

and associates proved that human malaria is transmitted by a particular
genus of mosquito, namely, Anopheles.
In 1900 Sambon and Low, at the suggestion of Manson, built a mos-
quito-proof hut in the Roman Campagna, in which they lived during the
most malarial months of that year without contracting malaria, having
taken precautions against mosquito bites by promptly retiring within
doors at sunset; otherwise they lived as did the natives. At this time
these investigators sent infected Anopheles mosquitoes from the Roman
Campagna to London, where Manson's son, Dr. P. Thurburn Manson,
and Mr. George Warren permitted themselves to be bitten by these
mosquitoes and "shortly afterwards both of these gentlemen, neither of
whom had been abroad or otherwise exposed to malarial influences,
developed characteristic malarial fever, and malarial parasites were
found in abundance in their blood, both at that time and on the oc-
currence of the several relapses of malarial fever from which they sub-
sequently suffered. The mosquito-malaria theory has now, therefore,
passed from the region of conjecture to that of fact."12
Bass and Johns " in 1912 were the first to cultivate successfully the
1
malaria parasite in vitro.

The plasmodia. The causal organisms of malaria belong to the genus
Plasmodium of the family Plasmodiidae, suborder Haemosporidiidea,
order Coccidiida, class Sporozoa, Phylum Protozoa. They are blood-
inhabiting microparasites, passing the principal portion of their asexual
cycle within the red blood cells of the host where they produce, with
the aid of hemoglobin, a characteristic malarial pigment (melanin).
If parasites are present in the blood they should be visible, after
proper staining, on careful thin-smear microscopic examination, as pig-
mented intra corpuscular bodies in the form of signet rings, ameboid
forms, segmenting forms, or as crescents in estivo-autumnal fever of 10
or more days' duration. An excellent manual by Wilcox14 is available for
the microscopical diagnosis of malaria in man. Of interest in routine
examination of malaria blood smears is the use of fluorescent dyes (0.1
per cent aqueous rivanol or berberine sulfate fluorochrome) and then
irradiation by ultraviolet ray (Metcalf, 194515 ).
RossI6 states that the paraSites "will not generally be numerous
enough to cause illness unless there is at least one parasite to 100,000
haematids; that is, 50 parasites in 1 cmm of blood; or 150,000,000 in a
man 64 kilograms (142 pounds) in weight. ... Such calculations dem-
onstrate the absurdity of supposing that there are no plasmodia present
in a person because we fail in finding one after a few minutes' search.
As a matter of fact, even if as many as 150,000,000 plasmodia are present
in an average man, the chances are that ten to fifteen minutes' search
will be required for each plasmodium found; while if we are careless
or unfortunate, we may have to look much longer." The time elapsing
from inoculation by the mosquito to appearance of infected erythrocytes
(presumably one or more exoerythrocytic cycles) varies with the species
of the plasmodium as well as with the strain and other factors. In
falciparum malaria the incubation period for the Coker strain ranged
from 6 to 25 days, while that for the Long strain ranged from 9 to 10
days.
a. Plasmodium falciparum (Welch 1897) ( Plasmodium praecox
Blanchard 1900) is the causal organism of estivo-autumnal fever (sub-
tertian) of the tropics and subtropics, the most severe form of malaria,
often resulting fatally. Although it is a tertian fever there is considerable
irregularity in the occurrence and duration of the febrile stage owing
to a corresponding irregularity in the sporulation of the parasites, schizo-
gony usually requiring about 48 hours, though often less. The infected
red corpuscles are usually normal in size, though some may be slightly
shrunken, often crenated, and rather dark green (brassy). The intra-
corpuscular parasite in all its stages is small (not over two-thirds the
size of a corpuscle) and fairly ovoid in outline; the pigment is darker
than in other forms; clumps early in coarse granules, and "Maurer's
dots" appear in the corpuscles in the later stages. The signet ring is thin
and small and the chromatin dot is commonly double and out of line
with the ring. There may be two and even four signet rings in one red
corpuscle, The segmented state, rarely if ever seen in the peripheral
blood, produces from 8 to 24 merozoites. Characteristic crescent-shaped
or kidney-shaped bodies appear in the peripheral blood in about 10 days
after infection; these are the sexual forms (gametocytes) and occur as
crescents in this species of Plasmodium only. The macrogametocyte or
female form, measuring from 10 to 15J,l shows the chromatin granules
well concentrated in the mid-region, while the micro gametocyte or male
form, measuring from 7 to 10 J.I. has a more hyaline appearance. A remnant
of the red blood corpuscle often remains slung from the opposite ends
of the crescent and forms the so-called "bib."
h. Plasmodium vivax (Grassi and Feletti 189() is the cause of tertian
fever of temperate climates, which occurs also abundantly in the tropics
and subtropics, with regularly recurrent paroxysms every 48 hours.
The parsitized corpuscles are distinctly enlarged, quite pale, and usually
contain fine pigment granules known as "Schiiffner's dots." The signet
ring is large and conspicuous and the dot is in line with the ring and
rarely double. The fully grown merocytes or schizonts are very irregular
and bIzarre in form. The number of elements, merozoites, in the sporu-
lating or segmented stage commonly seen in the peripheral blood is from
12 to 24 (usually about 16), and their arrangment is irregular. Sporula-
tion occurs regularly every 48 hours. There are no "crescents" in this
species; the gametocytes are round or oval in form, filling practically
the entire red cell when full grown. The macrogametocyte has the
chromatin arranged in a compact mass; the microgametocyte has the
pigment well distributed and presents a more hyaline appearance.
c. Plasmodium malariae (Laveran 1881) is the cause of quartan
fever, with recurrent paroxysms every 72 hours. This form of malaria
is much less common but coincides in distribution with estivo-autumnal
fever. The pigment is coarse and generally occurs in marginal streaks.
The parasitized corpuscles are usually normal in size, and the parasite
is small and more or less oval in shape though, when partly grown, it
frequently extends across the equator of the corpuscle in the form of a
band. The ring-forms have one vacuole and usually one dot. The gameto-
cytes are rarely seen. The segmenting stage gives rise to the typical
"daisy" form, each sporulated body radiating from the center. The num-
ber of bodies varies from 6 to 12, usually 8. Sporulation occurs every 72
hours. The gametocytes resemble those of Plasmodium vivax.
17
Rodhain has shown that P. malariae can be transmitted intrave-
nously from man to the chimpanzee and maintains its virulence and spe-
cific character for man; also the malariae type of plasmodium of the
naturally infected chimpanzee is transmissible to man, producing a
malaria fever of the quartan type.
d. Plasmodium ovale Stephens 1922 is the cause of a mild form of
tertian fever in East Africa and probably South America. The name
indicates the oval shape which is generally assumed by the parasite as
well as the parasitized erythrocyte. The infected corpuscles may become
somewhat enlarged. The pigment is dark and granular and "Schiiffner's
dots" are present in all stages. The merozoites range from 8 to 12 in
number.
Life cycle of the plasmodia. The life history of malaria parasites con-
sists of two cycles: first, the asexual, also known as the blood cycle,
cycle of Golgi,'B or schizogonic cycle; and, secondly, the sexual, also
known as the mosquito cycle, cycle of Ross, or sporogonic cycle. 19
The asexual cycle (Fig. 78), accomplished in the blood and other
tissues of man, begins with the injection of spindle-shaped sporozoites
(Fig. 78) into the circulation with the bite of an appropriate anopheline
mosquito.
Though most diagrams illustrating the life cycle of the malaria
parasite show the sporozoite entering the erythrocyte at once (including
Fig. 78), actually there has been much speculation for many years as to
what did happen in the life cycle of the Plasmodium between the in-
oculation of the sporozoite by the mosquito and the appearance of the
trophozoite or signet ring in the erythrocyte (Huff 20 ). That there are
pre-erythrocytic stages in the developmental cycle of Plasmodium elon-
gatum Huff and P. gallinaceum Brumpt and other bird malaria parasites
has been proved by Huff and associates;21 the first generation following
infection by sporozoites being called "cryptozoites" and subsequent
generations, "metacryptozoites." These stages "require a period of de-
velopment in macrophages and related kinds of cells before they invade
erythrocytes." Huff22 raises the question, "Do such stages really exist in
human or simian malarial infections?" He cites indirect evidence viz.:
"( 1) the existence of periods of inapparent parasitism followed by para-
sitemia in sporozoite- ~duced infections, and (2) the differences between
sporozoite-induced and blood-induced infections with respect to their
response to anti-malarial drugs. . ., The evidence is conclusive that
the sporozoites and the 3 or 4 generations of development following them
do not occur in the blood during this period"; i.e., subinoculations within
about half an hour of blood from a heavily sporozoite (mosquito) in-
oculated malaria patient may occasionally result in an infection; however,
220 ~lEDJCJ\L ENTO~I()LUCY
following this brief period there is a period af seven or more days (de-
pending apparently on the species of Plasmodium) when subinoculations
will not result in infection.
__-------------------IOdoys ond over------~ _________________
_ - - - - - - 48 hours---------------~-------..-
(trophozoite) Amoeboid stag~
Signet ri~g .stage at parasite
'.oJllhtn corpus'e1e /
0
V._)
(f1:::..,
~
PorOSlI. enlers / LIFE CYCLE '\
humon red
blood corpuscle
(2or3moyenter
0 (Asexual) Mature parasite
0EJIn corpuscle
I
l
some corpuscle) \ IN HUMAN BLOOD Developing gametocyte
,/ ~ found in human
I Merozoites enter new corpuscle s ) I """ spleen and bone
Q) and repeat cycle 6\\ morrow
..)~ Fully developed merozolles break ~
~~~~~o~tl~~~n'er r 0 G (!) e I?J out of old corpuscle
(somesell.uol,
some a se wol)
I ~ (:) ElG~
G~~~~\ _
fEJ. .
...-
~(merocyte.orSC~IZ0nt)
\s;;?Y Parasite beginS
, e (!:l i9J to divid e v
'\ ' Pa r asite co ntin ues dlvid il'\g
\., - . ' ( sporulating) ..-~'
~ ~ o~Gometocyte
'y
.-
Female Anopheles biting human, cI (crescent)
t , , " "inJU'lIiJt.r"p~C:;flOr.1'..<;.jtp5" "fr.J)m " itc:; _ ~ PCffC'511V.i1i,Ji,Jumcn
~ salii vary glands into human
I
" "'-~ blood stream
d /'
./ " /
" blood into st<?moch
f
0 mosqui 0
t
Sporoloites migra te
d2 \
Salivary
glands
forward to
salivary glands
of female mosquito I I
I Development
of gametocytes ~
1
LIFE CYCLE
0 1 in stomach cavity E
(5e,uol)
l
IN 0 1
\
MOSQUITO
Fertilizati:> .
Stomach of Anopheline
mosquito
__ ___ Parasite sparulotes

within cyst
- Division continues and
cyst enlarges
- - - - - - 5 days or more---------------------__________________
FIG. 78. The life cycle of the malarial parasite, Plasmodium falciparum, in
man and the anopheles mosquito. (For exoerthrycytic cycles, see text.)
Following the presumable exoerythrocytic stages the cycle continues

very much as is shown in the diagram (Fig, '78), i.e., the invasion of the
erythrocytes and parasitemia, except that there may be repeated exo-
erythrocytic cycles in addition and concurren.tly, which helps to explain
relapses as well as certain medicinal failures. The young trophozoite
having parasitized an erythrocyte the "signet ring" stage is reached,
growing rapidly until the erythrocyte is more or less filled, depending
upon the species of parasite, and it is then known as a merocyte. The
full-grown merocyte (also known as a schizont) now divides into a
larger or smaller number of bodies (also depending upon the species)
which are then liberated, and when free in the plasma are known as
merozoites. The time required for this sporulation is from 24 to 72 hours
according to the species. Merozoites, like sporozoites, presumably also
have an exoerythrocytic cycle, after which the resulting trophozoites
enter other red cells and again the cycle repeats itself until the infection
is great enough to produce a paroxysm, i.e., in from 6 to 12 days, com-
monly about 10 days. The paroxysms may be due to the release of toxins,
although there is no clear evidence concerning this.
The great majority of the merozoites are asexual, but some of them
are potential males and females, which require a longer time, probably
not less than 10 days, to develop to their full growth, and are then
known as gametocytes. In Plasmodium vivax the sexual forms are not
easily recognized; however, the following characters are useful: "( 1 )
their larger size, (2) more abundant pigment, (3) there is usually only
one fairly large chromatin mass, whereas in an asexual form (schizont)
of nearly equal size the chromatin has already begun to divide into
several portions (segmenting stage)" (Stephens and Christophers). In
P. falciparum the sexual individuals are in the form of crescents. The
female crescent (macrogametocyte) has the pigment collected at the
center, while the male crescent (micro gametocyte) has the pigment
scattered throughout and is known as a hyaline crescent.
With development of the gametocytes completed all is ready for the
next cycle (the sexual) which can only be accomplished within the body
of anopheline mosquitoes. In the meantime the asexual cycles are re-
peated until senescence or some other biological process ends the life
of the parasites or until quinine or other plasmodiacide destroys them.
The gametocytes are not easily destroyed, persisting in the body for
long periods, during which time the infected person is a carrier. A per-
son eventually removed from reinfection becomes rid of malaria probably
because of parasite senescence. Rejuvenation of the parasite brood and
consequent relapses even after many years of latency may perhaps result
from the exoerythrocytic cycle referred to earlier.
Sexual cycle, the cycle of Ross (Fig. 78), has been observed only in
female anopheline mosquitoes; in the stomach of which flagellation of
the male gametocyte takes place. After a peripheral arrangement of the
chromatin (in clumps corresponding to the number of flagella) there are
extruded from three to six long slender £laments (flagella), each of
which separates from the parent body (exflagellation), forming the male
FIG. 79. Stomach of female anopheles mosquito with numerous plasmodial
cysts. (Photograph by Mayne.)
222
MOSQUITOES AS VECTORS OJ;' DISEASE 223
gamete (microgamete) corresponding in function to the spermatozoon
of higher animals. The female gametocyte, now known as the mac1'O-
gamete, having been taken into the stomach of the mosquito with the
microgametocytes in the act of sucking bloo<1, also undergoes certain
changes (maturation), becoming rounded or oval in form with the
chromatin mass centrally located. In this C()ndition and still in the
stomach of the mosquito, the microgamete COnjugates with the macro-
gamete, producing the zygote, which soon becomes motile and is then
known as the ookinete or vermiculus, in which stage the epithelium
of the stomach is penetrated and a position is shortly taken up just be-
neath the peritoneal membrane. Based on his studies of the plasmodia
of birds HufP:; points out that this penetration of the stomach wall
FIG. 80. A bursting plasmodial cyst on stomach elf mosquito. Spindle-shaped

sporozoites being liberated. (Greatly magnified.) (Photograph by Mayne.)
is not a boring process, for this zygote has lost its I)ointed ends long before the
penetration begins. When the ookinetes are first found in the vicinity of the
stomach wall, they are lying parallel to it in the serous mucoid layer adjacent
to the cells of the stomach wall. As the parasite grows, it becomes relatively
thicker and gradually force two of the stomach tells apart. It gradually be-
comes more spherical and forces the stomach cells apart nearer and nearer the
outside of the stomach wall. The stomach cells I10W begin to come back to
their original positions on the inner side. Finally the parasite, now an oocyst,
comes to lie under the outer envelope of the stoma.ch.
In this position the parasite grows enormously, forming an oocyst (Fig.

79) in which many nuclei appear in from fott1' to five days. These tiny
nucleated bodies give rise to hundreds of spindle-shaped organisms

(sporozoites) which are shed into the body cavity of the mosquito in
from 24 to 48 hours (Fig. 80). The majority of the sporozoites eventually
collect in the salivary glands, remaining there until the mosquito bites
again, when many of them may be injected with the saliva into the
wound. The time required for the completion of the sexual cycle varies
from 7 to 10 days under favorable conditions. Once infected the mosquito
probably remains infected and infective for the rest of its life. In making
malaria surveys one should be proficient in detecting plasmodial infec-
tions in anophelines. 24
What makes a mosquito a good natural vector? Conclusive experi-
mental evidence indicates that the Plasmodia of human malaria do not
develop within the bodies of culicine mosquitoes even though game-
tocytes are ingested. Whether all anopheline species are infectable is,
of course, not known; many malariologists are of the opinion that labora-
tory tests tend to prove that it is a matter of degree only, that no an-
opheline is completely refractory to the plasmodia of human malaria.
There is not only a variation in the degree of mosquito species hospi-
tality to plasmodia but also a variation in the species of plasmodia. Thus
A. maculipennis Meig. and A. quadrimaculatus Say are known to be
carriers of three species of plasmodia (Beyer, et al.,25 Thayer,26 King27)
while A. punctipennis (Say) is a strong carrier of the tertian parasite
and a very weak one (probably negligible) for the estivo-autumnal
parasite (Darling 28 ). Moore, Young, Hardman, and Stubbs 29 tested the
infectivity of foreign malarias for four species of California anophelines,
A. freeborni Aitken, A. punctipennis ( Say), A. occidentalis D. and
K., and A. pseudopunctipennis franciscanus McG They all appeared to
have about the same susceptibility; all developed sporozites. Successful
transmission was effected with the first three species. The length of the
sporogonous cycle in the four species at 75° F was about 10 days; the
shortest cycle was 8 days in A. freeborni.
A good natural vector is an Anopheles species which is freely in-
fectable by the several species of human plasmodia, offers a favorable
environment for development to the sporozoite stage, breeds, success-
fully and abundantly, is a house invader, and takes human blood re-
peatedly, i.e., is anthropophilous. The maintenance of close and constant
contact between an anthropophilous mosquito and its source of food
supply is an important factor in endemicity. The concept of species
sanitation is based on a knowledge of the factors referred to above.
An anopheline may be common, even abundant, yet of little importance
as a vector for one or more reasons.
Hindle 30 points out that, "the first instance of an Anopheles being
shown not to transmit malaria was in the case of the common Indian
species Anopheles subpictus (rossi) Grassi. This species is found quite
commonly in very large numbers associated with every degree of preva-
lence of malaria, but it has not been shown to act as a transmitting agent
(in India) under natural conditions, though it can be infected experi-
mentally."
"Anophelism without malaria." Hackett (lac. cit. p. 3) points out
that because of the presence of anophelines in great numbers in areas
without malaria, various theories have been advanced to explain this.
"Anophelism without malaria" is an expression now commonly used in
the study of malariology. Hackett refers to such theories as (1) robust
mosquitoes produced under unusually favorable conditions are un-
susceptible to plasmodial infection (Alessandrini); (2) brackish water
breeds robust anophelines which easily become infected, live long, and
are therefore most dangerous (Grassi); (3) coumarin, the active princi-
ple in a type of clover, Melilotus altissima, present in clover honey, either
kills sporozoites in the mosquito glands or protects the mosquito against
infection (D'Herelle); (4) adaptation of the maxillary tooth formula
to feeding habits of the mosquito,31 i.e., increase in the number of teeth
might be an adaptation to tough skins of larger domestic animals (zo-
ophilism); when the average number of teeth (maxillary index) was
between 14 and 15 this race fed constantly on domestic animals and
did not feed on man, those feeding on human blood haVing an index of
14 or less (Roubaud); (5) gonotrophic dissociation, which refers to
undeveloped ovaries in the female anopheline, following a blood meal,
e.g., a strain of anophelines which passes the winter in warm stables
and houses where there is plenty of available food, yet is not stimulated
to oviposition (Swellengrebel). Normally the ovaries of anopheline
females develop following a blood meal; also the females usually under-
go true hibernation without a blood meal. Thus A. maculipennis variety
atroparvus van Thiel resolves itself into two strains, one of which shows
gonotrophic dissociation and is the explanation of so-called "malaria
houses," i.e., when this strain "takes up winter quarters in a house, it
lives upon the family; and if the mosquito should be infected or some-
one in the house should be a carrier, by spring most of the family will
have contracted malaria and will have passed it on to the rest of the
anophelines sheltering in the house." (6) Geographic subspecies dis-
tinguished by egg types, seven being recognized, indicating wide
divergence in feeding and breeding habits; barred eggs of cattle feeders,
and eggs of more uniform pattern of malaria vectors and brackish water
breeders (Falleroni; and Hackett, Martini, and Missiroli, lac. cit. p. 3).
Number of persons infected by one mosquito. Whether an Anopheles
once infected can infect more than one person without again feeding
on infective blood is a matter of interest. Country-school privies as the
writer has often observed, may be infested with Anopheles mosquitoes
which have the opportunity to bite different persons in quick succession.
In an instructive series of experiments conducted by Mayne (Mitz-
main 32 ) he reports that one mosquito proved to be the sole infecting
agent in three cases. Mitzmain used Anopheles punctipennis (Say) with
Plasmodium vivax. He also demonstrated in 11 experiments that short
exposure to bites (interrupted feeding) was sufficient to cause success-
ful transmission of the disease.
Effect of temperature on plasmodia in the mosquito. In spite of the
fact that all conditions appear to be favorable ie., presence of numerou~
anopheline mosquitoes together with ample human population with
carriers of plasmodial gametocytes, malaria may be wholly absent in
particular localities. An analysis of conditions will usually reveal the fact
that the average temperature is low because the nights are cool al-
though the days may be fairly warm, or because of prevailing cool fogs.
It is generally agreed that malaria gametocytes cannot develop success-
fully within the body of the mosquito host when the average tempera-
ture is below about 60° F. It is nevertheless a matter' of interest to
know that King33 observed the survival of the parasite of tertian malaria
in the mosquito host (Anopheles quadrimaculatus Say) at a temperature
of 30° F for a period of two days; at 31 ° F for four days; and at 46° F
for 17 days; and the parasite of estivo-autumnal malaria survived a
temperature of 35° F for 24 hours.
Knowles and Basu 34 ( 1943 ), working with the vector Anopheles
stephensi Liston, found that the heaviest salivary gland infection of
Plasmodium vivax was obtained at 80° F and 50 per cent relative
humidity. Using the same species of Anopheles they found that at a
temperature of 100° F and with all percentages of humidity between
50 and 100 no infection with any species of malaria parasite was ob-
tained. The sporozoite stage of Plasmodium vivax was reached in the
salivary glands of A. stephensi in 18 days at 60° F, 15 days at 70° F,
11 days at 80° F, and 9 days at 90° F. For Plasmodium falciparum also
at 50 per cent relative humidity the time was 14 days at 70° F, 10 days
at 80° F, and 9 days at 90° F. Plasmodium malariae has been reported
to require 30 to 35 days at 67° F, with relative humidity not given.
Hibernating anophelines not carriers. Hibernation of the anopheline
host presents the problem of the overwintering of the parasite. Mayne
( Mitzmain35) again comes forward with an excellent discussion of the
question, "Is mosquito or man the winter carrier of malaria organisms?"
He reached the following conclusions:
. . . hibernating Anopheles, collected in the region investigated (northwestern

Mississippi), did not harbor parasites of malaria. This was determined after
an examination of 2,122 dissected anophelines, of which 1,211 specimens
were examined before May 15, 1915. Among the remaining 911 specimens,
serving as a malaria indicator for the spring season, 3 mosquitoes, between
May 15 and May 26, were definitely shown to contain oocysts, indistinguish-
able from those seen in mosquitoes experimentally infected with human
malaria.
In the investigation of man as the responsible winter carrier, 1,184 persons,
residing on the plantations selected, were examined for malaria parasites. Four
hundred and ninety-two infections were identified microscopically; 317 cases
were of subtertian type, 8 were mixed infections, and the remainder were of
the simple tertian type with the exception of one quartan case.
In the consideration of these infections an important fact stands out: nearly
one-fourth (24.8 per cent) of the human carriers harbored gametocytes.
It was proved that from a group of 103 persons, examined in March, 1915,
8 of the 15 gametocyte carriers identified were similarly infected during the
preceding fall.
The incrimination of man as the sole winter carrier is emphasized by the
fact that 3 malaria-infected Anopheles quadrimaculatus were found in the
homes of these gametocyte carriers during May 15 to May 26, previous to
which time 1,180 specimens of Anopheles from this source were found to be
negative.
Anophelines overwintering in w~rm stables and homes, as explained

in the previous chapter under races of Anopheles maculipennis Meig.,
may nevertheless play an important though highly circumscribed role
in the transmission of malaria.
Vectors of human malaria. It has been abundantly proved that the
causal organisms (plasmodia) of human malaria are transmitted from
man to man only through the agency of mosquitoes belonging to the
genus Anopheles. Russell, Rozeboom, and Stone 36 (1943) give identi-
fication keys to 240 species and 78 subspecies of anophelines of the
world. Of this number Russell in Practical Malariology (1946, loco cit.)
named 54 species as the chief malaria vectors. These are distributed
over the world, according to various authorities, as shown in the follow-
ing tabulation:
TABLE I
0
IMPORTANT ANOPHELINE VECTORS OF MALARIA
COUNTRY AND SPECIES LOCALITY BREEDING PLACES
UNITED STATES AND CANADA

Anopheles quadrimacula- (Eastern, central, and Sunlit, impounded water,
tus Say southern United States; marshes, swamps, rice
Gulf to Ontario) fields
Anopheles maculipennis ( Rocky Mountains, New Sunlit seepage water, ir-
freeborni Aitken Mexico, Pacific coast) rigation ditches, rice
fields (in part)
.. See also Herms and Gray.IT
COUNTRY AND SpJ;;CIES LOCALITY BREEDING PLACES
MEXICO, CENTRAL AMERICA,

WEST INDIES
Anopheles albimanus (Mexico, Central America, Su~lit lagoons (brackish
Wied. West Indies, Colombia to and fresh) , swamps,
Venezuela) pools
Anopheles p. pseudo- (Southern United States to Sunlit streams with green
punctipennis Theo. Argentina) algae, seepage, pools,
etc.
Anopheles darlingi Root (Central America, Vene- Sunlit fresh water, marshes,
zuela to Argentina) etc.
Anopheles aquasalis ( Central America, West Brackish marshes, irriga-
Curry Indies, to Brazil) tion seepages, etc.
Anopheles punctimacula (Mexico, Central America, Shaded pools, swamps,
D. and K. West Indies, to Brazil) streams
Anopheles bellator D. ( West Indies to Brazil) Water at base of leaves of
and K. bromeliads
Anopheles cruzii D. (Brazil ) Water at base of leaves of
and K. bromeliads
Anopheles m. aztecus (Mexico) Sunlit ponds with green
Hoffmann algae
SOUTH AMERICA
Anopheles albimanus (Colombia, Ecuador, See above
Wied. Venezuela)
Anopheles albitarsus L.- (Argentina) Rice fields, marshes, over-
A. flows, etc.
Anopheles p. pseudo- (To Chile and Argentina) See above
punctipennis Thea.
Anopheles darlingi Root ( Venezuela to Argentina) See above
Anopheles gambiae Giles (Imported into Brazil, now All sorts of collections of
exterminated" ) fresh water
EUROPE
Anopheles 1. labranchiae (Italy, Sardinia, Sicily, Upland streams, rice fields,
Falleroni Corsica, Spain, N. W. brackish coastal marshes
coast of Africa)
Anopheles labranchiae (England, The Nether- Sunlit fresh or brackish
atroparvus van Thiel lands, Germany, Sweden, marshes, swamps, etc.
Spain, northeastern
Italy)
Anopheles messeae Fal- (Norway, Russia, Siberia, Fresh-water pools, ponds,
leroni Manchuria) marshes
Anopheles superpictus (Spain, southern Europe, Pools in stream beds, ir-
Grassi Greece, Asia Minor) rigation canals, seepages,
etc.
Anopheles sacharovi (Balkans, Russia) Sunlit coastal marshes,
(= elutus) Favr. fresh or brackish water
NORTH AFRICA AND MIDDLE
EAST
Anopheles claviger Meig. (Ukraine, Asia Minor, Rock pools, wells, cisterns,
North Africa) marshes
Anopheles 1. labranchiae (Northwest coast of See above
FaIL Africa)
Anonheles pharoensis (North Africa, Palestine) Rice fields, swamps
Thea.
Anopheles sachaTOvi (Balkans, central Russia) See above
Favr.
Anopheles sergenti Theo. ( North Africa, Palestine, Rice fields, irrigation

Turkey, Syria) canals, borrow pits,
seepage water
Anopheles superpictus (Asia Minor) Stream bed pools, irriga-
Grassi tion canals, seepage
water, hill district
CENTRAL AND SOUTH AFIUCA
Anopheles funestus Giles (Tropical Africa) Stream margins, swamps,
ditches, seepage water
Anopheles gambiae Giles ( Tropical Africa, Egypt, Puddles, pools, sluggish
Arabia) streams, all sorts of col-
lections of water
Anopheles moucheti ( Uganda, Congo, Came- Stream margins, swamps
Evans roons)
Anopheles pharaoensis (Widely distributed in See above
Thea. Africa, Palestine)
Anopheles nili Thea. (Widely distributed in Shade loving, stream mar-
central Africa) gins
Anopheles pretoriensis (Widely distributed in Sunlit rock pools, stream
Thea. central and south beds, hoof prints, ditches
Africa)
PHILIPPINE ISLANDS
Anopheles minimus (Many islands, Java, Foothill streams, ditches,
f/avirostris Lud. Celebes) wells
Anopheles mangyanus (Many islands) Stream beds, irrigation
Banks ditches
JAPAN, NORTH CHINA,
KOREA
Anopheles hyrcanus (Widely distributed) Open clean water, rice
sinensis Wied. fields, swamps, ponds,
slow streams
Anopheles pattoni ( N orthen China) Beds of hill streams, rock
Christ. pools
SOUTH AND CENTRAL CHINA,
BURMA, TAIWAN
Anopheles minimus Theo. (Hilly regions southern Sunlit, slow moving
China, Taiwan, Burma) streams, rice fields, ir-
rigation ditches
Anopheles hyrcanus (Plain of central China, See above
sinensis Wied. Burma)
Anopheles ;eyporicllS (Hong Kong area) Rice fields in hilly country
Anopheles ;eyporicus
Anopheles culicifacies (Burma) Stream breeder, irrigation
Giles ditches
Anopheles maculatus (Burma, Southern China) Stream and river bed
maculatus Theo. breeder, rice fields,
pools, lake margins
Anopheles philippinensis (Burma, Inio-China) Rice fields, borrow pits,
Lud. swamps, ditches, tanks
INDIA AND CEYLON
Anopheles culicifacies (India, Ceylon) See above
Giles
Anopheles stephensi (India) Well, cisterns, roof gutters,
Liston temporary water
receptacles
Anopheles maculatus (India, Ceylon) See above

maculatus Theo.
Anopheles fiuviatilis ( India) Foothill stream edges,
James springs, irrigation canals
Anopheles minimus (Eastern and northern See above
Theo. Ceylon)
Anopheles philippinensis ( India) See above
Lud.
MALAYA (East Indies,
Siam, Malaya
Anopheles aconitus (East Indies, Malaya, Rice fields, irrigation
Donitz Indo-China) ditches, pools in creek
beds, reservoirs
Anopheles culicifacies (Siam) See above
Giles
Anopheles fluviatilis (Thailand) See above
James
Anopheles hyrcanus (Malaya, East Indies) Rice fields, impounded
nigerrimus Giles water, borrow pits, slug-
gish stretlms
Anopheles maculatus (Thailand, Malaya, East Stream and river bed
maculatu8 Theo. Indies) breeder, seepage water,
lake margins
Anopheles subpictus sub- (Malaya, East Indies) Borrow pits, buffalo wal-
pictus Grassi lows, all sorts of tem-
porary or permanent col-
lections of water
Anopheles sundaicus (Thailand, Malaya, Eiast Sea-water lagoons and
( Rodenwaldt ) Indies) swamps
Anopheles umbrosus (Malaya, Cochin-China, Shaded jungle pools, man-
(Thea.) East Indies) grove swamps
Anopheles philippinensis (Malaya, East Indies) See above
Lud.
A USTRALIA-MELANESIA-
POLYNESIA
Anopheles farauti (= A. (New Guinea, Solomcms Fresh or brackish water,
moluccensis) Lav. New Hebrides) , all sorts of natural or
artificial collections of
water, . clear or polluted
Anopheles punctulatus ( New Guinea, Solomor1s) Rain pools, stream mar-
punctulatus Don. gins, hoof prints, etc.
Anopheles bancrofti (New Guinea, northern Shallow, slow moving
Giles Australia) streams
Malaria surveys. Although an inquiry iIUO the facts about malaria
in some particular area may be described as a survey, Christophers,
Sinton, and Cove1l39 point out that " . . . tt is far from being merely
routine or mechanical." These authors stitte that the circumstances
affecting malaria are so varied "with its triflle-linked chain of man, the
parasite, and the mosquito, as well as all thlcJ various factors influencing.
this chain, that in the present state of our knowledge a malaria survey
is almost always a true piece of research work Such a survey is intended
to guide policy and action. . . . " Elmendm:f40 defines a malaria survey
as a quantitative as well as a qualitative procedure directed toward
securing information which will determine the status of malaria in a
community. The survey should serve as a blueprint for guidance . . .
and should provide an original base-line from which all subsequent
measurements of the course of the disease are determined."
In general the survey staff, on a minimum basis, should consist of the
following personnel: a malariologist, an entomologist trained as a culi-
cidologist, a sanitary engineer with special training in hydraulics. The
staff should include a laboratory technician trained in parasitology, a
mapper, and a clerk. Headquarters and laboratory should be conven-
iently located and properly equipped. Having made the acquaintance,
and having been assured the cooperation, of civil authorities, particularly
the local health officer, the following data will be assembled, recorded,
mapped, and card-indexed as required:
1. Demographical data: composition of the population by race, age, and
sex; occupations, economic status, housing, etc.
2. Meteorological data: rainfall, temperature, humidity, winds, etc.
3. Topographical data: soil, vegetation, etc.
4. Agricultural data: crops, farming methods, irrigation, drainage, rice
culture, marshes, etc.
5. Malaria statistics: morbidity and mortality; incidence (age, seasonal,
etc.); distribution, occupation, and economic status of malarial population;
spleen census, etc.
6. Parasitological and entomological data: parasite index based on blood
smears; oocyte and sporozoite index; numerical prevalence of all species of
adult anophelines with habitat preference (houses, pig pens, cattle sheds,
etc.); feeding habits of adults ascertained by use of precipitin test; larval in-
cidence (use of dippers), showing kinds and distribution of breeding places;
with a little practice collections of eggs can be made very readily. As previously
stated, eggs are important in the identification of anopheline mosquitoes.
(Aitken," 1948).
Animal Malarias
Avian malaria. Blood protozoa were nrst reported from birds in 1885
by Danilewsky from the Russian Ukraine. Herman 42 lists 178 species
and subspecies of North American birds in whose blood protozoon
parasites have been found; 63 of these protozoon infections are listed as
plasmodial. The species of avian plasmodia, of which there are many, can
be divided roughly into two groups based on whether they produce an
elongate or a round type of gametocyte. Among those belonging to the
elongate type is Plasmodium elongatum Huff, and among those belonging
to the round type are the following well-known species, Plasmodium
gallinaceum Brumpt, P. cathemerium Hartman, and P. relictum
(Grassi et Feletti). So far as is known avian plasmodia are transmitted
in nature by culicine mosquitoes, among them Culex pipiens Linn., C.
quinquefasciatus Say, and C. tarsalis Coq. (Reeves, loco cit. p. 182).
Numerous species of mosquitoes are susceptible to infection in the
laboratory, including Anopheles quadrimaculatus Say. For laboratory
purposes Aedes aegypti is an excellent vector of Plasmodium gallinaceum
Brumpt, as is Aedes atropalpus (Coq.) recently (1946) reported by
Trembly.
Other malaria-like infections of birds are caused by Haemoproteus,
e.g., Haemoproteus columbae Celli and San Felice of pigeons and doves,
transmitted by a louse fly, Pseudolynchia canariensis (Macq.); also quail
malaria caused by Haemoproteus lophortyx O'Roke carried by Lynchia
hirusta Ferris, the louse fly of quail. (See Chapter XVIII.) A malaria-
like disease of ducks is caused by Leucocytozoon anatis Wickware and
is carried by a simuliid fly, Simulium venustum Say. (See p. 153.)
Simian malaria. Among the several species of the genus Plasmodium
peculiar to monkeys are P. knowlesi Sinton and Mulligan, P. cynomolgi
Mayer, P. kochi (Laveran), P. inui Halberstadler and Prowazek, and
P. brasilianum Gonder and Berenberg-Gossler. These plasmodia differ
but slightly from the human species, and when P. knowlesi is introduced
(as for paresis treatment) into the blood of man, a clinical vivax malaria
response obtains (Milan and Kusch 43 ). Anopheles quadrimaculatus Say
is reported to be highly susceptible to P. cynomolgi (Coggeshall, 1941).
Clark and Dunn,44 in Panama, were able to infect certain species of
Panamanian anophelines with this parasite.
Saurian and amphibian malaria. Plasmodia have long been known in
lizards of the family Iguannidae; among these are Plasmodium agamae
Wenyon, P. diploglossi Aragao and Neiva, P. giganteum Theiler., and P.
floridense Thompson and Huff. Bullfrogs and toads have recently been
shown to have plasmodial infections, viz.: P. catesbianca Fantham,
Porter, and Richardson in the former and P. bufonis F., P., and R. in
the latter.
Bats, squirrels, buffalo, antelopes, and many other animals are hosts
for plasmodial infections. There are practically unlimited opportunities
in this field of research.
B. YELLOW FEVER
Yellow fever derives its name from the jaundiced color of the skin,
mucouS membranes, and sclera, which usually develops about the third
or fourth day of the illness. The onset is very sudden. It begins with a
severe headache, backache, and fever, with frequent vomiting. The
vomit becomes very dark after two or three days because it contains
blood, whence the name "black vomit." In fatal cases death usually
]\10SQUlTOES AS VECTOnS OF DISEASE 233
occurs between the fifth and eighth day of the illness; if the patient
survives until the seventh day his chances for recovery are generally
said to be good. One attack confers lifelong immunity. The death rate
may reach as high as 50 per cent.
Carter 45 (1931, page 146) considers the epidemic in Yucatan,
Mexico, in 1648 as the first certainly recognizable (from its description)
epidemic of yellow fever. He also gives (page 190) 1649 as the date
for the earliest introduction of yellow fever into Havana, and on page
192 he refers to its disappearance from Cuba after 1655 and its rein-
troduction from Vera Cruz in 1761. From that date until 1901 it was
endemic in Havana with devastating results. The last major American
urban epidemic took place in Rio de Janeiro in 1928 and 1929 with 435
recorded deaths. For nearly three centuries yellow fever took an huge
annual toll of life. Outbreaks have occurred as far north as Quebec,
Canada, and as far south as Tocopilla, Chile. New York City had
epidemics of yellow fever many times, as did Philadelphia.
Walter Reed and yellow fever. Although Dr. Carlos Finlay (loc. cit.
p. 3) of Havana had quite early (1881) advanced a mosquito-transmis-
sion theory, and had carried on what we now know to have been in-
criminating experiments with nonimmunes, his theory was discredited
until renewed interest was given it by the work of the United States
Army Yellow Fever Commission headed by Major Walter Reed in 1900.
Reed'6 and his colleagues made a preliminary report in which they
state: "Since we here, for the first time, record a case in which a typical
attack of yellow fever has followed the bite of an infected mosquito,
within the usual period of incubation of the disease, and in which other
sources of infection can be excluded, we feel confident that the publica-
tion of these observations must excite renewed interest in the mosquito
theory of the propagation of yellow fever, as first proposed by Finlay."
The United States Senate Document No. 822 (Jan. 27, 1911) is con-
cerned with yellow fever and contains a compilation of various publica-
tions by the Commission as well as others. In this document, McCaw
gives the following account of the work of the Commission:
In June, 1900, Major Reed was sent to Cuba as president of a board to

study the infectious diseases of the country, but more especially yellow fever.
Associated with him were Acting Asst. Surgs. James Carroll, Jesse W. Lazear,
and A. Agramonte.
At this time the American authorities in Cuba had for a year and a half
endeavored to diminish the disease and mortality of the Cuban towns, by
general sanitary work, but while the health of the population showed distinct
improvement and the mortality had greatly diminished, yellow fever apparently
had been entirely unaffected by these measures. In fact, owing to the large
number of nonimmune foreigners, the disease was more frequent than usual
in Habana and in Quemados near the camp of American troops, and many
valuable lives of American officers and soldiers had been lost.
Reed was convinced from the first that general sanitary measures alone
would not check the disease but that its transmission was probably due to an
insect.
In June, July, and August, 1900, the commission gave their entire atten-
tion to the bacteriological study of the blood of yellow-fever patients and the
post-mortem examinations of the organs of those dying with the disease. In
24 cases where the blood was repeatedly examined, as well as in 11 carefully
studied autopsies, Bacillus icteroides was not discovered, nor was there any
indication of the presence in the blood of a specific cause of the disease.
Application was made to General Leonard Wood, the military governor
of Cuba, for permission to conduct experiments on nonimmune persons, and
a liberal sum of money requested for the purpose of rewarding volunteers
who would submit themselves to the experiment.
It was indeed fortunate that the military governor of Cuba was a man who
by his breadth of mind and special scientific training could readily appreciate
the arguments of Major Reed as to the value of the proposed work.
Money and full authority to proceed were promptly granted, and to the
everlasting glory of the American soldier, volunteers from the Army offered
themselves for experiment in plenty and with the utmost fearlessness.
Before the arrangements were entirely completed, Dr. Carroll, a member
of the commission, allowed himself to be bitten by a mosquito that 12 days
previously had filled itself with the blood of a yellow-fever patient. He suffered
from a very severe attack, and his was the first experimental case. Dr. Lazear
also experimented on himself at the same time, but was not infected. Some
days later, while in the yellow-fever ward, he was bitten by a mosquito and
noted the fact carefully. He acqUired the disease in its most terrible form and
died a martyr to science and a true hero.
No other fatality occurred among the brave men who, in the course of the
experiments, willingly exposed themselves to the infection of the dreaded dis-
ease.
A camp was especially constructed for the experiments about 4 miles from
Habana, christened Camp Lazear in honor of the dead comrade. The inmates
of the camp were put into most rigid quarantine and ample time was allowed
to eliminate any possibility of the disease being brought in from Habana.
The personnel consisted of three nurses and nine nonimmunes, all in the
military service, and included two physicians.
From time to time Spanish immigrants, newly arrived, were brought in
directly from the immigrant station; a person not known to be immune was
not allowed to leave camp, or if he did was forbidden to return.
The most complete record was kept of the health of every man to be ex-
perimented upon, thus eliminating the possibility of any other disease than
yellow fever complicating the case.
The mosquitoes used were specially bred from the eggs and kept in a
building screened by wire netting. When an insect was wanted for an ex-
periment it was taken into a yellow-fever hospital and allowed to fill itself with
the blood of a patient; afterwards at varying intervals from the time of this
meal of blood it was purposely applied to nonimmunes in camp.
In December, 5 cases of the disease were developed as the result of such
applications; in January, 3, and in February, 2, making in all 10, exclusive of
the cases of Drs. Carroll and Lazear. Immediately upon the appearance of the
Rrst recognized symptoms of the disease, in anyone of these experimental
cases, the patient was taken from Camp Lazear to a yellow-fever hospital, 1
mile distant. Every person in camp was rigidly protected from accidental
mosquito bites, and not in a single instance did yellow fever develop in the
camp, except at the will of the experimenters.
The experiments were conducted at a season when there was the least
chance of naturally acquiring the disease, and the mosquitoes used were kept
active by maintaining them at a summer temperature.
A completely mosquito-proof building was divided into two compartments
by a wire-screen partition; infected insects were liberated on one side only. A
brave nonimmune entered and remained long enough to allow himself to be
bitten several times. He was attacked by yellow fever, while two susceptible
men in the other compartment did not acquire the disease, sleeping there 13
nights. This demonstrates in the simplest and most certain manner that the
infectiousness of the building was due only to the presence of the insects.
Every attempt was made to infect individuals by means of bedding,
clothes, and other articles that had been used and soiled by patients suffering
with virulent yellow fever.
Volunteers slept in the room with and handled the most filthy articles for
20 nights, but not a symptom of yellow fever was noted among them, nor was
their health in the slightest degree affected. Nevertheless they were not im-
mune to the disease, for some of them were afterwards purposely infected by
mosquito bites. This experiment indicates at once the uselessness of destroying
valuable property for fear of infection. Had the people of the United States
known this one fact 100 years ago, an enormous amount of money would have
been saved to householders.
Besides the experimental cases caused by mosquito bite, four nonimmunes
were infected by injecting blood drawn directly from the veins of yellow-fever
patients in the Rrst two days of the disease, thus demonstrating the presence of
an infectious agent in the blood at this early period of the attack.
Even the blood serum of a patient, passed through a bacteria-proof RIter,
was found to be capable of causing yellow fever in another person:"
The conclusions reached by the Commission follow:
1. The mosquito (C. fasciatus = Aedes aegypti) serves as the intermediate

host for the parasite of yellow fever.
2. Yellow fever is transmitted to the nonimmune individual by means of
the bite of the mosquito that has previously fed on the blood of those sick with
this disease.
S. An interval of about 12 days or more after contamination appears to be
necessary before the mosquito is capable of conveying the infection.
4. The bite of the mosquito at an earlier period after contamination does
not appear to confer any immunity against a subsequent attack.
5. Yellow fever can also be experimentally produced by the subcutaneous
injection of blood taken from the general circulation during the first and second
days of this disease.
6. An attack of yellow fever, produced by the bite of the mosquito, con-
fers immunity against the subsequent injection of the blood of an individual
suffering from the nonexperimental form of this disease.
7. The period of incubation in 13 cases of experimental yellow fever has
varied from 41 hours to 5 days and 17 hours.
8. Yellow fever is not conveyed by fomites, and hence disinfection of
articles of clothing, bedding, or merchandise, supposedly contaminated by
contact with those sick with this disease, is unnecessary.
9. A house may be said to be infected with yellow fever only when there
are present within its walls contaminated mosquitoes capable of conveying the
parasite of this disease.
10. The spread of yellow fever can be most effectually controlled by
measures directed to the destruction of mosquitoes and the protection of the
sick against the bites of these insects.
11. While the mode of propagation of yellow fever has now been definitely
determined, the specific cause of this disease remains to be discovered.
The following is an excerpt from a letter which Walter Reed wrote

to his wife in Baltimore late in the evening of December 31, 1900. The
letter was written from the Barracks in Quemados, Cuba: "Only ten
minutes of the old century remain. Here I have been sitting reading
that most wonderful book, 'La Roche on Yellow Fever' written in 1853.
Forty-seven years later it has been permitted to me and my assistants
to lift the impenetrable veil that has surrounded the causation of this
most dreadful pest of humanity and to put it on a rational and scientific
basis. I thank God that this has been accomplished during the latter
days of the old century. May its cure be wrought in the early days of the
new! The prayer that has been mine for twenty years, that I might be
permitted in some way at some time to do something to alleviate human
suffering has been granted! A thousand Happy New Years! . . . Hark,
there go the twenty-four buglers in concert, all sounding 'Taps' for the
old year." (Herrick. 47 )
Etiology. The search for the causal agent of yellow fever was carried
on most assiduously for many years and various discoveries were an-
nounced from time to time. Sanarelli in 1897 declared the organism to
be Bacillus icteroides; this was amply disproved by the United States
Army Yellow Fever Commission in 1900. Seidelin in 1909 described
Paraplasma flavigenum as the causal agent, and in 1919 Noguchi came
to the conclusion that a spirochete, Leptospira icteroides Noguchi, was
the cause of yellow fever. This turned out to be the cause of Weil's
disease or infectious jaundice. Yellow fever is now classed among the
virus diseases.
The virus is believed to be present in the circulation only during
the first three days of the illness. Aedes aegypti (Linn.) reared from
eggs in the laboratory and fed by Stokes, Bauer, and Hudson (lac. cit.
p. 4) on infected monkeys on the first or second day of the fever, in-
variably became infective. They found that the mosquitoes were in-
fective 16 days after feeding on an infected animal and remained so until
death, one mosquito producing a fatal infection in two monkeys 85 and
92 days after the original infective meal.
Mechanism of transmission. The virus of yellow fever is not only
present in the circulating blood during the first three or four days of the
illness, but also at least six hours or possibly somewhat longer, before
the onset of the fever. During this short period the vector mosquito,
Aedes aegypti (Linn.), may at one feeding ingest several thousand in-
fective doses of the virus. The mosquito then becomes a dangerous
vector for the rest of its life, which may be over 200 days. The extrinsic
incubation period normally under field temperatures takes about 12
days (Carter, lac. cit.) but may be considerably shortened by increased
temperature, i.e., to 4 days at 98° F or extended to 18 days at 70° F
( Da vis 48 ). After a susceptible person· has been bitten and receives an
infectious dose from an infected mosquito an incubation period of three
or four days follows before the symptoms of the disease appear. Now
again another source of infection is ready for other aegypti vectors re-
stricted as before to the short period of three or four days. An epidemic
may quickly develop in the presence of hordes of this rapidly multiply-
ing species of domestic mosquito.
Summary of recent discoveries. Soper et al. 49 (1943) state that "when
the medical historian of the future comes to write the chapter on yellow
fever, he will be obliged to devote considerable space to the develop-
ments of the decade and a half from 1926 to 1940. This short period
showed how impossible of fulfillment was the dream of final eradication
of yellow fever, but it saw great advances in our understanding of the
etiology, epidemiology, and prophylaxis of the disease, including the
discovery of animal susceptibility; the rediscovery of the virus origin;
the demonstration that mosquitoes other than Aedes aegypti can and do
transmit the virus; the development of the protection test for determin-
ing immunity; the organization of viscerotomy for the diagnosis of un-
suspected fatal cases; the proof that unrecognized yellow fever has been
widespread in large silent endemic areas of South America and Africa;
the demonstration that the disease exists in many countries of South
America as one of jungle animals, independent of the distribution of
Aedes aegypti and of man; and finally the modification of the yellow
fever virus in such a way as to make mass vaccination practical. These
outstanding developments have tended to overshadow the more prosaic
improvements in the organization of measures against the aegypti mos-
quito in Brazil which have transformed expensive temporary aegypti
reduction campaigns for the eradication of yellow fever into economical
permanent services for the species eradication of Aedes aegypti itself
from the infested areas."
Aedes vittatus (Bigot) has been reported as responsible for a recent
( 1940) epidemic of yellow fever of modified virulence in the Anglo-
Egyptian Sudan. This species is said to breed abundantly in rock pools
as well as tree holes. Filling such pools with sand is recommended as a
partial means of control. (Horgan, Proc. Amer. Mosq. Control Assoc.,
Feb., 1949.)
Jungle yellow fever. Yellow fever is generally regarded as an urban
or house disease transmitted solely by Aedes aegypti (Linn.), a domestic
mosquito which breeds largely in artificial containers in and about
human habitations. The epidemiological pattern of urban yellow fever
is relatively simple; namely, "man to mosquito to man." Control seemed
simple enough with meticulous inspection and careful application of
proper control measures. The Rockefeller Foundation reported 50 that
prior to 1929 the belief was expressed that yellow fever was not only
fast disappearing as a human menace but that it had been practically
eliminated. ''In ... 1925 only three cases of yellow fever were reported
from the entire Western Hemisphere; in the 11 months following April,
1927, no cases were reported, and it was assumed that the battle, which
had cost the lives of research workers and millions of dollars, was
practically won. Then almost without warning, the South American
Jungle struck back (Soper and associates, 51 1933), and in a few years'
time the epidemiological strategy of the battle had to be completely
altered." It is pointed out that vast areas of the hinterland of both South
America and Africa are endemic centers of yellow fever. Burke 52 (1937)
studied an epidemic involving 201 cases of yellow fever of the jungle
type in the absence of Aedes aegypti (Linn.) on the Planalto of Matto.
Grosso, Brazil, during the seasons of 1934 and 1935. He reports that the
identity of the disease was definitely established, the only difference
being in the conditions under which infection occurs. "The paucity of
human population in the infected district and the scattered distribution
of cases in both time and space, together with the isolated circumstances
attending many cases argue against man being the only vertebrate host
involved. The sera from five Cebus monkeys captured for this study in
known infected districts all gave positive protection test results, indicat-
ing immunity naturally acquired in the jungle. All available evidence
points to infection occurring either in clearings next to uncleared jungle
or in the jungle itself, especially during working hours." The Rockefeller
Foundation (loc. cit.) points out that jungle yellow fever must be con-
sidered as a possible permanent source of virus for the reinfection of
cities and towns where high densities of Aedes aegypti (Linn.) mos-
quitoes are tolerated. In jungle yellow fever the epidemiological pat-
tern appears to be "monkey to mosquito to monkey." Man becomes in-
fected when he goes into the forest and is bitten by infected mosquitoes
and he brings the virus back with him when he returns to his com-
munity. There he may encounter "aegypti" mosquitoes and an epidemic
of yellow fever may ensue. Students concerned with jungle yellow fever
will need to consult "An epidemiological study of jungle yellow fever
in an endemic area in Brazil," by Taylor, Laemmert, et al. 53
Aedes Ieucocelaenus (D. and S.) and Haemagogus capricornii
(Lutz), both forest-inhabiting mosquitoes, were incriminated by Shan-
non, Whitman, and Franca'" during the 1938 outbreak of yellow fever
in the State of Rio de Janeiro, Brazil. The presence of the virus in these
mosquitoes as well as in certain Sabathine species taken in the Brazilian
jungle was demonstrated. Subsequent investigations indicate that
Haemagogus capricornii was probably H. spegazzinii (see below).
The 1947 report of the International Health Division of the Rocke-
feller Foundation states that Haemagogus spegazzinii Brethes is by all
odds the commonest and most widely distributed species of the genus
Haemagogus in Brazil and is generally found in the endemic areas of
jungle yellow fever in Colombia. As to reservoir jungle animals the
above-mentioned report states that of the various kinds of animals tested
only the howler monkey (Cebus) circulated virus in abundance.
Several mosquitoes have been incriminated as vectors of yellow fever
in the tropical jungles of East Africa, among these Aedes simpsoni Theo.
and Aedes africanus Theo., both widely distributed in endemic jungle
areas. The latter species is apparently highly important because of its
recently discovered presence both in the forest and in the plantations.
There is thus presented the possibility of introducing yellow fever to
human populations directly from endemic jungle foci through the agency
of one species of mosquito. 55
Other mosquito vectors. Prior to 1928 (see Chapter I) no experi-
mental animal (other than man) was known to be susceptible to yellow
fever. With the discovery that the Indian monkey, Pithecus rhesus, was
susceptible as well as a score of other species of monkeys, and particu-
larly with the work of Sawyer on white mice, experimentation with
various species of mosquitoes grew apace. (See Soper et al., 1933, loco
cit. )
There are evidently many species of jungle-inhabiting vertebrates

which harbor the virus of yellow fever, more importantly perhaps several
species of monkeys. Hedgehogs are highly susceptible animals and are
of great value in laboratory investigations. Several species of mosquitoes
other than those here referred to lend themselves well to 'laboratory
transmission by the bite; among them is a common and widespread
nearctic species Aedes triseriatus (Say) 5S in which the monkey, Macacus
rhesus, was used as the host. It is of interest to note that the more im-
portant species of mosquitoes incriminated as probable or proved vectors
of the virus of jungle yellow fever are tree-hole breeders thus sharing
(perhaps roughly) the same ecological niche with their simian hosts.
Besides those species mentioned, many others (probably more than
30) have given positive results by the bite in laboratory tests; among
these are Aedes scapularis (Rondani), A. fluviatilis (Lutz), A. luteo-
cephalus Newst., A. stokesi Evans, A. albopictus (Skuse), Culex tha-
lassius Theob., Eretmapodites chrysogaster Graham, Mansonia africana
(Theob.), Haemagogus equinus Theobald, and H. splendens Williston.
C. DENGUE FEVER
Dengue fever, also known as breakbone fever or dandy fever, is a

widespread disease of tropical and subtropical regions, but it also
occurs in temperate climates where suitable vectors are present. In an
epidemiC which occurred in the State of Texas in 1922, originating in
Galveston during the second week of June, spreading later to other
parts of the state, beginning to abate late in September, and ending
in late autumn, the number of cases of the disease was estimated at be-
tween 500,000 and 600,000. 57
The disease is characterized by its sudden attack, severe rheumatic
pains in the joints and limbs, intense headache, backache, and high
fever; a remission of two to three days follows the Rrst attack of three
days; the second attack lasts usually but a day and is accompanied by a
rapidly spreading rash. The "saddle back" type of fever, though quite.
common, is not constant. The entire course may be run in Rve to six
days. Although the disease is of much importance because of its de-
bilitating effects and prolonged convalescence, the death rate is very
low. It is caused by a Rltrable virus, as shown by Ashburn and Craig,
1907. 58 The virus is present in the peripheral circulation during the ini-
tial stages of the disease, i.e., the Rrst three to Rve days, also probably
for some hours previous to the onset; hence the mosquito vector must
bite the patient during this short period of time in order to become
infected.
Mosquito transmission. Transmission experiments conducted by
Chandler and Rice (loc. cit.) with Aedes aegypti (Linn.) were success-
ful in four out of six cases, the mosquitoes having fed on patients in
the second to fifth days of the disease.
Simmons, St. John, and Reynolds 55 (1931) found that all lots of
Aedes aegypti (Linn.) that fed on blood from experimental cases of
dengue during the first 48 hours of the disease became infected. The
mosquito is able to transmit the infection after an incubation period of
11 days, though Chandler and Rice (lac. cit.) state that mosquitoes suc-
ceeded in transmitting the disease in from 24 to 96 hours. Infected
mosquitoes remain infected as long as they live. The disease has been
transmitted 174 days after infection, and Aedes aegypti (Linn.) has
been kept alive for seven months. (Simmons et al., lac. cit.)
Graham 60 (1902) was the first to demonstrate that mosquitoes trans-
mit dengue by the bite.
The incubation period in experimentally infected cases varies from
three to eight days. Simmons et al. give the average incubation period
at 5.66 days; average duration of the fever, 4.8 days. The virus is not
transmitted from infected female Aedes aegypti (Linn.) through the
egg to the offspring, neither does contamination of the skin by dengue
virus from crushed Aedes aegypti (Linn.) result in infection. A brief
immunity is conferred by an attack of dengue. (A description of A.
aegypti, and of its breeding habits will be found in the preceding chap-
ter. )
The investigations of Simmons, St. John, and Reynolds prove that
Aedes albopictus (Skuse) is an important vector of dengue, also that
Culex quinquefasciatus (= fatigans) Wied., hitherto so regarded is not
a vector.
Aedes albopictus ( Skuse) has the mesonotum ornamented with a
central silvery stripe; white scales of the thoracic pleura are arranged
in irregular patches. Its breeding habits under urban conditions are very
similar to those of A. aegypti, i.e., the larvae live in fairly clear water
in artificial containers; however, when away from human habitations
the larvae occur in tree holes, bamboo stalks, plant axils, and stone
cavities in the forests (up to 2,000 feet in Hawaii). It is particularly
prevalent in depreSSions in monuments in Japanese cemeteries, as well
as in flower containers. The species is found in Oriental regions, Mada-
gascar, New Guinea, north Australia, and the Philippine and Hawaiian
Islands.
Aedes scutellaris (Walker) resembles A. albopictus (Skuse) closely;
the mesonotum has a central silvery stripe, white scales of thoracic
pleura arranged in two wavy parallel lines. The distribution of the
species overlaps that of A. albopictus in Micronesia and in Polynesia
except Hawaii. It is reported to be a vector of dengue in New Guinea.
D. ARTHROPOD-BORNE VIRUS ENCEPHALITIDES

Equine encephalomyelitis is a disease of wide distribution; the causal
agent is a virus with neurotropic properties, as shown by Meyer, Haring,
and Howitt,61 who describe symptoms as follows.
Preceding the onset of symptoms which attract attention, the temperature
0
may be found to vary from 103 F to 107° F. Not infrequently when the
horse shows signs of drooping of the head, sleepiness, and circling motion or
other psychic and motoric disturbances, the body temperature may be normal.
The pulse and respiration are usually accelerated. Quite often the animal rests
against the wall or corner and may show backward and sideways motions.
Muscular twitchings are quite common. Many of the horses are down on the
second or third day and mayor may not get up when pressed to do so.
Paresis of the lips and drooling are frequently noted. Mastication and swallow-
ing mayor may not be impaired, but grinding of the teeth is quite regularly
observed. The conjunctiva is always infected and frequently icteric or gray-
ish and studded with petechiae or ecchymoses. In the mild cases which were
able to rise, recovery was as a rule uneventful but about half were so severe
that they terminated fatally in 3 to 8 days or were destroyed for humane
reasons.
Believing that bloodsucking insects might be instrumental in the
transmission of the virus, the author conducted tests during the late
summer of 1932 (Herms, Wheeler, and Herms 1934).62 In these tests
horseflies, Tabanus punctifer O. S., and horn flies, Haematobia serrata
Desv., were used-all with negative results.
Early in 1933 Kelser63 proved that the disease can be transmitted by
Aedes aegypti (Linn.) not only from infected guinea pigs to normal
guinea pigs but also to a horse which contracted the disease and died
within five days after the onset of symptoms. Blood drawn from the
horse at the height of the fever and injected into a guinea pig produced
the disease, and mosquitoes fed on the horse during the period of high
temperature and subsequently fed on a normal guinea pig likewise pro-
duced the disease. The largest percentage of deaths among the guinea
pigs bitten by infected mosquitoes occurred on the sixth day following
the infective mosquito bite. The mosquitoes were found to be capable
of producing the disease as early as six days and remained infectious
for at least 36 days. KeIser pointed out that it is possible that the mos-
quitoes, when once infected, may remain infectious the rest of their lives
as is the case in yellow fever and dengue.
Following the work of KeIser the author conducted further tests,
using Anopheles maculipennis Meigen (A. freeborni Aitken) and Aedes
dorsalis (Meigen), a salt-marsh mosquito, bred from larvae taken from a
salt marsh. Again all results were negative (Herms, Wheeler, and Herms,
loco cit.). However, later tests (unpublished) made in October 1934
give evidence of six successful transmissions out of 29 (guinea' pig t~
guinea pig) secured with Aedes dorsalis (Meigen) bred from fresh-
water larvae. The elapsed time between initial feeding on an inoculated
pig and feeding on a normal pig was six days in four of the positive
cases and 12 days in the remaining two. From 4 to 50 mosquitoes were
used in each test. Investigations made by Madsen, Knowlton, and
Rowe 64 with A. dorsalis (Meigen) also indicate that this species may
be a vector of the virus; although most tests failed, 2.5 per cent of the
total trials (with guinea pigs) were positive, with incubation period
varying between 9 and 19 days. Tests made by these authors with Aedes
nigromaculis (Ludlow) gave 55.8 per cent positive transmissions, with
incubation period varying between 4 and 10 days. Merrill, Lacaillade,
and Ten Broeck65 have demonstrated in repeated tests that Aedes sollici-
tans (Walker), a common Atlantic coast salt-marsh mosquito, will trans-
mit both eastern and western strains of the virus from infected to nor-
mal guinea pigs. Transmission was obtained with the eastern virus 11
days after the initial feeding and at later periods. Aedes cantator (Coq.),
another salt-marsh breeder, was shown to transmit the eastern virus,
though less readily. These authors have demonstrated a 1,000- to 10,000-
fold increase of the virus within the bodies of both Aedes aegypti
( Linn.) (western strain virus) and A. sollicitans 'Walker) ( eastern
strain virus). They report 63 days as being the longest period during
which Aedes aegypti (Linn.) is capable of transmitting the western
strain. Madsen and Knowlton66 were successful in transmitting the west-
ern virus to guinea pigs by the bite of Aedes nigromaculis (Ludlow).
Simmons, Reynolds, and Corne1l67 demonstrated that Aedes albopictus
( Skuse) could transmit the western virus. Kelser 68 reported studies def-
initely proving the ability of Aedes taeniorhynchus (Wied.) to trans-
mit the "western" type of equine encephalomyelitis from guinea pig to
guinea pig.
In 1940 Kitselman and Grundmann 69 reported the isolation of equine
encephalomyelitis virus from naturally infected conenose bugs, Triatoma
sanguisuga Le Conte. Syverton and Berry70 (1936) were able to trans-
mit the western strain of the virus to a ground squirrel, Citellus richard-
soni (Sabine) through the bite of ticks, Dermacentor andersoni Stiles.
The animal died five days after the ticks, which had fed on an inocu-
lated guinea pig, were placed on the animal. Tyzzer, Sellards, and Ben-
nett n in 1938 reported the natural occurrence of the infection in the
ring-necked pheasant. Man was reported susceptible to the infection in
1938 by Howitt. 72
Reeves (William C.), who later joined Hammon (W. McD.) of the
Hooper Foundation for Medical Research in the study of the arthro-
podan encephalitides, began his research work under the direction of
the author in 1940 and completed his work for the doctorate in October,
1943, the title of his thesis being "Culex tarsalis and Other Mosquitoes
as Vectors of the Virus Encephalitides of Western North America." In
1941 Hammon, Reeves, Brookman, and IzumF" reported the isolation
of the viruses of both western and St. Louis encephalitis from Culex
tarsalis mosquitoes in the Yakima Valley of the State of Washington. It
should be recalled at this point that mosquitoes were strongly suspected
as the vectors in the 1933 St. Louis outbreak, in which there were over
1,000 cases with about 200 deaths. In 1942 Hammon and Reeves 74
reported, "We can safely conclude that this species [Culex tarsalis Coq.
referred to above] is a natural vector ... in respect to St. Louis en-
cephalitis virus . . . and in addition ... that the chicken may serve
as a satisfactory reservoir of the virus."
Hammon,15 in a paper read at the 1948 meeting of the California
Mosquito Contr01· Conference, points out repeated failure to find virus
in overwintering wild mosquitoes; a chronic or latent stage of the in-
fection was not observed in chickens or other vertebrates, and thus no
true reservoir was found. Then came announcement by Smith 76 and
co-workers and by Sulkin 77 that these viruses had been found in chicken
mites, and furthermore that the virus was transmitted through the mite
eggs from generatfon to generation. Here, then, appears, to be a true
reservoir for the infection. In 1947 Reeves, Hammon, Furman, McClure,
and Brookman78 announced the recovery of western equine encepha-
lomyelitis virus from wild bird mites Liponyssus sylviarum (Canestrini
and Fanzago ). The wild birds were the yellow-headed blackbird, Xantho-
cephalus xanthocephalus (Bonaparte). This discovery is significant in
that it brings a wild vertebrate reservoir animal (a bird) into the en-
cephalitis picture. Much more work needs to be done with mites in
relation to the virus in order to elucidate their role as vectors.
Recently Hammon 79 (1948) has pointed out that the hypothetical,
cycle now becomes a more complicated one. "Mites infected previously,
congenitally or by feeding on an infected bird, may initiate infection in
nestling birds or in newly hatched fowl in the spring. These birds in
turn are fed on by mosquitoes. After the essential extrinsic incubation
period, these mosquitoes may infect other birds. Several such cycles in
mosquitoes and birds will build up the number of infected mosquitoes
and infection may then appear in men or horses. Infection is usually
seen first in horses, and later in men, undoubtedly because of the greater
number of mosquitoes having access to the large, unprotected domestic
animals, and because of the mosquitoes' feeding preference."
Japanese "B" virus. According to Hammon so the most severe epidemic
of encephalitis ever described occurred in Tokyo in the summer of 1924,
with over 6,000 cases resulting in 3,797 deaths. In the Japanese epidemic
of 1948 there were about 8,000 cases, with a 30 per cent mortality. (La-
Casse, Froc. Amer. Mosq. Control Assoc., Feb., 1949.) That this virus
also occurs in horses was shown by Hodes, and Peck81 in Okinawa by
complement-fixing and neutralizing antibodies. Mitamura, J.,82 and as-
sociates report a small epidemic of encephalitis in the prefecture of
Okayama in 1938, which lasted about 40 days. The chief species of
mosquitoes taken were Culex pipiens var. pallens Coq., Culex tritaenior-
hynchus Giles, and Anopheles hyrcanus var. sinensis Wied. Females of
both species of Culex taken between July 25 and August 24 infected
mice on which they fed. It is also reported that 12 mice became infected
out of 511 bitten by female Culex pipiens pallens that had developed in
the laboratory from larvae and pupae taken in nature, hence had had
no opportunity of acquiring the virus in the adult stages. Forty species
of mosquitoes are reported for Japan. Culex annulirostris Skuse has
recently been reported by Hurlbut and Thomas as a potential vector in
the Caroline Islands.
Among other recently discovered mosquito-borne encephalitis viruses
are the following: West Nile, Semliki Forest, Rift Valley, and Mengo
strains.
E. FILARIASIS
Filariasis. Infestation with nematode worms belonging to the family

Filariidae is known as filariasis. The larval worms of this family are
commonly known as microfilariae and occur in the circulatory and lym-
phatic systems, connective tissue layers, and serous cavities of the verte-
brate hosts.
Bancroft's filaria, Wuchereria bancrofti (Cobbold), is a widely dis-
tributed parasite of man, being indigenous to Polynesia and Melanesia,
where advanced elephantiasis (see p. 247) is frequently encountered;
the disease is found in many tropical and subtropical areas of the globe,
such as central Africa, southeast coastal China, and the East Indies. In
the United States only one endemic area has been known in the past,
a small area including Charleston, South Carolina. 83 Puerto Rico is re-
ported to be one of the best-known endemic areas of filariasis in the
Western Hemisphere, although the disease is not regarded as a major
health problem there. 84 The infection also exists in the northeast coastal
portions of South America.
The sheathed microfilariae, first noted by Bancroft in 1876, measure
about .3 mm in length and from 7.5J..1. to 10,u in diameter (Fig. 81). As
observed by Manson in 1877-78, they occur in the peripheral blood,
particularly at night. In the daytime most of the microfilariae are con-
centrated in the pulmonary vessels, the capillaries of the heart, and
other visceral organs. This nocturnal periodicity85 enables night-flying
mosquitoes such as Culex quinquefasciatus Wied. to suck up the para-
sites while biting. Manson called attention to this adaptation and be-
lieved that mosquitoes served as "nurses" for the filariae, and that when
the mosquitoes dropped into water and disintegrated the organisms
were liberated and infection of human beings resulted from drinking
infected water.
Life cycle. The fully grown, very slender, sexually mature worms,
the females reaching a length of 80 to 100 mm and the males about 40
mm, are often found (post-mortem) in tangles in nodular dilations of
the distal lymphatics, in lymphocytic varices, and in the glands them-
FIG. 81. Wuchereria bancrofti, in

human blood. X 333.
selves or even in the thoracic duct. Here myriads of minute micro-

filariae are born, each in a sheath or sac. Although many remain in the
lymph, others following a normal circulatory course will reach the pe-
ripheral circulation. Brown 86 reports an infection of 23,240 microRlariae
per ml of blood in a Virgin Islander (except for a slight general glan-
dular enlargement, there were no symptoms referable to this infection).
In the so-called classic filariasis the microfl.lariae manifest a nocturnal
periodicity (fl.lariae sanguinis hominis nocturna), i.e., they are present
in the peripheral blood of infected persons prinCipally during the hours
from 10 P.M. to 2 A.M. There is, however, a nonperiodic form of
bancrofti in the South PaciRc area. In either instance the microRlariae
are available to appropriate species of mosquitoes, i.e., anthropophilous
night feeders in the case of "filaria nocturna," and day biters in the case
of "fl.laria diurna." Many species of mosquitoes belonging to four com-
mon genera, Culex, Aedes, Mansonia, and Anopheles, have been incrim-
inated as vectors, some species being more readily infective than others.
The more numerous the individual mosquito(\s in a large population of
infected human beings the more dangerous is the situation. It must be
remembered that this is an infection referred to earlier in which the
pathogen does not multiply within the body Qf the vector; the mosquito
must imbibe one microfilaria from the blood of an infected person for
every filarial worm transmitted to a new Vlctim. Having successfully
reached the stomach of the mosquito, the tiny worm now loses its sac-
like sheath and in a matter of hours forces its way through the stomach
wall and works its way through the thoracic muscles, becoming some-
what "sausage-shaped" by the fourth or fifth day. There are two molts,
and by the end of the twelfth to fourteenth day the digestive tract has
developed, and at least some infective larvtte, now about 1.5 mm in
length, settle in the region of the head an<:]_ labium of the mosquito
(many larvae are lost in the abdomen) awaiting an opportunity to slip
down the salivary channel when the mosql.lito bites. The number of
larvae per mosquito successfully reaching this stage is evidently quite
smarr (heavy infections may kiff the mosquito); Hu and Yen S7 report an
average of 3.9.
When the mosquito bites, the larvae qui(~kly slip down the labium
and escape from the labellum, coming to lie upon the warm skin of the
person bitten. The larvae now penetrate tbe skin, enter the lymph
stream, and are carried to the lymphatics, as mentioned in the begin-
ning, where development to sexual maturity is accomplished in a matter
of months; from 12 to 18 months is reported probable from the time of
infection to the maturity and appearance of q new generation of micro-
filariae in the peripheral blood.
Asymptomatic filariasis. In endemic areas a considerable percentage
of the population are infected with the parasite, as shown by blood
smear examination: thus M urray S8 reported 19.1 per cent positive for
microfilaria in a survey of 5,l44 Samoans. Btown (lac. cit.) states that
on physical examination the infected persoll, though unaware of the
infection, may exhibit a general glandular enlargement especially of the
inguinal lymph glands.
Inflammatory filariasis. The author has Observed cases of so-called
"mumu" in which there were recurrent attacks of inHammatory filariasis,
many months after exposure to infection in th!:J South Pacific area. Brown
(lac. cit.) suggests that these inHammatory reactions "may be an allergiC
phenomenon due to a sensitivity to the procl.ucts of the worms or to a
superimposed bacterial infection, usually streptococci."
Obstructive filariasis (elephantiasis). The obstructive types of filaria-
sis (grossly enlarged scrotum, vulvae, breasts, or legs) usually follow
years after the original long exposure to mosquito bites in endemic areas
and are the exception rather than the rule. Microfilariae are frequently
absent from the blood, although Murray (loc. cit.) states that he found
that, in American Samoa, persons afflicted with elephantiasis were much
more likely to have microfilariae than those without this condition.
Brug's filaria, Wuchereria malayi Brug, the causal organism of
"filariasis malayi" is very similar to that of "Bancroft's filariasis." It oc-
curs in India, Ceylon, Thailand, the East Indies, the Philippines, and
New Guinea. In some areas, such as portions of coastal China and some
islands of the East Indies, the two Wuchereria infections overlap; the
behavior of the worms in the mosquito vector, as well as in man, and
the effects on the human host are very similar. W. malayi, which is
essentially nocturnal in periodicity, differs from W. bancrofti only in
minor details. The morphology of the sheathed microfilariae differs
chiefly in the cephalic space, which is twice as long as broad, and the
posterior extremity of the worm has a slight bulb at the tip with two
minute terminal nuclei.
While there is little difference in the causal organisms, it is impor-
tant to note that the principal vector mosquitoes of W. malayi are mem-
bers of the Mansonia group in which the wrigglers attach themselves to
submerged aquatic plants such as Pistia and Eichhornia, as described
in a previous chapter. Mansonia (Mansonoides) annulifera Theobald
is considered to be the principal vector, although several other related
species as well as certain anophelines, e.g., Anopheles hyrcanus sinensis
Wiedemann, are believed to be implicated.
Mosquito species involved. Throughout the filarial belt of the world
(between 30° Nand 32° S latitude) some 60 species of mosquitoes be-
longing to half a dozen genera have been implicated in greater or lesser
degree as vectors of the infection. Culex qUinquefasciatus Say, a widely
distributed tropical and subtropical species, heads practically all lists
of vectors; it is, however, Significant that in the Samoan area experimen-
tal evidence disclosed that only 29 per cent of C. quinquefasciatus
could be infected from a single blood meal, whereas 90 per cent of
Aedes pseudoscutellaris (Theob.) became infected; also, it is reported
that only one per cent of the former developed microfilariae to the in-
fective stage in contrast to 18 per cent in the Aedes species. In several
island groups of the South Pacific (among them Solomon, New Hebrides,
Fiji, and Samoa) a primary vector is Anopheles farauti Lav. which
often becomes tremendously infected and dies as a result; as many as
96 eight- to ten-day-old larvae have been observed (Reiber). Only 11
per cent of the total number of infected mosquitoes survived long
enough to transmit larvae. Under natural conditions this percentage
would no doubt be still lower.
In the China area Anopheles hyrcanus sinensis Wiedemann and Culex
pipiens pallens Coquillett, are proven vectors, as they are also in Japan
together with Aedes togoi (Theobald); in west Africa Anopheles gam-

biae Giles is listed as a vector (see above for filariasis malayi).
In testing the susceptibility of North American species of mosquitoes
to Wuchereria bancrofti (Bora Bora strain) Eyles and Most 89 found
that development to advanced or infective stage occurred in the fol-
lowing species: Culex pipiens Linn., C. quinquefasciatus Say, C. errati-
GUS (Dyar and Knab), C. salinarius Coq., Anopheles walkeri Theob.,
A. punctipennis (Say), Aedes triseriatus (Say), A. aegypti Linn., Aedes
atropalpus (Coq.), and Mansonia perturbans (Walker). In the following
species development did not proceed beyond the first larval stage:
Anopheles quadrimaculatus Say, Anopheles freeborni Aitken, Aedes tor-
mentor Dyar and Knab, and Psorophora ferox Humboldt.
With only three specimens of Culex tarsalis Coq. tested, Scott,
Richards, and Seaman 90 found that one of these developed infective
stages of W. bancrofti in the thorax, head, and proboscis.
Heartworm of dogs. Dirofilaria immitis (Leidy), a cosmopolitan
species, occurs in dogs (in over 30 per cent in some parts of the southern
United States) and occasionally in cats, as well as in various wild
carnivores. The adult worms, females measuring in length from 25 to
35 cm and the males 12 to 18 cm, invade the heart (right ventricle) and
pulmonary artery of the host, where they often form tangled knots
and may cause death as the result of embolism, asphyxia, and dilation
of the heart. This parasite has become widely distributed in the United
States.
Like Wuchereria bancrofti the microfilariae of D. immitis are found
in the blood stream and manifest a similar periodicity, except that more
of the latter appear to be present in the peripheral circulation during the
daytime (Hinman). The larvae of D. immitis are without sheaths. Culex
pipiens Linn., Aedes aegypti (Linn.), as well as a number of other
species of mosquitoes, are known to be vectors. Development in the
mosquito is similar to that of Wuchereria as described above. Sexual
maturity of the worms in the body of the dog is said to require from 5
to 9 months. Short-haired dogs are believed to be more susceptible to
. heartworms than long-haired dogs, probably because they are less pro-
tected against mosquito bites. To prevent infection in endemic areas,
dogs must be protected from mosquitoes, patticularly at night.
Other filarial worms. Among other species belonging to the family
Filariidae are Mansonella ozzardi (Manson), said to be nonpathogenic
to man, transmitted by Aedes aegypti (Linn. ); Acanthoceilonema
perstans (Manson), transmitted by Culicoides austeni Carter, Ingram,
and MacSe (see Chapter X); Loa loa (Cobbold), causing calabar swell-
ings, transmitted by Chrysops dimidiata v. d. Wulp (see Chapter XIV);
Onchocerca volvulus Leuckart, transmitted by Simulium damnosum
Theo., Simulium ochraceum Walker, and other species of black flies (see
Chapter X).
F. RABBIT MYXOMATOSIS
Rabbit myxomatosis, also known as "big head" of rabbits, is caused
by a virus closely related serologically and immunologically to the virus
of rabbit fibroma (Shope). 91 Experimental observations by Philip"2 on
the latter virus indicate that the infection can be transmitted mechani-
cally in the laboratory by several species of conenose bugs (Family
Reduviidae), e.g., Triatoma protracta, Triatoma infestans, and Rhodnius
prolixus. It was presumed that the mechanical transfer of the infection
in interrupted feeding tests was by the contaminated mouth parts of
the bugs.
The virus of rabbit myxoma produces lesions in the skin, lymph
glands, tunica vaginalis, testicle, spleen, and lungs. The disease runs a
rapid course, and death ensues within a week or two. Rabbits are the
only known susceptible hosts. Recently Aragao"" (1'04;J) has reported
successful transmission of the virus by mosquitoes, Aedes scapularis
(Rondani) and A. aegypti (Linn,). Here, as in the tests made by Philip
with rabbit fibroma, the transmission of the virus was thought to be
mechanical; i.e., only the proboscides of the mosquitoes were contami-
nated. The infected mosquitoes were able to transmit the virus two or
three times up to 17 days after the infecting blood meal. Other mosqui-
toes as well as other bloodsucking arthropods have been incriminated in
transmission.
C. FOWL-POX
Fowl-pox, an important virus disease of poultry, while spread in
various ways, such as contact between diseased and healthy birds, may
also be spread, according to Brody,24 by Aedes stimulans (Walker) by
intermittent feeding, harboring the virus in or on its body for at least
two days following an infective meal, and by Aedes degypti (Linn.),
which can definitely transmit the virus more than once during its life.
The latter species is able to transmit the disease within one hour after
an infective meal and continues to be infective for at least about 40 days.
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80. ---, 1945. "The encephalitides of virus origin, with special reference
to those of North America," Clinics, 4:485-503 (with 57 references) .
81. Hodes, Horace L.; Thomas, Lewis; and Peck, John L.; 1946. "Com-
plement-fixing and neutralizing antibodies against Japanese B virus in the
sera of Okinawan horses," Science, 103:357-59.
82. Mitamura, T.; Kitaoka, M.; Watanabe, S.; Hosoi, T.; Tenjiu, S.; Seki,
0.; Nagahata, K.; Jo, K.; and Shimizu, M.; 1939. "Weitere Untersuchungen
tiber die Uebertragung der japanischen epidemischen Enzephalitis durch
Mticken." Tr. Soc. Path. Japan, 29:92-105.
83. Francis, Edward, 1919. Filariasis in Southern United States. U. S.
Public Health Service, Hygienic Lab. Bull., no. 117. 34 pp., IX plates.
84. Tampi, M. K., 1931. "A study of filariasis in Porto Rico," Porto Rico
J. Pub. Health & Trop. Med., 6:435-41.
85. Hinman, E. Harold, 1936. "Attempted reversal of filarial periodicity
in Dirofilaria immitis," Proc. Soc. Exper. Biol. & Med., 33:524-27.
86. Brown, H. W., 1945. "Current problems in filariasis," Am. J. Pub.
Health, 35:607-13.
87. Hu, Stephen M. K., and Yen, Chia-Hsien, 1933. "Studies on the
susceptibility of Culex pipiens Linn. var pallens Coq. to experimental infec-
tion with Wuchereria bancrofti Cobbold in Shanghai area," Chinese M. J.,
47:1359-66.
88. Murray, William D., 1948. "Filariasis studies in American Samoa,"
Naval Med. Bull., 48:327-41.
89. Eyles, Don E., and Most, Harry, 1947. "Infectivity of Pacific Island
Wuchereria bancrofti to mosquitoes of the United States," Am. ]. Trop. Med.,
27:211-20.
90. Scott, O. K.; Richards, C. S.; and Seaman, E. A.; 1945. "Experimental
infection of Southern California mosquitoes with Wuchereria bancrofti," J.
Parasitol., 31 :195-97.
91. Shope, R. E., 1932. "A filterable virus causing a tumor-like condition
in rabbits and its relationship to Virus myxomatosum," J. Exper. Med., 56:803-
22.
92. Philip, Cornelius B., 1942. "Mechanical transmission of rabbit fibroma
(Shope) by certain haemophagous bugs," ]. Parasitol. 28:395-98.
93. Aragao, H. B., 1943. "0 virus do mixoma coelho do mato (Sylvilagus)
sua transmissao pe los Aedes scapularis e aegypti," Mem. Inst. Oswaldo Cruz,
38:93-99.
94. Brody, Arthur L., 1936. The Transmission of Fowl-Pox. Cornell Univ.
Agric. Exp. Sta. Memoir no. 195. 37 pp.
CHAPTER XIII
._.
MOSQUITO ABATEMENT
Historical. Ancient Greeks and Romans believed there was some

etiologic connection between intermittent fevers and swamps, hence the
use of drainage methods in Greece and especially Rome to overcome
the "poisoning effects" of stagnant water. Lancisi, the greatest Italian
physician of his time, published a treatise on swamp fevers in 1717,
according to Russell, in which he suggests that since malaria disappeared
after drainage, the poison from the marshes was pOSSibly transmitted by
mosquitoes which also came from the swamps.
The use of larvicides dates back to about 1793 (Russell, et al., loco cit.,
p. 215) when oil (probably whale oil) was used in Philadelphia on rain
barrels to kill mosquito larvae.
L. O. Howard/ for many years (1894-1924) chief of the United
States Bureau of Entomology, records "an experiment against mosqui-
toes" which he conducted in 1892. He "sprinkled four ounces of coal oil
upon the surface of the pond" which "contained 60 square feet . . . .
The pool which upon the evening of the 5th (July) had been teeming
with animal life, contained no living insects during the following ten
days. The kerosene, curiously enough, seemed to exercise no deterrent
effect upon the adult female mosquitos. They still continued to attempt
to deposit eggs and in this attempt were destroyed. This is in my opinion
a most important point, and one which has hardly been anticipated."
This experiment was made in the Catskill Mountains of New York at an
elevation of about 2,500 feet.
Ronald Ross in his famous book, entitled The Prevention of Malaria/
states: "Since 1899 I had gradually become convinced that anti-mosquito
campaigns had better be conducted, not only against Anophelines but
against all kinds of mosquitoes at once." He refers to his advice given
on the spot in 1902 to continue general mosquito reduction at Ismailia
on the Suez Canal; within a year, he states, mosquitoes of both groups
(Culicine and Anopheline) were almost entirely banished and the ad-
missions for malaria reduced from about 2,000 a year to 214. Ross also
refers to the brilliant campaign commenced in 1901 by Malcolm Watson
in the Federated Malay States at Klang and Port Swettenham.
256
The remarkable work of Gorgas in controlling yellow fever in Havana
in 1901 and that of Gorgas and Le Prince 3 in the control of yellow fever
and malaria in the Panama Canal Zone attra<::ted widespread attention
to the subject of mosquito abatement. Ross (1902), in his book Mosquito
Brigades and How to Organize Them; gave ll)uch practical information
regarding methods of mosquito control. In 1939 Herms and Gray pub-
lished the first edition of their work on mosquito control (now in the
second edition). 5
L. O. Howard contributed a special article to Ross's book on malaria
prevention. In this article, which was entitled "Anti-malaria work in the
United States," Howard says" ... beginning with 1900 and gradually
increasing in amount year by year, a great dt:al of anti-mosquito work,
more or less well planned, has been done in different parts of the
United States; but, looking over the entire field it is astonishing to note
how little of this work has been done with q direct sanitary object in
~'iew." He r&'ll81'ks tll8t, '1t is FleMing, in t~ face of so much of this
sort of thing, to note a well-directed and rather large-scale bit of work
with a direct anti-malarial bearing which was begun at an early
date, and this is the work on Staten Island under Dr. A. H. Doty, the
health officer of the port of New York." This work was evidently started
in 1901.
Public interest in the United States was greatly stimulated by the
appearance, in 1904, of J. B. Smith's report On the mosquitoes of New
Jersey.6 Salt-marsh mosquito abatement on the Pacific coast began in
1904 at San Rafael, Marin County, California, under the direction of C.
W. Woodworth, and at Burlingame, California, in 1905. 1 In 19108 the
first specific malaria-mosquito control work in California was begun by
the author (Herms) at Penryn in Placer County. The campaign was
locally supported. Somewhat prior to 1910 the author called attention
to the danger from malaria due to anopheline breeding in seepage from
poorly constructed irrigation ditches and c~mals in California. "The
use of metal, cement, or tile irrigation ditches, which prevent lateral
flow except where wanted, will help greatly in lessening the vast number
of (malaria) mosquitoes now produced in 01' by poorly kept ditches."
Farmers of northern California were admonished. to "pay more attention
to the improvement of their irrigation methods."
In 19139 Carter of the United States Public Health Service began
to make mala"rveys in Virginia and North Carolina. In 1914 a
Federal appropnation of $16,000 was provided for malaria investigations
by the Public Health Service; 22 surveys and three malaria control dem-
onstrations were made in seven states in that year. From 1914 to 1928
the United States Public Health Service conducted directly or cooper-
atively malaria-mosquito control work in 343 communities in 17 states
and, including surveys and investigations, in 667 communities in 24

states.
Organization for abatement work. Before actual mosquito abatement
work is undertaken, it must be well planned and organized (Herms and
Gray, loco cit.). There must be an adequate preliminary survey. The
services of an experienced and practical mosquito-abatement expert
should be secured to make this survey. The expenses for the preliminary
survey may be defrayed from funds raised by public subscription or by
any other means.
The preliminary report should include data and recommendations
on the following matters:
1. The boundaries of the area which should be included within the
project.
2. The population, area, and assessed valuation of the proposed district.
3. The location and areas of the principal breeding situations such as
marshes, etc.
4. Tentative suggestions as to the best method of control for each prin-
cipal breeding area.
5. The amount and types of domestic breeding, and measures for abate-
ment.
6. The particular species of mosquitoes involved, with a brief discussion
of their breeding habits.
7. The probable organization that will be required, including personnel
and equipment.
8. Detailed preliminary estimates of cost, both for permanent work (in-
cluding capital outlays) and for regular maintenance. With these should be
submitted comparative costs for already operating districts of similar size and
conditions.
9. The economic losses caused by mosquitoes in the proposed district,
and the economic savings which should result from adequate mosquito abate-
ment.
10. Where a mosquito-borne disease is involved, statistical material of
epidemiological importance should be included (see malaria survey, p. 230, for
pattern) .
Personnel. The general supervision of organized community mosquito
control work, particularly as to poliCies and finance, is usually in the
hands of a board or commission appointed in various ways in different
states. The proponents of the undertaking should make the proper rep-
resentations to the appointing power or powers, so that only citizens of
outstanding character and ability are chosen. It is desirable that the
members of the board or commission serve without compensation, except
that expenses incurred in the performance of duty should be defrayed.
In making selections it is well to include an outstanding physician, a
public-minded attorney, a respected and able civil or sanitary engineer,
and a successful business man-all should have a deserved reputation for
unselfish public service.
The most important duty of the board or commission is the selection
of the executive officer, who for the larger districts should have at least
the following qualifications: (1) agreeable personality and honesty, (2)
successful experience in mosquito abatement work, (3) administrative
ability, (4) training in medical entomology and sanitary engineering.
For very small districts this officer may well be of the working-foreman
type.
A responsible group of citizens whose duty it is to administer the
expenditure of considerable sums of local tax money for mosquito control
is not likely to appoint as manager or executive officer a man who is
simply brimful of so-called personality on the assumption that he will
make a good "public relations" man. A good personality is an important
asset, but the tax-paying public is particularly concerned with the man's
ability to "deliver the goods." Thorough familiarity with the goods, in
this instance mosquito control, is essential to good public relations. 10
Personnel should be carefully selected so as not to freeze into the organ-
ization at the outset individuals who are unfit for this type of work.
In large districts it is necessary to subdivide the force into divisions
which can be handled by a responsible foreman or inspector and a crew
of laborers. The foreman should be a full-time employee, active, ener-
getic, interested in his work, and able to supervise his men in all sorts
of mosquito control operations. As he is in immediate contact with the
public, he must have a good personality, and must be able to get along
with people. He will require much "backbone," plus self-restraint and
patience. He must be physically equal to sustained activity in the field,
for the work is frequently arduous. The regular crews should be care-
fully selected and kept on a year-round basis, with temporary additions
during the peak of the breeding season as needed.
Adequate office facilities must be provided, inclusive of clercial and
telephone service, for public contact and business purposes. Detailed
maps of the district must be available, a reporting system, and a book-
keeping system in conformance with legal requirements must be de-
veloped. The office should maintain a skillful and continuous program
of publicity and public education.
Training of personnel and education of the public. In order to secure
a high level of performance in mosquito control operations the admin-
istrative and operational staff of a mosquito abatement district must be
properly trained in control techniques. Mosquito control involves highly
technical skills and complicated procedures which unless correctly ad-
ministered may injure the public health and in agricultural areas may
prove injurious to agriculture. Also in order to secure full cooperation of
260 . MEDICAL ENTOMOLOGY
the public there must be a program of public education. One or more
members of the staff should be trained to carry this out; in fact, the
nature of mosquito control is such that all members of the staff, and
laborers as well, are in constant contact with the public while on duty,
hence all have a measure of responsibility in this regard; even a minor
"slip up" may cause a "break down" in public relations.
Among similar agencies in other states, the California State Depart-
ment of Public Health through the Bureau of Vector Control maintains
a "Mosquito Control Training Program" designed to provide the opera-
tional personnel of the mosquito control agencies of California with the
fundamentals of mosquito control. Two training teams, each consisting
of one entomological instructor, one operational instructor, and an as-
sistant equipped with an outfitted station wagon, covered the state from
San Diego to Redding in 1948 holding 13 field training programs serving
39 mosquito abatement districts, 7 local health departments, 4 United
States Navy installations, and 5 other agencies during February, March,
and April. In addition to lectures and laboratory instruction, there were
field demonstrations in the use of insecticides and equipment. Attendance
certificates were given, and much informational literature was distributed.
Instructors used on these teams were required to have had field experi-
ence and top training, particularly since many of those receiving in-
struction were men of experience themselves.
Mosquito surveys. The Simplest form of mosquito survey on which
policy and action can be based has as its minimum objective the ac-
curate determination of all species of mosquitoes present in a given
locality, estimating their relative abundance, mapping their distribution,
and studying their respective breeding places. The survey will delimit
the boundaries of the area which should be included within the project;
it should ascertain the population and assessed valuation of the area;
it should specify in general the best method of control for the principal
breeding areas and the probable organization that will be required
(personnel and equipment); it should furnish estimates of cost both
for permanent control work and for regular maintenance.
Surveys must be done intelligently (i.e., by qualified persons) and
with adequate means; they are intended to guide policy and action,
usually involving considerable sums of public money. Adequate maps
are required to show topographical details. Surveys in greater detail
should secure the following information concerning the area under con-
sideration: ( 1 ) meteorological conditions (by the month), rainfall,
humidity, temperature, winds, etc.; (2) topography, soil, vegetation,
drainage, water courses, ponds, lakes, springs, swamps, salt marshes,
ditches, etc.; (3) agricultural crops, methods of farming, major crops,
irrigation practices, rice culture, etc.; (4) additional specific epidemiQ-
logical information with regard to malaria a.nd other mosquito-borne
infections, if present, such as sporozoite rate aud oocyst rate in dissected
anopheline mosquitoes; numerical prevalence of diHerent species of
adult anophelines in houses, privies, pig pens, barns, etc.; spleen rate
and parasite index in blood; malaria morbidity, incidence, etc.; presence
of arthropod-borne encephalitides calls for additional technical details
(see Chapter XII).
Essentials of mosquito abatement. Bearing in mind the fact that no
mosquito ever came into existence without water in which its larval
stages were completed, and that a very sma.ll quantity of water may
serve the purpose, the control of all collectious of suitable water avail-
able to mosquitoes is a matter of basic importance. The study of water
management is highly important. The objective of mosquito abatement
operations is the elimination of mosquito Pl:oduction. The abatement
method must fit the offending species of mO~quito. Marsh drainage or
the usual larvicidal treatments would be in(lffectjve ill the control of
tree-hole species such as Aedes varipalpus (Coq.) where specialized
treatment is required. Even under temperate-·zone conditions the work
of mosquito abatement is a continuous year-rOund program, with special
alertness in the spring to curb the first broo<l of larvae, and intensive
larviciding through the summer. At the end of the breeding season efforts
should be intensified to reduce the last brood as far as possible so as to
have fewer overwintering mosquitoes to start I:he following year's brood.
An intensive adulticiding program is in order. Use the winter months
for maintenance work on drainage systems; for construction of new
drains and permanent structures; and for planning the following year's
work. Permanent corrective work should be the aim.
Water management. Anyone concerned with mosquito control opera-
tions soon learns that much mosquito breeding is due to mismanagement
of water intended for useful purposes. Irrigatton systems so essential to
crop production in arid regions too often do not provide proper drain-
age, or the canals and ditches may be so sitltated or so constructed as
to permit seepage resulting in ideal breeding places for potent malaria
vectors such as Anopheles freeborni Aitken. Improper flooding of rice
fields or uncontrolled flooding, as well as imI)roper irrigation practices,
in the production of alfalfa and ladino clover may be productive of pest
mosquitoes. Irrigated pastures may be a SOltrce of many mosquitoes.
Water impoundment may result in the prodU(~tion of important malaria
vectors such as Anopheles quadrimaculatus Say in the middle and south-
ern United States. The Significance of such iIbpounded water, so far as
malaria is concerned, depends upon the presence or absence of a vector
Anopheles adaptable to such breeding conditions. Mosquito production
is due in large measure to man's carelessness and indifference, so well
illustrated by the ubiquitious borrow pits formed by the removal of earth
for highway and railroad embankments aDd fll1s; and by the draiDage
ditch improperly constructed with respect to slope or outlet (see Kingll ).
Irrigation. Where irrigation is properly practiced with due attention
to the economical use of water and good agricultural practice, there
need be little or no mosquito breeding and consequent malaria menace.
However, when seepage results from breaks in ditches and particularly
from side-hill canals, ideal swampY ' areas are produced and cattle and
FIG. 82. Breaks in the irrigation ditch are responsible for considerable
inundation, producing favorable breeding places for mosquitoes. The rapidly
running water in the ditch is unfavorable for mosquitoes.
horses leave water-filled hoof prints, an ideal breeding situation for
Anopheles freeborni Aitken in the western United States. Because of the
breeding habits of this important western malaria-bearing mosquito,
poorly constructed and improperly operated or leaking irrigation ditches
commonly account for malaria in the neighborhood (Fig. 82).
Wilson ' 2 points out that "malarial effects are not attributable directly
to the results of irrigation where it is economically and properly prac-
ticed-where care is taken to irrigate only land which has an open soil
and such slopes and natural draiDage as to prevent waterlogging, no
unhealthy effects will result. . .. It is desirable, in order to mitigate the
possible evil effects of irrigation, to keep the canal as much as possible
within soil so that its surface level may be low, and thus only raise the
sub-surface water plane to the least height practicable; that earth wanted
to complete embankments be never taken from excavations or borrow-
pits except where such localities admit readily of drainage."
It may be pointed out that there is a distinct difference between
agricultural drainage and mosquito abatement drainage as applied to
irrigation districts. Agricultural drainage is concerned merely with the
problem of lowering the ground water level to a point where crops can
be raised successfully. Frequently considerable quantities of mosquito-
breeding water remain, often in the drains themselves (Fig. 83). Such
FIG. 83. Drainage water resulting from faulty irrigation, a source of numerous
mosquitoes.
drainage consists usually of large deep main drains with comparatively
few laterals. Mosquito abatement drainage, on the other hand, is a matter
of more careful attention to detail, with great care to obtain uniform
grades and smooth bottoms for the drains, so as to avoid mosquito breed-
ing.
Impounded water. The storage of water in deep, clean, and steep-
walled reservoirs seldom presents a mosquito problem; neither does
storage of water in steep-sided reservoirs in rocky canyons. "The im-
poundage of flowing streams," Bishop, Hollis, et. al.,13 (1947) point
out, "profoundly alters the biology of the affected waters. One of the
changes may be a vast increase in the potential for anopheline mosquito
production unless appropriate control works and procedures are planned,
designed, and operated. The choice is, therefore, between the probability
of a disastrous 'man-made' increase in transmission of malaria or a suc-
cessful resolution of the continuing conflict between adverse biological
pressures and control practices." The magnitude of operations by the
Tennessee Valley Authority begun in 1933, based on water management
( control), is indicated in the fact that these involved 10,000 miles of
reservoir shore line in a region of widely varying topography. The im-
pounding of a lake in a region of flat topography within the Tennnessee
Valley produces a wide variety of breeding places for Anopheles quadri-
maculatus Say. There is here provided a huge food supply of micro-
organisms in the flotage and optimum protection against enemy fish and
other aquatic predators. One of the important ways to control mosquito
breeding in this situation is water level management. The effect of water
level manipulations on anopheline larvae is both direct and indirect;
direct in that as a result of drawing the water out of the marginal vegeta-
tion the larvae are exposed to an open and clean water surface wherein
few can survive; and indirect in that the water is removed from the
marginal plants, drift, and flotage which provide food and protection for
the mosquito larvae. One of the important elements in water level man-
agement is periodic water level fluctuations. Complicated biological
study and engineering features beyond the scope of this book are in-
volved in accomplishing the desired results.
The author has seen mosquito breeding swamps in California made
over into small recreational lakes by some dredging and impoundment
of open water unsuitable for mosquitoes.
Controlled reflooding may be successfully employed in the control
of salt-marsh Aedes mosquitoes. The eggs of salt-marsh Aedes do not all
hatch on anyone hatching; some eggs do not hatch until there have been
a number of successive wettings. It is at least theoretically possible ulti-
mately to hatch out all mosquito eggs in a given reclaimed marsh by
purposefully flooding the marsh at the monthly high tide, holding the
water on the marsh for several days until a brood of larvae appears, and
then draining the marsh rapidly before the larvae develop into adult
mosquitoes. The marsh is then allowed to remain unwatered until a
subsequent monthly high tide when the flooding is repeated. As no
adults have been permitted to emerge, no new eggs are deposited, and
thus by successive wettings all eggs should be hatched eventually with-
out additions of new eggs. Thus actually thousands of acres of salt marsh
on Suisun Bay near San Francisco have been freed of Aedes dorsalis
(Meig. ), so that these marshes can now be flooded or dried out at the
will of the owner without hatching hordes of mosquitoes. From 7 to 10
successive floodings and drainings were necessary to accomplish this.
Flushing. Several species of Anopheles mosquitoes, particularly A.
minimus fiavirostris (Lud.), the principal malaria vector of the Philip-
pine Islands, breed in the running water of small upland streams, the
larvae being able to hold fast against the current by means of small
MOSQUITO ABATEMENT 26.5
hooks; other anophelines breed in stream-bed pools. Flushing devices,
either automatic or hand-operated, can be installed at intervals along
the breeding length of such streams so as to release suddenly a relatively
large quantity of water accumulated behind a dam. The mosquito larvae
are swept downstream forcibly and are destroyed by stranding and crush-
ing. (See Herms and Gray, lac. cit.)
Salinification. Anopheles albimanus Wied., the principal malaria
vector of the West Indies, breeds prolifically in certain brackish coastal
lagoons. Hurlbut14 showed that where the salinity remains between 15
and 25 per cent sea-water breeding is abundant, while collections made
concurrently where the salinity was 70 to 80 per cent were entirely nega-
tive. Breeding of this species can be eliminated by increasing the salinity
to about 75 per cent sea water by making adequate sea connection, thus
subjecting the marginal lagoons to tidal action.
Russell (lac. cit.) reports that Knipe and lIackett used tide gates in
reverse position .'It Dur.'lzzo, in .4Joonm, to trans;form a. bra.ckish swamp into
a sea-water lagoon to inhibit the breeding of Anopheles sacharovi Favr.
Creeks and small streams. Except for flood. water left behind during
overflow from floods, great rivers themselves rarely afford opportunity
for mosquito breeding. Flood water left behir{d when rivers recede may
become a prolific breeding place for certain ~pecies of mosquitoes such
as Aedes vexans (Meigen), and in some insta1lces in appropriate regions
a malaria hazard may be created in that a brEleding place for Anopheles
quadrimaculatus Say may result. In many parts of California, as creeks
and smaller streams recede during the rainlesfl summer, numerous sunny
pools are left behind which soon become green with algae (e.g., Spiro-
gyra) among which vast numbers of larvae of Anopheles pseudopunc-
tipennis franciscanus McCracken occur; also quiet deeper pools occur
along the banks shaded by wild grapes, willow, and other vegetation, in
which Anopheles punctipennis (Say) find suiblble breeding places. Pools
along the banks can frequently be drained off or can at least be thor-
oughly larvicided.
Small streams or creeks that border COITlmunities or flow through
the town frequently become clogged with rubhish and eventually become
prolific mosquito breeders, and situations frequently result which are
favorable for Anophelines. Communities ShOl11d not permit rubbish to
be thrown into stream beds. By neatly channelizing such stream beds
and planting the banks with shrubbery, an eyesore and mosquito hazard
may be rendered attractive and harmless.
Drainage. The removal or control of collettions of water which pro-
duce mosquitoes presents a distinct problem in nearly every case. The
type of drainage required in many instances will generally not require
technical engineering skill. There are, howeveJ:, drainage projects of con-
siderable proportions for which engineering skill is essential; such skill

is eminently necessary for large-scale salt-marsh drainage operations
where land reclamation calls for dykes, drainage canals, pumps, tide
gates, and the like. Unless the executive officer is himself an engineer,
properly qualified engineering personnel must be employed to secure
effective control.
In the case of a swamp caused by springs, a system of deep circum-
ferential cut-off drains is recommended in order to intercept the seepage
water and conduct it around the wet area. Where streams debouch from
the hills onto a Rat plain or valley, water from heavy rains, particularly
in the spring, tends to spread out and leaves temporary pools which may
produce mosquitoes. Usually these temporary pools can be more eco-
nomically controlled by larviciding; but, in some cases at least, drains
.may be dug which lead back into the main stream at a lower elevation.
These ditches will usually require considerable maintenance.
Mosquito breeding in rolling country is commonly due to artificial
obstructions, particularly railroad or highway embankments with im-
properly placed culverts. Usually culverts are set too high so that pools
or swampy areas are formed on the upper side of the embankment. Cor-
rections are usually not speedily made if at all; hence heavy larviciding
is a necessary remedy.
In some swampy situations because of small or negative gradients,
drainage becomes very difficult or impossible. In such cases sumpage
ditches or sumpage wells may be constructed, and the collected water
may then be heavily larvicided. Surface water may also persist because of
hardpan (impervious subsoil, etc.), in which case vertical drainage may
be resorted to by digging sumpage wells, or by blasting.
Drain ditches. The purpose of laying out drain ditches is to secure
effective and economical drainage. Although open drain ditches are
most commonly used in mosquito control operations, subsurface drainage
with the use of tile, or bundles of saplings laid in the ditches may at
times be appropriate. Where the general slope of the ground is appreci-
able to the eye, this is usually a simple matter, i.e., following the low
points to a place where the drainage water can be disposed of into some
natural water course or other situation where there is sufficient fall to
carry away the water. Laterals are then run from the main drain by the
shortest distance to connect up with low spots or wet areas. The bottom
of the main drain ditch must be kept deep enough so that the laterals
can reach all the low spots in the area to be drained.
For most of the ordinary ditching for mosquito control transit and
level are unnecessary. All that one needs are a few long stakes, 500 feet
or more of stout chalk line or strong cotton cord, for line, and a hand
level with a 10-foot board marked in feet and inches. If drains of con-
siderable size and yardage are to be excavated, the usual surveying
methods are employed, and the work should be done under contract
with power machinery unless such equipment is owned or otherwise
available to the district.
Hand labor, with pick and shovel, serves many purposes very well.
As a rule the square-point long-handled shovel is to be preferred to the
round-point shovel for ditching. Under some conditions long-handled
spades may be satisfactory. Mattocks may be found useful in some soils.
In many cases, more or less dense vegetation has to be cut down and
cleared before ditching can be performed with any speed or economy.
Heavy grass or weeds may be cut with scythe, sickle, or machete. In
open fields a horse-drawn hay mower may be economical. For brush,
either axes, brush hooks, or machete may be used. The machete, how-
ever, is a dangerous tool in the hands of a man not accustomed to its use.
Dense grass and some forms of brush may be killed by applying stove
distillates or Diesel oils, which are toxic to vegetation, or standard weed
killers such as 2, 4 D. After killing and drying, the vegetation can be
burned. Arsenical weed killers such as sodium arsenite, or an acid solu-
tion of arsenious chloride, may be used where there is no danger to
cattle or other herbivorous animals. However, in any agricultural com-
munity the use of arsenical weed killers should be avoided in mosquito
control operations.
Weed burners, capable of throwing a flame several feet, can be used
for clearing dense vegetation, but the general experience seems to be
that they are more expensive than other methods of clearing. If burning
is to be done, due consideration must be given to the nesting season
and nesting habits of wild life.
Ditching with dynamite. Swampy ground may often be economically
ditched with dynamite. Special directions will have to be followed and
trial shots will be necessary as a rule to determine the correct depth of
holes, their distance apart, and the amount of dynamite per hole. The
most satisfactory results are obtained by using 50 to 60 per cent straight
dynamite, fired by self-propagating detonation.
Dynamite ditch construction is advantageous in that a ditch can be
blown through land with stumps, boulders, etc., without first removing
these obstructions; it is only necessary to place heavier loads at these
points.
Maintenance of ditches. After the drainage ditch is constructed, it
must be maintained in effective working order. Three main conditions
making constant maintenance necessary are:
(a) Growth of vegetation,
(b) Caving or sloughing of banks,
(c) Artificial obstructions.
Under some conditions growth of vegetation in and adjacent to
ditches is not a problem, but as a rule ditches will require clearing sev-
eral times a year in order to keep them free from obstructing growths.
In tropical or semitropical countries the problem of keeping drains free
from vegetation is most difficult, and the source of considerable main-
tenance expense.
While weed killers may be helpful under some conditions, depend-
ence must be placed in most cases on hand labor in cutting down and
clearing out vegetation. The frequency of clearing will depend, of course,
on local soil and climatic conditions, and it will be difficult as a rule to
estimate in advance what the annual cost of maintenance of ditches
will be.
Caving or sloughing of ditch banks is apt to occur in new ditches, for
the first year or two. After that the banks usually become fairly stable,
and but little further trouble is encountered, unless cattle are pastured
along the ditches. In that case the animals may break down the ditch
banks and cause some trouble.
Artificial obstructions are as a rule frequent only in the vicinity of
public roads near a city or town. It is surprising how many people will
haul refuse away from their homes out into the country and dump it,
usually in a ditch close to a road and thus block the ditch completely or
partially. The ditch thereupon becomes a mosquito breeder.
Frequent inspection of drainage ditches, say at least once every two
or three weeks during the breeding season, should be carried on and all
obstructions to f!.ow, whether natural or artificial, removed promptly.
During the winter all ditches should be gone over carefully and trimmed
to grade, and the proper side slopes remade, so that all will be in first-
class order at the beginning of each breeding season.
Salt-marsh drainage. Salt-marsh drainage requires special study and
experience because of tidal action, soil conditions, differences in behavior
of salt-marsh mosquitoes, and other factors. The rich reward in comfort
and reclaimed land has given incentive to salt-marsh mosquito abate-
ment. Strong public-spirited organizations have given marked support
to this work on the Atlantic coast as well as on the Pacific, particularly in
New Jersey and California.
These marshes include vast areas of tidal marshes affected by salt or
brackish water along the shores of oceans and particularly the various
bays, sounds, and estuaries. The effect of daily (diurnal) and spring tides
resulting in f!.uctuations of water level is the principal feature distinguish-
ing these from fresh-water marshes, although other characteristics, such
as the salt-marsh vegetation, are of importance.
While salt marshes appear to be flat, there is a gradual slope be-
tween low tide level and the adjacent dry land. For practical purposes
these marshes may be divided into two main areas, the area subjected to
daily tidal action, where mosquito breeding seldom occurs, and the area
between the elevation of mean high tidal water and the elevation of
the extreme high tides. It is in the latter area that practically all the
breeding of salt-marsh mosquitoes occurs.
Mosquito control operators concerned with projects involving salt
marshes must acquaint themselves thoroughly with tidal phenomena.
These vary in range and type in different parts of the world. The 80-
called "spring" tides, one or two of which Occur each month, are the
tides which fill pools along the upper portions of the marshes, and in
these pools the principal salt-marsh mosquito breeding takes place.
The dates, times, and heights of the monthly highest tides are im-
portant in their bearing on the approximate time of emergence of a new
crop of salt-marsh mosquitoes. It should be the invariable practice dur-
ing the salt-marsh mosquito breeding season to inspect all known or
suspected marsh breeding areas, beginning about two days after the
highest tide and completing the inspection within six or seven days. The
delay of two days is for the purpose of giving the larvae opportunity to
develop to a sufficient size to be easily seen with the unaided eye. As
under favorable conditions the time from egg hatching to emergence
may be as short as eight days, it is obvious that if a Hight is to be pre-
vented, the inspection and necessary control measures must be com-
pleted before the time of emergence.
Because salt-marsh mosquito control involves the location of dikes
and tide gates, the knowledge of the dates, times, and heights of both
the lowest tides and the highest tides, particularly of "storm tides," the
combined effect of "spring tides" and piling up of water on shore due
to high winds and river Hood waters as well, is important.
Marsh vegetation is often very dense and interferes with inspection
and larviciding; hence burning is usually recommended. But it must be
borne in mind that peat deposits commonly occur, and if the water
level has been lowered as a result of drainage, which is usually prac-
ticed, dry peat may ignite and cause a peat fire. Peat fires can be ex-
tinguished only by Hooding, which may be expensive. An additional
hazard which may follow drainage is the shrinkage of the soil and the
formation of "cracks." These cracks may be several feet in depth and
may contain water in which mosquitoes will breed in abundance. Such
cracks are difficult and expensive to larvicide though airplane applica-
tion of DDT is helpful. An area of marsh that has cracked due to drain-
age operations should be plowed so as to break up the surface and fill
in the cracks. An occasional disking after the initial plowing is recom-
mended. For mosquito abatement purposes it is desirable to lower the
water level only to the point where the water is drawn from breeding
areas in dense vegetation but not to the extent that cracking of the soil
occurs or a peat fire hazard is created. Drainage and burning must also
be done with due regard for wild life conservation.
Types of salt-marsh drainage. There are two principal methods of
marsh drainage, (1) open marsh drainage, and (2) reclamation. In open
FIG. 84. Salt-marsh drainage operations with drag line mounted on timber
mats to prevent sinking in mud. The bucket is about to cut into a breeding
pool which will be drained into ditch in foreground. (Photograph by H. F.
Gray.)
marsh drainage the marsh is opened up by ditches to the free ebb and
Row of tides so as to eliminate standing water suitable for mosquito
breeding. In the reclamation type the area to be drained is surrounded
on the low sides by a dyke which is pierced in one or more places by
outlet structures, tide gates, which permit water behind the dyke to
run out at low tides, but prevent the return flow at high tides. Suitable
drainage ditches are dug to conduct water to the outlets (Fig. 84). The
reclaimed marsh may be used for agricultural or industrial purposes.
Filling and pumping. In almost all mosquito abatement work low wet
areas will be encountered which cannot be economically drained. AI-
FIG . 85. Upper figure shows a pond adjacent to a railroad and caused by
obstructing the natural draina-ge, a source of many mosquitoes every year.
Larviciding, while serving the purpose of mosquito control, requires repeated
expenditure of time, labor, and money. The lower figure shows the same spot
after it had been permanently corrected by the railroad company.
though some such places may be ponded and the water stocked with
top minnows, usually the most satisfactory method is filling (Fig. 85).
Most smaller holes, such as borrow pits, can be filled in by hand shovel-
ing; larger holes may be filled by means of power scrapers. If sanitarily
handled, municipal garbage and refuse may be used in a "fill and
cover" method. Such fills are covered with earth so as to prevent fly
breeding and to some extent rat breeding.
Salt marshes may be filled by hydraulic dredges, which suck mud
and sand from the bottom of an adjacent bay and pump the mud and
water mixture through a pipe and discharge it on the marsh. Where
harbor or channel improvements are being made by hydraulic dredging,
very satisfactory arrangements may often be made to use the mud and
sand to fill a near-by mosquito-breeding marsh.
r::;:,"
FIG. 86. Rice field with roadside seepage ditch in which mosquitoes breed.
When the general land level is at or below the low-water level of
an adjacent river or bay, pumping has to be resorted to. Portable pump-
ing units are frequently of much value.
Mosquito breeding in rice fields. The control of mosquito breeding in.
rice fields is obviously a matter that must have the attention of those
concerned with rice culture (agronomists) as well as the mosquito con-
trol experts. Much of the breeding can be controlled by proper manage-
ment with respect to the habits of mosquitoes. Many species of mos-
quitoes are involved, some of which are malaria vectors. In the exten-
sive rice-growing areas of Arkansas, 98 to 99 per cent of the population
of noxious mosquitoes consist of four species, Psorophora confinnis
(L.-A. ) and P. discolor (Coq.) both flood-water breeders, constituting
83 to 94 per cent, and Culex erraticus D. and K. and Anopheles quad-
rimaculatus Say, a malaria vector constituting the rest (Horsfa1l15 ). In
some newly developed areas the introduction of rice culture on a large
scale is characterized by hasty, haphazard rrlethods (quick returns at
minimum cost), with utter disregard of saund agronomic practice.
Poorly constructed irrigation systems and even poorer drainage methods
result in a veritable bog adjacent to rice fields (Fig. 86). Too often as
much as half of the mosquito production is Jue to poor practices and
could be largely controlled by improved methods of water management.
Growing rice in very porous soil requires much replenishment of water,
and the continous freshening by replenishment may in certain areas in-
vite abundant breeding of malaria vectors, e.g., Anopheles freeborni
(Aitken) in California. Where rice growing is an old established prac-
tice great care is normally exercised in the matter of water management,
particularly so in the vicinity of communities. People must learn to live
in such an environment, and it is folly to endllre mosquitoes which may
be produced by rice fields.
The use of larvicides in rice fields is fraught with danger in that the
rice plants may be damaged. Rao and Sweee 6 found that the use of a
one per cent dilution of Paris green in road dust and wood ash produced
no ill effects on the rice or in straw yield. There was no indication that
dusting should be stopped during the period in which the paddy was
in Hower.
Several authors have recommended intermittent irrigation, e.g., Hill
and Cambournac l1 in Portugal, and Russell, 1<nipe, and Rao 18 in South
India. The latter recommend a cycle of five wet days and two dry days,
particularly for the control of the vector, Anopheles culicifacies Giles.
In 1948 Cambournac and Fonseca,t9 reporting on anopheline and
malaria control in the rice-growing regions of Portugal, point out that
the control of anopheline larvae in rice fields by the use of DDT emul-
sions is both economical and practical. They recommend 5 liters of a
one per cent emulsion per hectare, i.e., about 5.28 quarts to about 2.5
acres, poured in the irrigation ditches and thus automatically dispersed
in the plots.
In California experimental work is in prOgress in the rice-growing
area of the Sutter-Yuba Mosquito Abatement District in which the rice
seed is treated with DDT (wettable powdeI' which clings to the seed
in the hopper) and is then seeded by airplane. In this same district
studies were made during September, 1947, tJsing DDT applied in air-
plane thermal exhaust aerosols for the control of Anopheles larvae. Ac-
cording to State Board of Health reports, better than 95 per cent mor-
tality of Anopheles freeborni was obtained with as little as one pint of
DDT solution per acre with :;l. dosage of 0.025 pounds of DDT per acre.
The cost of treating 18,000 acres of rice fields was 30 cents per acre;
'this sum included cost of hiring the airplane pilot, ground crew wages,
DDT, and solvents.
Anopheline control in Bromeliads. Anopheles bellator Dyar and Knab
is regarded as the most important vector of malaria in Trinidad (West
Indies), where it breeds in the rain water collected among the leaves of
epiphytic Bromeliads, particularly the large so-called "tank plants,"
Gravisia aquilega and H ohenbergia stellata, and others. These plants
are found in great profusion on immortelle trees which are planted to
shade cacao; hence the problem of anopheline control is bound up with
the cacao industry. Manual removal of the bromeliads is costly and has
limited value. Gillette"° reports that the spray killing of bromeliads by
the mechanical application of copper sulfate is the method of choice.
He states that one application of 0.25 per cent to 0.5 per cent solution
may suffice for a 10-year period. The instance is cited of the treatment of
about 1,500 acres in a district of 30,000 population has resulted in a
spleen rate reduction from 28 per cent in 1945 to 5 per cent in 1947.
Sewer inlets and catch basins. In the newer types of street inlets little
opportunity is afforded for water to collect and remain standing for
mosquito breeding. Most of the old types of inlets and catch basins,
especially those connecting with a combined sewer (for domestic sewage
and storm water), are apt to produce mosquitoes, particularly Culex
pipiens Linn. Larviciding is most economically done by means of a
motorcycle sidecar (Fig. 87). One filling with 25 gallons of larvicide
with air pressure to 50 pounds per square inch, will suffice for a day's
work, i.e., from 200 to 300 catch basins.
Public utilities street vaults. In practically all urban areas the various
public utilities, such as telephone, power and light, gas, electric railway,
telegraph, and water, have numerous vaults in streets which are fre-
quently the source of a severe local infestation of Culex pipiens Linn.
during the breeding season. Heavy residual spraying with DDT will
give good control.
Cesspools, privies, liquid manure pits. Where pit privies are built in
wet areas, water collects and prolific mosquito breeding (Culex quin-
quefasciatus Say, Culex pipiens Linn., and C. tarsalis Coq.) may re-
sult. The use of DDT or cresylic acid larvicide is recommended. Where
liquid wastes are disposed of in leaching cesspools, mosquito production
may be very great if mosquitoes have access to them. Even small knot '
holes or v€mt pipes afford a ready means of entrance. If necessary re-
pairs are made so the egress is cut off, the larvicide need not be applied;
otherwise treatment at appropriate intervals of about 10 days must be
practiced.
Liquid manure pits in connection with greenhouses and plant nurs-
eries commonly produce prodigious numbers of Culex pipiens Linn.
and Culex tarsalis Coq. Larvicides must be used which are not dangerous
to plants; DDT and several other newer insecticides may be employed.
Tree holes. In wooded areas and on estates tree-hole mosquitoes, such
as Aedes varipalpus (Coq. ), often cause great annoyance. Tree holes,
particularly in oaks, collect water in which mosquito larvae thrive.
FIG . 87. Motorcycle sidecar equipped for larviciding. (Photograph by H. 1

Gray.)
Liberal application of DDT or cresylic acid larvicide is recommende(
the larvicide should be applied not only to the water but also liberall
swabbed on the wood above the water line to repel egg-laying fema
mosquitoes. The practice of tree surgery on estates has greatly reduce
mosquito breeding in tree holes. Heavy applications of DDT aeros'
have been effective in canyons or narrow valleys.
Cemetery urns. Urns and other types of receptacles intended f,
flowers on graves may stand partly filled with water for considerable
lengths of time without changing and may furnish breeding places for
many species of mosquitoes. Thus a mosquito survey of the cemeteries
within the boundaries of the Alameda County (California) Mosquito
Abatement District showed that 11 species were breeding in such re-
ceptacles, namely, Culex pipiens Linn. about 70 per cent, Culiseta in-
cidens Thom. about 25 per cent, Anopheles pseudopunctipennis fran-
ciscanus McCracken and A. punctipennis (Say) about 5 per cent. Culex
tarsalis Coq. and C. stigmatosoma Dyar larvae were only rarely found.
Of the 18 cemeteries the largest contained 80,000 graves with some
27,000 containers for flowers. During the summer season at least half of
these containers were breeding mosquitoes; in January, 20 per cent still
contained larvae. Excellent control was effected by the use of DDT
aerosol, 5 per cent emulsion in Diesel oil, applied at about three-week
intervals. Each treatment (made before 7 a.m.) required only about
one hour, whereas previously over 200 man-hours were required for
hand treatment (Aarons,21 1948).
FIG. 88. Tin cans, tubs, and barrels in which water may stand, furnishing
breeding places for mosquitoes.
Sundry nuisances. Water-holding receptacles of many kinds may

prove suitable for mosquito breeding and must not be overlooked by the
"mosquito man." However, it sometimes happens that an overemphasis
of the tin can results in attracting the attention away from more im-
portant matters, such as dripping hydrants, stagnant ditches, etc. Indeed
the water in tin cans unless in shady situations usually becomes too hot
for mosquitoes during most of the summer. (Fig. 88.) Heaps of broken
gourds commonly reek with mosquito larvae; tubs and barrels of water
frequently produce many of these pests, even certain species of anophe-
lines. Stagnant water in poorly constructed street gutters is often a
serious menace. Dripping faucets (Fig. 89) may result in pools of water
suitable for mosquito breeding.
Routine inspections for mosquito breeding. Inspections for the purpose
of finding breeding places in organized mosquito abatement districts are
made either upon the receipt of complaints or for routine purposes. The
inspector should be guided by the nature of the complaint: (1) mos-
quitoes annoying at night or affecting sleep, and (2) annoying during
the daytime or toward evening, while working in the garden or watering
the lawn. In the first instance it is probably a domestic fresh water
species, breeding on the premises or in the immediate vicinity. In the
second instance it is probably a salt-marsh species, if salt marsh is near,
or it may be a flood-water or tree-hole Aedes. In a previous chapter (p.
181) exceptions to this rule are discussed, particularly the spring dis-
persal flights of some species of Anopheles. Obviously the inspector must
have a thorough knowledge of the species of mosquitoes and be well
informed concerning breeding habits. 'When complaints are received,
the inspector should visit the premises and if possible capture mosquitoes
--,
I
I
FIG. 89. The dripping faucet with

the resulting pool of water is often a
constant menace in the entire absence
of other mQsquito-breeding sources.
for positive identification, so as to simplify inspection for larvae and to

ensure effective abatement.
In searching for adult mosquitoes for species determination it is
important to remerriber that few species al'e active during the day;
hence one must search for them in dark, co()l and moist places, under
houses, in basements and cellars, behind pic:tures, in closets and dark
corners. A flashlight is very useful in these operations. Aedes dorsalis
(Meigen) and other day fliers often hide in shrubbery and may be
found by shaking bushes and vines or kicking weeds. The mosquitoes
may be collected in small cyanide bottles or by means of a sucking tube
with an extension rubber tube.
Inspections, whether due to complaint or as a matter of routine,
must be thoroughly and intelligently carried out; the breeding may be
occurring in a rain barrel, a lily pond, or a concealed cesspool; it may be
taking place in a concealed chamber fed by a natural spring; in an
abandoned well, a broken or clogged sewer or drain; Hoor boards may
have to be removed.
The work of routine or house-to-house inspection must be properly
organized for efficient and economical coverage. The inspectors must
be intelligent, well trained technically, and capable of meeting all sorts
of people.
In inspecting large tracts of marsh to locate the precise producing
areas, it is always advisable to mark off the marsh into definite sec-
tions, which can be examined one at a time, so that no portion is over-
looked. Breeding areas, when located, are marked by setting up stakes
in the center of the breeding area for the crew of larviciders which fol-
low, or for future reinspection. Frequently both inspection and larvicid-
ing may be done concurrently by the same man.
Larvicides. All too commonly mosquito control is thought of as the
practice of spraying oil (larvicide) on the surface of stagnant water.
Larvicides have a useful place in mosquito abatement operations; how-
ever, all properly conducted operations should look upon larviciding as
secondary to the methods set forth previously in tbis chapter. The cor-
rect use of larvicides should be observed at all times. The older larvicid-
ing methods such as oiling22 and dusting with Paris green 23 have been
replaced by methods using such materials as DDT, TDE, BRC, or new
formulations.
Oil (petroleum) which is to be used on water to destroy the aquatic'
stages of mosquitoes must, of course, be lethal, at least in time, to all
larvae and pupae; it must spread well on the surface of the water and
must have lasting qualities. Mixtures suitable for larvicidal purposes
should have a specific gravity of 31 ° to 39° Baume (API) and a viscosity
of 45 or lower Saybolt at 100° F; they must be economical, easily ob-
tainable, not unpleasant to handle, and if possible nontoxic to animals.
In ordering petroleum oil for mosquito abatement purposes the fol-
lowing specifications should be designated: (1) it must be homogeneous;
(2) it must not separate into fractions on prolonged' standing; (3) it
must be free from granular and fibrous material which might cause the
clogging of spray nozzles; (4) other phYSical characteristics must lie
within the following maxima and minima, specific gravity at 60° F not
less than 0.830 nor more than 0.870; viscosity (Saybolt universal) at
100° F not less than 30 nor more than 45; boiling range from 330° F to
800° F.
Cresylic acid (boiling fraction 225°-250° C) must be emulsified with
soap. The original "Panama" larvicide was made with a rosin soap as
the emulsifying agent-200 pounds of powdered rosin with 30 pounds
of caustic soda added to 150 gallons of crude carbolic acid, heated and
stirred. Whale oil soap or other cheap soaps nlay be used as emulsifiers.
The preparation of this larvicide is best done by manufacturing chemists
who have proper equipment. The larvicide is applied in the proportions
of 1 part to 30,000 parts of water.
Pyrethrum extract is very toxic to mosquitoes in the larval as well as
adult stages. Vannote and Ginsburg24 prepared an emulsion of kerosene
extract (extracted from 1 pound of pyrethrum flowers per gallon of
petroleum solvent) emulsified with water and soap (66 per cent kero-
sene extract with 34 per cent water containing 3 per cent to 5 per cent
of soap). This stock emulsion was diluted 1 to 10 with water for the
field spray. Using in excess of 50 gallons of the dilution per acre Gins-
burg obtained a complete kill of larvae on salt marsh. Formulations
made of pyrethrins, which give a rapid knockdown, and insecticides
such as DDT with long residual effect and beavy kill are desirable.
Paris green can be successfully used against the surface-feeding
anopheline larvae. This material is suitable for airplane dusting on a large
scale as well as for application with hand dusters. The Paris green is
diluted with road dust, hydrated lime, finely powdered charcoal, finely
powdered talc or other inert finely powdered material. It apparently acts
as a stomach poison, the small particles of Paris green being ingested
with larval food. The poison is usually mixed with the diluent in the ratio
of 1 part of Paris green to 99 parts of diluent (by weight). For airplane
dusting a higher percentage of Paris green is recommended, namely
from 25 per cent to 33 per cent. The amount of Paris green per acre of
marsh varies according to various authors from slightly less than one
pound to two and even bUT pounds wbere v~g~hhon "is ~~ns~.
Concerning the dangers from the use of this poison Barber and
Hayne 23 state:
In our experiments we have used no precautions other than to stand to

the windward of the dust cloud-the place where one would naturally stand in
distributing dust-and we have experienced no harmful results whatever. How-
ever, even with the small quantities used in larvicide work, it is well to remem-
ber that one is working with a poison and that some precautions should be
taken at least until the matter is further investigated. It is probably sufficient
to keep to the windward of the dust clouds and to avoid inhaling the dust as
far as possible. In case a great deal of exposure is necessary, one should use
some precaution to keep any large amount of paris green from entering the
clothing or accumulating on the skin. The danger to domestic animals through
drinking treated water seems very remote . . . . We have never observed any
effect of the poison on culicine larvae or on any aquatic insect or animal, how-
ever delicate, other than the surface feeding anopheline larvae. In particular,
we have not observed any indication of harm to top-feeding minnows or to
any other natural enemy of larvae.
Dichloro-diphenyl-trichloroethane (DDT) has been rigidly tested by
many mosquito control agencies in large-scale field operations in prac-
tically all parts of the world. Both as a larvicide and adulticide it has
been proved a superior culicide. Its use is not "foolproof"; it is a poison-
ous chemical, and care must be exercised when using it in the presence
of food and food containers in the home and in barns and stables.
Furthermore, since DDT, a white crystalline substance, is practically
insoluble in water, solvents are employed in the preparation of DDT
sprays. The solvents are inflammable and explosive; hence when using
such sprays indoors or in enclosed places without emulsifying in water
reasonable caution must be exercised. Some of the solvents are toxic to
man.
Emulsions. An emulsion concentrate may be made of 25 per cent
DDT, 10 per cent Triton X-100, and 65 per cent xylene or other solvent.
Concentrates are available commercially. Emulsions are made by add-
ing the required amount of concentrate slowly to water with continuous
stirring. To prepare a 5 per cent emulsion of DDT, for example, one
volume of the concentrate is mixed with four volumes of water. When
used in the field as a larvicide the emulsions should be applied at the rate
of 0.1 to 0.2 pound of DDT per acre of water surface. Since DDT is not
a strong pupacide, treatments must be made with this in mind or formu-
lations must be employed which include pupacides.
DDT-oil solutions. Diesel oil, kerosene, and fuel oil will dissolve
DDT in about 24 hours. To make a 5 per cent solution add about 2
pounds (2Ys pounds for a true 5 per cent weight-per-volume solution)
of technical DDT to each five gallons of oil. Used as a larvicide, control
can be obtained with 5 quarts per acre of a 1 per cent DDT-oil solu-
tion. Oil without DDT for similar control would require from 15 to 25
gallons per acre. Using a 5 per cent DDT-oil solution as little as 1 to 2
quarts per acre will be effective when applied with pressure spray
equipment.
The use of a DDT-oil solution is well illustrated in the control of
Anopheles farauti Lav., vector of both malaria and filariasis in the New
Hebrides Islands, as reported by Yust. 25 This vector bred largely during
military operations in road ruts created by military vehicles. Native
huts treated with 5 per cent DDT in kerosene at the rate of one gallon
per 1,000 square feet were free of A. farauti adults for three months.
Breeding sites were sprayed weekly with a 5 per cent solution of DDT
in No. 2 Diesel oil at the rate of about two quarts per acre. Weekly
spraying was necessary because of the formation of new water pools.
DDT dust larvicide is employed in the same way and for the same
purpose as Paris green; however, used against the larvae of Anopheles
quadrimaculatus Say it is said to be about 25 times as toxic. It is applied
as a 10 per cent DDT dust in pyrophyllite, talc, or other inert dust at
the rate of 0.1 to 0.2 pounds DDT per acre, i.e., one pound dust mixture.
Where there is abundant vegetation the dosage must be increased. For
the control of Culex and Aedes species a larger dosage is also required.
Dichloro-diphenyl-dichloroethane (TDE, also referred to as DDD)
is an analog of DDT and in comparative tests made with fourth instar
mosquito larvae by the author and many other investigators it is at
least equal to DDT as a larvicide, except in lasting residual properties.
The more rapid dissipation may have its advantages in that there is less
accumulation in the soil or water after repeated treatments. TDE is also
less toxic to fish and to warm-blooded animals as well.
Benzene hexachloride (CoHo CIs, also known as 666) and chlordane
(C1oHsCls ) both have larvicidal as well as pupacidal properties which
deserve much more research, particularly with regard to formulations.
Larvicide drips. During the years when petroleum oil was the ac-
cepted larvicide, so-called "oil drips" were used to good advantage in
giving continuous drop-by-drop treatment to ditches and small streams
of moving water. This scheme is now followed with some modifications
in a similar use of DDT. Oil drums or other containers properly equip-
ped with a faucet permitting adjustment for automatic exit of small
quantities of fluid are placed at the head gate or point where irriga-
tion water enters the pasture or other large irrigated areas where there
is mosquito breeding. Emulsible DDT is used and the drip is regulated
according to the quantity of water to be treated. Larvicide drip dispens-
ers must be constantly serviced to prevent clogging and mechanical
difficulties.
Aerosols. Ginsburg 26 points out that the term "aerosol" is derived
from the word sol used in chemistry to denote colloidal dispersions of
solid particles in liquid, gaseous, or solid media; thus if the dispersion
medium is water the system is called hydrosol, if air it is called aerosol,
etc. The term aerosol has recently come into general use to specify cer-
tain insecticidal sprays, dusts, and fumes applied against mosquitoes and
other insects. Aerosols include a variety of natural and artificial systems
such as smoke, clouds, mists, fog, fumes, atomized oil in oil burners,
smoke screens, etc. in which air constitutes the continuous external
dispersion medium and the fine particles form the internal or dispersed
phase.
The main object of aerosol spraying is to disperse insecticides, such
as mosquito larvicides and adulticides, etc., in such a fine state of sub-
division and in such a way that they remain suspended or Hoative in the
air for long periods of time, thereby prolonging their effectiveness. The
work of Ginsburg27 in the development of pyrethrum aerosols for use
out of doors is well known. DDT aerosols are now widely used both
as larvicides and adulticides. Airplane thermal exhaust aerosols are com-

monly employed in rice-field and other large-scale mosquito control
operations. Numerous types of equipment are available for producing
aerosols, among them the engine exhaust of a "jeep," which is inexpen-
sive and effective (Fig. 90).
Residual treatment. Duration of toxicity of residual DDT. Using
a stock solution of 35 per cent technical grade DDT in xylene, with
Triton X-100 as an emulsifying agent, Weathersbee, Arnold, and Hop-
FIG . 90. Aerosol equipment in adulticiding operations. (Photograph by E. A.

Smith.)
kins,28 1948, prepared water emulsions of 1, 2, 3, 4, and 5 per cent and

applied each as a residual spray in five field tests on surfaces in stab1es,
a farmhouse, and doghouses, where Anopheles quadrimaculatus was
by far the predominant species of mosquito, as shown by preliminary
observations. The one per cent concentratioh kept the sprayed surface
free of mosquitoes for 27 days; the 2 per cent for 35 days; the 3 per cent
for 62 days; the 4 per cent for 101 days; and the 5 per cent for 135 days.
The duration, therefore, of toxicity to A. quadrimaculatus of DDT
emulsion applied as a residual spray resulting in a residue varying from
40 to 200 mg per square foot is proportionat~ to the concentration used.
The authors concluded that concentrations of 4 per cent and 5 per cent
(160 to 200 mg per square foot) DDT appcar to be more economical
and more efficient than lesser concentrations.
Indoor residual spraying to destroy adult mosquitoes is accomplished
by treating all the wall surfaces and ceilings, undersurfaces of tables,
beds, etc. with a 5 per cent DDT-oil emulsion spray using either a hand
or air-pressure sprayer. The object is to deposit a residuum of about 200
mg of DDT per square foot. One thorough treatment may last in ex-
cess of three months. Rooms should be ventilated during spraying, and
DDT should not be allowed to fall on food , dishes, or cooking utens ils.
FIG. 91. Knapsack spray pump for

mosquito control operations.
The residual effectiveness of DDT (190 mg of 5 per cent DDT
emulsion per sq ft) applied indoors to adobe walls and straw-thatched
roofs under subtropical conditions is shown in a report by Downs
et al., 29 who noted residual action 29 months after application.
Gray 30 points out that this residual effect of DDT and its analogs
gives us not only a completely new technique in mosquito control, but
an entirely new concept, particularly in the control of disease vectors.
Application of larvicides. Methods of applying larvicides will, of
course, depend upon the materials used, i.e., whether liquids or dusts,
and whether large areas are involved or only a few catch basins. For
relatively small areas or for numerous small and widely separated areas,
hlind application is both economical and convenient (Fig. 91) .
Hand spraying equipment such as the knapsack spray pump has had
almost universal use. This type of equipment partly atomizes the larvi-
cide, resulting in a better spread and a better coverage of the water
surface and is at the same time more economical in the use of larvicide
and in labor. Knapsack sprayers consist of closed metal cylinders of
three to five gallons' capacity, fitted with a brass plunger pump which
forces air into the container until a strong air pressure is built up. To
the outlet pipe, which extends to the bottom of the container, is attached
a piece of flexible hose and a spray nozzle. The spray nozzle is usually
attached to the end of a one-quarter-inch metal pipe (Fig. 92). In
FIG. 92. Knapsack spray pump in use in mosquito control.
operation the tank is filled about three-quarters full with larvicide. A

hand lever extending over one shoulder when the sprayer is strapped on
the back operates the pump plunger. The container should be provided
with a lip about one and one-half inches high extending around and
above the tip of the can to prevent the larvicide from spilling down the
back of the operator. One man with a knapsack sprayer can spray about
five acres per day of eight hours under open field conditions. Power
equipment has largely replaced the knapsack sprayer.
Paris green dusts may be broadcast by hand or by means of a me-
chanical blower, preferably the rotary type. In either case the dusting
should be done with the wind so that the dust floats away from the
workman, thus avoiding exposure to the arsenic which may cause a
dermatitis. Where anopheline marshes of great area are to be treated,
airplane dusting is to be preferred.
Oiled sawdust may be employed to good advantage. The practical
use of this material first came to the notice of the writer (Herms, lac.
cit.) while he was on military duty in 1918. A small detachment of
troops was camped near an abandoned sawmill, and a huge hill of saw-
dust was available for filling numerous small anopheline breeding pools
in the vicinity. It was observed that oil-soaked sawdust gave off a thin
.film of oil for many days. Thereafter liberal use of oil-soaked sawdust
was made whenever it was available. For small-scale projects sawdust
was moistened with fuel oil and broadcast by hand. A small Winding
creek much overgrown with shrubbery and weeds, with steep banks
and many pools, can be successfully treated in this manner.
Power equipment to apply eulicides. Enormous strides have been
made in the development of power equipment designed to give efficient
distribution of the newer insecticides in field practice. Many problems
have had to be met. It has been pOinted out that in order to capitalize
on the increased toxicity and the heretofore unheard-of residual proper-
ties of these products, a much finer break-up of the insecticides is de-
manded as well as a more widespread and uniform distribution. There
is new equipment for producing and distributing aerosols, sprays, and
dusts. Much ingenuity has been employed in adapting various types of
wartime vehicles such as "jeeps" and "weasels" to mosquito control
operations.
Aircraft in mosquito control. The use of aircraft in the distribu-
tion of insecticides for mosquito control became almost world-wide dur-
ing World War II,Dichloro-diphenyl-trichloroethane (DDT) in various
formulations was the insectlcide of choice in these operations. The
American Mosquito Control Association has recently (1948) issued a
valuable illustrated brochure 31 which presents practical information on
when, where, and how aircraft may be used to distribute mosquito in-
secticides, chiefly DDT, and on appropriate methods for appraising re-
sults. All agencies using aircraft for mosquito control purposes should
have this brochure available to the personnel concerned.
Historically the airplane in this connection was first used in Canada
to survey areas in which mosquitoes were breeding (Hewitt,32 1919).
The Alameda County (California) Mosquito Abatement District early
in its formation (1930) used aerial photographs to map its salt-marsh
mosquito breeding areas extending for some 25 miles along the shores
of San Francisco Bay. These aerial photographic maps were made on a
scale of 800 feet to the inch (Herms and Gray, loe. cit.). King and Brad-
ley 33 (1925) and Williams and Cook34 (1927) describe the earliest use
of the airplane in dusting operations against malaria-bearing mosquitoes,
the former near Lake City, Florida, and the latter in the vicinity of
Quantico, Virginia. Although the airplane had been used to apply spray
as early as 1930 (Ginsburg 35 ), it was not until 1944-1945, when DDT
became available, that spraying by airplane came into wide use (Fig.
9S ).
Much of the airplane work done in mosquito abatement districts is
with military trainer biplanes which carry from 80 to 90 gallons of liquid
insecticide or 650 to 750 pounds of dust. The helicopter is growing in
favor because it can be used at any time of day, as it provides a down-
ward thrust to the insecticide; the wing type plane is used principally
in the early morning when the cooling air carries the insecticide down.
The dusting equipment consists of a hopper, a wind-driven agitator,
feed conh'ol gate, and venturi spreader. Spray equipment consists of a
FIG. 93. Use of airplane in spraying operations. (Courtesy of Kern County,

California, Mosquito Abatement District.)
tank, liquid distributing lines (gravity or pressure), and a device for
discharging the liquid into the air. Various types of nozzles are used.
Effective dispersal of sprays has been accomplished by introducing the
liquid spray into the exhaust stream of the engine with resulting atomiza-
tion. Insecticide formulations must be carefully selected on the basis of
the intended purpose.
The use of aircraft makes it economically possible to treat extensive
breeding areas and wild terrain; rice fields may be readily and rapidly
treated, as may also extensive flooded areas caused by river overflow,
etc. The airplane is particularly useful in large-scale "prehatching"
treatments, such as the application of an appropriate insecticide, e.g.
DDT-fuel oil spray, to breeding areas while they are covered with
snow, which on melting in the spring automatically treats all pools har-
boring newly hatched boreal mosquito larvae. Preirrigation treatment
of irrigated pastures may also be carried out by airplane, using water
soluble DDT.
Airplanes are widely used in California in mosquito control opera-
tions. RepOl:ting on tbis type of work at the Sixteenth Annual Conference
of the California Mosquito Control Association (1948) Geib, superin-
tendent of the Kern Mosquito Abatement District, stated that spraying
3 to 4 quarts of a 5 per cent aqueous emulsion of DDT at rates of .3
to .4 pounds of DDT per acre gave satisfactory control of Aedes larvae
on pasture land and river bottom. Using Stearman PT-17 biplanes owned
. and operated by the district the average cost per acre during the 1947
season was 66 cents.
Repellents. There is little doubt that through the centuries man has
tried to protect himself from mosquito bites by using herbs and smoke
smudges, probably with little success. Oriental punk has long been
widely employed; considerable quantities were used in the tropics by
the armed forces of both sides during World War II. During World
War I troops who were forced to operate in mosquito-infested regions
were often advised to smear themselves with so-called "bamber oil"
(among other ointments), a mixture of citronella oil, kerosene, and
coconut oil, with a little carbolic acid added. Relief was probably not
marked. Early during the World War II period steps were taken to
develop repellents with some real effect. Granette 36 at Rutgers Univer-
sity (New Jersey) and Knipling and Dove 37 at the United States Bureau
of Entomology and Plant Quarantine, Orlando, Florida, laboratories, in
cooperation with the armed forces and other agencies (some commer-
cial), developed the widely used repellents known as Rutgers 612,
dimethyl phthalate, and indalone. Rutgers 612 is most effective against
Aedes species, while dimethyl phthalate is most effective against Anoph-
eles quadrimaculatus. Granette in a personal communication advised
the author that effectiveness of repellents varies considerably with the
various anopheline species. Indalone is an effective repellent against
Hies such as Stomoxys. All of these repellents are more or less effective
for three to five hours and they remain effective for a much longer time
(about a week) when sprayed liberally (50 to 100 cc) on clothing.
Screens to exclude mosquitoes. Where there is no control of mosqui-
toes or only incomplete control, particularly in the presence of malaria-
bearing species, sleeping quarters, living quarters, porches, etc., should
be adequately screened against intruding mosquitoes. The best size mesh
for all purposes is No. 18, i.e., 18 strands to an inch. The screens must
be suited to exposure and to the climatic conditions of the situation
where they are used. They must be protected or reinforced against
mechanical breakage at points of stress. Painting screens with a 5 per
cent oil emulsion of DDT is a good practice.
Duck clubs. In most states the duck-shooting season does not begin
until November first or later. By that time cold weather usually stops
mosquito breeding. However, many duck clubs start flooding their
duck ponds long before this, perhaps to have the ponds ready to attract
the earlier migrants. These slowly filled shallow ponds may prove to be
a mosquito menace. Some clubs keep their ponds well flooded through-
out the year, and if the banks are steep and top minnows have access
to all parts of the pond, there is no mosquito problem. The ponds which
are most difficult to handle are those that are drained off in January or
February after the duck season is over, and are allowed to remain dry
during the summer, being again flooded in late summer or early autumn
while the weather is still warm. Breeding is sure to occur. Aedes eggs,
such as those of Aedes dorsalis (Meig.), froIll preceding years promptly
hatch, and a plague of mosquitoes soon appears.
Neff (1935), of the United States Bureau of Biological Survey, states
in the Proceedings of the Sixth Annual Meeting of Mosquito Abatement
Officials in California, p. 16, "Impounding of water, or the maintenance
of a constant water level by means of tide gates that permit an equalized
flow of water through the salt marshes, seems to be the method of con-
trol which has aroused little or no adverse comment from biologists.
This method also offers opportunity for experimental work on planting
of better wild life food plants than may formerly have existed on the
areas with fluctuating water levels."
Although expensive, pyrethrum should be used instead of oil where
a larvicide becomes necessary. Neff (Zoc. cit.) states that pyrethrum larvi-
cide has "no known deleterious effect upon wild life."
The following rules in dealing with mosquito control in connection
with duck clubs are suggested:
1. Continuous all-year Hooding of ponds is permissible and approved, pro-
vided the ponds are stocked with "mosquito fish" at all times.
2. Intermittent maintenance of ponds is permissible, provided, (a) the
water is effectively removed early in the spring before breeding occurs, and
(b) the water is not put in in the autumn until the weather is cool enough to
prevent mosquito breeding.
3. Ponds must have sound, tight banks and bottoms to prevent wet areas
due to seepage.
4. Ponds must have sufficient depth throughout to permit mosquito fish to
penetrate freely all parts.
5. Ponds must not be overgrown with vegetation, especially along and
near the margins, so that "mosquito fish" may have free access to all parts.
6. If the water is pumped, the supply and equipment should be adequate
to fill the ponds with reasonable speed.
7. Duly authorized inspectors of mosquito abatement districts in which the
MOSQUITO ABATEMENT 2B9
ponds occur should be permitted to inspect the area frequently to determine

whether or not mosquito breeding is taking place.
B. If in spite of all precautions mosquito larvae do occur, a larvicide, pref-
erably a pyrethrum emulsion, should be applied, but only where breeding is
actually in progress.
Mosquito control and wild life conservation. Mosquito abatement
operations if intelligently conducted need not be detrimental to wild
life, though no doubt they have been so at times. In conducting control
operations iR suburban and rural areas an understanding of wild life
ecology is urged, and a modification of measures to suit the situation is
necessary. It is regrettable if wild life has been harmed; but there have
also been unfounded complaints on the part of misinformed and intol-
erant wild life conservationists which have made it distinctly difficult
for mosquito abatement officials to perform their proper function. It is
important that the seemingly divergent viewpoints of conservationists and
duck clubs and mosquito abatement officials be harmonized. No doubt
each group will need to make reasonable concessions.
Cooperation between responsible mosquito abatement officials and
the representatives of wild life interests is only made difficult when one
side or the other sets itself upon a pedestal. Our properly trained experts
in mosquito control are just as truly biologists as are wild life author-
ities, the only difference is usually in the fact that the former are trained
in the field of invertebrate zoology and the latter in the field of verte-
brate zoology. Both should be ecologists and have training in limnology.
Fortunately, most of our authorities in the field of mosquito control have
had training not only in the ecology of aquatic invertebrates but for
obvious reasons also in vertebrate ecology. Entomologists trained in
mosquito control and ecology, and wild life conservationists equally well
trained in ecology ought to be able to see "eye to eye" as biologists.
Working thus together as biologists on an equal footing, the aims and
objectives of both sides will be advanced.
The almost universal use of the newer insecticides, DDT and others,
in agricultural pest control operations as well as in mosquito control has
brought to a focus the problem of the tolerance of each wildlife species,
both aquatic and terrestrial, and the indirect effect as pertains to the food
of the several species involved in treated areas. Cope 38 states that "in the
case of DDT and wildlife (exclusive of fish), something over one pound
per acre seems to be a safe level, while for fish a figure of 0.2 pound per
acre would apply." For anopheline control 0.1 pound per acre of DDT
is usually adequate and somewhat larger doses for culicine larviciding.
For benzene hexachloride it is indicated that up to one pound per acre
may be safe for wildlife, fish, and fish food organisms. Chlordane appears
to be damaging to some fish at one pound per acre. TDE appears to be
about one-fifth as toxic to warm water fishes as DDT.
Hess and Keener 3D (1947) made field tests with a 20 per cent solu-
tion by weight of technical grade DDT in a methylated napthalene
(Velsicol NR-70) applied by airplane in the form of a thermal aerosol.
The rate of application was approximately 0.1 pound of DDT per acre.
They report the following results:
The effect of routine treatments on an area basis was almost complete

elimination of anopheline mosquitoes and surface Hemiptera and a significant
reduction in culicine mosquitoes with no evidence of reduction of other forms.
Observations in areas at the end of a full season of 16 applications at the rate
of 0.1 lb. DDT per acre indicated that surface Hemiptera had been almost
completely eliminated and mosquitoes were considerably reduced, but other
forms were not significantly affected. Surface Hemiptera are not important as
fish food and because of their predaceous habits may actually compete with
fish for food. On the basis of these studies, it is therefore concluded that DDT
applied in the manner indicated does not have any significant injurious effect
upon the total population of fish food organisms-it is also concluded that the
DDT treatment had no injurious effect upon the resident fish population.
Natural enemies of mosquitoes. A review and summary of the litera-

ture dealing with "Predators of the Culicidae" by Hinman 40 indicates
that, excluding fish, the chances of finding satisfactory predators is not
very encouraging. Among the natural enemies of mosquitoes few are so
frequently referred to as dragonflies (also known as "mosquito hawks"),
bats, and surface-feeding fish. Dragonflies, order Odonata, are preda-
ceous in both the nymphal and adult stage. The aquatic nymphs are
commonly found in quiet, shallow, permanent pools suitable also for
mosquito breeding, and both may flourish in the same pool in spite of the
fact that the dragonfly nymphs, usually relatively few in number, may
feed on mosquito wrigglers. Since the nymphs feed in the mud and debris
at the bottom, probably few wrigglers are captured. If the wrigglers are
easily available, the nymphs will feed on mosquito larvae voraciously.
Warren 41 reported a nymph of Pantala consuming 75 full-grown mos-
quito larvae by seven o'clock in the evening, which he had placed in a
glass half-full of water in the morning. Adult dragonflies are exceedingly
adept at capturing mosquitoes on the wing just before and at sunset.
However, here again the number of dragonflies, which also feed on
other insects, is no match for the mosquitoes. Dragonflies do not fly at
night when night-flying mosquitoes are on the wing.
Bats are insectivorous and feed freely on mosquitoes; as many as 250,
it is said, may be captured by one bat in a night, but with many other
species of crepuscular and night-flying insects available, bats are not
effective enough to be a large factor in control, even though one might
tolerate them in large numbers near the home. Bat roosts, however, have
been established to accommodate bats for the purpose of mosquito
control.
Fish of various species have been advocated for many years. In
Nature for December, 1891 (pages 223-224), there is this item: "An
Englishman living on the Riviera, according to a correspondent, having
been troubled by mosquitoes, discovered that they bred in the large
tanks kept for the purpose of storing fresh water, which is rather a rare
commodity at this Mediterranean resort. He put a pair of carp in each
tank and succeeded in this way in extirpating the insect pest."
Howard (1910, loco cit.) refers to the control of mosquitoes by gold-
fish in an ornamental aquatic garden near Boston:
I took from the pond a small goldfish about three inches long and placed it
in an aquarium where it could, if it would, feed upon mosquito larvae and still
be under careful observation. . . . On the first day owing perhaps to being
rather easily disturbed in its new quarters, this goldfish ate eleven larvae only
in three hours, but the next twenty-three were devoured in one hour; and as
the fish became more at home, the 'wigglers' disappeared in short order when-
ever they were dropped into the water. On one occasion twenty were eaten in
one minute, and forty-eight within five minutes. This experiment was fre-
quently repeated and to see if this partiality for insect food was characteristic
of those goldfish only which were indigenous to this locality experimented
with, some said to have been reared in carp ponds near Baltimore, Maryland,
were secured. The result was the same. . . .
The most useful of all fishes for mosquito control is the top minnow
or mosquito fish,'2 Gambusia affinis (Baird and Girard), a hardy, rapidly
breeding, prolific surface-feeding fish which within its range normally
inhabits shallow water suitable for mosquito propagation. It is vivip-
arous and may produce as many as six to eight broods in a season with
an average of 40 to a brood. The size of the fish ranges from 1 ~ inches
in length in the male to nearly 2 inches in the female. This fish is easily
propagated and adapts itself to a variety of conditions with ease. It has
been introduced into various parts of the world, even over great dis-
tances, for example, from Texas to the Hawaiian Islands and thence
to the Philippines. Transportation of top minnows can be done satis-
factorily in 10-gallon milk cans with tops punched with holes and water
kept below the point where the top of the can begins to narrow. Although
as many as 500 fish may be transported for an hour's trip with only
moderate loss, not over 200 young fish per can should be shipped on
longer trips, and special care must be exercised to remove dead fish at
intervals and freshen the water.
For garden pools 10 square feet in diameter, 20 top minnows will
be ample, and no artificial feeding will be necessary. The Gambusia
will more or less regulate their own numbers according to the food sup-
ply available.
Top minnows will evidently not feed on mosquito larvae when these
are motionless, hence are not markedly effective in the control of mos-
quitoes whose larvae are sluggish; e.g., the usually motionless larvae of
Anopheles quadrimaculatus Say do not attract the attention of top min-
nows as readily as do the more active larvae of A. m. freeborni Aitken;
hence the minnows are not so effective in control.
Other fish which have been found useful are Heterandria formosa,
Fundulus diaphanus, and Fundulus dispar for fresh wa,ter, and Cyprino-
don variegatus, Fundulus heterocliteus, Fundulus similis, Fundulus ma-
;alis, and Lucania para for salt or brackish water. To these may be added
a number of species as listed by various authors, notably Hegh,43 Rad-
cliffe/ 4 and Hamlyn-Harris. 45 The International Health Board of the
Rockefeller Foundation has issued ( 1924) a comprehensive treatise
entitled "The use of fish for mosquito control."
In Guayaquil, Ecuador, the problem of the yellow fever mosquito was
solved, according to Connor,46 by the use of fish. Connor states that the
domestic water supply is delivered to the houses daily and is stored in
tanks and other receptacles, there being at the time of his writing about
7,000 of the former and 30,000 of the latter, such as barrels, oil cans,
earthenware bowls, etc. In these various containers yellow fever mos-
quitoes developed in countless numbers. Experimentation with several
species of fish finally resulted in the selection of the "chalaco" (Dormi-
tator latifrons, family Gobiidae). These fish, furnished to the Yellow-
Fever Service by local fishermen, were placed in a specially prepared
well, the conditions of which approximated those of the stream from
which the fish were taken. After a few days the fish were removed to
a second well, the water of which was the same as that used by the city.
Connor writes further as follows:
The fish are then taken from the wells and placed in tins or pails and
delivered to the inspectors. Instructions have been given to each inspector that
every fresh-water container in his district is to be supplied with one fish, re-
gardless of the presence or absence of mosquito larvae in the container at that
time. The public is encouraged, personally, by notices in the newspaper, and
by the inspectors themselves, to exercise reasonable care in protecting the £sh.
The public ,of Guayaquil has responded in a whole-hearted manner to the
requests of the Yellow-Fever Service, and many families have in their posses-
sion at this time the identical fish which was given them to mosquito-proof
their water container nearly eighteen months ago.
More than 30,000 water receptacles have in this way been purged of
mosquito larvae in a relatively short time and at a minimum of expense. With
the continued use of fish it is believed that the yellow-fever mosquito can be
reduced to such small numbers that, should a few cases of the disease be in-
troduced into the community, it would not spread.
Transportation of exotic mosquitoes by airplane. The accidental im-
portation of exotic mosquitoes may result in severe epidemics of disease
with heavy loss of life and may necessitate expensive campaigns to root
out the evil. Particularly noteworthy is the malaria epidemic in Brazil
following the introduction of Anopheles gambiae Giles from Africa,
referred to in an earlier chapter as described by Soper and Wilson (lac.
cit. p. 228). Such dangers, particularly importation of the vector of yellow
fever, have long been recognized as possible through air transportation.
Various amendments have been made to the "International Sanitary
Convention for Aerial Navigation of April 12, 1933," signed by the United
States in April, 1934, according to Miller, Burgess, and Carpenter47
( 1947). Article 54, (a 1944 amendment) reads, "In view of the special
risk of conveying insect vectors of malaria and other diseases by aircraft
on international flight, all such aircraft leaving affected areas will be
disinsected . . . further disinsectization of the aircraft on or before
arrival may be required if there is reason to suspect the importance of
insect vectors."
There are said to be about 35,000 airplanes entering the United States
annually from foreign ports, 80 per cent of which are from the West
Indies and Latin America. Scheduled flight times from all parts of the
world are short enough to enable stowaway mosquitoes to arrive alive
in the United States. Southern and California ports of entry, with more
favorable year-round climatic conditions, are regarded as offering the
most likely points of invasion. Although the enormous wartime move-
ment of aircraft (Herms, 48 1946) showed a tremendous potential of
insect transport particularly at Honolulu, strict quarantine regulations in-
cluding airplane disinsectization administered by the United States Navy
proved very effective. Freon-pyrethrum-DDT aerosol "bombs"-con-
tinue to be used for disinsectization purposes. Automatic centrally con-
trolled devices for releasing aerosols in aircraft are now being tested.
Recently Rosen, Reeves, and Aarons 49 reported the presence of Aedes
aegypti (Linn.) on Wake Island. Reeves (1949) at the 8th Pacific Science
Congress presented a paper on "Possible Recent Introductions of Mos-
quito Vectors of Human Disease in the Central Pacific."
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15. Horsfall, William R., 1942. Biology and Control of Mosquitoes in the
Rice Areas. Fayetteville: Univ. Arkansas, in Agric. Exper. Sta. Bull., no. 427.
46 pp.
16. Rao, B. A., and Sweet, W. C., 1937. "Paris green and paddy," Records
Malaria Survey India, 7:185-89.
17. Hill, Rolla B., and Cambournac, F. J. C., 1941. "Intermittent irriga-
tion in rice cultivation, and its effect on yield, water consumption and
Anopheles production," Am.]. Trap. Med., 21:123-44.
18. Russell, Paul F.; Knipe, Fred. W.; and Rao, H. Ramanatha; 1942.
"On the intermittent irrigation of rice fields to control malaria in South India,"
J. Mal. Inst. India, 4:321-40.
19. Cambournac, F. J. C., and Fonseca da, A. E., 1948. "Experiments
on the control of anopheline larvae and malaria in the rice-growing regions
of Portugal," Proc. 4th Internat. Congo Trap. Med. & Malaria (Abstracts),
Washington, D. C.
20. Gillette, H. P. S., 1948. "The control of Btomeliad malaria in Trinidad,
B. W. 1.," Proc. 4th Internat. Congo Trap. Med. & Malaria (Abstracts),
Washington, D. C.
21. Aarons, Theodore, 1948. "Cemetery mosquito control by aerosol,"
Proc. and Papers 16th Annual Conf. Calif. Mosq. Control Assn., Berkeley,
pp. 84-85.
22. Le Prince, J. A., 1915. "Control of Malaria; oiling as an antimosquito
measure," U. S. Public Health Service, Pub. Health Rep., 30:599-608.
23. Barber, M. A., and Hayne, J. B.; 1921. "Arsenic as a larvicide for
anopheline larvae," U. S. Public Health Service, Pub. Health Rep., 36:3027-34.
24. Vannote, R. L., and Ginsburg, J. M., 1931. "Practical application of
pyrethrum mosquito larvicide," Proc. 18th Annual Meeting, New Jersey Mosq.
Extermination Assn., pp. 111-120.
25. Yust, Harold R., 1947. "DDT to control Anopheles farauti on Espiritu
Santo, New Hebrides Islands," J. Econ. Entomol., 40:762-68.
26. Ginsburg, Joseph M., 1943. "Aerosol sprays for killing and repelling
mosquitoes," Proc. 30th Annual Meeting, New Jersey Mosq. Extermination
Assn., pp. 211-17.
27. ---, 1935. "Larvicides and a method of temporary protection from
adult mosquitoes in limited areas," Proc. 25th Annual Meeting New Jersey
M osq. Extermination Assn.
28. Weathersbee, Albert A.; Arnold, F. T. Jr.; and Hopkins, Julian P.;
1948. "Observations on the duration of toxicity of DDT to Anopheles quadri-
maculatus Say under field conditions," J. Nat. Malaria Soc., 7:138-43.
29. Downs, W. G.; Iris, R. c.; and Gahan, J. B.; 1948. "Residual effec-
tiveness of DDT in the third season after application," Am. ]. Trop. Med.,
28:741-745.
30. Gray, H. F., 1948. "Some newer ideas in mosquito control," Bol. Ofic.
san. panam., 27:321-25.
31. American Mosquito Control Association, 1948. The Use of Aircraft
in the Control of Mosquitoes, in Bull. no. 1., ed. P. F. Russell, G. H. Bradley,
A. D. Hess, J. A. Mulrennan, and H. H. Stage. 46 pp. + 45 plates.
32. Hewitt, C. G., 1919. "The use of the aeroplane in entomological
work," Agric. Gaz. Canada,6:877.
33. King, W. V., and Bradley, G. H., 1925. "Airplane dusting controls
malaria mosquitoes," Aero Digest, 7:652-53.
34. Williams, L. L., Jr., and Cook, S. S., 1927. "Paris green applied by
airplane in the control of Anopheles production," U. S. Public Health Service,
35. Ginsburg, J. M., 1931. "Airplane application of larvicide on mosquito
breeding places inaccessible from land," New Jersey Agric. Exper. Sta. Annual
Rept, (1930-1931), 19:173-76.
36. Granett, Philip, 1943. "The significance of the development of mos-
quito repellents for the protection of military and civilian populations," Proc.
30th Annual Meeting, New Jersey Mosq. Extermination. Assn., 203-10.
37. Knipling, E. F., and Dove, W. E., 1944. "Recent investigations of
insecticides and repellents for the armed forces," J. Econ. Entomol., 37:477-80.
38. Cope, Oliver B., 1948. "Toxicities and tolerances of new insecticides
in relation to wildlife and fish," Proc. and Papers 16th Annual. Conf. Califor-
nia Mosq. Control Assn. (Berkeley), pp. 26-29.
39. Hess, A. D., and Keener, G. G. Jr., 1947. "Effect of airplane distributed
DDT thermal aerosols on fish and fish food organisms," J. Wildlife Manage-
ment, 11:1-10.
40. Hinman, E. H., 1934. "Predators of the Culicidae (mosquitoes),"
]. Trap. Med. & Hyg., 37:129-34, and 145-50.
41. Warren, Alfred, 1915. A study of the Food Habits of the Hawaiian
Dragonflies or Pinau with Reference to Their Economic Relation to Other In-
sects. "College of Hawaii Publications," Bull. No.3. 45 pp.
42. Seale, A., 1917. "The mosquito fish, Gambusia affinis (Baird and
Girard), in the Philippine Islands," Philippine]. Sc., 12:177-89.
43. Hegh, E., 1918. Comment nos planteurs et nos colons peuvent-ils se
proteger contre les moustiques qui transmettent des maladies. Report Minister
of Colonies, Service of Agriculture of Belgium, no. 4. 200 pp.
44. Radcliffe, L., 1915. Fishes Destructive to the Eggs and Larvae of
Mosquitoes. Washington, D. C.: Dept. of Commerce, in Bur. Fisheries, Cire.
17, pp. 1-19.
45. Hamlyn-Harris, Ronald, 1929. "The relative value of larval destructors
and the part they play in mosquito control in Queensland," Proc. Roy. Soc.
Queensland, vol. 40. 38 pp. + 8 plates.
46. Connor, Michael E., 1921. "Fish as mosquito destroyers." Natural
History, 21:279-81.
47. Miller, Albert; Burgess, Robert W.; and Carpenter, Stanley J.; 1947.
"Potentialities of transportation of exotic anophelines by airplane," J. Nat.
Malaria Soc., 6:227-43.
48. Herms, W. M., 1946. "Wartime aviation quarantine. Pests and their
control," J. Pest Control Industry, 14:24-25.
49. Rosen, L.; Reeves, W. C. and Aarons, T. 1948. "Aedes aegypti on
Wake Island," Froc. Hawaian Entomolog. Soc. (for year 1947), 13:255-56.
CHAPTER XIV
._.
HORSEFLIES, DEER FLIES,

AND SNIPE FLIES
A. HORSEFLIES
Family Tabanidae
Tabanidae. To this large cosmopolitan family of the Order Diptera,
comprising about 2,500 species, belong the avidly bloodsucking Hies
commonly known as horseHies, gaclRies, breeze Hies, greenheads, deer
flies, or mango Hies. These Hies are usually quite large and heavy-bodied
(Fig. 94), measuring in length from 7 to 10 mm in the smaller species
Flc. 94. The black horsefly, Tabanus atratus. (Left) male, (right) female.
X 1..5. (Photograph by Hine.)
to from 20 to 30 mm in the larger species. They are strong Hiers and

notorious pests of horses, cattle, deer, and many other warm-blooded
animals and are at times annoying to man, particularly the persistent
members of the genus Chrysops (Fig. 95). The males feed on vegetable
sap and some may suck the juices of soft-bodied insects, but they do not
bite. The eyes are very large and widely separated (dichoptic) in the
females, and contiguous (holoptic) in the male. The antennae are short
297
(though fairly long in some genera such as Chrysops) and are porrect,
consisting of three joints, the terminal segment or flagellum being elon-
gate and annulated, without an arista. The wing venation (Fig. 12B) is
simple and characteristic. The mouth parts of the females are blade-like
and func tion as cutting instruments. The metamorphosis is complete
(Fig. 96).
The family is divided into two subfamilies, the Pangoniinae, which
includes all Tabanidae with apical spurs on the hind tibiae and in which
ocelli are usually present, e.g. , the species belonging to the following
FIG. 95. A deer fly (Chrysops) in the act of

oviposition. Note also an egg mass farther down
on the leaf. X 1. (Photograph by Hine.)
genera: Silvius, Chrysops, Goniops, Apatolestes, and Pangonia; and the

subfamily Tabaninae, in which the apical spurs and ocelli are lacking,
as in Tabanus (Fig. 97) and Chrysozona .
Since most of the species are aquatic or semiaquatic in breeding
habit (some are able to develop in moist earth, leaf mold, and rotting
logs), the eggs are deposited in layers on objects over water or moist
situations favorable for the larvae, such as overhanging foliage, project-
ing rocks, sticks, and aquatic vegetation. The narrow cylindrical eggs
(1.0 to 2.5 mm long) vary in numbers from 100 to 1,000, and are de-
posited commonly in layers covered with a waterproof secretion which
also binds the eggs together tightly.
The larva (Fig. 98) has a slender, cylindrical, contractile body taper-
ing at both ends and consisting of 11 segments (interpreted also as a
head and 12 segments, the last very short). The head is small and ,re-
tractile, with pOinted mandibles capable of inflicting a sharp bite; at
the posterior end is situated a tracheal siphon which telescopes into the
anal segment. The pupa (Fig. 98), resembling naked Lepidoptera, is
obtect, abruptly rounded anteriorly, tapering posteriorly, with wing and
leg cases closely attached to the body; the abdominal segments are free
and about equal in length, segments two to seven each bear a more or
less complete ring of spines near the posterior third. The adult fly
emerges from the pupal case through a split or slit along the dorsum of
the thorax. The Tabanidae therefore belong to the Orthorrhapha, i.e.,
I
1
FIG .
\
96. Tabanus punctifer, Egg mass all willow leaf, larva , pupa, and adult
female fly.
the flies escape from the pupal skin through a T-shaped opening. In
contrast the adults of the Cyclorrhapha such as houseflies escape from
the pupal case through a round opening made by pushing off the anterior
end. In pushing off this end of the puparium the housefly and its allies
use a frontal bladder-like structure known as a ptilinum, absent in the
Orthorrhapha such as the Tabanidae.
Students concerned with the Tabanidae are referred to such mono-
graphic works as those of Osten Sacken,l Hine,2 Enderlein,3 Kr6ber,4
and Stone. s
Breeding habits and life history. The eggs, numbering up to nearly
a thousand, are deposited during the warmer months of the year in com-
pact layers on objects such as the leaves of willow and emergent aquatic
vegetation which grows from or overhangs swampy areas, ponds, etc.
(Fig. 95). The incubation period is greatly influenced by weather con-
ditions, but during midsummer the usual range is from five to seven
days. On hatching the larvae fall to the surface of the water, upon mud
or moist earth, in clumps and quickly drop to the bottom or burrow
individually into the wet or damp earth, where they begin feeding at
once on organic matter. Many species are predators, sucking the juices
of insect larvae, crustacea, snails, earthworms, and other soft-bodied
animals; cannibalism has been observed in several species. The larvae
of Tabanidae are commonly encountered buried in the mud along the
~PV.
/./ ' TO( Un. Emp.
FIG. 97. Morphological details of a femal horsefly, Tabanus pttllctifer.

Explanation of abbreviations: Ant., antenna; Ai., alula; Emp., empodium; Fm.,
femur; HI., haltere; L. Prsc ., lobe of pre scutum (notopleuron); Dc., occiput;
Pv., pulvilli; Sc., scutum; Scl., scutellum; Tar., tarsi; Th., tibia; Te., tergite;
Tg., tegula; Un ., ungues; Vx., vertex.
edges of marshy ponds, roadside ditches, and the overflow from rice
fields; the writer has found numerous larvae of Tabanus gilanus Town-
send in the mud at the edge of salt marshes. The larvae of certain species
may be found in moist leaf mould and debris.
The larvae grow rapidly during the summer and autumn, and very
slowly if at all during the winter in the single-brooded species, attain-
ing full growth in the following early spring. There is some uncertainty
as to the number of larval instars; it varies, according to some authors
from four to nine, the first molt evidently taking place shortly after hatch-
ing. An excellent account of the early stages of Tabanidae may be found
in the work of Marchand. s •
When the full-grown larva prepares to pupate, it moves into drier
earth, usually an inch or two below the surface, and in a day or two the
HORSEFLIES, DEER FLIES, AND SNlpE FLIES 301
pupal stage is reached. This stage requires from two to three weeks,
varying with the species. Stone 7 reports that most of the Chrysops species
emerge in less than two weeks, even in as short a time as five days. The
flies emerge from the pupa at the surface, the wings soon unfold, and
the insects take refuge among nearby foliage or rest on objects near at
hand; in a short time they begin to feed, the females seeking blood and
FIG. 98. Lateral view of larva (left) and

ventral view of pupa (right) of Tabanus
gilanus.
the males feeding on flower secretions and vegetable juices where one
may find them by sweeping with a net.
Much important information based on rearing experience with many
species may be obtained by consulting the various publications by
Schwardt, among them Horseflies in Arkansas.s In an earlier paper (Re-
search paper 219, Journal Series, University of Arkansas) Schwardt re-
ports life-history records based on 202 individuals of Tabanus lineola
Fabr., a common North American species of "greenhead," viz., average
incubation period 4 days, average larval period 48.8 days, average pupal
period 8.1 days, average preoviposition period 9 days, total develop-
mental period averaging 96.9 days.
In the Sierra Nevada and other mountain ranges horseflies breed in
great numbers at elevations of 8,000 to 9,000 feet in soggy ground caused
by springs, and water from melting snow in the summer. Deer and other
wild animals suffer much from the bites of these flies.
Bites. The horseflies have broad blade-like mouth parts (Fig. 24),
which inflict a deep, painful wound, causing a considerable flow of
blood; and owing to their intermittent biting habits there is danger of
infection.
Webb and Wells: working on T. phaenops O. S. in western Nevada,

estimated that eight flies feeding to satiety would consume a cubic centimeter
of blood. On this basis they calculated that 20 to 30 flies feeding for six hours
would take an average of at least 100 cc of blood. This would amount to
approximately a quart in ten days. Philip,'o working in Minnesota, derived a
larger estimate of blood loss. Basing his figures on a somewhat heavier infesta-
tion than that in Nevada, Philip placed the daily loss of blood for each animal
at 300 cc, or nearly one-third of a quart. Neither of these estimates includes
the blood which exudes from the bite after the fly leaves. Philip, however,
calls attention to this additional loss. The horseflies most abundant in Arkansas
are comparable in size to the species on which these estimates of blood loss
were made, and the infestation is often heavier than 50 flies per animal.
(Schwardt, 1936, loco cit.)
In describing an outbreak of gadflies in Kentucky, Garman l l writes:

Beef cattle had lost an average of 100 pounds as a result of the constant
annoyance from them. . . . On cattle I counted from ten to nineteen. On
mules and horses in harness they were a constant annoyance and even hogs
were not exempt. Seven of the flies were counted on the exposed side of one
of these animals lying in a puddle.
The persecuted stock appeared to have given up fighting their enemies
and allowed them to have their way. The switch of a cow's tail was observed
to pass over the backs of clinging flies without causing them to move. . . .
During the middle of the day animals suffered so much that they refrained
from grazing at all, either standing close together about the barn or else lurk-
ing singly in thickets or standing in pools formed by small streams.
Relation to anthrax. Anthrax, also known as malignant pustule or

carbuncle, wool sorter's disease, or charbon, is caused by Bacillus anthra-
cis. Nearly all species of domesticated animals and man are susceptible;
the herbivora and rodents are most liable to infection.
After the inoculation of the organism into the animal the incubation
period is from three to six days. The bacilli are seen in the blood stream
in advanced cases as chains of rod-shaped bodies. Entrance to the body
is gained mainly in one of three ways: (1) through lesions or pricks,
i.e., inoculation, producing local anthrax or malignant pustule; (2) by
inhalation of the spores, producing pulmonary anthrax; and ( 3 ) by
ingestion with food, producing intestinal anthrax.
The horseflies are decidedly intermittent in their biting habits and
inflict a dennite lancet-like prick from which blood exudes so that the
probOSCis becomes soiled. The flies will bite sick animals as well as
healthy ones.
Horseflies relate directly to the first mode of infection (inoculation),

and it is not altogether improbable that an epidemic of anthrax might
thus be started and could be spread. Nuttall cites Bollinger (1874), who
captured horseflies on a cow dead from anthrax and saw the bacilli in
preparations made from the stomachs and intestines of the insects. Two
rabbits inoculated therewith died of anthrax.
Mitzmainl~ (Mayne), in a series of experiments with Tabanus striatus
Fabr., showed that direct mechanical inoculation of guinea pigs could
be readily effected by the bite of the fly. He permitted the flies to feed
on an inoculated guinea pig shortly before its death and transferred
the flies soon thereafter (45 seconds to 30 minutes) to a healthy guinea
pig. The death of these animals followed in from three to three and
one-half days.
Instances are recorded in which apparently the simple bite of the
infected fly was all that was needed to produce malignant pustule in
FIG. 99. Trypanosoma c1)ansi, the

causal organism of surra. (After
Yutuc.)
humans. Several veterinarians have related instances to the writer in

which this had occurred, notably one case in which a man was in the
act of burying a cow dead of anthrax when he was bitten severely on
the back of the neck by a horsefly and in due time developed a malignant
pustule. Nuttall also cites a number of similar instances.
Surra is a highly fatal disease of equine animals in southeastern Asia
and many of the islands of the Malay Archipelago. It was first described
(1885) from India by Evans. The causal agent is Trypanosoma evansi
Steel. Yutuc,'" who reported on the first outbreak of surra in British
North Borneo, states that it is highly fatal to dogs; carabaos are evidently
not severely affected as a result of an infection and thus serve as reser-
voirs. In India camels also serve as reservoirs. In the Philippine Islands
and elsewhere virulent outbreaks of surra occur among bovines.
The trypanosomes (Fig. 99) are found in the blood of infected ani-
mals, and especially in the lymph of swollen glands, from the beginning
of the first symptoms. During the early stage of the disease, according
to Yutuc, practically no clinical manifestations are visible save a variable
appetite and an intermittent fever; there is progressive emaciation and
edema of the abdomen and genitalia. Guinea pigs, white mice, and
monkeys are highly susceptible laboratory animals.
Mitzmain H (Mayne) succeeded in transmitting the infection from
animal to animal through the agency of a horseHy, Tabanus striatus
Fabr.
In a series of experiments in which Tabanus striatus Fabr. was used,
he first allowed the flies to bite an infected guinea pig or horse for not
more than one minute, usually 45 seconds, and then transferred them to
a healthy animal where they were allowed to complete the meal without
interruption. An interruption of five seconds to three minutes was neces-
sary to transfer the Hies from animal to animal. The horses and mule
employed in these experiments were kept in a screened stabJe for from
six to eight months previously, and the monkeys, guinea pigs, and rabbits
in Hy-screened cages for about 90 days. In all cases the animals were
examined frequently (blood examinations were made) and declared
surra-free at the time the experiments began.
Flies bred from eggs were allowed to bite a guinea pig which had
been inoculated with blood from a carabao which had been infected with
surra for nearly a year previous to the test. Three Hies were applied
individually in tubes to the surra-infected guinea pig and allowed to
feed from 45 seconds to one minute and 30 seconds, after which they
were transferred to a monkey and allowed to feed until satisfied, i.e.,
from 5 to 21 minutes. The first rise in the monkey's temperature, 40.10 C,
occurred on the eleventh day, accompanied by the presence of a few
trypanosomes in the peripheral circulation; the trypanosomes increased
in numbers until the death of the monkey on the twenty-fifth day.
Blood from the heart of this monkey was inoculated into a horse
and two guinea pigs. The latter showed infection on the eighth and
ninth days respectively, and the horse on the seventh day. Two Hies were
permitted to bite this horse, the insects being interrupted in their biting
in from 40 to 45 seconds and then transferred to a healthy horse, where
the feeding was completed. The latter animal showed numerous tryp-
anosomes in its circulation on the ninth day. Thus positive results were
secured in both a monkey and a horse.
Blood from this newly infected horse was inoculated into a mule,
two monkeys, and two guinea pigs, all of which became infected in due
time, both monkeys dying on the fourteenth and fifteenth d!ys respec-
tively.
A second series of experiments was carried on with captured flies,
which were allowed to bite the surra-infected horse mentioned above
and later a healthy horse, similar feeding methods being observed. This
experiment also gave positive results, as did blood inoculations into
monkeys and guinea pigs.
In order to eliminate the possibility of hereditary transmission of
trypanosomes in the flies a further experiment was conducted with 74
flies, hatched from eggs of a fly which, 1?revious to egg depOSition, had
fed on a surra-infected monkey; the 74 flies were allowed to bite a
healthy monkey during a period of two weeks with negative results.
Mayne concludes that the « • • • contaminated Iabellum of the fly
does not appear to be a factor in the conveyance of infection. The maxi-
mum length of time that Trypanosoma evansi (Steel) has been demon-
strated microscopically in the gut of this species of fly after feeding on
infected blood is thirty hours; the organisms were found in the fly's
dejecta two and one-half hours after biting the infected animal; and
suspensions of flies, when injected subcutaneously, were found infective
for animals for a period of ten hours after the flies had fed on infected
blood."
In a letter to the writer under date of November 18, 1913, Mayne
states that" ... infection is not transferred by Tabanus striatus Fabr.
later than twenty minutes after the infective meal. The longest time I
have succeeded in inducing flies to transmit was fifteen minutes and all
results from twenty minutes to forty-eight hours were entirely negative.
This despite the fact that trypanosomes survive in the intestinal tract
of T. striatus Fabr. for a period of thirty hours." He believes this horsefly
to be the principal carrier of surra and that the stable fly, Stomoxys cal-
citrans (Linn.), is ruled out, which is indeed indicated by the long and
careful series of experiments conducted by that worker on both species
of flies.
Basu 15 (1945) reports that the occurrence of surra in India is sea-
sonal: the peak period for bovine surra is in August; for equine surra
it is in October; and for surra of camels and others it is in September. The
aggregate incidence reaches its peak in September. He also states that
the seasonal curve of surra and human malaria in the Punjab coincide.
Tularemia. In 1919 a disease of man of hitherto unknown etiology
occurring in the state of Utah (U.S.A.) was reported by Francis16 as
deer-fly fever or Pahvant Valley plague. It was later described by the
same author17 and given the name tularemia. It is a specific infectious
disease traceable· to Pasteurella (= Bacterium) tularensis (McCoy and
Chapin). It was described by Francis as a disease of rural populations
occurring during the summer months, coinciding with the prevalence of
the newly discovered vector, Chrysops discalis Will., a deer fly (Taban-
idae). Francis states, "Following the fly bite on some exposed surface of
the body (neck, face, hands or legs) the onset is sudden, with pains
and fever; the patient is prostrated and is confined to bed; the lymph
glands which drain the bitten area become tender, inflamed and swol-
len, and commonly suppurate, requiring incision. The fever is of a septic
type, lasting from three to six weeks, and convalescence is slow." The
disease occurs not only in the United States but also in Canada, Alaska,
northern Europe, Russia, Japan, and many other parts of the world. The
pathology of tularemia is described in great detail by Lillie and Francis."
In the acute form of the disease a primary ulcer (eschar) develops at the
site of the inoculation. Francis and his co-workers found that rabbits
constitute an important reservoir for the infection and that it is trans-
mitted from rabbit to rabbit by the Chrysops fly. The fly is undoubtedly
merely a mechanical vector as indicated by the experiments of Francis
and Mayne, et al. (lac. cit.). It has also been found that tularemia can
be transmitted from rabbit to rabbit by means of the rabbit louse,
H aemodipsus ventricosus (Denny). Cimex lectularius Linn., the com-
mon bedbug, was also found to be a successful vector in laboratory ex-
. periments with guinea pigs, as was the mouse louse, Polyplax serratus
( Burm. ), in the case of white mice. Mosquitoes and fleas have been
shown to be able to transmit the infection. Several species of ticks are
involved. Dermacentor andersoni Stiles, is an important vector, perhaps
the most important one because the infection is transmitted transovar-
ianly from generation to generation. Tularemia is now known to exist
in nature in many species of animals; among these are meadow mice,
ground squirrels, beavers, coyotes, sheep, and quail and other game
birds. Infection may be acquired not only by insect bites but also by
contact with infected insect feces, infected raw meat, and in rare in-
stances through contaminated water. Over 2,000 cases of tularemia were
reported in the United States in 1938, with a mortality rate of about 5
per cent.
Loiasis. The so-called mango fly, Chrysops dimidiata v. d. Wulp,
has been shown by Leiper (lac cit. p. 6), to be a vector of Loa loa
(Cobbold), the African eye worm of man in various endemic regions
in Africa, particularly the Belgian Congo. Chrysops silacea Austen has
been proved to be a carrier of the organism by Connall and Connall,19
who completely elucidated the life cycle not only in this fly, but also in
C. dimidiata v. d. Wulp. Microfilariae of Loa loa are found in the periph-
eral blood vessels during the daytime, showing a diurnal periodicity
which gave rise to the term Microfilaria diurna Manson. The larvae
measure about 300~ in length by 7.5fJ, in thickness, resembling
Wuchereria bancrofti (Cobbold) quite closely. In this stage they are
ingested by the Chrysops flies and undergo development similar to
that' of Wuchereria bancrofti in the mosquito. Metamorphosis .is com-
pleted in from 10 to 12 days, the larva increasing in length "tenfold."
When the infected fly bites, the mature larvae issue from the proboscis,
come to lie upon the skin of the host, and quickly disappear by bur-
rowing.
The adult worms, females measuring from 50 to 70 mm in length
and the males about half this length, inhabit the superficial subcutaneous
connective tissue and are known to move about from place to place
quite rapidly, giving rise to transient itching swellings known as Cala-
bar swellings. The parasites have been observed in many parts of the
body, such as the scrotum, penis, breast, eyelid, anterior chamber of
the eye, tongue, finger, and back. The worms may be most readily ex-
cised when they travel across the bridge of the nose or conjunctiva.
EI debab. EI debab is a trypanosomiasis of Algerian horses and camels
traceable to Trypanosoma berberum Edmond and Et. Sergent. This
disease is evidently spread by horseflies, Tabanus nemoralis Meig. and
T. tomentosus Macq. being considered the vectors.
Control. Comparatively little of a specific preventive nature has
been done, except for the notable work of Porchinski, reported by
Howard. 20 Porchinski observed that tabanids collect in great numbers
in the neighborhood of damp places and lower themselves to the surface
of pools to drink, actually touching the water with their bodies. It oc-
curred to him that a covering of kerosene on the water would endanger
the lives of the insects as they came in contact with the surface. Hence
a quantity of kerosene was applied to a given pool, with most gratifying
results. By the third day of the experiment, the "pool of death" was
covered with "floating islands" of dead tabanids. Porchinski recommends
that a favorite pool be selected, and that the oil be poured on so that a
thick uniform layer of oil is formed covering the entire pool. Such "pools
of death" apparently attract the tabanids from over a considerable
adjacent area. The oil must of course be applied as early as possible
during the season when the adult flies appear and begin to mate and
deposit eggs.
The author21 has called attention to the breeding of tabanid flies in
rice fields, particularly in roadside pools, the result of rice-field drainage.
Correction of drainage defects is an important procedure. Drainage of
breeding areas is good practice where this can be done without inter-
ference with agriculture. In the California rice fields the author found
characteristic egg masses of Tabanus punctifer O.S. attached to the
stems and blades of rice plants. These eggs were commonly heavily
parasitized by the hymenopterous egg parasite Telenomus (= Phanurus)
emersoni (Girault). This parasite has been shown by Parman 22 to be
a potent factor in the control of tabanids near Uvalde, Texas, where
artificial dissemination of the parasite was practiced.
The species of Tabanidae. There are about 2,500 species of Tabanidae
included in about SO genera, with over 1,200 described species in the
genus Tabanus alone. They are world-wide in distribution. Although
the author has examined much of the literature concerning the Taban-
idae, only a few species are briefly referred to in this work (see below);
Students concerned with the Tabanidae of given localities should con-
sult the work of specialists in that area, e.g., Fairchild's excellent articles
on the Panamanian species.
1. Tabanus atratus Fabricius, the black horsefly (Fig. 94), measures
from 16 to 28 mm in length. It is distributed over most of the United
States east of the Rocky Mountains and into Mexico. The whole insect
is uniformly black and the thorax and abdomen in well-preserved speci-
mens are thinly covered with a whitish "dust" which is easily rubbed
off if the specimens are not cared for properly.
2. Tabanus stygius Say is the black and white horsefly and is a
widely distributed species east of the Rocky Mountains. Its length is
20 to 22 mm. The third segment of the antennae is red at the base,
blackish at the apex; the first and second segments and palpi are dark;
the legs are black, often the tibia are reddish at the base; the wings
are yellowish brown with the posterior border approaching hyaline, a
brown spot on the bifurcation of the third vein, and the transverse vein
closing the discal cell margined with brown; the abdomen is a uniform
black; in the female the thorax dorsally is plainly whitish pollinose with
more intense longitudinal lines; the thorax of the male is dorsally a uni-
form grayish brown.
S. Tabanus punctifer O.S. (Fig. 96) is also a black and white horse-
fly resembling T. stygius Say except that the front tibiae are white on
the basal third and the thorax is uniformly white in both sexes; there is
usually a small dark spot near the tip of the wing. It is the largest and
best known species of horsefly in western North America, particularly
along the Pacific coast.
4. Tabanus vicarius Walker (= Tabanus costalis Wied.), the green-
head, is one of the most dreaded stock pests common throughout the
southern United States.
Length 12-14 mm. Palpi yellowish, antennae brownish with the annulate
portion darker; thorax including the scutellum uniformly grayish yellow
pollinose; legs largely black, base of front tibiae and the middle and hind
tibiae except at apex yellowish; wings hyaline with the costal cells yellowish,
veins yellowish; abdomen above alternately striped with black and grayish
yellow. In the female the frontal callosity black above, with a very much
narrowed prolongation, the part of which adjacent to the callosity is some-
times obliterated, leaving the upper part as a separate spot. The male is
much like the female and easily associated with it, but there is a tendency
toward obliteration of the distinct markings of the abdomen, the black of the
female is replaced by brownish and the stripes ma.y blend so that the whole
base of the abdomen is practically one color.
(Hine, loc. cit.)
5. Tabanus lineola Fabr., the lined horsefly, is also an important
stock pest widely distributed in eastern, centtal, and southern North
America.
Length 12-15 mm. Palpi white; antennae reddish, annulate portion of

third segment darker; thorax brown and gray striped, the latter color not
prominent; wings hyaline; legs reddish, apex of the front tibia plainly, apexes
of middle and hind tibiae faintly, and all of the tarsi dark brown; abdomen
aoove orown or Mack with three prominent, gra.y stripes. The males and
females of this species are easily associated. In the latter sex there is some-
times a confusion of colors; the dark is replaced by reddish but the gray mid-
dorsal stripe is always prominent in all well-preserved specimens.
(Hine)
6. Tabanus sulcifrons Macq. is known as th~) autumn horsefly.

Length 18-21 mm. Palpi brownish, antennae Tlearly black with the third
segment brownish at base; legs dark, bases of tihiae darker; wings with a
distinct brownish tinge, cross veins at the end of th«'l discal cell and bifurcation
of the third vein margined with.brown. Female front with parallel sides, frontal
callosity shining brown, not quite as wide at the front, nearly square and
with a linear prolongation above. Segments of the abdomen above with promi-
nent gray, hind margins which expand into large gray triangles in the middle;
usually a black mark on the anterior part of each of the second and third
segments at the apex of the gray triangle. In the Ihales the division between
the large and small facets of the eye prominent; head slightly more convex
than in the female but nearly of the same size, coloration of the whole body
the same as in the female.
(Hine)
7. Tabanus striatus Fabr. is said to be the most prevalent horsefly

of the Philippine Islands, and is known to be an important carrier of
surra. The following description is after Mayne (loc. cit.).
The male is very distinct from the female, b()ing smaller and having a
larger head and different color markings. Size: 11 to 15 millimeters. Wing
expanse: 25 to 28 millimeters. The distinctly clavate palpi are shorter than
in the female, only two-thirds as long as the labium; they are dirty white
and fringed with moderately long black hairs. The abdominal color markings
take the form of a T of pale cadmium yellow in a field of burnt sienna,
bordered with pale clay yellow. The area of the large facets of the eye is
colored Roman sepia surrounded by an elliptical band of ultra ash gray. The
field of small facets has a mauve fringe bounding an area of iridescent mauve
and Prussian green.
Female: The front is narrow, converges slightly anteriorly; the color is
golden, marked with a black callosity of irregular form. Size: 15 to 17 milli-
meters. Wing expanse: 26.5 to 29 millimeters. The head is considerably
smaller than that of the male; eyes iridescent mauve and Prussian green.
The palpi are prominently conical, as long as, or slightly longer than, the
labium; the color is the same as in the male, mottled with short black hairs.
The abdomen is alternately striped with Cologne earth and pale clay yellow.
The median stripe is pale clay yellow. In both sexes the thorax is indistinctly
striped with pale clay yellow and pale brown, and the wings are transparent
except the costal and subcostal cells, which are pale brown.
8. Chrysops callida O. S. is a widely distributed species, measuring

from 7 to 9 mm, and is black in color with large pale yellow spots on
the sides near the base of the abdomen.
9. Chrysops celer O. S. is black in color, the female with dense orange
pile on the pleurae. It measures from 8 to 11 mm in length. It appears
to have a more northerly distribution.
10. Chrysops discalis Williston is gray to yellow-gray in the female,
with black spots on the abdomen; in the wing picture the hyaline discal
cell and spot at the bifurcation of vein R. +5 are quite characteristic. In
the male the color is predominately black with yellow-gray spots on the
abdomen. Length 8 to 10.5 mm. It is reported from Utah, Nevada,
California, Oregon, Washington, Nebraska, North Dakota, Wyoming,
Montana, Colorado, also Manitoba and Saskatchewan.
11. Chrysops dimidiata v. d. Wulp is a southwest African species
measuring 8.5 mm in length. The face is dusty brownish yellow, the
thorax is piceous, and the abdomen is reddish ochraceous with a fulvous
pilosity.
12. Chrysozona americana Osten Sacken occurs from central Alaska to
New Mexico (type locality, Hudson Bay Territory) sharing generic
characters:
Small, slender, greyish species; eye somewhat pilose; frons very broad and
basal callus transverse; first antennal segment one and one-half to three times
as long as wide and usually swollen; third with four divisions, the first divi-
sion with no distinct angle above; wing brown or gray, with hyaline macula-
tions . . . knob of haltere white or pale yellow.
(Stone)
Key to the Tabanid Genera of Nearctic America"

(Arranged by T. H. C. Aitken after Brennan" and Stone.' Refer also to Surcouf.")
1. Hind tibiae with apical spurs; ocelli usually present.
Subfamily Pangoniinae (2)
Hind tibiae without apical spurs; ocelli usually absent, if present
rudimentary ............. . ............ . Subfamily Tabaninae (9)
2. Flagellum of antenna composed of eight annuli (3)
Flagellum with five distinct annuli .. (7)
3. Second anal vein sinuous .. . Bequaertomyia Brennan 1935
Second anal vein not sinuous.. (4)
4. Eyes of female acutely angulate above; wings darkened anter-
iorly .... . Goniops Aldrich 1892
Eyes of female rounded (normal); wings of uniform color.. (5)
5. Palpi short, stubby, about equal in length to proboscis, which is
conspicuously shorter than head . Apatolestes Williston 1885
Palpi slender, distinctly shorter than proboscis, which is often as
long as or longer than head.. (6)
6. Cell R, petiolate .. Esenbeckia Rondani 1864
Cell R, open. . ..... Stonemyia Brennan 1935
7. (2) Pedicel of antenna about half as long as scape .
Silvius Meigen 1820
Pedicel of antenna more than half as long as scape, often nearly
as long (8)
8. Wings evenly infuscated; abdomen globose, much wider than thorax;
antennae very slender and elongate; stump at bifurcation of vein
R.+, .... ....... ..Neochrysops Walton 1918
Wings irregularly infuscated, exhibiting a variety of patterns (en-
tirely hyaline in C. hyalina Shannon); abdomen normal; antennae
variable; bifurcation of vein R. +' without stump (rarely appear-
ing adventitiously) ..Chrysops Meigen 1803
9. (1) Scape of antenna considerably longer than thick; frons of
female widened below, broader than high, with a velvety-black
spot to each side at angle made by eye and subcallus; t flagellum
of antenna with four annuli; wing gray, with small white macula-
tions. Chrysozona Meigen 1800
Scape of antenna usually scarcely longer than thick; frons of female
not broader than high, without velvety-black spots; flagellum of
antenna usually with five annuli; wing pattern, if any, otherwise.. (10)
.. Philip" has omitted the genus Bequaertomyia from his recent catalog of Nearctic
Tabanidae, based on the elevation by Mackerras and Fuller (1942) of this primi-
tive group of flies to family rank (Pelecorhynchidae) in which Bequaertomyia
appears.
t That part of the frons below the level of the lower, inner angle of the eye and
above the antennae is termed the subcallus; the frons proper (just dorsad) usually
possesses two denuded calli, one at the lower margin, the basal callus, and the
median callus, usually narrow and frequently broadly jOined to the basal callus.
10. Eye bare; subcallus very swollen and shiny; genae denuded; dorsal
angle of flagellum small and blunt; wing at least partially infus-
cated ...................................... (11)
Without above combination of characters; if the subcallus is en-
larged and denuded, the eye is densely pilose ( 12)
11. Scape of antenna swollen, at least below; apical half of vein R.
turned abruptly forward; wing, at least anteriorly, infuscated with
a cresent-shaped, hyaline apex; tibiae somewhat swollen ...
Bolbodimyia Bigot 1892
Scape of antenna not noticeably swollen; vein R. not turned abruptly
forward; apex of wing not hyaline; tibiae not swollen.
Whitneyomyia Bequaert 1933
12. (10) Flagellum of antenna with no dorsal angle; frons of female
very narrow, the median callus a very slender line; no ocellar tuber-
cle; wing with at least a subapical brown spot; eye bare
Diachlorus Osten Sacken 1876
Not with this combination of characters· (13)
13. Basal portion of antennal flagellum with a prominent, forward-
projecting tooth reaching nearly to base of annulate portion;
eye pilose Dicladocera Lutz 1909
Basal portion of antennal flagellum with o. without a prominent
dorsal angle, but if this is produced forward the eye is bare (14)
14. Basal callus in female lacking or very much reduced, separated
from eye by a considerable space; neither palpus black nor abdomen
with a narrow dorsal stripe (15)
Basal callus in female as wide, or nearly as wide, as frons, or, if
narrowed, still considerably wider than median callus; either palpus
black or abdomen with a narrow dorsal stripe (17)
15. Eye distinctly pilose; no distinct ocellar tubercle (eye of female usu-
ally with a single, diagonal, purple line which often shows even in
dried specimens). . Atylottls Osten Sacken 1876
Eye bare or very sparsely pilose; ocellar tubercle present or absent
(frons of female about five times as high as width at base) . (16)
16. Basal callus a swelling at base of a slender raised ridge; a distinct
ocellar tubercle present in female; abdomen brownish, with white
bands, the apex compressed Leflcotabanlls Lutz 1913
No basal or median calli or ocellar tubercle present; bright green or
yellow species, the abdomen not distinctly compressed apically
.. .Chlorotabanus Lutz 1909
17. (14) Annulate portion of antennal flagellum hairy; no ocellar tuber-
cle; second palpal segment short and stout, with erect hair; pro-
boscis short. .Anacimas Enderlein 1923
Not agreeing entirely with above, the hair of antennal flagellum
very inconspicuous if present (18)
18. Rather small, species with bare or sparsely pilose eye, scarcely
any angle, and no dorsal excision on flagellum of antenna, and fre-
quently· a stump vein from vein R. . ...... Stenotabanus Lutz 1913
Eye bare or pilose, but if a stump vein from vein R. is present
either the dorsal angle of the antennal flagellum is distinct or the
eye is densely pilose, or both ................... .Tabanus Linne 1758
B. SNIPE FLIES
Family Rhagionidae (Leptidae)
Snipe flies belong to the dipterous family Rhagionidae formerly
known as Leptidae. The family comprises nonbloodsuckers as well as
bloodsucking species. Leonard 26 characterizes the family as follows:
Flies of moderate to large size usually more or less elongate and nearly
bare to moderately pilose, rarely rather densely hairy, never, however, with
distinct bristles. Males usually hoI optic; more rarely dichoptic. Empodium
pulvilliform, there being three pads of about equal size between the tarsal
claws. (Hilarimorpha has no visible empodia and no discal cell but is usu-
ally referred to this family.) Squamae small or vestigial. Antennae extremely
variable: (a) segments of flagellum distinct, sometimes as many as thirty
in number; (b) the segments not more than eight in number; more closely·
applied, without style or arista; (c) fewer in number with a differentiated
segmented style or arista, altogether not more than eight; (d) the third seg-
ment simple with or without a dorsal or terminal arista. Veins of the wings
distinct, not crowded anteriorly; third longitudinal cell furcate; basal cells
large; five posterior cells usually present.
One of the members of the family is the "worm lion," Vermileo; the
larvae excavate pits in sand as do "ant lions" (Myrmeleon). Among the
bloodsucking genera are Atherix, e.g., A. longipes Bell, a severe biter
in Mexico; and Symphoromyia.
The genus Symphoromyia 27 "includes leptid flies with five posterior
cells, the anal cell open; third antennal joint simple, rather deep vetti-
cally, attached above its middle, usually kidney-shaped (sometimes con-
cave in profile below the arista, then not quite kidney-shaped); arista
subapical; tibial spurs none in front, two in the middle, one behind, but
often quite weak in males." The females of several species are vicious
biters, behaving somewhat as do the tabanid flies belonging to the genus
Chrysops. They alight on the exposed parts of the body quite silently
and singly and inflict a sudden painful bite usually before their presence
is known. Among the severe biters are Symphoromyia atripes Bigot, a
western species measuring 5.3 to 8 mm in length, black, with reddish
legs; S. pachyceras Williston, particularly a Pacific coast species, measur-
ing 6 to 9 mm in length, wholly black except narrowly on the knees, the
inner proboscis, and stems of the halteres which are yellow; S. kincaidi
Aldrich, pile of the thorax and head black, of abdomen largely yellow,
front and middle knees narrowly red, a Pacific coast species. S. hirta
Johns. is shown in Figure 100.
The mouth parts of Symphoromyia evidently vary considerably. The
biting forms have a prominent stout retractile labial sheath which closely
ensheaths the functional chitinized piercing structures.
FIG. 100. A snipe fly, Symphoromyia hil'ta.

(Adapted after Rearle.)
Practically nothing is known about the breeding habits and life his-
tory of the species of Symphoromyia. The rhagionids as a group are
known to breed in moist soil, where there is decaying vegetation; the
larvae are predaceous.
BIBLIOGRAPHY
1. Osten Sacken, C. R., 1875. Prodrome of a Monograph of the Tabanidae
of the United States. Part I; Part II, 1876; Supplement, 1878, in Mem. Boston
Soc. Nat. Hist., voL 2, part iv, nos. 1, 4, 6.
2. Hine, J. S., 1904. "The Tabanidae of western United States and
Canada," Ohio Naturalist, 5:217-48.
3. Enderlein, G., 1925. "Studien an blutsaugenden Insekten. I. Grundla-
gen eines neuen Systems der Tabaniden," Mitt. a. d. Zool. Mus. Berlin, 2:253-
409 (5 text figures).
4. Krober, 0., 1927. "Die Chrysopsarten Afrikas," Zoologische lahrbil-
cher. Abt. f. Syst., akol., u. Geogr. der Tiere., 53:175-268 (5 plates).
5. Stone, Alan, 1938. The Horseflies of the Subfamily Tabaninae of the
Nearctic Region. Washington, D. C.: Dept. Agric., in Misc. Pub!., no. 305.
171 pp.
6. Marchand, Werner, 1920. The Early Stages of Tabanidae (Horse-
flies). New York: Rockefeller Institute for Medical Research, in Monograph
no. 13. 203 pp. + 15 plates.
7. Stone, Alan, 1930. "The bionomics of some Tabanidae (Diptera) ,"
Ann. Entomolog. Soc. Amer., 23:261-304.
8. Schwardt, H. A., 1936. Horseflies of Arkansas. Fayetteville: Univ.
Arkansas, in Agric. Exper. Sta. Bull., no. 332. 66 pp.
HORSEFLIES, DEER FLIES, AND SNIPE FLIES 31.5
9. Webb, J. L., and Wells, R. W., 1924. Horseflies: Biologies and rela-
tion to western agriculture. Washington, D. C.: Dept. Agric., in Bull., no.
1218. 36 pp.
10. Philip, C. B., 1928. "Methods of collecting and rearing the immature
stages of Tabanidae," ]. Parasitol., 14:243-53.
11. Garman, H., 1910. An Outbreak of Gadflies in Kentucky. Kentucky
Agric. Exper. Sta., in Bull., no. 151.
12. Mitzmain, M. B., 1914. "II. Summary of experiments in the trans-
mission of anthrax by biting flies." Washington D. C.: Govt. Print. Office, in
Public Health Bull., no. 94. pp. 41-48.
13. Yutuc, Lope M., 1938. "A report on the first outbreak of surra in
British North Borneo and its control measure," Philippine J. Animal Indust.,
5:501-15 (4 plates).
14. Mitzmain, M. B., 1913. "The mechanical transmission of surra by
Tabanus striatus Fabricius," Philippine f. Sc., 8:223-29.
15. Basu, B. C., 1945. "Distribution and seasonal incidence of surra in
India," Indian J. Vet. Sc., 15:277-79.
16. Francis, Edward, 1919. "Deer fly fever or Pahvant Valley plague,"
U. S. Public Health Service, Pub. Health Rep. 34:2061-62.
17. ---; Mayne, Bruce; and Lake, G. C.; 1921. Tularaemia Francis
1921: A New Disease of Man. Washington, D. C.: U. S. Public Health Service,
in Hygienic Laboratory Bull., no. 130. 87 pp.
18. Lillie, R. D., and Francis, Edward, 1936. Pathologoy of Tularaemia.
Washington, D. C.: U. S. Public Health Service, in Nat. lnst. Health Bull.,
no. 167. 217 pp.
19. Connall, A., and Connall, S. L. M., 1922. "The development of Loa
loa (Guyot) in Chrysops silacea (Austen) and in Chrysops dimidiata (van
der Wulp)," Tr. Roy. Soc. Trap. Med. & Hyg., 16:64-89.
20. Howard, L. 0., 1899. A Remedy for Gadflies. Porchinski's Recent
Discovery in Russia, with Some American Observations. Washington, D. C.:
Dept. Agric., in Div. of Entomol. Bull., no. 20, n. s.
21. Herms, W. B., 1927. "Tabanids breeding in rice fields." Pan-Pacific
Entomologist, 4:91-92.
22. Parman, D. C., 1928. Experimental Dissemination of the Tabanid
Egg Parasite, Phanurus emersoni Girault, and Biological Notes on the Species.
Washington, D. C.: Dept. Agric. In Circ. no. 18. 6 pp.
23. Brennan, J. M., 1935. "The Pangoniinae of Nearctic America. Diptera:
Tabanidae," Univ. Kansas Sc. Bull., 22:249-402.
24. Surcouf, J., 1921. Diptera, family Tabanidae. Brussels: Genera In-
sectorum de P. Wytsman, Fasc. 175.
25. Philip, Cornelius B., 1947. "A catalog of the bloodsucking fly family
Tabanidae (horseflies and deerflies) of the Nearctic region north of Mexico,"
Amer. Midland Naturalist, 37:257-324.
26. Leonard, M. D., 1930. A Revision of the Dipterous Family Rhagionidae
(Leptidae) in the United States and Canada, in Mem. Amer. Entomolog. Soc.,
No.7. 181 pp. (3 plates).
27. Aldrich, J. M., 1915. "The dipterous genus Symphoromyia in North
America," Proc. U. S. Nat. Mus., 49:113-42.
CHAPTER XV
._.
HOUSEFLIES
ORDER DIPTERA, SUPERFAMILY MUSCOIDEA

House-invading flies. Many species of robust flies belonging to various
families of Diptera are commonly found indoors; however, relatively few
of these species are of public health importance. Flies which habitually
enter the house, coming in contact with household food or drink, and
breeding normally in excrement and dead animal matter or feeding on
these, are a potential menace to the public health. Flies of this charac-
ter usually belong to the Muscidae, Anthomyidae, Sarcophagidae (flesh
flies), and Calliphoridae (blowflies). Curran (1934, loco cit. p. 143) has
combined the Anthomyidae with the Muscidae and places the flesh flies
(Sarcophagidae) and blowflies ( Calliphoridae), in the family Me-
topiidae, which he designates as the flesh flies. The blowflies include
the bluebottle and the green bottle flies, which deposit their eggs on dead
animals or garbage, also upon cold meat and other foods of man, which
when ingested may cause intestinal disturbances. The flesh flies deposit
living young (larviposit).
Family Muscidae. The family Muscidae, to which the true housefly
and several other house-invading species belong, is characterized by
Curran (1934, loco cit.) as including "flies of medium to small size,
usually dull colored, with the squamae large or of medium size, the
hypopleural bristles absent; the second antennal segment grooved
above. Arista plumose, pubescent, bare or pectinate, eyes approximate
or widely separated in the males, the front rarely narrowed in both
sexes; frontal bristles always present, intrafrontals frequently present;
orbitals developed but rarely in the males. Abdomen composed of four
segments in the male, five in the female. Male genitalia usually not
prominent but sometimes conspicuous; fifth sternal lobes sometimes
prominent."
The true housefly. Hewitt'sl description (translation from Schiner)
of Musca domestica Linn. (Fig. 101), the common housefly, is undoubt-
edly the best for our purpose:
Frons of male occupying a fourth part of the breadth of the head. Frontal
stripe of female narrow in front, so broad behind that it entirely fills up the
, 316
HOUSEFLIES 317
width of the frons. The dorsal region of the thorax dusty gray in color with
four equally broad longitudinal stripes. Scutellum gray with black sides. The
light regions of the abdomen yellowish, transparent, the darkest parts at least
at the base of the ventral side yellow. The last segment and a dorsal line black-
ish brown. Seen from behind and against the light the whole abdomen shim-
mering yellow, and only on each side of the dorsal line on each segment a dull
transverse band. The lower part of the face silky yellow, shot with blackish
brown. Median stripe velvety black. Antennae brown. Palpi black. Legs black-
ish brown. Wings tinged with pale gray with yellowish base. The female has a
broad velvety black, often reddishly shimmering, frontal stripe, which is not
broader at the anterior end than the bases of the anteimae, but becomes so
very much broader above that the light dustiness of the sides is entirely ob-
literated, the abdomen gradually becoming darker. The shimmering areas of
the separate segments generally brownish. All the other parts are the same
as in the male. Mature insect 6-7 mm. in length, 13-15 mrri. across the wings.
I
FIG. 101. Developmental stages of the common housefly, Musca domestica:
egg, larva, pupa, and adult female.
Why called the housefly. Of a total of 23,087 flies collected by

Howard 2 in dining rooms in different parts of the United States, 22,808
(98 per cent of the whole number) were Musca domestica Linn. Again,
of a total of 294 flies collected by the writer, representing the entire fly
population of one house, 202 (94.4 per cent) were Musca domestica
Linn. Thus the term common housefly is not misapplied. It is further-
more a cosmopolitan species.
Distribution of sexes. In order to determine the distribution of the
sexes of Musca domestica Linn. observations were made under two
different conditions: first, six sweepings with an insect net were made
over a pile of horse manure on which many flies had gathered (the re-
suits are shown in Table II); second, all but half a dozen flies were col-
lected in one house, giving a fairly representative lot for indoors, even
under screened conditions (Table III).
TABLE II
SHOWING RESULTS WITH REGARD TO SEX AND SPECIES IN SIX SWEEPINGS FROM
A HORSE-MANURE PILE ON MAY 19, 1909
_________ I_:r~1 ~cond

I I '
1_ Thir~ _Four~ Fifth _ Sixth !_Total
0' \l 10' <;>1[0' <;> 0' <;> 0' <;> 0' <;> 0' <;>
Housefly (Musca domestica) 71534 813 649 7742105 11232697
Muscina sp. 2 6 0 7 0 5 2 5 3 10 1 4 8 37
Blowfly (Calliphora sp.) 2 2 0 111 0 0 0 0 0 1 0 4 3
Lucilia sp. o 110 110 1 0 1 0 0 0 0 0 4
Other species 1 4 0 4 2 1 4 2 4 2[ 2 013 13
Totals 112
---1----'---'------'1------
166 4 9416 71115 85,11 222[ 9 11657 754
1
TABLE III
SHOWING NUMBER OF INDIVIDUALS COLLECTED IN A SCREENED DWELLING
JUNE 1, 1909, REPRESENTING THE ENTIRE FLY POPULATlON OF THE HOUSE
0' <;>
Housefly (Musca domcstica) 86 ll6
Muscina sp. 3 1
Homalomyia sp. 5 0
Calliphora sp. 1 2
Totals 95 ll9
These two tables give us some information as to the relative abun-

dance of the housefly, and the distribution of the sexes. Table III shows
clearly that of those flies which frequent both the manure pile and the
home, the common housefly (Musca domestica Linn.) composes 90 per
cent, and that of the total collected, over 95 per cent (95.4 per cent)
were females. Thus, it is clear that it is the "instinct" to oviposit (to
lay eggs) that mainly attracted these insects to the manure. In fact,
fresher parts of the manure pile are often literally white with housefly
eggs in countless numbers. Observations made in the near vicinity of the
manure piles proved that certainly the same percentage (over 95 per
cent) of the flies clinging to the walls of the stable, boxes, and so on
were males.
That males and females are normally about equal in number is
evidenced by the fact that of a total of 264 pupae collected indiscrimi-
nately and allowed to emerge in the laboratory, 129 were males and 135
were females.
Of the total number of houseflies (202) collected indoors (June,
1909), representing all but perhaps six of the total number in that partic-
ular house, 57 per cent were females, showing nearly equal distribution
HOUSEFLIES 319
for the sexes. This would, it seems, indicate that males and females are
equally attracted to the house by odors issuing therefrom.
Life history of the housefly. The housefly passes through a complex
metamorphosis (Fig. 101) i.e., egg, larva (maggot), pupa, and adult
or fully winged insect. Under warm summer temperatures the egg
stage requires about 20 hours, the larval stage about 5 days, the
pupa about 4 days, a total of about 10 days from egg to adult insect.
This allows for the development of from 10 to 12 generations in one
summer.
From 75 to 150 eggs are deposited singly, piling up in masses, and
there are usually several such layings at intervals of three or four days.
Female flies begin depositing eggs from 9 to 12 days after emerging
from the pupa case. Dunn,3 entomologist, Board of Health Laboratory,
Ancon, Canal Zone, reporting on his observations, states that as many
as 159 eggs may be deposited in one batch, that large batches are some-
times deposited at intervals of but 36 hours, and that one female may
deposit as many as 21 batches, or a total of 2,387 eggs, in 31 days after
emergence. He also states that oviposition may take place as early as 2~
days after emergence, and that copulation may occur within 24 hours
after emergence and one successful copulation seems to be sufficient to
fertilize the female for her lifetime. Under our laboratory conditions
houseflies reach sexual maturity in three or four days and begin deposit-
ing eggs on the ninth day after emergence from the puparium. Sun-
shine stimulates their breeding habits. Egg laying may continue through-
out the lifetime of a fly, i.e., for more than two months.
Influence of temperature on life history. While conducting an exten-
sive series of experiments in which many hundreds of houseflies were
used in all stages, a record was made of the temperature at which the
containers were kept. Ordinarily not more than one to three quarts of
manure were used for the growing maggots, hence the temperature of
the environment did not differ widely from that of the manure. The
temperature of an average manure pile to which material is added daily
varies from 18° C to 66° C. Young growing larvae are most numerous
at temperatures varying from 45° to 55°. Below 45° half-grown and
full-grown larvae occur; above 55° the temperature seems to become
too great.
From the following table (Table IV) it will be seen that tempera-
ture influences very materially the time required for the development
from egg to imago, but nevertheless with an average outdoor tempera-
ture of 18° C flies ordinarily require only from 12 to 14 days to pass
through the same stages; this is, of course, due to the higher tempera-
tur~ of the manure pile, as indicated above. The shortest time required
for complete metamorphosis of Musca domestica Linn. is seen to be
9% days, and that at 30° C.
TABLE IV
SHOWING INFLUENCE OF TEMPERATURE ON THE LENGTH OF LIFE HISTORY
OF MUSCA DOMESTICA
The insects were kept at the temperature indicated from egg to emergence of
the imago. The average temperature is here given, the variation from the
average was probably not more than ± 1 Temperature of the air and not of
0.
the manure is here considered.
16° C 18° C 20° C 25° C 30° C

-----
Min. Max.
~IMax. Min. Max. Min. Max. Min. Max.
---- f----
Egg stage 36 hrs. 40 hrs. 27 hrs. 30 hr,. 20 hrs. 30 hrs. 12 hrs. 20 hrs. 8 hrs. 12 hrs
Larval stage 11 ds. 26 ds. 10 ds. 14 ds. 8 ds. 10 ds. 7 ds. 8 ds. S ds. 6 ds.
Pupal stage 18 ds. 21 ds. 12 ds. 15 ds. 10 ds. 11 ds. 7 ds. 9 ds. 4 ds. 5 cis.
ds.I22~" ds. -
Total time required
from egg to imago 40 >~ d,. 48 % ds. 2H& ds. 30 % ds. 18% 14~~-ds.iI7%dS. 9~ ds. 11).~ ds.
Average time required

to develop from egg
to imago i 44.8 days 26.7 days 20.5 days 16.1 days 10.4 days
I
Excrement the preferred breeding place. Excrement, especially of
horses (Fig. 102), is the material upon which Musca domestica Linn.
prefers to deposit its eggs, and this is the material on which the larvae
feed. It seems quite safe to say that under rural conditions 95 per cent
of the houseflies are bred in horse manure. The eggs of the housefly
hatch in from 12 to 24 hours; the newly hatched larvae begin feeding at
once and grow rapidly.
To gain an estimate of the number of larvae developing in an aver-
age horse-manure pile, samples were taken after four days' exposure to
flies, with the following results: first sample (4 Ib) contained 6,873
larvae; second sample (4 Ib), 1,142; third sample (4 Ib), 1,585; fourth
sample (3 Ib), 682; total 10,282 larvae in 15 pounds. All of the larvae
were quite or nearly full grown. This gives an average of 685 larvae per
pound. The weight of the entire pile was estimated at not less than 1,000
pounds, of which certainly two-thirds was infested. A little arithmetic
gives us the astonishing estimate of 455,525 larvae (685 x 665), or in
round numbers 450,000. 4 •
The larval stage is the growing period of the fly, and the size of the
adult will depend entirely upon the growth that the larva attains. An
underfed larva will result in an undersized adult. The growing stage
requires from four to six days, after which the maggots usually crawl
away fro:rp. their breeding place, many of them burrowing into the loose
ground just beneath t~e manure pile, or under boards or stones, or into
HOUSEFLIES 321
dry manure collected under platforms and the like. One and three-
quarter pounds of dry manure, taken from beneath a platform, contained
2,561 pupae. The larvae spend three or four days in the prepupal or
migratory stage before actually pupating. In a given set of individuals
under similar conditions the various stages are remarkably similar in
duration: when one pupates, the rest will certainly follow in short order,
and when one emerges, the others quickly follow.
When the fly emerges from the pupa case the wings are folded in
tight pads, and change in size is due only to expansion and addition
in weight and not in growth. Stomach contents or development of eggs
in the female add to weight. This is why no young houseflies are seen,
FIG. 102. A typical rural fly-breeding place-the manure pile. The principal
menace is the fresh, warm manure added on top daily. The nearby vehicle is
a remnant of the "horse and buggy days." The manure pile is far from obsolete.
i.e., young in the sense of being small. Little flies are not "baby" flies;
they are either a different and smaller species or are undersized. One can
influence the size of the adult fly by underfeeding it in the larval stage
(see Herms, 5 1907).
With one adult fly depositing from 120 to 150 eggs per lot, with at
least six lots at intervals of from three to four days, Hodge 6 gives us the
following astounding statement: "A pair of flies beginning operations
in April may be progenitors, if all were to live, of 191,010,000,000,000,-
000,000 flies by August. Allowing on~-eighth of a cubic inch to a fly, this
number would cover the earth 47 feet deep."
Other breeding places. While horse manure is a favorite larval food
and is commonly regarded as the chief factor in the production of
houseflies under most rural and village conditions, situations may arise
in which other materials are vastly more important, particularly in this
day of the automobile and tractor. Cow manure if well mixed with
bedding is frequently an important factor in the development of flies.
Flies will also breed freely in hog manure, but the swarms of flies about
the pig pens usually originate in the waste feed, slops, etc. Chicken
manure is the most important factor in the breeding of flies in poultry
districts, and the pest of blowflies in such areas is the result of dead
birds buried in shallow pits or simply disposed of by throwing them
into a gully or in a corner. The dead birds (large or small) should be
burned or buried with crude oil or plenty of lime, or sprayed with
creosote oil. Human excrement is a very dangerous substance, and if
exposed to flies in open privies becomes a very prolific breeding place,
which emphasizes the need of flyproof privies or the application of other
means to prevent flies from breeding.
Great swarms of flies are often found around feed troughs; the ani-
mals (hogs and cattle) may be literally covered with them. An examina-
tion of the waste feed behind or beneath the troughs or in and about
the mixing vats will almost invariably reveal numerous maggots. Storage
receptacles for slops sometimes present a wriggling mass of maggots.
The correction of such fly breeding manifestly depends upon greater
care in handling the mash, wet or dry. Spraying fences, walls, and floors
with DDT is also good practice.
It frequently happens that brewer's grain or spent hops, bran mash,
and ensilage are only partly consumed by the animals and the waste is
thrown out into the fields in heaps. Such heaps of waste are commonly
a source of enormous numbers of flies (nearly all Musca domestica)
about dairies where otherwise conditions may be very good and where
no apparent reason for the swarms of flies exists. Such wastes should be
scattered in a thin layer so as to hasten drying and thus prevent fly
breeding.
Garbage heaps, particularly when fermentation and decomposition
begin, are commonly sources of many flies of several kinds. Heaps of
decaying onions and other vegetables, fruits, etc., as well as decaying
straw and weeds, may become infested with maggots, often the larvae
of the biting stable flies (Stomoxys calcitrans). Every household in every
community should be provided with a garbage can equipped with a
tight-fitting lid. All liquid matter should be drained from the kitchen
refuse at the sink; only solids should be placed in the garbage can, and
these should first be wrapped amply in paper. Household gar-
bage should not be buried or allowed to accumulate in heaps in
back yard or alley (Fig. 103). Rats as well as flies thrive in such an
environment.
HOUSEFLIES 323
In the country, in the absence of sewers or septic tanks, the dish-

water from the kitchen is frequently piped from the sink to a ditch in
the back yard. On many occasions these ditches become clogged and
vile smelling, and an examination will reveal numerous maggots develop-
ing in the muck-a source of flies which is commonly overlooked.
The writer has often seen septic tanks with open cracks and knot
holes which permit flies to enter and lay eggs. Maggots bave been found
in countless numbers in the soft sludge mat covering the liquid in the
defective tank. Crevices and knotholes are easily covered with tin, thus
FIG . 103. A poor excuse for a fly-tight garbage can. Care of garbage should
be regulated by ordinance.
preventing ingress and egress for flies and eliminating breeding within
the tanks.
Range of flight. Ordinarily under city conditions it may be safely
said that where flies are abundant they have been bred in the immediate
vicinity. The housefly can, however, use its wings effectively and may
also be carried by the wind, though it usually seeks protection very
quickly when there is a strong breeze. Where houses are situated close
together flies have the opportunity to travel considerable distances by
easy flights and they are often carried on meat delivery wagons and on
animals.
In an illuminating experiment Copeman et al! have shown that
houseflies may invade a community at a distance of from 300 yards to
17,000 yards from their breeding place, such as a garbage dump.
In a city in Montana 387,877 marked flies were liberated from a
release point, and a total of 1,056 of them were recaptured at 78 stations
situated between 50 and 3,500 yards from the point of release. 8
Longevity of flies. In order to determine the longevity of flies it is
necessary to keep the same individual under observation from the time
of emergence from the pupa to the time of death. The writer has done
this by keeping each pupa in a separate vial, noting the time of emer-
gence to the hour and spotting each fly lightly with Chinese white
dorsally on the thorax. The spots can be arranged singly and in com-
bination so that many different flies can be kept under observation at
the same time. After marking, flies were liberated in bobbinet-covered
cages (size of cages never more than 8 by 10 by 18 inches). Each cage
was provided with sugar water and a receptacle of horse manure. A
full set of experiments under sufficiently varying conditions indicate an
average life of close to 30 days, with a maximum life of something
over 60 days during the summer months. In hibernation flies may live
over winter, i.e., from October to April, in the eastern and central
United States. In California, flies emerge from their pupa cases through-
out the winter, and their life history is then considerably longer than
in summer.
Other house-invading flies. There are many species (more than 30)
belonging to the genus Musca, most of which have habits similar to
those of the cosmopolitan M. domestica, namely M. vicina Macq. the
common housefly of the Orient, also common in Hawaii; M. vetustissima
Walker, common in Australia; M. nebulo Wied., a tropical species; M.
sorbens Wied., reported to be abundant on the central and south Pacific
Islands.
Fannia ( H omalomyia ) canicularis (Linn.), known as the lesser
housefly, is frequently seen hovering in mid-air or flying hither and
thither in the middle of the room. Whereas the true housefly is en-
countered most abundantly in the kitchen or dining room, particularly
on food, the "little housefly" will be seen as frequently in one room as
another, and very seldom actually on the "spread" table. The writer
commonly observes a half dozen or more of these little flies dancing
weirdly in the center of the lecture room midway between the floor and
the ceiling. Various observers have estimated that this species consti-
tute from one to 25 per cent of the total population of flies in the average
house.
In size the species varies from 5 to 6 mm. Its color is grayish, re-
sembling that of the housefly very closely. Hewitt describes it as fol-
lows:
Head iridescent black, silvery white, especially around the eyes. The
antennae are blackish gray with non-setose arista. Palps black. The thorax
is blackish gray with three indistinct black longitudinal stripes; the scutellum
HOUSEFLIES 325
is gray and bears long setae; the sides of the thorax are lighter. . . . The
legs are black, and the middle femora bear comb-like setae below. The some-
what large squamae at the bases of the wings are white and the halteres are
yellow. . . . The head of the female is gray with a wide frons, black frontal
stripe and gray sides. The longitudinal stripes of the thorax are faint and the
abdomen, which is more pyriform than that of the male, has a slightly golden
attachment.
The eggs of this species are depOsited on decaying vegetable matter

and excrement, particularly of humans, horses, and cows. The larvae
emerge in about 24 hours and may be recognized as compressed, spiny
organisms about 6 mm long when full grown (Fig. 104). The pupal
period lasts about seven days under favorable conditions.
FIG. 104. (A) larva of Fannia

(= Homalomyia) canicularis, (B)
larva of Fannia (= Homalomyia)
scalaris. X 6. (Redrawn and adapted
after- Hewitt.)
A B
Fannia (= Homalomyia) scalaris (Fabr.), the latrine fly, is very

similar to the foregoing. In size the two flies are about the same, if any-
thing the latrine fly is somewhat the larger. The thorax and abdomen are
bluish black; the antennae, palpi, and legs are black. The abdomen
has a dark median stripe which, with segmentally arranged transverse
bands, produces a series of dorsal triangular markings. The middle tibia
is provided with a distinct tubercle.
The eggs of this fly are deposited on excrement of humans, horses,
cows, etc., also on decaying vegetable matter. The egg stage lasts about
24 hours, the larval stage about six days and over, and the pupal stage
about nine days.
While the larva of the "latrine fly" resembles the larva of the lesser
housefly in general, it is readily distinguished because its singie lateral
protuberances are distinctly feathered (Fig. 104).
Other species of Fannia with similar breeding habits, referred to as
"Old World" species but now widely distributed are F. manicata (Meig.)
and F. incisurata (Zett.). The genus belongs to the family Anthomyiidae.
Musca autumnalis DeGeer (formerly called Musca corvina Fabr.)
is described by Austen 9 whose valuable brochure on the "'housefly"
should be read by all sanitary officers interested in housefly problems.
He says:
. . . though agreeing approximately with the housefly in length, is a bulkier,

more compactly built and thick-set insect . . . often decidedly larger. In
the male the upper surface of the abdomen has a black base, from which there
is a backward prolongation in the shape of a longitudinal, median stripe, both
base and stripe being sharply defined, and presenting a well marked contrast
to the cinnamon-buff of the remaining ground-color; in the female the upper
surface of both thorax and abdomen is grey, with darker markings. In the case
of both sexes, however, the surest criteria for distinguishing Musca autumnalis
from M. domestica are those presented by the upper surface of the head.
Whereas in the male Musca autumnalis the eyes are so close together as to
be almost or actually in contact at one spot, in the male housefly the space be-
. tween the eyes is always much broader, and, as already indicated, may be
nearly equal to one-fourth of the total width of the head. As regards the oppo-
site sex, in the female of Musca autumnalis the black longitudinal area (frontal
stripe) in the center of the space between the eyes is approximately equal in
width to the grey border on each side, separating it from the corresponding
eye. In the female housefly, however, the frontal stripe is much broader, and
its width greatly exceeds that of the border, yellowish-golden in front and be-
low, blackish above, on each side of it. The resting position of the wings in
Musca autumnalis is the same as in M. domestica. In the autumn, in country
districts in the British Islands, Musca autumnalis frequently enters houses and
public buildings, sometimes in large numbers, and subsequently hibernates in
attics, roof-lofts, towers, in the folds of curtains in disused rooms, and in
similar retreats.
Major Austen reports that it breeds in cattle droppings scattered in the

field.
Muscina stabulans (Fallen) is larger and more robust than the true
housefly, varying in length from 7 to nearly 10 mm. Its general appear-
ance is dark gray. The head is whitish gray, the antennal arista bears
setae on both the upper and lower sides. The thorax is gray with four
longitudinal black lines: the abdomen is almost black in color, covered
with gray in places, giving it a blotched appearance. The legs are slender
and are reddish gold or cinnamon in color. The wings are folded like
those of Musca domestica Linn.; the "fourth longitudinal vein is not
elbowed and converges but slightly towards that of the vein before it"
(Austen). The eggs of this species are laid upon decaying organic matter
and excrement, inclusive of human feces and rotting cow dung, in which
the larvae develop. The complete life cycle is said to require from five
to six weeks.
HOUSEFLIES 327
Pollenia rudis (Fabr.), the cluster fly, may be distinguished from all
other houseflies in that the wings when not in use close over each other
at the tips scissors-like. According to Austen it measures normally about
one-third inch in length; it is thus as a rule a much larger insect than
the true housefly. It is more heavily built and slower in its movements.
Austen continues the description:
The upper surface of the dark greyish-olive middle region of the body
(thorax) is clothed with a thick coat of fine, silky, recumbent, yellowish or
golden-yellow hair, easily visible to the naked eye, and, though readily rubbed
off, still recognisable with the aid of a lens even in a much damaged specimen.
The iron grey upper surface of the posterior division of the body (abdomen) is
mottled with shimmering metallic patches of lighter grey. (In 1908) Dr. D.
Keilin, working in Paris, made the extraordinary discovery that the maggot of
the cluster-fly is an internal parasite of a small earthworm (Allolobophora
chlorotica, Sav.) [which like the fly itself, is exceedingly common and widely
distributed in Europe, North America, and elsewhere]. The popular name of
the insect (namely cluster fly) is due to the habit of this fly of clustering
together, sometimes in very large numbers like a swarm of bees, when hibernat-
ing in houses or other buildings.
Blowflies and flesh flies. The blowflies, comprising the bluebottles and
greenbottles, included in the family Calliphoridae 1o as well as the flesh
flies (family Sarcophagidae of most authors l l ), are placed in the family
Metopiidae by Curran who characterizes this family as follows:
Flies of medium to moderately small size, the abdomen usually dark

and tessellate or metallic green or blue. Front in both sexes broad, usually
somewhat. narrowed in the males, rarely very narrow; vibrissae present; anten-
nae long or short, the arista plumose, pubescent or bare. Abdomen composed
of four segments in the males, the fifth short in the females; abdominal bristles
usually strong, at least on the apical segments. Hypopleura with a row of
bristles; post-scutellum developed only in Mesembrinella. Apical cell usually
open, rarely closed and petiolate, usually ending far before the apex of the
wing. . . . The absence of the post-scutellum distinguishes the family from
the Tachinidae, while the presence of hypopleural bristles separates it from
the Muscidae.
The larvae of the blowflies and flesh flies usually feed on dead animals,
garbage, excrement, etc. They are principally scavenger in habit.
Calliphora. Among the several species of bluebottle flies (metallic
blue in color) two are quite common, namely, Calliphora vomitoria
(Linn.), having black genae with golden-red hairs, and C. vicina R. -D.
[ = C. erythrocephala ( Meig. ) ], with fulvous genae and black hairs
(Fig. 105). The eggs of these species hatch in from 6 to 48 hours; the
growing larvae feed on the flesh for from three to nine days, and after
328 ~IEDICAL ENTOMOLOGY
attaining full growth leave the food and bury themselves in loose earth
or debris. This period (prepupal period) lasts from two to seven days,
commonly four, after which pupation takes place. The pupal period
varies considerably according to temperature, lasting from 10 to 17 days,
commonly 11 clays. Thus the life history of the blowfly requires from 16
to 35 days, usually 22 days. The life of the adult is about 35 days on an
average.12
Phaenicia (= Lucilia) sericata (Meig. ) is of a yellow-green or
cupreous-green metallic color varying from metallic blue and green to
copper. It may occur indoors but is typically a scavenger. The palpi are
yellow. There are usually three (occasionally four) postacrostical bristles
present on each side. The second abdominal segment is devoid of mar-
ginal macrochaetae present in L. sylvarum Meig., which also has black
palpi. At a temperature of 80° ± 2° F, with beef lung or fish as food ,
FIG. 105. A common blowfly, Calli-

phora vomitoria.
the entire life history of Phaenicia sericata (Meig.) from the deposition
of the egg to emergence of the fly requires about 12 days: egg stage
(the egg hatching the same day if deposited during early morning)
about eight hours; larval stage (feeding period) about two and a half
days; prepupal stage (migrating larvae) about three days; pupal period
about six days. It is the most abundant species of the genus in North
America, particularly in northern United States and southern Canada
(Hall). P. sericata (Meig.) lends itself well to rearing in large numbers
for experimental purposes. Rearing procedures are described by Dor-
man, Hale, and HoskinsY
The size of the flies and the sex ratio" varies according to the amount
of food available during the larval or feeding stage. The sex ratio of 2.8
to 3.1 males to 6.9 to 7.2 females for flies resulting from larvae which fed
until they left the food voluntarily, Le., from 72 to 78 hours, is reversed
to 6.2 to 6.5 males to 3.5 to 3.8 females in flies in which the larvae were
permitted to feed only 30 to 36 hours, i.e., were underfed.
HOUSEFLIES 329
Lucilia illustris (Meig.) is a holaractic species, widely distributed
in North America; it is common in the midwestern portion of the
United States where it ordinarily deposits its eggs on carcasses of ani-
mals in competition with Phaenicia sericata (Meig.). It appears to be
an open woodland and meadow species. North American authors have
heretofore not fully recognized the fact that they were pretty surely
dealing with L. illustris and not with Lucilia caesar (Linn.) as they
often stated. The two species resemble each other very closely. Hall
(loc. cit.) states that Lucilia caesar (Linn.) does not occur in North
America. The thorax of L. illustris is metallic blue-green with bronze
and purple reflections; the legs are black; the normal number of lateral
bristles in Lucilia is three.
Phormia regina (Meig.), known as the black blowfly, is a Widely
distributed holaractic species; it is found throughout the United States
and as far south as Mexico City. It commonly deposits its eggs in the
wool of sheep. It is said to be a cold-weather blowfly, occurring most
abundantly during the early spring months and becomes less abundant
as hot weather approaches. The thorax is black with a metallic bluish-
green luster; there are darker black longitudinal stripes on the dorsum
extending somewhat beyond the suture; the legs are shining black; the
abdomen is olivaceous or bluish green to black, and ~hining; the length
of the fly is 6 to 11 mm.
Sarcophaga haemorrhoidalis (Fall.), one of the numerous species of
flesh flies, occurs throughout North America as well as Europe. It meas-
ures 10 to 14 mm in length; in color it is gray. The terminalia of the
female are red. It reminds one of an overgrown housefly, but it is lighter
gray, the eyes are brighter reddish brown in color, and it is larviparous.
The larvae have a wide range of feeding habits, being, however, prima-
rily scavengers. They feed on dead insects, carrion, mammalian excre-
ment, etc.
The larvae may be deposited on the hand of a person holding the
female fly. The life history in the presence of ample food and warm
temperature requires from 14 to 18 days. The growth of the larvae is
very rapid after extrusion when food such as carrion is available. The
larval stage may be completed in about three days, followed by the
prepupal or migratory stage lasting usually about three days. The pupal
stage requires from 8 to 10 days. Sarcophaga carnaria (Linn.), measur-
ing 10 to 16 mm in length, is another widely distributed and common
species of flesh fly.
Germ carriers. The common housefly, Musca domestica Linn., is by
accident of habit and structure an important and dangerous disease-
transmitting insect. In habit the housefly is revoltingly filthy, feeding in-
. discriminately on excrement, vomit, and sputum, and is, on the other
hand, equally attracted to the daintiest food of man. The housefly's

proboscis is provided with a profusion of fine hairs which serve as col-
lectors of germs and filth; the foot (Fig. 106) of the fly when examined
under the microscope presents an astonishing complexity of structure.
Each of the six feet is equally fitted with bristly structures and pads
which secrete a sticky material, adding thus to their collecting ability.
When the fly feeds it regurgitates droplets used in liquefying solid food,
and extrudes droplets of excrement as well. Its structure, added to its
natural vile habits, make the housefly an ideal transmitter of filth
diseases.
The common housefly has long been known to contaminate food,
but has, nevertheless, been regarded as a scavenger, and thus a beneficial
FIG. 106, Foot of the common house-

fly. (Much enlarged.)
insect; however, if there remains any doubt in the mind of the reader
as to its harmfulness, after pondering what follows, let him take the
time to make a few careful observations for himself.
In order to show that the housefly (Musca domestica Linn.) can
carry "germs" of a known kind, a simple test can be made with a cul-
ture of Staphylococcus aureus. After the fly is allowed to walk about in
the culture tube it is transferred to a sterile ~lgar plate upon which it is
permitted to walk for about three minutes. The plate is then incubated
for 24 hours. Figure 107 shows the trail of the fly in one of our tests;
every place that the foot touched is plainly marked by a vigorous bac-
terial growth.
Esten and Mason 15 in an article entitled "Sources of Bacteria in Milk"
state:
The numbers of bacteria on a single fly may range all the way from 550
to 6,600,000. Early in the fly season the numbers of bacteria on flies are com-
paratively very small, while later the numbers are comparatively very large.
The place where flies live also determines largely the number that they carry.
The average for 414 flies was about one and one-fourth million bacteria on
each. It hardly seems possible for so small a bit of life to carry so large a num-
ber of organisms. The objectionable class coliaerogenes type was two and one-
half times as abundant as the favorable acid type.
HOUSEFLIES 331
A significant study was made by Yao, Yuan, and Huie '6 in Peiping,
China. This was based on a total of 384,193 flies, of which 98.4 per cent
were Musca domestica Linn., 1.1 per cent Fannia canicularis (Linn.)
and F . scaIm'is (Fabr.), 0.31 per cent .Lucilia caesar (Linn.), 0.16 per
cent Calliphora vicina R. D. and C. vomitoria (Linn.), and 0.03 per cent
Sarcophaga carnaria (Linn.). They found an average of 3,683,000 bac-
teria per fly in the slum district, and an average of 1,941 ,000 for the
cleanest district. They found eight to ten times as many bacteria inside
the flies as on the outside.
FIG. 107. Cultures of Staphylococcus aureus transferred by a housefly to

a sterile agar plate upon which it was allowed to crawl for three minutes.
Incubation period was 24 hours.
The fly usually acquires infection by walking over infectious mate-

rials, both its feet and wings becoming contaminated. The intestinal
contents of flies also become charged with infection when feeding, and
this is dejected in the fly "specks," and vomit droplets. It seems plausible
that flies might become infected in the larval stage by developing in
infectious fecal matter and that the newly emerged and unfed flies
would be dangerous. Under experimental conditions Graham-Smith 17
has produced infected blowflies by feeding the larvae on meat infected
with spores of Bacillus anthracis. He found that the blowflies remained
heavily infected for at least two days after emerging and that the bacilli
I •

could be cultivated either from the legs and wings or intestinal contents
of flies more than 15 or 19 days old.
The opportunity for flies to become infected is so great in all com-
munities, even the most sanitary, that no fly should be trusted to alight
on food prepared for human consumption. The following quotation is
from Nuttall and Jepson. 18
It should be remembered that a fly may cause relatively gross infection

of any food upon which it alights after having fed upon infective substances,
be they typhoid, cholera or diarrhea stools. Not only is its exterior contami-
nated, but its intestine is charged with infective material in concentrated form
which may be discharged undigested upon fresh food which it seeks. Con-
sequently, the excrement voided by a single fly may contain a greater quantity
of the infective agents than, for instance, a sample of infected water. In poten-
tial possibilities the droppings of one fly may, in certain circumstances, weigh
in the balance as against buckets of water or of milk.
Leidy19 in 1871 expressed an opinion that flies were probably a

means of communicating disease to a greater degree than was generally
suspected. From what he had observed in one of the large military
hospitals, in which hospital gangrene had existed during the Civil War,
he thought flies should be carefully excluded from wounds. It was, how-
ever, not until the Spanish-American War in 1898 that the real menace
of the fly became evident as indicated by the following quotation from
an article by Veeder in the Medical Record of September 17, 1898:
To clinch the argument, and apparently to leave no loophole for escape,

I have made cultures of bacteria from fly tracks and from the excrement of
flies, and there seems to be not the slightest difficulty in so doing. Indeed the
evidence of every sort is so clear that I have about reached the conclusion that
the conveyance of infection in the manner indicated is the chief factor in
decimating the army.
Gastrointestinal diseases. The housefly is primarily a food contamina-

tor and vector of filth diseases because of its feeding and breeding habits,
as already explained. Pathogenic organisms are collected on feet and
mouth parts and ingested while feeding, then deposited mechanically \
while the fly is crawling on human food or deposited by regurgitation
or with the fly's excrement.
Of the hQusefly's ability to transmit typhoid bacilli, Jordan 20 writes:
Not only may bacilli stick to the legs and wings of these insects, but if
swallowed they may survive the passage of the alimentary tract. Typhoid
bacilli have been isolated from houseflies captured in houses in Chicago, in the
neighborhood of badly kept privy vaults used by typhoid patients, and it has
HOUSEFLIES 333
been shown experimentally that living bacilli may remain in or upon the body
of flies for as long as twenty-three days after infection.
Yao, Yuan, and Huie (loc. cit.) found Shigella (= Bacillus) dysen-
teriae (Shiga) in 15 of 50 batches of 100 flies tested. Alimentary tracts
of flies also revealed cysts of Endamoeba hist()lytica. The specific death
rate for gastrointestinal diseases roughly paralleled the number of flies
captured.
Faichnie21 (1929) in a study of the etiology of enteric fever came to
the conclusion that comparatively little typhoid is carried on the feet
of flies, but he found that both E. typhosa :'Ind Salmonella paratyphi
( Kayser) (B. paratyphosus A) multiplied in tbe intestines of flies fed on
infected excrement.
Cholera was among the first diseases in which the housefly was in-
criminated as a carrier. Tizzoni and Cattani in Bologna in 1886 isolated
cholera vibrios from flies caught in cholera wards. Simmonds in 1892
captured Shes in tne post-mortem morgue in Bamburg aml isoh'teo
cholera vibrios from these in large numbers. "Upon the surface of
vegetables and fruits kept in a cool moist place, experiments have shown
that the spirillum may retain its vitality for from four to seven days"
(Jordan).
In their study of the epidemiology of cholera, Gill and La}22 found
evidence to support the startling suggestion that possibly one phase of
the life cycle of the cholera vibrio may be passed in the body of the
housefly. The results of their work show that the vibrios disappeared
from the body of the fly after about 24 hours but reappeared on or about
the fifth day, at which time the fly was capable of infecting food by its
feces.
Yaws (Frambesia) is caused by Treponema pertenue Castellani. The
disease is widely distributed in the tropics. The spirochetes are found
in the superficial ulcers on the hands, face, feet, and other parts of the
body. The following statement by Nuttall and Jepson (1909, loco cit.)
presents evidence showing that Musca domestica Linn. is amply able to
transmit this infection:
Castellani (1907) tested the matter of the fly transmission of yaws by
experimental methods. He allowed M. domestica to feed (1) upon yaws mate-
rial (scraping from slightly ulcerated papules), and (2) upon semi-ulcerated
papules on the skin of these yaws patients. In both cases he was able to dis-
cover the Treponema (Spirochaeta) pertenuis in microscopic preparations made
from the flies' mouth parts and legs. Furthermore, he allowed M. domestica to
feed on· yaws material (1 and 2 as above) and afterwards transferred them
to scarified areas upon the eyebrows of monkeys. Of 15 monkeys thus ex-
perimented upon, three developed yaws papules itt the places which had been
contaminated by the flies.
Ophthalmia. In commenting on ophthalmia as carried by flies,
Howard (1911, lac. cit.) has the following to say:
Dr. Lucien Howe of Buffalo informed the writer [Howard] that in his
opinion the ophthalmia of the Egyptians is also transferred by Hies and pre-
sumably by the housefly, and referred the writer [Howard] to a paper which
he read before the Seventh International Congress of Ophthalmology at Wies-
baden in 1888. He referred to the extraordinary prevalence of purulent oph-
thalmia among the natives up and down the river Nile and to the extraordinary
abundance of the flies in that country. He spoke of their remarkable indiffer-
ence to the visits of flies, not only children, but adults, allOWing flies to settle
in swarms about their eyes, sucking the secretions, and never making any
attempt to drive them away. Doctor Howe called attention to the fact that
the number of cases of the eye disease always increases when the flies are
present in the greatest numbers and the eye trouble is most prevalent in the
place where the flies are most numerous. In the desert, where Hies are absent,
eyes as a rule are unaffected. He made an examination of the Hies captured
upon diseased eyes and found on their feet bacteria which were similar to
those found in the conjunctival secretions. Flies captured in Egypt swarm-
ing about the eyes of'" ophthalmia patients and sent to Washington, D. C.,
were identified as Musca domestica.
Poliomyelitis. Flies, biting and nonbiting, as well as mosquitoes have

long been under suspicion as vectors of the virus of poliomyelitis. The
virus of poliomyelitiS has been isolated from human stools and sewage
(Paul and Trask,23 1941). Various species of muscoid flies breed in excrec
ment and sewage, as has been pointed out earlier in this chapter. The
presence of the virus of poliomyelitis has been demonstrated in flies
collected in the field during both urban and rural epidemics by various
investigators. (Sabin and Ward,24 1941.) Sabin and Ward 25 (1942) state
that there is no longer any doubt that flies can carry the virus. The virus
was isolated from one of three batches of flies (in Atlanta), more than
95 per cent of which were Musca domestica. Isolation of the virus was
made from 7 out of 12 batches of flies (collected in Cleveland, Ohio,
where fresh meat was added to the banana and sugar); from 90 to 95
per cent of these flies were Calliphoridae (blowflies). Hall 10 who iden-
tified the flies points out that all of the positive samples of flies Cal\-
tained Phaenicia (= Lucilia) sericata (Meig.) and Phormia regina
(Meig. ). Just where and how these flies originally obtained the virus
is an unsolved problem, as Hall remarks. Positive experimental evidence
was secured by intraperitoneal inoculation into Cynomolgus monkeys
of etherized fly extract; unetherized material was given both intranasally
and by mouth. Precisely what role, if any, flies play in the transmission
of poliomyelitis to human beings is not clear; that flies might serve as a
link in the infection of animal reservoirs is of course not impossible.
HOUSEFLIES 335
Since it is known that muscoid flies may harbor the virus of poliomyelitis,
every effort should be made to control flies by any means possible; be
it by environmental sanitation or by the use of insecticides, which nor-
mally should be only supplemental to good sanitation. Fly control needs
encouragement. However, in the face of evidence that poliomyelitis can
be spread by human carriers and since evidence of transmission by
flies is incomplete, fly campaigns recommended as a means of poliomye-
litis control, particularly the community-wide spread of DDT or other
insecticide by airplane, only creates false hope and false security. It must
be borne in mind, too, that such treatment only affects the adult fly
population; the residual effects are not likely to influence in any large
measure the following broods of flies.
Tuberculosis. Investigations by Dr. Ch. Andre of the University of
Lyons were reported at the Anti-Tuberculosis Congress at Washington,
1908, as follows:
Flies are active agents in the dissemination of Koch's bacillus because

they are constantly going back and forth between contagious sputa and feces,
and foodstuffs, especially meat, fruit, milk, etc., which they pollute by con-
tact with their feet, and especially with their excretions.
The experimental researches of the author show the following:
1. Flies caught in the open air do not contain any acid-fast bacilli that
could be mistaken for the bacillus of Koch.
2. Flies that have been fed on sputum evacuate considerable quantities
of bacilli in their excretions. The bacilli appear six hours after ingestion of the
sputum, and some may be found as long as five days later. These flies, there-
fore, have plenty of time to carry these bacilli to a great distance, and to
contaminate food in houses apparently protected from contagion, because not
inhabited by consumptives.
3. Food polluted by flies that have fed on sputa contains infective bacilli
and produces tuberculosis in the guinea pig.
4. Flies readily absorb bacilli contained in dry dust.
5. Flies caught at random in a hospital ward produced tuberculosis in the
guinea pig.
Andre's conclusions are: "The sputa and feces of tuberculosis sub-

jects must be disinfected; flies should be destroyed as completely as
possible; foodstuffs should be protected by means of covers made of
wire gauze."
Eggs of parasitic worms. Extensive and careful work on the dispersal
of eggs of parasitic worms by the housefly has been done by Nicol}.2"
The following is a summary of his investigations. Flies feed readily on
excrement in which eggs from parasitic worms occur. Eggs may be con-
veyed by flies from excrement to food in two ways, namely on the ex-
ternal surface of tpe body and in the intestines. The latter is possible
only when the diameter of the eggs is under .05 mm. Eggs with a diame-
ter of up to .09 mm may be conveyed on the external surface; however, .
these adhering eggs are usually got rid of by the fly within a short time,
while those harbored in the intestine may remain there for several days.
It was found that material containing eggs of parasites, and in particular
ripe segments of tapeworms, remain a source of infection through flies
as long as two weeks.
The eggs of the following parasitic worms were shown experimentally
to be capable of transmission by Musca domestica Linn.: Taenia solium
Linn., Taenia pisiformis (Bloch), Taenia hydatigena Pallas, H ymeno-
lepis nana (v. Siebold), Dipylidium caninum (Linn.), Diphyllobothrium
latum (Linn. ), Enterobius vermicularis (Linn.), Trichocephalus (=
Trichuris) trichiurus (Linn.), both internally and externally, Necator
americanus (Stiles), Ancylostoma canium (Ercolani), Ascaris equorum
Goeze, Toxascaris leonina (v. Linstow), Hymenolepis diminuta (Ru-
dolphi), externally only. No trematode parasites were experimented
with, and the observations of Stiles that the larval fly can ingest ascarid
eggs and pass them on to the adult fly was not confirmed.
Intestinal protozoa. Roubaud 21 found that the cysts of Endamoeba
coli (Grassi), Endamoeba histolytica (Schaudinn), and Giardia lamblia
Stiles passed through the intestine of the fly uninjured, and that free
amebae (both coli and histolytica) when fed to flies were found dead
in the fly's intestine in less than an hour. It is generally believed that
the housefly plays an important role in the epidemiology of amebiasis
through the spread of cysts of Endamoeba histolytica. Root 28 found
motile Chilomastix mesnili (Wenyon) in a fly's feces seven minutes after
it had fed on an infectious stool.
Murrina, a trypanosomiasis of horses, mules, and burros in Panama,
is caused by Trypanosoma hippicum Darling. Darling29 showed that the
disease can be transmitted mechanically by houseflies by contact with
fresh blood from an infected animal and thence to an open wound of
another animal. The disease is, of course, transmitted in other ways,
vampire bats playing an important role.
Bovine mastitis. Sanders (loc. cit.) reports investigations made at
the Florida Agricultural Experiment Station which incriminated the
houseflies (Musca domestica) as well as Hippelates flies as vectors of
bovine mastitis. Houseflies (also Hippelates) were seen to alight
at the natural openings of calves, yearlings, pregnant heifers, and
lactating cows. They fed on lachrimal fluid, fatty body secretions, milk
droplets accidentally spilled, and on secretion at the tip of the teats of
animals in herds where mastitis prevailed. Exposure tests were made
with flies feeding alternately on infected material and the teat orifice;
also the teat orifice was exposed to flies taken directly from premises
HOUSEFLIES 337
wher~ mastitis prevailed. "Mastitis developed in each of the experimen-
tal animals by the exposure technique employed."
Cutaneous habronemiasis. An examination of certain persistent ulcera-
tions, summer sores, on the lower portions of the bodies of horses (the
sores have a tendency to disappear during colder weather) may reveal
the presence of larval nematode worms belonging to the genus Habrone-
FIG. 108. Segment of intestine (inside out) of fowl infested with numerous
tapeworms, Choanotaenia infundibulum.
ma measuring from 1 to 1.5 mm in length. The presence of these worms

in sores on the eyes is termed habronemic con;unctivitis. Adults of
Habronema muscae (Carter) , measuring from 8 to 14 mm in length in
the male and 13 to 22 mm in the female, occur in the stomach of the
horse, where they lay their eggs, which pass out with the feces. The
newly hatched larvae find their way into the bodies of fly larvae, which
are evidently true intermediary hosts, and in these further development
occurs. Flies resulting from these larvae may contain a number of the
worms (from 1 to 20) in their bodies, often in the hood. According to
Ransom 3o the infection of horses with H. muscae is apparently brought
about by the swallowing of infected flies or infection by the worm larvae
migrating from the insect as it feeds. The sores above-mentioned are
evidently the result of larvae entering lesions while the animals are lying
down in infested manure.
Fowl taeniasis. Domestic fowls are commonly infested with tape-
worms and the extent to which this may occur is well illustrated in Figure
108. Several species of tapeworms inhabit the intestines of fowls. Al-
though Grassi and Rovellpl as early as 1886 had made observations con-
cerning the development of these parasites, it was not until 1916 that
experimental evidence was published by Gutberlet,32 proving the house-
fly (Musca domestica Linn.) to be an intermediate host.
The most important of the fowl tapeworms is Choanotaenia infun-
dibulum (Bloch). It measures from 50 to 200 mm' in length. The scolex
is small and rounded, measuring about 0.4 mm in width. The rostellum
is armed with a single row of 16 to 20 hooks. The cysts found in the
housefly are oval in shape and measure about 200p. in length by 120p.
in diameter. Gutberlet infected the adult flies by feeding them on liquids
which were infested with tapeworm eggs, and it is assumed that fly
larvae breeding in infested fowl droppings would become similarly in-
fected. Reid and Ackert 33 in experiments with this tapeworm recovered
proglottids in fowl feces at the end of the seventh week. These authors
point out that buttermilk is very attractive to both flies and chickens and
that chickens devour the flies eagerly.
Raillietina cesticillus (Molin), according to Gutberlet, also has the
housefly as its host; and Hymenolepis carioca (Magal.) has Stomoxys
calcitrans (Linn.), the stable fly as its host.
FLY CONTROL
Fly control. Effective fly control is based on a knowledge of the habits
and life history of the particular offending species. First find the hreeding
places and then apply the appropriate remedy. The presence of flies
always denotes defective sanitation, particularly in the disposal of ma-
nures, garbage, sewage, slops, food wastes, ensilage, brewer's grain,
spent hops, wet mash, dead animals, etc. The prevention of fly breeding
requires good housekeeping practices in a broad sense.
Rural fly control. Since the principal rural breeding places of house-
flies are usually in and about barns and stables, particular attention must
be paid to these as well as to barnyards and corrals, with speCial reference
to the disposal of manures and the prevention of accumulations of
manure in the stable. Concrete floors permit proper cleaning of stalls.
In constructing a concrete floor, provision must be made for carrying
away the animals' urine and the water used in cleansing the floors and
HOUSEFLIES SS9
stalls; the stall Hoors should be given a one-inch drop from the manger
to the manure gutter, which latter should be "6 inches deep and 14
inches wide. In order to facilitate the drainage of the liquids a 3-inch
U-shaped channel is sometimes made in the bottom of the gutter next
to the manure alley, but this is not necessary where a slope is given the
gutter bottom. The gutter should be given a uniform fall of 3 inches to
100 feet, and the Hoor of the manure alley should have a slope towards
the gutter of 1 inch to 10 feet. A small watertight liquid manure cistern
may be provided outside the barn into which the gutter drains, but if a
manure shed is used, the cistern should be in the shed. The gutter should
be connected to the cistern by means of a drain pipe effectively trapped
like the soil pipe in a house and so arranged that the trap may be easily
cleaned."34
Concrete stall Hoors should be covered with wood to prevent animals
from coming in direct contact with the concrete. The super-Hoar should
be so made that it can be lifted while the concrete is being cleaned. If
the crevices of the wood Hoor are not also frequently cleaned, Hy larvae
will develop in these also.
Manure wastage. Piling manure in a barnyard results in a loss of
manurial value due to leaching and fermentation, estimated at from 25
to 50 per cent. The Cornell University Experiment Station has carried
on investigations which show the loss of valuable plant food when
manure is disposed of in the usual exposed manner for six months. When
the manure was tested, it was found that the horse manure had lost 57
per cent and the cow manure 49 per cent in gross weight, and the loss
in value based on plant food (nitrogen, phosphorus, and potassium)
amounted to 65 per cent for the horse manure and 23 per cent for the
cow manure. When manure wet with urine is thrown from the stable
on to the heap it contains about 75 per cent water which holds most of
the plant food. Exposed to leaching rains and weather the liquid sinks
into the ground beneath or Haws away. Thus not only does the barnyard
manure pile result in flies but also in a loss to the farmer.
Manure disposal. Wherever manure is piled up and accessible to flies,
these insects are afforded opportunity to breed. As stated before, it
requires only about four days for the larvae to reach full growth, after
which they begin to migrate into the drier portions of the heap and
crawl out into nearby debris, beneath platforms, etc. It is therefore
imperative, if fly breeding is to be prevented, that manure be properly
handled at once.
Under ordinary rural conditions it would seem possible to remove
the manure to the fields every two or three days. For this purpose a cart
may be backed up against the stable doorway, and the manure may be
thrown in, carted away to a field, and scattered. This saves much time in
handling and is sound agricultural practice. Since moisture and warmth
are both necessary for the production of fly larvae, the scattered manure
cannot serve this purpose.
According to the Wisconsin Bulletin No. 221 "Manure is never so
valuable as when perfectly fresh, for it is impossible under the best sys-
tem of management to prevent all loss of its fertilizing ingredients. For
this reason, whenever possible, the manure should be hauled directly to
the field and spread. The system saves time and labor as it involves
handling but once. The manure will be leached by the rain and snow,
nevertheless the soluble portion will be carried into the soil, where it is
needed. When spread in a thin layer, it will not heat, so there will be no
loss from hot fermentation, and where manure simply dries out when
spread on the ground, there is no loss of valuable constituents."
A manure spreader is a valuable part of farm equipment. Farmers
and gardeners who wish to use "rotted" manure for fertilizing purposes
should compost it; few flies breed in "rotted" manure. Fresh manure may
also be placed in trenches and covered with lime and earth or it may be
stored in fly-tight composting pits.
Composting pits. Composting pits are frequently maintained on coun-
try estates and truck gardens where quantities of rotted manure are used
for fertilizing purposes. Such pits are usually made of concrete and
covered with wood, all carefully constructed to exclude flies and mos-
quitoes, which latter may breed in the liquids collected in the sump.
A properly constructed composting pit makes it possible to preserve the
urine, which is very valuable in addition to the more solid excreta.
A cross section of a composting pit is shown in Figure 109. In this
case a pump is indicated by means of which the urine and water are
pumped out of the sump and returned to the manure from time to time.
Water should be applied to the manure occasionally to prevent burning,
which may destroy much of the value of the fertilizer.
The size of the pit depends also on the number of horses stabled.
A pit such as is shown in the figure, with a length of 60 feet, ought to
store the manure from 10 horses for a period of six months.
Close packing. The following description of close packing is from
Austen (loe. cit.), who states that the essence of this method is the
utilization of the natural heat of fermenting manure for the destruction
of the eggs, maggots, and pupae (if present) of the housefly:
For close packing (a method introduced in 1915 by Lieut.-Colonel S. A.
Monckton Copeman) an area of hard, level ground, at least three or four
feet greater in extent each way than the ultimate size of the intended dump,
must be selected or prepared to receive the manure. On this, each day's
manure is utilised in forming or adding to a compact rectangular block, which
may be of any desired dimensions horizontally, but for convenience of treat-
HOUSEFLIES 341
ment should not exceed five feet in height. Each load of manure 011 being
added to the dump must be pressed down firmly with shovels, and if the
weather be dry should be sprinkled slightly with water; finally the sides,
which should be somewhat sloping, must be beaten and smoothed down with
the shovel. . . . It was found by Colonel Copeman that, four inches beneath
the surface of a heap of fresh stable manure treated in this way, the heat pro-
0
duced by fermentation may be as much as 169 F, though housefly maggots
0
are speedily killed at temperatures only slightly above 114.8 F. It should
'",
.'"
, Pit 24" sqlJore 24"deep
FIG. 109. Cross section of a concrete flyproof composting pit. (After

Rosenthal in Advisory Pamphlet on Camp Sanitation and Housing, Commis-
sion of Immigration and Housing of California.)
be noted that the fertilizing value of close packed manure is greater than that
of loosely stacked manure .
Although the success of close packing as a preventive of housefly breeding
in horse-manure has been proved in England, the method has yet to be tested
in warmer climates. In the event of failure from some unforeseen cause, the
portion of the dump (both sides and top) in which maggots are seen should
be covered over with a layer of sacking (old coal sacks, if without holes,
answer well) , soaked in heavy oil and secured by means of large stones; the
sacking need only be allowed to remain for one week, after which if required
it may be employed on another part of the dump.
Chemical treatment of manure. Although there is a strong popular
demand for some chemical which, if applied to the manure, will result
in fly prevention, the writer is not enthusiastic about this method of con-
trol for two reasons; first, this scheme may be used as a substitute for
sanitation-cleaning up and keeping clean; and second, owing to the
necessity for frequent application, there will certainly be neglect.
Ordinary applications of the insecticides are of little or no avail,
because of the hardiness of fly larvae, and furthermore, the larvae cannot
be reached easily, buried as they are in the bedding or manure.
Borax is recommended by the United States Department of Agricul-
ture at the rate of 0.62 pound to 8 bushels of manure. The New Jersey
Agricultural Experiment Station recommended the use of borax, stating:
One ounce of borax to one cubic foot of manure will kill 90 per cent or
more of the larvae and inhibit the eggs from hatching, provided it is applied
in solution, and the larvae and eggs come in contact with it. It was figured out
that the average output from one horse is one and one-half cubic feet of
manure per day and that two and one-half gallons of water containing one and
one-half ounces of borax should penetrate all parts of this amount of manure.
In making treatments, it was noted that wet manure which had been soaked
by rains, or when located in low wet areas, was difficult to treat, for it was
next to impossible to make any additional water (by sprinkling) penetrate the
wet manure. When manure is in this condition, particularly in low wet areas,
and contains numerous larvae, it should be put on higher ground in order
that it may dry out somewhat; or a thick layer of gravel or cinders placed
under the manure in wet low areas will help considerably. Manure treated
with borax is detrimental to some plants when the manure is used as a fer-
tilizer, but, as far as known, it will not injure plants provided one does not
use over 15 tons of treated manure (not over one pound of borax for every
16 cubic feet) to the acre.
As a practical matter 11 ounces of borax in 10 gallons of water is

sufficient to treat the daily accumulation from 10 animals. Borax is an
excellent chemical to use on dirt floors of stables, in privies, and on ac-
cumulations of decomposing matter which will not be used as fertilizer.
Dust has considerable value in fly control. The author found that
very few flies bred in the manure from cavalry horses on duty during the
summer in arid dusty regions when the manure was thoroughly mixed
daily with road dust and disposed of in windrows a short distance from
camp.
Use of DDT in rural fly control. Gunderson, extension entomologist,
Iowa State College, who played an important role in the successful fly
control campaign in Iowa ("Make Iowa Fly Free in 1947") sets up the
following program of action:
1. Clean up fly breeding places, such as manure piles, hay and straw
stack bottoms, weed piles, and other decaying plant and animal materials.
Scatter on the fields every 3 days for greatest fertilizer value.
HOUSEFLIES
2. Treat outdoor toilets and manure piles with borax (11 ounces ill 10
gallons of water on each day's accumulation for 10 animals). These places
may be sprayed every 7 days with a 2lh per cent DDT spray to accomplish
similar results.
3. Spray inside walls of an all farm out-buildings housing livestock or live-
stock products with 2lh per cent DDT (2 pounds 50 per cent DDT wettable
powder in 5 gallons of water). Apply at the rate of one gallon per 1,000
square feet of surface. Dairymen should check with local milk inspector be-
fore applying this formula in the milk house. Some inspectors require DDT
emulsion or oil spray in such places.
4. Spray cattle every two weeks or as often as necessary. Use one pound
of 50 per cent DDT wettable spray powder in ten gallons of water for early
sprays. Increase to two pounds of 50 per cent DDT wettable powder in five
gallons of water when flies are found on legs and bellies of cows. Use 1 to 2
quarts of spray per animal.
Dairy barns. Michelbacher and Smith35 (1946) report excellent re-

sults obtained with DDT in the control of Hies in dairy barns. They
recommend DDT water suspension sprays in preference to DDT-kero-
sene solution for several reasons: (1) Based on the actual amount of
DDT used, water suspension sprays proved to be more effective, prob-
ably because kerosene carries part of the DDT into the sprayed surface
where it is lost, while a water suspension spray deposits DDT on the
surface. (2) Some fire hazard is involved where a kerosene spray is
used, thus vitiating insurance contracts. (3) Water suspension is a much
safer material to use since it is not harmful to man or animals if some
of the spray is aCCidentally deposited on the skin. Water suspension
sprays were used at the rate of 2 to 4 pounds of actual DDT to 100
gallons of water. The 4-pound dosage was preferred because of its longer
lasting effect. It was applied, at pressures ranging from 100 to 400
pounds, to the interior surfaces of the barns, including alleyway, wash-
room, and milk-cooling room, also to ceiling and walls of the feed storage
room provided the feedstuffs were protected from the spray. Frequently
the calf barn and the bull pen were also treated. To do a thorough job
40 to 60 gallons of dilute spray were needed to treat a 24-cow barn.
During the summer this treatment lasted for about six weeks; three to
four spray applications a year are suggested. The following precautions
must be taken before spraying begins: (1) All milk cans and utensils
put under cover or removed; (2) milk coolers and sinks carefully cov-
ered with heavy paper or other material; (3) all feedstuffs removed or
carefully covered to avoid wetting with spray; (4) feed troughs hosed
clean just before and after the spray is applied in order to reduce spray
residue which might collect in them. The authors point out that the but-
terfat production per cow increased from 394 to 429 pounds in dairies so
treated; other factors may, of course, have played a part in bringing
about this increase, but fly control was, no doubt, an important factor.
Feed wastes. The presence of numerous flies about the premises of
a certified dairy, which upon inspection was found to be very clean and
otherwise in excellent condition with no evident reason for the presence
of the flies, was a matter of no small concern. The trouble was located
in a field a few hundred feet from the dairy bam where numerous heaps
of waste brewer's grain (used as cattle feed during late summer and
autumn) had been hauled and dumped. Each heap was a veritable
wriggling mass of housefly maggots. Liberal application of a larvicide
(fuel oil in this case) to the heaps soon destroyed the source, and further
waste feed was spread out in a thin layer to facilitate rapid drying and
thus defeat fly breeding. The epidemic of flies was soon brought under
control. An adulticidal spray such as DDT will quickly eradicate the
infestation. Fly breeding about dairies is frequently traceable to accumu-
lations of waste feed about the troughs, the waste becoming mixed with
feces and urine. Thus bad conditions may arise in spite of the proper
disposal of manure.
Community fly control. Among the earliest successful community-wide
fly-control campaigns was one conducted by the author4 in Berkeley,
California, in 1909-1910 during the "horse-and-buggy" days when there
was really a fly problem. The population of Berkeley was about 40,000
at that time. A letter from the then secretary of the Chamber of Com-
merce dated April 8, 1909, reads in part, " . . . it is my desire to enlist
your aid for the relief of Berkeley. . . . This office is located between
two livery stables that breed myriads of flies." The campaign was started
at once with the full cooperation of all city officials and the Board of
Health through the health officer. Much newspaper publicity was given
to the work and the Board of Health distributed bulletins written by the
author and entitled "Essentials of Housefly Control" and "Fight The
Fly, Why, When and How." Twenty students from the class in medical
entomology were offiCially appointed sanitary inspectors. A detailed
house-to-house inspection was made to locate breeding places, and ad-
vice was given as to means of control. Police officers assisted when diffi-
cult situations arose. Although insecticides were used to some extent, the
campaign was primarily based on sanitary procedures.
After a few years of community interest in housefly control there
followed a lull in interest, perhaps due in large measure to conversion
of livery stables into garages, a change over from the horse to the auto-
mobile. During the past three years there has been a widespread popular
demand for fly-control programs, owing no doubt to the popular associa-
tion of flies with the spread of poliomyelitis, and to the enthusiastic
approval of the new miracle insect killer, DDT.
HOUSEFLIES 345
Rowe of the United States Public Health Service, in a modem
36
presentation (1948) points out that in the conduct of the most effective
and economical community fly-control programs both preventive and
insecticidal measures are essential. He states that too much emphasis
cannot be placed upon preventive measures. These are incorporated in
what are known as standard environmental sanitation practices. How-
ever, such practices must be specific and directed toward the abatement
of all fly-breeding situations in so far as possible. In agricultural com-
munities stables, corrals, and feed pens may be fly-breeding hot spots;
the outdoor toilet (privy) may be a contributing factor; garbage and
community garbage dumps may also contrib\lte flies; these situations are
all amenable to relatively simple sanitary and insecticidal procedures.
However, according to Rowe, some of the most vexing fly problems in a
community are created by the haphazard dllmpage or the careless dis-
posal of industrial end-products and wastes. Maior fl.y-breeding foci are
often created around stockyards, stock feeding pens in connection with
beet-sugar refineries, abattoirs, hide and tallow plants, canneries, food
and milk processing plants, et cetera. Vast numbers of flies-houseflies,
blowflies, pomace flies, etc.-may be produced in the accumulated wastes
and contaminated soil around the plants. Special study must be given to
each situation and a practical remedy should be employed that is adapt-
able to each; the program should aim at elimination of the problem
whenever possible rather than at insecticidal treatment. Fly problems
are man-made.
DDT has been the insecticide of choice for community Hy-control
programs in the United States, primarily because of its residual value in
the destruction of adult flies. The housefly lends itself particularly well
to the lethal effect of residual DDT in that it spends much of its life
resting or walking on surfaces which may be readily treated. The lethal
dose of DDT for an average size housefly is a tenth of a microgram,
according to Douglas.37
Three types of DDT sprays have been used, namely, emulsions, solu-
tions, and wettable powder suspensions, with little difference in effective-
ness. Because of the so-called "fire hazard" kerosene or other oil solutions
are seldom used. For all-around uses, inside and outside, the emulsion
type seems to be most suitable. The residuill efficiency of emulsions in
general is rather high as compared with solutions, and they leave little
objectionable residue. There should be a minimum residue of 200 milli-
grams per square foot. Douglas (loc. cit.) believes SOO milligrams is
better and in some situations even 400 to 500 milligrams is actually a
desirable residue. He states that a 200-milligram residue has an effective
residual life of 95 days, and 400 milligrams 110 days. One gallon of 5
per cent DDT spray to 1,000 square feet of surface gives the minimum
of 200 milligrams of residue. Slick surfaces require an increase of the
spray to 7.5 per cent. Application should be with a hand-compression
sprayer or power sprayer giving a wetting spray in a uniform fan-
shaped pattern. The principal resting places of flies should be covered,
such as ceilings, around cracks and crevices, along edges of ceilings,
doorways, and furniture. Electric and switch cords should be painted or
sprayed. The United States Public Health Service warns: Keep food
and water containers covered after spraying the room. Flies fall into them
as DDT takes effect.
DDT-resistant houseflies. Evidence in support of observations made
in many parts of the country that houseflies may be developing resistance
to DDT has been accumulating steadily. Lindquist and Wilson 38 by
selective breeding produced a strain of houseflies that was more resistant
to DDT spray than were flies from their regular stock. About 10 per cent
of the Hies survived the initial "fine-mist spray" treatment, and these
were used as the parent stock in establishing a new "special" colony.
Each of 14 generations of flies was similarly exposed to treatment, and
the survivors were allowed to propagate. The average mortality as the
result of tests with the 14th generation was 69 per cent for the original
strain of flies and 34 per cent for the selected flies.
That this resistance of the flies is not due to a deterioration of DDT
under normal storage conditions has also been proved by tests. It is also
unlikely that flies have changed their habits of alighting on surfaces;
flies have been observed to alight on treated surfaces without toxic effect.
Several other factors may enter into the probable cause of increased
fly populations in spite of DDT treatments, e.g., apparent failures came
during extremely hot weather; DDT is said to be most effective in cool
weather. Also careless use of DDT or inadequate treatment may be a
factor. Certainly the wide use of DDT has resulted in a relaxation of
sanitary practices, thus causing increased populations of flies.
No doubt satisfactory fly control will be achieved in many areas
where proper DDT formulations are carefully applied and sanitation
practices are rigidly adhered to. Much experimental work is being done
with other insecticides, particularly now that fly control has been widely
demonstrated. Among the promising insecticides is benzene hexachloride
wettable powder, 12 per cent concentration at a dosage of 17 pounds
in 100 gallons of spray (Marsh and Metcalf).
Garbage cans. Where garbage cans are used, and certainly every
household should possess a garbage receptacle that is kept tightly closed
against flies, it is strongly urged that all liquids be drained from the
refuse before disposing of it and that the solids be wrapped in a news-
paper before placing in the can. Keep the can clean. In this way fly
breeding in garbage cans may be effectually prevented, and an act of
HOUSEFLIES 347
mercy is done the scavenger and other handlers of garbage. It is a good
practice to spray the outside of the garbage can and the rack on which
it stands every week or two with a 5 per cent DDT oil solution.
Use of manure on lawns. Veritable swarms of flies may suddenly make
their appearance on porches and windows and in the house after com-
paratively fresh manure has been spread on lawns as fertilizer. Such
manure is commonly infested with full-grown maggots or pupae which,
in a few days after the fertilizer has been applied, give rise to a pest of
flies. It is, no doubt, wise to use old composted manure for this purpose
or subject the manure to a thorough steaming or drenching with hot
water at 195 0 F (nearly boiling) before applying it to the lawn.
Two objections are commonly raised against the steaming or drench-
ing method of treatment: first, that the useful bacteria are destroyed,
i.e., the manure may be rendered sterile; and second, that all other de-
sirable constituents are leached out by the water. Not all of the useful
bacteria, by any means, are destroyed by the hot water, and those re-
maining quickly multiply and soon render the manure as good as ever
in this respect. In the second place, the leachings may be preserved quite
easily by placing the manure to be treated in a shallow, tight box similar
to those used by plasterers for mixing mortar, and adding a spigot in a
hole with plug from which the leachings can be drawn and used as
liquid manure.
This method is also useful to mushroom growers who must use rotted
manure in which certain species of fly larvae, mites, etc., may occur in
great numbers.
Railroad cars laden with manure are often sidetracked in or near
communities and are responsible for swarms of flies. The writer has on
several occasions recommended that the periphery of the entire carload
be subjected to a treatment of live steam from the locomotive. This has
been done with good results.
Lawn clippings. Grass clippings from lawns around public buildings
are often dumped in heaps on the premises. Hot weather soon produces
a vile-smelling, decomposing, warm mass of grass which is very attractive
to flies, and within a few days numerous maggots, particularly those of
the housefly, may be found infesting the mass. Thus, again, in spite of the
absence of manure and garbage there may be a veritable plague of flies.
The method of prevention is obvious: the clippings should be spread
out thinly in order to dry out, and they may then be burned or otherwise
disposed of.
Flies from septic tanks. Small invasions of flies may be traced to
nearby septic tanks covered with knotty or imperfect wood superstruc-
ture which permit ingress of flies. Flies may be seen coming and going
through these apertures, and examinat!on of the top sludge will reveal
countless maggots and pupae on the surface. Closing the knot holes or
other apertures with tin or otherwise will soon end the trouble.
Sewage treatment plants. Flies of many species, notably the common
housefly and blowflies, may be attracted to sewage treatment plants be-
cause of odors; and occasionally under improper management countless
numbers of flies originate in sludge beds at the end of the treatment
process. There may also be much fly breeding in wet sludge when this
is applied to the soil as a fertilizer. The control of a breeding source of
this kind may be accomplished by deep plowing and, of course, dis-
continuance of the use of wet sludge as a fertilizer.
FIG. 110. A sanitary privy: front view to left, rear and side view to right.
(After Stiles and Lumsden.)
Sewage treatment plants of modern construction do not as a rule
breed flies. However, under all circumstances raw sewage must not be
accessible to flies because of possible transport of pathogenic organisms
of fecal origin. Sewage-works engineers suggest that fly breeding may be
effectively prevented by quick drying of sludge. Chemical treatment of
sludge is not generally favored, although spraying screenings with a
solution made of one and a half pounds of pure carbolic acid and 40
pounds of caustic soda in 60 gallons of water has been recommended to
stop fly breeding. The use of DDT as suggested in community fly control
is recommended.
Privies. In the absence of modern plumbing particular attention
should be paid to the location and construction of a box privy with
receptacle or dug pit. No doubt many gastrointestinal disorders are
traceable to insanitary privies. Two important precautions are involved:
HOUSEFLIES 349
first, selection of a location which will avert pollution of wells or other
water supply; and second, choice of a type of construction which will
prevent flies from gaining access to the excreta and will insure privacy.
A sanitary privy (Fig. no) must meet the following requirements,
according to Stiles and Lumsden: 39
(1) The excreta must not touch the ground; hence some kind of water-
tight receptacle (box, pail, tub, barrel, tank, or vault) for the excreta must
be used under the seat. (2) Domesticated animals must not have access to
the night soil; therefore the privy should have a trapdoor in the back to ex-
clude them. (3) Flies and other insects must not have access to the excreta;
therefore the entire privy must be made rigidly flyproof, or some substance
must be used in the receptacle to protect the contents from insects.
Where the excreta are deposited in a pit or cesspool great care must
be exercised in banking up around the outside of the building so as to
prevent flies from gaining access to the pit. Borax applied to the excreta
as suggested for manure is an added precaution to kill fly larvae. Paradi-
chlorobenzene and orthodichlorobenzene are similarly useful. Also the
liberal use of a 5 per cent DDT-oil emulsion spray is urged for applica-
tion to all parts of the privy, inside and out, particularly the inside walls
of the box seat. DDT does not kill the fly larvae.
If the privy is built on skids or can be otherwise easily moved, in
addition to the treatment mentioned above, the accumulated excreta
should be burned from time to time by adding straw and crude oil and
setting them aflame. If sufficient water is available, the country home
should be equipped to receive the sewage.
Fly traps. Fly traps, properly baited, are useful in making fly surveys
for fly density data; however, other more valuable techniques to measure
active fly populations have been devised, such as "bait cards" and "fly
grills."40
BIBLIOGRAPHY
1. Hewitt, C. Gordon, 1910. The Housefly, Manchester, England: The
University Press, xiii + 195 pp.
2. Howard, L. 0., 1900. "A contribution to the study of the insect "fauna
of human excrement," Proc. Wash. Acad. Sc., 2:541-604.
3. Dunn, L. H., 1923. "Observations on the oviposition of the housefly,
Musca domestica, in Panama," Bull. Entomolog. Research, 13:301-5.
4. Herms, W. B., 1911. "The housefly in its relation to the public health."
Berkeley: Univ. Calif., in Agric. Exper. Sta. Bull., no. 215. pp. 513-48 (16
figs.).
5. - - , 1907. "An ecological and experimental study of Sarcophagidae,
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6. Hodge, C. F., 1911. In Nature and Culture, July, 1911.
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Carriers of Infection," in Reports to the Local Government Board on Public
Health and Medical Subjects, London, n.s. no. 53, Report no. 4.
8. Parker, R. R., 1916. "Dispersion of Musca domestica Linn. under city
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9. Austen, E. E., 1926. "The housefly: Its life history, etc.," London:
British Museum (Natural History) in Economic Series no. lA 2nd Ed., pp. 5-
52. .
10. Hall, David G., 1948. The Blowflies of North America. Thomas Say
Foundation of Entomolog. Soc. Amer. vol. 4, 477 pp.
11. Aldrich, J. M., 1916. Sarcophaga and Allies in North America. Thomas
S'lY Foundation of Entomolog. Soc. Amer. 302 pp. (16 plates).
12. Herms, W. B., 1911. The Photic Reactions of Sarcophagid Flies, etc.
Cambridge: Harvard Dniv. in Contributions from Zool. Lab., Museum Compo
Zool., No. 217.
13. Dorman, S. C.; Hale, W. C.; and Hoskins, W. M.; 1938. "The
laboratory rearing of flesh flies and the relations between temperature, diet, and
egg production," ]. Econ. Entomol., 31:44-51.
14. Herms, W. B., 1928. "The effect of different quantities of food during
the larval period on the sex ratio and size of Lucilia sericata Meigen and
Theobaldia incidens Thom.," ]. Econ. Entomol., 21:720-29.
15. Esten, W. N., and Mason, C. J., 1908. "Sources of Bacteria in Milk,"
Storrs Agric. Exper. Sta., Bull., no. 51, pp. 94-98.
16. Yao, H. Y.; Yuan, I. c.; and Huie, Dorothy; 1929. "The relation of
flies, beverages, and well water to gastrointestinal diseases in Peiping," Nat.
Med. ]. China, 15:410-18.
17. Graham-Smith, G. S., 1911. "Further observations on the ways in
which artificially infected flies carry and distribute pathogenic and other bac-
teria," in Reports to the Local Government Board on Public Health and
Medical Subjects, London (n.s., no. 53, further reports no. 4) pp. 31-48.
18. Nuttall, G. H. F., and Jepson, F. P., 1909. "The part played by Musca
domestica and allied (non-biting) flies in the spread of infective diseases,"
in Reports to the Local Government Board on Public Health and Medical
Subjects, London n.s., no. 16.
19. Leidy, Prof., 1871. "Flies as a means of communicating contagious
diseases," Proc. Acad. Nat. Sci. of Phila., p. 297.
20. Jordan, Edwin 0., 1908. A Textbook of General Bacteriology, Phila-
delphia: W. B. Saunders Co. 557 pp.
21. Faichnie, N., 1929. "The etiology of enteric fever: Personal views
and experiences," ]. M. A. S. Africa, 3:669-75.
22. Gill, C. A., and Lal, R. B., 1931. "Epidemiology of cholera, with
special reference to transmission: A preliminary report," Indian]. Med. Re-
search, 18:1255-97.
23. Paul, J. R., and Trask, J. D., 1941. "The virus of poliomyelitis in stools
and sewage," ]. A. M. A., 16:493-98.
24. Sabin, A. B., and Ward, R., 1941. "Flies as carriers of poliomyelitis
virus in urban epidemics," Science, 94:590-91.
HOUSEFLIES 351
25. ---, and ---, 1942. "Insects and epidemiology of poliomyelitis,"
Science, 95:300-301.
26. Nicoll, William, 1911. "On the part played by flies in the dispersal
of the eggs of parasitic worms," in Reports to the Local Government Board
on Public Health and Medical Subjects, London n.s. no. 53, further reports
(No.4) on flies as carriers of infection.
27. Roubaud, E., 1918. "Le role des mouches dans la dispersion des amibes
dysentriques et autre protozoaires intestinaux," Bull. Soc. path. exot., 11 :166-
71.
28. Root, F. M., 1921. "Experiments on the carriage of intestinal protozoa
of man by flies," Am. J. Hyg., 1:131-53.
29. Darling, S. T., 1912. "Murrina, a trypanosomal disease of horses in
Panama, and the means used in controlling an outbreak," Tr. 15th Internat.
Congo Hyg. & Demog., Washington, 5:619-31.
30. Ransom, B. H., 1913. The Life History of Habronema muscae (Carter),
a Parasite of the Horse Transmitted by the Housefly. Washington, D. C.: Dept.
Agric., in Bur. Animal Indust. Bull, no. 163. 36 pp.
31. Grassi, B., and Rovelli, G., 1889. "Embryologische Forschungen an
Cestoden," Centralbl. f. Bakt., 5:370-77, and 401-10. Cited by Gutberlet.
32. Gutberlet, J. E., 1916. "Morphology of adult and larval cestodes of
poultry," Tr. Am. Micr. Soc., 35:23-44 (2 plates).
33. Reid, W. M., and Ackert, J. E., 1937. "The cysticercoid of Choano-
taenia infundibulum (Bloch) and the housefly as its host," Tr. Am. Micr. Soc.,
56:99-104.
34. Dolve, R. M., 1912. Barn Plans. State College: North Dakota Agric.
ColI., Govt. Agric. Exper. Sta. of North Dakota, in Bull. no. 97. 57 pp.
35. Michelbacher, A. E., and Smith, Gordon L., 1946. "Control of flies in
dairy barns," Berkeley: Univ. Calif., in Agric. Exper. Sta. Circular, no. 365,
pp. 94-96.
36. Rowe, John A., 1948. "Fundamentals of community fly control," Proc.
and Papers, 16th Ann. Conf. Calif. Mosq. Control Assn. (Berkeley), pp. 11-14.
37. Douglas, James R., 1948. "Controllable noxious arthropods, other than
mosquitoes, affecting man and animals," Proc. and Papers 16th Ann. Conf.
Calif. Mosq. Control Assn. (Berkeley), pp. 14-16.
38. Lindquist, Arthur W., and Wilson, H. G., 1948. "Development of a
strain of houseflies resistant to DDT," Science, 107:276.
39. Stiles, C. W., and Lumsden, L. L., 1911. The Sanitary Privy. Washing-
ton, D. c.: Dept. Agric., in Farmers' Bull., no. 463. 32 pp.
40. Scudder, H. I., 1947. A new technique for sampling the density of
housefly populations. U. S. Public Health Service, Pub. Health Rep. 62:681-
86.
CHAPTER XVI
.~.
BLOODSUCKING MUSCOID FLIES

Tsetse Flies, Stomoxys Flies, Horn Flies
ORDER DIPTERA-FAMILY MUSCIDAE
A. TSETSE FLIES
Introduction. The genus Glossina, a member of the family Muscidae,
comprises the tsetse flies of Africa. The genus was established in 1830
by Wiedemann, and in the same year Robineau-Desvoidy described
Glossina palpalis. Bequaertl states that the word "tsetse" was introduced
into the English language by R. Gordon Cumming in 1850 in his "Five
Years of a Hunter's Life in the Far Interior of South Africa," and David
Livingstone in 1857 "focussed the attention of the scientific world upon
the ravages of the fly."
That tsetse flies enjoyed a wide distribution in geological times is
evidenced by the fact that several species of fossil Glossina flies from
the Miocene shales of Colorado, U.S.A., have been described. Today
the tsetse flies are restricted to continental Africa south of the Tropic
of Cancer, an area known as "the fly belt." Buxton 2 states that the tsetses
hold an area of about 4~ million square miles and are an immense
obstacle to the development of tropical Africa.
Adult tsetse flies (Fig. Ill), both male and female, depend on blood
for their existence, and while they feed on a wide variety of animals,
there are host preferences among the different species. Although man is
freely attacked, he is not considered a favored host. In areas where large
wild mammals have been decimated, the flies tum freely to man. Gloss-
ina palpalis (R.-D.) favors reptiles, particularly crocodiles and monitor
lizards.
Students concerned with tsetse flies will consult the publications of
2
Buxton, Austen and Hegh 3 (1922), Newstead 4 (1924), Hegh 5 (1929)
6
Swynnerton (1936), as well as those other authors which cover particu-
lar segments of the subject, e.g., the work of Bequaert (1946)7 which
deals with the tsetse flies of Liberia.
General characteristics. The tsetses are medium-sized flies, ranging in
352
BLOODSUCKING MUSCOID FLIES 353
size from that of a housefly to that of a blowfly. They are brownish in
color; the body is wasp-like, and the wings when at rest are crossed
scissors-like and extend well beyond the tip of the abdomen. The wing
venation is characteristic in that the fourth longitudinal vein (MH 2)
bends suddenly upward before it meets the anterior transverse vein,
which is very oblique (Fig. Ill). The dis cal cell is remarkably "cleaver-
like" in outline.
The palpi are nearly as long as the proboscis, which points bayonet-
like in front of the head. The antenna 1 arista (Fig. 112) bears a series of
long bilaterally branched and regularly arranged hairs only on the
upper surface. Griinberg 8 attached taxonomic value to the aristal hairs.
FIG. Ill. (Left) Glossina palpalis, (right) Glossina morsitans.

(After N ewsstead. )
The mouth parts consist of the labium which ensheathes the two slender
piercing stylets, the labrum-epipharynx, and the hypopharynx. A charac-
teristic "onion-shaped" bulb is situated at the base of the haustellum
(Fig. 113). Both sexes are avid bloodsuckers, feeding usually in broad
daylight and outdoors. They are particularly attracted to moving ob-
jects.
Life history. The female tsetse fly gives birth to full-grown larvae
which are extruded singly at intervals of about 10 to 12 days during
the lifetime of the mother. During the intrauterine state (there are
three larval stages) the larvae feed on fluid from special glands com-
monly referred to as "milk glands." The newly extruded larvae are
creamy white to pale yellow and have a pair of intensely black, shining
lobes at the posterior extremity. After each larviposition at least one full
meal of blood is needed before the next larva begins to develop, eight
to ten being produced during the life of the female.
The larvae are unable to crawl as do other muscoid larvae because
of the reduced cephalopharyngeal armature. Lewis" paints out that the
larva moves and burrows by peristaltic movements and longitudinal
contractions of the whole body. Coarse pebbly sand favors the larvae
in burrowing, although a depth of only a few centimeters is reached .
The behavior of the larva at this time, Lewis points out, is of great
importance in determining its chances of survival. "If it is slow to pene-
trat e the soil, it is exposed for a longer period to the possible attack of
predators and parasites." Pupation takes place within an hour of larvi-
112 Ilg
FIG. 112. Antenna of a Glossina fly, shOWing arista with branched hairs.
(Much enlarged.)
FIG. 113. Mouth parts of a Glossina fly. X 17.
position. The pupa rapidly darkens to a blackish brown color with the
posterior lobes and general form as shown in Figure 114. The pupal
stage lasts from three to four weeks and longer, depending much on soil
temperature and soil moisture . A great deal of work has been done on
the ecology of the puparium (Nash'O).
The fly emerges from the puparium by breaking loose the end of the
pupal case by pressure from the ptilinum. The flies are said to have a
striking dislike for excrementous matter, and the . larvae are ordinarily
depOSited in the root tangles of the mangrove and shade of other vegeta-
tion where the soil is not too dry and is loose. The presence or absence
of organic matter seems to be immaterial (Nash).
Trypanosomiasis. The tsetse flies are important vectors of trypano-
somes. The term trypanosomiasis applies to all infections with flagellate
protozoan parasites of the genus Trypanosoma and includes African
sleeping sickness. The Trypanosoma belong to the class Mastigophora,
order Protomonadida, family Trypanosomidae; they invade the blood,
lymph, cerebrospinal fluid, and various organs of the body, such as the
liver and spleen of many species of vertebrate animals, from fish to man.
Many species of trypanosomes are regarded as nonpathogenic. Nearly
all of the Tryanosorna require an intermediate insectan host.
The first trypanosome was discovered by Valentin in 1841 in the
blood of the salmon. The name Trypanosoma l ! was given to these organ-
isms by Gruby in 1842-43. Attention was not called to trypanosomes of
mammals until the work of Lewis in 1878 on the parasites [Trypanosoma
lewisi (Kent) J of the blood of the rat in India." After that followed the
FIG. 114. Pupae of the Glossina fly. x4.8.
discovery of other more important pathogenic trypanosomes, e.g., in

1880 Evans discovered the trypanosome, Trypanosoma evansi (Steel),
of surra; in 1895 Bruce discovered T. brucei Plimm. and Brad£., the
causal organism of nagana, the tsetse-fly disease of cattle and horses
of Africa, and in 1897 he ' 2 showed that Glossina morsitans Westwood
transmitted the infection. In 1901 Dutton' :!,l.! discovered trypanosomes
in a blood smear prepared by Dr. R. M. Forde from a European patient
in Gambia; he also found them in the blood of a native child. Following
this discovery the organism was named Trypanosoma gambiense Dut-
ton. In 190,'3 Bruce and Nabarro '5 proved Glossina palpalis (R.-O.) to
be the carrier by feeding freshly caught flies ranging in number from
31 to 9 on a black-faced monkey daily, sometimes twice daily, beginning
May 13. Trypanosomes were present in the monkey's blood on May 27.
In 1903 Castellani ' 6 reported trypanosomes in the cerebrospinal fluid
of negroes in Uganda suffering from sleeping sickness. The trypanosomes
o Trypanosoma lewisi is transmitted from rat to rat by the rat flea, NosopsylltlS
fasciafus (Bose) , and the rat louse, Polyplax spinulosus (Burm.).
found by Castellani were supposed to be a different species from that of
Dutton (Trypanosoma gambiense Dutton) and were called T. ugan-
dense Castellani, 1903. Kruse later gave to this trypanosome the name
T. castellanii. The important discoveries of Dutton and Castellani were
quickly confirmed by David Bruce, who found these trypanosomes 38
times out of 38 in the cerbrospinal Huid obtained by lumbar puncture
in natives of Uganda suffering from sleeping sickness, and 12 times out
of 13 in the blood. According to the rules of priority, the last two species
names mentioned must give way to Trypanosoma gambiense Dutton. A
second species of trypanosome producing sleeping sickness in Rhodesia,
Nyasaland, and adjoining territory was described in 1913 by Stephens
and Fantham (loc. cit., p. 5) as T. rhodesiense.
Sleeping sickness is widely distributed in Africa, extending along the
west coast from Senegal (in part) to Angola (in part) and eastward to
the valley of the upper Nile. It has been estimated that between 1896
and 1906 from 400,000 to 500,000 natives perished from this pestilence.
Dutton and Todd found that in some villages from 30 per cent to 50 per
cent of the population was infected. In many parts of Africa the num-
ber of cases remains high; on the other hand as Buxton (1948) points
out there are a number of areas in East and West Africa where there
are healthy people and cattle living permanently on ground from which
tsetse Hies have been exterminated.
Age does not affect the distribution of the malady, since children
as young as 18 months to 2 years have been known to be infected. Sex
does not inHuence the disease. Occupation and social position, however,
do show a marked inHuence. The majority of the cases observed are
among the agricultural and lower classes, where the degree of exposure
to tsetse-Hy bites is greatest. In areas where Glossina palpalis occurs,
inland fishermen in particular may be exposed to the bites of this species.
The bite of an infected tsetse appears to produce more local reaction
than that of the uninfected fly. The usual duration of the incubation
period is ten days to three weeks, but clinical symptoms may be delayed
for as long as two to five years. During the first phase of the disease,
which may continue for many months, the trypanosomes are in the
blood, the trypanosomiasis stage; this phase is characterized by an irre-
gular fever, glandular enlargement, debility, and languor. In the second
phase, the sleeping sickness stage, the trypanosomes are constantly found
in the cerebrospinal fluid; a constant accompaniment is the enlargement
of the posterior cervical lymph nodes, Winterbottom's sign; there is
speech impairment and tremors of the tongue; there is nervousness,
pronounced languor and drowsiness, which gives way to lethargy, and
finally the victim falls into a comatose state, wasting rapidly, due largely
to starvation, until death ensues. Although authors commonly refer to
Gambian sleeping sickness as caused by Trypanosoma gambiense and

the Rhodesian sleeping sickness as caused by Trypanosoma rhodesiense
there is as yet no uniformity of opinion in the matter; there may be but
one causal species.
Transmission. The natives of French Guinea long attributed the
transmission of sleeping sickness to flies; it had already been shown by
Bruce that nagana, a disease of cattle and horses, was transmitted by
tsetse flies, when Dutton and Todd gave attention to the biting flies of
Gambia. These investigators found that of the flies which bite both man
and animals, Tabanus dorsovittata Walker and Glossina palpalis (R.-D.),
were the most important. The latter is very common in western Africa,
where it abounds in the mangroves which line the inland rivers during
the warmer months when these insects are very troublesome.
Experiments made by these workers, however, gave negative results.
It was Bruce and his collaborators (loc. cit.) who subsequently went
over the matter and showed that Glossina palpalis (R.-D.) is a vector.
Animal experimentation showed that these flies can transmit the trypano-
somes mechanically for a period of less than 48 hours; the organisms
become more and more attenuated after the fly has bitten the infected
individual, and it loses the power of infection in less than 48 hours. Thus
the tsetse fly may be a mechanical vector for a few hours, during which
time its soiled proboscis is involved, i.e., trypanosomes are introduced
into the wound produced by the bite before the proboscis is completely
cleaned. Interrupted feeding would thus be a factor. Mechanical trans-
mission from man to man in nature is believed to be very uncommon,
if it occurs at all.
KleineH in 1909, working with Trypanosoma brucei, causal organism
of nagana, was evidently the first to demonstrate its development in the
tsetse fly, Glossina palpalis. Robertson 18 working with the causal organ-
ism of sleeping sickness, Trypanosoma gambiense, reported in 1913 that
it is first established in the posterior part of the mid-gut of the insect,
where multiplication occurs and trypanosomes of varying sizes are pro-
duced. From the tenth or twelfth day onward slender, long forms are
to be found in increasing numbers. These finally move forward to the
proventriculus and are the dominant type: The proventriculus becomes
infected as a rule between the twelfth and twentieth days. The salivary
glands become infected by the slender (proventricular ) type which
reaches the glands by way of the hypopharynx; arriving in the glands,
they become attached to the wall and assume the crithidial condition.
Multiplication takes place, and trypanosomes are formed which closely
resemble the blood type. The development in the salivary glands takes
from two to five days before the metacyclic forms are infective. The
fly is never infective until the glands are invaded. The trypanosomes
are never attached to the wall of the alimentary canal, and there is no
intracellular multiplication in the gut cycle. The metacyclic forms be-
come mature and multiply by longitudinal fission in the circulatory
system of the mammalian host. The Sleeping Sickness Commission found
that infectivity lasted at least 96 days. Only a small percentage of the
tsetse Hies become infective. The life of a female G. palpalis (R.-D.) in
captivity has been observed to be about four and one-half months.
The problem of sleeping sickness is greatly complicated in that many
species of game animals harbor the causal trypanosomes and may thus
serve as natural reservoirs of the infection. Furthermore the affinities
of Trypanosoma rhodesiense have not yet been definitely determined.
There are some who contend that this is a varient of T. gambiense;
others contend that it is a human strain of T. brucei and considerably
less adapted biologically to man and his domestic animals than to the
wild game. This might account for the greater virulence of this species
in relation to man. Glossina morsitans and G. swynnertoni are vectors.
The principal vectors of African sleeping sickness, each in its respec-
tive zone, are Glossina palpalis (R.-D.), G. morsitans Westwood, and
G. swynnertoni Austen.
Nagana. Trypanosoma brucei Plimm. and Bradf. is the causal or-
ganism of nagana, long known to be a fatal tsetse-Hy disease of African
horses, mules, and camels, less rapidly fatal to cattle, sheep, and dogs.
Many other mammals are susceptible to the infection. Bruce (loc. cit.)
found that many species of wild game animals harbor the trypanosome
and thus form reservoirs. The disease is characterized by progressive
emaciation, fever, edema of the abdomen and genitalia, and marked
depression. The trypanosomes are found in the blood and especially
the lymph-gland swellings from the beginning of the first symptoms.
Glossina morsitans West., G. longipalpis (Wied.), G. pallidipes Aus-
ten, G. tachinoides West., and G. austeni Newstead relate to its trans-
mission in practically the same way that Glossina palpalis (R.-D.) and
other Glossina Hies relate to sleeping sickness of man, i.e., the Hies are
infective for a day or two after feeding on an infected animal, then
become noninfective for a period of about three weeks, when they
again become infective and remain so for the rest of life. The incuba-
tion period after inoculation into the body of the host is said to be
about 10 days.
Glossina species. N ewstead (loc. cit.) recognized twenty species, one
subspecies, and five varieties belonging to the genus Glossina. These
he divides into three groups, (1) the fusca group, which includes the
10 largest species, viz., Glossina brevipalpis Newstead, G. fusca (Walk-
er), G. fusca var. congolensis N ewst. and Evans, G. fuscipleuris Aust.,
G. haningtoni Newst. and Evans, G. longipennis Corti, G. medicorum
Aust., G. nigrofusca .Newst., G. schwetzi Newst. and Evans, G. severini
Newst., and G. tabaniformis West.; (2) the palpalis group, which in-
cludes Glossina caliginea Aust., G. pallicera Bigot, G. palpalis (Rob.-
Desv.) and two varieties, var. wellmani Austen and var. maculata Newst.,
also one subspecies G. palpalis subspecies fuscipes Newst., and G.
tachinoides West.; (3) the morsitans group, which comprises Glossina
longipalpis (Wied.), G. morsitans West., G. morsitans var. pallida
Shirc., G. morsitans var. paradoxa Shirc., G. submorsitans Newst., G.
pallidipes Aust., G. swynnertoni Aust., and G. austeni Newst.
Glossina palpalis (R.-D.) (Fig. Ill) is the most important vector of
Gambian. sleeping sickness. It covers an enormous area in Africa, but it
occurs chiefly in the Congo and in west Africa. It is usually found on
the shores of rivers and lakes, but it may occur quite far back from
them; and as Swynnerton points out, it requires a combination of several
types of country one of which must be relatively massive wooding or
thicket of more or less evergreen type (see BequaerC). It feeds mainly
on large reptiles such as crocodiles and monitor lizards, but it can live
on the blood of mammals as well. Although man is not regarded as one
of its favored hosts, it will nevertheless feed freely on persons available
in its riparian habitat.
Glossina morsitans Westwood (Fig. 111) is a most efficient vector of
both Rhodesian sleeping sickness of man and nagana of animals. It has
a wide distribution in Africa; it is of importance in the Sudan, northern
and southern Rhodesia, the Belgian Congo, and many other localities.
This species requires "savanna of sufficient shade value, and with suffi-
cient logs, rocks, or tree rot holes to form a good rest-haunt and breed-
ing-ground, and relatively open glades or plains in which to hunt for
its prey." It is typically a "game fly," but attacks human beings readily,
hence is a most dangerous tsetse. .
Glossina swynnertoni Austen, like G. morsitans West., is a strong
vector of both Rhodesian sleeping sickness and nagana. It is largely
confined to the northern part of Tanganyika, according to Swynnerton,
who describes it as "the fly of the driest and most open areas and ap-
parently unable to inhabit the more mesophytic savannas. It breeds
normally in thicket, though rock suits it as well. . . . It utilizes open
spaces as feeding grounds . . . . It is primarily and essentially a 'game'
fly." It attacks human beings with readiness, and like Glossina morsitans
West. is a very dangerous tsetse.
Tsetse-fly control. Since the memorable discoveries of Bruce and
others that tsetse flies are responsible for the transmission of nagana
and sleeping sickness, few insects have been so minutely studied by the
most capable investigators. The practical and extended control of breed-
ing places offers serious difficulties, not the least of these being the fact
that the larvae are retained within the body of the female until fully
grown, hence are not directly dependent upon an external food supply.
The monumental work of the late C. F. M. Swynnerton gives ample
testimony to the tremendous ramifications of the tsetse-fly control prob-
lem. Among the many possible modes of attack there are the following:
(a) direct attack, involving the use of Rytraps; the direct effect of tem-
perature and moisture on pupae; the use of natural enemies, etc.; (b)
indirect attack by modification of cover, redUCing or expelling game
animals, thus depriving the flies of food supply; introduction of fly bar-
riers by setting up clearings or thickets according to the species of fly
to be dealt with; reclamation and appropriate agricultural practice. Be-
cause of the wide divergence in the ecological requirements of the
several important species of tsetse flies, the utilization of appropriate
control measures is a matter of long and tedious investigation.
Buxton (1948, loco cit.) states that it is not difficult to exterminate
Glossina palpalis from a linear fly belt along water; it is done by clear-
ing or hand catching. In large areas of bush, such as those inhabited
by Glossina morsitans, satisfactory results have been achieved by bush
clearing, by attacking the restricted permanent habitats, and also by
game destruction and human settlement. Buxton also suggests that the
use of DDT "smoke" gives some promise.
B. STOMOXYS FLIES
Family Muscidae, Genus Stomoxys
General characteristics. Owing to similarity in color and size the
stomoxys fly is often mistaken for the common housefly Musca domestica
Linn. However, the former is more robust and has a broader abdomen .
. In color it is brownish gray with a greenish-yellow sheen; the outer of
the four longitudinal thoracic stripes are broken, and the abdomen is
more or less checkered. The wings when at rest are widely spread apart
at the tips, and are distinctly iridescent; the apical cell is open. When
resting the fly has its head thrown well up and the wings slope decidedly
toward the surface upon which it has settled. The probOSCis protrudes
bayonet-like in front of the head. The antennal aristae, unlike those of
the housefly, bear setae on the upper side only. Stomoxys calcitrans
(Linn.) enjoys practically a world-wide distribution.
Habits. Although the stomoxys fly, Stomoxys calcitrans (Linn.), is
commonly called the "stable fly," it occurs much less abundantly (often
absent) about stables than does the housefly. It is also called the "biting
housefly," since it may occur indoors, especially in the autumn and
during rainy weather, and bites human beings Viciously. It is often very
annoying along the sandy vegetation-strewn shores of lakes. Recently
the name "dog fly" has become attached to this species. The stomoxys
fly is typically an out-of-door day-biting fly and is usually to be found
in abundance during summer and autumn \\There large numbers of
domesticated animals occur, horses and cattle affording an abundant
food supply. This species does not breed in eXCrement as freely as does
the housefly. Sunny fences, walls, light-colored canvas coverings, and
FIG. US. Feed racks for dairy cattle aHord ali ideal breeding place for
stomoxys flies in that the moist lower layers of material in the trough furnish
abundant food for the larvae.
light objects in general, when in the proximity of: animals, are abundantly
frequented by stomoxys flies.
The stomoxys fly is a vicious "biter" which (haws blood quickly and
fills up to full capacity in from three to four mi~1Utes if undisturbed; but
ordinarily even when undisturbed it changes pC)sition frequently or flies
to another animal, where the meal is continued. This fly feeds readily
on many species of warm-blooded animals, fC)r example, rats, guinea
pigs, rabbits, monkeys, cattle, horses, and marl. Both sexes are blood-
sucking. The flight of the stomoxys fly is direct and swift. It may travel
many miles.
Breeding habits and life history. Although the stomoxys fly can be suc-
cessfully reared in the manures of horses, cattle, sheep, etc. , it may be
safely said that it does not breed commonly in excrement uuder field
conditions unless this is well mixed with straw or hay. Very good breed-
ing places are afforded by the leftover soggy hay, alfalfa, or grain in the
bottoms of or underneath out-of-door feed racks (Fig. 115) in connec-
tion with dairies, feed lots, etc. This material becomes soggy and fer-
ments, and here practically pure cultures of stomoxys larvae may be
found. The material must be moist; dryness prevents larval development.
Piles of moist fermenting weeds and lawn cuttings also furnish fairly
good breeding material. Piles of decaying onions have been found by
the writer to harbor myriads of larvae late in the autumn. Old straw
piles that remain in the field through the year may produce an abun-
dance of stable flies in the moist fermenting straw near the ground,
Pupa Adult
:\ ot in sa me scale
FIG. 116. Life cycle of the stable fly, Stomoxys calcitrans. x2. (Photograph by
H. F. Gray.)
particularly if cattle have access to it and moisten it with urine. Moist
piles of fermenting peanut litter produce large numbers of these flies, as
do beach deposits of grasses along the Florida coast. The author has
also noted this along the southern shores of Lake Erie.
The larvae of stomoxys and of the housefly can readily be differenti-
ated by the form, size, and position of the posterior spiracles; otherwise
they resemble each other closely. The pair of posterior spiracles of the
stomoxys larva are roughly triangular, widely separated, and situated
near the periphery, while in the housefly larva they are elliptical, quite
large, close together, and more central in position (Fig. 119).
The eggs (Fig. 116) of the stomoxys fly are about 1 mm long, curved
on one side, and straight and grooved on the opposite side. In depositing
her eggs the female fly often crawls far into the loose material, placing
them usually in little pockets in small numbers, often in pairs. Egg
depositions range in number from 23 to 102, usually between 25 and
50, and there are ordinarily four or five layings. Mayne (Mitzmain19)
found in his observations made in the Philippine Islands that the maxi-
mum number of eggs produced by a single Stomoxys is 632 and possibly
820 and that there may be as many as 20 depositions during the lifetime
of the female.
The incubation period varies from two to five days, commonly three
days, at a temperature of 26° C. Higher temperatures result in a shorter
incubation period. The newly hatched larvae bury themselves in their
food at once, thus protecting themselves against light and dryness. At
a temperature of from 21° to 26° C, the larvae reach full growth in
from 14 to 26 days. Mayne found that the larval stage averaged 12 days
at a room temperature of 30° to 31°C.
Before pupation the larvae usually crawl into the drier parts of the
breeding material, where the chestnut-colored pupae are often found in
enormous numbers. The pupae are from 6 to 7 mm long and may be
recognized by the posterior spiracles as in the larva. The pupal period
also varies, dependent largely on temperature. At a temperature of from
21 ° to 26° C, this period varies from 6 to 26 days, with the greatest fre-
quency between 9 days and 13 days.
At an average temperature of 29° C, Mayne found the pupal period
to average five days.
If not handicapped, the imago emerges with astonishing rapidity,
crawls away, unfolds its wings, and is ready to flyaway in less than
half an hour. The fact that the proboscis is temporarily attached be-
neath the thorax gives the newly emerged insect a very peculiar appear-
ance, and it may then be easily mistaken for a housefly.
Summarizing the life history of the stomoxys fly it may be said that
at a temperature of 21° to 26° C, the shortest periods are: egg, 2 days,
larva, 14 days, pupa, 6 days, total, 22 days; the average, egg, 3 days,
larva, 15 days, pupa, 10 days, total, 28 clays; and the maximum, egg, 5
days, larva, 26 days, pupa, 26 days, total, 57 days. The total time at
21 ° C, from the laying of the eggs to the emergence of the adults, was
from 33 days to 36 days, as observed in five individual cases. Mayne
reports the development of this fly in 12 days under optimum condi-
tions.
Copulation takes place within a week, and egg deposition begins in
about 18 days, after emergence from the pupa cases at a temperature
of from 21° to 26° C. Higher temperatures undoubtedly decrease this
time. In warmer parts of the world the stomoxys fly continues to breed
throughout the year, slowing up the cycle during the colder months.
Longevity. With approximately 4,000 flies under continuous daily
observation in glass quart jars, 50 flies to a set, the writer has found
that the average length of life of the stomoxys fly under favorable labora-
tory conditions of feeding (i.e., daily feedings on monkeys or rabbits)
is about 20 days. The maximum life under these conditions was found
to be 69 days and several hours in a female.
Mayne has found the maximum for a female stomoxys fly to be 72
days and for the male 94 days.
The writer has observed that a set of flies which fed only on sugar
water deposited no eggs, although many of them lived 20 days or longer,
while control flies fed on blood did lay eggs. Hence it seems apparent
that the flies must have blood in order to develop eggs.
As a cattle pest. Bishopp20 regards this fly as one of the most impor-
tant sources of annoyance to livestock. Injury is brought about in various
ways, e.g., worry due to the attacks of myriads of flies, loss of blood,
and loss of flesh.
Freeborn, Regan, and Folger 21 have shown that the reduction in milk
production caused by the stable fly amounted to 9.26 per cent, which for
a five months' period means a loss of 50 gallons of milk, amounting to
$10 per cow per season. The total loss occasioned by the three dairy
cattle pests-houseflies, stable flies, and horseflies-amounted to 14 per
cent.
Poliomyelitis. The following is quoted from a report by Rosenau on
poliomyelitis, presented at the 1912 meeting of the International Con-
gress on Hygiene and Demography.22
When I first began to study the disease [poliomyelitis], I regarded it prob-

ably as one which is spread by direct contagion, by contact, either directly or
indirectly, from person to person. The first circumstance which shook my faith
that we were dealing with a contagious disease was the fact that we had
eighteen negative results in attempting to prove the presence of the virus in
the secretions from the nose and throat. I could not help asking at the time
if it were not possible to find the virus, which is so potent, in the secretions of
the nose and throat of persons who have the disease and those who are con-
valescing from the disease. These results were confirmed at the same time
by Strauss, of New York, who had negative results in a large series and by
Neustaedter's recent results and by other results, all of the examinations
having proven negative excepting one recently reported by Kline, Patterson
and his associates at this congress and in the literature recently.
A second circumstance which led me to believe we were not dealing with
a contagious disease was the fact brought out by Dr. Richardson. Children in
all stages of this disease were crowded into schools, institutions, tenement dis-
tricts and other places where there was every chance for the spread of the dis-
ease, but it did not spread there, but it continued to spread in the rural, thinly
scattered districts where one would not expect to find contagious disease. There
was a resemblance to rabies. All those who have worked with this virus in
laboratories were at once struck with the resemblance between poliomyelitis
and rabies. The latter being a wound infection, there is some analogy between
it and poliomyelitis, and poliomyelitis might be transmitted through some sort
of wound. I was fortunate enough to have had experience with yellow fever,
both in the investigation of it and the sanitary measures against it, before the
mosquito period, and I was much struck with many analogies which came to
me between that disease and certain features of poliomyelitis.
The work I bring to your attention consisted of taking a number of flies
-Stomoxys calcitrans-caught in a net and bred for the purpose; you can catch
several hundred of these flies in a stable in a very short time. We placed these
flies in a large cage and exposed monkeys to their bites, the monkeys having
been purposely infected with the virus of poliomyelitis. Care was taken to
place the monkeys in the cages in all stages of the disease, before and after. In
fact, a monkey would be exposed to the bites of the flies on the same day he
was infected, so that the flies could drink the blood of the monkey during all
stages of the period of incubation of the disease, for we do not yet know in
what stage of the infection the virus appears in the blood at its maximum, or
the best period for infecting these flies. Following this we exposed healthy
monkeys to the bites of the same flies, and after several weeks' time these
healthy monkeys came down with a disease which in all essential respects re-
sembles anterior poliomyelitis. Out of twelve healthy monkeys so exposed, six
of them now have symptoms of the disease, three of them in the virulent form.
Of the other three monkeys, two are coming down, but one seems to have a
milder infection than the other. This mild infection consists of trembling and
weakness of the hand, and some weakness of the jaw which lasted about a
week or so and then passed away. We cannot be sure whether that is true
poliomyelitis or not until we are able to test the monkey subsequently. If it
were poliomyelitis, that monkey would be "immune." In three of the six cases
that came down with the disease, having been bitten by flies, there was some
diarrhea. The disease in the monkey resembles more closely that which we see
in children, rather than the disease we produce purposely experimentally by
bringing the virus in direct association with the central nervous system. Of
course, that may be only a coincidence, but it is interesting.
The work of Rosenau was repeated and confirmed during October,

1912, by Anderson and Frose 3 ,24 who summarize as follows: "Three
monkeys exposed daily to the bite of several hundred stomoxys, which
at the same time were allowed daily to bite two intra cerebrally inocu-
lated monkeys, developed quite typical symptoms of poliomyelitis eight,
seven and nine days from the date of their first exposure."
In order to test the findings of the above-mentioned investigators and
to secure further biological evidence if possible, the author in coopera-
tion with W. A. Sawyer,25 undertook a special investigation of the prob-
lem, beginning in October, 1912. Believing it unwise to use flies col-
lected out of doors, these insects were reared for the purpose in an in-
sectary. The importance of this precaution is m.ade evident by the fact
that Hies captured out of doors in Berkeley were shown to transmit a
pathogenic organism to a rabbit, infection undoubtedly having been
acquired in nature. This infection, resulting in abscess, was successfully
transmitted from rabbit to rabbit through the agency of the stomoxys
fly.
Seven carefully planned experiments were conducted over a period
of nearly a year; about 4,000 laboratory-reared flies, a large number of
monkeys, rabbits, and other experimental animals were used; the results
were entirely negative.
Control of stable flies. The more important breeding places of the
stomoxys can be controlled by removing moist feed wastes from feeding
troughs and from feed lots, stalls, stables, etc., and scattering the wastes
to hasten drying. Moisture is necessary for the development of the larvae;
therefore dry material is not suitable. Weeds, lawn cuttings, decaying
onions, vegetation washed up on lake shores in the immediate vicinity
of summer resorts, etc., should not be allowed to accumulate in piles
long enough to ferment and decay.
Bishopp (lac. cit.) has shown that loosely piled straw stacks (oats
and wheat) are important breeding places of the stomoxys fly, hence
he recommends "that the straw for feeding and bedding purposes be
baled and stored under cover. Where this is not practicable the stacks
should be rounded up so as to make the top largely rain proof and the
sides nearly vertical." DDT as recommended for housefly control can
also be used successfully in the control of stable flies.
C. THE HORN FLY
Family Muscidae
Introduction. The so-called horn fly (also called Texas fly), which
torments cattle and horses (the latter to a less extent) in many parts of
North America is classified by some entomologists as Haematobia stimu-
lans Meigen (= irritans R.-D.) and by others as Lyperosia irritans Linn.
[= Haematobia serrata (R.-D.)]. In his description of the habits of
these flies in Europe, Hammer 26 states that Haematobia stimulans extends
farther north in Scandinavia and Russia than does Lyperosia irritans, the
latter occurring only in the southern portions of this area. Hammer re-
marks that L. irritans has adapted itself to milder climates with higher
temperatures. He also describes this species as attaching very closely to
cattle, only leaving the animals to oviposit; it spends both day and night
on the cows whether they are in the byre or in the field. H aematobia
stimulans leaves the animals at night and hides away among vegetation.
The horn fly with which we have dealt in California, where it is a severe
cattle pest, sticks close' to the cattle at all times; it also deposits large
reddish-brown (or yellowish-brown) eggs, while Hammer states that
Haematobia stimulans deposits white eggs. This insect is known as the
horn fly because it rests on the horns of cattle much of the time, particu-
larly at night, probably because they afford a safe resting place.
The horn fly was introduced into the United States from Europe,
where it has been an important cattle pest for many years. (See Ham-
mer, Zoc. cit.) According to the United States Bureau of Entomology and
Plant Quarantine it was first reported in the fall of 1887 from Camd~n,
N. J., appearing during the following year in Maryland and Virginia,
probably having appeared in Philadelphia in 1886, and by 1892 was
found over the entire continent from Canada to Texas and from Massa-
chusetts to the Rocky Mountains. California cattlemen state that it made
its appearance in this state in about 1893-1894. It appeared in Honolulu,
Hawaii, in 1897.
Characteristics. The horn fly is about half the size of the common
housefly, i.e., about 4 mm long. It has much the same color and in most
FIG. 117. Side view of head of the horn fly.
other respects resembles the stomoxys fly. The mouth parts (Fig. 117)
are like those of stomoxys except that the labium of the horn fly is rela-
tively heavier and the palpi, almost as long as the proboscis, are flattened
and loosely ensheath this structure. The arista is plumose dorsally. The
wing venation is as in stomoxys.
Horn flies appear early in the spring and become most abundant in
late summer and autumn. When the horn fly is at rest on an animal .or
elsewhere its wings lie flat on its back and fold rather closely, but
when it bites, the wings are spread and the insect stands almost per-
pendicularly, hidden between the hairs of the host. Apparently the
habit of resting at the base of the horns is only done when flies are
overabundant.
Life history. The horn fly deposits its eggs chiefly, if not exclusively,
on freshly passed cow manure. The fly is seen to dart from the animal
and deposits its eggs in groups of four to seven, or singly, on the surface
of the dung. The eggs are relatively large (1.3 to 1.5 mm), larger than
the eggs of stomoxys; they are reddish brown in color, hence not easily
seen on the cow dung. Under laboratory conditions, at least, few eggs
are deposited by the females, rarely over twenty. At a temperature of
24° to 26° C, the eggs hatch in 24 hours.
The larvae burrow beneath the surface of the droppings, reaching
full growth in from three to five days, when they crawl underneath into
drier parts and pupate. The pupal period requires from six to eight days.
Hence the entire life history (Fig. 118) from the egg to the adult
requires from 10 to 14 days at a temperature of from 24° to 26° C.
Damage done. The damage occasioned by the horn fly is chiefly
through irritation and annoyance, which in dairy animals results in dis-
turbed feeding and improper digestion causing loss of flesh and reduced
milk production. Dr. James Fletcher estimated the loss in Ontario and
~ --~-~ -~ ----
-_.-:
,
FIG. 118. Life cycle of the "horn fly," Haenwtobia serrata.
Quebec, Canada, at one-half of the product of meat and milk. Range

animals literally run themselves thin in trying to get away from these
pests.
The actual loss of blood must be considerable when literally thou-
sands of these flies attack an animal. From 10 to 25 minutes are required
for the fly to fully engorge itself; during this time it withdraws and re-
inserts its proboscis in the same puncture many times as in a pumping
motion. Much undigested blood is discharged from the anus of the fly
while in the act of feeding.
Control. The most effective method to prevent the multiplication of
the horn fly is to scatter the droppings from cattle with a rake or other
implements or Simply by dragging a branch of a tree over the field. Hogs
allowed to run with the cattle serve this purpose very well. The manure
thus scattered dries out quickly, and the larvae that are present perish,
owing to the fact that they require much moisture for development. The
writer has seen this method applied successfully in various parts of Cali-
fornia where the dry summer favors such means of handling the situa-
tion. On wide ranges this method is impracticable, but in connection with
dairies it is entirely feasible. The use of DDT cattle sprays, as already
described, gives good control on the animals.
Lyperosia exigua (de Meij.), commonly known as the "buffalo fly,"
is particularly important to the cattle and dairy industries of Australia.27
Among the animals which it attacks are buffalo, cattle, horse, dog,
and man. The fly oviposits in fresh dung from buffalo and cattle in
particular.
Other species of bloodsucking muscoid flies. The genus Philaema-
tomyia represented by the single species, P. insignis Austen, is of particu-
lar interest because of the form of the probOSCis, which is intermediate
between the biting and nonbiting muscid type. P. insignis Austen is a
widely distributed African and Oriental species resembling Musca domes-
tica Linn. in size and general appearance. According to Austen 28 the
proximal portion of the proboscis is a strongly swollen, polished, chitinous
bulb, while the distal portion is soft and fleshy and folded back under
the distal end of the bulb when not in use; when in use its terminal
section, consisting of a tubular extension, is protruded from between the
labella; it is surrounded at the distal extremity with a circlet of stout
chitinous teeth. When it is not being used the entire proboscis can be
retracted within the buccal cavity. Austen states that the fly probably
feeds by cutting through the epidermis with the teeth at the end of the
tubular extension and then sucks up the blood.
The Ethiopian genus Stygeromyia (S. maculosa Austen, and S.
sanguinaria Austen) is said by Austen (loc. cit.) to be in some respects
intermediate between Stomoxys and Lyperosia. It resembles Stomoxys
in general appearance and form of the body but is distinguished "by the
relative stoutness of the short, chitinous, horizontal proboscis, and by the
palpi being equal to the proboscis in length, large, expanded towards the
tips, and curved upwards."
Stomoxys nigra Macquart, S. omega Newstead, and S. inornata Grun-
berg are all Ethiopian species and resemble S. calcitrans in feeding habits.
BIBLIOGRAPHY
1. Bequaert, J., 1930. "Tsetse Hies-past and present (Diptera: Mus-

coidea)." Entomological News, 41 :158-64, 202-3, and 227-33.
2. Buxton, P. A., 1948. "The problem of tsetse Hies," Proc. 4th Internat.
Congo Trop. Med. and Malaria, Washington, D. C. (Abstracts).
3. Austen, E. E., and Hegh, E., 1922. Tsetse-flies, Their Characteristics,
Distribution, and Bionomics. London: Imperial Bur. Entomol. 188 pp.
4. Newstead, R, 1924. Guide to the Study of Tsetse Flies. Liverpool:
School Trap. Med. in Memoir, no. 1. n.S. 272 pp. (28 plates, 4 maps).
5. Hegh, Emile, 1929. Les tse-tses, vol. 1. Brussels: A. J. Engelterzi. xiv
+ 742 pp. (327 figs. + 15 colored plates).
6. Swynnerton, C. F. M., 1936. "The tsetse flies of East Africa." Tr.
Roy. Entomolog. Soc. London, vol. 84. xxxiv + 579 pp. (7 maps, 22 plates).
7. Bequaert, Joseph C., 1946. "Tsetse-flies in Liberia: Distribution and
ecology; possibilities of control," Am. /. Trap. Med. 26 (Suppl.) :57-94.
8. Griinberg, Karl, 1904. Die Blutsaugenden Dipteren. Jena: Gustav-
Fischer. xi + 188 pp. (127 figures) .
9. Lewis, D. J., 1934. "The behavior of the larvae of tsetse flies before
pupation," Bull. Entomolog. Res., 25:195-99 (1 plate).
10. Nash, T. A. M., 1939. "The ecology of the puparium of Glossina in
northern Nigeria," Bull. Entomolog. Res., 30:259-84.
11. Laveran, Alphonse, et Mesnil, Felix, 1904. Trypanosomes et Try-
panosomiasis, Paris. xi + 417 pp. Translated by D. N. Nabarro, 1907. Chicago
Medical Book Co.
12. Bruce, D., 1897. Further Report on the Tsetse-Fly Disease or Nagana
in Zululand. London: Harrison and Sons. 69 pp.
13. Dutton, J. E., 1902. "Trypanosoma in man," Brit. M. 1. 1:42.
14. ---, 1902. "Note on a Trypanosoma occurring in the blood of man,"
Brit. M. J., 2:881-84.
15. Bruce, D., and Nabarro, D., 1903. Progress report on sleeping sick-
ness in Uganda. Rept. Sleeping Sickness Comm. Roy. Soc. London, no. 1.
88 pp. (10 plates) .
16. Castellani, A., 1903. "Trypanosoma in sleeping sickness," Brit. M. J.,
1:1218.
17. Kleine, F. K., 1909. "Positive Infectionsversuche mit Trypanosoma
brucei durch Glossina palpalis," Deutsche med. Wochnschr., 35:469-70.
18. Roberston, Muriel, 1913. "Notes on the life history of Trypanosoma
gambiense, with a brief reference to the cycles of Trypanosoma nanum and
Trypanosoma pecorum in Glossina palpalis," Phil. Tr. Roy. Soc. London (Ser.
B), 203:161-84 (5 plates).
19. Mitzmain, M. B., 1913. "The bionomics of Stomoxys calcitrans (Lin-
naeus); a preliminary account," Philippine J. Sc., 8 (Sec. B) : 29-48 ..
20. Bishopp, F. C., 1939. The Stable Fly: How to Prevent its Annoyance
and its Losses to Livestock. Washington, D. C.: Dept. of Agric., in i'armers'
Bull., no. 1097. 18 pp. (revised).
21. Freeborn, S. B.; Regan, W. M.; Folger, A. H.; 1925. "The relation of
flies and fly sprays to milk production," J. Econ. Entomol., 18:779-90.
22. Rosenau, M. J., 1912. In "Society Proceedings," J. A. M. A., 59:1314.
Report on poliomyelitis presented at meeting of International Congress on
Hygiene and Demography, 1912.
23. Anderson, John F., and Frost. Wade H., 1912. "Transmission of polio-
myelitis by means of the stable fly (Stomoxys calcitrans)," U. S. Public Health
Service, Pub. Health Rep., 27:332-35.
24. ---, 1913. Poliomyelitis: Further attempts to transmit the disease
through the ag~ncy of the stable fly (Stomoxys calcitrans). U. S. Public Health
25. Sawyer, W. A., and Herms, W. B., 1913. "Attempts to transmit polio-
myelitis by means of the stable fly (Stomoxys calcitrans)," J. A. M. A., 41:461-
66.
26. Hammer, Ole, 1941. "Biological and ecological investigations on flies
associated with pasturing cattle and their excrement," Videnskabelige med-
delelser fra Dansk naturhistorisk forening, vol. 105, 257 pp.
27. Kriggsman, B. J., and Windred, G. L., 1933. Investigations on the
buffalo fly, Lyperosia exigua de Meij. Commonwealth of Australia: Council
for Sc. and Ind. Res., Pamph. 43. 40 pp.
28. Austen, E. E., 1909. Illustrations of African bloodsucking flies other
than mosquitoes and tsetse flies. London: Mus. Nat. Hist. 221 pp. + xiii plates.
CHAPTER XVII
._.
MYIASIS
Myiasis is a term meaning an infestation of the organs and tissues of

man or animals by fly maggots and the disturbances resulting therefrom.
Such invasions may be benign in effect or may result in more or. less
violent disturbances, even in death. When the intestinal tract is invaded
it is called intestinal myiasis; when the stomach is concerned it is gastriC
myiasis; invasion of the urinary tract is called urinary myiasis; invasion
of the nasal passages, nasal myiasis; invasion of the ears, auricular myiasis
or otomyiasis; of the eyes, ophthalmomyiasis; when wounds or ulcera-
tions of the skin are infested by maggots the term traumatic dermal
myiasis is applied; invasion of the skin is also known as cutaneous my-
iasis, etc. When maggot infestations are traceable to species which are
normally scavengers, coprobionts, or saprobionts, the term accidental
myiasis is usually employed; when the species of maggot is a necrobiont
or facultative sarcobiont (involving fresh wounds) the term semispecific
is used; and when the infestation is traceable to obligatory sarcobionts
such as the warble flies and botflies the term obligate myiasis is used.
All who are concerned with the subject of myiasis must become thor-
oughly familiar with The Flies That Cause Myiasis in Man by Maurice
T. James. 1 1>
ACCIDENTAL MYIASIS
Accidental myiasis may be caused by a considerable number of species
of Diptera belonging to several families such as the Calliphoridae (blow-
flies, comprising the bluebottle and greenbottle flies), Sarcophagidae
(flesh flies, scavenger flies), and Muscidae (houseflies, scavenger flies).
The larvae of these flies normally feed on decomposing animal and vege-
table matter, garbage, dead animals, and manures. Infestations in man
are usually traceable to the ingestion of fly eggs or larvae with con-
taminated food or water.
Identification of fly maggots. Maggots, the larvae of muscoid Diptera,
are footless, more or less cylindrical, tapering anteriorly, and truncated
posteriorly; they are distinctly segmented, with ordinarily 11 or 12 visible
" Also see Hall, David G., 1948. The Blowflies of North America. Thomas Say
Foundation. 477 pp. 5 color + 46 black and white plates. "
372
MYIASIS 373
segments (Fig. 119) . Fully grown larvae differ greatly in length ac-
cording to the species, ranging from 5 mm to 35 mm.
At the blunt or posterior end are found the spiracles (Fig. 119)
POSTERIOR
SPIRACLES
PHAENICIA
SERICATA
CEPHALO-PHARYNGEAL SKELETON
PHAENICIA SERICATA
~ POSTERIOR SPIRACLES
~ CALLIPHORA VOMITORIA
CEPHALO-PHARYNGEAL SKELETON
PHORMIA REGINA
POSTERIOR SPIRACLES
PHORMIA REGINA
~
~
POSTERIOR CEPHALO-PHARYNGEAL
SPIRACLES SKELETON
MUSCA MUSCA DOMESTICA

DOMESTICA
POSTERIOR
SPIRACLES
STOMOXYS
CALCITRANS
FIG. 119. Taxonomic details used in the classification of muscoid fly larvae.
(Redrawn after various authors. )
which afford useful diagnostic characters. There are two stigmal plates
more or less separated from each other, within which are situated spir-
acles one to three in number, either slit-like, sinuous, or more or less
circular. A prominence known as a "button" is best seen in certain slit-
like forms, as in Phaenicia (Lucilia) sericata (Meig.) (Fig. 119). The
button may be absent or variously situated depending upon the species,
hence has taxonomic value.
In using the posterior spiracles for purposes of classification the fol-
lowing characters are to be noted: (1) diameter of the stigmal plate, the
space occupied by one stigmal plate on a line drawn through the center
of both; (2) length, when slits are absent, the space occupied by a plate
on a line drawn dorsoventrally through the center of the plate; or, when
slits are present, the space occupied by a plate along a line drawn from
the lower edge of the button (or space if button is absent) through the
longest slit (middle slit) to the margin of the plate; (3) width, along
a line drawn at the middle of the plate at right angles to the length line;
( 4) distance between the plates; (5) general form of the plates; (6)
shape of spiracles; (7) presence or absence of button; (8) general struc-
ture of plate.
Intestinal myiasis. According to Braun and SeiferF at least 30 species
of fly larvae had been reported (1926) from cases of intestinal myiasis.
No doubt, other species have been added since. They are principally
members of the families Muscidae, Calliphoridae, and Sarcophagidae,
which commonly deposit their eggs or larvae on cold meat, cheese, and
other foods of man and are thus ingested. It is also suggested that the
flies may deposit their eggs in or near the anus, particularly in the use
of old-fashioned open privies. The larvae on hatching are believed to
make their way into the intestine.
Hoeppli and Watt (1933),0 experimenting with the larvae of C hry-
somya megacephala (Fabr.) and Phaenicia (Lucilia) sericata (Meig.),
secured results that agree with those obtained by Desoil and Delhaye
(1922) 4 with larvae of Calliphora vomitoria (Linn.) showing that fly
larvae of certain kinds as well as the eggs are able to resist the influence
of temperature, gastric juice, hydrochloric acid, and ferments in all the
concentrations occurring in the human stomach, but that the larvae are
very susceptible to mechanical injuries and to the obstruction of their
stigmata. Ample food for the larvae is provided by the intestinal contents
as well as the intestinal mucosa itself. On the basis of experiments with
dogs and cats which were fed fly larvae with their food, with the result
that no living larvae of any of the species were found either in the large
intestine or in passed feces of these animals, Causey5 (1938) remarks
that" . . . it is time to revise our discussion of intestinal myiasis in ac-
cordance with known facts."
The clinical symptoms of intestinal myiasis depend on the number
as well as the species of fly larvae and on their location in the intestine.
No doubt many instances occur in which living fly larvae are passed in
the stool without having caused any intestinal disturbance. In severe
MYIASIS 375
infestations there will be nausea, vertigo, and more or less violent pains
in the abdomen; diarrhea with discharge of blood may occur as the result
of injury of the intestinal mucosa by the larvae. Living and dead larvae
are expelled with either the vomit or stool, or with both.
An obstinate case of intestinal myiasis was reported by Herms and
Gilbert (1933). 6 The patient, Z. W., female, age 38 years, was first seen
April 26, 1930. Her chief complaints were attacks of nausea, vomiting
and diarrhea, nervousness, and joint aches. There were recurring attacks
of nervousness, vomiting, and diarrhea, and apparently rather frequent
hemorrhages from the bowels. The patient was considerably depressed at
times, and treatment was difficult because of lack of cooperation except
after she had had a bad spell. Because of the difficulty of obtaining stool
specimens, especially during the acute attacks, and in view of the fact
that it was felt that there must be other reasons for her condition, early
in the spring of 1931 during an attack of nausea with vomiting and
diarrhea the patient was kept for one entire day in the office under ob-
servation and stool and vomit specimens were obtained, both containing
the first larvae which were studied. During these attacks it was difficult
for her to obtain relief with fairly large doses of opiates. Following this
observation she was given santonin by mouth and colonic irrigations
containing thymol. Many larvae were recovered after this, all of which
were dead. Following the attacks of diarrhea the patient had a number
of severe hemorrhages. Tetrachlorethylene capsules were given by
mouth, but they caused gastric distress. In the hospital a duodenal tube
was passed and tetrachlorethylene was injected beyond the stomach. For
a few weeks there was apparent improvement, but the attacks recurred,
with the passage of larvae by vomiting and bowel discharge.
Three lots of fly larvae were studied in the laboratory, i.e.: March 31,
May 12, and July 28, 1931. Adult flies belonging to three genera, Call-
iphora, Phaenicia (Lucilia), and Sarcophaga, were reared from these
larvae.
The recurrence of violent symptoms with evacuation of larvae in
vomit and stools would ordinarily point to repeated infections, but the
fact remains that the patient as observed by her physician, the co-author,
lived in a way that would seem to preclude repeated infestations, and
the exposure of stools and vomit to flies is ruled out because of circum-
stances indicated in the case report. The authors have advanced a possi-
ble explanation based on the pedogenetic reproduction of fly larvae as
suggested by observations made by Parker (1922), 7 viz.: "The increases
led me to believe that Calliphora erythrocephala occasionally multiplies
in an unusual way, and that this way is not polyembryony but pedo-
genesis." In the case described there were certainly broods of very young
larvae at intervals, at which time also full grown larvae were present.
Cheese skippers as agents. The larvae of the cheese fly, Piophila casei
(Linn.) of the family Piophilidae, frequently cause intestinal myiasis, as
they are able to pass through the digestive tract without injury. Sim-
mons s cites a number of instances indicating the frequency of their oc-
currence in the digestive tract of man. The adult flies measure from 2.5
to 4 mm in length; superficially they appear shining black, with reddish-
brown eyes and wings held flat over the dorsum when at rest. The eggs
are deposited on cured meats, old cheese, dried bones, smoked fish, and
many similar materials. The eggs hatch in from 30 to 48 hours at a
temperature of 65° F; the larval stage requires about 8 days, the pupal
about 12 days. These stages are greatly influenced by temperatures. The
larvae have the peculiar habit of curving the ends of the body together
and then suddenly springing to a distance of from three to six inches.
Soldier fly. A case of intestinal myiasis caused by the larvae of a
soldier fly, H ermetia illucens (Linn.) (Family Stratiomyidae ), is reported
by Meleney and Harwood. 9 This fly feeds mainly on flowers, and the eggs
are deposited on decaying fruits, vegetables, and animal matter. The
source of infection according to the authors was apparently raw fruit
or vegetables. The symptoms were local irritation in the stomach and
rectum, and spells of fainting. The patient was a boy of ten years. M. A.
Stewart informed the author that he observed a patient in Houston,
Texas, who continued to expel these larvae for several months.
Gastric myiasis. The frequency with which the "rat-tailed" larvae
of the drone fly Tubifera tenax (Linn.), occur in liquid excrement must
lead to extreme caution in accepting reports that these larvae have been
evacuated with discharges from the bowels. The writer has on several
occasions received specimens of "rat-tailed" larvae which were said to
have been evacuated by the "double handful" and the patient was said
to have "steadily improved" thereafter.
There are, however, several recorded cases which seem to be incon-
trovertible, notably the case reported by Hall and Muir,ID who also
brought together all information then available relative to Tubifera and
myiasis. The case referred to was that of a boy aged five years "who had
been ailing for about ten weeks and who had been under medical treat-
ment for indigestion and obstinate constipation for about five weeks at
that time. The child was emaciated and anemic. Very striking symp-
toms were the constant and pronounced twitching of the eyelids and
other nervous movements. He gritted his teeth in his sleep at times,
and made convulsive movements of the limbs. When awake he com-
plained of pain in the limbs and headache. The emaciation seemed to be
due to the fact that the boy had for some time vomited almost
everything he ate. The breath was very bad, 'worse than rotten eggs,'
according to his parents. On the basis of the nervous and digestive
MYIASIS 377
disturbance and the general debility, a diagnosis of worm infestation
was made."
With this diagnosis in mind the mother of the boy gave him a dose
of a proprietary worm remedy, resulting in the discharge of an object
wriggling round vigorously in the feces and urine. The slop jar into which
the stool was passed was in regular use and had been previously rinsed
with tap water and allowed to dry during the day. The specimen was
identified by the authors as one of the "rat-tailed larvae" measuring 3.2
cm. in length, including the long "tail." A second larva was said to have
been discharged the following day. The case is believed by the authors
to be probably a genuine case of "gastric myiasis."
After the ,passage of the larvae the child is said to have improved in
health and become normal; the nervous symptoms and vomiting disap-
peared.
Three chances for infection were painted out; namely, first, the eating
of "overripe" or probably decaying peaches in which "rat-tailed" larvae
might have occurred; secondly, the drinking of "ditch" water polluted
with kitchen refuse, etc.; lastly, stable manure in a neighbor's yard where
the child played.
The authors offer the following comment relative to the gastric dis-
turbances:
A larva supplied with the stigmatic apparatus of Eristalis would appar-
ently be fitted for life in a stomach with a small amount of food and plenty
of the atmospheric air which is swallowed in eating and drinking and at other
times. Such a condition would simulate the normal life conditions fairly
closely. That the stomach would not fill to the point where it would drown the
larva might be insured by the vomiting, perhaps automatically, the activity of
the larva increasing as the stomach filled to where it threatened to cover the
rising stigmatic tube, and so setting up an irritation leading to vomiting. The
mother states that the child's stomach 'was extremely intolerant of milk and
that drinking milk was promptly followed by vomiting. This suggests that
milk, usually taken in long drinks and considerable quantities, quickly threat-
ened the larva with drowning and set up such activity as promptly to cause
vomiting.
HaHn has added the records of other cases, giving a total of 17 claim-
ing the presence of syrphid larvae in the digestive tract of man, one
claiming their presence in the nostrils of man, and two claiming their
presence in the diseased vagina of cattle.
Tubifera (= Eristalis) tenax (Linn.), the drone fly (Fig. 120), is a
large insect, somewhat larger than a honeybee and resembling the drone
bee very closely; indeed it is commonly referred to as its mimic. The fly
deposits its eggs on liquid manure or other fllthy liquids in cans, slop
jars, privies, septic tank effluent, etc. The larvae are known as ,"rat-tailed
larvae" (Fig. 121); these also occur occasionally in heaps of horse

manure.
The family Syrphidae includes a very large group of flies, varying
greatly in size, many of which are brightly colored. They are nearly aU
flower loving, feeding on nectar mainly. Only one genus needs to be con-
sidered here, namely Tubifera, the larvae of which have a long anal
breathing tube, i.e., "rat-tailed," and the adults of which are commonly
called drone flies.
Urinary myiasis. As in intestinal myiasis the symptoms of urinary
myiasis depend on the number and kind of larvae, and their localization.
There may be obstruction and pain; pus, mucus. and blood in the urine;
12(J 121
FIG. 120. The drone fly, Eristalis tenax, -whose larvae are commonly called
"rat-tailed larvae." X 3.5.
FIG. 121. The "rat-tailed larva" of Eristalis tenax, drone Hy. x2.
and a frequent desire to urinate. Larvae are expelled with the urine.
ChevriP~ reports that Fannia canicularis (Linn.) (see Chapter XV) is
most frequently found in urinary myiasis, although Fannia scalaris
(Fab.) and Musca domestica LinnY have been encountered. ~ppli
and Watt (lac. cit.) believe that albumin and sugar in the urine may
provide food, as may mucus and leucocytes; the lack of oxygen presents
the chief difficulty, although very small amounts of oxygen are needed by
the larvae.
Infection is probably usually accomplished at night in warm weather
when persons may sleep without covering. The flies deposit their eggs
MYIASIS 379
around the urethral opening; these hatch in a few hours, and the larvae
enter the urethra.
Traumatic dermal myiasis, the invasion of wounds or ulcers of the
skin by fly larvae, is of common occurrence in warm, humid climates. A
large number of species of flesh flies are responsible for this type of
myiasis, particularly the screwworm flies, green bottle flies, and related
species.
The following description of a case caused by Phormia regina (Meig.)
reported by StewartI4 will serve to illustrate this form of myiasis:
The dermatitic area was not large at first but it continued to spread after
hospitalization. An extremely offensive odor was given off, but aside from the
dermatitis and irritation of the sores, the patient appeared to be feeling well;
appetite, digestion and egestion were good. At first treatment was applied only
to the area around the ears, but on the night of the patient's admission to the
hospital the discovery was made that the scalp was a mass of pus and a super-
saturated sulphur wash was applied. The hair was parted to allow the wash
to penetrate freely to the scalp and a towel was tied about the head, coming
below the ears. The supersaturated sulphur wash was applied every two hours.
After the second treatment was applied to the scalp the patient became
very restless, working the fingers into the palms of her hands and alternately
putting her hands to her ears. Soon she began to scream, acted frantic, and
became nearly delirious. She was given a sedative without effect.
On taking the towel from the patient's head the nurse observed fly larvae,
which had been forced into activity by the treatment, crawling over the towel,
hair and down the cheeks. The nurse estimates that she killed twenty-five or
thirty larvae in the hour and a half she spent in removing them and still the
hair and scalp remained full of them. Back of the ears the mass of living larvae
was so great that they could almost have been spooned out. At this time the
patient complained of a buzzing in the ears similar to that occurring when the
ears are full of water, and said that she could not hear. The nurse then used
toothpick swabs to remove the great quantity of larvae found in the pinnae of
the ears; in so doing most of the larvae were killed, but some were kept alive
and placed on raw beef in vials so that they might complete their larval growth
and pupate.
As soon as pupation occurred the puparia were removed to fresh vials and
covered with fine dry soil until they emerged as adults, when they were iden-
tified as Phormia regina Meig.
Mter the removal of all visible larvae had been completed the patient's
hair was clipped, the supersaturated sulphur wash treatment was continued,
and the scalp was bandaged. To the original area of dermatitis around the ears
was applied a paste consisting of salicylic acid, 2 gm.; zinc oxide, 24 gm.;
starch, 24 gm.; petrolatum, sufficient to make 100 gm.
It is obvious that an adult female fly had been attracted to the suppurating
scalp sores by the foul odor given off and had oviposited in one or more of
these sores. The larvae were driven from the scalp to the pinnae of the ears
by the application of the supersaturated sulphur wash.
Chrysomya megacephala (Fabr.) was responsible for much traumatic

myiasis in the South Pacific Islands during World War II. StewarP5
describes a new treatment for traumatic dermal myiasis:
A new douche, composed of 15 per cent chloroform in light vegetable oil ,
has been employed in the treatment of seventeen cases of traumatic dermal
myiasis. In every case all the maggots were removed with a single treatment,
extending over a period of thirty minutes.
The new douche has advantages over the commonly used chloroform-milk
solution in that chloroform is entirely soluble in vegetable oil and only slightly
soluble in milk; in that the chloroform-vegetable oil solution is very stable and
can be kept indefinitely in closed containers, whereas the chloroform-milk mix-
ture has to be made up fresb for each application, and in that the vegetable oil
is very soothing to the raw tissue of the infested wound.
SEMI -OBLIGATE MYIASIS
Callitroga (= Cochliomyia) americana C. & P. (Fig. 122) is the

name recently given to the New World screWWorm fly by Cushing and
Patton!G to separate this dangerous form (the primary screwworm),
FIG. 122. Callitroga (= Cochliomyia) americana, the Texas screwworm fly,

larva, and pupa. (After Bishopp.)
which feeds on the tissues of living animals, from Callitroga (= Coch-

liomyia) macellaria (Fabr.), the secondary S~rewworm fly (Fig. 123),
which is more particularly a scavenger fly. C. americana is an obligate
parasite and according to Laake, Cushing, and Parish initiates the great
majority of cases of screwworm infestations in man and animals in the
United States and probably in the entire Neotl'opical region. It is known
to be the cause of nasopharyngeal myiasis in man. The following is a de-
MYIASIS 381
tailed description of a case which terminated fatally (from Osborn
[1896] quoted by Richardson in the Peoria, Ill., Medical Monthly for
February, 1883):
While traveling in Kansas in the latter part of last August, a citizen of
this place had the misfortune to receive while asleep a deposit of eggs from
this fly. He had been troubled for years with catarrh, hence the attraction to
the flies. He returned horne a few days after the accident and shortly after be-
gan complaining of a bad cold. Growing rapidly worse, I was called to attend
FIG. 123. Callitroga (= Cochliomyia) macellaria just emerged from the

pupa cases. A dead animal near by furnished food for the larvae; pupation
took place in the sand underneath the carcass. The newly emerged flies have
crawled up on the grass and will soon be ready to flyaway. Note characteristic
resting attitude, with head down.
him . Monday, my first day, his appearance was that of a man laboring under a
severe cold. Had slight congestion of the lungs, and moderate fever. His nose
seemed greatly swollen and he complained of a smarting, uneasy feeling in it,
and general misery through the head. Gave him treatment to relieve the con-
gestion and fever. Tuesday, saw him again. His nose and face were still
swollen, and in addition to the other symptoms he was becoming slightly de-
lirious and complained a great deal of the intense misery and annoyance in his
nose and head. A few hours after, I was sent for in haste with the word that
something was in his nose. I found on examination a mass of the larvae of this
fly (or 'screw worms' as they are commonly caned in the South) completely
blocking up one nostril. On touching them they would instantly retreat en
masse up the nostril. Making a 20 per cent solution of chloroform in sweet milk
I made a few injections up both nostrils, which immediately brought away a
large number, so that in a few hours I had taken away some 125 of them. By
Wednesday evening erysipelas had begun, implicating the nose and neighbor-
ing portions of the face. Another physician was called. By continual syringing
with a strong antiseptic solution of salicylate of soda, bicarbonate of soda, and
carbolic acid we hoped to drown out the remaining larvae. But they had by
this time cut their way into so many recesses of the nose and were so firmly
attached that we were unnable to accomplish much. Finally we resorted to the
chloroform injections, which immediately brought away a considerable number.
Friday I was able to open up two or three canals that they had cut, extracting
several more that had literally packed themselves, one after another, in these
fistulous channels. His speech becoming suddenly much worse, I examined the
interior of his mouth and found that a clear-cut opening had been made en-
tirely through the soft palate into his mouth and large enough to insert the end
of a common lead pencil. Saturday the few remaining larvae began changing
color and one by one dropped away. On Sunday for the first time hemorrhage
from both nostrils took place, which continued at intervals for three days, but
was not at any time severe. On this day the patient began to improve, the de-
lirium and erysipelas having subsided, leaving but little or no annoyance in his
head. In a few days he became able to go about home, and even to walk a dis-
tance of half a mile to visit a friend and return. But while there he began com-
plaining of a pain in the neighborhood of his left ear, apparently where the
eustachian tube connects with the middle ear. It proved to be an abscess. Being
already so reduced by the first attack he was unable to withstand the second,
and died after an illness of nearly three weeks, completely exhausted by his
prolonged sufferings. Three days before his death the abscess discharged its
contents by the left nostril. The quantity of pus formed was about 2lf.l ounces
(78 grams).
In all about 250 larvae were taken away from him during the first attack,
and, as the visible results, not only had they cut the hole through the soft
palate, but had also eaten the cartilage of the septum of the nose so nearly
through as to give him the appearance of having a broken nose. The case occu-
pied, from the first invasion of the fly to its final result, nearly two months. He
doubtless would have recovered but for the formation of the abscess, which
from all the symptoms, was caused by one or more of the larvae having found
their way up the left eustachian tube.
Callitroga (= Cochliomyia) americana C. & P. is strongly attracted
to the wounds and sores of animals. Laake17 estimates the loss occasioned
by this fly in the southwestern United States at $5,000,000. He found the
following predisposing causes of attack: among sheep and lambs, wounds
caused by needle grass take first rank; among goats and kids, shear cuts
take first rank; among cattle, injuries by the horns of other cattle; among
calves, exposed tissue at birth; and among horses and mules, wire cuts.
Laake points out that the more common causes of screwworm attack
MYIASIS 383
are due to farm practices that can be corrected. He stresses particularly
care in shearing, dehorning, removing and disposing of old barbed wire
from dismantled fences, also the timing of dehorning, castrating, and
branding so as to expose the wounds as little as possible to flies during
the season of abundance.
An epidemic of human myiasis, 81 cases in 5 provinces of Chile oc-
curring during the months of December to April, 1945-1946, is reported
by Gajardo-Tobar and Honorat018 who also describe the biology of the
fly concerned, Callitroga americana (C. and P.) = [Cochliomyia hom-
inivorax (Coq.)].
Cutaneous myiasis. The larvae of Wohlfahrtia vigil (Walk.), a mem-
ber of the Sarcophagidae in eastern North America, frequently cause
cutaneous myiasis. Walker19 describes the cases of a five-month-old boy
thus:
Most of the lesions were clustered together on the left side of the neck
under the angle of the jaw, one being on the left cheek. They had been first
noticed by the mother 24 hours earlier, and when seen by the writer they were
already secondarily infected with pus organisms, the child being in a poor gen-
eral condition and suffering from an intestinal disorder. They were similar to
the lesions observed in the previous cases, each being a boil-like sore with an
external opening, and from these openings five or six larvae had already been
expressed. Only three additional larvae were obtained, these measuring 5 to
7 mm. in length. Each was placed on raw beef in a separate test-tube, plugged
with cotton wool. In 24 hours they reached a length of 12 to 13 mm., and in
another 24 hours they were full-grown, each measuring about 17 mm. in
length.
On the third day after their removal from the child the larvae were placed
with the meat in a jar of earth and immediately burrowed into the latter. Next
day they were at the bottom of the jar and two of them had begun to contract.
Three days later they were dug up and all had transformed into puparia.
On September 27, 18 days later, a male Wohlfahrtia vigil emerged. I
waited for the others to appear until October 5, but neither having emerged by
that time I opened one of the puparia on that day and the other a week later,
and in both I found pupae which had evidently died some time before the
proper time for emergence, as they were quite colorless. In this case, like the
previous ones, the child recovered rapidly after the removal of the maggots.
Two new cases of cutaneous myiasis in Panama were reported re-
cently (1948) by Calero,20 viz.: Case I. Removed second-stage larvae of
Stephanostoma (~ Sarcophaga) haemorrhoidalis (Fall.) from fistulous
tract in the region of the mentum. Case II. Removed 166 larvae of Call-
itroga (= Cochliomyia) hominivorax (Coq.) from right parietal region
of the cranium; three points of perforation of the periosteum were found
upon removing the larvae. The patient did not present either meningeal
symptoms or encephalic irritation.
Life history of the screwworm fly. The adult fly of Callitroga
(= Cochliomyia) americana (C. & P.) has a deep greenish-blue metallic
color with yellow, orange, or reddish face, and three dark stripes on the
dorsal surface of the thorax. It is difficult, unless one is experienced, to
separate this species from Callitroga (= Cochliomyia) macellaria
(Fabr. ). Laake, Cushing, and Parish state that the females of C. mac ell-
aria (Fabr.) may usually be distinguished from C. americana C. & P.
by the fact that the basicostal scale (a small sclerite at the base of the
wing) of the former is of a yellowish color, whereas in the latter (C.
americana C. & P.) it is black. Also C. macellaria (Fa br.) is covered on
the mid-line of the underside of the abdomen by a dense white pruinosity
absent in C. americana C. & P. The species can be easily and accurately
determined by the use of the characters exhibited by the male termi-
nalia. 21
Individual females of Callitroga (= Cochliomyia) americana C. & P.
according to Laake, Cushing, and Parish, may lay as many as 2,853 eggs,
the eggs being deposited in characteristic batches of 10 to 393 eggs each,
and the laying of as many as 300 eggs may be completed in from four
to six minutes. The incubation period of the eggs on wounds in animals
ranges from 11 to 21.5 hours, under natural conditions. The larval feed-
ing period ranges from 3.5 to 4.5 days; the prepupal period from a few
hours to about three days (7 hours to 76 hours); the pupal stage lasts
about seven days. The prepupal and pupal stages are greatly influenced
by temperature and moisture. The life history from egg to adult under
optimum natural conditions requires about 11 days.
Screwworm control. The screwworm is a true parasite and lives only
in the living flesh of warm-blooded animals; it is not found in snakes,
lizards, or other cold-blooded animals, nor in carcasses, dead fish, de-
caying meats, or decaying vegetable matter. The maggots found in dead
animals are not true screwworms (Bruce and Sheely22). In the light of
this information much serious damage by screwworms may be prevented
by good herd management, which involves such matters as controlled
breeding and proper utilization of climate and natural growth of native
grasses; having the calves dropped to come along with the grass; elimi-
nating nonbreeders and shy breeders; separating the breeding herd from
the steers; feeding bulls during the winter and having them in good
condition in spring; castrating bull calves when they are young by the
bloodless operation; nubbing the sharp points of horns; marking and
branding during the winter season and using a repellent smear on these
parts; using common sense in handling and driving cattle; eliminating
the use of the whip and catch dogs; avoiding the jamming of cattle
into pens and chutes; eliminating unnecessary wounds and bruises of all
kinds. In addition, Bruce and Sheely point out that livestock should be
--- -_ ~---------- __- -
MYIASIS 385
examined frequently for wounds and injuries and that all cases of screw-
worm should be treated promptly; in treating animals one should avoid
making wounds bleed; animals should be castrated, marked, branded,
and dehorned during the winter months; bloodless emasculaters should
be used for castration; all wounds should be treated with a repellent
smear; ears of cattle, sheep, and goats should be inspected for ticks;
breeding should be controlled so calves will be dropped in the early
spring.
SUNDRY MAGGOT INFESTATIONS

Tumbu fly and Congo floor maggot. Several tropical African species
of calliphOrine Hies are commonly referred to in the literature on myiasis,
among them Cordylobia anthropophaga (E. Blanchard), the "tumbu
fly." Austen describes it as being a "thickset, compactly built fly, of an
average length of about 9Yz mm . . . . Head, body, and legs are straw
FIG. 124. Congo floor maggot, Auch-

meromyia luteola. (After Blacklock, in 1\lar-
lini's Zoonosen der Haut in wiirmeren Liill-
dern.)
yellow." According to Blacklock and Thompson"" the eggs are deposited
in excrement-polluted sand and soil. The incubation period may be as
short as 24 hours. If contact is made with the skin of man or other ani-
mals the larvae penetrate the unbroken skin, forming furuncular swell-
ings, or where multiple and contiguous infection occurs, extensive
'''sloughing and gangrenous" conditions result. In 8 to 10 days the full-
grown larvae, measuring 13 to 15 mm in length, leave the host and
pupate in a few days. Wild rats are looked upon as the main reservoir of
the infection in nature.
In the same locality with the "tumbu fly" there may also occur Auch-
meromyia luteola (Fabr.), the larva of which is a bloodsucker and is
known as the "'Congo floor maggot" (Fig. 124). The Hy is commonly
found in and about human habitations. The eggs are deposited in small
clusters in various situations, such as on sleeping mats spread on the
ground in huts, in dusty crevices, in dry sand, situations where the larvae
may readily find suitable food. According to Roubaud,24 whose treatise
on this insect should be read by all interested in this subject, the eggs
hatch in 36 to 40 hours., The larvae are remarkably resistant to extreme
dryness and lack of food. They are nocturnal in their feeding habits,
sucking the blood of sleeping persons, producing a wound by means of
powerful buccal hooklets. They feed for 15 to 20 minutes, detach and
hide in the crevices of mats, etc., during the day, repeating the attack
almost nightly if hosts are available. The larval period may be as short
as two weeks or, in the absence of food, perhaps as long as three months
when the larvae pupate in protected situations. The pupal stage lasts
from 11 to 12 days.
Wool maggots. Blowflies, inclusive of the screwworm fly already dis-
cussed, were undoubtedly at one time solely scavengers feeding in the
maggot stage on carrion and animal wastes, but with the introduction of
herds of domesticated animals they have acquired the habit of attacking
living animals. The term "blowfly strike" refers to cutaneous myiasis
produced by the development of blowfly maggots on living sheep. Frog-
gatt 25 writes:
Blowflies have always been in Australia, and under natural conditions be-
fore such disturbing elements as cattle, horses, sheep, and rabbits were intro-
duced, they were simply scavengers or flesh-flies that deposited their eggs in
animal matter that happened to be festering in the sun. In the study of eco-
nomic zoology the acquisition of new habits by an insect is no uncommon oc-
currence. In the case of the blowfly we know that first some one species learned
the habit of blowing live wool from the practice (already acquired) of blowing
the wool of sheep that had died from some other cause. The smell of the dead
wool taught them that the damp or soiled wool on a live sheep was of a similar
character, and once the maggots set up decomposition of the yolk and fibre the
smell attracted other flies. Thus other species became sheep-maggot flies until
all our common blowflies had learned the habit.
To upset the balance of nature is always a dangerous undertaking, and
there is not much doubt that it has been the destruction of the natural means
of control of the fly and the provision of a much larger number of suitable
breeding-grounds that has caused the remarkable increase in the number of
flies. After great droughts landowners have had to fight millions of rabbits
breeding all over the West, and poisoned water and poison-carts have been
put to considerable use. Sometimes the dead rabbits were stacked up and left
to rot; sometimes they filled up the water-holes; sometimes they were piled
up feet high along the wire-netting fences. It can be easily understood that
under such conditions the blow-flies must have increased a millionfold. Again,
the poisoning of dingoes and rabbits has caused the death of many of our
hawks and carrion crows, and I feel quite sure that one of these birds in tear-
ing up a carcass will often kill far more maggots than a dozen of the smaller
insectivorous birds that capture the flies, but often only after they have de-
posited their eggs on the sheep.
The next, and perhaps the most important, factor in the development of
the sheep-maggot pest is the work of sheep-breeders themselves. Forty years
ago there were many thousands of Merino sheep of the bare-belly, bare-legged
MYIASIS 387
type, which did not produce a third of the weight of wool of the modern,
improved Merino. The ambition of every sheep-breeder has been to make
every inch of the sheep's skin grow wool, and in the case of some classes of
Merinos to produce a wrinkled skin, giving even more wool-bearing surface.
A sheep clothed with such a mass of thick, close, fine wool, fitting closely over
the rump and round the tail, is sure to get more or less stained and damp round
the crutch, and to attract flies. This artificial increase in weight, quantity and
fineness of wool is accompanied, too, by an increased secretion of yolk, which
rising from the skin and spreading all through the wool fibre, forms an addi-
tional attraction for the flies, and supplies food for the maggots.
The following species of flesh flies are listed by Froggatt and other
authors as attacking live wool: Phormia regina (Meig.), known as the
black blowfly because of its blackish-blue color and regarded as the most
important wool-maggot fly in the United States; Phaenicia (= Lucilia)
sericata (Meig.), an important sheep maggot fly in the United States,
Australia, and other parts of the world; Chrysomya albiceps (Wied.)
[Chrysomya (Pyconosma) rufifacies (Macq.)], common in Australia
and India; Callitroga (= Cochliomyia) americana (c. & P.), a menace
to sheep raising, particularly in the southern United States, Mexico, and
elsewhere in the range of this insect; Chrysomya bezziana (Villen.)
[Chrysomya flaviceps (Wlk.) = Chrysomya dux (Esch.)], an important
Australian and Oriental sheep maggot fly; Wohlfahrtia magnifica
(Schin. ), the principal sheep maggot fly of southern Russia, also causes
human myiasis in Egypt.
Control of wool-maggot flies. The following measures are recom-
mended for the prevention and treatment or «blowfly striKe" by various
workers, among them Babcock and Bennett26 of the Texas Agricultural
Experiment Station, and Belschner 27 of the New South Wales Depart-
ment of Agriculture. (I) Carcass burning: dig a trench along the back
of the carcass nearly as wide as the animal and as long, and 12 to 14
inches deep; fill trench with wood (one-quarter cord of wood is sufficient
to burn cow or horse) or cow chips; start fire at windward end and entire
carcass will be wholly consumed within 12 to 24 hours. (2) Poisoning:
a freshly killed animal is very attractive to Hies, and if treated with an
arsenic solution will kill every fly that feeds on it, or if a carcass is al-
ready alive with maggots, a similar treatment will kill these also. For
this purpose arsenite of soda at the rate of 3 pounds to 30 gallons of
water is recommended. (3) Crutching, if properly done, prevents the
attack of Hies by reducing the opportul1ity for the wool of the crutch,
rump, and Hanks to become wet and soiled and thus attractive to Hies.
It consists particularly of shearing the wool away from the breech, over
the tail and down the back of the hind legs. (4) Jetting: applying an
arsenic solution (one pound of arsenic to 40 gallons of water) by means
of a single jet under pressure of 125 pounds to the rump of the sheep;
in this way the arsenic is forced through the wool to the skin, where it
dries and protects the animal for a longer time than spraying. (5) Para-
sites: the use of parasites against the flies.
Foot maggots of animals. A lameness that varies in degree and is
traceable to myiasis commonly occurs in Philippine cattle, carabaos, and
goats and is caused by the larva of Booponus intonsus Ald. Woodworth
and. AshcrafF8 state that the eggs of this fly are attached to the hairs
of the lower portions of the legs of the host animal, the incubation period
varying from three to five days. The young larvae work their way down
to the foot and enter the flesh, leaving their posterior ends exposed, and
when full grown at the end of two to three weeks they leave the flesh
and drop to the ground where they bury themselves and pupate, the
pupal period requiring about 10 days.
The method of treatment suggested by these authors consists of clean-
ing the affected area with soap and water, removing as many maggots
as possible and applying a chloroform pack, followed by heavy applica-
tions of pix liquidae every third day until the lesions heal. As a pre-
ventive means, daily inspection is recommended of all cattle, carabaos,
and goats, especially during the dry season when the animals should
have access to plenty of water and mud as wallows. I
Toxic effect of ingested fly larvae. A disease known as "limberneck"

in chickens is believed to be traceable at least in part to the ingestion of
large number of fly larvae such as those of Lucilia caesar (Linn.), Phae-
nicia sericata (Meig.), and no doubt other species of flesh flies, or of
meat infested with type C Clostridium (= Bacillus) botulinum (Van
Ermengem ) .29 It is believed that the organism multiplies in the un buried
bodies of dead animals, as flesh is a favorable medium for growth, and
that the flesh flies developing in the carcasses become infected, and in
J:urn chickens eating the maggots (or the flesh) acquire the infection .
. This is another good reason why dead animals should be speedily and
. safely disposed of, preferably by incineration.
Bloodsucking maggots of birds. In a study of bloodsucking fly larvae
in birds' nests Plath30 found an average of 61 per cent of a total of 63
nests examined to be infested with an average of 47 maggots per nest.
The species of birds were the Nuttall sparrow, California purple finch,
greenback goldfinch, willow goldfinch, and California brown towhee.
In a later paper the same author 31 adds several other species, namely,
the rusty song sparrow, cliff swallow, Oregon towhee, yellow warbler,
western robin, russet-backed thrush, and cedar waxwing. The species
of flies responsible for the maggots were found to be Proto calliphora
azurea (Fallen), Phormia metallica Town., both blowflies, and Hylemyia
nidicola Ald., an anthomyid fly. Storer32 reports taking 76 larvae and 24
MYIASIS 389
pupae from the nest of the Bailey mountain chickadee. These were identi-
fied by Aldrich as Protocalliphora splendida Macq. variety near hirudo
S.&D.
Plath concludes that from 5 to 10 per cent of the parasitized nestlings
die from loss of blood, and some of them which do become full-Hedged
are so weakened by the loss of blood that they fall an easy prey to rapa-
cious animals.
Guberlet and Hotson 33 ( 1940) reported larvae of Protocalliphora
hesperia S. and D. infesting a young nestling English sparrow (Passer
domesticus) at Seattle, Washington. These authors describe the larvae
and reared adult Hies which emerged in 14 days after the maggots had
been removed from the bird and placed in soil to pupate.
OBLIGATE MYIASIS
Oestrid flies, now segregated into four families, were heretofore placed
in one family, the Oestridae. The larvae of all oestrid Hies are obligate
parasites in the digestive tract or other parts of the body of many mam-
mals. Bots are the larvae of members of the family Gasterophilidae,
medium-sized, 9-18 mm long, stout, pollinose Hies; the eyes are large;
the antennae are depressed in facial grooves, the arista is bare; mouth
parts are rudimentary; the thorax is pilose; the wings are large, the apical
cross vein is absent, the squamae are small. Warbles are the larvae of
the family Hypodermatidae, large robust,' hairy, bee-like flies; with
arista bare, palpi small or absent; mouth parts rudimentary; wings large,
apical cell open. The term "'warbles" is also applied to the larvae of the
family Cuterebridae, large hairy Hies often black and white in color.
As in other oestrids the mouth parts of these Hies are nonfunctional al-
though there is a large oral orifice; the palpi are very small; the arista
is naked or plumose; wings and squamae are large. Head bats refer to
the larvae of the Oestridae (in a restricted sense), bee-like flies known as
"nose Hies"; with eyes naked and small, antennae depressed in sunken
facial furrows, mouth parts vestigial, wings and squamae large; the fe-
males are larviparous.
The horse botflies belong to the family Gasterophilidae with only one
genus, Gasterophilus,34 and four North American species, (1) Gastero-
philus intestinalis (DeGeer), with cloudy patches near the center and
apex of the wings and possessing a prominent spur on the third tro-
chanter; (2) G. inermis (Brauer), which also has wings with cloudy
patches, but the trochanter is without a spur; (3) C. haemorrhoidalis
(Linn. ), without cloudy patches on the wings, and with, the anterttJr
basal cell markedly shorter than the discoidal cell and the tip of the
abdomen reddish; and (4) G. nasalis (Linn.), also with hyaline wings
and anterior basal cell equal or nearly equal in length to the discoidal
cell. The Hies of this genus are somewhat smaller than honeybees, the
mouth parts are rudimentary, the antennae are very small and sunken
in pits, the arista is bare, and according to Curran the apical cross-vein
is absent; the vein closing the discal cell is also absent and the fourth and
fifth veins evanescent apically; the squamae are small; the ovipositor of
FIG. 125. Eggs of the horse botfly, attached to a

hair of the host. X 20.
the female is large and protruberant. They are strong Hiers. The larvae
live in the stomach and intestines of horses.
Gasterophilus intestinalis (DeGeer) [Gasterophilus equi (Clark)]
is the common horse botHy or nit Hy, a widely distributed, nearly cosmo-
politan, species commonly seen in the United States during midsummer
A B c
FIG. 126. Horse botflies. (A) Gasterophilus intestinalis, (B) G. haemorr,
llOidali&, and (C) G. nasalis. (Adapted after Hearle.)
to early autumn, June to September. The light yellow eggs (Fig. 125)
are firmly attached to the hairs of the forelegs, belly, Hanks, shoulders,
and other parts of the body of the horse, but chieHy on the inside of the
knees where they are accessible to the tongue, teeth and lips. The female
Hy (Fig. 126A) hovers from two to three feet away from the animal,
MYIASIS 391
darting swiftly and repeatedly at the horse, each time attaching an egg
to a hair. Wells and Knipling34 report one fly placing 905 eggs in 2%
hours. Friction and moisture from the tongue of the horse seem necessary
for the hatching of the eggs; the incubation period is from 7 to 14 days,
but may be greatly prolonged. by cool weather so that viable eggs may be
found unhatched on the hair of the horse until late autumn, long after
the flies have disappeared. Eggs kept in dry cartons may remain viable
at room temperature for at least three months and hatch when moistened
with saliva. The larvae on hatching (Fig. 127) are provided with an
127 128
FIG. 127. Newly emerged larva of the horse botfly . X 60.

FIG. 128. Larva of the horse bot, Gasterophilus intestinalis. X 4.
armature which enables them to excavate galleries in the subepithelial

layer of the mucous membrane. W ehr, :< 5 who has studied the behavior of
the larvae, states, "Many very small , thread-like subepithelial burrows,
ramifying in every direction, were visible on the anterior half of the
tongue, while those on the posterior half of the tongue were larger in
size. Larvae were visible at the terminations of many of these galleries."
Wehr found that newly hatched larvae when placed on the tongue of a
freshly killed rabbit almost immediately began burrowing and within
one minute nearly all became entirely embedded in the mucous mem-
brane. From the mouth in the normal host the larvae apparently pass
rapidly to their preferred site in the alimentary canal, the left sac or
esophageal portion of the stomach, where third (even second) instar
and the final instar larvae remain fixed with little or no change in
position until the following spring and early summer, when they detach
themselves and pass out of the intestine with the droppings. They are
then from 1.5 to 2 cm in length (Fig. 128). Pupation takes place shortly
thereafter in loose earth or in dry droppings. The pupal stage varies con-
siderably, depending upon moisture and temperature, but the usual time
is from three to five weeks when the winged botfly emerges. Copulation
soon takes place and egg laying begins in early summer. The life history
requires about one year.
Gasterophilus haemorrhoidalis (Linn.) (Fig. 126B) is a North
American and European species. It is commonly known as the "nose
fly," because the female fly forceably "strikes" the animal in the region
of the nose, where it attaches its black eggs to the fine hairs of the lips
or may even thrust the screw-like stalks, with which the eggs are pro-
vided, directly into the tender skin. Because of the orange-red terminal
segments, this fly is also known as the "redtailed bot."
The fully grown larvae have the habit of moving from the stomach
during the early spring and attaching close to the anus before finally
dropping to the ground.
Gasterophilus nasalis (Linn.) [G. veterinus (Clark)] (Fig. 126C) is
the chin fly or throat botfly, also a widely distributed species said to be
especially abundant in the Rocky Mountain region. This fly is very
annoying to horses, since its eggs are attached to hairs under the jaws,
and when the fly darts at the throat, it causes the animals to throw their
heads up as though struck under the chin. Egg deposition takes place
during late spling and early summer. Unlike G. intestinalis (DeGeer)
moisture is not required for the liberation of the larvae. The larvae hatch
in from four to five days. The newly hatched larvae travel along the jaw
and enter the mouth between the lips. There seems to be no tendency
to burrow through the skin of the throat. From the mouth of the horse
the larvae travel to their preferred site in the alimentary tract, the pyloriC
portion of the stomach and the anterior portion of the duodenum, where
they are found in groups and remain for 10 to 11 months, i.e., until they
are mature. Pupation takes place in a few hours after the larvae are
voided with the manure during the early summer. The pupal stage
requires about three weeks.
Gasterophilus inermis (Brauer) is a European species recently re-
ported from North America (Illinois) by Knipling. 36 The eggs are de-
posited on the hairs of the cheeks of the host and, according to Knipling,
when hatched the larvae penetrate the epidermis and work their way
under it until the mouth is reached, thence after molting in the epithelial
layer of the cheek they migrate to the rectum, where they remain until
fully mature. The larvae drop to the ground and pupate as do other bots.
The pupal period is 21 days in the case of Knipling's material. The adult
is small, and "densely covered with silvery to yellowish hair, contrasting
MYIASIS 393
with the more or less orange-colored hair in G. haemorrhoidalis, G.
nasalis and G. intestinalis."
Gasterophilus pl~corum (Fabr.) is a European and African species,
which does not occur in the United States. It is said to deposit its eggs
on the food of the host animal and on nearby objects. The larvae bur-
"!\)'W intI) the mucl)U'i, ill~illh1?We 1)\ th~ ill<;mt\\, ill\'e,\:?ti\\.'e, 'i,1)1)\\' tl) the
stomach and rectum.
Pathogenesis. While a moderate infestation of bots will give no out-
ward indications, a heavy infestation will be indicated by digestive dis-
FIG. 129. Horse bots, Gasterophilus intestinalis, attached to mucous lining of

the stomach of a horse. )(.75. (Photograph by Wherry.)
orders (which may of course be traceable to other causes as well). The
discovery of bots it) the manure is sufficient evidence. A light infestation
is probably of no consequence; there are indeed some individuals who
erroneously maintain that a horse must have at least a few bots in order
to be well.
The injury which bots produce is: (1) abstraction of nutriment, both
from the stomach and its contents; (2) obstruction to the food passing
from the stomach to the intestine, particularly when the larvae are in or
near the pylorus; (3) irritation and injury to the mucous membrane of
394 1lEDlCAL ENTOMOLOGY
the stomach (Fig. 129) due to the penetration of the oral hooklets; (4)
irritation of the intestine, rectum, a,n d anus in passage.
Treatment. Although carbon disulphide had been in use for many
years in Europe as a remedy for horse bots, no wide use was made of
the chemical for this purpose in the United States until after the experi-
mental work of Halp7 in 1917. The treatment should be administered
only by veterinarians. After preparation of the animal by fasting it for
18 hours (water may be allowed), the chemical is administered in gela-
tine capsules at the rate of 1.5 fluid drams for each 250 pounds of weight.
The bots begin to appear in the animal's droppings in Bve or six hours.
Purgatives should not be used in this treatment.
FIG . 130. The common ox warble

fly or heel fly, Hypoderma lineata,
(Adapted after Hearle.)
Cattle grubs (ox warbles) are the larvae of flies belonging to the
family Hypodermatidae, genus Hypoderma, the heel flies. Although the
normal host is cattle, horses and humans are occasionally parasitized.
Persons dealing with cattle are familiar with the tumorous swellings on
the backs of cattle during the late winter and early spring, and most
stockmen have squeezed out the large grubs which inhabit these tumors.
There are two well-known species: Hypoderma lineata (de Villers),
widely distributed in the United States, as well as in Europe and Asia;
and Hypoderma bovis ( DeGeer ), less widely distributed and more
northerly in its occurrence 38 in the United States; it is found particularly
in the New England states, but it does occur in California.
Hypoderma bovis (DeGeer) is the larger of the two species, meas-
uring about 15 mm in length, while H. lineata (de Villers) measures
about 13 mm. The former has the thorax covered with dense yellow
hairs in front and black ones behind, with the terminal yellow hairs on
the abdomen, while the latter, H. lineata (de Villers) (Fig. 130), has a
fairly uniform hairy covering of mixed brownish black and white with
four prominent smooth and polished lines on the thorax, the hairs of
MYIASIS 395
the terminal segment of the abdomen being reddish orange. The fuII-
grown larvae are easily distinguished by examination of the spiny arma-
ture: H. bovis (DeGeer) has the last two segments entirely devoid of
spines, while H. lineata (de Villers) has only the last one smooth. It may
also be said that the full-grown larva of the former measures from 27
to 28 mm in length and that of the latter about 25 mm (Fig. 131).
Life history and habits. The eggs of both species are laid on the hairs
of cattle, H . lineata (de Villers) attaching as many as a dozen in a row
to a single hair, while H. bovis (DeGeer) is said to attach but a single
egg to a hair. As many as 800 eggs, it is stated, may be laid by a female
of either species (Warburton <D ). The eggs are eVidently deposited by
preference on the legs from the hock to the knee of the standing animal,
FIG. 131. Larva or grub of the ox warble

fly, Hypoderma lineata. X 1.3.
but in recumbent animals the eggs may be attached to the hairs of other
parts of the body close to the ground. Although no pain is inflicted at
the time of oviposition, cattle become terror-stricken when the fly is
discovered and gallop madly for water or shade in which to stand to
escape the enemy. This is termed "gadding" and often spreads to the
whole herd.
The eggs of both species hatch within a week and the tiny armored
larvae crawl down the hairs of the host and bore either directly into the'
skin or into the hair follicles .4 0 Knipling'sl1 studies indicate that there
are only three stages, although others have suggested there might be
four or five.
Bishopp, Laake, and Wells 42 state that the eggs are ordinarily de-
posited only on sunny days, although H. bovis (DeGeer) may continue
to oviposit during cloudy periods. A stiff breeze apparently deters the
flies, although egg deposition was observed at temperatures as low as
40° F. The eggs hatch in from three to four days, and the larvae pene-
trate the skin causing considerable irritation. The larvae then work up-
ward between the muscles and in a few months thereafter are found in
the abdominal and chest cavities of the host. The above-mentioned
authors state further, "During the following seven or eight months they
constantly burrow about over the surface of the paunch, intestines,
spleen, and other organs. Grubs are especially numerous between the
muscular and mucous layers of the oesophagus or gullet. The grubs in
these situations are slender and their length ranges from about one-
tenth to about two-thirds of an inch. In the fall, winter, and spring the
grubs migrate through the muscular tissues of the back and in a short
time reach the under surface of the skin. During this last journey some
of them enter the spinal canal and may burrow along the spinal cord
for considerable distances. Soon after the skin is reached the grub cuts
a minute hole through to the surface. At this time it is still slender and
white and about two-thirds of an inch long, and is smooth except for
small spines at each end. From one to five days later the grub molts
for the third time." Upon emerging "from this molt the skin is closely
set with spines. The body of the host now begins to isolate the invading
parasite by forming a pocket or cyst around it. The growth of the grub
from this time on is rather rapid, and a fourth molt occurs about 25
days after the third. In this last stage of its development the color grad-
ually darkens, first becoming yellow, then brown, and finally almost
black. During this entire development beneath the skin a breathing hole
is kept open to the surface, and the grub lies with its two breathing
pores, which are located on the posterior end, applied rather closely to
the opening in the skin. As growth proceeds the hole in the skin is
gradually enlarged." In late spring and early summer "at the end of
the period of development in the back, which requires from 35 to 89
days, growth is complete, and the repulsive, spiny grub works its way
out and falls to the ground." There the larvae crawl away into the loose
earth or debris, becoming rapidly dark brown to black in pupation, and
in from four to five weeks emerge as warble flies. The complete life
cycle requires about a year.
The warbles begin to appear in the backs of cattle in some parts of
California about October first, and Warburton reports mid-February as
the time when the indications of newly forming dorsal tumors are most
numerous in England.
In a summary of his work in Anatolia (Turkey) Kurtpinar43 (1947)
reports that of a total of 7,025 cattle examined 67.3 per cent were in-
fested with Hypoderma bovis (DeGeer). The larvae appeared in the
backs of cattle in January and were found there until the last of June;
adults were found from early May until the end of June. Kurtpinar also
reports that 75 per cent of 1,200 goats examined were infested by a
different species, namely, Hypoderma aegagri Brauer.
MYIASIS 397
Injury done. The injury done by the warbles is first that of irritation
caused by their migrations in the body of the animal and later in their
emergence from beneath the skin; secondly, the escape of the larva
from the tumor leaves an open, running wound which persists for a
long time and is attractive to screwworm flies and other tormenting in-
sects. The direct pathogenesis is of minor irilportance, however, in the
face of the economic loss produced by this irlsect.
Economic losses. The economic losses produced are: (1) Reduction
in the milk secretion of cattle, which is estin1ated at from 10 to 20 per
FlG. 132. A piece of sale leather 21 X 31.5 em, showing work of ox warble.
x.S.
cent of the normal yield. (2) Loss of flesh due to the wild endeavor of
the animals to escape from the flies and the irritating larvae (which is
pointed out by Holstein: "A cow quietly grazing will suddenly spring
forward, throw up her tail, and make for the nearest water at a head-
long gait. Seemingly deprived at the moment of every instinct except
the desire to escape, she will rush over a high bluff on the way, often
being killed by the falL This, with miring in water holes and the fact
that cattle are prevented from feeding, causeS the loss"). (3) Deprecia-
tion of the value of the carcass as flesh, which becomes greenish yellow
and jelly-like in appearance at the points where the grubs are located,
and is not fit for consumption. (4) Injury produced to the hide which
becomes "grubby," full of holes, where the grubs have emerged (Fig.
132).
391>, MEDICAL ENTOMOLOGY
The following is quoted from Tanners' Work for October, 1913:

The case is recorded by Boas of Denmark of a cow which remained in poor
condition and gave 33 pounds of milk per day. Forty-six grubs were extracted
from this animal and eight days later she was giving 44 pounds of milk per
day, continued to do so most of the summer and was in good flesh and condi-
tion in the fall. In this case the loss of milk due to the grub infestation was 25
per cent. The loss in flesh on account of grubs has been variously estimated at
from $1.00 to $5.00 or more per head. If we assume that 25 per cent of all of
the cattle in the United States are more or less infested with grubs, a quite con-
servative estimate, 50 per cent probably being nearer the actual percentage,
the loss in flesh on account of grubs amounts to from $15,000,000 to $75,000,-
000 a year, the total number of cattle in the United States being calculated as
approximately 60,000,000. If we also assume that infested milch cows lose 10
per cent in milk production and that 25 per cent of the 20,000,000 milch cows
in the United States are affected, there should be added to the account a loss
of not less than $30,000,000 per year.
As to the loss in hides it is stated by European tanners that a grubby hide
is, on the average, less in value by one-third than a perfect hide, but for this
country I have no definite information other than that grubby hides in the
green state are commonly valued at one cent a pound less than perfect hides.
On this basis the depreciation in value of a hide of average weight of 65
pounds, if grub-infested, would be 65 cents and the depreciation in the value
of the estimated 15,000,000 grubby cattle of the United States so far as their
hides are concerned thus amounts to $9,750,000. It is, however, quite probable
that the actual loss in the value of hides when made into leather is much
greater than this.
Without including the loss on account of the direct damage to beef car-
casses from the presence of grubs, we may, on the basis of the foregOing, esti-
mate the total loss from grubs in the United States in round numbers at from
$55,000,000 to $120,000,000 per year.
Treatment. Although hand extraction of the warbles is possible in the
case of small herds, the most eHective method is the application of derris
dust or derris washes in various formulations. Where wet applications
are undesirable, dry powder may be used, as follows: equal parts of wet-
table sulfur (325 mesh) and finely ground (200 mesh or finer) cube or
derris powder containing 5 per cent rotenone. One pound of the mix-
ture applied with a dusting can or jar is sufficient to treat from 12 to
18 cattle. In preparing the mixture for application the cube or derris
and wettable sulfur must be thoroughly mixed (Laake 44 ).
For large-scale treatment of cattle, the use of rotenone suspensions
applied by means of power sprayers is recommended by Wells!5 A spray
is made of cube or derris powder (5 per cent rotenone), 5 pounds; wet-
table sulfur, 10 pounds; and water, 100 gallons.<I Application is made
" The most efficient control is obtained with 7112 pounds of 5 per cent rotenone to
100 gallons of water, plus a small amount of a wetting agent. (Douglas, J. R., and
Furman, D. P., 1949. J. Ec. Ent., 42:884-87.)
MYIASIS 399
with a power sprayer, about 350 pounds pressure, directed downward
to the backs of cattle at a distance of about 15 inches. The spray is ap-
plied as the animals pass through a chute. It is estimated that with one
nozzle a herd of 1,000 cattle could be treated in one day.
Hearle 46 reports that derris as a wash has proved effective in large-
scale experiments in several countries, including Canada. The formula
recommended is standardized derris powder, one pound; soft soap one-
quarter pound; water one gallon. The soft soap is boiled in a quart of
water, and when cooled a little is poured into the derris powder in a
bucket and mixed into a paste. Cold water and the remainder of the
soap solution are then added slowly while stirring, to make up one
gallon, and the mixture is ready for use. Standardized derris warble-fly
powders ready for use are sold commercially. Before application the
derris wash must be agitated frequently to ensure a good mixture. Al-
though the keeping qualities are good if the liquid is placed in a well-
stoppered container, it is advisable to prepare only an amount sufficient
for immediate application. Where infestation is heavy, the wash should
be liberally applied to the backs of the animals with a soft cloth or a
worn stable brush, care being taken to cover completely the area affected
by the grubs. In many cases, however, it is more economical to pour a
little derris wash from a bottle on to each cyst, and to rub it in with the
fingers. In the case of animals that are not stall-tied, a crush or dehorn-
ing chute is an aid to handling and treating them.
The date for the first application of the derris wash, varying in dif-
ferent parts of Canada, is in early spring when the swellings in the backs
of the animals caused by the grubs first become conspicuous. In the in-
terior of British Columbia this treatment is given in mid-February; in
the prairie provinces and eastern Canada, about the third week in March.
The second and third applications are made after intervals of 28 days,
and the fourth after a further interval of about 35 days. A fifth dressing
35 days after the fourth is necessary in milder regions such as the in-
terior of British Columbia, where the first application is made in mid-
February. The intervals between the third and fourth, and fourth and
fifth dressings are longer than between the preceding ones, being timed
to accord with the larval development of Hypoderma bovis (DeGeer).
With regard to the treatment of beef herds for H. lineata (de Vil-
lers) during winter and early spring, the main objection of many
ranches is that working cattle through a chute endangers the calf crop.
In our experience these fears are baseless. Ice is a more serious menace,
but the danger to stock from slipping may largely be overcome by sand-
ing the yards. The April treatment can be combined with dehorning, if
this is practiced.
Kurtpinar (lac. cit.) describes a treatment consisting of derris ellip-
tica (Cooper's extract) and 12 per cent hydrogen peroxide plus one per
cent methylene blue which killed the second-stage warbles in one to
two minutes and the third instar in two to five hours; 2,664 cattle having
a total of 17,232 larvae were treated. The average larval kill was 98.2
per cent.
The Caribou warble fly, Oedemagena tarandi (Linn. ), is widely
distributed over the range of its host both in northern Europe and
northern North America. Rearle (lac . cit.) states that the fly is yellowish
orange in color and has a bee-like appearance. The life history resembles
that of the warble fly of cattle.
Read maggots, grub in the head, or sheep bots are the larvae of
Oestrus avis Linn., a very widely distributed species belonging to the
, family Oestridae. The fly (Fig. 133) is abOllt 12 to 14 mm in length,
FIG. 133. The sheep botfly or

head maggot fly, Oestrus avis. (Adap-
ted after Rearle.)
smaller than a honeybee, which it resembles; it is yellow to brownish
gray in color, and hairy. The abdomen is variegated with brown and
straw yellow; the feet are brown. It is further described by Osborn as
follows:
The under side of the head is puffed out and white. The antennae are
extremely small and spring from two lobes which are sunk into a cavity at
the anterior and under part of the head. The eyes are purplish brown, and three
small eyelets are distinctly visible on the top of the head. It has no mouth
and cannot, therefore, take any nourishment. The wings are transparent and
extend beyond the body, and the winglets (calypteres) which are quite large
and white, cover entirely the poisers. It is quite lazy and, except when attempt-
ing to deposit its eggs, the wings are seldom used.
Life history. The "head maggot fly" deposits living young during
early summer to autumn in the nostrils of sheep and goats and may also
attack human beings. One female fly may deposit as many as 60 larvae
in an hour. The larvae at once begin to move up the nasal passages,
working their way into the nasal and frontal sinuses often as far as the
base of the horns in rams and attach themselves to the mucous mem-
MYIASIS 401
branes. Here numbers of these whitish grubs may be found wedged in
closely in various conditions of development. The posterior ends of the
larvae present conspicuous spiracles. The grubs reach full growth with
a length of from 25 to 30 mm by the following spring, a larval period
of from 8 to 10 months. At the end of this time they work their way out
of the nostrils (they are usually sneezed out), fall to the ground, bury
themselves in the earth, and pupate in a few hours. The pupal period
lasts from three to six weeks and over, 19 to 34 days according to Fallis47
who found that reared flies lived on an average of 16 days, one surviving
as long as 28 days. Fallis also found that the complete development of
the parasitic stage in spring lambs in Texas and New Mexico required
from 2Y2 to 3Y2 months.
Symptoms. In the presence of the fly the sheep (or goats) are very
much excited, shake the head, rush with their noses between their
fellows, push their noses into the dust, snort, and otherwise indicate
that they are trying to escape something that persists in entering the
nostrils. Once infected there is a purulent discharge from the nostrils,
vigorous shaking of the head, and perhaps the occasional discharge of a
maggot, loss of appetite, grating of the teeth; and, when the animal
walks, the fore feet are lifted in a pawing movement. The great majority
of the cases do not result fatally, but death often comes in a week or
less after the appearance of aggravated symptoms.
Grub-in-the-head is distinguished from "gid," caused by a larval
tapeworm, Multiceps multiceps (Leske) [= Coenurus cerebralis
(Batsch) ], in that the former is always associated with purulent dis-
charges from the nostrils, absent in the latter, that the symptoms of the
former appear during the summer, and that the latter occurs ordinarily
in lambs and yearlings only. There is no undue sneezing or rubbing of
the nose in gid. Because of the mucus exuding from the nostrils of the
sheep, the infection is commonly known as "snotty nose."
Treatment. Materials such as snuff, pepper, etc., may be introduced
into the nostrils or sprinkled among the flock, to induce violent sneezing,
which causes the expulsion of many of the larger grubs. Law recom-
mends the injection of benzine, lifting the sheep's nose somewhat and
pouring a teaspoonful of the remedy into each nostril. The lower nostril
into which the benzine is poured is held shut for thirty seconds; the
other side is then turned and the treatment repeated. The application is
repeated daily or more often until the maggots are all expelled.
Prevention. "Salt log~" are used in sheep pastures by some sheep
raisers. These logs are made by boring 2-inch holes at intervals of about
6 inches along the length on top. Salt is placed in the holes, which are
kept about half full, and in turn the edges of the holes are repeatedly
smeared with pine tar, or other repellent material. In endeavoring to
reach the salt the sheep involuntarily smears its nose with the sub-
stance, which protects it to some extent against the head maggot Hy.
Head maggot of horses. An important species of head maggot attack-
ing horses in Russia and parts of Europe and in Egypt is Rhinoestrus
purpureus (Brauer). Its habits are said to be similar to those of Oestrus
ovis Linn. Like other species of related genera it may attack man either
in the nose or eye.
Head maggot of deer. Deer, reindeer, elk, and other related wild
animals are commonly infested with head maggots (Fig. 134); among
these are the European species Cephenemyia stimulator Clark in the
roe deer, C. rllfiu({rbis Meig. in the red deer, C. ulrichii Brauer in elk,
and C. trompe Linn. in reindeer, also in Alaskan reindeer. Among the
!
'r
f
FIG. 134. Head maggots attached to tissue in nasal sinuses of the deer. X .6.
known species in the United States besides C. trompe are C. phobiger

Clark (a robust black species with yellow pile on thorax; legs black)
from the white-tailed deer (Odocoileus virginianus), eastern United
States; C. pratti Hunter from black-tailed deer (Odocoileus hemionus)
in the western United States, often at high elevations; C. fellisoni Town.
from the PaciRc coast black-tailed deer (Odocoileus hemionus colum-
bianus ).
Dermal creeping myiasis of man is commonly caused by the wander-
ing larvae of flies belonging to the family Cuterebridae, particularly
Dermatobia hominis (Linn.); and also Hypoderma lineata (de Villers)
and Hypoderma bovis (DeC.), both known as ox warble flies, belonging
to the family Oestridae.
Dermatobia hominis (Linn.) (Family Cuterebridae,) is commonly
found in Central and South America and Mexico. The larva is known
in its early stage as ver macaque and in its later stages as torcel or berne.
The fly measures from 12 to 16 mm in length and is entirely brown in
color. This fly parasitizes a large number of species of mammals and
even birds. It has been found in cattle, pigs, dogs, mules, monkeys, man,
MYIASIS 403
and various wild animals. In man the larva llas been reported from
various regions of the body, mainly head, arm, \)ack, abdomen, scrotum,
buttocks, thigh, and axilla.
Although it is not certain that this Dermatdbia does not deposit its
eggs directly on or in the human skin, it is rlOW known that several
species of mosquitoes, bloodsucking flies, anet ticks, particularly the
mosquito Psorophora (Janthinosoma) lutzii (Tlleo.) and possibly other
insects act as carriers of the eggs. The femalf.). Dermatobia is said to
oviposit on the undersides of the bodies of the :mosquitoes so that when
the latter suck blood, it is possible for the eggs to come in contact with
the warm-blooded host where either contact Of warmth stimulates the
larvae; rapid emergence results and entrance to the skin of the host is
effected. The larval period in the body of the host is said to require
about two and a half months when, like the Hypoderma species, the
larvae leave the tumorous swellings they hav~ produced, drop to the
'C>'di\\l, 'd1).~ P'd':>'5 \\lmllg'D. a pllpa\ pei)'\)~ iel.l~;''.i;''i'q, \'.i';Y.i'i', I-.\>mrc I-,~ c:';s..
weeks.
Dunn48 (1930) has described the life history of the human botfly
most accurately as the result of an infestation which he suffered in the
Panama Canal Zone. In his case the fly Limnophora, not a bloodsucker,
was the vehicle for the eggs. Two larvae we(e observed to enter the
skin of his arm, requiring 42 minutes for the Brst and one hour and 35
minutes for the second. Dunn experienced "absolutely no sensation
caused by the entrance of the (first) larva until after the first 30 min-
utes. Then, as the posterior end was being drawn inside, a sharp prick-
ing, which lasted for about two minutes, was experienced." He states
that there was at first a sharp itching at night, and by the end of two
weeks the lesions had the appearance of small boils and by the end of
three weeks these were excruciatingly painful. fit the end of 46 days and
15 hours and 50 days and 1572 hours respectively the larvae emerged
from the skin, causing "absolutely no pain or sensation." The pupal
periods were from 22 to 24 days.
The foregOing account of an infestation of human bats supports the
opinion of other authors that the larvae remain in a relatively fixed spot
in the subcutaneous tissues. The larvae of the ox botfly on the other
hand have the tendency to migrate in the subcutaneous tissues often for
considerable distances.
The author49 ( 1925 ) described the migrations of the larvae of
Hypoderma bovis (DeC.) (Family Hypodermatidae) as follows:
Mr. C. is a ranch superintendent, spending much of his time on horseback.
On a Sunday toward the end of July, 1924, he had ridden to a point known as
Mission Ridge near San Jose, when feeling tired, he dismounted and lay down
on the ground in the open and slept. He distinctly remembers that his shirt had
rolled up above his belt, exposing his skin, but felt no irritation at the time.
Whether this exposure was taken advantage of by the fly can only be a matter
of conjecture. Several days later, exact time not remembered, soreness was ex-
perienced and a slight swelling in the region of the right groin appeared. In
about a week the swelling had increased to the width of a hand with no dis-
coloration. The swelling then crept downward toward the left side affecting the
scrotum, thence downward along the left leg to the knee and calf, thence back
up the left leg following about the same course to the left groin, thence across
to the right groin and back again to the left and upward along the left side of
the body, slightly anterior to the shoulder, thence downward to the upper
right arm to near the elbow, when the arm could not be raised without great
pain, thence the swelling travelled upward again to the neighborhood of the
shoulder blade where a "hive-like" local swelling was formed without any
itching sensation. Mr. C. stated that at this point he was "bothered" all night,
and while rubbing his arm and manipulating his shoulder muscles a larva of
some insect "popped" out. This emergence took place about the end of Octo-
ber (1924). The larva was placed in a vial for shipment but was lost in transit.
Relative to the second larva which was delivered to the writer in person
on the day following its emergence, Mr. C. states that since October when the
first larva emerged, no further swellings were observed, but soreness in the
region of the thigh and lower abdomen, similar to severe strain, persisted.
However, on January 28 (1925) he experienced a severe "soreness" in the
region of his right thigh which gave much distress, particularly when walking.
By that night a swelling had developed and the following day the muscular
soreness continued to spread; by January 31 a hernia-like swelling had devel-
oped which enlarged upward and outward to the region of the belt-line, the
lower hernia-like swelling gradually disappearing. Sunday night, February 1, a
hive-like swelling as observed in the case of the first larva began to form, en-
larging to an area of about four by eight inches. Tuesday evening, February 3,
lymph exuded from a small opening near the middle of the swollen area. About
a tablespoonful of lymph stained with blood was pressed out and in the proc-
ess of manipulation a larva similar to the first "popped" out. This specimen
was delivered to the writer February 4 in good condition and identified as a
third-stage larva of Hypoderma bovis DeG. The larva was milky white in
color, about 12 mm. in length by 2 mm. in width at the middle, tapering
bluntly at both ends. Very little swelling and practically no discoloration were
visible on examination, although the point of emergence was clearly seen.
Gasterophilidae as human parasites. The larvae of the horse botflies
burrow freely, as already explained, and may cause a form of creeping
cutaneous myiasis in human beings. Gasterophilus intestinalis (DeGeer)
is the species usually involved. 50 The course made by these creeping
larvae is quite tortuous and plainly visible. The infection causes severe
itching. The larvae, which measure from 1 to 2 mm in length, can be
easily extracted surgically.
Ophthalmomyiasis of man is more particularly traceable to the larvae
of he~d-maggot flies of sheep, deer, and related animals. Cases traceable
MYIASIS 405
to Oestrus ovis Linn. [Cephalomyia ovis (Linn.)] of sheep and Rhin-
oestrus purpureus (Brauer) are frequently reported in European horses.
Three first-stage larvae of Oestrus ovis Linn. measuring about 1 mm in
length that had been removed from the eye of a patient in Honolulu by
Dr. R. Faus were studied by the author 51 (Fig. 135). The attending
physician reported that the three larvae were buried in the sclera and
were extremely adherent to the conjunctiva, causing acute conjunctivitis,
larchrymation, ulceration, and neurosis.
The larvae of Oestrus ovis Linn. at times invade the nasal cavities
of man as is normally the case with sheep. Severe frontal headaches
result.
Rodent bats. The larvae of the family Cuterebridae are commonly
parasitic upon rodents, wild and domestic rabbits, and mice, which may
FIG. 135. Larva of Oestrus ovis from eye of human.
be severely infested with dermal tumors in which the large grubs lie.
The adult flies are robust and bumblebee-like, having the scutellum
elongate, the arista plumose or pectinate, the oral opening large, and
the palpi small. There are four genera in this family, one of which in-
cludes the human botfly, Dermatobia hominis (Linn.) previously des-
cribed; the other three, Cuterebra, Pseudogametes, and Rogenhofera,
are parasitic on rodents and rabbits, and also on some other animals, in-
cluding cats ~nd dogs. Cuterebra horripilam Clark and C. buccata Fabr.
usually infest rabbits; C. peromysci Dalmat infests white-footed mice
(see Dalmat52 for life cycle); C. tenebrosa Coq.; C. beameri Hall para-
sitizes pack rats; C. emasculator Fitch is the emasculating scrotum-
inhabiting bot of squirrels.
Surgical maggots. Although now largely discontinued in favor of
other treatments, the use of sterile maggots, maggot therapy, in the dis-
infection of osteomyelitis and other infected wounds :was introduced
into professional medical practice by Baer shortly after the end of

World War 1. Baer53 had noticed that when men wounded in battle had
been lying out on the ground for some time before being carried into
dressing stations, their wounds were infested with maggots. He noticed
particularly that these men whose wounds were crawling with maggots
did not develop infections, as did the men whose wounds had received
early treatment. It was discovered that the maggots were eating the dead
tissue in which the bacterial infection throve; the maggots actually
served as a "viable antiseptic." Baer's work attracted a great deal of
attention and much experimentation followed, resulting in numerous
publications by many investigators.
In 1932 Livingston and Prince 54 reported that filtered, uncontami-
nated products derived from the bodies of larvae in culture, when
brought into contact with pyogenic organisms in petri dishes, destroyed
the cultures.
The fly larvae used in earlier osteomyelitis treatment apparently be-
longed indiscriminately to the following species, namely, Phaenicia
sericata (Meig.), L. caesar (Linn.), and Phormia regina (Meigen). It
was assumed that all these species fed only on dead tissues. Stewart55 has
shown that even Phaenicia sericata (Meig.) larvae, which have been most
commonly used in practice, will establish themselves in and feed upon
normal healthy tissue, although they prefer necrotic tissue. He warns
that they, and probably the larvae of Phormia regina (Meigen), Lucilia
caesar (Linn.) and Wohlfahrtia nuba (Wiedemann), are potentially
dangerous to normal tissue and must be utilized with care by an experi-
enced person.
Stewart56 also came to the conclusion that not only do the scaveng-
ing activities of the maggots play an important role in the successful
results obtained, but the calcium carbonate, which was found to be con-
stantly exuded by the larvae, is also of importance because of its prop-
erty of alkalinizing the wound and of markedly increasing phagocytosis.
Robinson in his later investigations discovered that allantoin and urea
are present in maggot excretions, and that both have good effect in the
treatment of osteomyelitis; however, because of its low cost and high
solubility urea is now generally used, thus largely disposing of the use
of maggots.
BIBLIOGRAPHY
1. James, Maurice T., 1947. The Flies That Cause Myiasis in Man. Wash-
ington, D. C.: Dept. Agric., in Misc. Publ., no 631. 175 pp.
2. Braun, M., und Seifert, 0., 1926. Die tierischen Parasiten des Men-
schen. Leipzig: Curf Kabitzsch. x + 608 pp.
3. Hoeppli, R., and Watt, John Y. C., 1933. "Experiments on resistance
MYIASIS 407
of ~dipterous larvae in connection with the problem of intestinal and urinary
myiasis," Chinese MI, 47:1298-1306.
4. Desai!, P., et Delhaye, R., 1922. "Contribution a la pathogenie des
myiases intestinales par l'etude de la resistance des oeufs et larves de Calli-
phorees aux agents physiques et chimiques intervenant dans Ie tube digestif."
Compt. rend. Soc. de bioi., 87:1098-1100.
5. Causey, O. R., 1938. "Experimental intestinal myiasis," Am. J. Hyg.,
28:481-86.
6. Herms, W. B., and Gilbert, Q. 0., 1933. "An obstinate case of in-
testinal myiasis," Ann. Int. Med., 6:941-45.
7. Parker, G. H., 1922. "Possible pedogenesis in the blowfly, Calliphora
erythrocephala Meigen," Psyche, 29:127-31.
8. Simmons, Perez, 1927. The Cheese Skipper as a Pest in Cured Meats.
Washington, D. C.: Dept. Agric., in Dept. Bull., no. 1453.55 pp.
9. Meleney, H. E., and Harwood, P. D., 1935. "Human intestinal myiasis
due to the larvae of the soldier fly, Hermetia illucens Linne (Diptera, Stratio-
myidae)," Am. J. Trap. Med., 15:45-49.
10. Hall, M. C., and Muir, J. T., 1913. "A critical study of a case of
myiasis due to Eristalis." Arch. Int. Med., 2:193-203.
11. Hall, M. C., 1918 .• "A note regarding myiasis, especially that due to
syrphid larvae," Arch. Int. Med., 21:309-12.
12. Chevril, R., 1909. "Sur la myiase des voies urinaires," Arch. de
Parasitol., 12:369-450.
13. Mumford, E. P., 1926. "Three new cases of myiasis in man in the
north of England," Parasitology, 18:375-83 .
., 14. Stewart, M. A., 1929. "A case of dermal myiasis caused by Phormia
regina Meig.," J.A.M.A., 92:798-99 .
.- 15. ---, 1934. "A new treatment of traumatic dermal myiasis," J.A.M.A.,
103:402.
16. Cushing, E. C., and Patton, W. S., 1933. "Studies on the higher Dip-
tera of medical and veterinary importance. Cochliomyia americana sp. nov.,
the screwworm fly of the New World," Ann. Trap. Med., 27:539-51.
17. Laake, E. W., 1936. "Economic studies of screwworm flies, Coch-
liomyia species (Diptera, Calliphorinae), with special reference to the pre-
vention of myiasis of domestic animals," Iowa State Call. J. Sc., 10:345-59.
18. Gajardo-Tobar, R., and Honorato, Armando, 1947. "Anotaciones acerca
de una epidemia de miasis humana," Hospital de Viiia del Mar, Chile, 3:5-14.
-./ 19. Walker, E. M., 1922. "Some cases of cutaneous myiasis, with notes
on the larvae of Wohlfahrtia vigil (Walker)," J. Parasitol., 9:1-5 (3 plates).
v 20. Calero, Carlos, 1948. "Cutaneous myiasis in Panama," ]. Parastiol.,
34:343-44.
21. Cushing, E. C., and Hall, D. G., 1937. "Some morphological differ-
ences between the screwworm fly, Cochliomyia americana C. and P., and
other closely allied or similar species in North America (Diptera: Calli-
phoridae)," Proc. Entomolog. Soc. Wash., 39:195-98 (2 plates).
22. Bruce, W. G., and Sheely, W. J., 1944. Screwworms in Florida. Gaines-
ville: Univ. Florida in Agric. Extension Servo Bull., no. 123, 28 pp.
,.r 23. Blacklock, B., and Thompson, M. G., 1923. "A study of the tumbuBf,
Cordylobia anthropophaga Griinberg, in Sierra Leone," Ann. Trop. Med.,
17:443-501 + 4 plates.
24. Roubaud, E., 1913. "Recherches sur les Auchmeromyies," Bull. scient.
France et Belgique, 47:105-202.
25. Frogatt, W. W., 1922. Sheep-Maggot Flies. Dept. Agric., New South
Wales, Farmers' Bull., no. 144.32 pp.
26. Babcock, O. G., and Bennett, D. H., 1921. The Screwworm and the
Wool Maggot. Texas Agric. Exper. Sta., in Circ. no. 27. 15 pp.
27. Belschner, H. G., 1937. A Review of the Sheep Blowfly Problem in
New South Wales, (Studies on the Sheep Blowfly Problem). Sydney: Dept.
of Agric., New South Wales: pp. 1-60.
28. Woodworth, H. E., and Ashcraft, J. B., 1923. "The foot maggot,
Booponus intonsus Aldrich, a new myiasis-producing fly," Philippine]. Sc.,
22:143-56 (8 plates).
29. Bengston, Ida A., 1923. "A toxin-producing anaerobe isolated prin-
cipally from fly larvae," U. S. Public Health Service, Pub. Health Rep., 38:340-
44.
30. Plath, O. E., 1919. "A muscid larva of the San Francisco Bay region
which sucks the blood of nestling birds," Univ. Calif. Pub. in Zool., 19:191-
200.
31. ---, 1919. "The prevalence of Phormia azurea Fallen (larva parasitic
on nestling birds) in the Puget Sound Region and data on two undescribed
flies of similar habit," Ann. Entomolog. Soc. Amer., 12:373-81.
32. Storer, T. I., 1929. "Protocalliphora in the nest of a mountain chick-
adee," The Condor, 31:227.
33. Guberlet, John E., and Hotson, H. H., 1940. "A fly maggot attack-
ing young birds, with observations of its life history," The Murrelet, 21:65-
68.
----
34. Wells, K. W., and Knipling, E. F., 1938:, "A report of ~9me recent
studies on species of Gastrophilus occurring in horses in the United States,"
Iowa State Coll. J. SC. 12:181-203.
35. Wehr, Everett E., 1933. "The life history of Gastrophilus larvae of
the horse and lesions produced by the larvae," Cornell Veterinarian, 23:254-71.
36. Knipling, E. F., 1935. "Gastrophilus inermis Brauer, a species of horse
bot not previously recorded from North America," Entomological News,
46:105-7.
37. Hall, M. C., 1917. "Notes in regard to bots, Gastrophilus spp.," J.
Am. Vet. M. A., 51 n.s., 5:177-84.
38. Bishopp, F. C.; Laake, E. W.; Brundrett, H. M.; and Wells, R. W.;
1926. The Cattle Grubs or Ox Warbles: Their Biologies and Suggestions for
Control. Washington, D. C.: Dept. Agric., in Dept. Bull., no. 1369. 119 pp.
or' 39. Warburton, Cecil, 1922. "The warble flies of cattle," Parasitology,
14:322-41.
• 40. Hadwen, S., 1915. "Warble flies-a further contribution on the biology
of Hypoderma lineatum and Hypoderma hovis," Parasitology, 7:331-38 (2
plates) .
MYIASIS 409
41. Knipling, E. F., 1935. "The larval stages of Hypoderma lineatum
de Villers and Hypoderma bovis DeGeer," J. Parasitol., 21:70-82.
42. Bishopp, F. C.; Laake, E. W.; and Wells, R. W.; 1929. Cattle Grubs
or Heel Flies with Suggestions for Their Control. Washington, D. C.: Dept.
Agric., in Farmers' Bull., no. 1596. 22 pp.
43. Kurtpinar, Hasip, 1947. Anadolu ehU hayvanlarinda goriilen Hypo-
derma nevileri, iktisadi onemi ve miiscadelesine dair en uygun tedbirler
iizerine ara§tirmalar. (Species of Hypoderma found in domesticated animals
of Anatolia; their economic importance and most suitable control measures.)
Qali§malar: 153, T. C. Tarim Bakanligi Ankara Yiisek Ziraat Enstitiisii. 60 pp.
44. Laake, E. W., 1942. "Dry application of cube or derris combination
with wettable sulfur for the control of cattle grubs," J. Econ. Entomol., 35:112-
13.
45. Wells, R. W., 1942. "The use of power sprayers in the control of cattle
grubs," ]. Econ. Entomol., 35:112-13.
46. Hearle, Eric, 1938. Insects and Allied Parasites In;urious to Livestock
and Poultry in Canada. Dominion of Canada, Dept. Agric., in Publ., no. 604.
108 pp.
47. Fallis, A. Murray, 1940. "Studies on Oestrus avis L.," Canad. J. Re-
search, 18:442-46.
48. Dunn, L. H., 1930. "Rearing the larvae of Dermatobia hominis Linn.,
in man," Psyche, 37 :327-42 (1 plate).
v 49. Herms, W. B., 1925. "A case of human myiasis caused by the ox-
warble, Hypoderma bovis DeC.," J. Parasitol., 11:149-50 .
./50. Bedford, C. V.; Williams, D. H.; and Newton, M. V. B.; 1933. "Creep-
ing eruption, with special reference to cutaneous myiasis and report of a case,"
Canad. M. A. ]., 28:377-82.
51. Herms, W. B., 1925. "Ophthalmomyiasis in man due to Cephalomyia
(Oestrus) ovis Linn.," J. Parasitol., 12:54-56.
52. Dalmat, Herbert T., 1943. "A contribution to the knowledge of the
rodent warble Hies (Cuterebridae) ," J. Parasitol., 29':311-18.
53. Baer, W. S., 1931. "The treatment of chronic osteomyelitis with the
maggot (larva of the blowfly)," J. Bone & Joint Surg., 13:438-75.
54. Livingston, S. K., and Prince, L. H., 1932. "The treatment of chronic
osteomyelitis; with special reference to the use of the maggot active principle,"
J.A.M.A.,98:1143.
55. Stewart, M. A., 1934. "The role of Lllcilia sericata Meig. larvae in
osteomyelitis wounds," Ann. Trop. Med., 28:445-60.
56. ---, 1934. "A new treatment of osteomyelitis: Preliminary report,"
Surg. Gynec. & Obst., 58:155-65.
CHAPTER XVIII
._.
LOUSE FLIES
FAMILY HIPPOBOSCIDAE
Order Diptera (Series Pupipara)
Characteristics of Hippoboscidae. Three families of flies constitute the
Pupipara: (1) Hippoboscidae (louse flies), (2) Nycteribiidae (bat tick
flies), and (3) Streblidae (bat flies). The bloodsucking parasitic flies
belonging to the family Hippoboscidae are readily recognized as Diptera
when winged. The larvae are retained within the body of the female,
being nourished by special glands within the mother until time for pupa-
tion is reached; they are then extruded and pupation quickly follows,
whence the term, "pupipara." The adult flies are described as follows by
Williston: "Head Battened, usually attached to an emargination of the
thorax; face short, palpi forming a sheath for the proboscis, not pro-
jecting in front of the head; antennae inserted in pits or depressions
near the border of the mouth, apparently one-jointed, with or without
a terminal bristle or long hairs. Eyes round or oval, ocelli present or
absent. Thorax Battened, leathery in appearance; scutellum broad and
short. Halteres small or rudimentary. Abdomen sac-like, leathery in ap-
pearance, the sutures indistinct. Legs short and strong, broadly separated
by the sternum; tarsi short; claws strong and often denticulated." The
members of this. family are all parasitic in the adult stage upon birds or
mammals. There are about 400 species widely distributed throughout the
world. They range in size from 2.5 to 10 mm.
The sheep "tick" or ked, Melophagus ovinus (Linn.), is a wingless
bloodsucking species, reddish brown in color, about 5 to 7 mm in length.
It is a world-wide parasite of sheep and goats. The head is short and
sunken into the thorax, the body sac-like, leathery, and spiny (Fig. 136).
Life history. The eggs are retained and hatch within the body of the
female ked, where the larvae develop in about seven days and are
extruded fully grown ready to pupate. The extruded larva pupates dur-
ing the course of a few hours, becoming chestnut brown in color; the
secretion with which it is covered hardens and serves to glue the pupa
firmly to the wool of the host. The pupae are commonly found on in-
410
LOUSE FLIES 411
fested animals in the region of the shoulders, thighs, and belly. Pupae
may be found on sheep at all times of the year, though the time re-
quired for development in the winter is longer than in the summer.
Swingle," who has observed this insect very carefully, states that pupae
require from 19 to 23 days to hatch in the summer, whereas 19 to 36
days are requ~ the winter on sheep kept in the barn and prob-
ably 40 to 45 days on sheep out of doors. The time required for the
females to reach sexual maturity is from 14 to 30 days and over, when
they begin extruding young at the rate of one about every seven to eight
days. Swingle (loc. cit.) considers about four months as the average life
of the insect, during which time from 10 to 12 pupae are deposited.
The entire life of the ked is spent on its host; when off the sheep
the insects die in from two to eight days, the majority in about four days.
FIG. 136. The sheep "tick" or

louse fly, Melophagus ovinus. (Left)
pupa, (right) adult. X 4.5.
Damage done. The presence of a few louse flies on the body of a
sheep does not materially affect it. Ordinarily the presence of the insect
is indicated by the fact that the animal rubs itself vigorously, bites the
wool, and scratches. Badly infested animals show emaciation and gen-
eral unthriftiness. The injury to lambs is especially marked.
Control. Since the principal time for migration from the sheep to the
lambs is at shearing when the insects are removed from the hosts with
the wool, it is wise to take particular pains at this time to store the wool
at some distance from the lambs. Inasmuch as the "ticks" die within a
week when away from the host and cannot well crawl any great distance,
this suggestion is well worth considering. Swingle states that "sheep
free from 'ticks' can be kept for months beside a heavily infested one
with a tight partition only three feet high between them without be-
coming infested. . . . A bunch of females placed in the wool of a sheep
will be found in practiGaIly the ~ame place for two days. Males, how-
ever, are most inclined to migrate." A flock of sheep once freed from
"tick" can therefore be kept clean unless infested animals are introduced.
The writer has reasons to doubt the efficiency of "lime-sulphur"
sheep-dip for the sheep "tick." Tobacco dips when used in 0.07 per cent
solution will eradicate sheep "tick" according to Imes,2 if two dippings
are given with an interval of 24 to 28 days between them.
Louse flies of deer. Lipoptena depressa (Say) and Lipoptena subulata
Coq. (= L. ferrisi Bequaert) are common parasites of deer in North
America. These species are smaller than Melophagus ovinus (Linn.), but
otherwise resemble it; they are wingless when established on the host
but have well-developed filmy wings on emergence from the pupal
stage (Fig. 137). The parasites have been found in chains, three or four
individuals attached to one another, the first fly drawing blood from the
host, the second with its proboscis thrust into the abdomen (dorsally)
of the first, the third drawing on the second, and so on to the last in-
r"'·""-·~·-
I
FIG. 137. Louse fly of the deer,

Lipoptena depressa, showing wingless
I _ _J and winged forms. X 5.
dividual. Lipoptena cervi (Linn.), known as the "deer ked," is reported
to be a common species on European deer, and according to Bequ~ert3
has become naturalized in a few localities of the northeastern United
States on the Virginia white-tailed deer. Lipoptena mazamae Rondani
occurs on deer in South and Central America and in the southeastern
United States.
Concerning Lipoptena depressa (Say), Hare 4 states that it is a com-
mon parasite of deer (genus Odocoileus) in western North America. The
pupating larvae are smooth and clean, dropping from among the hairs
of the host to the ground, where the entire pupal period is passed. At
emergence the imago possesses fully developed though fragile wings
and flies among the trees in the woodland haunts of the host. Upon
reaching a deer these winged forms (volants) immediately crawl be-
tween the hairs and begin sucking blood. Here they remain as permanent
parasites for the rest of their lives, soon loosing the wings by a simple
process of wear. Under natural conditions the volants of L. depressa
may be expected to live and remain active without a host for only one
to eight days. Bloodsucking beginS the first day after contact with the
host, and sexual maturity is reached after about 12 days of feeding.
After copulation the first larva is deposited by the female on the sixteenth
to nineteenth day and thereafter singly at three-day intervals, up to 35
mature larvae. Adults of both sexes live up to four months on the host.
LOUSE FLIES 413
By actual count (HermanS) 1,350 Hies have been reported from one
host. Hare states that his observations indicate that the seasonal peak in
California may extend into late fall in some localities; he found these
Hies abundantly on the wing on October 5 (1948) in Monterey County.
THE GENUS HIPPOBOSCA
Nine species of the genus Hippobosca are recognized as valid by
Bequaert. 6 The wings are always well developed in the genus and are
functional throughout adult life. With the exception of the ostrich louse
By, Hippobosca struthionis O. E. Janson, the species of this genus are
ectoparasites of mammals. Except for H. struthionis O. E. Janson, an
abundant parasite of the ostrich in South Africa, host speci:6.city is not
pronounced.
Hippobosca equina Linn. is a common species in England and is
known as the "forest-Hy." It is usually found on horses, mules, and don-
keys, sometimes on cattle and other animals. H. rufipes v. Olfers is also
primarily a parasite of equines and occurs in South Africa.
Hippobosca longipennis Fabr. (= H. capensis v. Olfers) is a louse
Hy reported by Bequaert to be commonly found on domestic dogs,
espeCially on the pariah dog of India. It is also common in many parts
of the Mediterranean region.
Hippobosca variegata von M iihlfeld (= H. maculata Leach) occurs
on domestic cattle and equines and is widespread in distribution. H.
fulva Austen off the hartebeest is known only from its type locality,
northeastern Rhodesia. H. hirsuta Austen is reported to be a parasite of
the water bucks and allied antelopes of Africa.
H. camelina Leach is a parasite of the camel and dromedary. H.
martinaglia Bedford, found on antelopes in South Africa, is recognized
as valid by Bequaert.
LOUSE FLIES OF BIRDS
The pigeon Hy, Pseudolynchia canariensis (Macq. ) [= Lynchia
maura (Bigot) = Olfersia maura Bigot = Lynchia lividicolor Bigot],
is an important parasite of domestic pigeons throughout the tropics and
warmer regions of the world. It is found throughout the southern
United States and California. The dark brown Hies have long wings, 6.5
to 7.5 mm, and are able to Hy swiftly from the host but usually alight
near by. They move about quickly among the feathers of the host and
bite and suck blood from parts that are not well feathered.
The mature larvae, at first pale yellow and later jet-black in color,
are deposited on the body of the bird while it is quiet, but they soon
roll off and collect in the nests. Bishopp7 gives the duration of the pupal
stage at from 29 to 31 days when the mean daily temperature is about
414 MEDICAL ENTOMOLOGY"
73° F. Thus the thorough and regular cleaning of the nests at intervals
not to exceed 25 days is probably the most important single step in con-
trol. The pupae are very resistant, hence ordinary insecticides are of
little use. Bishopp states that "one of the most effective and easily
applied treatments for squabs is fresh pyrethrum powder, one to three
pinches (depending upon the size of the squab) scattered among the
feathers."
In addition to its evil effects as a bloodsucking parasite, the pigeon
fly is the vector of pigeon malaria caused by Haemoproteus columbae
Celli and San Felice. 8
Bequaert9 reports that Microlynchia pusilla (Speiser) is probably the
most widely spread hippoboscid fly of doves. It is a strictly New World
species; Stilbometopa podopostyla Speiser, also a New World species,
has been taken from doves. Ornithoctona erythrocephala (Leach) has
been reported from both wild doves and the domestic pigeon, as well
as from birds of prey and others.
Pseudolynchia brunnea (Latreille) is regarded as a distinct species
by Bequaert and is a parasite of birds of the nighthawk family. It is very
dark brown in color, often nearly black.
Lynchia hirsuta Ferris is a common and abundant parasite of the
Californian valley quail, Lophortyx californica californica Shaw, and has
been shown by O'Roke10 to be a vector of quail malaria caused by
Haemoproteus lophortyx O'Roke. Lynchia fusca (Macquart) is a para-
site of the owl, Bubo virginianus pacificus Cassin, in California and has
been experimentally shown to be a vector of quail malaria by Herms
and Kadner.ll The flies feed readily on quail and deposit their mature
larvae freely on these birds. The incubation period of the infection in
the fly was found to be from 9 to 13 days and in the quail about 25 days.
Stilbometopa impressa (Bigot) is also a parasite of the Californian
valley quail. 12 Lynchia americana (Leach) is characteristically a parasite
of owls in North AmericaY
Bat flies are pupiparous bloodsucking parasites belonging to the
family Streblidae; they are all parasitic on bats in tropical and sub-
tropical climates. The members of the family may be separated from
the Hippoboscidae by the large leaf-like palpi which project in front
of the head and do not form a sheath for the proboscis. They differ from
the Nycteribiidae (the spider-like bat flies) in that they do not have the
head resting in a groove on the dorsum of the thorax. Little is known
about the life history of these insects. The species of the family Stre-
blidae have been reviewed by Kessel. 14
Spider-like bat flies belong to the family Nycteribiidae. They are very
small (2 to 3 mm long) wingless spider-like parasites of bats. Except
for a very few species described from North and South America, they
LOUSE FLIES 415
are primarily parasites of Old World bats. Ferris 15 has reviewed the New
World species.
BIBLIOGRAPHY
1. Swingle, Leroy D., 1913. The Life History of the Sheep Tick, Melo-
phagus ovinus. Laramie: Univ. of Wyoming, Agr. Exper. Sta. Bull., no. 99.
2. rmes, Marion, 1932. The Sheep Tick and Its Eradication by Dipping.
Washington, D. C.: Dept. Agric., in Farmers' Bull., no. 798. 22 pp.
3. Bequaert, J., 1937. "Notes on Hippoboscidae, 5, The American species
of Lipoptena," Bull. Brooklyn Entomolog. Soc., 32:91-101.
4. Hare, John Edward, 1945. "Flying stage of the deer louse-Hy, Lipa-
ptena depressa (Say), in California," Pan-Pacific Entomologist, 21 :48-57.
5. Herman, Carlton M., 1945. "Hippoboscid Hies as parasites of game
animals in California," California Fish and Game, 31:16-25.
6. Bequaert, J., 1939. "Notes on Hippoboscidae, 13. A second revision of
Hippoboscinae," Psyche, 46:70-90.
7. Bishopp, F. C., 1929. "The pigeon Hy, an important pest of pigeons
in the United States," J. Econ. Entomal., 22:974-80.
8. Huff, C. C., 1932. "Studies on Haemoproteus of mourning doves,"
Am. J. Hyg., 16:618-23.
9. Bequaert, J., 1939. "Hippoboscid flies from North American doves,"
Science, 89:267-68.
10. O'Roke, E. C., 1930. "The morphology, transmission, and life history
of Haemoproteus lophortyx O'Roke, a blood parasite of the California valley
quail," Univ. Calif. Publ. in Zool., 36:1-50.
11. Herms, W. B., and Kadner, C. C., 1937. "The louse fly, Lynchia fusca,
parasite of the owl, Bubo virginianus pacificus, a new vector of malaria of the
California valley quail," J. Parasitol., 23:296-97.
12. Herman, Carlton M., 1944. "Notes on the pupal development of
Stilbometopa impressa," J. Parasitol., 30:112-18.
13. Ferris, C. F., 1927. "Some American Hippoboscidae (Diptera: Pupi-
para) ," Canad. Entomologist, 59:246-51.
14. Kessel, Q. C., 1925. "A synopsis of the Streblidae of the world," J.
N. Y. Entomolog. Soc., 33:11-34 (4 plates).
15. Ferris, C. F., 1924. The New World Nycteribiidae (Diptera: Pupi-
para). Entomological News, 35:191-99.
CHAPTER XIX
._.
FLEAS
ORDER SIPHONAPTERA
Structural characteristics. "No part of the external anatomy of an
adult flea could possibly be mistaken for that of any other insect. The
head, the mouth parts, the thorax, the legs, the abdomen, the external
genitalia, all present features that are not elsewhere duplicated among
the hexapods." Snodgrass! (1946) continues to marvel, as well he may,
"there are numerous peculiarities that strain the imagination for a plau-
sible explanation, and the complexity of the male intromittent apparatus
is almost beyond belief." Yet internally, except for the array of spines
in the proventriculus, the flea is a fairly generalized insect. '
Fleas constitute the order Siphonaptera (Siphunculata of some au-
thors ); they are laterally compressed, highly sclerotized, totally wing-
less, minute bloodsucking ectoparasites of warm-blooded vertebrate
animals. The commoner species vary from 1.5 to 4 mm in length. The
males are as a rule smaller, often considerably smaller, than the females;
both sexes are bloodsuckers. The posterior pair of legs is strikingly
adapted for leaping; some species, such as the chigoe fleas, are able· to
burrow partly into the skin of the host and are practically sessile.
The head is a highly specialized cranial capsule; it is set closely
against the notum of the pronotum with a very short neck which pre-
cludes much movement of the head, but does allow some. On the sides
of the head are depressions (grooves) that contain the tiny knobbed
antennae (the knobs are segmented), and in front of these are the simple
eyes when these are present. The position of a conspicuous bristle (ocular
bristle) in certain position in front of the eye, may be useful in classify-
ing fleas; also useful in classincation is the presence in some species of
a conspicuous comb of bold spines, ctinidium, located just above the
mouth parts-the oral or genal ctenidium (Fig. 138).
The mouth parts of the adult flea include a minute labrum, a long,
slender, 'unpaired epipharyngeal stylet, a pair of maxillae with paired
maXillary stylets, a small hypopharynx, and a simple labium. Mandibles
are lacking in the adult flea, though present in the larva in the form of
toothed jaws.
416
MALE
CTENOCEPHALIDES fEUS
MALE
HYSTRiCHOPSYUA.IIPPIEI
FIG. 138. Showing the structural details used in the classification of Siphonap-
tera, and alsQ the life history.
417
The thorax of the flea is compact, consisting of the pro-, meso-, and
metathorax .. The pronotum lies immediately behind the head, and at its
posterior margin in many species there is a ctenidium of spine-like proc-
esses known as the pro notal ctenidium, which is useful in rough classi-
fication. The mesotlwrax is the middle segment of the thorax; the meso-
notum, the middle segment, is a simple arched plate. The metathorax is
highly developed and is specially fitted to sustain the jumping mecha-
nism. The several segrrwnts of the thorax and abdomen as well are made
up of plates (sclerites); the dorsal plates are known as tergites, the
ventral plates are the sternites, and the lateral plates are pleurites.
The abdomen consists of ten (actually eleven according to Kessel")
segments, which like the thoracic segments are made up of plates (scle-
APPENDIX OF SPERMATHECA
DUCT OF SPERMATHECA UTERUS

FIG. 139. Copulatory organs of the female Nosopsyllus fasciatus. (After Fox.)
rites), except that the pleurites are concealed. There are numerous
backward-pointing spines. On the apical edge of the seventh tergite are
the antepygidial bristles; the ninth tergite consists of a peculiar pin-
cushion-like structure known as the pygidium, probably a sensory organ .
The male terminalia are particularly important in classification. Among
the parts to be observed are the claspers, !povable and nonmovable por-
tions, and the manubrium (Fig. 138). In cleared specimens the spring-
like penis may be seen lying in the region of the fifth and sixth segments
which in copulation projects out from between the upper and lower
claspers. The females possess a sacculated spermatheca (Fig. 139), situ-
ated in the region of the eighth or ninth segment and easily visible in
cleared specimens. Some species have two spermathecae. This organ is
characteristic for many species and is, therefore, an important taxonomic
character (Fig. 138).
The legs consist of five jOints: the coxa, the joint nearest the body;
the trochanter, a very small segment; the femur; the tibia (strongly
FLEAS 419
spined); and the five-jointed tarsus terminating in a pair of ungues or
claws which may be considered as a sixth segment.
Digestive tract. As soon as blood begins to flow from the wound
made by the protraction and retraction of the maxillary laciniae, it is
drawn up into the pharynx by the action of both the cibarial and the
pharyngeal pumps. By means of powerful muscles the blood is aspirated
from the wound and on relaxation it is carried to the long narrow esoph-
agus which begins in the region of the brain and passes through the
FIG. 140. Life cycle of a flea: (upper left) egg, (center) larva, (lower left)
pupa, (upper right) female, (lower right) male.
circum esophageal ring. The esophagus opens into the stomach through
the bulbous proventriculus, which is provided internally with radially
arranged (seven rows) hair-like chitin-covered processes (Fig. 141)
which, when the encircling bands of muscles contract, cause them to
meet and form a valve, thus preventing regurgitation from the stomach.
The stomach is a capacious distensible organ nearly as long as the abdo-
men, emptying into the short intestine which in hIm empties into the
wide rectum with its six rectal glands. Where the stomach joins the in-
testine, four filamentous Malpighian tubules arise.
Life history. The eggs of a flea (Fig. 140) are comparatively large
(5 mm long), glistening white, and rounded at both ends. Relatively few,
from 3 to 18, are deposited at one laying; however, during the entire life-
time of a female the number may be quite considerable. Bacot (1914)3
records a total of 448 eggs over a period of 196 days deposited by a
single female flea, Pulex irritans Linn. Most species deposit dry eggs
which do not become attached to the hairs of the host even though
oviposition takes place on the host. Fleas seldom oviposit among the
hairs of the host, preferring the nests of the hosts where flea excrement
occurs. Captured fleas will readily oviposit in glass vials or other re-
ceptacles in which they are trapped. If deposited on a dog or cat the
eggs fall off readily when the animal stretches and shakes itself; thus
myriads of eggs may be found on the sleeping-mat of a flea-infested
animal. Temperatures of 65° to 80° F when combined with a fairly high
humidity, 70 per cent and over, appear to favor egg laying. The incu-
bation period varies from 2 to 12 days.
High mean temperature from 35° C to 37° C inhibits development,
which may account for the fact that the eggs do not hatch well on the
host. At a temperature of from 17° C to 23° C, Mayne (Mitzmain4)
found that the egg stage lasted from seven to nine days; at from 11 ° C to
15° it lasted about 14 days. Others have found that this stage may be
completed in from two to four days in Ctenocephalides felis (Bouche).
The best growth and survival of this species was obtained at 65 to 90
per cent relative humidity (Bruce 5 ).
The embyro is provided with a sharp spine (egg burster or "can
opener") on the head by means of which the eggshell is cut into shreds
by a tumbling motion of its inhabitant, which is thus liberated. (Kessel,
lac. cit.) The larvae (Figs. 138 and 140) are very active, slender, 13-
segmented, yellowish-white maggots, with segmentally arranged bristles.
The mouth parts are of the biting type, and the newly hatched larvae
of some species, e.g., Nosopsyllus fasciatus (Bosc), may subsist wholly
on the feces of the adult fleas. Very little food seems to be necessary for
their development, though excrementous matter, e.g., feces from rabbits,
rats, squirrels, and other rodents, and also dry blood may be used as
food. Excessive moisture is certainly detrimental to the life of the larvae,
although a high percentage of moisture in the air is needed. The larvae
are frequently found in houses in the crevices of the floor under the
carpet or matting, and also in stables, coops, kennels, nests of rodents,
pig pens, etc. When conditions are favorable, the time required for the
larval period may be but 9 to 15 days; if they are unfavorable, it may
extend over 200 days. At the end of the active feeding period when full
growth has been achieved, the larva enters a quiescent stage, spins a
cocoon, and pupates. The cocoon is whitish in appearance and so loosely
spun that one may see the pupa within it.
The pupal period (Figs. 188 and 140) is influenced by temperature
and varies greatly, from as short a time as seven days to nearly a year.
The life cycle (egg to adult) accordingly may vary from as short a time
as 18 days to many months. Under laboratory conditions at a tempera-
FLEAS 421
ture of 24° C Kerr 6 (1946) reports the life cycle of Ctenocephalides
felis (Bouche) to be 20 to 24 days (larval stage 11 to 12 days). His
cultures were from fleas fed on a cat.
Mayne (Mitzmain, 19107 ) observed one individual of the squirrel
flea, Diamanus montanus (Baker) (Ceratophyllus acutus Baker), from
the moment the egg was laid to the emergence of the adult flea, securing
the following data: incubation period of the egg, 8 days; first instar larva,
6 days; second instar larva, 10 days; third instar larva, 12 days; cocoon
(pupal stage), 31 days; total, 67 days.
Longevity of fleas. Bacot (loc. cit.) states that with nearly saturated
air at 45° to 50° F fleas can live for many days unfed. He reports that
Pulex irritans Linn. survived for 125 days, Nosopsyllus fasciatus (Bosc)
for 95 days, Xenopsylla cheopis (Roth.) for 38 days, Ctenocephalides
canis (Curt.) for 58 days, and Ceratophyllus gallinae Schrank for 127
days. If fed on their natural host, P. irritans Linn. may live upward of
513 days, N. fasciatus (Bosc) for 106 days, and X. cheopis (Roth.), fed
on man, 100 days. Ct. canis (Curt.) and C. gallinae Schrank have lived
for periods of 234 and 345 days respectively when fed on man. Thus
Bacot indicates that the maximum possible length of life of the various
species mentioned is 966 days for Pulex irritans Linn., 738 days for
Ctenocephalides canis (Curt. ), 680 days for Nosopsyllus fasciatus
(Bosc), 481 days for Ceratophyllus gallinae Schrank, and 376 days for
Xenopsylla cheopis (Roth.). In a moist medium such as sprouting wheat
grains and sawdust Mayne (Mitzmain, 1910, loco cit.) has kept squirrel
fleas alive for 38 days in one case and 65 days in another, the former a
male, and the latter a female. Male rat fleas fed on human blood alone
averaged 21f2 days (maximum 17) of life, and the females 34% days
(maximum 160).
Hosts and occurrence of species. As will be seen later in this chapter,
the rodent fleas are important from the public health standpoint, and
ready transfer of fleas from host to host of different species adds much
to the danger of disease transmission.
While it is apparent that ordinarily a certain species of flea predom-
inates on a given species of host, e.g., Ctenocephalides canis (Curt.) on
the dog, and particularly the cat, Nosopsyllus fasciatus (Bosc) on the rat
in Europe and the United States, Xenopsylla cheopis (Roth.) on the rat
in Asia, Ctenopsyllus segnis (SchOn.) on the mouse, and Pulex irritans
Linn. on the human, etc., host specificity in fleas is not strongly marked
in many species.
In an unpublished report to the writer on the species of fleas found
on rats in San Francisco, Rucker states that a great preponderance of
the rat fleas recovered in San Francisco were Nosopsyllus fasciatus
(Bosc) as based on 10,972 specimens as follows:
PEII CENT
Nosopsyllus fasciatus (Bosc) 68.07
Xenopsylla cheopis (Roth.) 21.36
Pulex irritans Linn. 5.57
Ctenopsyllus segnis (SchOn.) 4.48
Ctenocephalides canis (Curt.) 0.52
The following tables (Tables V to XI) adapted after McCoy8 throw
mnch light on the interchange of hosts and the predominance of species:
TABLE V
FROM BROWN RATS [Rattus r. norvegicus (ERXLEBEN)]
I N.Jasciatus X. cheopis P. irritons C. segnis Ct. canis

No. of rats I~_ _-,-_ _ _ _----,-_ __
~mbed ~I Female ~ Female ~ Female Male Female Male Female
I
606 570 I 1,252 790 1,146 225 425 44 137 13 15
TABLE VI
FROM BLACK RATS [Rattus rattus rattus (LINN.)]
!
N. fasclatus X. cheopis P. irritans C. segnis Ct. cams
No. of rats - - - - - -
combed
I -------
Male Female Male Female I Male I Female Male Female Male Female
------ --- --- --- ---
11 7 32 6 5 0 0 4 17 0 2
I
I
I
TABLE VII
(Mus musculus LINN.)
FROM MICE
From an unknown number of Mus musculus

I
N. fasciatus X. ChlO,hij P. irritans I C. $fgr/is Ct. canis
~ ~:~ ~ Female Ma: Female I~ I Female ~ Female
-
1 5 2 0 0 0 i
3 I 10 o o
I I i I
TABLE VIII
FROM CALIFORNIA GROUND SQUIRRELS [Citellus beecheyi beecheyi
( RICHARDSON) J
Diamanus Hoplopsyllus
No. of montanus anomalus
squirrels
combed
Male Female Male Female
132 2,065 2,306 86 140

I
FLEAS 423
TABLE IX
FROM THE DOG (Canis familiaris LINN.)
Ct. canis P. irritans Ct·felis D. montanus

No. combed
Male Female. Male Female Male Female Male Female
--- --- --- --- --- --- -_ - - -
4 10 44 8 17 0 1 1 0
I !
TABLE X
FROM THE CAT (Felis domestica LINN.)
Ctenocephalides felis
No. combed
Male Female
2 5 15-
TABLE XI
FROM MAN (Homo sapiens LINN.)
P. irritans Ct·felis Ct. canis D. mantanus

No. of
individuals
Male Female Male Female Male Female Male Female
--- --- --- --- --- ------
29 117 220 1 0 1 0 1 2
I
Plague. The first recorded pandemic of plague according to Wu I
et al. 9 was that of Justinian in the sixth century, starting in Egypt in 542
A.D. and spreading to Constantinople. It' lasted 50 to 60 years, and its
victims are estimated at 100,000,000. The second plague pandemic, the
"Black Death," took place in the middle of the fourteenth century in
Europe and claimed 25,000,000 victims or about one-fourth of the popu-
lation. In Great Britain from half to two-thirds of the people perished.
The great plague epidemic of London, 1664-1666, is said to have killed
70,000 persons out of a total population of 450,000. Plague disappeared
from England in about 1680, having been almost continously present
for nearly 140 years, with five epidemics.
Gradually this infection receded from Europe and the Near East,
and as Wu et al. (lac. cit.) points out "the existence of epidemic foci,
comparable to stagnant pools left behind by the lowering tide, was
recognized. . . we now know a whole series of endemic plague foci,
usually with epizootics among the wild rodents situated near or even
contiguous with Central Asia . . . the whole of this vast territory with
its hosts of wild rodents might be compared with a heap of embers
where plague smoulders continuously and from which sparks of infec-
tion may dart out now and then in various directions." The present
pandemic is believed to have originated in a wild hibernating rodent,
the tarabagan (Arctomys sibirica Schreber) in the interior of China and
began as an epidemic in Hongkong in 1894 and was transported along
world trade routes to many parts of the globe. The rat, as transported
in commerce, constitutes the chief means of spreading the diseases, the
infection being carried from rat to rat by means of rat fleas. For this
reason plague may appear in a city far removed from the original focus
of infection.
The first recorded appearance of plague on the North American con-
tinent, according to Hampton lO occurred in San Francisco, California,
on March 6, 1900, when the body of a Chinese who had died of plague
was discovered in the Chinese quarter. Rat infection was not demon-
strated until 1902. Human cases of plague continued to appear in San
Francisco, and 12~ cases with ll3 deaths were reported up to February,
1904, when the last case in the first outbreak was recorded. In May,
1907, plague was again discovered in San Francisco, and the last of this
series with 159 cases and 77 deaths occurred in June, 1908.
The next outbreak of plague in the United States, according to
Hampton, occurred in New Orleans in 1914 with 30 cases and 10 deaths
reported from June 21 to September 8. The Los Angeles outbreak of
pneumonic plague-November 1, 1924, to January 5, 1925-resulted in
41 cases with 34 deaths. The outbreaks traceable to plague-infected rats
and their fleas were quickly stamped out. However, as Hampton points
out, large areas in our Pacific coast states and adjacent territory harbor
plague-infected wild rodents such as ground squirrels and their ecto-
parasites (fleas) presenting a problem fraught with potential danger.
The answer to the question, "Is the disease in man and rodents
identical?" was not forthcoming until 1894 with the work of Yersin and
Kitasato in Hongkong. The former found the organism in the corpses of
dead rats and according to Wu et al. gave the first detailed and accurate
description of Pasteurella pestis, Yersin11 calling it Bacille de la peste.
To Kitasato, Wu states, we owe the earliest account of the organism, as
he found the plague bacilli in the "finger blood of a patient with axillary
bubo."
The disease in man. Wu et al. (lac. cit.) give the period of incuba-
tion from 2 to 10 days; the onset usually occurs within a period of 3
FLEAS 425
days. Fox in -Insects and Disease of Man/ 2 page 294 (P. Blakiston's Son
& Co., by permission), describes the disease as follows:
0
It develops suddenly with a rapid rise of temperature, reaching 103 or 104 F.
0
in two or three days, after which it is more or less irregular. There is headache,
the eyes are injected and the facies are characteristic of extreme illness. Prostra-
tion is profound and comes on early. Delirium also appears early. The charac-
teristic lesion of the disease, the bubo, usually is sufficiently pronounced by the
second day to be readily detected. The most common site for the bubo is the
femoral or inguino-femoral region, then the axillary region, cervical, iliac and
popliteal. Over the enlarged glands oedema appears and pressure elicits great
tenderness. The individual lymph nodes cannot be palpated. This swelling
forms the primary bubo. Secondary buboes may appear in other parts of the
body. In these, the glands are not matted together as in the primary bubo.
Four forms of skin eruption may be described-a petechial eruption, ecchy-
moses, a subcuticular mottling, and the so-called plague pustule . . . a bulb-
ous-like formation containing thin, turbid material teeming with plague bacilli.
It is believed to indicate the original point of inoculation, the flea bite. Extend-
ing from this to the nearest lymphatic glands faint red lines indicating lym-
phangitis may be observed. A secondary pneumonia due to the deposit of
plague bacilli in the pulmonary tissues may occur. In about a week if the
patient survives, the bubo breaks down leaving an ulcer which heals slowly.
Plague is an acute infectious disease caused by Pasteurella (= Bac-

terium) pestis (Lehman and Neumann), (Atlas u. Grund. d. Bakt.,
1896, p. 194). It is essentially a disease of rodents, usually transmitted
by rodent fleas, but it may under certain conditions cause serious epi-
demics among human beings. The term bubonic plague is applied when
inflammation of lymph glands results from the infection and buboes are
formed; these are the first foci and may remain so localized and cause
little discomfort. The buboes vary from 2 em to 10 em in diameter and
are usually located in the groin (femoral glands) and axilla (axillary
glands). When invasion of the blood stream occurs, a secondary pneu-
monic plague results. This form of plague mayoe transmitted from man
to man as droplet infection and is not dependent upon either rodents
or fleas. Pneumonic plague is almost invariably fatal.
A third type is known as septicemic plague, a fulminating type, due
to invasion of the blood stream, which runs a very rapid course; death
results before local signs are evident. Wu et al. (lac. cit;) state that "this
fulminating type of plague is met with in pneumonic as well as in bu-
bonic epidemics. Fulminating instances in pneumonic outbreaks are
often peculiar to the final stage and are presumably instrumental in
bringing about a spontaneous decline of the epidemic as droplet infec-
tion is absent, the patients succumbing before cough develops."
Fleas as vectors. Ogata 13 in 1897 came to the conclusion on epidemio-
logical grounds that fleas were the agents of transmission, pointing out
that fleas leave the rat as it becomes cold after death and so may trans-
mit the virus direct to man. He pointed out that the flea can ingest
plague bacilli while feeding, having produced plague in mice by in-
jecting an emulsion of crushed fleas taken from plague rats.
Simond" 4 in 1898 was the first to succeed in transmitting plague from
a sick rat to a healthy rat through the agency of infected fleas. Simond's
work was discredited for several years, but was successfully repeated
by Verjbitskj15 in 1903.
Liston" 6 in 1904, working in Bombay, came to the following conclu-
sions: (1) There was one flea infesting rats in India far more commonly
than did any other, viz., Xenopsylla cheopis (Roth.); (2) that these fleas
when feeding on a plague rat harbored the plague bacilli in their bodies
and that these multiplied therein; (3) that where fatal plague occurred,
many of these infected fleas were at large, and (4) that after a local
epizootic of rat plague, man was also found to harbor these rat fleas
and might become infected as had the guinea pigs used in the experi-
ment.
The following is a very brief summary of experiments conducted by
the Indian Plague Commission before and after its organization in 1905.
In the first instance healthy rats were confined in close proximity to
rats which, inoculated with plague, were beginning to succumb to that
disease and were artificially infested with rat fleas [X. cheopis (Roth.)].
The separate confinement of the rats in each case was so arranged that
both contact with and access to all excreta were excluded, although it
was provided that the fleas could pass from the inoculated to the healthy
rats; this transfer actually did take place, and in many cases these fleas
contained virulent plague bacilli; and when healthy nonimmune rats
were thus infected they died of plague to the extent of 79 per cent; this
extent of infection fell to 38 per cent, when partly immune rats of local
origin were employed.
That the plague had ol'iginated in the healthy rats through the inter-
mediary of rat fleas was further demonstrated when these were actually
transferred from artifiCially plague-infected to healthy English rats, and
the disease developed in 61 per cent of the latter.
Further, on constructing a series of miniature houses so as to repro-
duce the conditions pertaining to ordinary domiciles, it was found that
whenever these were so constructed as to admit rats to their roofs, but
not to their interiors, guinea pigs confined therein became successively
infested with rat fleas and infected by plague, but that in those houses
to which rats could not gain access plague was originated in guinea
pigs living therein, either by transferring rat fleas to them, derived from
plague-infected guinea pigs, or by an accidental admission of rat fleas
FLEAS 427
from other sources. Also, when so confined, guinea pigs had under these
conditions died of plague; healthy flea-free guinea pigs, subsequently
introduced, became infected, and the infection remained in the place
in proportion as the test animals were accessible to, and were found to
be infested with, fleas: in other words, that "if the fleas be present, the
rate of progress is in direct proportion to the number of fleas present."
Further, that when healthy guinea pigs were confined in one of the
houses, to the interior of whose roof fleas could not gain access, they
became flea-infested and infected when running on the ground, but to
a less extent when the cage was placed two inches therefrom, and not
at all when it was suspended two feet above it. The fact that infection
took place where pigs were located two inches above the ground indi-
cates that contact with infected soil is not necessary for plague to orig.
inate, and that "an epizootic of plague might start without direct con-
tact of healthy with infected animals."
To demonstrate that this communication of plague from guinea pig
to guinea pig was through the intermediary of fleas, rat fleas were taken
from a morbid guinea pig and allowed to feed through muslin on
healthy animals. The results were positive.
The state of affairs that existed in actual domiciles in which plague
occurred or had existed was next inquired into, advantage being taken
of the fact that plague-susceptible guinea pigs would serve as hosts as
well as for the collection of fleas.
Guinea pigs free from fleas were introduced into rooms in which
persons had died of plague, or from which plague-infected rats had been
taken. They were allowed to be at large in these rooms for periods of
from 18 to 24 hours. These guinea pigs not only collected the fleas on
their bodies, most of which were rat fleas, but 29 per cent of them con-
tracted plague and died of plague within a few days after being restored
to ordinary confinement. As before, many of the fleas which they yielded
harbored plague bacilli in their stomachs and were capable of infecting
additional animals.
Further, after first washing the floors and walls of the rooms with
mercuric chloride and so adequately disinfecting them for plague, but
not for fleas, and then introducing guinea pigs, these became plague-
infected when rat fleas were present.
That the infection was actually due to fleas was also shown by the
positive results when fleas collected from rats occurring in plague-
infected houses were transferred to healthy rats or guinea pigs in the
laboratory. These in due course became infected and died of plague.
Similarly fleas taken from the clean guinea pigs allowed to run in
plague-infected houses, and transferred to fresh animals, communicated
plague to them in 8 out of 40 tests.
In the next pl~ce plague-free white rats, guinea pigs, and monkeys
were placed in enclosures, which precluded contact as well as soil in-
fection in plague-infected rooms, pairs of one animal or another being
used in each of the 42 experiments of this class conducted, one in-
dividual being confined to a flea-proof receptable and the other to an
adjacent one accessible to these insects (one animal being thus a con-
trol). In the latter case plague resulted in four instances, or 10 per cent
gave postive results.
As a variation of the same experiments the enclosures for individual
animals, while protected from soil or contact infection, were surrounded
as a screen to fleas by two and one-half inches of "tangle foot" or were
unprovided with this protection, the "tanglefoot" being replaced by
sand. (Twenty-nine experiments were conducted.) In the latter case
the animals became infested with fleas, one having as many as 20; seven
became fatally infected with plague. In the former, individual fleas were
found on only three of the rats and no animals became plague-infected.
Examining the fieas entrapped, 247 in number, it was found that
147 were human fleas, 84 were rat fleas, and 16 cat fleas. Moreover, a
large proportion of each kind was examined. No plague bacilli were
found in the cat fleas, one only in 85 of the human fleas was infected, and
no less than 23 out of 77 of the rat fleas harbored plague organisms.
It was also shown that, when rats in the course of an epizootic died
of plague, the pathological features manifested in their bodies corre-
sponded to those exhibited by artificially rat-flea-infested animals, and
hence it was inferred that in nature and under experimental conditions
the animals had alike succumbed to a single agency. This similarity
especially related to the site in which buboes arose, as in both instances,
where the place of inoculation could be observed, it was the same.
Further observations. Blue t7 reports a number of observations made
in San Francisco during 1906: Two small boys found the body of a
dead rat in an unused cellar; the rat was buried with unusual funeral
honors and in forty-eight hours both were taken ill with bubonic plague.
Again, a laborer picked up a dead rat with the naked hand and threw
it into the bay. He was taken ill with plague three days later. The case
of a physician's family is also cited in which foul odors pervaded their
second story apartment over a grocery store. On removing the wainscot-
ing around the plumbing to ascertain the cause of the odor, two rat
cadavers were found in the hollow wall. In two or three days thereafter
the two members of the family who used the room sickened, one dying
on the fifth day of cervical bubonic plague. Blue believes that the re-
moval of the wainscoting set free infected rat fleas.
The following instance is reported in the United States Public Health
Reports (November 7,1913, page 2356): A fatal case of plague occurred
FLEAS 429
in Manila (P. 1.) in the person of an American, editor of the Manila

Daily Bulletin. A plague rat had been found on September 6 in the
block adjacent to the one in which the newspaper offices were located.
The editor was admitted to the hospital September 19 and died at the
Plague Hospital three days later. A mummified rat was found in the
desk of the late editor, together with live fleas, Xenopsylla cheopis
(Roth. ). Both the fleas and rat revealed bipolar staining organisms, and
inoculations into healthy laboratory rats produced typical cases of
plague terminating fatally.-
That the mummified rat must have been dead at least two weeks and
that the live fleas contained plague bacilli suggests "strong proof that
plague might be introduced into a country without either the importa-
tion of human or rat cases of plague and that fleas might be alone con-
cerned."
Role of the flea in plague transmission. The Indian Plague Commis-
sion showed that the average capacity of a flea's stomach [Xenopsylla
cheopis (Roth)] was .5 cubic millimeter, and that it might receive as
many as 5,000 Pasteurella pestis while imbibing blood from a plague
rat. The Commission found that the bacillus multiplies in the stomach
of the flea and that the percentage of fleas with bacilli in the stomach
varied with the season of the year. In the epidemic season the percentage
was greatest for the first four days, and on one occasion the stomach
was found filled with the organisms on the twentieth day. In the non-
epidemic season no plague bacilli were found in the stomach after the
seventh day. They also found that in the epidemic season fleas might
remain infective up to 15 days, while in the nonepidemic season but
seven days, and in the latter case the percentage of infection in animals
was much less than in the epidemic season. They showed that while one
flea was occasionally able to carry the infection, this was not usual. It
was found that both males and females were capable of transmitting
the disease. As to the manner of dissemination the Commission found
bacilli in the stomach and rectum only, never in the salivary glands or
body cavity, and but rarely in the esophagus, and then only when the
flea was killed immediately after feeding. After digestion, the blood in
the stomach passes into the rectum and is ejected as a dark red or tarry
droplet, containing virulent plague bacilli, which if rubbed into recent
flea-bites resulted in infection of the animal. The actual inoculation
therefore, it was believed, was accomplished indirectly by the flea-bitten
person's scratching or rubbing the site of the bite after the infected
flea had discharged fecal material upon the skin. That there is a great
deal of difference in the consistency and other characteristics of the
fecal deposits of the various species of fleas is easily observable, e.g.,
the human flea, Pulex irritans Linn., defecates freely while feeding but
is not a ready vector of bubonic plague, while the rat flea, Xenopsylla
cheopis (Roth.) seldom defecates while feeding and is a potent vector.
Bacot and Martin 1S (1914) demonstrated a mechanism of infection
which is based on observations made by SwellengrebeP9 (1913) in Java
that Xenopsylla cheopis (Roth.) seldom defecates when feeding and
showed that infection resulted when the flea's only contact with the
experimental animal was by means of the proboscis, i.e., the infection is
introduced with the bite directly. That this mode of infection was in the
manner of regurgitation due to a temporary obstruction at the entrance
to the stomach was discovered by Bacot and Martin (lac. cit.), who
state that on
examining the contents of the stomach of a flea a day or two after it has fed
upon infected blood, clusters of minute brown specks darker in colour and
firmer in consistency than the rest of the contents are visible. . . . Later the
stomach and proventriculus show jelly-like masses of a brown colour. These
masses are possessed of considerable cohesion and are with difficulty teased out
so as to make a film suitable for microscopical examination. The plague-cul-
ture grows in the proventriculus as well as in the stomach. Owing to its gelat-
inous consistency, it not infrequently leads to incompetence and even com-
plete blocking of the proventricular valve. . . . Although with the proventric-
ulus obstructed in this manner fresh blood cannot find its way into the stom-
ach, this does not prevent the insect sucking, as the pump which aspirates
blood up the sucking tube and propels it into the stomach is situated in the
pharynx. On the contrary the flea suffers from thirst and is persistent in its
efforts to satisfy this appetite, but only succeeds in distending the oesophagus.
The blood in the distended oesophagus may flow out again on cessation of the
sucking act, and we have seen drops of blood escape from the mouth parts of
"blocked" fleas when the insect withdrew its proboscis. . . . Given the oppor-
tunity, the insects suck blood again and again and if the pharyngeal pump
ceases for a moment, some of the blood will by the elastic recoil of the
oesophageal wall be driven back into the wound and carry with it plague
bacilli.
Bacot and Martin found that infected fleas lived as long as 50 days at
from 10° C to 15° C and 23 days at 27° C and died infected. Working
with two species of rat fleas, Xenopsylla cheopis (Roth.) and N osop-
syllus (= Ceratophyllus) fasciatus ( Bosc ), fed on septicemic blood,
Bacot and Martin conclude that they "can transmit plague during the
act of sucking and that certain individuals suffering from a temporary
obstruction at the entrance to the stomach were responsible for most of
the infections obtained, and probably for all." In the course of time the
plague culture forming the proventricular plug undergoes autolysis and
the normal passage of blood is re-established.
Figure 141A shows the position of the spine-like epithelial cells in the
proventriculus when at rest, the opening into the stomach being free for
FLEAS 431
the passage of blood. Figure 141B shows the opening into the stomach
closed against the outward passage of blood on contraction of the
muscular bands. Bacot20 points out that it is the lodgment and growth
of the bacilli among the spines that is the initial stage of the blockage.
In describing the mechanism of plague transmission by fleas, Eskey
and Haas 21 (1940) show numerous microphotographs of blood-distended
stomachs of fleas after feeding. The elapsed interval between an infec-
tive blood meal and an infective bite (transmission) for Xenopsylla
cheopis (rat flea) averaged about 21 days (shortest interval 5 days,
longest 31 days); for Diamanus montanus (ground-squirrel flea) the
average was 53 days; the longest interval in the series was 130 days.
The average length of life of fleas after being plague infected was 17
days (maximum 44 days) for X. cheopis, and 47 days (maximum of 85
days) for D. montanus.
Eskey 22 has shown that virulent plague organisms are more con-
stantly present in the feces of some species of fleas than in others. He
FIG. 141. Spine-like epithelial cells ~K

in the proventriculus of Xenopsylla
cheopis: (A) at rest, (B) showing
opening into stomach closed.
reports that plague followed every inoculation of feces deposited by
infected Diamanus montanus (Baker), while less than one-third of the
fecal inoculations of Nosopsyllus fasciatus (Bosc) gave positive reac-
tions. He also reports that the feces of Xenopsylla cheopis (Roth.) gave
positive reactions, but these fleas did not survive long enough to deter-
mine whether or not the results would be constant for any length of
time. Eskey points out that there seems to be danger of infection from
virulent plague organisms present in the feces of all plague-infected fleas.
Still another possible mode of transmission, which applies, however,
only to transfer from radent to rodent, has been suggested by various
workers, namely, that of crushing infected fleas with the teeth, with
infection through the mucosa of the buccal cavity resulting in lymph-
node involvement in the region of the neck.
Plague in field rodents. In California plague was demonstrated in
ground squirrels, Citellus beecheyi beecheyi (Richardson), under nat-
ural conditions in 1908 by McCoy. According to that author at the time
of his writing a report on this finding (1910) ,23 about a dozen persons
had contracted the disease under circumstances that pointed conclusively
to squirrels as the cause. The two species of fleas commonly infesting
the ground squirrel in California are Diamanus montanus (Baker), (=
Ceratophyllus acutus Baker) and H oplopsyllus anomalus Baker, of
which the former is far more numerous. McCoy proved the first-named
species a carrier as follows: He inoculated a ground squirrel subcuta-
neously with a broth culture of P. pestis derived from a human case of
plague. This squirrel died on the £fth day, but three days before its
death, 100 fleas, D. montanus (Baker), were put in the cage with it. The
dead animal was removed from the cage while warm, and 27 live fleas
taken from its body. Smears made of the crushed bodies of two of these
fleas showed an abundance of pest-like bacilli in each. The remaining
25 fleas were put into a clean cage with a healthy squirrel. This animal
died of subacute plague 10 days later, the buboes being in the region
of the median, posterior inguinal, and pelvic glands. A pure culture of
P. pestis was obtained from the liver. McCoy states that the experiment
is conclusive in showing that D. montanus (Baker) may convey plague
from a sick to a healthy squirrel. The squirrels used in the experiment
were kept in quarantine for at least a month prior to their being used,
which was necessary to exclude any naturally infected ones. McCoy
found the bacilli in squirrel-flea feces four days after removal of the
fleas from the host.
Sylvatic (selvatic) plague. The designation selvatic plague was pro-
posed by Ricardo Jorge (1928) (see p. 4) for the plague of field
rodents. Fleas play an important role in transmission from rodent to
rodent and consequently in the endemicity of the disease. It is now
known that under certain ecological conditions in vacated squirrel bur-
rows fleas may continue to harbor virulent P. pestiS for many months
thus providing a virtual insectan reservoir for the infection under sylvatic
conditions. Fleas have been known to survive though starved for more
than six months (196 days).
Aside from the matter of flea transmission it is important to bear
in mind that the great epidemic of plague in Manchuria resulting in
60,000 deaths in 1910-1911 was of the pneumonic type and sprang from
the wild tarabagan, Arctomys sibirica Schreber (Siberian marmot),
which was hunted for its valuable reddish brown fur by numerous
Chinese hunters unfamiliar with its dangers. The mountainous portions
of Central Asia, i.e., portions of Siberia, Mongolia, Tibet, and Manchuria,
are regarded as the original home of plague, and the tarabagan as well
as its flea parasites play an important role as reservoirs of the infection
in this area. These large rodents are about half a meter in length with·
a bushy tail about 15 cm long. It is pointed out that the low body tem-
perature of the tarabagan during hibernation enables the animal to sur-
vive and thus to carryover the infection from one season to the next,
and the flea, Oropsylla silantiewi (Wagner), as well as perhaps other
bloodsucking ectoparasites, transmit the infection from animal to animal.
Comparable endemic foci of sylva tic plague occur in South Africa,
where the gerbilles (Muridae, Berbillinae) belong to three genera,
FLEAS 433
particularly Tatem, e.g., Tatera lobengulae De Wint.; the multimammate
mouse, M astomys coucha (A. Smith) ( M uridae, M urinae ), and their
flea parasites play the leading role. 24 In the Russian steppes the susliks,
Spermophilus refuscens and other species (Sciuridae), and their flea
parasites, Citellophilus tesquol'um (vVagner) and N eopsylla setosa
Wagner, play a similar role. In North America, as already explained,
ground squirrels (Citellus spp.) (Sciuridae) and their fleas, e.g., Dia-
manus montanus (Baker) ( = Ceratophyllus acutus Baker), may be
important reservoirs of sylva tic plague. In South America, the cavy,
Cavia aspel'ea Pallas, and its fleas, e.g., Rhopalopsyllus cavicola (Wey-
FIG. 142. A California ground squirrel, Citellus beechyi. (Photograph by

R. Maynard, California State Department of Public Health.)
enb.), playa similar role. Burroughs Z" gives further consideration to the
matter of animals (other than rats) nahlrally infected with plague.
Sylvatic plague remains localized or at best spreads slowly, and in
each endemic region a particular native animal or group of animals
(rodents) maintains the infection, and when other small house-invading
rodents such as mice and rats come in contact with such a focus, the
infection may be carried to human habitations, and human cases may
result; or likewise if humans invade the territory of sylva tic plague, in-
fection may occur. Under such circumstances man may contract the in-
fection by direct traumatic contact with infected rodents, or possibly
by flea bites, which is less likely. Meyer 26 points out that by 1915 at
least 24 cases with 15 deaths were directly attributed to exposure to
ground squirrels in six California counties. Meyer 27 (1947) lists 38
species of wild rodents and rabbits which have been found plague-in-
fected in 14 states of the Western United States. Three groups of rodents
constitute the primary natural reservoirs of plague in that part of the
country: ( 1) ground squirrels (Citellus) , Pacific coast and northern
intermountain area (Fig. 142); (2) wood rats (N eotoma) , southwest
desert areas; (3) prairie dogs (Cynomys), plateau regions of Arizona
and New Mexico.
Although there is some evidence to support the hypothesis setting
forth the probable presence of plague among wild rodents in North
America, particularly ground squirrels, Citellus, long before the demon-
stration of plague in ground squirrels in California by McCoy (lac. cit.)
in 1908, and therefore long before the discovery of plague in rats in San
Francisco in 1902; nevertheless Meyer states that the available evidence
is not adequate to warrant a decision. There appears to be some pale-
ontological evidence as well connecting the genus Citellus of North
America with ancestors on the Asiatic high plateau which is also the
home of plague.
Meyer in "The Ecology of Plague" (1942, lac. cit.) points out that
many of the unknown factors in sylva tic plague are intimately related
to the influence of the climatic conditions on the life and longeVity of the
insects involved (fleas and lice). Too little attention has been given to
the host-parasite relationships of this as well as other diseases involving
arthropod vectors. The factors influencing the low transmissibility of
sylvatic plague are as yet little understood. In the meantime Meyer
(1947, lac. cit.) urges the maintenance of so-called "rodent free" belts
around towns and "research and more research into the ecology of
sylvatic plague."
Wild rodent fleas. Dunn and Parker 28 investigated the flea population
of a variety of species of wild animals in the Bitter Root Valley of
Montana. Oropsylla (= Ceratophyllus) idahoensis (Baker) was found
infesting a large percentage of the 94 ground squirrels [Citellus colum-
bianus (Ord)] examined, the average per animal being 3.86. While O.
idahoensis (Baker) was by far the most common species of flea on this
species of ground squirrel, six other species were taken in order of abun-
dance, viz.: Opisocrostis tuberculatus (Baker), N eopsylla inopina Roth.,
M onopsyllus eumolpi (Roth. ), C ediopsylla inequalis (Baker), and
Monopsyllus vison (Baker) (one specimen). It is of interest to know
that these authors took Oropsylla idahoensis (Baker) from the follow-
ing species of wild animals: cottontail rabbits, Sylvilagus n. nuttallii
(Bachman); snowshoe rabbits, Lepus bairdi Hayden; pine squirrels,
Sciurus hudsonicus richardsoni Bachman; woodchucks, Marmoto f.
flaviventer (Aud. and Bach.); and the bushy-tailed woodrat, N eotoma
c. cinerea (Ord). The woodchuck showed a heavy infestation (average
FLEAS 435
15.47 per animal) of Thrassis acamantis (Roth.). Observations made on

the marmot in California also show a heavy flea infestation, averaging
26.57 fleas per animal according to unpublished data by Stewart.
Pearse 28 in his study of fleas on rodent hosts in Nigeria concludes
that "the ecological factors which are associated with a high degree of
infestation are dry climate or habitat, the occupation of a more or less
permanent home by the host, and large size of the host. Factors as-
sociated with low degree of infestation are wet climate or habitat, lack
of permanent abode of host, small size of host, and wandering or
arboreal habits of host."
While California ground squirrels, e.g., Citellus beecheyi beecheyi
(Richardson), have several species of fleas infesting them, among these
Diamanus montanus (Baker), Hoplopsyllus anomalus Baker, and
Malaraeus telchinum (Roth.), there is usually a preponderance of the
first-named species. Wood rats (Neotoma) commonly have several
species, among them Ctenopsyllus segnis (Schon.), Orchopeas sexden-
tatus sexdentatus (Baker), Anomiopsyllus nudatus Baker, and Malaraeus
telchinum (Roth.).
Infected and infective fleas. In relation to the spread of sylvatic
plague Meyer 30 calls attention to certain paradoxical observations: (1)
that despite active reservoirs with hundreds of infected rodents very
few cases of human plague were diagnosed on the North American
continent; (2) that plague-infected fleas are taken from animals which
had anatomically been declared n,oninfected; (3) squirrel hunters and
plague-survey crews are commonly covered by fleas and are bitten by
squirrel fleas yet are not infected.
Meyer points out that in sylva tic plague man becomes infected with
bubonic plague by immediate contact with the sick or dead rodents, flea
transmission being infrequent. The danger represented by individual
fleas appears therefore more limited than was originally believed. Wild
rodent fleas serve as "preservers" of plague infections in suitable rodent
burrows for many months and under such natural conditions, while
infected, harbor bacilli which are of low virulence or avirulent. These
"preserver" fleas are believed to be "nonblocked."
Meyer continues,
Rodents with latent infections will hibernate only to develop acute plague
early in spring (March and April). Since the flea population is as a rule simul-
taneously very high, a great reservoir of infected vectors is thus created. The
cadavers of the dead rodents are rapidly and effectively removed by the larvae
of the Lucilia flies, while the fleas pe~ist in the nests. With the migration of
the young squirreL; and chipmunks into the empty abandoned burrows and
nests, highly susceptible hosts are thus brought in contact with infected and in-
fective fleas. They may bring the vectors to the surface and some may thus
436 MEDICAL ENTO'MOLOGY
contribute to the intensity and the expansion of the virus. These events are
probably accompanied by a variable degree of subclinical immunization
favored by factors of age and reduced metabolism due to the approaching
hibernation. Again latent infections and infected fleas in the burrows furnish the
chain which connects the epidemic of one year with that of the next. Thus
sylvatic plague smoulders for years and is everlasting. Suppressive measures
against sylvatic plague in order to be effective must by necessity be directed
against the hosts, the rodents, and the vectors, the various species of fleas. In
selecting the procedures to reduce the rodent populations chemicals, preferably
gases which are also insecticidal, must be chosen. [See methyl bromide fumi-
gation, p. 458.]
Laboratory tests indicate that many species of Siphonaptera (more

than 30) can be infected experimentally, but relatively little has been
done on vector efficiency except for the very significant work of Wheeler
and Douglas 31 (1941). These investigators point out that the vector
efficiency of a species of arthropod incriminated in the transmission of
an infectious agent must necessarily take into consideration the follow-
ing factors: (1) the infection potential, (2) the vector potential, and (3)
the transmission potential. The first is based upon the percentage of a
given species in which the infection becomes established; the second,
upon the percentage of infected individuals which become infective, and
the third, is the mean number of transmissions effected by a group of
infective individuals. Thus these workers demonstrated that Diamanus
montanus (Baker) has an infection potential of .85, a vector potential
of .52, and a transmission potential of 2.58 with a vector efficiency of
1.14, compared with an infection potential of .98, a vector potential of
.29, a transmission potential of 1.44, and a vector efficiency of .39 for
Xenopsylla cheopis (Roth.).
Reporting on the results of his vector efficiency studies in relation
to sylvatic plague, Burroughs (loc. cit., 1947) states that experimental
evidence was obtained that different strains of a species of flea taken
in different geographical areas may differ markedly in their biological
vector capacity. This observation was based primarily on experiments
with Diamanus montanus, which proved much less efficient than Xenop-
sylla cheopis in striking contrast to the results obtained by Wheeler and
Douglas (loc. cit.), whose D. montanus came from a widely separated
area.
Eskey and Haas (loc. cit., 1940) conducted transmission experiments
with individual fleas collected from wild rodents in areas known to be
sylvatic plague foci. Many of the infective fleas (proved on guinea pigs)
transmitted plague to more than one experimental animal (guinea pigs).
Both male and female fleas transmitted the infection, although males
proved to be much less efficient on the whole; however, one Xenopsylla
FLEAS 437
cheopis female infected 10 guinea pigs, and one male ground-squirrel

flea, Opisocrostis labis (Jordan and Rothschild) infected 11.
Murine (endemic) typhus fever. For many years a mild form of
typhus has existed in seaport communities in warmer climates in many
parts of the world. This infection was long believed to be of murine
origin. Mild typhus was first reported in the United States (New York)
in 1898 and was referred to as Brill's disease by some authors and by
others as classical typhus (see Zinsser). It was later reported in Atlanta,
Georgia; Charlotte, North Carolina; and Galveston, Texas. In 1931
Mooser, Castaneda, and Zinsser 32 reported rats as the carriers of typhus
fever in Mexico. The causal organism received the name Rickettsia
mooseri Monteiro in 1931. In 1932 Dyer et al. 33 reported transmission of
endemic typhus by rubbing crushed infected fleas or infected flea feces
into wounds. The mechanism of transmission, including infection by
means of dry flea feces, is essentially the same as in the classical louse"
borne typhus previously described. Several species of rat fleas have been
incriminated, including Xenopsylla cheopis and Nosopsyllus fasciatus,
as well as (experimentally) the sticktight flea, Echidnophaga gallinacea
(Westw.) (Alicata,34 1942). Dyer and associates report that the incuba-
tion period in one experimental animal, a guinea pig, after inoculation
of a flea emulsion was 10 days; also the virus remained viable in the flea
for at least 52 days, showing the importance of the flea as a reservoir
under natural conditions.
Zinsser 35 (1937) held to the belief that both types of typhus may be
either endemic or epidemic. He states: "Although the murine disease
reaches man first from infected rats by flea vectors, this virus can also,
like the European, pass from man to man by the louse . . . capable of
epidemic dissemination of the murine as well as of the classical typhus .
. . . Brill's disease is an imported classical typhus, endemically estab-
lished in cities with large immigrant populations."
Recently (1948) Castaneda,36 reporting on flea-borne and louse-
borne typhus in Mexico, states that the geographic distribution of both
types of typhus in Mexico follows the climate of each zone. Pure murine
typhus is found in warm regions and is flea-horne. Classical typhus oc-
curs in cold and mild zones, where louse-borne infection is frequent. In
these zones murine typhus also occurs and may become louse-borne and
cause important epidemics of high mortality. In Mexico City he reports
that from 1927 to 1931 it was demonstrated that murine typhus was the
prevalent type, while in 1938 most of the isolated strains from typhus
patients were classic. He continues, " . . . during the last seven years
murine strains have been found only on rare occasions; complement
fixation tests have corroborated the prevalence of classic typhus in the
cold and temperate zones where typhus is endemic." Castaneda states
that rat campaigns are advisable in zones where there is danger of louse-
borne murine typhus.
Beck and van Allen 37 (1947) state that two epidemiologic periods
have occurred in California. The first period was probably of epidemic
type and louse-borne, 1916 to 1917, consisting of an isolated epidemic
of 32 cases and 4 deaths with source of infection in Mexico. The second
period of the murine type and flea-borne was from 1919 to 1945 inclu-
sive, representing 362 cases and 17 deaths. Cases during the latter part
of the second period occurred during every month of the year with a
slight increase during late summer and autumn. Rattus norvegicus, R.
rattus, R. alexandrinus, and Mus musculus were found naturally in-
fected. Ground squirrels, Citellus beechyi, were experimentally infected
with a murine strain of typhus.
THE COMMONER SPECIES OF FLEAS
Systematic. There are said to be more than 900 described species of

Siphonaptera; for North America and the West Indies, Ewing and FOX 38
( 1943) list 209 species and 63 subspecies and varieties in 61 genera and
14 subgenera. The order is divided into six families as follows: (1)
Hectopsyllidae, (2) Pulicidae, (3) Dolichopsyllidae, (4) Ischnopsylli-
dae, (5) Hystrichopsyllidae, (6) Macropsyllidae. An extensive index to
the literature of Siphonaptera of North America has been published by
Jellison and Good. 390
Key to the Families of Siphonaptera
(Modified by Stewart after Ewing)
1. The three thoracic tergites together longer than the first abdominal
tergite ...... ........ ............ .... .............. .................. 2
The three thoracic tergites together shorter than the first abdominal
tergite ................................................................ ............ Hectopsyllidae
2. No vertical suture from dorsal margin of head to bases of antennae;
frontal region almost evenly rounded along margin 3
A vertical suture passing upward from the bases of the antennae to the
dorsal margin of head; margin of frontal region usually most strongly
curved at vertex......................................................................................... 4
3. Abdominal tergites with but a single row of setae .......... Pulicidae
Abdominal tergites with at least two rows of setae ....... Dolichopsyllidae
4. Head without a pair of dark anteroventral flaps on each side. 5
Head with a pair of dark anteroventral flaps on each side ..
Ischnopsyllidae
5. Occipital region without dorsal incrassation; frontal region entire
H ystrichopsyllidae
Occipital region with dorsal incrassation; frontal region divided, the
anterior part bearing a border of spines.. ..Macropsyllidae
.. Also see Holland, C. P., 1949. The Siphonaptera of Canada. Ottawa: Dominion
of Canada Dept. Agric. Pub. 817, Tech. Bull. 70. 44 maps + 42 plates.
FLEAS 439
Family Pulicidae. Pulex irritans Linn. (Fig ..143) is commonly known

as the human flea. It is cosmopolitan in distriblltion and occurs on many
domesticated animals, particularly swine. This species has neither oral
nor pronotal ctenidia. The meta coxae have a roW or patch of short spine-
lets on the inner side; the mesosternite has an internal rod-like incrassa-
tion extending dorsoanteriorly. The maxillary hlcinae extend about half-
way down on the fore coxae, which distinguishes this species from Pulex
dugesii Baker (mandibles extending at least three-fourths the length
of the fore coxae), also known as a human flea but restricted to Mexico
and the border states of the United States. Pulex irritans Linn. transmits
FIG. 143. Pulex irritans, the human flea; male (right), female (left). X 17.
plague under laboratory conditions and may be the chief vector of two
unusual types of plague, e.g., viruola pestosa (a vesicular form) and
angina pestosa (a tonsillar form) found in ECllador.
Ctenocephalides canis (Curtis) and Ctenocephalides felis (Bouche)
are the dog flea and cat flea respectively. Both species attack cats and
dogs as well as man. Both have the oral ctcnidia consisting of eight
spines and the pronotal comb of 16 spines. They may be separated as
follows:
In the female the head is fully twice as long as high (seen from side); two
or three bristles on metathoracic episternum; bristles on metathoracic epimeron,
first row, four to eight, second row, five to seven; seven to ten bristles on inner
side of hind femur.. ....... ......... . .............. ·felis
In the female the head is less than twice as long as high (seen from the
side); three or four bristles on metathoracic episternum; bristles on meta tho-
racic epimeron, first row, seven to eleven, second row, seven to nine; ten to
thirteen bristles on inner side of hind femur .......... · .................................... canis
Xenopsylla cheopis (Rothschild) (Fig. 144) is the Oriental or Asiatic

rat flea. It habitually inhabits buildings and bites man freely. It resem-
bles Pulex irritans Linn. in that both the oral and pronotal ctenidia are
absent. The ocular bristle is in front of and just above the middle of the
eye; there are two bristles on the gena; oral bristles placed low down
just above the base of the maxillae; each abdominal tergite has but one
row of bristles; the hind femur has a row of about eight bristles. The
mandibles reach nearly to the end of the anterior coxae. Incrassation of
mesosternite consists of one rod extending anterodorsally and one rod
extending upward nearly perpendicularly (Fig. 138).
Mellanb y 40 (1933) has performed experiments proving that X. cheo-
pis (Roth.) can complete its life history between 18° C and 35° C in
moist air. Between 18° C and 29° C air with a relative humidity of 40
per cent is unfavorable, while with 60 per cent pupation takes place suc-
cessfully. Pupation at 18° C required eight days, at 22° C it required
d "
/!¥,/ ~ ~~"~~
~) \~
-/;>/
(:;:5:;;V'
~
';)
FIG. 144. Xenopsylla eheopis, the Oriental rat flea; male (left) , female
( right). X 17.
six days, and at 29° C to 35° C it required four days. The develop-
mental zero for pupation is about 15° C.
Xenopsylla brasiliensis (Baker) is an African species, the predomi-
nant rat flea in Uganda, Kenya, and Nigeria. It has spread to South
America and certain areas in India. It is regarded as a more important
vector of plague than X. cheopis (Roth.) in Kenya and Uganda, since it
is "the flea of the hut" while the latter infests rats of stone and brick
buildings.
Xenopsylla astia Roth. has a restricted distribution
being found mostly along the low-lying coast of Ceylon, the east coast of India,
and along the opposite coast of Bengal . . . while X. astia may be the respon-
sible vector [of plague] in certain circumscribed and isolated outbreaks, the
available evidence . . . points to its inferior position in the epidemiological
picture. . . . Moreover astia outbreaks, if and when they do occur, are not
known to carry over from one season to another.
(Wu, et al., loe. cit.)
FLEAS 441
Xenopsylla hawaiiensis Jordan is a common flea of the Hawaiian rat,
Rattus hawaiiensis. According to Eskey reported by Jordan 41 this species
of flea has a very peculiar distribution.
It has not been found in Honolulu or vicinity, while it is quite common on rats
caught about nine miles away on the opposite side of the island. It is essen-
tially a flea of field rats and rarely found on rats caught in buildings.
Family Dolichopsyllidae. Nosopsyllus (= Ceratophyllus) fasciatus
(Bosc) is the European rat flea (Fig. 145). It is widespread over Europe
and America, being less common in other parts of the world. It has been
recorded on rats, house mice, pocket gophers,. skunk, man, and many
other host animals. It has but one ctenidium, the pronotal, which has
FIG. 145. Nosopsyllus fasciatus, the rat flea; male (left), female (right). X 17.
18 or 20 spines. There are three bristles in front of the eye; and in the
female two bristles, and in the male four in front of these. There are
three or four hairs on the inner surface of the hind femur. N. fasciatus
(Bosc) is regarded as a negligible factor in the causation of natural out-
breaks of plague.
The genus Nosopsyllus may be distinguished from the genus Dia-
manus by the fact that in Diamanus there are long, thin bristles on the
inside of the mid and hind coxae from the base to the apex, while in
Nosopsyllus such bristles occur at most in the apical half.
Diamanus montanus (Baker) ( Ceratophyllus acutus Baker) (Fig.
146) is a common species of squirrel flea described from California.
This species may be recognized by a spine at the tip of the second joint
of the hind tarsus longer than the third joint and reaching over on to the
fourth joint; the abdominal tergites have each two rows of bristles; the
male claspers are very large and long, and sickle-shaped.
Ceratophyllus niger Fox was originally described from specimens
taken from man and from Rattus r. norvegicus in California.
Ceratophyllus gallinae Schrank is commonly known as the European
hen flea although it has a wide range of hosts. See previous pages.
Family Hectopsyllidae. Tunga penetrans (Linn.) [Dermatophilus
(= Sarcopsylla) penetrans (Linn.)] is commonly known as the chigoe
flea.
Echidnophaga gallinacea (Westw.) is commonly known as the stick-
tight flea of poultry and other animals.
FIG. 146. Diamanus montanus, the squirrel flea; male (right), female (left).
x17.
Family Hystrichopsyllidae. Ctenopsyllus segnis (SchOn.) (Leptopsylla
musculi Duges) is the cosmopolitan mouse flea (Fig. 147). It is com-
monly found on rats. It bites man reluctantly and is regarded as a weak
vector of plague; its role in human outbreaks is considered negligible.
Rat fleas on ships and at seaports. The United States Public Health
FIG. 147. Ctenopsyllus segnis, a mouse flea; male (right), female (left). x17.
Service has conducted a number of rat-flea and rat surveys at various
seaports. Williams 42 reports that on a two-year survey of ships at the
Port of New York 1,913 ships produced 18,265 rats, an average of 9.6
rats per ship. The ship rat is almost exclusively the black rat, Rattus
r. rattus (Linn.) and the roof rat, Rattus rattus alexandrinus (Geoffroy-
St. Hilaire and Audouin), constituting 99.65 per cent of all rats. Because
FLEAS 443
of the climbing habits of these rats, they are more likely to get into cargo
and aboard ships than the Norway rat. The report indicates that the
majority of ships carry few rats, and only about 50 per cent of arriving
ships constitute about 90 per cent of the potential plague menace.
The dead rats collected after ship fumigations (hydrocyanic acid)
were examined for fleas. A total of 7,886 fleas were taken from 18,265
rats, an average of 0.43 per rat, which was about 30 per cent of the
expectation of fleas from live rats. Of the total number of fleas, 6,992
(88.68 per cent) were Xenopsylla cheopis (Roth.) and 786 (9.97 per
cent) Nosopsyllus (= Ceratophyllus) fasciatus (Bosc). The remaining
number of rodent fleas were Ctenopsyllus segnis (Schon.), 63, Xenop-
sylla brasiliensis (Baker), 4. Pulex irritans Linn. appears as a single
.specimen and the cat and dog flea numbered but 7.
The rat-flea survey at Norfolk, Va. (Hasseltine, 1929),43 resulted in
the capture of 1,561 rats, of which 883 harbored fleas; 4,898 fleas were
taken. Of these fleas 81.6 per cent were Xenopsylla cheopis (Roth.),
and 17.7 per cent were Nosopsyllus (= Ceratophyllus) fasciatus (Bosc).
The Norfolk survey was based on caged, trapped rats taken under favor-
able conditions for harborage and propagation. Consequently, the num-
ber of fleas per rat (the living rats were chloroformed) was much higher
than the average per ship rat on fumigated vessels, i.e., 5.5 against 0.43;
also Rattus r. norvegicus (Erxleben) constituted all but four of the total
number of rats taken at Norfolk.
At the port of New Orleans, Fox and Sullivan44 report that from 3,839
live rats 17,559 fleas were taken, of which 6,566 were Xenopsylla cheopis
(Roth. ), 10,269 were Ctenopsyllus segnis (Schon.), (Leptopsylla musculi
Duges ), and 724 were Nosopsyllus (= Ceratophyllus) fasciatus (Bosc)
-an average of 4.83 per rat. These authors state that it is quite significant
that at the port of New Orleans, where plague has actually existed,
Xenopsylla cheopis (Roth.) is the predominant rat flea present during
every month of the year.
Inspection of vessels for rat infestation has two principal objects ac-
cording to Williams: 45 first to determine the presence or absence of rats
so as to determine whether or not a vessel shall be fumigated or other-
wise treated to kill fleas and rats; and second, to determine the location
of rats when present so as to apply fumigation effectively and to main-
tain proper rat-proofing.
The chigoe flea. Tunga (= Sarcopsylla) penetrans (Linn.) (Fig.
148) belongs to the family Hectopsyllidae and is known as "chigoe,"
"jigger," "chigger," "chique," "nigua" or "sand flea." The head of this flea
is definitely angular and is usually larger proportionately than the heads
of other fleas; there are no ctenidia on the head or pronotum; the mouth
parts are conspicuous; the palpi are four-segmented. Tunga penetrans
(Linn.) is a tiny burrowing flea found in the tropical and subtropical

regions of North and South America, also in the West Indies and Africa.
Its introduction into Africa is said to have Occurred as late as 1872. The
chigoe is reddish brown and measures about 1 mm in length except that
the impregnated female may become as large as a small pea. The adult
fleas are intermittent feeders but adhere closely to the host. The female
when impregnated proceeds to burrow into the skin of the host, fre-
quently between the toes and into the soles of the feet, causing nodular
swellings which ulcerate. The eggs are deposited either in the ulcer
or drop to the ground when discharged from the body of the female,
the tip of whose abdomen protrudes slightly from the nodule. The larvae
FIG. 148. The chigoe flea, Tunga

penetrans.
which emerge in a few days from the eggs are typical flea larvae. Those
hatching in the ulcer usually drop to the ground to develop under con-
ditions similar to those having hatched on the ground. Faust and Maxwell
(1930) 46 report a case in which the eggs had hatched in or on the body
around the sites of the burrows of the gravid females, and the larvae
had thrived and grown there. The larval peric)d under favorable condi-
tions probably requires not more than 10 to 14 days and the cocoon
or pupal period about a like number of days.
Pathogenesis. The chigoes commonly attack the bare feet, these being
nearest the ground, infesting the skin between the toes and soles; but
no part of the body is exempt from attack. The burrowing female flea
causes extreme irritation; the area surrounding it becomes charged with
pus, producing a distinct elevation. The ulcera.tions due to the presence
of numerous chigoes become confluent. Weltman in a personal com-
munication to the author attributes the commonly observed autoampu-
tation of toes of natives in Angola to the work of the chigoe. Tetanus
and gangrene frequently result.
FLEAS 445
Where the chigoe flea occurs, walking in bare feet should be avoided.
Parts of the body attacked by the fleas should receive immediate at-
tention. Repeated bathing of the infested part with Lysol gives good
results. The insect can be removed quite easily by means of a sterile
needle or very fine-pointed knife blade. The wounds caused by this
treatment must be carefully dressed and allowed to heal.
The sticktight flea. Echidnophaga gallinacea (Westw. ), also known
as the "sticktight," is a serious poultry pest in many parts of subtropical
America. It commonly attacks poultry of all kinds, also cats, dogs, rab-
bits, horses, and man. It differs from Tunga penetrans (Linn.) in having
the angle of the head acutely produced, while in T. penetrans (Linn.)
the head is obtuse instead of rounded and the eyes and antennae are
in the posterior half of the head. It is from 1 to 1 y:! mm in length.
Before copulation both sexes are active, hopping about much as do
other species of fleas. Shortly after feeding, the females attach themselves
firmly to the skin of the host and begin, to burrow. At this time the sexes
are in copulation, The burrowing fema1es deposit their eggs in the ulcers
which have been produced by the infestation, The usual incubation
period according to Parman 47 is from six to eight days at a temperature
average of 76° F. If the eggs are deposited in the ulcer, the larvae crawl
out and drop to the ground, where they grow rapidly under favorable
conditions, feeding on nitrogenous matter, dry droppings, etc. The full
grown larva, which is not unlike other flea larvae, is about 4 mm in
length, reaching this stage in about two weeks. The larva then spins a
cocoon, pupates, and in about two weeks (9 to 19 days) emerges as a
fully developed flea, The life history requires from 30 to 60 days, Eggs
are also deposited in the dust or dry droppings of poultry or in old
nests, etc.
The fleas are most likely to attack the skin around the eyes, the
wattles and comb, and the anus or other bare spots, The ulceration and
wart-like elevations around the eyes often become so aggravated that
blindness occurs, the host is unable to find its food, and death results.
Since this flea also lives on dogs, cats, rats, quail, blackbirds, and spar-
rows, suitable precautions should be taken to exclude these from chicken
pens,
The western hen flea, Ceratophyllus niger Fox, is considerably larger
than the sticktight and does not attach except when feeding and then
only for a brief period. It readily attacks man and cats and dogs. It
breeds primarily in fowl droppings. The European hen flea, Ceratophyl-
lus gallinae Schrank, which has habits similar to C. niger Fox, is also at
times a serious pest of poultry (Stewart, 1927).48
Fleas in the household. Very few species of fleas are annoying to house-
hold pets, Among these are particularly the dog and cat fleas. Cteno-
cephalides canis ( Curt. ) and Ctenocephalides felis (Bouche) (Fig.
149), and the so-called human flea, also known as the sand flea, Pulex
irritans Linn. While the common name might imply that there is a
specific host relationship, this is not the case, since interchange of host
species is quite usual. Cat and dog fleas readily attack humans, and the
human flea is often remarkably abundant on swine.
Fleas in the house generally indicate that cats or dogs or both are
present or have been present fairly recently. Fleas may be carried on
clothing into the house from pig pens or from flea-infested public meet-
ing places. The exclusion of cats and dogs or their proper management
is necessary to prevent flea infestations. Cats and dogs (unless properly
FIG. 149. Ctenocephalides felis, the cat flea; male (left), female (right). X 17.
bedded), as well as rats, must be excluded from the basement and
from beneath the house.
Ordinarily fleas lay their eggs on the infested animal, but because
the eggs are dry, they drop off when the host shakes itself. For this
reason mats should be provided upon which the animals may sleep at
night, and these should be shaken off every day or two over fire or into
kerosene (see flea control). The eggs are minute glistening white objects.
The incubation period varies considerably, but they usually hatch in
from five to six days, sometimes less, and the worm-like sparsely haired
larvae emerge. The larvae feed on particles of dry blood, fecal matter,
and various organic substances collected in corners and crevices. The
larvae are quite active and in two to four weeks reach a length of about
one-fourth inch and then spin a crude cocoon in which they pupate. The
flea emerges from the cocoon in about a week; thus three to four weeks
are required for the entire life history of a common house flea under favor-
able conditions. Undisturbed mats, rags, and carpets favor the develop-
ment of fleas. All carpets or matting tacked down and covering the floor
should be dispensed with and smaller rugs substituted. Houses that have
been vacant for several weeks may be badly infested with fleas because
these insects are able to live without food for several weeks.
FLEAS 447
Fleas as intermediate hosts of cestodes. Although Melnikoff in 1867
showed that the biting louse of the dog, Trichodectes canis DeGeer,
serves as an intermediate host of the double-pored dog tapeworm, Dipyl-
idium caninum (Linn.); it has since been shown by other workers that
fleas play a more important role in the transmission of this tapeworm,
particularly the cat and the dog flea, Ctenocephalides felis (Bouche) and
Ct. canis (Curt.). Although Dipylidium caninum (Linn.) is a tapeworm
of dogs, cats, and certain wild carnivores, it also occurs in man, particu-
larly in young children. The tapeworm measures from 20 to 70 cm in
length; the mature proglottids are shaped like pumpkin seeds, and each
has a double set of reproductive organs with a genital pore on each side.
The scolex has a rostellum which is armed with three to seven circlets
of spines and has four deeply cupped oral suckers. The embryonated
eggs of the tapeworm are discharged in the fecal material of the host
and are ingested by the larval flea and develop into cysticercoids in the
body cavity of the insect. Thus the mature flea is infected and when
ingested by a cat or dog or human, the cysticercoids are liberated and
develop into tapeworms in the animal's digestive tract.
A common tapeworm of rats and mice, rarely of man, Hymenolepis
diminuta (Rudolphi), has numerous intermediate arthropod hosts, among
them Nosopsyllus fasciatus (Bosc) and Xenopsylla cheopis (Roth.).
Flea control. With a substantial background of knowledge concerning
the parasitic habits, the life history, and particularly the ecology of fleas,
control of these pestiferous, disease-bearing insects can be accomplished.
Management of host animals and the practice of good sanitation in the
household and in public places are fundamental principles which must
not be neglected· granting the effectiveness of the newly discovered in-
secticides, the proper use of which must be understood as well as their
limitations.
Dichloro-diphenyl-trichloroethane (DDT) may be applied either as
a 10 per cent dust (particularly when used in murine typhus fever control
operations) or as a 5 per cent DDT-kerosene solution for application as
a spray to floors and beneath rugs in the house and sleeping places of
cats and dogs. One gallon of 5 per cent DDT-kerosene solution sprayed
lightly over areas of 1,000 to 2,000 square feet will completely eradicate
fleas. DDT is not a repellent, and it kills slowly. It retains its killing
effect against fleas for about two months.
When DDT is used as a dust it is mixed with pyrophyllite, Fuller's
earth, calcium dust, or walnut shell flour and is applied with a hand
duster or with a hand shaker. That DDT dust kills fully grown flea
larvae [Ctenocephalides felis (Bouche)] as well as adults was shown
by Kerr (loc. cit.), who used among other rates a 5 per cent DDT-
pyrophyllite dust. All larvae died within 24 hours after treatment. Adult
fleas remained normal in behavior for about 10 minutes after exposure
to toxic doses, then followed a period of increasing activity, with the fleas
jumping vigorously in 15 to 20 minutes; within an hour they were in-
capable of movement except a slow waving of the legs.
Wherever pets sleep or are accustomed to lie down DDT dust
(10 per cent recommended) is applied where fleas will come in contact
with it. The dust should be forced into cracks and crevices and in all
places where fleas may breed. Excess powder if objectionable after a few
days can be wiped up with a cloth or removed with a vacuum cleaner.
Flea-infested basements as well as yards and other out-door areas should
be Similarly dusted.
Dogs may be safely dusted with DDT; often only a line of 10 per
cent dust along the middle of the back is sufficient to kill all the fleas
and to prevent reinfestation for several weeks. Since DDT agitates fleas
before killing, this activity causes the dog to scratch and bite vigorously
until the insects cease crawling about.
Although DDT dust is not absorbed through the skin of mammals,
cats should not be treated with DDT because of their cleaning habits;
however, treating the cat's bed is usually sufficient. Flea powders con-
taining rotenone and pyrethrum are safe to use on cats.
In the use of DDT dust (10 per cent DDT) in murine typhus con-
trol, the United States Public Health Service points out that this is aimed
at the flea link in the typhus chain. Proper dusting of rat runs and har-
borages with DDT combats fleas both on and off their rodent hosts.
With regard to the use of DDT for the control of murine typhus
fever the United States Public Health Service points out that rats in
passing over the dusted places, pick up a considerable amount of dust
on their feet, bellies, and tails, and fleas on the rats come in contact with
the DDT dust and, if exposed to it for a sufficient length of time, are
killed. Thorough residual dusting (not too heavy) of rat runways, both
inside of buildings and in outside areaways, is essential not only to
ensure killing fleas on the rats but also to kill the fleas (off the animals)
in or near rat runs, nests, or harborages. Treatment of premises and build-
ings should be repeated at intervals of two or three months throughout
the flea breeding season. Success depends upon a good knowledge of the
habits of rats and careful inspection.
Flea repellents. Tests made by Lindquist, Madden, and Yates 49 show
that insect repellents used during World War II, dimethyl phthalate,
Rutgers 612, and indalone, are effective flea repellents giving an average
repellent period of IS9-260 minutes, the first giving the longest. Tests
made with a mixture of one per cent pyrethrins and 2 per cent IN-9S0
in mineral oil gave even a longer repellency.
Rodent control. Officials responSible for the prevention and control
FLEAS 449
of plague, murine typhus, and other infections traceable to rats and

other rodents must recognize the fact that prevention and eradication of
such diseases depends on intelligent rodent control, a function that
should be placed on a much higher plane than is usually implied in the
duties of the so-called "rat catcher." The position of rodent control officer
calls for a high degree of intelligence, much technical training, and a
substantial understanding of the whole problem; this person deserves
the same respect and credit that is accorded any other individual who
functions in the field of public health. The rodent menace must be met
with intelligence; rats in particular must not be tolerated; they constitute
not only a sinister menace to health, sufficient in itself to condemn them,
but if further incentive to their extermination is necessary it is reported
on good authority that a rat eats $2.00 worth of food each year and
destroys ten times this amount by fouling, gnawing, and otherwise dam-
aging grain, stored products, and other articles; 200,000,000 bushels of
grain are reported to be destroyed or contaminated by rats annually
in the United States; thus rats cause an aggregate annual loss of over
$2,500,000,000 in the United States alone.
Rats. Together with the house mouse, Mus musculus Linn., the cos-
mopolitan species of rats belong to the order Rodentia (gnawing mam-
mals ), family Muridae. Rattus rattus norvegicus (Erxleben), the brown
rat or sewer rat, better known as the Norway rat, is the largest of
the common house rats. Full-grown adults measure about 16 inches
in total length (tail TYz inches), and weigh 11 to 12 ounces (exception-
ally 24 ounces). The nose is blunt; the ears are of moderate size and
slightly hairy; the tail is scaly, nearly naked, and blunt-ended. The body
fur is coarse and generally brown, with scattered black hairs; the under
parts are pale grey to yellowish white (Fig. 150). Rattus rattus rattus
(Linn.) is the black rat, which is almost solidly blue-black in color; it
is smaller and more slender than the Norway rat. The nose is sharply
slender, and the ears are noticeably large, thin-membraned, and naked;
the tail is slender and longer than the body and head. Rattus rattus
alexandrinus (Geoffroy-Saint Hilaire and Andouin) is called the Alex-
andrine or roof rat, and also the tree rat. In color it is grayish brown
above and white or yellowish white below (hence sometimes called
yellow-, gray-, or white-bellied tree or roof rat). It is usually classified
as a subspecies of the black rat, as it has the same general characteristics
in form and habits. Both can climb readily and may nest in trees, palms,
and attics of houses. Rattus rattus frugivorus Rafinesque is called the
white-bellied frugivorus rat and is said to be an outdoor rat that does
not live close to man. It is usually regarded simply as a color phase of
R. r. alexandrinus. Rattus hawaiiensis Stone is now a common species
of rat on Maui and other Pacific Islands. It is called the Polynesian rat
and is a small species averaging about 60 grams in weight. It lives in
grass and brush areas away from human habitations. Doty 50 states that
it is very destructive to sugar cane. In color it is "cinnamon brown above,
cinnamon buff on the sides, and light buff or buffy white below, strongly
mixed with black hairs on the back and sides. The feet are nearly white
above."
Rat biology. Since successful control is based on a full knowledge of
the biology, particularly feeding habits and nesting habits, of the several
\ species of rodents involved in a given problem, much attention must be
given to this subject. Sufficient literature is available to serve as a guide,
FIG. 150. House rats. (A) Rattus r. rattus, the black rat; (B) Rattus n. norve-
gicus, the brown rat; and (C) Rattus r. alexandrillus, the roof rat. X .13.
but there is no substitute for field observation and experience. Each

species of rat must be studied carefully inasmuch as their habits may
differ conSiderably; a certain procedure which results successfully in
one case may fail utterly in another.
Most species of rats are active at night; however, Rattus hawaiiensis
feeds commonly during daylight hours; also unlike other species which
are omnivorous, the Polynesian rat lives on grass and weed seeds along
with some vegetative parts of plants such as sugar cane (Doty, lac. cit.).
Likewise this latter species is said to be quite docile, while the cos-
mopolitan species listed above are as a rule vicious. Norway rats are
cannibalistic and will devour young rats in nests, eat dead rats, and
attack one another under stress. A knowledge of food habits and feeding
habits is particularly important in the use of poisoned baits and pre-
baiting.
The Norway rat stays mainly at the ground level and seldom goes
FLEAS 451
above the first floor of a building. It burrows underground a foot or two

and makes tunnels several feet in length with several hidden exits or
"bolt holes." Crude nests are made in burrows under buildings, beneath
piles of lumber, in garbage dumps, along banks of ditches and streams.
It favors pigpens, slaughterhouses, barns, chicken yards, and sewers.
Roof rats, Alexandrine and black rats, climb readily and often live in
attics and upper stories of buildings; they commonly nest in trees, es-
pecially tops of palms and in dense hedges or vines growing on fences.
Like the Norway rat, the roof rat can swim readily. Alexandrine and
- black rats are far more common on ships than the Norway rat.
According to Storer51 female Norway rats average 8 to 9 embryos
(extremes 2 and 17) at a pregnancy and may breed four or more times
during a year. Pregnancy lasts 21 to 22 days. The young are able to
shift for themselves when about 3 weeks old. A female may first breed
at the age of 4 to 5 months. Although it is not known how long wild
rats live (some authorities believe about three years), the population is
probably largely replaced by new·individuals each year. One pair of rats
may produce a population of more than 1,500 by the end of a year.
Early spring and summer are the periods of greatest production, though
young rats are found during any month of the year.
There are nine signs made by rats which are useful in detecting
infestations (Rats and their Control; Rat and Mosquito Control Com-
mission, Chamber of Commerce of Honolulu, 1943): (1) droppings,
(2) runways, (3) tracks' over dust or soft surfaces, (4) burrows, (5)
gnawings, (6) live rats, (7) dead rats, (8) nests, (9) rat odor. Storer
(lac. cit.) lists in addition sound, i.e., squeals, scampering in an attic;
smears along beams, etc., made by contact of the body of the rat; and
urine stains. With adequate information as here suggested, a fairly
accurate rat census can be made.
Methods of rat control. One should aim at permanent control as the
only satisfactory solution of the rat problem. There should be good
sanitation the year round. Good housekeeping is good insurance against
rats; cleanliness is the rat's worst enemy. The United States Fish and
Wildlife Service (Circular 13, 1948) points out that food and shelter
are the two most important factors in a rat's existence. Any program of
rat control to be successful must incorporate these four major phases:
( 1) elimination of rat harborages, (2) elimination of food supply for
rats, (3) rat proofing of buildings, and (4) destruction of rats.
Rat proofing and rat stoppage. Silver, Crouch, and Betts 52 state that
the rat problem will have been largely solved when rat proofing becomes
the regular practice. Storer (lac. cit.) points out that rat prooling (ex-
clusion) is not enough; such construction must be frequently inspected
and kept in good repair. Rat proofing can be done at the time of con-
struction at slight extra cost, but when applied to old buildings it is
usually an expensive procedure. However, exterior rat proofing, i.e.,
rat stoppage can be done inexpensively (Holsendorf and Clark53 ). Loss
occasioned by rats on farms in the United States is reported to amount
to $63,000,000 annually, much of which could be averted by ratproofing
barns, corncribs, granaries, poultry houses, and pig pens (building out
and starving out rats) as described by Silver, Crouch and Betts (lac. cit.).
Cutting off the rat's food supply (starving out the rat) refers not only
to ratproofing so as to make food inaccessible; it also refers to the sani-
tary disposal of refuse from the household, garbage, slaughter-house
refuse, feed refuse in connection with piggeries, poultry houses, barns,
feeding lots, etc. Cleanliness, neatness, good housekeeping in general, and
efficient farm management will aid materially in the control of rats and
other rodents.
Destruction of rats. In the use of rat poisons a system of pre baiting
and baiting is employed. Since the rat poison is usually mixed with rat
food, one should have some knowledge of the food habits of these ro-
dents. Rolled oats and other cereals, corn meal, bread crumbs, diced
bread, raw meat (horse meat is acceptable to rats), and raw fish are
the commonest baits used. Pre baiting is a practice based on the fact that
rats are suspicious and avoid new objects, even a new food. If a clean
prebait is offered they will overcome their fear in a few nights (two or
more) and will accept the new food. Small piles of clean unpoisoned
baits are placed at selected sites or in special containers for four or five
nights replenishing the supply whenever the bait is eaten. On the fifth
or sixth day the unpoisoned prebait is replaced with poisoned bait. If
the poisoning is successful the above-described routine may not need
to be repeated for several months; otherwise a follow-up campaign may
be undertaken in a couple of weeks.
Among the commoner less dangerous rat poisons are the following:
red squill, antu, zinc phosphide, and barium carbonate; but 1080 and
thallium sulphate should only be used by trained professional operators.
(1) Red squill is obtained from an onion-like plant (Urginea maritima)
native to the Mediterranean area. The bulbs are sliced, dried, and
ground to a fine reddish powder. Different batches of this powder vary
in toxicity; hence squill of lower toxicity is fortified until its toxic
strength reaches the point where rats are killed by 500 to 600 milligrams
per kilogram of live rat (such tests are known as bioassays). For use
one part by weight of red squill powder is well mixed with 9 parts of
bait such as raw hamburger steak (horse meat is acceptable to rats)
mixed with rolled oats or bread crumbs wet with water. A convenient
way of handling the bait is to place a small quantity about the size of a
marble in a paper square and twist in the form of a "torpedo." A little
FLEAS 453
glycerin mixed with the bait prevents hardening and sodium sulphite
may be used as a preservative. Red squill is the safest rat poison for
general use since experiments show that cats, dogs, chickens, pigeons,
and hogs usually will not eat it, probably because of its disagreeable
taste, or will quickly vomit any which may }lave been eaten. Red squill
is a poison, hence should be treated accordingly and be placed out of
reach of children and pets.
(2) Antu is an abbreviation for the chemical alpha-napthyl-thiourea,
a light, gray powder, quite insoluble in water, highly stable and non-
irritating to the human skin. It is highly toxic to the Norway rat in con-
centrations of 1 to 1 liz per cent, but much l(~ss toxic to Alexandrine and
black rats. This deficiency can be corrected 1:)y increasing the percentage
of "antu" in baits to 5 per cent. It kills rats by causing an accumulation
of body fluids within the chest cavity. Sublethal dosage quickly builds
up a tolerance. Antu is particularly toxic to dogs and young chicks; it
can also poison pigs. Monkeys tolerate large quantities, hence it may not
be dangerously toxic to human beings; however, antu should be guarded
like any other poison to prevent children, pets, domestic animals, or
foodstuffs from coming in contact with it. Keep antu out of the mouth;
do not eat or smoke when handling it. As a rat poison it is used in baits
in concentrations referred to above, or it cafl be used as a dust on run-
ways or pumped into burrows in a mixture of 20 per cent antu, 2 per
cent DDT, and 78 per cent pyrophyllite. This formula tends to control
rat fleas as well as the rats.
(S) Zinc phosphide is a heavy dark-gray powder with a faint odor
of phosphorus. It is used in baits at a percentage of from 2l1z to 5 per
cent.
(4) Barium carbonate is a fine white powder. The proportions used
are: bait, 4 parts by weight, and barium ~arbonate, 1 part by weight.
It is slow acting and although regarded as mild it may still be fatal to
chickens, dogs, and cats; hence great care I11ust be exercised in its use.
It must be inaccessible to children and to pets and other animals. It is a
pOison.
(5) Compound 1080 (sodium fiuoroacetate) is a violent poison
(Ward and Spencer54 ). Water poisoned with liz ounce of 1080 per gallon
will kill monkeys on the basis of about 3 Oullces of the solution to a 150-
pound animal. It is highly fatal to man. Cats and dogs are almost sure to
be fatally poisoned if rats, mice, or other rodents killed with 1080 are
eaten. Compound 1080 should be used only by carefully instructed, re-
liable personnel competently supervised. It should not be used in dwell-
ings.
(6) Thallium sulfate is a heavy-metal poison obtained from smelters.
It is used by governmental agencies for the control of both rats and neld
rodents. For thallium, as for 1080, there is no certain antidote. It is a
slow acting poison in contrast to 1080 which acts very quickly.
Toxic gases such as calcium cyanide, sulfur dioxide, methyl bromide,
and carbon monoxide (from automohile exhaust) are commonly forced
into rodent burrows by means of special pumps to kill rats and other
burrowing rodents.
Use of traps. In trapping rats the proper placement of the trap is far
more important than the selection of a bait. Bacon strips, a piece of fresh
fish, or bacon-scented oatmeal is better than cheese as a bait according
to the "Fish and Wildlife Service." Such baits should be tied firmly to
the trigger of the trap.
Dead rats resulting from poisoning, gassing, trapping, or any other
cause should be treated copiously with kerosene, boiling water, or an
insecticide before disposal, to destroy any fleas still present on the body.
Incineration or deep burial of dead rats is good practice. Deodorants
such as powdered activated charcoal may be used where rats have died
within walls. Storer (loc. cit.) refers to a commercial product, Hyamine
1622, as useful to reduce such odors.
Field rodents. Among the more important field rodents are the ground
squirrels belonging to the genus Citellus, family Sciuridae. They are
important reservoirs of sylva tic plague and tularemia and may cause·
losses to cereal crops amounting to from 10 to 15 per cent or more. They
materially reduce the forage in pastures and may damage ditch banks
and levees by burrowing.
Storer 55 (1947) states that there are seven varieties or subspecies of
ground squirrels in California, each occupying a separate area. The most
widespread is the dark-colored Douglas ground squirrel, Citellus beechyi
douglasii ( Richardson), with much black between the shoulders; it
occurs northward from San Francisco Bay throughout the region west
and north of the Sacramento and Feather rivers. The brownish Beechey
ground squirrel, Citellus beechyi beechyi (Richardson) (Fig. 142) oc-
cupies coastal California from the Golden Gate and Carquinez Strait
south nearly to San Diego. The gray-toned Fisher ground squirrel,
Citellus beechyi fisheri (Merriam), inhabits the greater part of central
California from the Feather and Sacramento rivers south to the southern
end of the San Joaquin Valley. The Oregon ground squirrel, Citellus
oregonus (Merriam), a short-tailed brownish-gray species of the north-
eastern part of California, is essentially an inhabitant of grasslands. The
Belding ground squirrel, Citellus beldingi (Merriam) is known as the
picket-pin; it is also bob-tailed and is found in the high Sierra at eleva-
tions of 6,500 feet to 12,000 feet.
Every ground squirrel has two thin, internal cheek pouches, opening
inside the lips, one on each side of the mouth, which are used to carry
FLEAS 455
food. Storer points out that this characteristic is important in control
because ground squirrels may be killed by absorbing poison on baits
through the lining of the pouches.
Habits. Ground squirrels find their food on the surface of flat country,
hillsides, or embankments. They are active by day during the warmer
season of the year and on warm days in winter. All species 'dig burrows
for safety, for shelter during very hot or rainy weather, for hibernation,
for storage of food, and for the rearing of young. Burrows are about 4
inches in diameter, within 2112 to 4 feet below the surface, and may
reach a length of 30 feet; occasionally they are much longer. Entrances
to the burrows are always open. These tunnels, often branched, are made
in hillsides, in ditch banks, in railroad embankments, and in Hat lands.
Storer describes a complicated system of tunnels with a total length of
471 feet and 33 opening.
During the rainy months ground squirrels feed chiefly on green
herbage and seeds on the ground; when seed crops ripen these animals
collect unhulled seeds of both wild and cultivated plants in their cheek
pouches and hide these in caches in the ground and in various crevices
for later use.
Estivation and hibernation. In the lowlands ground squirrels are
reported to estivate as early as mid-May, whereas in the high mountains
they do not begin hibernation until as late as early August. Emergence
occurs in late winter or early spring. Storer (lac. cit.) points out that
estivation explains why old breeding adults suddenly appear in spring,
after all squirrels active in a field during autumn had been killed, and
there seemed no chance for migration from surrounding fields. It is
extremely important to carryon intensive control in the spring and
early summer when all squirrels are active. During hot summer weather
and in the autumn and early winter, some of the squirrels may be under-
ground and out of reach. There is some doubt whether dormant animals
are killed by gassing their burrows with carbon disulfide.
Breeding. Ground squirrels probably produce only one litter a year,
of which about 5 or 6 are said to survive long enough to appear above
ground. The breeding season is during the months of December to April.
The young are seen in greatest numbers during May and June, when
they scatter to new territory or move to unoccupied old burrows. Control
operations, therefore, should be espeCially intensive in late winter and
early spring.
Ground-squirrel control. Storer (lac. cit.) states that there are six
general means of controlling ground squirrels, viz.: (1) poison bait, (2)
poisonous gases, (3) trapping, (4) shooting, (5) exclusion, and (6)
encouragement of natural enemies.
Strychnine-coated barley. In the use of poison for squirrels several
important factors must be considered; namely, it must not be distaste-

ful to the rodents and must enter the circulation readily; the poison
must be applied to food readily eaten by the squirrels and at a time
when the usual green food is at its minimum. It has been found that
strychnia sulphate (the pure alkaloid should not be used) is most effec-
tive, but the bitter taste must in some manner be concealed. The use of
whole barley, i.e., threshed but still retaining the husk, is recommended.
In this form the barley is not eaten by birds and is most acceptable to
the rodents; it is also cheaper in this form. Wheat is very acceptable to
the rodents but when poisoned is very destructive to birds, particularly
quail, doves, and other grain eaters.
Piper has devised a formula whereby the bitter taste of the strych-
nine is delayed about two minutes, thus enabling the squirrel to fill
its cheek pouches before the bitter taste is noted. The formula is as
follows: .
Formula for strychnine-coated barley. Following is a government formula
for preparing poisoned barley for California ground squirrels.
Barley (clean whole grain) 16 quarts
Strychnine (powdered alkaloid) ............ . 1 ounce
Bicarbonate of soda (baking soda) ............................ . 1 ounce
Thin starch paste .. ... . .. .. .... .. .... .. ... . ... .. ................ . % pint
Heavy corn sirup ................ . ............................ . ;.4 pint
Glycerin ....................................................... . 1 tablespoonful
Saccharin Ylo ounce
Mix thoroughly one ounce of powdered strychnine (alkaloid), and one
ounce of common baking soda. Sift this into three-fourths pint of thin, hot
starch paste and stir to a smooth, creamy mass. (The starch paste is made by
dissolving one heaping tablespoonful of dry gloss starch in a little cold water,
which is then added to three-fourths pint of boiling water; boil and stir con-
stantly until a clear, thin paste is formed.) Add one-fourth pint of heavy corn
sirup and one tablespoonful of glycerin and stir thoroughly. Add one-tenth
ounce of saccharin and stir thoroughly. Pour this mixture over 16 quarts of
clean barley and mix well so that each grain is coated.
Caution.-All containers of poison and all utensils used in the preparation
of poisons should be kept plainly labeled as POISON and out of reach of
children, irresponsible persons, and livestock.
Grain poisoned with strychnine placed in proper containers will re-
tain its poisonous character and remain effective for an indefinite period,
but heavy dews and rain may remove the poison and destroy the effec-
tiveness of the bait. Therefore, this method is applicable during the dry
season only. Simpson56 reported that 30 kernels in the cheek pouches of
squirrels rapidly prove effective, whereas 60 or 90 or more in the stomach
may produce only a few convulsions and recovery ensues. He says, "This
FLEAS 451
fact should be remembered in placing poison, for by scattering the
grain a few kernels here and there near the burrow the squirrel is in-
duced to store the grain temporarily in the cheek before a sufficient
quantity is obtained for a meal. . . . " It should be scattered where the
squirrel is accustomed to find food, and will probably be found most
efficient if placed early in the morning, between the hours of 3 A.M.
and 7 A.M.
Other poisons such as zinc phosphide may be used with such baits
as oat groats, whole oats, and whole barley; mineral oil, petroleum jelly
or lecithin hold the poison to the bait.
1080. Field rodents such as ground squirrels and prairie dogs are
very susceptible to 1080 poisoning. Effective control has been obtained
by the use of concentrations of 1 to 2 ounces mixed with 100 pounds
of grain; however, the control of field rodents is a complex procedure
and should only be undertaken by experienced persons authorized by
law to do so. The careless use of 1080 will endanger domestic livestock
and beneficial wild life found in the same habitat. The secondary hazard
to dogs, cats, pigs, and carnivorous wild life follOWing the use of 1080
in field rodent control is considerable.
Gassing with carbon bisulfide. Carbon bisulfide is an effective fumi-
gant for ground squirrels and is commonly employed. Carbon bisulfide
is obtained commercially in the form of a liquid, which is readily vapor-
ized or is converted chemically into other gases. While it is a useful
material against ground squirrels, there are some objectionable features;
namely, it is very inflammable, must be kept in tightly closed containers,
and, under certain conditions may explode; furthermore, during the dry
season if "exploded" in the burrow there is danger of igniting dry grass
or other inflammable material in the vicinity. If handled with as much
I care as gasoline, for example, the danger is not great. The advantages
in its use are these: it is readily converted into a poisonous gas, diffuses
quickly, destroys life rapidly, and can be used most readily during the
rainy season, when green food is abundant, and the use of poisoned grain
is not practicable.
Carbon bisulfide is applied in two ways: with a special pump to
force the fluid or gas into the burrow or by soaking waste balls in the
fluid and then placing them in the burrow. The balls of waste may be
ignited, but this is hazardous. In either case it is suggested that from
one to three days prior to the application of the poison all squirrel bur-
rows in the area to be treated should be stopped with earth. The holes
found opened indicate the burrow in which there are squirrels.
The method of applying carbon bisulfide by the ignition method is
as follows: to handle a large area to the best advantage two men should
work together.
One man is provided with a supply of "waste," "sacking," or other ab-
sorbent material, divided into a number of small balls about half the size of
the fist. The bisulfide is carried in an ordinary one-gallon oil can, and re-
filled from time to time from a supply kept in a cool place out of the sun.
He is supplied with matches. (Matches are dangerous, hence other methods of
exploding the gas should be used.) His "pardner" carries a mattock or long-
handled shovel. On arrival at an opened squirrel burrow, a ball of "waste"
is saturated with two ounces of bisulfide, dropped deeply in the burrow and
then a match applied. After a moment's time the man with the shovel stops
with earth this burrow and all other burrows near from which the gas escapes.
On subsequent inspection of the field all opened burrows will indicate holes
lacking affective treatment.
Exploding the bisulfide causes considerable gas to escape, but

the ignition produces a violent chemical reaction and as a result sufficient
oxygen from the air combines with carbon and sulphur elements to produce a
volume of gas three times that which the original bisulfide would produce on
evaporation. The gases produced, carbon dioxide and other sulphur dioxide,
in the proportion of one to two, seem just as effective as bisulfide of carbon,
and the method is superior in that the explosion produced drives these gases
deeply into the burrow.
Two ounces of 60 cc of the bisulfide produce about 12 gallons of gas.
To use the gas unexploded, simply omit igniting it.
A much cheaper and more efficient method of destruction with car-
bon bisulfide was early devised by Long,57 namely, a pump with a
device measuring the quantity of liquid and serviceable at all seasons of
the year. This pump loaded with nine pints of refined bisulfide weighed
25 pounds. Only one-half ounce (15 cc) is required for each hole, against
two ounces by the ignition method, and it is claimed that the men using
the pump have been able to treat from 200 to 250 holes with each gallon
of the bisulfide, against 50 to 60 holes per gallon with the waste ball
method above described.
The use of Long's pump is described as follows: "Insert the hose in
the squirrel hole at least one foot; then run one-half ounce of bisulfide
from the reservoir into the measuring cup; then turn cock with handle
down to allow liquid to run into vaporizing chamber, meanwhile cover-
ing the hole with dirt with the aid of a mattock. Then pump thirty
strokes (in cold weather use one ounce with forty strokes). This equals
12 cubic feet or 1.5 per cent bisulfide gas. Withdraw the hose, close hole
opening by stamping in the dirt with the heel and proceed to the next
hole." Twenty minutes' to a half hour's treatment with air containing 2
per cent of carbon bisulfide is certain to be fatal. Improved carbon bisul-
fide pumps are on the market.
Gassing with methyl bromide. With the discovery that rodent fleas
FLEAS 459
may serve not only as vectors of sylvatic plague but also as pseudo reser-
voirs, the necessity of flea control as well as rodent control is indicated
in directing sylvatic plague suppressive measures. Carbon bisulfide is
very uncertain and inefficient in its lethal action on adult fleas. Berry 58
found that methyl bromide, CH3 Br, 10 cc of liquid per burrow opening,
may be used effectively against ground squirrels in wet or dry soils and
at various temperatures. Methyl bromide is colorless, practically odor-
less, and noninflammable, but it is a dangerous fumigant and should be
used only by persons well informed concerning its hazards.
Stewart and Mackie 59 (1938) have shown that ground-squirrel fleas
of all stages are susceptible to this fumigant but that the adults are
more easily killed than those in immature stages. These authors also
found that the dosage used as a rodenticide, approximately 10 cc per
burrow opening, is sufficient to kill the fleas in all stages of development.
They point out that this fumigant has a peculiar characteristic in that
"it does not kill immediately but manifests its toxicity in delayed kill
without, however, exciting either the rodent or the flea before stupefy-
ing them."
Trapping and shooting. Where ground squirrels are digging into
ditch banks, and in other cases where they must be disposed of promptly
at any cost, special means must be adopted. Trapping and shooting are
valuable under such circumstances. Both of these methods can be used
at any season.
Natural enemies. Among the natural enemies of ground squirrels
listed by Storer are coyotes, badgers, weasels, wildcats, red-tailed hawks,
golden eagles, and gopher snakes. Badgers, weasels, and snakes cap-.
ture the ground squirrels in their burrows. Wildcats and coyotes lie in
wait near the burrows until the squirrels venture forth in search of food.
"Dixon examined 186 stomachs of wildcats from 40 different localities
in California; 26 contained ground squirrels and these with other rodents
were found to constitute more than half of the food. Hawks and eagles
swoop down on the squirrels from the air. The importance of preserving
as many as possible of these native enemies of the ground squirrels is
evident." When poison is used in the control of ground squirrels, reason-
able precaution should be exercised to prevent killing their natural
enemies.
Recently several investigators have called attention to the possible
relationship of flesh-eating birds to the epidemiology of sylva tic plague.
Jellison 60 states, "The abundance and variety of rodent fleas found in
the nest of a burrowing owl (Speotyto cunicularia) suggest that this
species may prove of particular interest. qasts from predatory birds fed
plague-infected guinea pig tissue were consistently infectious." Wheeler,
Douglas and Evans 61 report collecting sticktight fleas, Echidnophaga
gallinacea, from the same species of owl in Kern County, California,

during a plague epizootic. Seventy individuals of this flea taken from
one owl were mass-inoculated into a test guinea pig which proved posi-
tive for plague. This is probably the first record, according to these
authors, of a bird host as a carrier of plague-infected parasites, and the
first demonstration of natural plague infection of this species of flea.
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39. Jellison, Wm. L., and Good, Newell E., 1942. Index to the Literature
of Siphonaptera of North America. Washington, D. C.: U. S. Public Health
Service, in Nat. Inst. Health Bull., no. 178. 193 pp.
40. Mellanby, Kenneth, 1933. "The influence of temperature and humid-
ity on the pupation of Xenopsylla clzeopis," Bull. Entomolog. Research, 24:197-
202.
41. Jordan, Karl, 1932. "A new Xenopsylla from Hawaii," Novitates
ZoOlogicae, 38:264-66.
42. Williams, C. L., 1929. "A rat and rat-flea survey of ships at the port
of New York," U. S. Public Health Service, Pub. Health Rep., 44:443-76.
43. Hasseltine, H. E., 1929. "Rat-flea survey of the port of Norfolk, Va.,"
U. S. Public Health Service, Pub. Health Rep., 44:579-89.
44. Fox, Carroll, and Sullivan, E. C., 1925. "A comparative study of rat-
flea data for several seaports of the United States," U. S. Public Health Serv-
ice, Pub. Health Rep., 40:1909-34.
45. Williams, C. L., 1932. "Rat infestation inspection of vessels," U. S.
Public Health Service, Pub. Health Rep., 47:765-800.
46. Faust, E. C., and Maxwell, J. A., 1930. "The finding of the larvae of
the chigoe, Tunga penetrans, in scrapings from human skin," Arch. Dermat.
& Syph., 22:94-97.
47. Parman, D. C., 1923. "Biological notes on the hen flea, Echidnophaga
gallinacea," ]. Agric. Research, 23:1007-9.
48. Stewart, M. A., 1927. "A means of control of the European hen flea,
Ceratophyllus gallinae Schrank," ]. Econ. Entomol., 20:132-34.
49. Lindquist, A. W.; Madden, A. H.; and Watts, C. N.; 1944. "The
use of repellents against fleas," ]. Econ. Entomol., 37:485-86.
50. Doty, R. E., 1945. "Rat control on Hawaiian sugar cane plantations,"
Hawaiian Planters' Record, vol. 49, no. 2, 239 pp.
5!. Storer, Tracy I., 1948. Control of Rats and Mice, Berkeley: Univ.
Calif., in Agric. Exten. Servo Circ., no. 142. 37 pp.
52. Silver, James; Crouch, W. E.; and Betts, M. C.; 1942. Rat-proofing
Buildings and Premises. U. S. Fish and Wild Life Service Washington, D. C.:
Conservation Bull., no. 19. 26 pp.
53. Holsendorf, B. E., and Clark, P. W., 1937. "The rat and rat-proof
construction of buildings," U. S. Public Health Service, Pub. Health Rep.,
Suppl. no. 131. 68 pp.
54. Ward, J. C., and Spencer, D. A., 1947. "Notes on the pharmacology
of sodium fluoroacetate-Compound 1080," J. Am. Pharm. A., 36:59-62.
FLEAS 463
55. Storer, Tracy I., 1947. Control of Field Rodents in California, in Calif.
Agrie. Ext. Cire., no. 138. 51 pp.
56. Simpson, F., 1911. "Ground squirrel eradication," Calif. State Bd. of
Health Bull., 6:507-12.
57. Long, John D., 1912. "A squirrel destructor," U. S. Pub. Health Serv-
ice, Pub. Health Rep., 27:1594-96 (4 plates).
58. Berry, C. E., 1938. "Methyl bromide as a rodenticide," Bull. Dept.
Agric., State of Calif., 27:172-80.
59. Stewart, M. A., and Mackie, D. B., 1938. "The control of sylvatic
plague vectors," Am. ]. Hyg., 28:469-80.
60. Jellison, William L., 1939. "Sylvatic plague: Studies of predatory
and scavenger birds in relation to its epidemiology," U. S. Public Health Serv-
ice, Pub. Health Rep., 54:792-98.
61. Wheeler, C. M.; Douglas, J. R.; and Evans, F. C.; 1941. "The role
of the burrowing owl and the sticktight flea in the spread of plague," SCience,
94:560-61.
CHAPTER XX
._.
TICKS AND TICK-BORNE DISEASES
CLASS ARACHNIDA, ORDER ACARINA,

SUPERFAMILY IXODOIDEA
Introduction. Probably all species of vertebrate animals higher than

fishes are subject to attack by ticks, but particularly mammals, whose
warm blood is highly attractive to these parasites. The food of ticks
consists entirely of blood and lymph, and both males and females are
bloodsuckers, as are all immature stages. Hunters have long observed
tremendous infestations on the bodies of wild animals. Stockmen suffer
enormous losses due to ticks on cattle, horses, and other stock; poultry
is often severely parasitized. Hunter and Hooker/ United States Bureau
of Entomology, reported that as many as two hundred pounds of blood
may be withdrawn from a large host animal by ticks in a single season.
Woodward and Turner/ using the common cattle tick, Boophilus annu-
latus (Say), found that tick-infested cows under experimental condi-
tions gave only 65.8 per cent as much milk as tick-free cows. Further-
more, the tick-free cows gained 6.1 per cent in body weight during the
time of the experiment, while the tick-infested animals gained 3.6 per
cent. Death due to exsanguination by ticks is believed to be possible.
Jellison and Kohls 3 found that an adult female Dermacentor andersoni
Stiles withdrew from 1.7 to 2 gm of blood in the act of engorgement,
and they concluded that "tick-host anemia is not only an experimental
disease but occurs with some frequency in nature, and may be the
immediate cause of death in animals." During the months of October
to March, 1935-36, one riding academy in Alameda County, California,
lost 83 horses, whose deaths were reported to be due to loss of blood
( exsanguination) through huge infestations of Dermacentor albipictus
(Packard). Autopsies, blood examinations, and inoculation of blood into
other horses produced no symptoms not attributable to simple secondary
anemia.
There are many disorders and diseases of man and animals traceable
to ticks, among these are (1) dermatosis, inflammation, itching, swelling,
and ulcerations at the site of the bite, also skin ulceration and lesions
464
TICKS AND TICK-BORNE [>ISEASES 465
resulting from improper or partial removal of tick mouth parts; ( 2)

envenomization, inoculation of toxic salivary fluids at the site of the bite
often resulting in severe systemic disturbances; (3) exsanguination, a
serious matter when an animal is badly infesfed with ticks, resulting in
a secondary anemia and possibly death; (4) tick paralysis, an acute as~
cending, flaccid type of paralysis, often fatal, c~aused directly by the bite
of certain species of ticks, affecting children ~nd young animals such as
lambs, calves, colts, pigs, and dogs; (5) otpacariasis, invasion of the
auditory canal by ticks; (6) predisposition to myiasis and secondary
bacterial infections through tick-bite injuries. Infections transmitted by
ticks may be classified in part as follows: babesiosis, piroplasmosis,
rickettsiosis, and spirochetosis; they may also assist in the spread of
tularemia and certain viral and helminthic infections.
High vector potential of ticks. Some of the factors which account for
the potency of ticks in the spread of diseases of man and animals are
the following: they are first of all persistent bloodsuckers-they attach
firmly while feeding and cannot be dislodgea easily; the later stages
(nymphs and adults) are highly chitinized, pence very resistant to en-
vironmental stresses; they are relatively free from natural enemies; most
species have a wide host range, thus ensuring a relatively certain source
of blood; very few species of ticks are dependent upon a single host
species; usually many species are available. Cooley reported 28 species
of hosts for the adult of Dermacentor andersoni Stiles, and 32 for the
immature stages. Other factors adding to the vector potential of ticks are:
longevity-the life span of some species may }Je over five years; the long
duration of the life cycle in many species, due to longevity of the several
developmental stages, is an important factor in that, for example, the
tick vector of relapsing fever may be a dangerOUS carrier of the infection
for several years having received the infectioo through the egg from the
previous generation; the reproductive potential of ticks is quite great-
some species may deposit as many as 18,000 eggs; ticks apparently have
the power to regenerate lost parts such as amputated legs, and the ability
to repair mutilated mouth parts.
Man not a natural host for ticks. Ticks llre in general parasites of
animals in nature; they parasitize man and }lis domesticated mammals
fortuitously. However, man has set the stage for enormous infestations
of his herds of cattle, horses, and other anirtJ als . Certain tick-borne in-
fections in wild animals have persisted for many centuries, and a host-
parasite relationship has developed resultin~ in a benign (nonpatho-
genic) condition of infection rendering the wild animal a true reservoir.
Thus rabbits (several species) harbor the ric).{ettsiae of Rocky Mountain
spotted fever, which infection is transmitted from rabbit to rabbit by a
relatively host-specific tick. There is thus mttintained in nature a con-
tinuous reservoir of infection; man is not an essential factor. Other
species of ticks, more particularly Dermacentor andersoni Stiles, which
bite other animals as well as man, pick up the infection from an infected
rabbit, and if man intervenes (a fortuitous host) he will suffer a rickett-
sial infection often of great severity, probably because he is an inappro-
priate host. Spirochetal infections are described later in this chapter.
No doubt, other infections not yet described will be revealed.
Diseases limited to animals, i.e. not transmissible to man may also
possess similar tick-host relationships. Thus the causal agent of bovine
anaplasmosis is harbored by wild deer without causing apparent symp-
toms, but when it is transmitted to domestic cattle by ticks a severe
disease results.
Historical. Although ticks were referred to {is "disgusting parasitic
animals" by Aristotle in the fourth century B. c., the orderly classifica-
tion of these parasites dates from the publications of Linnaeus in 1746
(Fauna Suecica). Linnaeus placed them among the Acari in the genus
Acarus specifically, according to Nuttall (Monograph of the Ixodoidea,
Cambridge University Press, May 1911). Pliny is said to have used the
word ricinus to Signify a "tick" because the fully engorged female re-
sembles the seed of the castor-oil plant (Ricinus). The scientific nomen-
clature of ticks began with Latreille in 1804 when he set up eleven
genera, among them Ixodes; therefore, the species of Linne became as
we have it today, namely, Ixodes ricinus (Linnaeus). It was not until
1844 that C. L. Koch separated the ticks from the Acari, which in-
cluded both ticks and mites. In 1896 Neumann placed the ticks in the
order Acarina and divided them into two subfamilies, namely, (1)
Argasinae and (2) Ixodinae. Finally, in 1901, Salmon and Stiles raised
the two subfamilies of Neumann to family rank and placed them in the
superfamily Ixodoidea established by Banks in 1894. Thus, systematic
knowledge of the ticks grew slowly, and knowledge of their biology was
exceedingly scant. Nevertheless, when Nuttall and associates published
their bibliography on ticks in 1911 there were already 2,004 titles.
Theobald Smith (see p. 2) was the first (1893) experimentally to
incriminate ticks as vectors of disease. He had previously (1889) dis-
covered that Texas cattle fever was caused, like malaria, by a blood-
corpuscle-invading protozoon, Babesia bigemina. Earlier discoveries
pertaining to malaria were, no doubt, helpful. Smith and his associates,
of course, also knew well the habits of the bloodsucking ticks, Boophilus
annulatus (Say) which abundantly parasitized the diseased as well as
healthy cattle. They knew that this species is host-specific, hence needs
but one host for its complete development. The astonishing fact was
revealed by Theobald Smith that the etiological agent of Texas cattle
fever was carried from the infected mother tick to her offspring (genera-
tion to generation) through the eggs. This discovery gave the answer
to the puzzling question as to how this tick could infect other cattle
when it was well known that once it had dropped off its host (the female
deposits her eggs on the ground and then dies) it never again resumes
its parasitic position on another host animal. Transovarian (generation
to generation) transmission explained this readily-the newly hatched
ticks were already infected, hence infectious, when feeding began.
In 1903 Marchoux and Salimbeni proved fowl spirochetosis, caused
by Borrelia anserina (= Spirochaeta gallinarum Blanchard), to be tick-
borne. They proved the common fowl tick, Argas persicus (Oken), to be
a vector. This was apparently the first demonstration that a tick can
transmit spirochetosis. In the meantime, the puzzling so-called "tick-
bite" fever of man in the African Congo was nearing a solution. This
fever was long associated with the bite of certain ticks by the natives.
David Livingstone, the explorer, in 1857, was the first to report upon
the evil effects following the bite of a tick. Murray in 1877 named this
tick, Ornithodoros moubata. It was not until 1904 that Ross and Milne,
Dutton and Todd in 1905 (reporting from Uganda and the Congo),
and simultaneously Robert Koch reporting from (German) East Africa
proved that this tick, Ornithodoros moubata (Murray), was a vector of
the causal organism, a spirochete; the "tick-bite fever" of the natives,
therefore, proved to be a tick-borne relapsing fever. Generation-to-gen-
eration transmission through the egg was also demonstrated. These were
epoch-making discoveries and quickened the interest of workers in the
fields of parasitology and epidemiology everywhere. In the brier period
of a dozen years both families of ticks, the Ixodidae, hard ticks, and the
Argasidae, soft ticks, had been incriminated as disease vectors involving
both man and his domesticated animals.
Characteristics of ticks. With the few wingless insectan forms well in
mind, such as the sac-like Pupipara, particularly the sheep ked, ticks are
easily distinguished from insects, in that the body is not definitely
divided; i.e., there is a strong fusion of the thorax and abdomen produc-
ing a sac-like leathery appearance. A discrete head is lacking, but the
mouth parts, together with the basis capituli in many species (Ixodidae)
form a structure known as the capitulum. Like other Arachnida the
mature ticks bear four pairs of legs; the larvae are hexapod.
All ticks bear a pair of spiracles, situated lateroventrally on the
abdomen, one on each side near the third and fourth coxae. A pair of
simple eyes may be present. Many species of ticks are eyeless.
Ticks vary considerably in size according -to species but rarely ex-
ceed 15 mm in length in the fully engorged females. The females are
capable of great distention, and when fully engorged "are seed-like in
form.
Tick mouth parts and feeding habits. The capitulum (head) bears
the mouth parts and accessory external structures. (Fig. 30, Chapter VI.)
The basal portion is known as the basis capituli, from which projects a
pair of protrusible chelicerae. The distal portions (digits) of the cheli-
cerae are divergent and are provided with recurved teeth. Projecting
fonvard and situated ventraJJy and medianJy on the basis capituli is the
hypostome bearing many recurved teeth. Laterally are located the palpi
(one pair), consisting of four articles, of which two or more maybe
used; commonly three are visible.
When sucking blood both the hypostome and the chelicerae are in-
serted into the skin of the host. The impression that the mouth parts are
formed like a corkscrew and may be removed by "unscrewing" is, of
course, erroneous. By "unscrewing" one is more likely to leave the mouth
parts in the flesh. Because of the recurved teeth the tick is enabled to
hold so fast to the host that it is difficult to remove it without tearing the
capitulum from the body of the tick. The tick itself, however, withdraws
its mouth parts quickly and apparently with little effort by slipping the
hood-like portions of the capitulum over the relaxed mouth parts and,
with a quick jerk, drops off and escapes.
One does not usually feel the tick when it is biting. As Cooley 4 has
so well said, "a person is always completely surprised when he finds a
tick attached." The best way to remove a tick, and this should be done
without delay, is to take hold of it with the fingers and pull it off slowly
with a firm straight pull without jerking.
The length of time that a tick remains attached in the act of uninter-
rupted feeding depends on the species and the stage of development.
The seed ticks commonly feed for a number of days; the nymphs and
adults differ greatly in this respect, thus the common poultry tick, Argas
persicus (Oken) feeds nightly and intermittently, while the nymphs and
adults of the southern cattle tick, Boophilus annulatus (Say) feed from
six to eight days before becoming fully engorged. Other species of ticks,
notably the California relapsing fever tick, Ornithodoros hermsi Wheeler,
is able to engorge fully in from 15 to 20 minutes. Both male and female
ticks suck blood; however, in the case of the Ixodidae, only the females
become greatly distended when engorged, while both sexes become
distended in the Argasidae.
Life history. Under natural conditions a few species of ticks show
a marked host specificity, e.g., Boophilus annulatus (Say), the southern
cattle tick, and D. parumapertus Neum. on jackrabbits. However, most
species have a fairly wide range of hosts, e.g., Dermacentor occidentalis
Neum. The life histories of ticks vary considerably for the several
species hence it is quite i~possible to generalize, except that it may be
said that all species of ticks pass through four stages (egg, seed tick,
nymph, and adult) in from six weeks to two years, e.g., the Rocky Moun-
tain spotted fever tick, Dermacentor andersoni Stiles, requires normally
about two years to complete its life history. The fully engorged female
ticks usually deposit their eggs on the ground, the number varying from
100 in some species to 18,000 in others. The newly hatched larvae,
known as seed ticks, are hexapod (six-legged) and remain in this condi-
tion until the first molt is completed. The nymph emerges from the first
molt with its fourth pair of legs present, and remains in this stage until
the second molt, from which the adult tick emerges. As many as seven
molts may occur in certain Argasine ticks. Copulation takes place after
the last molt, when the females engorge and then deposit eggs. In the
majority of species the ticks drop off the host animal to molt, but in
several species, notably the Texas cattle fever tick Boophilus annulatus
( Say), the molting takes place on the host. There may be two or pos-
sibly three generations of ticks in one year under very favorable climatic
conditions in species which molt on the host.
The seed ticks emerging from the eggs on the ground commonly
climb up grasses and other low vegetation in order to come within easy
reach of grazing or passing animals. The nymphs and adults employ
the same method. Wild lilac (Ceanothus) is a favorable shrub for the
purpose, in fact in some localities in California it is known as a "tick
bush."
The larvae haVing reached the body of a host, there follows a
sequence of feeding and molting until maturity is reached. When this
sequence is completed on one animal, as for example, in the case of the
cattle fever tick, Boophilus annulatus (Say), the species is said to be a
one-host tick. When the sequence is completed on two host animals, as
with the African red tick, Rhipicephalus evertsi Neum., the species is
said to be a two-host tick. The larvae of this two-host species hatch on
the ground like other ticks, then proceed to attach themselves to the
ears (often inside) and flanks of cattle, where they become fully en-
gorged and molt while on the host. The nymphs then engorge and
drop off to molt, after which the adult tick emerges and must now find
a second host upon which it engorges itself and then drops to the
ground, where the females lay eggs.
When the tick species requires three different hosts to complete its
cycle it is called a three-host tick, as for example, Dermacentor ander-
soni Stiles, the Rocky Mountain spotted fever tick. In this species the
larvae select smaller animals such as ground squirrels upon which en-
gorgement is achieved after which the larvae drop to the ground to molt
and reach the nymphal stage. In this stage the second host is usually a
larger animal such as a rabbit or coyote. After engorgement on this host
the nymph drops to the ground and with its last molt it reaches the adult
stage and once more finds a host animal (the third host) upon which
it feeds and thereafter drops to the ground, where the females lay their
eggs.
In such species of ticks as Ornithodoros hermsi Wheeler, the vector
of relapsing fever in California, several individual host animals are re-
quired; such species are known as many-host ticks. There are usually
five molts in this species, each of which is completed off the host, hence
at least five host animals are needed to complete the cycle.
Longevity. The longevity and hardiness of ticks is something truly
remarkable, a matter not to be overlooked in applying control measures.
Furthermore, chemicals which kill most insects in a few minutes act
very slowly on these arachnids.
Unfed larval ticks of the species, Argas persicus (Oken), remain alive
quite readily for more than a month; nymphs survive a longer time and
the adults even longer than the nymphs. Nuttall and Warburton 5 cite
cases in which nymphs of this species survived two months, and adults
(unfed) "a little over two years." Graybill 6 reports considerable varia-
tion in the longevity of the Texas fever tick, depending on the season of
the year; unfed larvae survived from 7 to 85 days (average 38.6) for
July, and 30 to 234 days (average 167.4) for October. Nuttall and War-
burton (loc. cit.) mention cases in which the unfed larvae of Ixodes
ricinus (Linn.) survived 19 months, unfed nymphs 18 'months, and unfed
adults 15 to 27 months. Unfed adult Dermacentor andersoni Stiles sur-
vived 413 days. The author had a female Ornithodoros megnini Duges
remain alive without food in a pill box for two years and seven months.
A specimen of O. coriaceus Koch remained alive similarly over six years
with an average of two blood meals annually.
Classification. The superfamily Ixodoidea, which includes well over
450 species of ticks, is divided into two families, namely: (1) Ixodidae,
also known as the hard-bodied ticks, comprises scutate ticks with a
terminal capitulum; sexual dimorphism is marked; the males have a
scutum which almost covers the dorsum and are incapable of great
distention; in the females the scutum is a small shield immediately
behind the capitulum; the females are capable of enormous disten-
tion. (2) Argasidae, also known as the soft-bodied ticks, includes ticks
without a scutum (nonscutate); sexual dimorphism is not marked; the
capitulum is ventral and the palpi are leg-like; the eyes when present
are lateral and situated on the supracoxal folds; the spiracles are very
small.
The two families may be readily separated by means of the following
table (Table XII), and Figure 151.
IXODIDAE. TICKS ARGASIDAE.
Dermacentor iY1dersonl Stiles Omithodoros hermsi w..H..M.
MALE FEMALE MALE FEMALE
..J
«
en
'o"
o
.~
,~ I
;~~. ---+--_.- ·····_·_-t

,...~ !
z
w
>
"ry
j
6\
"'0...'
......:'
!":....
; \'
.. '\
en
UJ
<)
0::'
oco
...J
,...«
Z
W
<!>
en
w
,...
«
...J
0..
-'
«
:::.
t!)
i=
en
FIG. 151. Structural details of the two families of ticks, Ixodidae and Argasidae.
471
472 MEDICAL ENTOMOLOGY'
TABLE XII
DIFFERENCES BY WHICH THE TWO FAMILIES OF THE IXODOIDEA MAY BE
SEPARATED (ADAPTED AFTER NUTTALL)
Argasidae Ixodidae
Sexual dimorphism Slight Marked

Capitulum Ventral Anterior
Base No porose areas J Porose areas in-
Palpi Leg-like, with subequal articles Relatively rigid, of very varied
form
Body
Scutum Absent Present
Festoons Absent Generally present
Eyes (when present) Later&l on supracoxal folds Dorsal on the sides of the
scutum
Legs
Coxae Unarmed Generally armed with spurs
Tarsi Without ventral spurs Generally armed with 1 or 2
ventral spurs
Pulvilli Absent or rudimentary Always present
FAMILY IXODIDAE (HARD-BODIED TICKS)

The family Ixodidae. Nuttall and Warburton (1911, loco cit.) include
nine genera.:> in the family Ixodidae; namely, Ixodes, Haemaphysalis,
Dermacentor, Rhipicentor, Rhipicephalus, Margaropus, Boophilus,
H yaloma, and Amblyomma, divided into five groups as follows (see also
Fig. 152 showing typical capituli) ;
Ixodidae
I
Prostriata Metastriata
I
Brevirostrata Longirostrata
I I
Group 1 Group 2 Group 1 Group 2
I I \ \
1 Ixodes 2 Haemaphysalis 3 Dermacentor 8 Hyalomma 9 Amblyomma
4 Rhipicentor
5 Rhipicephalus
6 Margaropus
7 Boophilus
Genus Ixodes. This genus, which comprises about 60 species, is
clearly separated from all other genera of the family Ixodidae by the
.. Colley, 1938, added a new genus, Otocentor, which becomes a monotypic
genus with Dermacentor nitens Neumann the genotype. The horse and related
animals are evidently the favored host.
anal groove surrounding the anus in front (Prostriata) and the absence
of festoons. The remaining genera fall naturally into two divisions (see
diagram), the one characterized by a comparatively short capitulum,
and the other by a comparatively long one. Eyes are absent; spiracles are
round or oval; palpi and basis capituli are of variable form; coxae either
unarmed, trenchant, spurred, or bifid; tarsi without spurs; sexual dimor-
FIG. 152. Characteristic capituli of the several genera of ticks belonging

to the family Ixodidae: (A) Boophilus, (B) Ixodes, (C) Dermacentor, (D)
Amblyomma, (E) Haemaphysa1is, (F) Rhipicephalus, and (G) Hyalomma.
(Adapted after Cooley.)
phism is pronounced, especially with regard to the capitulum; in the
male the venter is covered by nonsalient plates; one pregenital, one
median, one anal, two adanal, and two epimeral plates; e.g., Ixodes
ricinus (Linn.), commonly known as the European castor bean tick. It
is very widely distributed and feeds on a wide variety of hosts. The
species widely referred to as Ixodes californicus Banks (Fig. 153) is now
FIG. 153. A common deer and

cattle tick of the Pacinc coast, Ixodes
pacific liS; (left) female, (right) male.
xS.5.
described as Ixodes pacificus Cooley and Kohls s (1943). This is a com-
mon deer tick in California but flourishes on cattle as well. It bites
human beings freely and often causes severe disturbances. Among the
ticks referred to as bird ticks is Ixodes howelli Cooley and Kohls (1938)
taken from the Sierra Nevada rosy finch, Leucosticte tephrocotis dawsoni
Grinnell, at an elevation of 12,000 feet.
Genus Dermacentor. Usually they are ornate ticks with eyes; festoons
11 in number; palpi short, broad or moderate; basis capituli rectangular
dorsally. In some species coxae I to IV of the male increase progressively
in size; in all species coxa IV is much the largest; the male shows no
ventral plates or shields. Coxa I is bifid in both sexes. The spiracles are
suboval or comma-shaped. The genus includes about 20 species. Students
concerned with species of the genus Dermacentor must consult the
publications of R. A. Cooley (e.g., 1938, loco cit.).
Dermacentor variabilis (Say) is the principal, if not the only, vector
of Rocky Mountain spotted fever in the central and eastern portions of
the United States. It is also an important vector of tularemia and bovine
anaplasmosis. It may cause canine paralysis and is a common pest of
dogs, which are the preferred host of adult ticks of this species. It also
freely attacks horses and many other animals, including man. The imma-
ture stages feed almost exclusively on small rodents, very largely on
meadow mice, [Microtus pennsylvanicus pennsylvanicus (Ord) in the
Atlantic states]. It is a widely distributed North American species. It
is commonly referred to as the "American dog tick" (Smith, Cole, and
Gouck 9 ).
The fully engorged females drop from the host and in four to ten
days lay eggs (4,000 to 6,500) on the ground. The incubation period
during the summer months is about 35 days; however, temperature in-
fluences this stage greatly. The larvae remain in clumps on the ground
or on low-growing vegetation while awaiting a host, usually a field
mouse. The period of larval engorgement varies from 3 to 12 days, after
which the larvae drop to the ground and molt in about a week, though
often greatly prolonged. The nymphs, having reached a host, again
usually a field mouse, engorge in from 3 to 11 days, and once more drop
to the ground and molt in from three weeks to several months. The un-
fed adults may live for more than two years; however, having reached
the adult stage the ticks usually attack dogs and other large host animals
including man. Engorgement of the females requires from 5 to 13 days.
Mating takes place on the host. Like the unengorged adults the imma-
ture stages have a remarkable longevity in the absence of suitable hosts,
which may prolong the life history two or more years, although under
favorable conditions the life cycle from egg to adult may not require
more than three months.
Dermacentor andersoni Stiles, the Rocky Mountain wood tick (Fig.
154) is a widely distributed and very common species throughout the
Rocky Mountain region of the United States including Idaho, Wyoming,
Montana, parts of Utah, Colorado, Nevada, Oregon, Washington, Cali-
fornia, and British Columbia. "It is most abundant in regions or localities
where the predominating vegetation is low, brushy, and more or less
open, i.e., in areas where there is good protection for the small mam-
malian hosts of the larvae and nymphs and sufficient forage to attract
the large hosts (either wild or domestic) of the adult ticks. It is rela-
tively scarce in heavily timbered areas or country of a strictly grassland,
prairie type" (Parker et al. 10 )
This species may be recognized by comparison with Fig. 154. The
adult ticks feed mostly on large animals, such as horses, cattle, and
sheep, and also on deer, bear, and coyote; the larvae prefer to feed on
rodents such as ground squirrels, pine squirrels, woodchucks, and chip-
munks; the nymphs feed on the same host species as the larvae; all three
stages may feed on animals of intermediate size such as jack rabbits,
marmots, and badgers. In the wilder parts of its range where there are
no domesticated animals, the adult ticks feed on deer, elk, bear, moun-
tain goat, and coyote.
The life cycle of the Rocky Mountain wood tick is flllly described
by Cooley,ll and from his description the following summary is largely
FIG. 154. The Rocky Mountain spotted fever tick, Dermacentor al1de'l'soni;
(left) male, (right) unengorged female. X 3.5.
taken. Copulation takes place on the host, and when fully fed, the
greatly distended female drops to the ground. The preoviposition period
is about a week, after which egg-laying begins, continuing over a period
of about three weeks. If undisturbed, the eggs pile up ahead of the
female in one large mass, averaging some 6,400 eggs. The incubation
period of the eggs requires about 35 days, when the young hexapod seed
ticks emerge and find suitable rodent hosts, feed for three to five days,
drop off, and molt in from 6 to 21 days, emerging as nymphs with four
pairs of legs. These nymphs, the progeny of overwintered adults, go
into hibernation to come up for feeding the next spring when they seek
larger-sized hosts as already explained, to which they attach for feeding
over a period of from four to nine days. When fully engorged, the
nymphs drop to the ground and in 12 to 16 days or more, molt for the
second time, emerging as adults. Some of these adults may find hosts
in the same summer in which they have emerged as adults, but by the
time they have emerged, Cooley explains, the season has generally be-
come hot and dry, making it necessary for them to seek protection under
waste and vegetation. The "normal cycle" is therefore two years. The
larvae are found feeding through the summer months, and while the
adults disappear by about July 1, the nymphs continue in diminishing
numbers until late summer. Since man is usually bitten only by the adult
ticks, danger from this source exists only from early spring to about
July 1.
Like other species of ticks, Dermacentor andersoni Stiles is remark-
ably resistant to starvation. Hunter and Bishopp'2 report that all unfed
seed ticks hatching from a mass of eggs usually die within one month
after the first eggs hatch if food is not available. However, in one in-
stance a period of 117 days elapsed between the beginning of hatching
of the eggs and the death of the last seed tick. (A later record noted by
these investigators exceeded 317 days.) Unfed nymphs have been found
to survive one year and 11 days, and adults collected on vegetation dur-
ing the spring months may survive for 413 days without food. Students
concerned with the anatomy of this important species of tick will need
to consult the work of Douglas 13 (1943).
Dermacentor albipictus (Packard) is commonly known as the winter
tick, elk tick, or horse tick. It is a widely distributed North American
species. It is a one-host tick and does not occur on the host during the
summer months, it is distinctly a winter tick. The eggs are laid during
the spring months and hatch in from three to six weeks. The seed ticks
then bunch tightly together, remaining in a torpid condition until the
first cold weather in autumn, when they become very active and seek
host animals. Molting takes place on the original host animal. The
females reach maturity with the final molt and engorgement usually in
about six weeks after the seed ticks have become attached. Although
the females drop off the host after final engorgement as do other ticks,
egg laying is delayed until spring, often after an interval of several
months.
Hearle (1938, loco cit., p. 399) states that heavy infestations of horses,
cattle, moose, elk, and deer may result in death from "tick poverty" due
to the drain on the vitality of the infested hosts. In this respect, Hearle
states, it is the most important species occurring in Canada. This "tick,
it is reported, caused a loss of at least 20 per cent of the moose popula-
tion in Nova Scotia. A disease of moose (Alces americana americana
Jardine) is described by Thomas and Cahn 14 as occurring in northeastern
Minnesota and the adjacent region. It is reported that guinea pigs and
rabbits infested with the tick, Dermacentor albipictus (Packard), from
diseased moose have reproduced in detail the symptoms of weakness,
anemia, paralysis, excessive blind activity, and death as exhibited by in-
fected moose. The causal bacterium is described as Klebsiella paralytica
"because of the paralysis it causes" (Cahn, Wallace, and Thomas 15 ).
Dermacentor occidentalis Marx has a narrow Pacific coast distribu-
tion, being particularly abundant in southwestern Oregon, rare north-
ward and southerly. Adults of this species have been taken from cow,
horse, mule, ass, deer, rabbit, sheep, dog, man, and the immature stages
from many species of smaller animals such as ground squirrel, rabbit,
skunk, and field mouse. Dermacentor andersoni and D. occidentalis,
though distinct species, are very closely related and have been hybrid-
ized.
Key to the adults of the genus Demacentor
in the United States (after Cooley)
1. Spurs on coxa I widely divergent .. ............... 2
Spurs on coxal I with proximal edges parallel or only a little diver-
gent .... ................. 3
2. Scuta with deep large punctations parumapertus Neum.
Scuta with the larger punctations shallow and moderate in size (known
from peccary in southern Texas) . . halli McIntosh
3. Spiracular plate oval, without dorsal prolongation and with goblets
few and large .. ............ .. .... albipictus (Packard)
Spiracular plates oval, with dorsal prolongations, and with goblets
many or of moderate numbers 4
4. Cornua" long.. ..occidentalis Marx
Cornua short or of moderate length. 5
5. Spiracular plate with goblets very numerous and small
variabilis (Say)
Spiracular plate with goblets moderate in size and number 6
6. The larger punctations of the scuta very large and deep
andersoni Stiles
The larger punctations on the scuta moderate in size and depth (known
from Rocky Mountain sheep in southern Arizona) hunteri Bishopp
Genus Haemaphysalis. The members of this genus, numbering about

45 species, are usually small and but slightly chitinized, and the sexes
are very similar. They are inornate and eyeless, but have festoons; the
second segment of the usually short conical palpi projects laterally be-
yond the basis capituli, forming an acute angle. The spiracles of the
males are ovoid or comma-shaped; in the female rounder or ovoid. The
genus is commonly found on mammals but may severely parasitize
ground-nesting birds.
Haemaphysalis leporis-palustris (Packard) is a widely distributed
North and Central American species. Although it is commonly known
as the rabbit tick, it has been taken on a number of species of birds and
occasionally on domestic animals such as the horse, cat, and dog. It
"Cornua: caudad projection extending from the lateral posterior dorsal angles of
the basis capituli.
rarely bites man but is important in the spread of Rocky Mountain
spotted fever and tularemia among wild animal reservoir hosts.
Haemaphysalis leachi (Audouin), the dog tick of Africa, is common
in Asia, Australia, and Africa. It is usually found on wild and domestic
carnivores, frequently on small rodents and rarely on cattle. This tick
is a vector of malignant jaundice in dogs.
Genus Rhipicentor. This small and relatively unimportant genus of
ticks is inornate, with eyes and festoons present; the palpi are short,
with basis capituli hexagonal dorsally and having very prominent lateral
angles. Coxa I is bifid in both sexes; coxa IV is much the largest; there
are no ventral plates or shields; spiracles subtriangular in the female
or comma-shaped in the male. The genus is represented by Rhipicentor
bicornis Nuttall and Warburton.
Genus Rhipicephalus. This genus comprises about 30 species. They
are usually inornate and possess eyes and festoons; the palpi are short
and the basis capituli is usually hexagonal dorsally. Coxa I is bifid. The
male possesses a pair of adanal shields and usually a pair of accessory
glands; some males when replete show a caudal protrusion. The spiracles
are bluntly or elongatedly comma-shaped.
Rhipicephalus sanguineus (Latr.) is the cosmopolitan brown dog
tick; its principal host is undoubtedly the dog, although it is known to
attack numerous other animals. It is a widely distributed tick occurring
primarily in tropical and subtropical climates and is widely known as
a vector of malignant jaundice of dogs. The adult ticks are most often
found in the ears and between the toes of dogs, and the larvae and
nymphs in the long hair at the back of the neck. McIntosh 16 states that
the eggs are deposited in cracks and crevices of the kennel or other
quarters frequented by the dog. The ticks have a strong tendency to
crawl upward, hence are often found hidden in cracks in the roofs of
kennels or in the ceilings of porches. It may become a very great nui-
sance. The eggs hatch in from 20 to 30 days and over, depending upon
temperature. The life cycle corresponds to that of other species of Ixodine
three-host ticks.
Genus Margaropus. This genus has an obsolete anal groove, and no
ornamentations or festoons; the palpi are short and the capitulum is
highly chitinized and similar in shape to that of Boophilus. The coxae
are conical and unarmed except for a small spine posteriorly on coxa 1.
The male has a median plate prolonged into two long spines projecting
beyond the anus on both sides; when replete it shows a caudal pro-
trusion. It may be separated from Boophilus, which it closely resembles,
by the presence of greatly thickened posterior legs and by the prolonged
median plate.
Margaropus winthemi Korsch, the Argentine tick, is a native of South
America introduced into South Africa; it is frequently found on horses
and occasionally on cattle. When engorged the females may easily be
mistaken for B. decoloratus (Koch) but may be distinguished by the
dark bands at the joints of the legs. This is a one-host tick, usually present
in larger numbers during the winter.
Genus Hyalomma. Ornamentations are absent or present and may
be at times confined to the legs; eyes are present, and there mayor may
not be festoons. The palpi are long and the basis capituli is subtriangular
dorsally. The female approaches Amblyomma. The male has a pair of
adanal shields, with or without accessory adanal shields and two posteriOl
abdominal protrusions capped by chitinized points. Coxa I is bifid.
Spiracles are comma-shaped. The genus includes less than half a dozen
species.
Hyalomma aegyptium (Linn.). The "bont-Ieg" tick, with its two
varieties, is fairly widespread over much of southern Europe, Asia, and
FIG. 155. The lone star tick, Ambly-

omma americanum. X 3.5.
Africa. The adults are parasitic on cattle, horses, sheep, goats, wild
mammals, and occasionally birds, while the larvae and nymphs are
found on rodents, hares, and birds, frequently infesting domestic poultry.
It is usually a two-hosted tick, the larvae and nymphs feeding on the
same host, though the larvae may drop off and seek a new host.
This tick is very hardy and exists under extremes of heat and cold.
The adults attach by preference around the anus and genitalia and may
produce severe lesions. Lameness is frequently encountered in sheep
as a result of tick bite.
Genus Amblyomma. These are generally ornate ticks, with eyes and
festoons; the palpi are long, article 2 especially so; the basis capituli is of
variable form. The male is without adanal shields, but small ventral
plaques are occasionally present close to the festoons. Spiracles are sub-
triangular or comma-shaped. There are about 90 species in this genus.
Amblyomma americanum (Linn.) is the '1one star" tick (Fjg. 155) of
the southern United States, particularly Oklahoma, Texas, and Louisiana.
Its range appears to extend considerably northward, and southerly into
Mexico, Central America, and South America (Brazil). This species has
a wide variety of hosts, including wild and domestic animals, birds, and
man.l7 It takes high rank as a pest and vector of disease, e.g., Rocky
Mountain spotted fever, Bullis fever, and tularemia. It is a three-host tick.
Amblyomma caiennense (Fabr.) is reported as occurring in the
southern United States, Central America, the West Indies, and South
America. It commonly attacks man, horses, cattle, sheep, dogs, pigs, and
many other animals. This species appears to be abundant in a few coun-
ties in Texas (Cooley and Kohls, loco cit.).
Genus Boophilus. Members of this genus have neither festoons nor
ornamentations, but eyes are present. The palpi and hypostome are
short, compressed, and dorsally and laterally ridged. The basis capituli
is hexagonal dorsally and slightly chitinized. Coxa I is bi£d; the anal
groove is obsolete in the female and faintly indicated in the male. Unfed
FIG . 1.56. The Texas fever tick,

Boophilus annulatus; (left) female,
(right) male. x 3 ..5.
adults are small; the scutum of the female is quite small and the spiracles
are circular or oval in both sexes. There are less than half a dozen species.
The Texas cattle fever tick, Boophilus annulatus (Say) (Margaropus
annulatus Say) (Fig. 156) is normally restricted to North America south
of the 37° latitude into parts of Mexico. It is typically a cattle tick, al-
though it occurs at times in smaller numbers on deer, sheep, horses ,
mules, and other animals. The bison is evidently also a suitable host.
Boophilus annulatus microplus (Can.) (= australis Fuller) is referred
to as the tropical cattle tick (Tate 18 ).
Fully engorged females range in length from 10 to 12 mm, while
the male ranges from 3 to 4 mm. The body of the female is about equally
rounded both posteriorly and anteriorly, with a slight median incurving.
The anterior pair of legs is set well out on the shoulders away from the
capitulum (in Dermacentor close to the capitulum) . The palpi are
short, and the capitulum is inconspicuous. The relatively small (about
1 mm long, scutum is solid chestnut brown in color.
Economic importance. Tremendous strides have been made in the
last few years in the control of the Texas fever tick. It is of interest to
note that as late as 1906 it was estimated that the annual losses to south-
ern United States occasioned by the "cattle tick" directly and indirectly
amounted to $130,500,000. 19 This sum included (1) death, from Texas
fever, of pure-bred cattle imported from the North for breeding pur-
poses; (2) death, from Texas fever, when cattle reared in isolated tick-
free areas were unintentionally or accidentally placed with ticky cattle,
or on tick-infested areas; (3) death of native cattle from excessive para-
sitism and fever, occasioned by the ticks; (4) universal loss of weight by
all tick-infested cattle, and their failure to gain flesh at a rate great
enough to make beef production profitable; (5) the lower price which
"Southern" cattle brought upon the market, regardless of how perfect
their condition; (6) sterility induced in high-grade cattle by tick in-
festation; (7) the expense of maintaining the Federal quarantine for the
protection of the North against invasion by the tick, and the added
expense of maintaining quarantine pens for southern cattle shipped North
for slaughter; (8) the discouraging effect on the breeding of pure-bred
cattle in the South by reason of southern breeders not being allowed
to exhibit in northern show rings; (9) by no means least, the potential
loss in fertility of southern farm lands due to a one-crop system which,
with the tick eradicated, would quickly give way to a diversified agri-
culture thus tending to conserve and increase the fertility of soils; (10)
shrinkage in milk production of tick-infested cattle.
From 1906, when cooperative tick-eradication work was undertaken
between the United States Bureau of Animal Industry and state authori-
ties, to December, 1918, a total of 458,529 square miles of territory was
released from quarantine against Texas cattle fever, which speaks well
for the methods employed.
Life history of Texas fever tick. Boophilus annulatus (Say), is a one-
host tick. The life history may be divided into two phases: ( 1) the
parasitic phase during which the tick is attached to the host and which
terminates when the mature tick drops to the ground after fertilization;
(2) the nonparasitic phase when the tick is on the ground. After the ma-
ture female tick drops to the ground, there is a pre oviposition period of
three or four days to perhaps a month. Oviposition usually begins about
72 hours after the tick drops and continues usually for eight or nine days
but may be greatly prolonged if the temperature is adverse.
The maximum number of eggs deposited by a female tick according
to Graybill (lac. cit.) was 5,105, the minimum 357, with an average
ranging from 1,811 to 4,089. The incubation period, also dependent on
temperature, ranged from 19 days in summer to 180 days in the early
autumn, with the average of 43.6 days for April, 26.3 days for May, 24.5
days for June, 20.5 days for July, 21.2 days for August, and 35.9 days
for September. The hatching period depends on the time when the eggs
are laid, the eggs first deposited ordinarily hatching first. The seed ticks
(Fig. 157) on hatching are very active; they climb onto blades of grass
or other objects which they ascend to the top, remaining clustered there
until a suitable host animal brushes against them to which they can
attach themselves. The time during which the seed ticks remain alive,
i.e., longevity of the newly hatched ticks, again varies considerably, de-
pending on temperature; the longevity for April ticks was found to be
65.1 days, May ticks 62.3 days, June ticks 65.1 days, July ticks 38.6 days,
August ticks 84.9 days, October ticks 167.4 days. The total average
ranged from 86.9 days for June to 279.6 for October.
The three stages considered in the parasitic period of the ticks are
larval (seed tick), nymphal, and adult. As Graybill has well said, "The
FIG. 157. (Left) eggs and (right) larva of the Texas fever tick, Boophilus
anntllatus. X 50.
duration of each of these stages and the duration of a single infestation
upon cattle during different portions of the year are of great practical
importance. Upon the duration of an infestation depends the time ani-
mals must be kept on the tick-free fields in order to become free from
ticks." After the seed tick has attached itself to the host, the larval
period ranges from 5 to 10 days, the nymphal period from 5 to 20 days,
and the life of the adult female from 5 to 35 days, with a total period of
infestation, including the time for molting twice, which is accomplished
on the host, from 20 to 65 days. The entire life cycle may be completed
in about 40 days under most favorable conditions, usually nearer 60
days under natural conditions.
DISEASES CARRIED BY IXODINE TICKS
Texas cattle fever. Red water, splenic fever, bloody murrain, Mexican
fever, tick fever, etc., are names given to a widely distributed disease
of cattle, being endemic in southern Europe, Central and South America,
TABLE XIII
COMPARISON OF THE LIFE CYCLE OF A WOOD TICK AND THE
TEXAS CATTLE FEVER TICK
Texas cattle fever tick

Wood tick (Dermacentor sp.)
(Boophilus annulatus)
I. Adult tick becomes engorged on host Adult tick becomes engorged on host animal
animal and drops to ground and drops to ground
II. Adult tick begins egg laying (3,000 Adult tick begins egg laying (3,000 ± eggs)
± eggs) after 3-5 days after 3-5 days
III. Seed ticks hatch from eggs in about Seed ticks hatch from eggs in about 30 days
30 days
IV. Seed ticks bunch on grass and await Seed ticks bunch on grass and await coming
coming of host animal, from one, of host animal from one day to several
day to several weeks weeks
V. After feeding 7-12 days seed ticks

drop to ground and molt
After feeding 7-12 days seed ticks molt on
VI. Nymphs crawl up on grass and await host animal
coming of second host animal from
one day to several weeks
VII. Ticks get on second host animal and

feed 5-10 days, then drop to ground
and molt second time Nymphs feed 5-10 days, then molt second
time on host animal and the newly emerged
VIII. Mature un engorged ticks crawl up on mature ticks mate
grass and await coming of third
host animal from one day to several
weeks
IX. Adult ticks mate and feed 5-8 days to Adult ticks feed 4-14 days to engorgement,
engorgement, then drop to the then drop to the ground and lay eggs
ground and lay eggs
parts of Africa, Mexico, the Philippines, and the southern United States
where it has heen known for more than a century, having been intro-
duced into this country probably from Europe. The causal organism is
Babesia begemina (Smith and Kilbourne).
The name Texas fever became attached to the disease in the United
States because of the large herds of cattle which were driven northward
from Texas and gave a certain disease in some mysterious manner to
northern cattle that crossed the trail of the southern cattle. The first
account of the disease was given by James Mease in 1814 before the
Philadelphia Society for Promoting Agriculture. In 1879 Salmon began
an investigation of the disease; and in 1889 Theobald Smith made his
epoch-making discovery of the intracorpuscular protozoan parasite in-
habiting the blood of the diseased cattle. Immediately thereupon fol-
lowed the experiments of Smith and Kilbourne, on suggestion of Salmon,
which proved the disease to be tick-borne, a suspicion held as early as
1869 according to Smith and Kilbourne. Until that time (1889) infection
was variously attributed to saliva, urine, or feces.
The disease may assume either an acute or a chronic form, the acute
form occurring during the summer months and the chronic during the
autumn and early winter. The symptoms of the acute form are as follows:
The temperature often rises to 106° to 108° F within 24 to 48 hours.
The sick animal leaves the herd, stands with arched back, head lowered,
and ears drooping, the muzzle dry, appetite lost, and rumination stopped.
There is constipation during the first stage of the disease, which may give
way to diarrhea later. The manure is frequently stained with bile and
may be tinged with bloody mucus; the urine is often very dark red or
coffee-colored. The blood becomes thin and watery, so that when making
an incision into the tip of the ear and allowing the blood to flow over
the hand it does not stick to the hand as does the blood from a healthy
animal.
Vast numbers of red blood corpuscles are destroyed by the parasites,
which accounts in a measure for the reddish color of the urine through
the elimination of hemoglobin by the kidneys; and it is believed that the
excessive work that the liver has to perform in attempting to trans-
form the excess of destroyed corpuscles into bile, causes this organ to
become deranged in function, and eventually a complete stagnation
may result with fatal termination. Mortality ranges from 50 to 75 per
cent.
The chronic form of the disease according to Mohler 20 shows all the
symptoms of the acute type, but in a milder degree. The temperature
usually remains about 103° and never exceeds 105° F. There is loss of
appetite, stoppage of rumination, constipation, and albumin in the urine.
An anemic condition of the blood, as indicated by the pale and bloodless
mucous membranes, is also present, but hemoglobin is not usually ex-
creted by the urine; hence the red-water symptom is absent. There is also
excessive loss of flesh, and before the end of the attack the affected ani-
mal is greatly emaciated. Although death rarely occurs, the value of the
animal is much reduced.
Experimental evidence. In 1888 an "investigation into the nature,
causation and prevention" of the disease was undertaken by the United
States Department of Agriculture, Bureau of Animal Industry, under
the direction of D. E. Salmon. The work was done by Theobald Smith
and F. L. Kilbourne 21 and marks a most important epoch in our knowl-
edge of protozoon diseases and in the history of preventive medicine.
During a period of about four years of almost continuous investiga-
tion, the problem was exhaustively studied in both the field and the
laboratory. The field experiments were carried out along three different
lines:
(1) Ticks were carefully picked from Southern animals so that none could
mature and infect the ground. The object of this group of experiments was to
find out if the disease could be conveyed from Southern to Northern stock on
the same inclosure without the intervention of ticks. (2) Fields were infected
by matured ticks and susceptible cattle placed on them to determine whether
Texas fever could be produced without the presence of Southern cattle. (3)
Susceptible Northern cattle were infected by placing on them young ticks
hatched artifically, i.e., in closed dishes in the laboratory (Smith and Kilbourne,
1893, lac. cit.).
Healthy native cattle (Washington, D. C.) were exposed to sick
native cattle free from ticks for months without contracting the disease,
proving that the excretions had nothing to do with the transmission of
the disease. In the absence of ticks, sick animals are harmless. Again
several thousand, mostly full-grown ticks, collected from cattle in North
Carolina, were scattered over the ground in a field on September 13.
Four native cattle were placed in the field September 14; of these ani-
mals three contracted Texas fever. This experiment was repeated with
five experimental animals, and a newborn calf, all of which contracted
the fever. A yearling heifer was plac~d in a box stall and a number of
young ticks, hatched artifiCially in glass dishes, were placed on the ani-
mal at intervals. The heifer contracted Texas fever. A repetition of this
experiment on various occasions always gave similar results. It was
definitely concluded that "Texas fever in nature is transmitted from
cattle which come from the permanently infected territory to cattle out-
side this territory by the cattle tick, Boophilus bovis, and that the infec-
tion is carried by the progeny of the ticks which matured on infected
cattle, and is inoculated by them directly into the blood of susceptible
cattle." The authors above cited state that the contents of· the bodies
of ticks in various stages of growth were examined microscopically with
considerable care, but that the abundant particles resulting from the
breaking up of the ingested blood corpuscles obscured the search so
that nothing definite was discovered. "The very minute size of the micro-
organism renders its identification well-nigh impossible, and any at-
tempt will be fraught with great difficulties."
Other tick carriers of the protozoon are Boophilus microplus (Cane-
strini) and B. decoloratus (Koch) within their range-the former Aus-
tralia, Philippine Islands, South America, and the latter Africa.
The causal agent. Babesia bigemina [= Piroplasma bigeminum
(Smith and Kilbourne)], the causal protozoon parasite of Texas cattle
fever, was discovered by Theobald Smith in 1889 and was named Pyro-
soma bigeminum.
Rees 22 reports that the piroplasms described and illustrated by Smith
and Kilbourne (1893) included two species: (1) Babesia bigemina and
(2) B. argentina Lignieres, which is of minute size and possesses other
characteristics as well. He was able to cultivate the latter in vitro but
could not cultivate the former.
The life cycle of Babesia begemina (Smith and Kilbourne) in the
tick was fully elucidated by Dennis (1932).23 He summarizes his results:
When blood which is infected with B. bigemina is taken into the gut of
the tick, many of the intracorpuscular parasites are soon freed. Certain of these
normal-appearing parasites become transformed into gametes through growth
and slight structural modification. The gametes are motile vermicule-like bodies
which show no differentiation between the sexes. The gametes become asso-
ciated in pairs, the individuals of which eventually fuse to form the zygote.
The zygote becomes a motile ookinete which passes through the thin wall of
the gut and penetrates the contiguous reproductive organs. The ova of the tick
are invaded by the ookinetes which round up and grow to form sporonts. The
sporont secretes a cyst within which it divides to form naked sporoblasts. The
sporoblasts form multinucleate sporokinetes which migrate and are carried by
cell proliferation throughout the tissues of the developing tick; some of the
sporokinetes come to occupy the anlagen of the salivary glands. The sporoki-
nete undergoes fragmentation to form the minute infectious sporozoites" (Fig.
158).
Rocky Mountain spotted fever has been known in the Bitter Root
Valley of Montana (U.S.A.) since 1872.24 It is also known as "tick
fever," "black fever," "blue disease," and "black measles." The most
characteristic and constant symptom is the rash which appears about the
second to the fifth day on the wrists, ankles, and less commonly on the
back, later spreading to all parts of the body. Parker 25 st'ates that the rash
is sometimes preceded by a mottled appearance of the skin. The symp-
toms most often complained of at the outset are frontal and occipital
headache, intense aching in the lumbar region, and marked malaise. The
incubation period is from two to five days in the more severe infections
and from 3 to 14 days in the milder ones. The fever rises rapidly in the
more virulent infections to 104 0 and 106 0 F. In fatal infections death
usually occurs between the ninth and fifteenth day. Both mild and viru-
lent strains of spotted fever occur in the same locality. Because errors in
diagnosis are easily made, laboratory tests are advised. One of the com-
mon tests consists of the intraperitoneal injection of blood (1 cc whole
blood) into male guinea pigs. In positive tests the guinea pigs show
scrotal swelling, reddening, and sloughing of the skin.
The causal agent was discovered by Wolbach 26 in 1919; he named
7@ @
8 ~~ ~~
CYCLE IN THE TICK

9 ~ ~
1 I
to ~
~
A
" V
I
FIG. 158. A schematic diagram of the life cycle of Babesia bigemina. 1-6,
the cycle in the blood of the bovine host, showing binary fission ; 7, parasite
just taken into gut of tick; 8, freed trophozoites in the gut of the tick; 9,
vermicule-like isogametes; 10, beginning of syngamy, association of the
gametes in pairs; 11, completion of syngamy; 12, motile zygote, or ookinete;
13-14, ookinete passing through wall of the gut of the tick, through the ovi-
duct, and entering the ovum; 15, sporont formed by the rounding-up and
growth of the ookinete; 16-17, formation of sporoblasts; 18, sporokinetes in
one of the large cells which are destined to form part of a salivary acinus; 19,
sporozoites in the salivary gland (a single acinus shown) of the larva of the
tick, whence they are transferred into the blood of a new host. (After Dennis.)
487
it Dermacentroxenus rickettsi in honor of Dr. Howard T. Ricketts, who
made great contributions to our knowledge of both Rocky Mountain
spotted fever and typhus fever and lost his life in the conduct of in-
vestigations in Mexico. The several rickettsioses transmitted by lice,
fleas, mites, and ticks are caused by infectious agents known as rickett-
siae; hence the causal agent of Rocky Mountain spotted fever is gen-
erally referred to as Rickettsia rickettsi (Wolbach).
Rocky Mountain spotted fever is endemic in the Unital States, in
some parts of Canada (western), in Mexico, and in some parts of SO"!lth
America where it is known as Sao Paulo fever and by other names (see
below). The greatest number of cases occur in populations engaged in
outdoor occupations, principally agriculture. Both sexes and all ages
are subject to the disease. In the western part of the United States most
of the cases occur in men, while in the eastern part of the United States
more women and children contract the disease. Parker suggests that this
is probably due to the fact that the eastern vector is a tick, Dermacentor
variabilis (Say), which infests the dog, a household animal. The Rocky
Mountain wood tick, Dermacentor andersoni Stiles, is found far less
frequently on dogs.
Tick transmission of spotted fever. In 1902, after a preliminary investi-
gation, Wilson and Chowning 27 advanced for the first time the theory
that a tick ("wood tick") acts as the natural vector of the disease. Ac-
cording to Ricketts (in the 48th Biennial Report of the Montana State
Board of Health, p. 106) as recorded by Hunter and Bishopp28 "the first
experiments which resulted in the proof of the transmission of spotted
fever by a tick were conducted by Drs. McCalla and Brenton of Boise,
Idaho, in 1905. In these experiments a tick which was found attached to
a spotted fever patient was removed and allowed to bite a healthy per-
son. In 8 days this person developed a typical case of spotted fever. The
experiment was continued by allowing the same tick to bite a second
person. In this case again a typical case of spotted fever resulted."
The famous experiments of Dr. H. T. Ricketts 29 began in April, 1906.
The more important published work of this lamented investigator has
been brought together in a memorial volume 30 from which the following.
summary is made of his reports on spotted fever. First of all it was
shown that the disease could be transmitted to guinea pigs by direct
inoculation and that the duration of the fever and cutaneous phenom-
ena resembled very closely the conditions as observed in humans. Hence,
knowing the susceptibility of this species, it was used for further experi-
mentation.
On June 19, 1906, a small female tick was placed at the base of the
ear of a guinea pig inoculated intraperitoneally June 11 and S cc of
defibrinated blood of a spotted fever patient. The tick fed for two days
TICKS AND TICK-BORNE OISEASES 489
on this animal and was then removed and kept for two days in a pill
box and on June 23 placed at the base of the t)ar of a healthy guinea pig,
the former animal dying on the same day with characteristic symptoms.
On June 28 the second guinea pig showed d~cided rise in temperature,
which continued high until July 5 and became normal on July 7. Proper
controls were conducted and two guinea pigs which were in the same
cage with the tick-bitten guinea pig for two weeks did not become in-
fected, indicating that mere association did npt result in contracting the
disease. It will be noticed that Ricketts. called the wood tick which he
used Dermacentor occidentalis Neum. Evidently the species was actually
Dermacentor andersoni Stiles.
In addition to many other successful experiments during the follow-
ing year Ricketts found that the disease can be transmitted by the male 31
as well as by the female tick and that "one ~ttack of the disease estab-
lishes a rather high degree of immunity to supsequent inoculation." Fur-
tbermore a collection ot tiCKS taKen in tbe field transmitted the disease
to a guinea pig in the laboratory, indicating the fact that infective ticks
occur in nature in small numbers.
It was also ascertained that "the disease may be acquired and trans-
mitted . . . by the tick during any of the c1ctive stages . . . and that
the larvae of an infected female are in soJJ1e instances infective. . . .
The disease probably is transferred through the salivary secretion of the
tick, since the salivary glands of the infected adult contain the virus."
The transmission is believed to be "biological rather than purely me-
chanical."
Experiments conducted by Moore (Ricketts, 1911, lac. cit., pp. 428-
436) show that the "minimum duration of feeding necessary for a tick to
infect a guinea pig was one hour and forty-five minutes. The average
time necessary seems to be about ten hours, while twenty hours were
almost constantly infective." Maver (see Ricketts, 1911, Zoe. cit., pp.
440-444) in a series of experiments with other species of ticks found that
spotted fever can be transmitted from infected to normal guinea pigs
by nymphal Dermacentor variabilis (Say) iflfected as larvae, by adult
Dermacentor parumapertus marginatus Banl<s and nymphs of Ambly-
omma americanum (Linn.). Ricketts showed transmission by adult
Dermacentor albipictus (Packard) infected <}s nymphs.
The infection in nature. Parker points out that field observations made
in eastern Montana in 1916 and 1917 suggested that under the "epi-
zootilogic" conditions concerned, some ageI1t other than Dermacentor
andersoni Stiles was likely involved in the flatural maintenance of the
virus. In 1923 Parker32 established the fact that the rabbit tick, Haema-
physaliS leporis-palustris (Packard), is capable of transmitting the in-
fection from rabbit to rabbit and also that infected rabbit ticks occur in
nature. A third important fact was established, namely that the infec-
tion is transmitted by infected female ticks to the egg as in the case of
Dermacentor andersoni Stiles. While the rabbit tick only rarely bites
man, it is important indirectly in that it is a potent vector under natural
conditions and is furthermore the only known vector which occurs in
all parts of the United States. The infection carried by this tick is re-
ported by Parker to be extremely mild. Rabbits of all species studied are
hosts of both wood ticks and rabbit ticks. Rabbits are natural reservoirs
of the infection in western North America, field mice in other parts. Dogs
are believed to be important reservoirs of the infection in the closely
related disease, boutonneuse fever of Europe.
The American dog tick, Dermacentor variabilis (Say), was proved
to be a carrier of the eastern type of Rocky Mountain spotted fever in
1931 by Dyer, Badger, and Rumreich,33 who used larvae bred from eggs.
The larvae were fed on a guinea pig infected with eastern spotted fever
and after engorgement were allowed to molt, and the nymphs were fed
to engorgement on a noninfected guinea pig and were then ground up
and injected into fresh guinea pigs, thus establishing a strain of virus
in guinea pigs. The results of these investigators confirmed the early
work of Ricketts and Maver (1911). They also proved that transmission
is transovarian. Parker,34 1937, reports successful stage-to-stage and
generation-to-generation transmission with Dermacentor occidentalis
N eum., the Pacific coast tick, and Rhipicephalus sanguineus (Latreille),
the brown dog tick. In the latter species virus continuity was shown
from larval ticks of one generation through six successive stages to adults
of the next. In Amblyomma americanum (Linn.), Maver (1911) had
already reported larva-to-adult continuity, and transmission from female
to larvae was accomplished by Parker. With Amblyomma caiennense
(Fabr. ), the Cayenne tick, transmission from larvae to adults has been
shown, and for Dermacentor parumapterus marginatus Banks, a rabbit
tick, transmission from nymphs to adults was shown by Maver (1911),
and continuity from larvae to nymphs as well as survival of the virus in
adults was shown by Parker (1937). Parker states that these data are
considered sufficient to indicate that each of these six additional species
is a possible natural carrier of spotted fever virus and that four of these,
D. occidentalis Neum., R. sanguineus (Latr.), A. americanum (Linn.),
and A. caiennense (Fabr.), are possible present or future agents of
transmission to man. Actually Amblyomma americanum Linn., the lone
star tick, is now known to be the vector of spotted fever in Oklahoma
and Texas (Parker, Kohls, and Steinhaus 35 ); and Amblyomma caiennense
(Fabr.) is the vector of the Sao Paulo fever (spotted fever) of Brazil
and of Tobia petechical fever (spotted fever) of Columbia.
Mechanism of infection. The infection is picked up with the bite from
the reservoir animal by a tick in any stage of its life history and is passed
on from stage to stage; e.g., infected blood is ingested by the larva, the
infection is passed on to the nymph (which may infect), and thence to
the adult which in turn may infect, and at least some infected female
ticks will pass the virus on through their eggs to the larvae of the next
generation.
When normal adult ticks are nrst fed on an infected host, just long
enough to ensure ingestion of the virus, and are then transferred directly
to a normal host (the tick-feeding thus being interrupted but essentially
continuous) Parker's37 tests showed a period of from 9 to 12 days be-
tween the ingestion and the transmission of the virus. Except under
experimental conditions infection is not transmitted by the same stage
of the tick that acquires it, but by the next and subsequent stages. The
infection is transmitted by the bite of the tick. The great majority of
persons with cases of Rocky Mountain spotted fever give dennite history
of "tick bite from 2 to 10 or 12 days before onset." The percentage of
ticks that contain infectious virus is reported to be very small: it may
be less than one per cent and is rarely as high as nve.
Parker 37 states that a minority of perhaps 5 per cent of infected per-
sons deny the possibility of having been infected by the tick bite. "It has
long been known that blood-virus of experimental animals would not
pass the unabraded skin, nor would it infect if ingested. The possibility
that tick virus would act differently was not considered until a short
time ago. In an exceptionally clear-cut series of tests it was shown that
tick virus would infect if merely dropped among the hairs on the unab-
raded skin of guinea pigs; also if dropped into the eye. In another
series of tests it was demonstrated, contrary to previously accepted
ideas, that tick excrement is frequently infectious, but tests to determine
whether it will cause infection by contamination of the unabraded skin
have proved negative."
The virus survives the winter in infected nymphal or adult ticks. At
the end of winter the virus in these ticks is present according to Parker
(lac. cit., 1933) as a nonsymptom-producing and frequently immunizing
virus, and does not produce symptomatically recognizable infections
until its level of virulence is raised, either by heat or by the ingestion
of blood. This phenomenon of reactivation has direct bearing on the fact
that the bites of hibernated ticks in nature very early in the spring do
not result in frank infection. It also explains why tick bites grow more
dangerous as the season advances. Parker states, "In the early spring
when the virus in the recently emerged tick is at its lowest ebb of in-
fectiousness, and while the days and the nights are still cool, it takes so
long for the virus to become reactivated after a tick has become at-
tached that it is usually detected and removed before the virus has be-
come virulent." It further gives force to the advice so seriously given
by all authorities on Rocky Mountain spotted fever to detach ticks at
once. It may be a matter of some hours and not infrequently a day or
more after the tick attaches itself before reactivation occurs. The shortest
recorded interval is one and three-quarters hours (Ricketts). If a tick
which has become attached is removed within a few hours, the danger
of infection is materially minimized. The danger from infection in the
Atlantic coast states is later in the summer.
Control and prevention. The control of Rocky Mountain spotted fever
over a wide area by the control of the tick vectors has been discourag-
ing. According to Parker (loc. cit., 1933) definite local good has been
accomplished by dipping, grazing control, and the control of rodents,
but lack of permanence of results and lack of knowledge as to the rela-
tive importance of the various factors of the local complex have all
militated against substantial gains.
In 1926 Cooley obtained parasitized nymphal ticks from a tick
parasite colony established by Larrousse on Naushon Island. The para-
site, introduced originally from France, is named lxodiphagus caucurtei
du Buysson, a chalcidoid parasite of the subfamily Encyrtinae (Hymen-
optera ), which is said to attack only ticks. Cooley 38 introduced these
parasites into Montana, where they were reared in quantity under lab-
oratory conditions, and a field release of 381,190 parasites was made
during the summer of 1928. The parasite has become established but
it is believed unlikely that it will prove useful as an agency of tick con-
trol.
The avoidance of tick bites is urged by keeping out of known tick-
infested areas during spring and early summer; or by wearing suitable
clothing, such as high boots, leggings, puttees or socks outside the trou-
ser legs. The advantage of early removal of ticks that may have escaped
attention and have become attached has already been pointed out.
Parker39 reported in 1935 on the results of the Public Health Serv-
ice vacci~le for the prevention of Rocky Mountain spotted fever and in
1935 (loc. cit.) he comments as follows:
The vaccine has definite protective value. The degree and the duration
of protection vary with the person vaccinated and with the degree of virulence
of the infecting strain of spotted fever virus.
The average person vaccinated in the spring retains marked immunity
during at least the remainder of that year. The immunity is usually' sufficient
to afford full protection against the relatively mild strains of spotted fever
but is apparently progressively less effective as the virulence of infecting strains
is increased.
Colorado tick fever. Parker, in 1937, reported a febrile diseases of
frequent occurrence in many parts of the Rocky Mountain region fol-
lowing the bite of Dermacentor andersoni Stiles. The disease is now
known as "Colorado tick fever." There is a close similarity between
this disease and dengue (see Chapter XII) both clinically and in the
particle size of the respective viruses; however, the two diseases do not
give cross immunity hence "appear to be distinct disease entities."
(Florio, Hammon et al.) 40 Parker describes Colorado tick fever as a
"disease of a remittent type and is commonly characterized by the occur-
rence of two febrile periods, each of two to four days' duration, with
a remission period of one to several days between. The onset is sudden
and the climax is often reached within the first 24 hours. There is no rash.
Symptoms other than fever are malaise, chilly sensations, severe head-
ache, conjunctivitis, photophobia, and generalized muscular and joint
pains with particularly severe aching in the lumbar region. The malaise
is usually intense. Constipation is the rule. The temperature often
reaches 104° to 105° F or over, but may not exceed 101° to 102,0 F. The
pulse rate is frequently 120 to 130. In most instances, though not al-
ways, the symptoms are more severe during the first febrile period. . . .
It is non-fatal."
Tick transmission of tularemia. Parker 41 states that tularemia infection
in ticks was supected in numerous instances during the seasons of 1922
and 1923 on account of the gross lesions at death in guinea pigs into
which such ticks had been injected. Confirmation was made by cultiva-
tion of Pasteurella (= Bacterium) tularensis from guinea pigs in which
the tick strain had been propagated. Dermacentor andersoni Stiles col-
lected from nature proved infective; also experimentally, infection ac-
quired by immature ticks was passed on to subsequent stages of the
same generation. Later Parker and Spencer 42 (1926) demonstrated con-
genital transmission. This is believed to be the first record of "hereditary
transmission" of a known bacterial infection. Probably several species of
ticks are able to transmit the infection; Dermacentor occidentalis Neum.
and Dermacentor variabilis (Say) have been incriminated. Haemaphy-
salis leporis-palustris (Packard), the so-called rabbit tick, is largely
responsible for the maintenance of the infection in nature. Pasteurella
tularensis has been recovered both from infected sage hens in Montana
and from the tick, Haemaphysalis cinnabarina Koch, taken from the
same birds. Davis and Kohls 43 (1937) discovered evidence indicating
that Ixodes pacificus Cooley and Kohls may also be a carrier of tularemia
to human beings.
Tick paralysis. A paralysis of sheep and calves attributable to ticks
has been known in Australia according to Henning,5B since 1843. Paralysis
reported as "acute ascending paralysis" associated with tick bite was
described in 1912 by Temple H in Oregon. The case reported was that of
a child in which there was a complete paralysis of the motor and sensory
nerves extending to the knees, causing inability to stand in the morning
after retirement in apparent good health. On the third day the paralysis
had involved the nerves of the throat and the child was unable to
swallow or speak. Upon removal of two fully engorged ticks from the
occipital region recovery was rapid and complete within a week. The
ticks were not positively identified, though they were presumably
Dermacentor andersoni Stiles. In 1913 Hadwen 45 reported cases of paraly-
sis in sheep following the bite of Dermacentor andersoni Stiles ( D.
venustus Banks). He also cites excerpts from letters (Canad. M. A. J.,
1912) from physicians in British Columbia indicating frequent occur-
rence of paralysis in children following tick bite. The ticks were com-
monly removed from the nape of the neck.
In 1913 Hadwen and Nutta1l 46 report having produced paralysis ex-
perimentally in the dog by means of Dermacentor andersoni Stiles (D.
venustus Banks). The paralysis was the same as in sheep. In the experi-
ment reported, paralysis was caused by a single tick in eight days. The
authors state: "On the hypothesis that the symptoms are due to toxins
given off by the tick, 'the period of incubation' might be explained on
the supposition that it is only when the tick commences to engorge or
feed rapidly, some days after it has becbme attached, that it gives off
the hypothetical toxin in its saliva in sufficient quantity to produce patho-
genic effects."
Tick paralysis is widely known in the western United States and
Canada adjacent to the Rocky Mountains and coincides with the distri-
bution of Dermacentor andersoni Stiles. Hearle, 1938 (loc. cit. p. 399),
states:
As an index of the wide distribution of this trouble in British Columbia,
incomplete returns from medical practitioners indicated that nearly 150 human
cases had been noted in the province. . . cases in sheep have been par-
ticularly numerous, and many deaths have resulted. Cattle are usually less
susceptible, but trouble from tick paralysis has been noted from time to time,
and in the spring of 1930 a serious outbreak in steers was investigated; over
100 paralysis cases, sixty of them fatal, being noted in one herd. We know
of only one equine case. In sheep districts where this trouble is prevalent,
flock masters are obliged to examine their animals frequently for the purpose
of removing the offending ticks from sheep showing symptoms of weakness or
staggers. . . . When animals become paralyzed all ticks showing engorgement
should immediately be removed; in searching for them special attention should
be paid to the back of the head, the region of the spine and along the neck,
since this is where they mainly become attached.
Ixodes holocyclus Neumann causes paralysis in calves, sheep, and

dogs in Australia. According to Ross47 this condition is caused only by
the mature engorging female tick two days before it is ready to drop
from the host. He used the dog in his experimental work. Several mild
cases of paralysis in children have been reported to the author in Cali-
fornia, evidently as the result of attachment by Ixodes pacificus C. and
K. Dermacentor variabilis (Say) may also be incriminated.
Bullis fever, also known as "lone star fever" referring to the vector,
the lone star tick, is a virus infection, resulting in an unusually low
white blood cell count and a severe postorbital and occipital headache
and lymphadenitis. The onset is usually abrupt, with an initial chill or
chilliness; fever soon follows; and there is pronounced lassitude and
prostration, with nausea. The fever lasts from 4 to 14 days. Con-
valescence is protracted. (Woodland, McDowell, and Richards,18 1943.)
This disease was first observed at Camp Bullis, Texas, in 1942. Anig-
stein and Bader,49 as well as the above-mentioned group, express the be-
lief, based on both epidemiological and experimental evidence, that the
vector of this infection is Amblyomma americanum (Linn.), known as
the "lone star tick"; it is believed to be at least one of the carriers of
Bullis fever in nature.
Q fever was first reported from Australia (Queensland) in 1937,50
where most of the infected persons were abattoir workers; the causal
organism is Coxiella (= Rickettsia) burneti (Derrick) (1939). The
disease is characterized by an acute onset with chills, prostration, and
a continuous fever, which lasts from a few days to two to three weeks;
death seldom results. In 1937 the. finding of a rickettsial infection was
reported in the tissues of animals in western Montana (U.S.A.); the
organism was later named Rickettsia diaporica by COX 51 and the disease
was called "Nine Mile Fever," "Nine Mile Creek" being the place in
Montana where Dermacentor andersoni ticks were collected and proved
naturally infected with Rickettsia diaporica,52 now shown to be the same
as Coxiella burneti. The first known human infection in the United
States, a laboratory infection, was reported by Dyer (1938), who pOinted
out its relationship to Australian Q fever (see Steinhaus, 53 1946, pp. 295-
302). In 1943, Parker and Davis 54 reported the Rickettsia (Coxiella) of
American Q fever in the lone star tick, Amblyomma americanum Linn.
In Australia Haemaphysalis humerosa Warburton and Nuttall, which
does not bite man, is considered an important vector of the infection
among bandicoots (Isodon spp.), serving as an important factor in main-
taining endemicity of the disease. Rats are also regarded as important
reservoirs. Ixodes holocyclus Neum., which bit_es man and marsupials,
may be a factor in areas where the disease exists in marsupials, in that
the infection may be deposited on the skin with tick feces or may be
otherwise transmitted. Infection may result through the unabraded skin
when it is contaminated with tick feces. Actual transmission by the bite
of ticks has not been satisfactorily demonstrated. Transovarial passage
in ticks has been observed. Several species of ticks other than those
mentioned can at least be experimentally infected.
Philip 55 (1948) reports that feces from infected Dermacentor andersoni
are very rich in rickettsiae, which remain viable in storage as long as
586 days. Infection from tick feces emulsion penetrated the unbroken
skin as well as the abraded skin of test animals; also powdered feces
dusted in the nostrils, eyes, and mouth produced infection. Philip also
recovered the infection from houseflies, Musca domestica, caught at
large in an animal room in which animals infected with Q fever were
kept. Attempts to transfer the infection through these flies to normal
animals failed.
In their Q fever studies in California, Huebner and Jellison and as-
sociates 56 recovered Coxiella burneti from raw milk, and they conclude
that raw milk may serve as a source of infection to man by some mode
as yet undetermined. Available evidence did not indicate that the drink-
ing of raw milk was the cause of the majority of cases thus far studied.
Lymphocytic choriomeningitis is an infectious disease caused by a
virus which is discharged in nasal secretions and in the urine of naturally
infected mice. The manner in which naturally infected mice contract the
disease is unknown. Experimental infection of all stages of Dermacentor
andersoni with the virus was demonstrated by Shaughnessy and Milzer 57
( 1939) by feeding the ticks on infected guinea pigs. They report stage-
to-stage transmission of the virus from larvae to nymphs and nymphs to
adults, as well as generation-to-generation transmission from adults to
eggs. Typical symptoms were produced in all test animals, and typical
pathology was shown by those that had died; surviving animals were
immune to inoculation of the virus. Feces from feeding adult ticks ap-
plied to the lightly scarified skin of guinea pigs caused typical symptoms.
East Coast fever is a highly fatal disease of cattle along the East Coast
of equatorial Africa. The mortality, it is said, may run over 90 per cent.
The disease is caused by the protozoon, Theileria parva (Theiler), and
is therefore a form of piroplasmosis ( theileriosis) though, unlike red
water, it is not readily transmitted by means of blood inoculations, nor
is it accompanied by jaundice or hemoglobinuria. A very characteristic
symptom is swelling of the superficial lymphatic glands.
The incubation period varies from 9 to 19 days and is usually from
10 to 15 days. The disease is transmitted by several species of ticks as
reported by Lounsbury as early as 1906. He proved the adult brown
tick, Rhipicephalus appendiculatus Neum., to be a vector, and later he
showed that the disease may also be transmitted by the Cape brown
tick, Rhipicephalus capensis Koch, and the red tick, Rhipicephalus evertsi
Neumann.
Henning 58 states that unlike red water, East Coast fever is not trans-
mitted from the adult female tick to the larva through the egg, but only
by an adult tick which became infected during its nymphal stage, or by
a nymph that became infected during its larval stage. While RhipiC-
ephalus appendiculatus Neum., R. capensis Koch, and R. simus Koch
are three-host ticks, R. evertsi Neum. is a two-host tick, hence as the tick
remains on the same host during both its larval and nymphal stages
transmission of the infection is pOSSibly only during the adult stage. The
larvae of the two-host tick, R. evertsi, hatch from eggs on the ground
as do other ticks, then proceed to attach themselves to the ears (often
inside) and the Hanks of cattle. The larvae become fully engorged and
molt while on the host. The nymphs then engorge and drop off the host
to molt and give rise fa the adult. The adult then attaches itself to a
second host, engorges, and drops to the ground where the females lay
eggs. Lounsbury and Theiler both found that ticks of the three-host
species which have sucked infected blood during their larval stage can
transmit the infection only ouring their nymphal stage; i.e., whether the
nymph feeds on either a susceptible or a non susceptible host, infectivity
is lost. A single tick can transmit the infection only once, and that during
the stage that follows the one in which it had the infectious meal.
Equine piroplasmosis. At least two types of piroplasmosis are found
in horses, mules, and donkeys, namely true equine piroplasmosis, trace-
able to Babesia caballi (Nuttall), occurring in Africa, Russia, Trans-
caucasia, and probably Siberia, and secondly a similar though distinct
disease traceable to Nuttallia equi (Laveran) occurring in Transcaucasia,
Italy, Africa, India, and South America (Brazil). Babesia caballi is
transmitted by Dermacentor reticulatus (Fabr.) in Russia; and Nuttallia
equi is transmitted by Rhipicephalus evertsi N eum. in South Africa.
Canine babesiosis (piroplasmosis), also known as "malignant jaundice"
of dogs, is prevalent in southern Europe, Asia, South Africa, and more
recently in the United States (Florida). The causal organism is Babesia
canis (Piana and Galli-Valerio); the carrier is Rhipicephalus sanguineus
(Latr.) in India, southern Europe, and the United States; Haemaphysalis
leachi (Audouin) is a South African vector, and Dermacentor reticulatus
(Fabr.) and Ixodes ricinus (Linn.) transmit the infection in southern
Europe. Brumpt and Larrousse 59 have shown that Dermacentor andersoni
Stiles can carry the disease. The infection is hereditary in the tick, but
transmission to the dog is effected by the bite of the adults, but not by
the larvae and nymphs, according to Brumpt. 60 The incubation period
varies from 10 to 20 days. Sanders 61 (1937) reported that R. sanguineus
(Latr.) is by far the most common species encountered in kennels and
on animals affected with canine babesiosis in Florida.
Sanders (lac. cit.) states that the acute form of this infection is not
difficult to diagnose; "the high temperature, increased pulse and respira-
tion, progressive anemia, jaundice, the history of tick infestation and the
demonstration of the causal organism are usually observed."
Heartwater, a dreaded disease of South African sheep, goats, and
cattle is caused by Rickettsia ruminatium (Cowdry). It is carried by the
"bont" tick, Amblyomma hebraeum Koch.
Henning62 states that the bont tick may subsist on any warm-blooded
animal; it has been found during all its stages on man, all species of
domestic animals, and several species of antelopes. Bovines are its fa-
vorite host. This tick may lay as many as 18,000 eggs. The tick becomes
a vector of the infection only after it has sucked blood from a diseased
animal; i.e., hereditary infection does not occur, hence the newly hatched
larvae are not infectious. Amblyomma variegatum (Fabr.) is able to
transmit the infection in the adult stage.
Bovine anaplasmosis. Anaplasmosis is an important and practically
world-wide infection of cattle caused by minute punctiform blood paril-
sites described by Theiler in 1910 as Anaplasma marginale with the
organism at or near the periphery of the red cell and Anaplasma mar-
ginale variety centrale, a somewhat smaller body, located approximately
in the center of the infected corpuscle.
Anaplasmosis is described by Stiles 63 as an acute, subacute, or chronic,
febrile, infectious, protozoon disease, characterized by loss of Hesh,
labored breathing, suspension of milk How, anemia, jaundice, and
marked degenerative changes in the red blood corpuscles owing to the
activity of the microscopic parasites. The average mortality ranges from
30 to 50 per cent in the animals affected. Mechanical transmission of the
infection by several species of Tabanid Hies has been reported, and Stiles
records a total of seventeen species of ticks which have been incriminated
by various investigators in transmission, among them Boophilus annulatus
(Say), B. decoloratus (Koch), B. microplus (Canestrini), Rhipicephalus
simus Koch, R. bursa Canestrini et Fanzago, Ixodes ricinus (Linn.),
Hyalomma lusitanicum (Koch), Rhipicephalus sanguineus (Latreille),
Dermacentor variabilis (Say), D. andersoni Stiles, and D. occidentalis
Neum.
In 1936 Herms and Howe1l6{ reported five cases of tick transmission,
two, nonhereditary, with Dermacentor albipictus (Packard) and D.
andersoni Stiles, and three in which the infection passed through the
eggs of the western dog tick, Dermacentor occidentalis Neum, the off-
spring being infective in both the larval and nymphal stages. In the
congenital transmissions from the time the infective larval ticks were
applied to the host animal to the time that marginal bodies first ap-
peared in the blood the elapsed time was 37 days, 32 days, and 123
days respectively. In the two nonhereditary transmissions the elapsed
time was 28 and 29 days. Howell, Stiles, and Moe 65 have demonstrated
transovarian transmission by Dermacentor andersoni Stiles.
That deer, namely the southern black-tailed deer, Odocoileus hem-
ionus columbianus (Richardson) and the Rocky Mountain mule deer,
Odocoileus hemionus hemionus (Rafinesque), may serve as reservoirs
for anaplasmosis was proved by Boynton and Woods in 1933. 66
COMBATING TICKS (IXODINE)

Ticks on live stock. Rotenone-sulphur dips are commonly used for
the control of ticks on the bodies of livestock. The dip is prepared as
follows: 10 pounds 5 per cent rotenone, 10 pounds wettable sulphur
(325 mesh or finer), 1 pint wetting agent, 100 gallons water. This dip
is nontoxic to the animals and according to Stewart and Furman (mimeo.
series, Univ. of California, Agric. Exp. Sta.) generally gives effective
control. This formula is also effective against lice, cattle grubs, and sheep
keds.
Arsenical dips are likely to be more effective but are also more toxic.
These dips are made as follows: 24 pounds sodium bicarbonate (sal
soda), 8 pounds arsenic trioxide, 1 gallon pine tar, 500 gallons water.
This dip is particularly useful in the control of the Texas cattle fever
tick, Boophilus annulatus. 67 From the time the arsenic is purchased to
the final disposal of the old dip, great care must be exercised in storing,
handling, and using it because of its very toxic nature. The animals
should always be watered a short time before they are dipped so that
they will not be thirsty and drink the dip. After the animals emerge from
the dipping vat, they should be kept on a draining floor until the dip
ceases to run from their bodies; then they should be herded into a place
free of vegetation until they are entirely dry. If they are allowed to drain
in places where pools of dip collect from which they may drink or if
they are turned at once to pasture where the dip may run from their
bodies on grass or other vegetation, serious effects may result. Also
crowding animals before they are dry should be avoided, and they
should not be driven far within a week after dipping, especially in hot
weather.
After the animals are watered, they are driven from the pen one by
one into the chute and thence into the dip. The plunge will wet the
animal all over, but a second plunge of the head by using a forked
stick is necessary as the animal proceeds through the dip. After emerging,
the animals should be driven to a tick-free pasture (see above precau-
tions ); the dipping process should be repeated in from 7 to 10 days.
Dipping vats contaihing arsenical dip must be protected when not
in use; also when vats are emptied for cleaning the dip must not be
allowed to flow on land or vegetation to which cattle or other animals
have access; seepage into wells or springs must also be guarded against.
Other dips. Benzene hexachloride has been reported as effective
against several species of ticks on live stock. A 0.5 per cent suspension
of BHC (of 10-13 per cent gamma isomer) is suggested. Lactating dairy
cattle must not be given a benzene hexachoride dip since the milk may
have an extraneous odor for a few days following dipping. Likewise
cattle that are to be slaughtered within 60 days should not be dipped
in this solution. However, no harm to cattle results from its use.
DDT has been reported as effective in controlling the winter tick,
Dermacentor albipictus, when used as an emulsion of 0.8 per cent DDT.
Information concerning this dip is incomplete.
To free dogs of ticks a thorough application of derris powder or wash
is recommended by Bishopp and Smith. 68 Derris when used as a powder
should have a rotenone content of at least 3 per cent and should be
applied lightly next to the skin at intervals of two or three days. A dip
or wash can be made by dissolving an ounce of neutral soap in a gallon
of tepid water and mixing two ounces of derris power of which the
rotenone content is 3 per cent. The dip should be applied at intervals of
five or six days. The dog may be put in a tub or a brush may be used.
The dip is allowed to dry on or if necessary surplus liquid may be
removed with a towel. Do not permit the dip or powder to get into the
eyes of the animal.
The application of the powder is simpler and kills the ticks with
which it comes in contact, but a more thorough covering is secured when
a wash or dip is applied. The wash also has a longer repellent action.
Kennels in which the dogs sleep should be thoroughly sprayed with a
coal tar creosote solution, but this should not be allowed to come in
contact with the dogs. Since the dog is the principal host upon which
the adult Dermacentor variabilis (Say), the vector of eastern Rocky
Mountain spotted fever, feeds, it is important in these areas to keep dogs
tick-free. It is also important that humans exercise every precaution to
avoid getting ticks on the body in order to prevent infection.
Control of ticks on vegetation. As has already been explained, when
Ixodine (hard) ticks reach maturity on the host they drop to the ground
where the eggs are deposited and the seed ticks (larvae) emerge, climb-
ing up low shrubbery, weeds, and grass ready to cling to any suitable
passing animal. Enormous numbers of larvae may thus be encountered,
particularly along cattle and deer paths in more or less wooded areas
and on the well-marked paths of meadow mice and other rodents in
less wooded situations. As has also been explained, in the commoner
species of hard ticks the nymphs likewise drop to the ground to molt.
In some areas large numbers of nymphs and young adult ticks (before
engorgement) climb up the branches and twigs of low shrubbery, where
they are in an excellent position to attach themselves to larger animals,
including man, as these brush against the vegetation; wild lilac (Cean-
othus) where abundant and thus infested in commonly called "tick
bush." Manifestly much good could be accomplished if an economical
insecticidal treatment were available both for use on the ground and on
vegetation along such pathways.
Smith and Gouck,69 1945, report marked success with both DDT
sprays and dusts against Amblyomma americanum and Dermacentor
variabilis. A spray was used containing one avoirdupois ounce of DDT
(technical grade), 5 fluid ounces of soluble pine oil, and water to make
15 gallons; the entire surface of the ground and all vegetation up to 3
or 4 feet above ground were covered. No injury to plants was apparent
in any of the tests. At the rate of one pound of DDT per acre highly
satisfactory control was maintained for a period of about two months,
proving pronounced residual effect on ticks. Dust containing 5 per cent
DDT in pyrophyllite was effective at 3 pounds per acre.
FAMILY ARGASIDAE (SOFT-BODIED TICKS)

The family Argasidae includes the so-called soft-bodied or nonscutate
ticks, in which sexual dimorphism is slight. The integument of all stages
except larvae is leathery, wrinkled, granulated, mammillated, or with
tubercles. The capitulum is either subterminal or distant from the anterior
margin; in larvae the subterminal or terminal capitulum lies in a more
or less marked depression, the camerostome. The articulations of the
palpi of all stages are free; the porose areas are absent in both sexes.
There are about 60 species. Cooley and Kohls 70 (1944) provide us with
an excellent account of this family. They recognize four genera, namely,
Argas, Ornithodoros, Otobius and Antricola.
GENUS ORNITHODOROS
The genus Ornithodoros. In this genus the capitulum is either sub-

terminal or distant from the anterior margin; the hypostome is well
developed. In the integument discs and mammillae commingle in a
variety of patterns; hood, camerostome, cheeks, and eyes are present
or absent; dorsal humps and subapical dorsal protuberances on legs are
progressively more prominent in the successive nymphal stages. The
body is more or less flattened but strongly convex dorsally when dis-
tended; the integumental pattern is continuous over the sides from dorsal
to ventral surfaces. (Cooley and Kohls, loco cit.). The genus includes
about 50 species, some 20 of which occur in North America, Central
America, and Cuba.
Ornithodoros moubata (Murray) occurs only in Africa where it is a
widely distributed vector of relapsing fever, hence is known as the Afri-
can relapsing fever tick (Fig. 159). It occurs in native huts, hiding in
dust and thatch. It feeds chiefly at night and engorges rapidly. It is an
eyeless species with a specific arrangement of the "humps" on the pro-
tarsus of the nrst pair of legs, being "subequal and tooth-like." The adults
measure from 8 to 11 mm in length and about 7 mm in breadth. The
color varies from dusty hrown to greenish brown in living specimens.
Eggs are deposited in batches of from 35 to 340 at intervals after blood
meals during the lifetime of the female. The maximum number of eggs
laid by one female was 1,217 according to Jobling. 71 The incubation
FIG. 159. Mrican relapsing fever tick, Ornithodorus

moubata. X3.
period lasts from 7 to 11 days and over, depending on temperature.

Davis72 calls attention to the fact that the larvae are completely quiescent,
with legs closely applied to the body, and within a few hours molt in the
split egg shell to the first nymphal stage. The form that many authors
refer to as the newly hatched larva is actually the nrst nymphal stage,
which attaches itself to its first host animal and remains there for about
a week feeding on lymph, then disengages itself and molts for the second
time. There then follow several other feeding periods and molts, six to
nine in all before sexual maturity is reached. Young females may molt
even after sexual maturity. This species will feed on a wide range of
host animals besides man including pigs, dogs, goats, and sheep; also
on rabbits, mice, rats, monkeys, and fowls in the laboratory. It appears
to be essentially a parasite of man, however, and is a man-to-tick-to-man
vector of relapsing fever.
Since the discovery that African relapsing fever is transmitted by
Ornithodoros moubata (Murray), many other species of Ornithodoros
have been found naturally infected with spirochetes infective to man;
indeed some believe that any species of this genus is capable of trans-
mitting all strains of relapsing fever normally transmitted by ticks belong-
ing to this genus. With this the author cannot fully agree, since he and
his students have been unable to transmit the California strain by means
of Ornithodoros coriaceus Koch; indeed tick-borne spirochetes seem to
possess a tick fastness or specificity to a high degree.
Ornithodoros talaje (Guerin-M(meville) is a South and Central
American (south to Argentina) and Mexican species, occurring also in
Florida, Texas, Arizona, Nevada, and Kansas, and has been taken in
several California counties,13 It feeds on wild rodents, also on swine,
cattle, horses, man and other mammals. It inflicts a very painful bite.
It is the vector of relapsing fever in Panama, Venezuela, and Colombia.
Bates, Dunn, and St. John 74 proved this tick a vector of relapsing fever
in Panama by human experimentation in 1921; in one instance the bites
of a naturally infected tick resulted in infection. This species has been
reported from New York by Matheson,75 who remarks, "How this tick
reached there (Ransomville, N. Y.) can only be surmised. It probably
maintained itself in the heated house and fed on the occupants or more
probably on mice or rats or the domestic cat and dog, though none of
these is listed as its normal host."
Ornithodoros venezuelensis Brumpt is a Central and South American
species. It transmits relapsing fever in Colombia, Venezuela, and Panama.
Dunn76 collected 4,880 ticks of this species in 68 homes in twenty villages,
towns, and cities in various parts of Colombia. Ticks infested with relaps-
ing fever spirochetes were present in nearly 28 per cent of the homes
in which collections were made. The altitude mentioned for one of the
localities, a barracks at Muzo, was 2,700 feet. Like O. moubata this
species appears to be essentially a parasite of man, although it is known
to feed on other animals.
Ornithodoros erraticus Lucas occurs in Spain and northern Africa.
It is an important vector of relapsing fever in northern Africa. O. pap-
illipes Birula [0. tholozani (Lab. et Megnin) 1 is a vector in Central
Asia. 77 O. normandi Larrousse is a very small Tunisian species, reported
to be a vector of relapsing fever.
Ornithodoros parkeri is described by Coolei s from ground squirrels
(Citellus sp. ), jack rabbits (Lepus), and the cottontail in Wyoming and
Washington.
Tick-borne relapsing fever. Although we are told that the natives in
many parts of Africa dreaded tick bites for many generations, David
Livingstone, the explorer, was the nrst to report (1857) upon the evil
effects following the bite of a tick which was later named Ornithodoros
moubata (Murray, 1877). It was not, however, until 1904 that Ross and
Milne,so and in 1905 Dutton and Todd,S! reporting from Uganda and the
Congo, and almost simultaneously Robert Koch reporting from German
East Africa, gave us the knowledge that these evil effects were due to
relapsing fever, a spirochetosis, and that the tick so dreaded by the
natives, was actually the vector.
The symptoms of the disease are described by Nuttall and Warburton
(loc. cit., 1908), as follows: "Headache (especially at the back of the
head), vomiting, abdominal pains and purging, with severe fever, a pulse
of 90 to 120, dry hot skin, congested eyes and shortness of breath. After
a period of fever lasting about two days, there is a fall of temperature,
but a fresh attack soon follows. These relapses occur more frequently
than in European (louse-borne) relapsing fever, being usually 5 or 6 in
number, but there may be more. The attacks leave the patient in a weak
condition for a long time after recovery, which usually follows, but death
occurs in about 6 per cent of the cases."
How infection is transmitted. Dutton and Todd (lac. cit.) proved
that the infection is not only introduced with the bite of the tick but is
also transmitted to the offspring of the female tick through the egg. The
newly hatched ticks were proved to be infective. Once infected the tick
remains so, and the infection may be transmitted from generation to
generation without renewal for at least three generations. The infection
is transmitted by the bite of either the male or the female tick during
all of its active stages. The coxal fluids are believed to play an important
role since they contain spirochetes which are washed into the tick feces
rendering these infectious as well. The attack of fever takes place in the
human in from 5 to 10 days after the tick has bitten. Feng and Chung82
report that "shortly after the spirochaetes are ingested by the ticks they
penetrate the stomach wall and reach the body cavity as evidenced by
finding spirochaetes in the legs six hours after the infective feed. . . .
The spirochaetes gradually disappear from the stomach and reach the
body cavity with the result that from the twelfth day on no more spiro-
chaetes could be found in the stomach contents. . . . From the body
cavity the spirochaetes invade the salivary glands, the coxal glands.
. . . The constant presence of numerous spirochaetes in the acini of the
salivary glands and the finding on several occasions of spirochaetes
actually inside the small salivary ducts suggests that besides the coxal
fluid the bite alone may also be infective. . . . [The spirochetes] multi-
ply by transverse division after they have penetrated the stomach wall
and reached the body cavity and other organs of the tick. . . . Mice in-
oculated . . . with feces remained sterile."
Tick-borne relapsing fever in the United States. The earliest known
focus of endemic (tick-borne) relapsing fever in the United States in
believed to have been in Colorado, where cases were reported by
Meador83 in 1915; this focus was in the mountains near Denver. In 1921
Briggs84 reported two cases of relapsing fever in which the infection had
been acquired at Polaris, Nevada County, California, at an elevation of
about 5,750 feet. It would seem that Briggs suspected lice since he
remarks, "Many tramps, put off trains at Truckee, find a day or so of
employment here, only to move on. It is quite evident, therefore, that
there are great opportunities for the dissemination of vermin by a no-
TICKS AND TICK-BORNE DISEASES 50.5
madic population." According to a letter to Briggs from Mark F. Boyd,
ticks collected some years previously in the vicinity of Lake Tahoe
(vicinity of Briggs's cases) in northern California were sent by Boyd
to Nathan Banks, United States National Museum and were identified
as Ornithodoros tala;e (Guerin Meneville), a species known to be a vector
of relapsing fever in Panama. This was evidently an incorrect identifica-
tion since diligent search for that species in this neighborhood proved
futile. However, in searching for a vector during the summers of 1931-
1933, an Ornithodoros tick, new to science, was discovered by Wheeler
(1935) and was given the name Ornithodoros hermsi Wheeler.85 A re-
examination of one of the ticks sent by Boyd to Banks proved it to be a
specimen of the new species described by Wheeler.
It is of interest to note that prospectors and others who worked in
the Sierra Nevada at altitudes of 5,000 feet and over frequently reported
suffering from a malaria-like disease which they called "squirrel fever."
In the light of the discoveries described here and particularly the ex-
perience of C. M. Wheeler noted below, it is obvious that this infection
was relapsing fever, infection having been due to contact with the blood
of spirochete-infected squirrels or due to tick bites not then suspected.
Porter, Beck, and Stevens86 state that, since the cases reported by
Briggs, no further cases of relapsing fever came to the attention of the
California State Department of Public Health until September, 1930,
when Dr. George Stevens reported a case of a school teacher who had
lived at Big Bear Lake (San Bernardino County) during July and August.
In 1930 Major V. H. Cornell reported a case in which infection had been
contracted at Lake Tahoe (Eldorado County) during July. In June, 1932,
C. M. Wheeler, an assistant on field duty, contracted relapsing fever
at Packer Lake (Sierra County) by accidentally smearing his fingers with
blood from a Sierra chickaree squirrel (Sciurus douglasii) which with
four othe;s he had shot a few minutes previously.81 The blood from the
squirrels proved positive for spirochetes (Spirochaeta recurrentis
Lebert). Seven days after the accident Wheeler came down with relaps-
ing fever.
On August 12, 1931, three specimens of mature, undescribed Ornitho-
doros ticks were taken in a cottage near Brockway on Lake Tahoe (El-
dorado County) at an elevation of approximately 6,000 feet, where cases
of relapsing fever had occurred about a month previously. In August,
1934, more ticks of the same undescribed species were collected at Big
Bear Lake at an elevation of about 5,700 feet in a cabin where there had
been cases of relapsing fever. Other specimens were taken in various
localities of the three counties already named (Eldorado, Placer, and
San Bernardino) at elevations of from 5,000 to 8,000 feet. This new
species of tick was named Ornithodoros hermsi by Wheeler (loc. cit.)
in 1935, and at the same time Wheeler, Herms, and Meyer88 reported
transmissions to a monkey and white mice by the bite of this tick.
The first demonstration of the vector capacity of this new species
was made in October, 1934, on a monkey bitten by ticks collected in an
endemic area. (Wheeler, Herms, and Meyer.) The monkey showed the
first characteristic symptoms 16 days after being bitten by the ticks.
Proof that this species is capable of transmitting the infection to human
beings was secured by Wheeler 89 in May, 1937, when an experimental
infection resulted in a series of tests made on human subjects. From
1921 to 1944, inclusive, 283 cases of relapsing fever were reported in
California, with epidemiological evidence pointing to their origin in the
Sierra Nevada at elevations of 5,000 to 10,000 feet. The same hitherto
undescribed tick was collected in significant numbers in the vicinity of
Big Bear Lake in southern California (San Bernardino County) at an
elevation of 6,700 feet during 1933 and 1934. This tick, Ornithodoros
hermsi, has since been taken in all endemic areas as high as 10,000 feet
elevation.
It should be added here that a more northerly focus of human relaps-
ing fever was reported in 1940 at the foot of Mount Lassen, Manzanita
Lake (California), elevation about 5,850 feet. Clinical human cases were
recognized, and ticks Ornithodoros hermsi taken in chipmunks' nests
in the infected cabin proved positive when allowed to bite white mice.
Ticks of this species have now been taken as far north as Kamloops,
British Columbia.
Weller and Graham 90 reported infected ticks in central Texas in
1930; they found a cave in the Colorado River Valley of central Texas
which was "literally alive with ticks, a handful of sand yielding thirty
or forty of different sizes." The cave, it was reported, is "frequented by
goats and sheep, also probably wild animals such as bats, foxes, skunks
and rabbits." Some of the ticks were applied to three rabbits, allowed
to feed for 15 minutes, and then crushed and rubbed into the abrasions.
Spirochetes were later observed in the blood of the rabbits.
Ornithodoros hermsi Wheeler (Fig. 161), the vector of relapsing
fever in California, transmits the infection by the bite of both male and
female ticks and in all stages of development. The proportion of infective
larvae in hereditary transmission appears not to exceed one per cent.
Wheeler 91 reports that from 35 per cent to 48 per cent of noninfective
ticks when allowed to feed as larvae on infected laboratory white mice
were able to acquire the spirochetes and transmit them to normal ani-
mals in some one or all of the subsequent developmental stages. One
female tick has caused four infections in white mice during a period of
about four months. Larvae from a presumably noninfective female tick
produced infections, and conversely the larvae of an infective female may
not be infective although occasionally an infection may result from bites
of later stages. The spirochetes are usually present in the blood of white
mice about five days after the animals have been bitten by an infective
tick. The time elapsing before infected ticks transmitted the infection
depended, of course, on the time when the ticks were ready to feed again
with molts intervening; this elapsed period was 15 to 20 days at the
shortest.
The life history of Ornithodoros hermsi Wheeler (Fig. 160) as de-
scribed by Herms and Wheeler79 is as follows: The very tiny light amber-
colored eggs are deposited at intervals in batches of 12 to 140 from May
to October and range well over 200 per female. A specific example fol-
A B c D
FIG. 160. Immature stages of Omithodorus hermsi, (A) larva, (B) first nym-
phal stage, (C) second nymphal stage, and (D) third nymphal stage.
lows: Female tick, no. 5, taken as a last-stage nymph August 17, 1934,
deposited a total of 232 eggs in four batches, 98 (April 8), 73 (May 19),
49 (May 26), and 12 (June 12) with but one feeding (i.e., after first
batch was laid) between egg-layings, and died October 21, 1935. Under
natural conditions the eggs are deposited in the hiding places of the
ticks: in summer cabins (Fig. 162) the eggs are laid in such corners and
crevices as afford protection to the ticks.
The incubation period at a temperature of 75° F and 90 per cent
humidity ranges from 15 to 21 days. The number of eggs and the per-
centage of larvae hatching seems to grow less in the later egg layings,
decreasing from as high as 95 per cent for the first batches to less than
50 per cent in the last. The first molt is usually accomplished within the
egg; however, the larva (seed tick) remains hexapod until after the
second molt. After about three days the larvae is ready to feed, remain-
ing attached to the host for only about 12 to 15 minutes as is the case
in later stages, although this attachment in the latter may be for from
one-half to one hour in many cases. The larvae when fully engorged
increase as much as three times in size and acquire a bright red color
due to the imbibed blood. In this condition these tiny ticks have been
referred to as a "strawberry seed insect" by persons living in relapsing
fever areas. Molting takes place in about 15 days after feeding. With
this molt the fourth pair of legs appears, and this stage is termed the first
nymphal instar. Ticks in the first nymphal stage may feed within a few
hours after molting, and again a period of 11 to 15 days elapses before
the third molt and the appearance of the second nymphal instar. Then
follows the third feeding and again an elapsed period, in this case about
FIG. 161. Ornithodoros hermsi. (Top left) mature female, (top right) mature
male, (bottom) female despositing eggs.
10 to 32 days, before the fourth molt and the appearance of the third
nymphal instar, or even the adult may appear with this molt. Usually a
fifth feeding and a fifth molt are necessary before sexual maturity is
reached. Egg laying may begin in about 30 days after the last molt,
fecundation taking place in a few days after maturity is reached. The
cycle from egg' to egg under our laboratory conditions required about
four months-e.g., from April 29, when eggs were laid, to August 24,
when eggs were laid by a female from the April 29 hatch of eggs.
The life cycle may be greatly prolonged in the absence of food be-
cause of the ability of these ticks to withstand starvation; thus larvae
may live as long as 95 days without food; unfed nrst-molt nymphs may
live as long as 154 days; unfed second-stage nymphs may live as long as
79 days; third stage, as long as 109 days, and adults well over 7 months.
Adult ticks have been kept alive in pill boxes with occasional feedings
for a period of over four years.
The mature female tick (Fig~ 161) measures from 5 to 6 mm in
length by 3 to 4 mm in width. The male resembles the female closely in
general appearance but is slightly smaller. This species is described as
ovoid, conically painted anteriorly, broadly rounded posteriorly. The
anterior dorsal portion of the hood is visible from above. Unengorged
specimens are of a light sandy color with the black of the intestinal
diverticulae visible through the integument ()f the dorsal surface; freshly
engorged specimens are of a dun, deep garnet shade with a grayish
sheen over the body. The anterior conical point is whitish. The color
changes to a grayish blue a few days after feeding. Legs and hood are
pale yellow. In newly molted forms the body and legs are lighter but
gradually assume the light sandy appearance and the leg darken cor-
respondingly.
Tick vectors in various parts of the world. The known vectors of
relapsing fever in the United States92 are Ornithodoros turicata (Duges)
in Texas and Kansas, and O. hermsi Wheeler in California, Colorado, and
Idaho. O. talaje (Guerin-Meneville) has been conected in Arizona and
California, and also O. parkeri Cooley,93 the only known species in a
large area from which a number of cases of relapsing fever have been
reported; however, the latter species has not been adequately tested on
humans.
In Mexico the vector is Ornithodoros tflricata (Duges). In Central
America O. talaje (Guerin-Meneville) and O. venezuelensis Brumpt are
vectors; in South America, O. venezuelensis and O. talate; in Africa, O.
moubata (Murray), O. savigngi (Audouin), and O. erraticus Lucas; in
Asia, '0. asperus Warburton, O. tartakovskYi Olenev, and O. tholozani
(Laboulb(me and Megnin) (= O. papillipes Birula; in Europe, O.
verricosus Olenev Sassuchin and Fenik and O. erraticus Lucas (= O.
moracanus Velu).
Spirochetes of endemic relapsing fever. The causal organism of tick-
borne Central African relapsing fever was named Spirochaeta duttoni
by Navy and Knapp in 1906. The South and Central American strain
was called Spirochaeta venezuelense by Brumpt in 1921, and the Texas,
U.S.A., strain was called Spirochaeta turicatae by Brumpt in 1933.
Other strains, such as S. kochi Novy, 1907, S. hispanica Sadi de Buen,
1926, S. persica Dschunkowsky, 1912, and S. neotropicalis Bates and St.
John, Central and South America, have been recognized.
FIG. 162. (Top) a Sierrian summer cabin in which relapsing fever was con-
tracted. (Bottom) interior view of the cottage.
That these are probably all strains or local varieties of one widely
distributed species, Borrelia (Spirochaeta) recurrentis Lebert, is believed
to be the case by various authors. Students should consult Publication
No. 18 (1942), American Association for the Advancement of Science,
entitled "A Symposium on Relapsing Fever in the Americas."
Porter, Beck, and Stevens (loc. cit., 1932) report finding spirochetes
in blood smears from a chipmunk shot near Polaris, the location of the
first endemic case reported by Briggs (1921), and also from a squirrel
killed at Packer Lake, another endemic focus . Beck (1937) described
three of the California foci (Big Bear Lake, Lake Tahoe, and Packer
Lake) and gives an account of the rodent population in each. The re-
sults of her laboratory inoculations of white mice indicate that of a total
of 183 inoculated, 13 were found to be infected with spirochetes re-
sembling Borrelia (= Spirochaeta) recurrentis. Herms and Wheeler (loc.
cit., 1936) proved this spirochete to be the causal organism of relapsing
fever in man. Only chipmunks and Tamarack squirrels were discovered
harboring this spirochete in these areas. Three varieties of chipmunks are
FIG. 163. Spirochetes of relapsing

fever in a blood smear.
listed as hosts, mainly Eutamias quadrimaculatus; the other principal host
of the spirochete is listed as the Tamarack squirrel or Sierra chickaree,
Sciurus douglassi albolimbatus. Beck reports 13 rodent strains and 6
human strains, those of the chipmunk being most virulent.
The spirochetes of relapsing fever in man and rodents in California
(Fig. 163) have been carefully studied by Miss M. Dorothy Beck,94 She
reports 13 strains of spirochetes resembling Borrelia (= Treponema)
recurrentis from rodents in the field. Only chipmunks (Eutamias sp.)
and Tamarack squirrels or Sierra Nevada chickarees (Sciurus douglasii
albolimbatus Allen) were found to harbor spirochetes in the districts
surveyed. She reports long periods of latency in mice, up to 114 days.
She also found that the spirochetes of both human and animal origin
show remarkable resistance to freezing and remain viable in defibrinated
sheep blood for at lease 195 days. She concludes that the rodent and
human strains are identical in morphology and that laboratory animals
show a similar susceptibility. "These strains are undoubtedly the same
since the rodent strains are directly transmissible to man."
The spirochetes show a high degree of tick-host specificity. Based on
experimental evidence, Davis 95 proposes the name Borrelia (Spirochaeta)
hermsi for the infection transmitted by Ornithodoros hermsi and Borrelia
parkeri for the infection transmitted by O. parkeri, a species which re-
sembles O. hermsi very closely and an experimental vector of spirocheto-
sis. Although O. parkeri bites man freely and may be a vector of relaps-
ing fever to man, though experimental evidence to that effect is lacking,
its elevational range is below the endemic areas of relapsing fever, and
its wild host relationships seem to differ. Nevertheless one human case
is attributed to O. parkeri by Davis, Wynns, and Beck96 (1941) at an
elevation of about 250 feet, far below the usual elevation of known
endemic areas.
Eradication of the infection from summer cabins (Fig. 162) known to
harbor the infection because of the origin of cases in the cabins is ren-
dered somewhat difficult because the manner of their construction per-
mits chipmunks and smaller rodents to gain entrance and establish nests
inside or between walls. By driving these animals out and keeping them
out by securely closing all crevices, knotholes, etc., to prevent their
return is good practice; however, it must be remembered that this de-
prives the ticks of their natural food, and they will turn avidly to the
human occupants for blood. These ticks are also extremely "hardy" and
may go without food for many months and continue to harbor the causal
organisms (spirochetes) of relapsing fever so that a long vacancy does
not ensure freedom from ticks; they do not starve easily.
Some of the measures useful in the control of ticks and of chipmunks
and other small animals infesting summer cottages are the following:
( 1) When a new cottage is being built, it should be rodent-proofed-
this is a simple and inexpensive procedure known to all carpenters in
such areas; (2) all crevices, knotholes, and other points of ingress
should be closed; all doors should be tight fitting, and the chimney
should be covered with wire netting to keep out bats and rodents; (3)
yards, woodpiles, etc., should be cleaned up and kept in order to prevent
chipmunks and rodents from concealing themselves and building their
nests; (4) rodent reproduction should not be encouraged by feeding and
making pets of these animals; the rodent population should be kept at
a low level; (5) old tree snags in the vicinity which may harbor nesting
squirrels and chipmunks should be destroyed; nests in such snags have
been found to harbor numerous relapsing fever ticks in all stages of
development; (6) firewood covered with loose bark beneath which ticks
may be concealed should not be brought into the house or woodshed;
(7) although only the younger stages of the ticks are certainly affected,
it is of value to apply a 10 per cent DDT dust to all surfaces inside the
cabin making sure that there is a visible layer of the dust throughout;
TICKS AND TICK-BORNE l)ISEASES 513
treatments early in the summer and again in the autumn are recom-
mended; (8) a 5 per cent solution of residual DDT spray should be
applied thoroughly to beds, mattresses, rugs, under rugs, on walls and
floors, under tables and chairs, in closets, porches, cracks in the walls
and along baseboards, etc. (food and dishes should be protected from
the DDT powder and spray); (9) in an area-wide program of relapsing
fever control, the elimination of breeding niches for chipmunks (and
other rodents) by complete destruction by fire (or other means) of
"snags," partly dead or dead trees is strongly urged-this is an important
aspect of forest sanitation in recreation areas.
Genus Otobius. In adults the integument is granulated; nymphs have
spines. The sexes are similar. The capitulum is distant from the anterior
margin in adults and near the margin in nymphs. The hood and eyes
are absent; the hypostome is vestigial in adults but well developed in
nymphs. Cooley and Kohls recognize two species O. mcgnini (Duges)
FIG. 164. Spinose ear tick, Otobius

megnini. X 3.5.
and O. lagophilus Cooley and Kohls,'" the latter a parasite on the face
of rabbits in western United States and Canada.
The spinose ear tick, Otobius megnini (Duges), is a widely distrib-
uted species, being found in warmer parts of the United States, South
America, South Africa, India, and no doubt other parts of the world. It
receives its name from the fact that the nymph is covered with spines
(Fig. 164), and in all stages the tick invades the ears of cattle, horses,
mules, sheep, cats, dogs, other domesticated animals, and man, as well
as deer, coyotes, rabbits, and other wild animals. Rather large dark eggs
are depOSited by this species on the ground; under laboratory conditions
at a temperature of about 21 0 C the incubation period is from 18 to 23
days. In the field newly emerged larvae crawl up weeds and other vege-
tation like the larvae of other ticks, coming in contact with suitable host
animals such as cattle and gradually working their way to the shoulders,
neck, and head, and thence to the deeper inner folds of the outer ear of
the host, where they assume a peculiar sac-like form. After molting in
the ear the nymphs attach themselves and remain attached for long
periods of time; this, the second nymphal stage, is the stage in which this
species is most easily distinguished from other ticks. Individual ticks
may remain in the ear as long as 121 days, as observed in our tests. On
detaching they crawl out of the ear, drop to the ground and molt again
(there are three molts), after which maturity is reached. Copulation
takes place within a day or two after the final molt, and oviposition
occurs in from 14 to 42 days, with a maximum oviposition period of 155
days in the individuals observed, during which time 562 eggs were laid.
FIG. 165. The poultry tick, Argas

persicus, dorsal view.
The longevity of unfed larvae at room temperature ranged from 19 to

63 days, with an average of 44 days.98
Damage done. The writer has received many complaints from various
cattle-grazing districts in California relative to the "ear tick." Ears are
occasionally sent in thoroughly infested with these pests in all stages. It
is commonly asserted that this tick is responsible for much deafness in
domesticated animals, and it is also believed to be responsible for illness
and even death, particularly in calves.
Treatment. Owing to the position occupied by the ticks on the host,
only local treatment is of any avail. A mixture of two parts of pine tar
to one part of cottonseed oil, injected with a warm metal syringe about
112 ounce in each ear (cattle and horses), is recommended by Imes. 99
The ear should be manipulated during the injection. This mixture kills
the ticks and affords protection against the invasion of others for about
thirty days. Another mixture for similar treatment is as follows : benzene
hexachloride (15 per cent gamma isomer) 5 per cent, xylol 10 per cent,
pure pine oil, q.s. ad. 100 per cent.
Genus Argas. The members of this genus are distinctly flattened, with
the margins quite thin even when the tick is fully engorged. The integu-
ment is leathery, minutely wrinkled in folds, often intermingled with
small, rounded "buttons," each with a pit on top and often bearing a
hair in the pit. Eyes are absent; sexes are similar. There are only two
species in North America.
Argas persicus (Oken) (Argas miniatus Koch, Argas americanus
Packard), a cosmopolitan fowl tick, is one of the most important poultry
parasites in existence (Fig. 165). In addition to "fowl tick," this pest is
commonly called "adobe tick," "tampan," or "blue bug." In color it
varies from a light reddish brown to a dark brown, depending on the
stage of engorgement. In size the obovate, flattened adults average
about 8.5 mm long by 5.5 mm wide in the female, and 6.5 mm long by
4.5 mm wide in the male. When unengorged their thickness is about .75
mm and when fully engorged may be nearly 3 mm at the thickest part.
The marginal border is always distinct even when the tick is engorged.
The sexes are not easily distinguishable; the males are smaller, and
though they may be as large as smaller female individuals, they taper
slightly more anteriorly, i.e., are more obovate. The genital orifice of
the male is "half-moon-shaped," while in the female it is "slit-like" and
situated farther forward, i.e., immediately behind the capitulum as in
other argasines. The capitulum has four long hairs, two hypostomal,
and one near the articulation of each palp, all directed forward. The
palpi are about twice as long as the hypostome, second article longest,
the others equal in length. The hypostome, apically rounded, has six or
seven fine denticles on each half distally, followed by stout teeth 2/2,
the numbers increasing to 3/3, 4/4, 5/5, basally, the teeth decreasing in
size, neither attaining the external border nor extending beyond half the
length of the hypostome (Nuttall).
Life history and habits. The nymphs and adults of Argas persicus
(Oken) are strikingly active at night, migrating long distances to find
their host, and hiding in an inactive condition during the day. The
writer has observed this pest in vast numbers hiding beneath the loose
barR: of the eucalyptus tree in California. Occasionally specimens are
sent in with the inquiry, "Are they parasites of the tree or do they attack
roosting chickens? The chickens seem to do very poorly, yet we find
nothing on them." At night if one observes somewhat closely, one may
see hordes of these ticks climbing up the sides of the chicken coop to
the roosts and upon the fowls, filling up leisurely with blood and before
daybreak departing for their hiding places. The females deposit their
large reddish-brown eggs in the crevices occupied during the day. The
eggs are laid in masses of from 25 to 100 and over, and there are usually
several layings, each preceded by a meal of blood, with a total of seven
hundred eggs per female. Egg deposition takes place in almost any sort
of receptacle in which the ticks may be kept for observation. Hatching
takes place in from 10 days to 3 or 4 weeks. The larvae (Fig. 166) are
hexapod and very active, attacking a host ilpparently as readily by day
as by night. Once attached the larvae feed for about five days, occasion-
ally longer, remaining firmly attached during this time. When fully
engorged they appear like little reddish globules, causing severe irrita-
tion. At the end of this feeding period the larvae detach themselves, hav-
ing become rather flattened in the meantime, and then crawl away from
the host, hiding in some convenient crevice near by. The larvae molt in
about a week, when the fourth pair of legs appear and they are now in
FIG. 166, Larva of the poultry tick,

Argas persicus, X 30.
the first nymphal stage, looking like miniature adults, Nocturnal feed-
ing now takes place, and in 10 or 12 days another molt occurs and the
second nymphal stage is reached. Again the tick attaches itself, being
now able to engorge itself in about an hour; again, after the expiration
of something over a week, a third molt takes place (there may even be
a fourth molt) and then the adult stage is reached. The adults are able
to engorge themselves in from 20 to 45 minutes. Under favorable condi-
tions the adult stage is reached in about SO days. Absence of hosts to
feed upon may greatly prolong the life history.
Since eggs are deposited mainly during July in California, the adult
stage mayor may not be reached before the rainy season begins, and
the overwintering stage may be in the second nymphal condition or as
adults , appearing in pestiferous numbers early during the following sum-
mer. Hence there is ordinarily one generation of ticks per year under
normal conditions. In the absence of a host this species can live more
than two years without food.
The species will bite man. Instances are recorded in which transient
lahorers occupying long-vacated but renovated poultry houses have been
badly bitten by the poultry tick. It might perhaps under certain circum-
stances become involved in the transmission of human spirochetosis.
Damage done. Each tick when engorging requires considerable
blood to become replete, hence, when myriads of these parasites attack
fowls great quantities of blood must be extracted. The writer has known
of chickens being picked up under the roost in the morning with no
apparent cause for death, and believes this to have been due directly to
the work of ticks. Weakened and unthrifty condition of a flock may be
traceable solely to ticks. Poultry suffering from ticks have dull, ragged
plumage, suffer from diarrhea, are weak, and lay poorly. Turkeys are
most likely to suffer.
A vian spirochetosis. A very dangerous disease, known as "fowl spiro-
chetosis," is traceable to Borrelia (Spirochaeta) gallinarum Blanchard
(Spirochaeta marchouxi Nuttall), occurring in India, Australia, Brazil,
Egypt, and Persia, and is no doubt very Widely distributed. The disease
attacks chickens, geese, turkeys, guinea fowls, and other birds. The
symptoms are described as follows:
The disease begins with diarrhea, followed by loss of appetite, the birds
appearing somnolent; the feathers being ruIRed and the eomb pale. The birds
cease to perch, lie down with the head resting upon the ground and death takes
place during a convulsive attack. At times the disease runs a slower course, the
legs become paralyzed, then the wings, and the bird grows thin and dies in
eight to fifteen days. Recovery may take place, but it is rare after paralytic
symptoms have appeared. At autopsy, during the acute period of the disease,
the spleen appears much enlarged and the liver swollen with more or less fatty
degeneration; at times the liver is dotted with focal necroses. In chronic cases
both these organs may appear atrophied. The blood is fluid and dark. Spiro-
chaetes are plentiful in the blood until shortly before death, and they disappear
as recovery sets in (Nuttall).
Argas persicus (Oken) was proved to be a vector of this spirochete

infection by Marchoux and Salimbeni, Balfour, Nuttall, and others.
These investigators have found that when the tick sucks blood from an
infected fowl the spirochetes multiply within the body of the tick
when it is kept at from 30 0 to 35 0 C, and it is capable of transmitting
the disease; but when it is kept at from 15 0 to 20 0 C it fails to do so.
However, if the tick is later kept at the higher temperature it becomes
infective. The spirochetes are transmitted by the bite and by fecal con-
tamination; the tick is said to be infective for six months or more. The
infection is carried over from one generation of ticks to the next through
the egg. The incubation period in the fowl is from four to ni\le days.
Recovery from the disease is followed by immunity.
Seddon1oo reported several mild cases of avian spirochetosis in Aus-
tralia (N.S.W.) in 1926. He states that the mildness of the attack may
be attributed to the fact that the cases had occurred during the winter
when the cold weather would inhibit the activity of the ticks. The disease
is now commonly found in the southern parts of New South Wales and
in several other parts of Australia.
Combating the fowl tick. Henhouse roosts should be painted thor-
oughly with kerosene or a solution of creosote and put in position with
the ends in cups of crude oil or creosote or embedded in oil-soaked waste,
or suspended by wires from the ceiling. Roost poles must be free from
bark. All old nests and rubbish should be burned, and the interior,
especially crevices, sprayed liberally with kerosene. Boiling water or
steam may be used instead of kerosene emulsion. A repetition of the
procedure once every five or six weeks during the tick season is recom-
mended. The use of considerable crude oil or creosote oil in and about
the houses is very desirable. Fowls should not be permitted to roost in
trees, because of the hiding places afforded the ticks beneath the bark,
particularly when this is loose.
If the henhouses can be made tight, fumigation with sulphur is use-
ful; about five pounds per 1,000 cubic feet of space should be employed.
For the treatment of fowls infested with larval ticks, an ointment of
kerosene, lard, and sulphur is advised.
Argas reflexus (Fabr.), commonly known as the "pigeon tick," differs
from A. persicus (Oken) in that the body narrows rather suddenly to-
ward the anterior end and that the thin margin is flexed upward. The
capitulum has "two long post-hypostomal hairs ventrally, directed for-
wards. Palps with articles sub-equal, the third the shortest, denticulated
hairs dorsally. . . . Hypostome apically notched, some small denticles
at the tip, followed by 2/2 stout teeth merging into 3/3 to 6/6 pro-
gressively smaller teeth" (Nuttall).
Other species of Argas are the following: A. brumpti Neumann, the
largest known species of the genus, measuring 15-20 mm in length by
10 mm in width. It is known to attack man in Africa. A. vespertilionis
(Latreille) occasionally attacks man in Africa. A. mianensis Brumpt
occurs in human habitations in Iran (Mianeh) where it is believed to be
a vector of a spirochete infection in man known as Mianeh fever.
Genus Antricola. The dorsal walls are flattened and marginated; be-
low the flattened dorsum the body is convex and deep; the integument
is semitranslucent and the surface smooth, shining, and with tubercles;
the mouth parts are adapted for quick feeding and not for clinging to
the host.
Antricola coprophilus (McIntosh) feeds on bats in Arizona and
Mexico.
Antricola marginatus (Banks) is found in bat caves in Cuba and
other parts Of the West Indies.
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and laboratory investigation," M. J. Australia, 24:281-99.
51. Cox, H. R., 1939. "A filter-passing infectious agent isolated from
tick. III. Description of organism and cultivation experiments," U. S. Public
52. Parker, R. R., and Davis, Cordon E., 1938. "A filter-passing infec-
tious agent isolated from ticks. II. Transmission by Dermacenter andersoni,"
53. Steinhaus, Edward A., 1946. Insect Microbiology. Ithaca: Comstock
Publ. Co., Inc. x + 763 pp.
54. Parker, R R., and Davis, Gordon E., 1943. "American Q fever: the
occurrence of Rickettsia diaporica in Amblyomma americanum in eastern
Texas," U. S. Public Health Service, Pub. Health Rep., 54:1510-11.
55. Philip, Cornelius B., 1948. "Observations on experimental Q fever,"
J. Parasitol., 34:457-64.
56. Huebner, R. J.; Jellison, W. L.; Beck, M. D.; Parker, R. R; and
Shepard, C. C.; 1948. "Q fever studies in southern California," U. S. Public
57. Shaughnessy, H. J., and Milzer, Albert, 1939. "Experimental infec-
tion of Dermacentor andersoni Stiles with the virus of lymphocytic chorio-
meningitis," Am. J. Pub. Health, 29:1103-8.
58. Henning, M. W., 1932. Animal diseases of South Africa. "South
African Agriculture Series," 11 :298-329.
59. Bmmpt, E., et Larrousse, F., 1922. "Transmission de la Piroplasmose
canine fran9aise par Ie Dermacentor venustus," Bull. Soc. path. exot., 15:540-
45.
60. ---, 1919. "Transmission de la piroplasmose canine tunisienne par
Ie Rhipicephalus sanguineus," Bull. Soc. path. exot., 12:757-64.
61. Sanders, D. A., 1937. "Observations on canine babesiasis (piroplas-
mosis)," ]. Am. Vet. M. A., 90, n.s. 43:27-40.
62. Henning, M. W., 1932. Animal Diseases of South Africa. "South
African Agricultural Series," 2:545.
63. Stiles, Geo. W., 1939. Anaplasmosis in Cattle. Washington, D. C.:
Dept. Agric., in Circ., no. 154 (revised). 10 pp.
64. Herms, W. B., and Howell, D. E., 1936. "The western dog tick,
Dermacentor occidentalis Neum., a vector of bovine anaplasmosis in Califor-
nia," J. Parasitol., 22:283-88.
65. Howell, D. E.; Stiles, Geo. W.; and Moe, Lewis H.; 1941. "The
hereditary transmission of anaplasmosis by Dermacentor andersoni Stiles,"
Am. J. Vet. Research, 11:165-66.
66. Boynton, W. H., and Woods, Gladys M., 1933. "Deer as carriers
of anaplasmosis," Science, 78:559-60.
67. Ellenberger, W. P., and Chapin, R. M., 1940. Cattle-fever Ticks
and Methods of Eradication. Washington, D. C.: Dept. Agric., in Farmers'
Bull., no. 1057. 27 pp.
68. Bishopp, F. C., and Smith, C. N., 1938. The American Dog Tick,
Eastern Carrier of Rocky Mountain Spotted Fever. Washington, D. C.: Dept.
Agric. in Circ., no. 478. 26 pp.
69. Smith, Carroll N., and Gouck, Harry K., 1945. "DDT to control
ticks on vegetation," ]. Econ. Entomol., 38:553-55.
70. Cooley, R A., and Kohls, Glen M., 1944. The Argasidae of North
America, Central America, and Cuba. Notre Dame:- The University Press,
in American Midland Naturalist. 152 pp.
71. Jobling, B., 1925. "A contribution to the biology of Ornithodoros
moubata Murray," Bull. Entomolog. Research, 15:271-79 (1 plate).
72. Davis, Gordon E., 1947. "A note on the larval stage of the argasid
tick, Ornithodoros moubata (Murray) 1877," J. Parasitol., 33:495-96.
73. Aitken, T. H. G., 1939. "Ornithodoros talaje on the California main-
land," Pan-Pacific Entomologist, 15:12-13.
74. Bates, L. B.; Dunn, L. H.; and St. John, J. H.; 1921. "Relapsing
fever in Panama," Am.]. Trop. Med., 1:183-210.
75. Matheson, R., 1931. "Note on the Ornithodoros talaje (Guer.-Men.),"
Parasitology, 23:270.
76. Dunn, L. H., 1937. "Studies on the South American tick, Ornitho-
doros venezuelensis Brumpt in Colombia," J. Parasitol., 13:249-59.
77. Hindle, E., 1935. "Relapsing fever: some recent advances," Trop.
Dis. Bull., 32:309-27.
78. Cooley, R. A., 1936. "Ornithodoros parkeri, a new species on rodents,"
79. Herms, W. B., and Wheeler, C. M., 1936. "Ornithodoros hermsi
Wheeler as a vector of relapsing fever in California," J. Parasitol., 22:276-
82. .
80. Ross, P. H., and Milne, A. D., 1904. "Tick fever," Brit. M. ]. 2:1453-
54.
81. Dutton, J. E., and Todd, J. L., 1905. The Nature of Human Tick
Fever in the Eastern Part of the Congo Free State, with Notes on the Distribu-
tion and Bionomics of the Tick. Liverpool: School Trop. Med. Memoir, no.
17. 18 pp.
82. Feng, Lan-Chou, and Chung, Huei-Lan, 1936. "Studies on the de-
velopment of Spirochaeta duttoni in Ornithodoros moubata," Chinese M. J.,
50:1185-90.
83. Meador, Charles N., 1915. "Five cases of relapsing fever originating
in Colorado, with positive blood findings in two," Colorado Med., 12:365-68.
84. Briggs, LeR. H., 1922. "Relapsing fever in California," J.A.M.A.,
79:941-44.
85. Wheeler, C. M., 1935. "A new species of tick which is a vector of
relapsing fever in California," Am. J. Trop. Med., 15:435-38.
86. Porter, G. S.: Beck, M. Dorothy; and Stevens, 1. M.; 1932. "Relapsing
fever in California," Am. J. Pub. Health, 22:1136-40.
87. Herms, W. B., and Wheeler, C. M., 1935. "Tick transmissions of
California relapsing fever," J. Econ. Entomol., 28:846-55.
88. Wheeler, C. M.; Herms, W. B.; and Meyer, K. F.; 1935. "A new
tick vector of relapsing fever in California," Proc. Soc. Exper. Biol. & Med.,
32:1290-92.
89. ---, 1938. "Relapsing fever in California-attempts to transmit
spirochaetes of relapsing fever to human subjects by means of the bite of the
vector, Ornithodoros hermsi Wheeler," Am. J. Trop. Med., 18:641-59.
90. Weller, B., and Graham, G. M., 1930. "Relapsing fever in central
Texas," J.A.M.A., 95:1834-35.
91. Wheeler, C. M., 1938. Progress of spirochaete infection in the de-
velopmental stages of the host tick, Ornithodoros hermsi Wheeler. Am. J. Trop.
Med., 18:413-19.
92. Davis, Gordon E., 1940. "Ticks and relapsing fever in the United
States," U. S. Public Health Service, Pub. Health Rep., 55:2347-51.
93. ---, 1941. "Ornithodoros parkeri Cooley: observations on the
biology of this tick," ]. Parasitol., 27 :425-33.
94. Beck, M. Dorothy, 1937. "California field and laboratory studies on
relapsing fever," J. Infect. Dis., 60:64-80.
95. Davis, Gordon K, 1942. "Species unity or plurality of the relapsing
fever spirochaetes," A Symposium on Relapsing Fever in the Americas. Wash-
ington, D. C.: Amer. Assoc. Advancement Sc., Publ. no. 18, pp. 41-47.
96. ---; Wynns, Harlin L.; and Beck, M. Dorothy; 1941. "Relapsing
fever: Ornithodoros parkeri a vector in California," U. S. Public Health Service,
97. Cooley, R. A., and Kohls, Glen M., 1940. "Two new species of
Argasidae (Acarina: Ixodoidea) ," U. S. Public Health Service, Pub. Health
Rep., 55:925-33.
98. Herms, W. B., 1917. "Contribution to the life history and habits
of Ornithodoros megnini," ]. Econ. Entomol., 10:407-11.
99. Imes, Marion, 1918. The Spinose Ear Tick. Washington, D. C.:
Dept. Agric., Bur. Animal Indust., in Farmers' Bull., no. 980. 8 pp.
100. Seddon, H. R., 1926. "A note on spirochaetosis in fowls," New South
Wales: Dept. of Agric., in Science Bull., no. 26, Veterinary Res. Rept. no. 2.
pp. 17-19.
CHAPTER XXI
MITES
CLASS ARACHNIDA, ORDER ACARINA

Characteristics. In the mites, as in the ticks, the unsegmented abdo-
men is broadly joined to the cephalothorax with little or no evidence
of separation. All but a few species are minute, i.e., just about visible to
the naked eye. Mites, like other arachnids, have four pairs of legs as
adults but only three pairs (exceptionally, fewer) in the larval stage.
The mouth parts are quite varied but follow the general pattern of the
ticks. The chelicerae in the parasitic species are piercing structures (see
Snodgrass). One ot more pairs of simple eyes are usually present. The
respiratory system is in most species similar to that of the ticks, i.e.,
tracheal, while others absorb oxygen through the general body surface,
which in these is quite soft. Nearly all species deposit eggs; however,
there are a few which are ovoviviparous, among them, Pediculoides
ventricosus (Newport). From the egg there emerges the hexapod larva,
which usually soon molts and then presents its fourth pair of legs. The
life history of many species requires less than four weeks; in some it is
as short as eight days.
An infestation of mites is termed acariasis. Those species which bur-
row into the skin, producing channels in which their eggs are depOSited
are said to cause sarcoptic acariasis, e.g., Sarcoptes scabiei var. hominis
(Hering) of human scabies; while those species which deposit their
eggs at the base of the hairs of the host or on the skin, producing scabs
are the cause of psoroptic acariasis, e.g., Psoroptes communis var. ovis
( Hering) of sheep scab. Other forms of acariasis are described in this
chapter.
Although Banks! recognized 29 families of mites in 1915, and Vitz-
thum 2 in 1943 recognized 80, actually only a few of these affect man
and his domestic animals. Students concerned with the body of mites
will need to consult the monumental work of Oudemans 3 (1937), also
the excellent publications of Ewing, particularly his Manual of External
Parasites. 4 Radford5 has published a very helpful list of "Genera and
species of parasitic mites," which gives the authority, year of validity,
and~ where pOSSible, the original host.
525
MANGE, OR ITCH MITES-SARCOPTIC ACARIASIS

Family Sarcoptidae
Characteristics. All members of the family Sarcoptidae, commonly
known as the itch mites, mange mites, or scab mites, are very small
(just about visible to the naked eye), whitish, and somewhat hemis-
pherical in form. Banks describes this family as follows:
The body is entire, and the surface transversely striated and provided
with a few bristles, often short, stout and sharp painted. The legs are short
and stout, arranged in two groups. The anterior legs are usually larger than
the others. The tarsi commonly terminate in a stout claw. There is generally
a long pedicellate sucker, sometimes with a jointed pedicel. The claw or sucker
may be absent and in its place' a long bristle. The legs often show a chitinous
framework of rings, both transverse and oblique. On the front of the body
is a prominent beak. The palpi are small, three-jointed, and appressed to the
sides of the beak beneath. . . . There are frequently sexual differences. Some
males have the third pair of legs very large and long, while the fourth pair
is very small. Sometimes there are plate-like lobes at the tip of the male ab-
domen, and the tarsi may terminate differently in the two sexes.
The family Sarcoptidae includes a number of important genera,
among them Sarcoptes, Psoroptes, Notoedres, Chorioptes, Otodectes and
Cnemidocoptes, each producing a particular type of acariasis.
Sarcoptic mites. The mange or itch mites belong to the genus Sar-
coptes; they have very short legs, the posterior pair not extending be-
yond the margin of the nearly circular body; suckers are present on the
first and second pair of legs. The sarcoptic mites burrow in the skin,
where they produce definite burrows in which the females deposit their
eggs.
The species of Sarcoptes inhabiting the skin of mammals are ordi-
narily regarded as varieties of Sarcoptes scabiei (Linn.) (Fig. 167). They
differ very slightly, and many of them may interchange hosts, e.g., Sar-
coptes scabiei var. suis Gerlach, is parasitic on swine and may be tem-
porarily parasitic on man; Sarcoptes scabiei var. equi Gerlach of the
horse is also temporarily parasitic on man. The species appropriate to
a given host, however, ordinarily exist only for a limited time on a
different host species.
Human scabies or itch. The. itch mite attacking man is known as
Sarcoptes scabiei var. hominis (Hering). The infection is known as
"scabies," "seven-year itch," or "Norwegian itch." It is universal in dis-
tribution. The female mite measures 330JJ. to 450JJ. in length and 250JJ.
to 350JJ. in breadth; the male is slightly more than half as large. The
mite attacks by preference the thin skin between the fingers, the bend
of the knee and elbow, the penis, the breasts, and the shoulder blades,
MITES 527
although any part of the body is subject to attack. An almost intolerable
itching results due to toxic secretions and excretions. The sinuous bur-
rows which the mite makes in the epidermis may reach 3 cm in length
(Warburton6 ), and tiny vesicles and papules are formed on the surface.
Scratching causes weeping and bleeding, which favor spread of the
mites. It is also a means of secondary infection. Transmission is due to
intimate personal contact such as hand shaking or sleeping with an in-
fested person; clothing, towels, bedding, etc., are much less involved.
Infection is usually effected by the adult fertilized female. 7 By tracing
a mite burrow to its end with a hand lens, the adult female mite can
usually be removed with a needle or scalpel and after treatment with
Fic. 167. Sarcoptes scabiei.
KOH on a glass slide can be readily seen under a microscope. Long-

standing chronic cases show few parasites.
Life history of itch mite. The female mites deposit their rather large
oval eggs (150JL X 100JL) at intervals in the tortuous tunnels which they
have made in the epidermis. From 10 to 25 eggs are deposited by each
individual during a period of about two weeks. The female, having de-
posited her complement of eggs, dies at the end of her burrow. The
hexapod larvae hatch in three or four days. In this stage the area of in-
fection is most rapidly increased. Maturity is reached in 10 to 12 days,
during which time there are three molts.
Treatment for human itch. Inasmuch as the mites are protected in
their tunnels in the epidermis, the skin must be thoroughly softened
by scrubbing vigorously with green soap and hot water before a remedy
is applied. Sulphur ointments (10 to 15 per cent precipitated sulphur)
give very good results if overnight applications are repeated at intervals
of three or four days. Benzyl benzoate ointment is also highly recom-
mended. Underclothing coming in contact with the parts affected should
be boiled, steam sterilized, or baked.
Swine mange. Mange of swine is caused by Sarcoptes scabiei var.
suis Gerlach (Fig. 168). Mange attacks swine commonly about the top
of the neck, shoulders, ears, and withers, and along the back to the root
of the tail. A microscopical examination of deeper tissue from beneath
scabs will usually reveal the mites. Suckling pigs and young shoats suffer
most. The affected animals scratch and rub vigorously, which may, how-
ever, be due to lice, but if the skin is cracked and thickly encrusted with
heavy scabs, and the hair stands erect, an examination for scab mites
FIG. 168. Life history and general char-

acteristics of a typical sarcoptic (mange or
itch) mite: (lower right) egg, (lower left)
larva, (upper left) male, ( upper right)
female. Sarcoptes scabiei var. suis, the itch or
mange mite of swine. x57.
should be made. The life history and habits of the swine mange mite
correspond in every respect with those of the itch mite of humans.
Treatment for swine mange. In the treatment of swine mange it is
necessary to apply external remedies, in addition to sanitary precautions
to prevent spread and the reinfection of treated animals. Remedies are
best applied in the form of solutions such as "lime and sulfur" dip be-
cause all parts of the animal's body are easily reached in the dipping
process. Hand dressing, scrubbing, and application of ointments may be
practiced where dipping is not practical; however, all parts of the animal
should be thoroughly treated. A suitable dip may be made as follows:
8 pounds of fresh lime and 24 pounds of Howers of sulphur to 100 gallons
of water, slaking the lime with sufficient water to form a thick paste,
sifting in the sulphur, and mixing with a hoe. This mixture is placed in
a kettle with 25 to 30 gallons of water and boiled for one hour at least,
two hours being better. The entire mass is used for swine but must not
be used for sheep. The dip is used warm at a temperature of from 100°
to HO° F.B
Lime and sulphur dips may be purchased ready made and are com-
monly used at the rate of one part of the solution to 15 parts of water;
however, care should be exercised to use the dip as directed, owing to
MITES 529
variation in constituents. Coal-tar dips are also used extensively and are
effective if given properly.
Dipping vats may be made of wood or concrete and are usually set
in the ground at a slight elevation to ensure drainage away from the vat.
A convenient size for a vat is "ten feet long on top, eight feet long on
the bottom, one foot wide on the bottom and two feet wide at the top.
The end where the hogs enter should be perpendicular and the other
end inclined, with cleats, so that the hogs can emerge after passing
through. The entrance should be by a slide. For pigs and small shoats
that can be easily handled, a barrel serves the purpose well; the pigs can
be caught, plunged in the dip and held there the required time. Some
successful swine raisers build cement bathing places or wallows for
swine and keep these filled with a watery solution of some dip or disin-
fecting solution." The time to treat young pigs, and this is important, is
at weaning time. Dipping twice as for older animals is necessary, and
if placed in uninfected quarters they ought to remain clean.
Mangy swine should be hand dressed with a stiff brush before
dipping in order to loosen up scabs and then kept in the dip long
enough to permit the solution to soak through the scabs, certainly not
less than two minutes. All the animals must be dipped a second time in
eight or ten days, in order to destroy the mites which have hatched from
the eggs which are not destroyed.
May0 8 recommends a disinfecting whitewash to be applied to pens,
etc., "Fresh lime, 25 pounds, flowers of sulphur, 15 pounds; mix the
sulphur with a little water, to a paste, add SO gallons of water and cook
for an hour, then add water sufficient to make 50 gallons and apply with
a spray pump, using a 'Bordeaux' nozzle."
Equine mange. Sarcoptic acariasis in horses, mules, and asses is
caused by Sarcoptes scabiei var. equi Gerlach. This species is also trans-
missible to man and is said to be the chief cause of a transitory itch of
cavalrymen and others handling horses extensively.
The most reliable diagnostic character is the discovery of the mite,
as already explained. The usual symptoms are first of all a strong tend-
ency to rub some circumscribed part such as the head, root of the
mane or tail, withers, or back, due to an incessant itching. If a person
scratches the affected parts, the animal moves its lips as though it were
nibbling. The skin of these parts also shows an eruption of "fine conical
papillae." The hair stands erect and bristly, much having dropped out,
but totally bare spots where there are no isolated hairs apparently do
not occur in mange but do in ringworm. The affected parts are at first
scurfy, then become covered with yellOWish scabs, which later exude
matter due to the rubbing and inflammation, and finally there are formed
scabs and crusts, often with deep crevices. During the first 14 days the
progress of the disease is usually slow, but by the sixth week the ravages
of the disease become extensive and there is rapid progress.
The life history and habits of Sarcoptes scabiei var. equi Gerlach
correspond in every respect with those of the species already described.
Treatment for equine mange. Before applying a remedy for mange
the animal should be clipped if possible so as to disclose all points of
attack which might otherwise be hidden by hair. The clipped hair must
not be blown away by the wind but should be burned. The parts affected
are next thoroughly lathered and left for a short while to soften, after
which warm water is applied and the scabs rubbed off as far as possible
with wisps of hay or straw which should also be burned. The affected
parts are now ready for a parasiticide, which should be applied by
hand.
Various remedies may be obtained for mange, all of which have more
or less value, but the writer has found that those containing sulphur are
the most effective. Cameron,9 Veterinary Director General, Health of
Animals Division, Department of Agriculture, Dominion of Canada,
prescribes for hand treatment of horses (and cattle as well, but not
sheep) a mixture of sulphur, 2 pounds, oil of tar, 8 ounces, and raw
linseed oil, 1 gallon. These ingredients are gradually heated together
but not boiled. This dressing should be very thoroughly applied over the
whole surface of the body and extremities of the animal, including the
ears and tail, at a temperature not less than llOO F and not more than
1200 F. Two dressings are necessary. Forty-eight hours before the second
application (twelfth to fourteenth day inclusive) the first dressing should
be washed off with hot water and castile soap. Ten days after the second
application the animals should again be washed to remove the dressing.
In severe cases it may be necessary to repeat the treatment many times.
For dipping animals Cameron (lac. cit.) uses a mixture of flowers of
sulphur, 24 pounds; fresh un slaked lime, 10 pounds; and water, 100 gal-
lons. The lime should be carefully slaked and made into a paste, and
the sulphur added and thoroughly incorporated with the paste; the
whole should be added to sufficient water, preferably boiling water, and
thoroughly boiled for at least two hours and while boiling must be
frequently stirred. The mixture should then be a rich chocolate-brown
color. It should be allowed to stand for a few hours or overnight, then
the liquid should be drained off without disturbing the sediment and
sufficient water should be added to make 100 gallons. The dipping vat
(cage or swimming) is filled with the solution in the proportions men-
tioned and heated to not less than llOo F or more than 115 0 F, main-
taining this temperature while the animals are held in the vat for at
least two minutes, during which period all crusts and scabs must be
loosened with a stiff brush. A second dipping must be undertaken be-
MITES 531
tween the twelfth and fourteenth days after the first dip in order to
destroy the new generation of mites (eggs ar.e not killed by the dip)
before they mature.
Stalls in which mangy animals have been :kept must be disinfected
with live steam or boiling water or be treated with an acaricide such as
lime and sulfur. Brushes, scrapers, "rubbers:! etc., should be boiled;
harnesses should be rubbed thoroughly with a strong disinfectant.
Bovine mange. Sarcoptic acariasis, or maDge, in cattle caused by
Sarcoptes scabiei var. bovis Robin is not so (:ommon as the psoroptic
form (scabies) but is far more difficult to cure. It usually attacks the
parts of the body where the hair is short, namely the brisket and around
the base of the tail. Treatment is the same as for horses described above.
Canine mange. The common mange of do~S is caused by Sarcoptes
scabiei var. canis Gerlach, closely resembling t:he swine parasite. Mange
in the dog appears first on the muzzle, arour1d the eyes, on ears and
b,=-~~"'t, ~Rd. l~t~,=- "''t>,=-~~d.", t~ t"h.~ b~~l,., ~bd.~W>.~-, lLlJn.. pj_cm\¥.bf'.J..~. S:'}TJJ1}-
toms are much as in swine and horses.
Infected long-haired dogs should be clipped before treating. The
following treatment is recommended:
The whole skin may be covered with a soluti.on of equal parts of green
potash soap and alcohol and just enough carboliC acid to give it the odor.
This is washed off next day and the surface is covered with the following:
Napthalin, 12 oz.; Vaseline," 212 oz.; oil of lavender, 8 drops. This may be
applied daily until a cure is established. Another very effective dressing and
*
equally safe is sulphur, 1 oz.; carbonate of potash, oz.; lard, 4 oz. For house
dogs balsam of Peru or styrax, ~ oz. to Vaseline," 2 oz., or alcohol, 1 pint,
makes a most agreeable, if somewhat expensive dressing, which though slow
is effective. (Law'·)
Notoedric mange. Mange of cats is caused by Notoedres minor var.
cati (Hering) (Sarcoptes minor var. felis Gerlach), smaller and more
circular than Sarcoptes but otherwise quite siJ1lilar. Notoedric mange of
cats begins at the tips of the ears and gradUi-llly spreads over the face
and head. After clipping the hair in the region of the affected parts,
apply ointments only, such as sulphur two parts, potassium carbonate
one part, and lard eight parts. N. minor var. cuniculi Gerlach causes a
severe mange of rabbits, beginning at the muztle and spreading over th.e
whole body.
N. muris (Meigen) occurs on rats, and recently, 1934, Skidmore l1
describes an acariasis of white rats caused by a mite for which he pro-
poses the name M yobia ratti (Family Cheyletidae).
Other mange mites. Sarcoptes scabiei var. ovis Megnin of sheep oc-
curs on the face primarily and causes "black nlUzzle." Repeated dipping
for several weeks in lime-sulphur is recommended. S. scabiei var.
" "Vaseline" Petroleum Jelly.
auckeniae Railliet occurs on llamas, and S. scabiei var. caprae Fursten-
berg on goats.
Scaly leg mite on poultry. The legs of domestic fowls (chickens, tur-
keys, pheasants, etc.) are frequently attacked by a microscopic burrow-
ing mite, Cnemidocoptes mutans (Robin and Lanquentin) (Fig. 169),
which causes a lifting of the scales and a swollen condition of the shank
with deformity and encrustation. The mites burrow and live in the skin,
depositing their eggs in channels as do the mange mites. Scaly leg (Fig.
170) is easily transmitted from fowl to fowl; hence the infested birds
should be segregated and treated.
A B
FIG. 169. (A) photograph of a portion of scale from a fowl affected with
"scaly leg," sllOwing mites (Cnemidocoptes mutans) in situ; (B) enlargement
of an individual mite. X 170.
In treating scaly leg the legs should be soaked and manipulated with
the hands in soap and warm water in order to soften the scabs. Then
when dry, "apply a coating of balsam of Peru or an ointment, containing
2 per cent of carbolic acid," or "a mixture of one part of oil of caraway
with five parts of Vaselineo" (United States Department of Agriculture,
Farmers' Bulletin 530). A simple treatment consists in dipping the legs
of the fowl in a mixture of equal parts of kerosene and raw linseed oil;
the oil must not touch the feathers on the legs; otherwise the skin may
suffer. This dipping process is best done while the birds are roosting,
lifting each bird from the roost, dipping, and then replacing it. The
process should be repeated in about a week.
" "Vaseline" Petroleum Jelly.
MITES 533
Depluming mite. Cnemidocoptes laevis var. gallinae (Railliet), known

as the depluming mite, is closely related to the "scaly leg mite," but
attacks the skin of the fowl near the base of the feathers. The mites
themselves do not cause the bird to lose its plumage, but the intense
itching caused by the mites impels the host to pluck its feathers in an
atte:llpt to reduce the itching.
A B
FIG. 170. (A) normal leg and claw of a fowl; and (B) one affected with
sarcoptic mites, Cnemidocoptes mutaM, causing scaly leg.
No satisfactory treatment is available, but some benefit may be ob-
tained by repeatedly rubbing the skin with a sulphur ointment made by
mixing three parts of flowers of sulphur with one part of lard.
FOOT AND TAIL MANGE-CHORIOPTIC ACARIASIS
Chorioptic or symbiotic scabies. Mites belonging to the genus
Chorioptes produce a mange which is restricted to certain parts of the
body such as the feet, tail, and neck. Chorioptes symbiotes Verhey en
[= C. equi (Gerlach)] attacks the lower parts of the legs of horses,
particularly those with long hairs on the fetlocks. The infection is known
as "aphis foot" in Australia. 12 A mixture of one part carbolic acid to 15
to 20 parts of linseed oil, or equal parts of kerosene and linseed oil, is
recommended as treatment. Several applications are needed to eHect a
cure.
Tail mange in cattle caused by Chorioptes bovis (Gerlach) is
localized on the tail or legs and is uncommon. Foot mange in sheep is
caused by Chorioptes ovis (Railliet).
SCAB MITES-PSOROPTIC ACARIASIS
Characteristics of psoroptic mites. The psoroptic or scab mites belong
to the family Sarcoptidae as do the itch and mange mites, hence par-
take of the family characteristics. However, in the psoroptic mites the
legs are long and slender, all four pairs extending beyond the margin of
the body, which is elongate. The "pedicel of the suckers is jointed" and
the "mandibles styliform, serrate near tip" and suited for piercing. The
FIG. 171. Psoroptes communis.
psoroptic mites do not burrow, as do the sarcoptic mites but live at the
base of the hairs of the host, piercing the skin and introducing a toxic
saliva which causes inflammation. An exudate follows which partially
hardens, forming a scab. As the mites multiply bites increase as well as
itching; more serum oozes out to form a crust of loose humid matter.
The parasitized area increases, and the skin becomes hardened and
thickened. This condition is known as scabies or scab. The eggs are
deposited among the piled-up scabs. Owing to the loose condition of the
scabs and the hardihood of the mites, this form of acariasis becomes
quickly and easily distributed from animal to animal by contact and
by rubbing against fences, trees, and the like.
The commonest scab mites belong to the genus Psoroptes (Fig. 171)
of which Psoroptes communis var. ovis (Hering) of sheep is best known.
Other varieties of this species infest cattle and horses.
Ovine scabies (sheep scab). Psoroptes communis var. ovis (Hering)
MITES 535
is the causal organism of scabies in sheep. This is by far the most impor-
tant species of scab mites. However, with the widespread use of dips
and rigid quarantine regulations, scabies in sheep is gradually being
controlled.
The sheep scab mite is easily visible to the naked eye. The adult
female measures about "one-fortieth" of an inch in length by "one-six-
tieth" of an inch in breadth, and the male "one-fiftieth" by "one-eightieth"
of an inch. As in all psoroptic species the mites are found on the surface
j
!
I,
FIG. 172. Showing life history and general characteristics of a typical

psoroptic or scab mite. (Lower left) egg, (lower right) larva, (upper left)
female, (upper right) male. Psoroptes communis. X 85.
of the body among the scabs at the base of the hairs. The parts of the
body most thickly covered with wool are chiefly affected. Scabies13 is
indicated by a "tagging" of the wool; the coat becomes rough, ragged,
and matted at the points affected. Tags of wool are torn away by the
sheep or are left attached to rubbing posts and other objects against
which the animal rubs. The sheep scratches vigorously and shows signs
of intense itching.
Life history of the scab mite. The female scab mite deposits an aver-
age of about 15 (maximum 30) eggs, one at a time, and the period of
oviposition often lasts several days, after which the female evidently
dies. The eggs (Fig. 172) are either attached to the wool near the skin
or deposited directly upon the latter. The hexapod larvae hatch in from
two to three days when next to the skin but take longer (up to six or
eight days) when on wool not close to the body; the first molt takes
place in three or four days, when the fourth pair of legs appears;
second and third molts take place within the next four or five days. The
males are said to molt but twice, and the f(~male is fertilized after the
second molt with a third molt before egg de}?osition. The length of time
elapsing from egg to egg averages about rline days. The infection is
spread by contact with infected sheep.
Treatment for sheep scab. Several kinds of dips with variations are
used for sheep scab, among them lime and sulphur, nicotine and sulphur,
nicotine cresol, coal-tar products, sodium silicofluoride, and derris. If
proprietary dips are used, extreme care must be exercised in following
the directions. The dip should have the aperoval of the United States
Department of Agriculture. All dips must be repeated in ten to twelve
days in order to destroy the mites newly hatched from eggs, which are
not killed by the dip. The dips commonly used and permitted by the
United States Bureau of Animal Industry for official dippings of sheep
for scabies are lime-and-sulphur and nicoti(le solutions. See Imes 13 for
directions for dipping sheep.
Psoroptic mange of horses is caused by psoroptes communis var. equi
Gerlach. It generally makes its appearance at or near the root of the
mane or tail. It is regarded as a serious disease. Treatment is the same
as for sarcoptic mange.
Bovine scabies (Psoroptic mange). Scabies in cattle is caused by
Psoroptes communis var. bovis (Gerlach) and is a comparatively com-
mon infection. The disease usually appears at the root of the tail, thighs,
neck, and withers and spreads rapidly to other parts of the body. Heavy
losses may result from this disease. '
Treatment for scabies in cattle is the same as for horses, namely lime
and sulphur as previously described.
AURI CULAR MITES-OTOACARIASIS
Auricular mites. A comparatively commo(l affection of cats, dogs, and
faxes, known as otoacariasis or parasitic otitis, is traceable to Otodectes
cynotis (Hering), which resembles Psoroptes very closely. These mites
belong to the family Sarcoptidae. They literally swarm in the ears of the
host causing much discomfort, tenderness of the ears, auricular catarrh,
loss of appetite, wasting, torticollis, "fits," etc.
Cleansing the ears first with soapsuds and warm water and then ap-
plying a sulphur ointment or a 10 per cent solution of tincture of iodine
in glycerin, or a one per cent solution of c~rbolic acid in linseed oil is
recommended. Banks recommends injecting olive oil containing one-
MITES 537
tenth part of naphthol. The hutches or kennels must be thoroughly dis-
infected with a strong lime-and-sulphur solution or other effective solu-
tion to prevent further contagion.
FOLLICLE MITES-FOLLICULAR MANGE
Family Demodicidae
Characteristics of follicle mites. The Demodicidae comprise the follicle
mites, very minute (.3 to .4 mm) mites with an elongated (vermiform)
transversely striated abdomen and four pairs of "stubby" three-jointed
legs. They parasitize the skin of mammals. All stages of the life history
are often present together in a follicle.
The follicle mite of man (Demodex folliculorum Simon) (Fig. 173)
inhabits the hair follicles and sebaceous glands "causing inflammation of
FIG. 173. A follicle mite. Demodex folliculorum. X 110.
the gland (comedones); their agglomeration in the meibomian glands

(in Il,lan) sets up inflammation of the margins of the eyelids" (Braun).
While the follicle mites may, under certain conditions, produce acne-
like conditions, it is hardly probable that many cases of "blackhead" if
any, may be traceable to these mites. They are nevertheless very com-
mon-said to occur in 50 per cent of mankind in all parts of the world.
In most animals the follicle mites are found in the region of the
muzzle, and the infection is known as follicular mange, characterized by
a raw appearance in advanced cases. Among those listed by Hirst 14 are
the following cosmopolitan species: Demodex canis Leydig, which
parasitizes dogs; D. cati Megnin, parasite of cats; D. equi Railliet, horses;
Demodex bovis Stiles, which causes swellings about the size of a pea in
the hides of cattle (shoulder and neck), affecting their value; Demodex
phylloides Csokor, which infests the skin of swine, producing white
tubercles.
Owing to the fact that the follicle mites occur so deeply in the skin,
treatment is very difficult. Penetrating materials are necessary, for exam-
ple, benzine, one part, and olive oil, four parts, or applications of tincture
of iodine. Frequent applications must be made until a cure has been
effected, which is doubtful. The following treatment for follicular mange
in dogs is said to be of value-wash the dog with soap and water, then
cleanse thoroughly and empty the pustules, after which soak the parts
affected (or dip the animal) for five minutes in 5 per cent Formalin or
2 per cent formaldehyde, following this with an application of sulphur
ointment. Repeat the treatment every three or four days until four or
five treatments are given. The use of Ichthyol prepared with lard or
lanolin in the proportions of one to seven is suggested by various authors.
The chances for the cure of follicular mange are slight, and valuable
dogs should be placed under the care of a skilled veterinarian where
facilities are available for the production and administration of auto-
vaccines.
LIPONYSSID MITES
Family Macronyssidae
Family Macronyssidae. The important group of Liponyssid mites was
heretofore included in the family Dermanyssidae Kolenati, 1859 until
1931, when Vitzthum (loc. cit.) erected the family Liponissidae. In
1936 Oudemans proposed Macronyssidae as the family name for this
group. Da Fonseca 15 calls attention to the fact that the type species of
Liponyssus (Lipimyssus setosus Kolenati), 1858, is lost; hence the
generic characters cannot be ascertained. It is, therefore, an unscientific
procedure to accept Liponyssus with unknown characters as a type for
a family, and Macronyssus Kolenati is proposed as a substitute. "Ma-
cronyssidae," therefore, becomes the new family name. Fonseca sum-
marizes the diagnosis of the new family as follows: "Mandibles' with
fixed and moveable arms devoid of any armature in the female. Palps
with a bifid seta at the fifth segment. Dorsal shield of adults entire or
divided in females. Sternal plate with two or three pairs of setae. Anal
plate more or less piriform, never enlarged or prolonged. Holoventral
shield of males entire or divided. Haemotophagus; oviparous or ovovi-
parous; sometimes facultative parthenogenetic; permanent ectoparasites
on mammals, birds and reptilia."
All representatives of this group (Liponyssids) were heretofore in-
cluded in the genus Liponyssus Kolenati, 1858; now, however, Fonseca
(loc. cit.) lists 17 genera of which 5 are new. Liponyssus is retained as
a monotypic genus Liponyssus setosus (Kolenati); the familiar tropical
Liponyssid rat mite and several other species are placed in a new genus
MITES 539
Bdellonyssus Fonseca 1941. LiponysStls bacoti (Hirst), therefore, be-
comes Bdellonyssus bacoti (Hirst) (Fig. 174) .
The genus Bdellonyssus Fonseca includes a homogeneous group of
mostly tropical species in which the "dorsal shield is undivided; sternal
plate of female with three pairs of setae and two pairs of slit-like pores;
genital shield of female tapering to a more or less sharp point, not scale-
like and with only a pair of setae; tibia I more than 1 Y2 times as long
as broad in the female; coxae of females and of nearly all males devoid
of spines other than the dorsal one of coxa II; first p~lpal segment with
a spur; holoventral shield of male undivided and undilated at the ven-
tral surface" (Fonseca).
Tropical rat mites. According to Fonseca (loc. cit. ) the triad of
species Bdellonyssus (= Liponysstls) bacoti ( Hirst), B. brasiliensis
FIG. 174. The tropical rat mite,

B. bacoti; (left) dorsal view, (right)
ventral view. (Mter Dove and Shel-
mire.)
(Fonseca), and B. nagayoi (Yamada) forms so homogeneous a group

that a correct identification will always be difficult. All three species have
been taken on the Norway rat as well as other animals, and all have
been reported to be annoying to man.
The tropical rat mite was first recorded from rats (Rattus r. norvegi-
cus) in Egypt by Hirstl 6 and described as Leiognathus bacoti. This mite
is now reported from many parts of the world as irritating to man;
Bishopp17 reported it from the southern United States in 1928, and it is
now known to be Widely distributed. Bishopp states: "The bite is dis-
tinctly painful at the time the mouth parts are inserted. A sharp itching
pain is usually experienced. Usually there is more or less irritation and
itching at the site of the bite for several hours along with the develop-
ment of a small haemorrhagic area."
Dove and Shelmire 18 (1932) report having experimentally transmitted
the Texas strain of endemic typhus through bites of the tropical rat mite,
B. bacoti (Hirst) from guinea pig to guinea pig. Various other workers
have been unable to confirm the work of Dove and Shelmire.
Williams and Brown 19 (1945) have demonstrated that B. bacoti acts
as an intermediate host of a filariid worm, Litosomoides carinii (Trav. ),
a parasite of the cotton rat, Sigmodon hispidus. Bertram, Unsworth, and
Gordon 20 were also successful. Recently Philip and Hughes 21 (1948)
have shown by preliminary tests that B. bacoti can transmit the causal
agent of rickettsial pox, Rickettsia akari, from mouse to mouse by the
bite. Transovarial passage to at least some of the mite offspring was also
demonstrated.
Life cycle. Bertram, Unsworth, and Gordon (1946, lac. cit.) sum-
marize the life cycle of B. bacoti (Hirst) as consisting of adult male and
female, the egg, the nonfeeding larva, the bloodsucking protonymph,
and the nonfeeding deutonymph. Two engorgements, one by the pro-
tonymph and one by the adult, are required to complete the life cycle in
from 10 to 12 days at a temperature of 25° C. These authors state that
unfertilized females reproduce parthenogenetically. Adult females were
kept alive for 33 days on six engorgements, eggs up to minimum of 12
being laid between meals.
The nymphs and adults are very active and readily leave the nests
and harborages of the hosts and travel freely for some distance to at-
tack persons in restaurants, warehouses, offices, the home, and other
situations where rats abound or have recently been.
Control of tropical rat mites is essentially one of rat control. In the
absence of food the mites perish in about two weeks; therefore, if no
other measures are taken, the pest will abate itself in about that time.
However, with the elimination of the appropriate murine hosts they
may become more annoying because of their enforced search for a blood
meal. Along with rat control the author has recommended that tables,
desks, chairs, cabinets, and woodwork be rubbed with a cloth moistened
with kerosene. Bishopp recommends a mixture of anthracene oil, one
part, and kerosene, two parts, for floors. Dusting rat harbors, floors, and
other surfaces on which these mites crawl with 10 per cent PDT also
gives good results.
The tropical fowl mite, Bdellonyssus (= Liponyssus) bursa (Berlese,
1888) is a widely distributed poultry pest reported from South America
(Argentina, Brazil), India, Africa, China, and more recently (1917,
Wood) from the United States. It appears to be a widespread parasite
of the English sparrow, Passer domesticus, as well as of the chicken,
Gallus domesticus. Although man is frequently bitten by this mite, no
symptoms are reported other than a slight local irritation. This mite is
a very tiny, fast-moving species measuring barely 1 mm in length.
Wood22 reports that the mites tend to accumulate on a few feathers
about the vent of the fowl but when abundant may be quite generally
distributed over the body. The fluffy down of the hen feathers lining the
nest is a favorite location. This species feeds both by day and by night;
the adults remain on the host and feed intermittently. The eggs of this
mite are laid either on the fluff feathers of the host or in the nest. The
incubation period according to Wood is three days; the minute larva
MITES 541
does not feed but molts in about 17 hours and is ready to feed in this,
the first nymphal, stage. The life cycle is said to require from 8 to 12
days.
The northern fowl mite, Leiognathus" (= Liponyssus) sylviarum
(Canestrini et Fanzago, 1877), resembles the tropical fowl mite very
closely; in fact, Hirst, according to Cameron,23 while pointing out the
slight differences suggests that the tropical mite may be but a variety
of the northern species. However, Fonseca places it in a different genus,
Leiognathus characterized as follows: "Dorsal shield undivided; sternal
shield with two pairs of setae; genital plate not scale-like, tapering to a
point posteriorly and provided with only a pair of setae; coxae devoid
of spines; holoventral shield of the male undivided." While reports of
damage, often considerable, to chickens (Gallus domesticus Linn.), are
from northerly regions, the first record came from Pisa, Italy, and severe
infestations are reported from California and Florida. Furthermore, the
host distribution as shown by Cameron (loe. cit.) indicates that it is a
widespread parasite of wild birds, sparrows, starlings, and blackbirds
being frequently mentioned.
On the fowl, these minute, very active mites prefer the region of the
vent and accumulate on a few feathers like the tropkal fowl mite and
similarly spread to all parts of the body when the infestation is heavy.
They also spread to the nests, roosts, and crevices of walls, but evidently
deposit their eggs only on the birds. Several authors believe this species
aestivates during the summer months; however, mites do occur on
wild birds and in their nests during the summer months.
The egg stage lasts about 30 hours; the larva, which is quite slug-
gish, does not feed; the average duration of this stage is 9 hours. Informa-
tion concerning the life cycle of this species of mite is very fragmentary
as it is also for B. bursa. The cycle from egg to adult is believed to re-
quire from 8 to 10 days.
Control. The following recommendations for the control of these
mites are made by Cameron (1938, lac. cit.). About 20 minutes before
the birds go to roost during frosty weather (dropping boards, nest boxes,
and roosts having been previously scraped clean) smear the roosts and
roost supports and the front edge of each nest box with nicotine sul-
phate, applying about 2 oz for every 50 birds and treat every pen on
the same evening, making sure that every bird goes to roost. Single
birds may be hand-treated with a few drops of nicotine sulphate smeared
around the vent, under thighs and wings, and on the neck. Make a prac-
tice of treating all birds as soon as they enter winter quarters in the fall,.
since they may have picked up infestations from wild birds. Prevent
wild birds from nesting near poultry houses. The use of extremely fine
.. This generic name is preoccupied and will, no doubt, have to be changed.
sulphur applied to the birds around the vent, under the wings, and on
other feathered parts is also recommended. This treatment must be re-
peated in 10 days.
DERMANYSSID MITES
Family Dermanyssidae
The family Dermanyssidae is characterized by Ewing24 as follows:
"Mouthparts adapted for piercing, the chelic~rae being either devoid of
one or both chelae or of all true teeth; but, however, recurved 'holdfast'
hooklike structures are sometimes present. Integument somewhat leathery
and distensible, but covered to a large degree by one or more dorsal
shields and one or more ventral plates. Each tracheal trunk opening
FIG. 175. The poultry mite. Der-

manyssus gallinae. ·
through a peritreme situated on the side of the body. Sexual dimorphism

evident and at times marked. Parasitic in habits and on vertebrates."
In the genus Dermanyssus the female chelicerae are long and needle-
like, being composed of two closely adhering elements; in the male the
chelicerae are not needle-like but are variously formed, both arms being
always present and easily recognized. The dorsal shield is large and
undivided in both sexes; the legs are rather slender, especially the front
pair. The rim of anus is greatly thickened behind, and the anus itself
usually situated in the posterior half of anal plate. (Ewing.)
The common red chicken mite, Dermanyssus gallinae ( DeGeer ) ,
also known as the roost mite (Fig. 175), is the most important member
of the family Dermanyssidae. It appears to be a pest of chickens through-
out the world. Other poultry such as turkeys as well as pigeons are in-
fested. English sparrows and some other wild birds may serve as hosts.
MITES 543
While man is commonly annoyed by crawling rnites, instances of actually

having been bitten appear to be quite rare. Tile size of the mites varies
from .6 to .8 mm in length; the males average slightly less in length.
They are somewhat pear-shaped and are light grey in color when un-
engorged and a dark red when engorged. Ewing25 describes the female
of Dermanyssus gallinae (DeGeer) as follows:
Rather large species, with a long body and long legs; no shoulders present.
Dorsal plate.-Covering considerably less than half of the dorsal surface of the
body, broadest above the second pair of coxae, with lateral margins converg-
ing posteriorly behind widest part until truncate end is reached. Each pos-
terolateral corner of dorsal plate is occupied by an oval light spot. Ventral
cephalothoracic characters.-The posterior pair of sternal pores is represented
by two minute circular openings on the posterior margin of the sternal plate
near the posterolateral corners. Anal plate.-Sligp.tly longer than broad, an-
terior margin slightly out-curved. Anal opening jUore than twice as long as
broad, with lateral margins parallel Anal covers very -long, occupying more
than half of the anal opening; behind them in the anal opening a minute,
transverse, sclerotized bar. Paired anal setae situated at a level slightly in
front of middle of anal opening and at about one-third the distance from rim
of anal opening to lateral margins of anal plate; unpaired anal seta situated
at a point about one-third the distance from posterior rim of anal opening
to posterior margin of anal plate.
Habits and life history. During the daytime the mites remain hidden
in crevices of the henhouse and roosts, under 'boards and debris. In these
hiding places the mites deposit their eggs. At night the pests swarm out
from their hiding places and attack the birds upon the roosts. Their
attack is not restricted, however, altogether to the nighttime, for swarms
of mites may be found on setting hens and lllying hens during the day
if these are nesting in dark places.
Bishopp and Wood,26 and Wisseman and Sulkin27 have made im-
portant contributions to a knowledge of the life history of this mite.
Adult female mites usually begin ovipositing in from 12 to 24 hours
after feeding and lay as many as 7 eggs, which hatch in from 48 to 72
hours at summer temperatures. The hexapod larvae are sluggish and
do not feed. The first-stage nymph (eight-legged and active) appears in
from 24 to 48 hours and, after a blood meal, molts and gives rise to the
second nymphal stage. After feeding and a second molt the adult stage
is reached. Occasional instances of a third nymphal stage are reported.
The entire life cycle may he completed in a, short a period as 7 days.
Damage done and control. The poultry mite is a serious pest in many
parts of the world. The damage which this mite causes is very consider-
able and may be summarized as follows: Egg production is greatly re-
duced or entirely prevented; setting hens are often caused to leave their
nests or perish; newly hatched chicks perish in large numbers in the
presence of these mites; chickens lose flesh, are unthrifty, and are un-
profitable for marketing; loss of blood and reduced vitality cause birds
to be easily susceptible to disease. Inasmuch as the mites are seldom
found on the bodies of the birds during the day, except in the first feed-
ing period when they sometimes remain attached for a night and a day,
or in dark nest boxes, control measures are directed most advantageously
when they are in their hiding places. A thorough clean-up of the premises
to which the birds have access, together with the elimination of every
useless article therein, such as boxes, coops, boards, etc., is the first step.
Old. nesting material should be burned and if the infestation is severe,
roosts and nests should be dismantled to be replaced by construction
that will facilitate future clean-ups. Methods must now be directed
against the cracks and crevices of the floors, walls, and even the roof of
the poultry house. In heavy infestations the mites sometimes migrate to
the outside of the house when the inside is sprayed. They should be
looked for in cracks on the outside and if present the outside should
also be treated. The most efficient method of treating poultry quarters
is by the use of liquid insecticides applied by "knapsack" spray pump.
A coarse spray is most effective and should be applied to each area from
several different angles to ensure penetration into all hiding places. Many
of the existing sprays are efficient. Any of the dips used on domesticated
animals made up in a slightly stronger solution than directed for the
dipping of animals will give fairly satisfactory results. The most satis-
factory of all applications, however, are the wood preservatives that have
coal-tar product anthracene oil as a base. These have the advantage of
being effective over a long period of time and soak into the wood rapidly
without leaving a greasy residue to soil the feet, feathers and eggs of the
birds. They should be diluted about one-half with kerosene to facilitate
spraying. One thorough treatment with this type of spray is generally
sufficient. Whitewash containing 3 to 5 per cent of crude carbolic acid
kills large numbers of mites but is not nearly so effective as anthracene
oil. Nicotine sulphate used on the roosts, as for lice, is effective in killing
young mites feeding on the birds and in protecting roosting birds. Used
as a spray at the rate of three tablespoonfuls per gallon of water to which
is added one-half teaspoonful of baking soda it gives very satisfactory
results if applied carefully and in drenching quantities.
Prevention. The chicken mite is introduced into clean flocks in many
cases in contaminated shipping coops. In some cases the introduction
may be accomplished by a few young mites that are engorging for the
first time on the introduced fowls. To overcome this possibility, newly
acquired birds should be kept in special coops for two or three days
before being placed in clean houses. These coops should then be de-
MITES 545
stroyed or disinfected with bOiling water or with one of the sprays sug-
gested above. Shipping coops from other poultry plants should not be
left in or near clean houses, nor should second-hand equipment be in-
troduced unless properly disinfected. Mites will live for from three to
five months without food, a fact which should be considered when
vacant buildings are to be occupied by clean flocks. In cases where the
control of the mite is impossible owing to the character of the quarters
or lack of them, some relief is afforded by wrapping the ends of the
roosts and other points of contact with rags soaked in crude oil to prevent
the mites from gaining access to the fowls after they have gone to roost.
To make this procedure effective crowding should be discouraged, and
the back roosts should not be near enough to the wall to permit the
movement of the mites to the plumage of the birds.
Other Dermanyssid mites. Dermanyssus americanus Ewing was de-
scribed from English sparrows in Washington, D. C. The palpi of the
female extend beyond the tips of the anterior femora; chelicerae are
needle-shaped; the peritreme very short, not extending beyond the third
coxa; the anus is situated almost entirely behind a middle transverse
line; the paired setae are slightly behind the level of the anterior margin
of the anus; the legs are very short and stout, the last pair falling far
short of the tip of the abdomen. The length of the mite is 0.70 mm, the
width, 0.39 mm.
Allodermanyssus sanguineus (Hirst), a mouse mite, the vector of
rickettsialpox of man, is described by Ewing (lac. cit.) as follows:
"Palpi slender, reaching to the tips of anterior femora; chilicerae showing
plainly the needle-like elements representing both arms. Dorsal shield
divided; anterior shield broadest at the shoulders, lateral margins behind
the shoulders concave; posterior shield circular, minute. Sternal plate
squarish, lying entirely between the second coxae, with three pairs of
sub equal marginal setae. Anal plate egg-shaped in outline, anterior
margin broadly rounded; anus situated centrally, rim very thin in front
and on the sides, but enormously thickened behind; paired setae situated
at the level of the center of the anus; median seta situated about two
thirds its length behind anus; caudal area reaching about halfway to the
base of median seta. Legs very long and slender. Length, 0.91 mm;
width, 0.46 mm."
Rickettsialpox of man was first observed in New York City during
the summer of 1946, the causal organism, Rickettsia akari, being isolated
by Huebner, Jellison, and Pomerantz. 28 The epidemiology of the disease
is described by Greenberg, Pellitteri, and Jellison;29 the infection is trans-
mitted by the rodent (mouse) mite, Allodermanys8U8 sanguineus (Hirst),
and the house mouse, Mus musculus, serves as a reservoir.
Pulmonary acariasis of monkeys is traceable to mites of the genus
Pneumonyssus which live in the lungs of the host, e.g., Pneumonyssus
simi cola Banks. Halarachne zalophi Oudemans occurs in the nasal pas-
sages of the California sea lion; and Sternostomum rhinolethrum Troues-
sart is said to produce catarrhal inflammation of the respiratory tract of
fowls. Lung mites commonly infest the respiratory system of snakes.
(Turk. 30 )
The role of bird mites in the epidemiology of arthropod-borne en-
cephalitides. In 1942 Hammon et al. 31 demonstrated that a large per-
centage of wild birds as well as domestic fowl possess neutralizing anti-
bodies for the western equine and St. Louis encephalitis. This discovery
gave impetus to an investigation already under way concerning the part
that avian bloodsucking ectoparasites may have in the infection chain
of these viruses. (See Chapter XII, Mosquitoes as Vectors of Disease.)
In 1944, Smith, Blattner and Heys,32 reported the isolation of the
St. Louis encephalitis virus from chicken mites, Dermanyssus gallinae
( DeGeer ), in nature during a nonepidemic period. Evidence was pre-
sented that this mite is capable of transferring the virus of St. Louis
encephalitis congenitally to its offspring and "that a colony of mites once
infected probably remains infected indefinitely." Sulkin33 ( 1945) re-
covered the virus of the western type of equine encephalitis in the same
species of mite, also in nature, but at the time of an outbreak of the
disease among horses in Texas Howitt and associates 34 (1948) reported
the isolation of the virus of the eastern type of the disease from D.
gallinae. It had previously been shown (pp. 242 to 244) (as early as
1942) that mosquitoes belonging to several species are capable of acquir-
ing, and transmitting to animals, the virus of both the St. Louis and the
western equine encephalitides. However, these viruses appear not to be
transmitted transovarianly in the mosquito, nor have they been recovered
in hibernating mosquitoes; hence poultry and/or wild birds mites serving
as reservoirs would constitute a strong link in the infection chain of the
arthropod-borne encephalitides. In 1948 Smith and associates 35 reported
having transmitted the virus of St. Louis encephalitis to hamsters by
mosquitoes, Culex pipiens, at periods varying from 4 to 27 days after
these insects had fed on chickens having viremia as a result of the bite
of infected mites. Thus the concept is supported that two bloodsucking
vectors may be involved, one the chicken mite which does not bite man
and in which the virus is maintained in nature by transovarian passage,
and the other, a mosquito, which transmits the infection from birds to
other vertebrates, including man. Two other species of bird mites are
involved, perhaps even more plausibly (Hammon 36 obtained negative
results with D. gallinae) , namely, Leiognathus (= Liponyssus) sylvi-
arum (c. and F.) from which Reeves and associates 37 isolated the virus
of western equine encephalomyelitis in California (mites were from
MITES 547
=
nest of English sparrow), and Bdellonyssus ( Liponyssus) bursa
(Berlese) from which Sulkin and IzumP8 isolated the western equine
virus in Texas.
While apparently complicating the infection chain, the discoveries
implicating mites do furnish the factor of maintenance of the virus in
nature. Thus mites infected previously either C()ngenitally or by feeding
on an infected bird may initiate infection, as Hammon (lac. cit., 1948,
p. 244) points out, in nestling birds or in newly hatched fowl in the
spring. The young birds are then fed on by nlOsquitoes, and after the
essential extrinsic incubation period, these mosquitoes may infect other
birds. Hammon continues to reason that several such cycles in mosquitoes
and birds will build up the number of infected mosquitoes, and infection
may then appear in men or horses. Infection is llsually seen first in horses
and later in man, no doubt, because of the greater number of mosquitoes
having access to these large unprotected animals and because of the
zoophilism of these insects.
CHIGGER MITES
Family Trombidiidae (Subfamily Trombiculinae)
Family Trombidiidae. The family of mites (Trombidiidae) to which
the chiggers (larval Trombidids) belong is characterized by Banks
(loc. cit.) as follows (adult characters):
the body being divided into two portions, the antE:rior (cephalothorax) bear-
ing the two anterior pairs of legs, the palpi, mouth parts, and eyes; the poster-
ior (abdomen) is much larger and bears the two posterior pairs of legs. The
mandibles are chelate; at least there is a distinCt jaw or curved spine-like
process. They are always red in color, some, however, being much darker
than others. The body is covered with bristles or feathered hairs according
to the species. The palpi are nve-jointed, quite prominent, often swollen in
the middle, the penultimate joint ending in one Or two claws, the last joint
(often clavate) appearing as an appendage or "thumb" to the preceding joint.
The legs are seven-joirited; the tarsi terminate in two small claws. The legs
are clothed in the same manner as the body. There are two eyes upon each
side of the cephalothorax, quite frequently borne on the distinct pedicel.
The chiggers belong to the subfamily Trombiculinae, in which "each

tarsus is 3-clawed, outer claws nearly always equal, forming a pair, mid-
dle claw usually longer than others; pseudosti~matic organs, when sim-
ple, long and more or less flagelliform" (Ewing39). There are about 50
recognized genera and well over 100 species.
Brennan40 has made an effort to clarify the terms used in describing
chiggers; the arrangement of the setae is considered important, but the
dorsal formula is not always constant and cannC)t be relied upon entirely
in the diagnosis of a species. Recently Wharton l l (1948) has shown that
the chaetotoxy of the legs is an important aid to the taxonomic study of
larval trombiculids. He points out that four types of modified setae are
found on the legs and palps as follows:
1. Blunt striated sensory setae, one on the dorsal side of each tarsus of
the first two pairs of legs and one at the base of the palpal tarsus on the ventral
surface.
2. Pointed striated sensory setae, on the genua and tibiae of all legs, the
tarsus of leg I, and the pretarsus of legs I and II. There may be one on tarsus
III and the palpal tarsus, but this is unusual.
3. Microsensory setae, may be present on the genua, tibiae, and tarsi of
legs I and II.
4. Whip-like setae, one or more, may be present on tela-femur, genu,
tibia, and tarsus of leg III.
The literature pertaining to the mite family Trombidiidae has grown

enormously during the past few years as shown by the bibliography
published by Williams 42 in 1944 consisting of 375 references.
Among the Trombiculid mites causing a severe dermatosis are the
European species, Trombicula autumnalis (Shaw), known as the harvest
mite. Eutrombicula differs from Trombicula in having bifid rather than
trifid claws in the larva. There are at least three species of chiggers
known to attack man in the Americas (Jenkins 43 ); namely, Eutrombicula
alfreddugesi (Oudemans), E. batatas (Linn.), and E. masoni (Ewing).
=
E. alfreddugesi Trombicula thalzahuatl (Murray); early mention
of Leptus irritans Lucas also probably refers to this species of larva; it is
the common chigger of the United States, ranging from New England
to Nebraska, south to Florida and Texas, and west to California; it also
occurs in Mexico, Central America, and South America, where it is
known as bete rouge. "It occurs abundantly in second growth cut-over
areas, especially blackberry patches, forest edges, and river valleys where
many species of birds, mammals and reptiles are often found" (Fig.
176).
E. masoni, according to Jenkins (loc. cit.), is distributed along the
Atlantic coast from Florida to Massachusetts and along the Gulf coast
to Texas, with isolated "islands" in Michigan and Minnesota. It is the
most abundant species in Florida and parts of Georgia and is often
present in large numbers when E. alfreddugesi is rare or absent.
E. batatas is a tropical species and has been collected in the warmer
southeastern part of the United States, and recently (Doetschman and
Furman, 1948, in MS) it was reported from California. It is said to attack
man rarely in the United States but is known to infest man freely in
Panama and other tropical areas.
MITES 549
The bright red adult mites are found on the surface of moist ground
or to a depth of 2 or 3 inches in drier soil, being often seen in the bases
and root stocks of clumps of grass, in sphagnum moss, and under logs.
In central Florida, Jenkins (loc. cit.) found adults of E. alfreddugesi
and E. masoni in largest numbers in winter an_d early spring; they be-
came less common in summer and fall. Their food is said to be insect
eggs.
According to Michener," the spherical pale-dull orange-colored eggs
of Eutrombicula batatas are laid singly on the Surface of moist soil. The
FIG. 176. Chigger (mite) or harvest mite. Adtllt (left), larva (right) of
Eutrombicula alfreddugesi. (From Ewing's Mantlal of External Parasites.
Courtesy of Charles C. Thomas, Publishers.)
incubation period ranges from 4 to 5 days, when the egg shell breaks
into two portions, anterior and posterior, which become separated; but
the nearly fully developed larva remains insidE! this "deutovum" for six
or seven days more. Having completed the (~eutoval stage the fully
formed red hexapod larva emerges and crawls rapidly about in search of
a host. The larvae can live for about two weeks (probably longer) with-
out feeding. The opportunity to find a suitable host should be good
since it is known that close to a hundred species of vertebrates, mammals,
birds, amphibians, and reptiles may serve as bosts. Ewing (1944, loco
cit.) reports rearing E. alfreddugesi from box nIftIes, Terrapene carolina
(Linn.), and from young toads, Bufo species, 9.s well as from snakes. 45
Keay 46 reports numerous hosts for T. autumnalis, among these the dog,
horse, shrew, fowl, and sparrow. Engorgement of the chiggers (see next
section explaining feeding habits) takes from 2 to 10 days, after which
they drop to the ground and remain quiescent (protonymph) within
the dead larval skin for 5 to 7 days, emerging with eight legs (the
nymph). The nymphal stage lasts from 21 to 52 days, including the pre-
adult stage, when another molt liberates the adult. The life cycle of E.
hatatas, according to Michener's (lac. cit.) summary requires from 39
to 85 days. The adult may live up to 45 days.
Chigger dermatitis. In many parts of the world, particularly in the
warmer portions and during the late summer months in temperate
climates, persons who have walked among tall weeds and grass, bram-
bles, and low scrub often suffer an intolerable itching, which begins
3 to 6 hours after exposure and is followed by a severe dermatitis con-
sisting of pustules and wheals. When, after a trip through tall weeds
or grass or a patch of berry brambles, itching begins around the ankles,
the knees, and the waist, a careful examination will pretty surely reveal
at least one (there will be others) bright red mite barely visible to the
naked eye either traveling fast or about to attach itself to the skin. A
microscopic examination will show that it has three pairs of legs and no
doubt is the larva of a trombiculid mite-a chigger.
Chiggers do not burrow into the skin as is commonly believed.
Ewing47 observed and studied daily 26 chiggers on his own skin; 21
were attached to the smooth surface of the skin, while five were attached
at the bases of hairs, each having the capitulum thrust into the mouth
of the hair follicle; not a single one had penetrated a pore or hair follicle.
The chigger does not suck blood but injects a digestive fluid (a proteo-
lytic enzyme) when firmly attached. This fluid dissolves the tissues so
that the resulting liquid may be utilized by the chigger. The skin of the
host becomes hardened, and there is formed a tube (stylostome) in
which the chigger lies and continues to imbibe the nourishing liquid until
it is replete; then it retreats and drops to the ground. According to
Michener and Michener 48 it is presumably the action of the digestive
fluid which causes the "bite" to itch after a few hours. Williams 49 points
out that histological preparations of chigger bites on rabbit ears show
that the epidermis is completely penetrated. A tube lined with stratum
germinativum cells is formed which extends to the derma and subcutis.
This tube appears to represent a reaction of the host to the secretion of
the chigger, and its inner layer of cells are necrotic and give evidence
of digestion.
To free infested areas of chiggers, lawns and gardens may be treated
with very fine dusting sulphur applied with a hand duster at the rate of
one pound per 1,000 sq ft, or 50 pounds per acre. Apply at intervals of
MITES 551
two weeks during the chigger season, May to August. The ground should
be carefully treated; the sulphur dust will not injure vegetation, but
bright metal objects are easily tarnished. Dusting with DDT as recom-
mended for tick control is also suggested by some authors. The use of
benzene hexachloride (12 per cent gamma isomer) applied as a dust or
spray at the rate of 4 pounds per acre has recently been reported as a
successful treatment by the United States Bureau of Entomology and
Plant Quarantine. Keeping down weeds and mowing the grass short so
as to expose chiggers to the sun and dry the soil are good preventive
measures.
Having been exposed to chiggers one should immediately take a bath
lathering the body with a sulphur soap and allow the lather to remain
about ten minutes before rinsing off.
Chigger repellents such as dimethyl phthalate and benzyl benzoate
are available for the impregnation of clothing, particularly about the
ankles and at the waistline. Relief from the "bites" may be had by
applying various remedies such as a 5 per cent solution of benzocaine in
alcohol.
Tsutsugamushi disease, also known as "Japanese Hood fever," "Japa-
nese river fever," "kedani fever," and more recently "scrub typhus" and
"Mossman fever" (Australia), is an ancient disease (Blake et al.,50 1945)
first described from Japan. It was long said that the infection is trans-
mitted to man by the larval trombiculid (chigger) mite, Trombicula
akamushi (Brumpt, 1910) from rodent reservoirs (voles, Microtus monte-
belli Sado). It was not, however, until 1916 that positive experimental
evidence to that effect was secured by Miyajima and Okumura 51 in tests
with monkeys. The specific rickettsial nature of the infection was deter-
mined in 1930 by Nagayo et al.,52 who described Rickettsia orientalis.
However, as early as 1920, Hayashi53 reporting on the etiology of tsut-
sugamushi disease gave the name Theileria tsutsugamushi to what he
then believed to be the causal agent. It is contended by Steinhaus 54 and
others that, since it cannot be proved that Hayashi did not see the causal
agent of this disease, the specific name which he gave should not be
ignored. Under the rules of priority the specific name of the causal
organism would then be Rickettsia tsutsugamushi (Hayashi), which
name is accepted by Philip 55 in his account of tsutsugamushi disease in
World War II.
The disease has an incubation period of 6 to 12 days and over. Dur-
ing the first 5 to 7 days it is characterized as follows (Blake, et. al., 1945,
loco cit.): ''headache (postorbital), apathy and general malaise, fever
( chills), relative bradycardia, anorexia, conjunctival congestion, lympha-
denitis, often regional, and an eschar." The eschar is the primary lesion
which originates at the point of "chigger" attack (ankle, shin, groin,
waistline, or axilla) in the great majority of cases. It is a painless papule
at first, usually unnoted by the patient. It slowly enlarges to a diameter
of 8 to 12 mm, the center becoming very dark and necrotic; a shallow
ulcer may result eventually, leaving a scar. Between the fifth and eighth
day in nearly all cases a characteristic dull red macular or maculopapular
rash appears on the trunk and may spread to the extremities involving
the palms, soles, face, and scalp. This may persist for about two weeks,
often fading in a few days. Enlargement of the spleen, nervous dis-
turbances, delirium, and prostration are common symptoms; in many
cases there is deafness. The majority recover by lysis in three to five
weeks. In extremely severe cases "symptoms and signs of more severe
pneumonitis and encephalitis are constant. Evidences of peripheral cir-
culatory collapse are common and signs of myocarditis may appear.
. . . Thromboses and cerebral or gastrointestinal hermorrhage may take
place." Mortality ranges up to 40 per cent; death results from cardiac
failure, usually in about three weeks. The rickettsiae may readily be
recovered from the blood of patients during the acute stage of the disease
by the intraperitoneal inoculation of 0.2 to 0.3 ml of blood in white mice.
(Blake et al., loco cit.)
Although the greatest number of cases occur at low elevations (sandy
bottom land overgrown with grasses and scrub), infections occur at
elevations of 2,000 to 3,000 feet and, in Formosa, as high as 6,500 feet.
The "chigger" mites have been reported as high as 8,000 feet in India.
The infection is endemic in many parts of southeastern Asia and ad-
jacent islands in the Indian Ocean and southwest Pacific (Ceylon, Japan,
Philippines, Netherlands Indies, Taiwan), and the coastal area of North
Queensland, Australia.
The importance of scrub typhus (tsutsugamushi disease) as a "medi-
cal casualty producer in some areas during the Asiatic-Pacific operation,
1941-1945," according to Philip (loc. cit.), "was second only to malaria
and was more dreaded by the men." Interference with actual combat
operations was greatest, Philip states, on the Assam-Burma front where
"eighteen per cent of a single battalion got scrub typhus in two months
and in that time 5 per cent of the total strength had died of it."
"American Task Force operations in the Schouten Islands resulted
in a thousand cases in the first two months on Owi and Biak, reaching
a total of 1,469 casualties in 6 months time, while at Sansapor beach-
head the curve for weekly admissions on a thousand-per-year basis shot
up to over 900 at the end of the second week, a rate higher for an indi-
vidual episode than any yearly rate for all causes in the entire American
Army in all theatres. . . . These two disasters alone provided a potential
estimated loss of over 150,000 man days to the American Sixth Army."
The vector. There are a number of species of chiggers in endemic
MITES 553
areas which attack man and cause "scrub itch" but are not vectors of
tsutsugamushi disease. In spite of much research during the late war
years, Trombicula akamushi (Brumpt) and Trombicula deliensis Walch
are the only proved vectors, as Philip (1948) points out. Furthermore,
Philip states that he has specimens of supposed T. akamushi from Japan
that are indistinguishable morphologically from T. deliensis; hence there
is good reason to suspect that the latter may really be but a variety of
T. akamushi (Philip and Kohls, 194856 ).
The adult mites measure from 1 to 2 mm in length, are generally
reddish in color, often pale; they are ground-inhabiting and nonparasitic
and are said to feed on insect eggs and minute arthropods; many authors
refer to them as scavengers. The winter is spent in this stage. They are
said to deposit their eggs in late spring or early summer in loose top
soil under leaves, etc., in damp places. The hexapod larvae (0.32 to 0.43
mm in length) hatch in about 3 weeks under favorable temperature
conditions (about 20° C) and wait the coming of a suitable vertebrate
host-mammal or bird (man is an accidental host). Having successfully
encountered a host the chigger attaches itself and begins to feed. Numer-
ous hosts are available, but continuous propagation of the infection is
dependent upon susceptible so-called reservoir animals (many of the
host animals are suitable for the maintenance of the vector species in-
cluding birds and reptiles), the first of which was demonstrated to be a
vole, Microtus montebelli (see above). Wartime studies as reported by
Philip and Kohls (1948, loco cit.) indicate that rats play an important
widespread role in the disease cycle in nature. "Six kinds of rats (among
them Rattus concolor browni and R. flavipectus yunanensis) indigenous
to New Guinea, Burma, or India, plus a species each of field mouse and
tree shrew (Tupaia belangeri versurae) have been added to two murid
species already demonstrated to carry natural infection in Japan and
Malaya, and to others under strong indictment in Formosa, Sumatra, and
Australia." Infection has been demonstrated in the circulation of English
sparrows, pigeons, and turkeys (Philip).
The larval mite (akamushi) is said to remain on the host, commonly
in the ears, feeding for three or four days, then drops to the ground
where in five or six days it molts and the nymphal stage (eight-legged)
is reached. This stage like the adult stage, which it is said to reach with
another molt in about ten weeks, is nonparasitic (Miyajima and Oku-
mura, loco cit.). Experimental evidence is complete showing that larval
mites (chiggers, i.e., akamushi) hatched from eggs layed by such female
mites may transmit the infection to experimental animals. The proportion
of infected larvae is said to be perhaps less than one per cent. That there
is a chigger mite in the United Stales which is a member of the "aka-
mushi" group is a matter of some interest. Wharton 57 states that the
larva of Trombicula myotis Ewing is more closely related to T. akamushi
than to any other species, and could easily be confused with it.
Control. During the early months of warfare in the southwest Pacific
area little or nothing was known about the ecology of scrub typhus,
information needed to plan successful measures for the control and pre-
vention of the disease. Thus because of the high incidence of infection
among troops bivouacked or maneuvering in the abundant tall kunai grass
warning against such practices were issued; however, severe epidemics
occurred under the "coconut canopy" along the beaches. There were
outbreaks under various ecological conditions, causing considerable con-
fusion. There appear to be several factors, however, that do provide some
basis for control: the mites are extremely sensitive to lowered humidity.
Therefore, clearing of grass and scrub (clean camp sites) is a practical
procedure, with the use of flame thrower and bulldozer suggested; sec-
ondly, rat harborage and a plentiful supply of food for native rats pro-
vide an abundance of hosts for the larval mites, hence rat control is an
important chigger-control procedure.
Protection of the person against chiggers by repellents (dimethyl
phthalate) applied to clothing was achieved during the latter part of
World War II. Dipping uniforms in a dibutyl phthalate-benzyl benzoate
emulsion gives good protection.
LOUSE-LIKE MITES
Family Tarsonemidae
Characteristics of Tarsonemidae. This is a small family of soft-bodied
mites having in the female a characteristic "prominent clavate organ of
uncertain use" between the first and second pairs of legs. The third and
fourth pairs of legs are separated from the first and second pairs by a
long interspace. Sexual dimorphism is marked in the several species. The
piercing, sucking mouth parts are provided with slender needle-like
stylets. Many of the species are predaceous or parasitic on insects, while
others suck the juices of plants.
Pediculoides ventricosus (Newport) (Fig. 177) is a widely distrib-
uted predaceous mite which attacks the larvae of a number of species
of insects such as the Angoumois grain moth [Sitotroga cerealella
(Oliv.)], the wheat joint-worm [Harmolita grandis (Riley)], the peach
twig borer (Anarsia lineatella Zell.), the cotton-boll weevil (Anthonomus
grandis Boh.), the bean and pea weevils [M ylabris quadrimaculatus
(Fabr.) and M. obtectus (Say)], etc. It is, therefore, normally a bene-
ficial mite, but unfortunately it also attacks man, producing a very dis-
agreeable dermatitis commonly called "straw itch."
MITES 555
While the male mite is very tiny, just about visible to the naked eye,
the female when pregnant becomes enormously swollen, measuring
nearly a millimeter in length, the abdomen presenting a globular appear-
ance, the cephalothorax and appendages barely visible.
Within the enlarged abdomen of the female may be found rather
large eggs which hatch internally, and the young mites develop to ma-
turity within the body of the mother before being extruded. The number
of young produced by the female is said to range from a few to nearly
300. The female mites are often fertilized within the body of the mother.
FIG. 177. Pediculoides ventriCOSllS.

(a) Male; (b) female; (c) gravid
female, showing developing eggs.
(Redrawn after various authors.)
A number of epidemics of dermatitis have been traced to these

mites, infection having been brought about by sleeping on straw mat-
tresses or while laboring in grain fields at harvest time. The infection
has been confounded with hives, scabies, and even chickenpox; the
neck, chest, abdomen, back, arms, and legS-in fact, the whole body-
may be involved, and the itching is very intense. The eruption, which
appears in 12 to 16 hours after the attack, is commonly accompanied
with some fever.
According to Goldberger and Schamberg 58 the itching subsides under
continuous exposure in from three to seven weeks. They recommend
treating the infection with an ointment of beta naphthol, sulphur, ben-
zoate, and lard.
To destroy mites in the straw of mattresses or in other situations,
fumigation with sulphur, or steaming, or dry heat is recommended.
As to prevention, Webster59 suggests burning the grain stubble dur-
ing the fall, winter, or spring, and also states that threshing direct from
the shock resulted in the control of the grain moth and consequently of
the parasitic mites.
FLOUR AND MEAL MITES, GROCER'S ITCH

Family Tyroglyphidae
Characteristics of Tyroglyphidae. This family includes a small group
of very tiny mites, ordinarily about 0.5 mm or less in length. Several of
the species attack grain, flour, meal, dried meat, hams, dried fruits, insect
collections, etc. Their development is so rapid that literally millions of
them may appear in a stored product in a few days.
The metamorphosis of this group involves a peculiar stage known as
the hypopus, appearing after the larval and nymphal stages, very unlike
either of these and very different from the adult. In this stage the mites
are said to attach themselves, nonparasitically, to flies and other insects,
which disseminate them.
Persons handling stored products of various kinds, cereals, flour,
meal, etc., may be attacked temporarily by the mites and experience a
severe dermatitis, known as "grocer's itch," "copra itch," etc.
Tyroglyphus (= Acarus) siro (Linn.) is the grain mite or cheese
mite, found in grain, stored products, and cheese; this mite causes a
rash known as "vanillism" in vanilla-pod handlers; T. (= Alcurobius)
farinac (DeG.) is known as the flour mite and is common in flour mills
and granaries; T. americanus Banks is known as the cereal mite and is
widespread and abundant on cereal products, seeds, stored prunes and
other fruits; Tyroglyphus longior Gerv. attacks grains, cereals, dry seeds,
etc., and causes "copra itch" among workers handling copra. T. farinae
(DeG.) and T. longior Gerv. have been reported from the urinary tract
and T. longior Gerv. from the intestinal tract (intestinal acariasis60).
RED SPIDERS
Family Tetranychidae
Characteristics of Tetranychidae. To this family belong the "web-
spinning mites," most commonly infesting vegetation and destructive to
fruit trees and other plants. The term "red spiders" is ordinarily applied
to the group. Tctranychus bimaculatus Harvey, the two-spotted mite,
attacks many species of plants as does the common red spider, T. tclarius
(Linn. ).
Persons employed in picking hops and harvesting almonds, etc., often
complain of itching produced by the red spiders, but this soon disappears.
MITES 557
QUILL MITES
Family Cheyletidae
Syringophilus bipectinatus Heller, the quill mite of poultry, lives in
the shafts of the primary wing feathers. Rebrassier and Martin61 report
that this mite caused a peculiar molt; the loss of feathers extending over
half of the body in most cases and in many instances over the whole body.
The birds were reported to be apparently in good physical condition.
S. columbae Hirst is a quill mite of pigeons.
AIR-SAC MITES
Family Cytoleichidae
Cytoleichus nudus (Vizioli) is known as the air-sac mite of poultry
because of its habitat in the air passages. Laminosioptes cysticola
(VizioU) occurs in the subcutaneous tissue but is considered of no eco-
nomic importance.
THE PENTASTOMIDS
Tongue Worms
The Class Pentastomida, also known as the Linguatulida, compre-
hends the so-called tongue worms, parasites of numerous species of
vertebrate animals, from fishes to man. Because of certain resemblance
to the phytoptid mites the tongue worms have been classed with the
Acarina (Sam bon, 62 1922). Heymons63 points out that the difference
between the pentastomids and annelids is far less than the difference
between them and the arthropods.
These endoparasites inhabit primarily the lungs and air passages of
their hosts. The body consists of a head or cephalothorax and a not
distinctly separated abdomen. The head possesses five ventral openings:
the median opening is the mouth, and the remaining two pairs lead to
hollows from which chitinous hooks protrude. They are legless, vermi-
form, and pseudo-annulated. Development is indirect, in that an inter-
mediate host is required.
The class Pentastomida is divided into two orders ( 1 ) Cephalo-
baenida and (2) Porocephalida. The order Porocephalida consists of two
families (1) Porocephalidae, which includes among others the genus
Porocephalus and the genus Armillifer; (2) Linguatulidae, which in-
cludes the genus Linguatula, to which the tongue worm Linguatula
serrata Frolich helongs.
Pentastomiasis of man is commonly encountered, the cause being
either linguatulids (linguatulosis) or porocephalids (porocephaliasis).
Linguatulosis infection is primarily due to ingestion of the eggs of
Linguatula serrata Frolich (= L. rhinaria Moniez), the adult of which
normally inhabits the nasal and frontal sinuses of the dog and wolf,
occasionally also those of the fox. The adult female reaches a length of
80 to 120 mm and a breadth of 8 to 10 mm (Fig. 178). The males are
hardly half this size. The eggs are discharged from the nostrils and mouth
of the infected animal and are ingested by the intermediate host with
food and water. Intermediate hosts under natural conditions are as a
rule wild rabbits in whose body cavities the larvae travel freely, invading
FIG . 178. Linguatula serr.ata.

Larva (left), adult (right). (Redrawn
after Brumpt.)
the liver and lungs; if infested rabbits are eaten by a dog, wolf, or fox
the larvae develop into the mature form in these animals. Faust6 .! has
reported upon the presence of the immature form (encysted nymph) in
the liver and lungs of a laboratory rabbit in Peking. He states that the
material from the Peking rabbit averaged 4 mm in length by 0.8 mm in
breadth.
Hobmaier and Hobmaier 65 report natural infections of brown rats
(Rattus norvegicus) and artificial infections of ground squirrels (Citellus
beechyi) , both of which may contribute as intermediate hosts to the
cosmopolitan distribution of Linguatula rhinaria.
Another linguatulid of man reported by Faust is Armillifer monili-
farmis (Diesing) from the liver of a Tibetan. This species as an adult
MITES 559
described by Hill 66 is parasitic in the respiratory tract of pythons and
has for nymphal host, among many others, such animals as mongooses,
monkeys, ground squirrels and rats. Leiperia gracilis (Diesing) and a
very closely related species, Porocephalus crocodili (Wheeler), occur in
the crocodile. .
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28. Huebner, Robert J.; Jellison, William L.; and Pomerantz, Charles;
1946. "Rickettsialpox-a newly recognized Rickettsial disease. IV. Isolation
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1947. "Rickettsialpox, newly recognized rickettsial disease. III Epidemiology."
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30. Turk, F. A., 1947. "Studies of Acari. IV. A review of the lung mites
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31. Hammon, W. McD.; Lundy, H. W.; Cray, J. A.; Evans, F. C.; Bang,
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certain vertebrates as part of an epidemiological study of encephalitis of west-
ern equine and St. Louis types," J. Immunol., 44:75-86.
32. Smith, Margaret C.; Blattner, Russell J.; and Heys, Florence, M.;
1944. "The isolation of the St. Louis encephalitis virus from chicken mites
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33. Sulkin, S. Edward, 1945. "Recovery of equine encephalomyelitis virus
(western type) from chicken mites," Science, 101:381-83.
34. Howitt, B. F.; Dodge, H. R.; Bishop, L. K.; and Corrie, R. H.; 1948.
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"Virus of eastern equine encephalomyelitis isolated from chicken mites
(Dermanyssus gallinae) and chicken lice (Eomenacanthus stramineus)," Proc.
Soc. Exper. Bioi. & Med., 68:622-25.
35. Smith, Margaret G.; Blattner, Russell, J.; Heys, Florence M.; and
Miller, Albert; 1948. "Experiments on the role of the chicken mite, Dermanys-
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]. Exper. Med., 87:119-38.
36. Hammon, W. McD., 1947. "The arthropod-borne virus encephalitides,"
"The Charles Franklin Craig Lectures," 1947, Am. J. Trop. Med., 28:515-25.
37. Reeves, W. C.; Hammon, W. McD.; Furman, D. P.; McClure, H. E.;
and Brookman, B.; 1947. "Recovery of western equine encephalomyelitis virus
from wild birds (Liponyssus sylviarum) in Kern County, California," Science,
105:411-12.
38. Sulkin, E. S., and Izumi, E. M., 1947. "Isolation of western equine
encephalomyelitis virus from tropical fowl mites, Liponyssus bursa (Berlese),"
Proc. Soc. Exper. Bioi. and Med., 66:249-50.
39. Ewing H. E., 1944. "The trombiculid mites (chigger mites) and
their relation to disease," ].Parasitol., 30:339-65.
40. Brennan, James M., 1947. "New species of chiggers (Acarina, Trom-
biculidae) from bats of the Nearctic region," J. Parasitol., 33:245-52.
41. Wharton, G. W., 1948. "Four new Peruvian chiggers (Acarina,
Trombiculidae)," Psyche, 55:87-100.
42. Williams, Roger W., 1944. "A bibliography pertaining to the mite
family Trombidiidae," Am. Midland Naturalist, 2:699-712.
43. Jenkins, Dale W., 1948. "Trombiculid mites affecting man. I. Bionomics
with reference to epidemiology in the United States," Am. J. Hyg., 48:22-35.
44. Michener, Charles D., 1946. "Observations on the habits and life
history of a chigger mite, Entrombicula batatas (Acarina: Trombiculinae) ,"
Ann. Entomolog. Soc. Amer., 39:101-18.
45. Miller, A. E., 1925. "The native host of the chigger," Science, 60:345-
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46. Keay, Gladys, 1937. "The ecology of the harvest mite (Trombicula
autumnalis) in the British Isles," I.Animal Ecology, 6:23-35.
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Washington, D. C.: Dept. Agric., in Bull. no. 986.19 pp.
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Natural History, 56:231-35.
49. Williams, Roger W., 1946. "A contribution to our knowledge of the
bionomics of the common North American chigger, Entrombicula alfreddugesi
(Oudemans), with a description of a rapid collecting method," Am. 1. Trap.
Med., 26:243-50.
50. Blake, Francis G.; Maxcy, Kenneth F.; Sadusk, Joseph, F., Jr.; Kohls,
Glen M.; and Bell, E. John; 1945. "Studies on Tsutsugamushi disease (scrub
typhus, mite-borne typhus) in New Guinea and adjacent islands: Epidemiol-
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7:53-68.
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Microbes Associated with Insects and Ticks with Special Reference to the
Biologic Relationships Involved. Ithaca, N. Y.: Comstock Publishing Co., Inc.
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in World War II," J. Parasitol., 34:169-91.
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Internat. Congo Trop. Med. and Malaria. (Abstracts) Washington, D. C.
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"Akamushi" group," J.Parasitol., 33:260-64.
58. Goldberger, J., and Schamberg, J. F., 1909. "Epidemic of an urticarioid
dermatitis due to a small mite (Pediculoides ventricosus) in the straw of mat-
tresses." U. S. Public Health Service, Pub. Health Rep., 24:973-75.
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Washington, D. C.: Dept. Agric., in Bur. Entomol. Circ., no. 118. 24 pp'.
60. Hinman, E. H., and Kampmeier, R. H., 1934. "Intestinal acariasis due
to Tyroglyphus longior Gervais," Am. J. Trap. Med., 14:355-62.
61. Rebrassier, R. E., and Martin, E. D., 1932. "Syringophilus bipectinatus,
a quill mite of poultry," Science, 76:128.
62. Sambon, L. W., 1922. "A synopsis of the family Linguatulidae," J.
Trop. Med., 25:188-206, and 391-428.
63. Heymons, R., 1935. Pentastomida. Dr. H. G. Branns. Klassen und
Ordnungen des Thierreichs. 5 Band: Arthropoda. iv. Abteilung Arachnoidea.
1 Buch Pentastomida. 268 pages (148 text figures) .
64. Faust, E. C., 1927. "Linguatulids (Order Acarina) from man and
other hosts in China," Am. J. Trop. Med., 7:311-22 + 2 plates.
65. Hobmaier, A., and Hobmaier, M., 1940. "On the life-cycle of Lingua-
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South. Calif. Acad. Sc., 33:117-22.
CHAPTER XXII
._.
VENOMOUS AND URTICARIAL ARTHROPODS
Insect venoms. Insect venoms, like other animal venoms, are toxic
principles probably not greatly unlike the bacterial toxins, but about
which we know comparatively little. Unlike many of the bacterial toxins
which reach toxic proportions only after a period of elapsed time sub-
sequent to the introduction of the infection into the body, the animal
venoms take effect almost instantly, i.e., as soon as introduced and
without incubation.
The venoms act in one or more ways when introduced into the body:
( 1) they may act directly on the blood corpuscles (hemolytic); (2)
they may act directly on the nervous system, producing shock or inhibit-
ing reflexes (neurotoxic); (3) they may produce an infiltration and con-
gestion of blood (hemorrhagic) often in the vicinity of the wound or in
deeper tissue, such as the mesenteries, etc. A given arthropod venom
may act in one or more of these ways.
It is a matter of common observation verified by various investigators
that repeated inoculation of minute or attenuated quantities of a venom
may lead to a degree of immunity, so it is also with the venoms or poi-
sons of bees, bedbugs, mosquitoes, fleas, conenoses, etc. Newcomers to
a flea-infested community may suffer great misery until they have ac-
quired an "immunity"; this may require many months and may never
be acquired by some individuals. Promising results have been obtained
by administering a course of immunization consisting of injections (six)
of a flea antigen1 [extract of Pulex irritans (75 per cent) and Ctenocephal-
ides canis (25 per cent)] given subcutaneously one week apart.
In the ants, bees, and wasps (aculeate Hymenoptera) there are two
poison-secreting glands, one of which is said by some authors to secrete
formic acid, and the other an alkaline fluid. The combination of the two
agents in certain proportions is believed to be necessary to produce
lethal effect on other insects. Koszalka 2 points out that "bee venom is
considered to be a complex organic compound, containing lecithin and
an albumin-free sapotoxin, which produces histamine-like effects and
has a potential hemolytic action."
The scorpion (an arachnid) secretes a large quantity of colorless
563
acid-reacting liquid soluble in water and heavier than water. According
to Calmette, less than 0.0005 gm from Buthus afer Leach will kill a
white mouse in about two hours.
How the venom is introduced. Arthropod venoms are introduced into
the body of man and other animals in one of three ways: (1) by con-
tact e.g., by means of urticarial hairs of certain caterpillars, such as the
brown-tail moth, Nygmia phaeorrhoea Don. [= Euproctis chrysorrhoea
( Linn. ) ], producing a condition similar to nettling, or with vesicating
fluids of the blister beetles (Meloidae), particularly Lytta vesicatoria
(Linn. ), resulting in a vesicular dermatitis; (2) by the bite, thrust of
piercing mouth parts, as in the conenoses (Reduviidae), or penetration
of the chelicerae of spiders; (3) by the sting, as in the bees and wasps
(aculeate Hymenoptera) and the scorpion. The operation and structure
of the sting of insects and arachnids differ considerably.
Stinging insects. The stinging insects belong to the order Hymen-
optera, suborder Aculeata, the ants, bees, and wasps, in which the
females of all species are provided with a specialized ovipositor known
as a sting, more or less well developed for piercing the skin of higher
animals or other insects. The sting is used either as an organ of defense
or offense; in the latter case often to procure food for the young. The
venom apparatus of the aculeate Hymenoptera, such as bees, wasps,
hornets, and bumblebees, does not differ greatly in structure.
The principal aculeate Hymenoptera are divided into the following
super-families, viz.: Formicoidea, the true ants; Sphecoidea, the digger
wasps; Vespoidea, the true wasps; and Apoidea, the bees.
Morphology of bee sting. The sting of the worker bee is regarded as
a specialized ovipositor; it originates from the seventh and eighth seg-
ments and lies between the oviduct and the rectum above. The darts
of the sting follow the ventral line of the abdomen and are held in
place by the sheath situated above, while the barbs of the darts point
oownward and outward. In a space above the sheath lie the fleshy palpi
(Fig. 179). The delicate attachment between the sting and the organs
of the abdomen accounts for the ease with which the sting is torn from
the abdomen when the barbs become embedded after the darts are
thrust into the skin. The sting can be easily extracted either by separating
the segments of the abdomen fIJJID it by means of dissecting needles, or
by squeezing the live bee between forceps, which causes it to protrude
the sting. The sting can then be grasped with other forceps and drawn
out. After extraction, the sting can be best examined when the parts are
floated out in a few drops of glycerin. The sting may be divided into
three parts, viz.: the piercing apparatus; the lateral plate and append-
ages; the poison sac and glands.
The piercing apparatus itself consists of three parts, one the so-
called sheath, the other two lying within the ~heath, and partially sur-
rounded by it. In appearance the sheath is yt~llowish and translucent.
The darts, which present concave surfaces to one another, are highly
chitinous. 'fbe dista1 one-tbird of t11e dart possesses a series of sharp
l
J
J
/r ,
1
(t)-:,'
"
"
?~~
i "
I.'
FIG. 179. Sting of a honeybee, Apis mellifera: a, the two serrated darts;
b, b', sting palpi; c, venom (poison) sac; d, verJ 0m gland; e, e', triangular
plates or levers; f, 1', semilunar plates or levers; g, g', lateral plates or levers ;
h, h', Y-shaped darts; i, i', points of attachrJ1 ent for darts to levers;
e, e', i, r,
pOints around which levers rotate; k, k', points of attachment for
lever ct, f'). X 17.5.
barbs, whose shape has been aptly compared to the tip of a crochet
needle. Cheshire states that each dart has fron'l three to six barbs, other
writers seem doubtful as to the number. MaflY darts have been care-
fully examined by the writer, but in no instance were less than ten barbs
distinguished on tbe outer edge of each dart (Fig. 179). Several writers
state that poison pores are to be found at the base of each dart, from
which poison exudes. In this matter the writer agrees with Snodgrass, as
he has failed to observe the exit of poison elsewhere than between the
darts at their tip.
Proceeding upward on the dart from the tiny barbs, the darts are seen
to form a "Y" as they lie within the sheath. The arms of the "Y" grad-
ually bend laterally. Plates are attached to the npper edges of these
laterally bent arms. One of the most remarkable portions of the darts is
the poison valve with which each is provided. At the point of separa-
tion the darts each present a delicate cup-shaped valve, whose closed
portion is directed downward toward the tip of the sting. This is formed
of the same chitinous material as the darts, and each is free to move
with the movement of the dart. In order to accommodate this enlarge-
ment of the darts, the sheath at this point expands to about five times its
smallest diameter, which is at the tip of the sting. For at least one-third
of its length the sheath at this portion is expanded into a symmetrical
oblong body providing ample room for the movement of the darts and
valves within.
A curious structure, said by many writers to be found on the sheath,
consists of two delicate, but strong, chitinous tracks or guide rails on
which the darts, correspondingly grooved, fit and move back and forth.
Since the sheath does not sufficiently surround the darts to direct their
course, this guide-rail system which Carlet has observed, and which is
accepted by other authors, probably explains why the darts move
smoothly and accurately within the sheath.
The lateral appendages are of three kinds: semilunar, triangular, and
lateral, according to shape or position. Both the semilunar and triangular
plates are attached to the bent ends of the Y-shaped darts. The triangular
plates are attached to the arms of the darts almost at their extremities,
while the semilunar ones are connected for about one-third of the dis-
tance from the ends of the arms. The apex of the triangle is attached
to the extremity of the dart. The other two points are directed outward
and downward, and serve as points of attachment for two elevated
edges on the lateral plates which hang thus suspended. As they hang,
half of their surface lies above and covers the dorsal surface of the semi-
lunar plates just beneath them. Continuing in the same straight line with
the semilunar plates and attached at their extremity to them, lie the
fleshy palpi covered with delicate hairs.
The third set of structures which completes the sting are the venom
sac and glands. In order to understand these it is necessary to know
that Hymenoptera are divided into two groups, those which kill their
prey by stinging, and those which only paralyze it. The former are the
more complicated, for they possess two poison glands; the acid gland,
which opens directly into the great poison sac, the larger of the two,
and the other, the alkaline gland, which is comparatively small and is
situated at the base of the poison sac. It is the combination of the acid
and alkaline fluids from the two glands that results in the death of the
attacked insect, or that causes the extreme pain and resulting reactions
in humans.
The formic acid gland alone is found in those Hymenoptera which
only paralyze their prey by their sting. This fact has led various observ-
ers to make chemical tests of both the formic acid and alkaline substance.
The result, according to Carlet and others, has been to show that neither
substance by itself is effective except to paralyze, but when combined
the substances have deadly effects upon other insects. Carlet's experi-
ments to prove this were made on houseflies and blowflies by injecting
each substance singly and then introducing both into the body of a fly.
Operation of the sting. The sting may be observed in operation by
confining the bee on its back and then prodding it until its sting is
angrily thrust in and out. This process shows three things, viz.: that the
sharp-pointed sheath always appears first when the thrust is made;
second, that the darts inside the sheath work back and forth alternately
and quite independently of the sheath or of one another; third, that the
poison exudes in droplets from the tip of the sting between the darts.
Cheshire (Bees and Bee-Keeping, London, 1886) states that
The sheath has three uses: first, to open the wound; second, to act as an
intermediate conduit for the poison; and third, to hold in accurate position the
long-barbed darts. The sheath does not inclose the darts as a scabbard, but is
cleft down the side which is below, when the sting points backward. The darts,
as soon as their ugly barbs establish a hold, first one and then another drive
back and forth by successive blows. These in turn are followed by the sheath,
when the darts again plunge more deeply, until the murderous little tool is
buried to the hilt. But these movements are the result of a muscular apparatus
yet to be examined. The dovetail guide-rails of the sheath are continued far
above its bulbous portion, and along with these the darts are also prolonged up-
ward, still held to the guides by the grooved arrangement; but both guides and
darts, in the upper part of their length, curve from each other like the arms of
the Y, before mentioned, to points C, C' (Fig. 179) where the darts make
attachment to two levers (i, i'). The levers, or plates, as they are called (Kl and
K'ln), are provided with broad muscles, which terminate by attachment to the
lower segments of the abdomen. These, by contraction, revolve the levers
aforesaid round the points f, f, so that without relative movement of rod and
groove, the points c, c' approach each other. The arms of the Y straighten and
shorten, so that the sheath and darts are driven from their hiding place to-
gether and the thrust is made by which the sheath produces its incision and
fixture. The sides being symmetrical, we may, for simplicity's sake, concentrate
our attention on one, say the le~t in the figure. A muscular contraction of a
broad strap joining K and D (the dart protractor) now revolves k on 1, so that
a is raised, by which clearly c is made to approach d; that is, the dart is sent
forward, so that the barbs extend beyond the sheath and deepen the punc-
ture. The other dart, and then the sheath, follow, in a sequence already ex-
plained, and which G is intended to make intelligible, representing the entrance
of the sheath, b the advance of the barbs, and c the sheath in its second posi-
tion. The barb retractor muscle is attached to the outer side of i, and by it a
is depressed and the barbs lifted. These movements, following one another
with remarkable rapidity, are entirely reflex, and may be continued long after
the sting has been torn, as is usual, from the insect. By taking a bee under
the microscope and forcing the sting into action, the sting movement will be
seen to be kept up by continued impulses from the fifth abdominal ganglion
and its multitudinous nerves, which penetrate every part of the sting mechan-
ism and may be traced even into the darts. These facts will show why an ab-
domen separated many hours may be able to sting severely.
Reaction to bee stings. The painful effect of the sting of bees is prob-
ably not due to the simple thrusts of the barbs already described, but
more particularly to the introduced venom. The severity is greatly in-
creased in the case of multiple stings; however, one sting in a previously
sensitized person or one inHicted at a vital point (e.g., a vein) may
result fatally. Pawlowsky, according to Martinj,3 estimates 500 bee stings
within a short time as a lethal dose for man. Tolerance to bee stings
appears to be developed in beekeepers, long exposed to stings; how-
ever, this tolerance seems to be lost when exposure to stings is discon-
tinued.
Allergic reactions are generally recognized as due to bee protein
and pollen adhering to the bee. Death may result from anaphylactic
shock.
Treatment for bee sting. If the sting remains in the wound, it must
be removed quickly and carefully so as not to force additional venom
into the puncture. Repeated and long-continued application of a bicar-
bonate of soda paste (soda with. water enough to make a paste) is
recommended. Eau de Javelle 1 to 100 is also recommended, as well as
calcium chloride in water 1 to 60; application of ammonia or alcohol will
also give some relief. If symptoms of envenomization arise, a phYSician
should be consulted. Kozalka (see above) has recently (1949) reported
rapid recovery from multiple bee stings attributable to the parenteral
use of calcium gluconate.
YeHow jackets or hornets such as Vespula ( = Vespa) diabolica
(Sauss. ), Vespula (= Vespa) maculata (Linn.), and Vespula pennsyl-
vanica (Sauss.) are members of the family Vespidae (Order Hymenop-
tera) . There are also several European species, among them Vespa
crabro Linn., Vespa vulgaris Linn., and Vespa germanica Fabr. All of
these species build nests (Fig. 180) of pulp made by masticating wood
fiber. Nests are built in hollow trees, among the branches, under eaves
of houses, and in holes in the ground. The somewhat elongate aerial
nests may reach nine to ten inches in diameter.
The young fertilized queens hibernate throughout the winter am]
start the nest in the spring. In the control of these insects one should
seek out the new nests under construction and destroy them by night
Cw . 180. Aerial nest of yellow jacket; side and top view of the insect, Vespula
diabolica; and longitudinal section of nest below.
when the queen and her progeny are at home. The aerial nests may be
cautiously and quickly enveloped at night with a tight drawstring bag
and then soaked with kerosene, scalding water, or burned. The under-
ground nests should be located by day and the entrance marked, then
treated by pouring an ounce or two of carbon bisulfide into the hole at
night and quickly pressing the entrance shut with the foot. An old-
fashioned yet effective and safe way to handle a yellow jacket problem
is to place a gallon jug, quarter filled with water, near the nest. As these
insects investigate this new object, the buzzing sound echoing from the
jug prompts them to enter, and as a result fall into the water and are
drowned. In this way one by one the entire colony is lured into the jug
(Melander) .
The small nests with exposed cells made by such forms as Polistes
bellicosus Cresson may be removed at night by quickly placing a tin can
or box with open end over the nest and then slipping a sheet of tin be-
tween the surface and the open end of the can so that the nest is cut
away. The surface from which the nest has been removed may be
daubed with creosote to repel other female hornets from starting a nest
on the same site.
Mud daubers or thread-waisted wasps such as Sphex (Psammophila)
luctuosa Smith, Sphex violaceipennis Lepeletier, Chalybion coeruleum
(Linn. ), and Pelopaeus californicus Sauss., make their nests, usually
quite small, of mud and provision them with insects and spiders to serve
as food for the larvae. These mud nests may be removed as suggested
above for Polistes.
Ants. The true ants belong to the family Formicidae (Order Hymen-
optera), which comprises about 8,000 described species, divided into five
or more subfamilies, e.g., Formicinae, Ponerinae, Dorylinae, Myrmi-
cinae, and Dolichoderinae.
The subfamily Myrmicinae includes several formidable stinging
species of pugnacious nature. Because of their numbers a mass attack
may result seriously. Among the more dangerous species are: (1) the
California or Mexican fire ant, Solenopsis xyloni var. maniosa Wheeler,
the workers of which have a yellowish-red head and a black thorax and
abdomen. They measure from 1.6 to 5.8 mm in length. (2) The Texas
harvester or agricultural ant, Pogonomyrmex barbatus (F. Smith), in
which the head, thorax, and legs are black and the abdomen red; the
workers reach from 7 to 9 mm in length; (S) the California harvester
ant, PogonomYl'mex californicus (Buckley), with a light rusty-red body,
and legs somewhat more yellowish. These ants will readily attack
humans and smaller animals. Young pigs may be killed by the stings of
this species. A small pig may walk leisurely upon an ant mound and
suddenly begin to kick and squeal, due to the terrific attack of ants
rushing forth from the nest. The animals commonly topple over with
legs outstretched and death may ensue.
Ants belonging to the subfamily Ponerinae also have well-developed
stings and a potent venom. This is apparently particularly true of the
Central and South American species, Paraponera clavata (Fabr.), which
is common in high tropical rain forests. Weber4 states that the ants of
this species boil out of their nests in large numbers when disturbed and.
rush for the intruder. He states that the workers are fully an inch in
length and blackish brown in color. The ant is greatly dreaded by the
natives.
For the control of the fire ant, Mallis 5 recommends the use of a mix-
ture of one part of carbon bisulfide and three parts of carbon tetrach-
loride injected into the nest openings by means of an oil can with an
eight-inch curve-tip spout. A few drops of the liquid in each opening
are sufficient. After treatment the openings must be covered. A large
measure of control may be effected by applying kerosene to the nests,
using a funnel or hollow rod to reach the deeper parts; potassium
cyanide in liquid form may also be used, but great care must be exer-
FIG. 181. A velvet ant, Mutillidae, also known

as a "cow killer." x2.2.
cised both in its preparation and application owing to its very poisonous
nahlre.
Mutillid wasps. Among the less known stinging insects are the mutillid
wasps belonging to the family Mutillidae (Order Hymenoptera). Mem-
bers of this family are commonly known as velvet ants, woolly ants, cow
killers, mule killers, etc. (Fig. 181). The mutillids are covered with a
velvety pubescence; many are brightly colored with orange or red or
yellow. The females are apterous, good runners, and may inflict a pain-
ful sting. They are parasites of bees and other wasps. There are very
many species, some of the commoner forms measuring from 12 to 1
inch in length. Our knowledge concerning these interesting insects has
been greatly advanced by Mickel. 6 A very common species in the cen-
tral United States is Sphaerophthalma occidentalis (Linn.), a black
species with a scarlet band. This species is very common on the beach
sands of Lake Erie, causing barefoot bathers much distress.
Stinging Eypris. In 1927 von Geldern 7 reported a tiny wasp from
Yolo County, California, identified as belonging to the genus Eypris
which inflicts a severe sting.
The wasps appear in fairly great numbers in the fall after a warm spell and
invade the house where they get into the bedding and clothing, and sting
when brushed or crushed by clothing or sheets against the skin. . . . The sting
is distinctly felt as a fairly sharp prick, decidedly less intense than a bee
sting. . . . In the oldest and youngest child no further manifestations occur,
but in the parents and second child a decided systemic disturbance follows.
A few minutes after being stung, there is felt a numbness, often at the site of
the sting, but at other times beginning at the finger tips. It remains localized for
a few minutes and then gradually spreads and involves the entire body. In
the mother there is an intense itching of the vulva and in the father an itching
of the pubes. This is followed by a marked diarrhoea, not painful in the father,
but resembling severe uterine cramps in the mother. The diarrhoea and cramps
last for about ten minutes. The mother, who is an asthmatic, experiences no
respiratory difficulty, but with the father, who has never had an attack of
asthma, wheezing occurs occasionally. Accompanying these symptoms there is
marked prostration, weakness and sweating. The duration of the attack is
about half an hour. The second child becomes drowsy and is awakened with
difficulty and wheezing occurs. He also recovers in about the same time as the
parents.
EssigB in 1932 reported a number of instances of Epyris stings all

from the same county in California as were the cases reported by von
Geldern. The species of wasp was identified as Epyris californicus (Ash-
mead) belonging to the proctotrupid family Bethylidae. It measures
barely over 5 mm in length and is black in color. Essig states, "concern-
ing the life history and habits of this particular species, beyond its pro-
pensity for stinging, absolutely nothing is known."
BITING (PIERCING) INSECTS

Insects that pierce the skin with their mouth parts are usually nor-
mally bloodsuckers and the act of biting or piercing is simply a part of
the act of food-getting. There are noteworthy exceptions, as later ex-
plained. The pain caused by the mechanical insertion of such tiny
mouth parts would be, no doubt, relatively benign, particularly if only
one or only a few individual insects were concerned in the attack; how-
ever, in perhaps every instance a venom of salivary origin is introduced.
These venoms apparently differ among the various species, as evidenced
by the resulting reactions, local and systemic, which are generally speCific
enough so that one who is experienced may be able to determine the
cause, Le., whether the offender was a bedbug, a flea, a mosquito, or a
black fly (simuliid). Rubbing the bite with cotton saturated with chloro-
form or carbon tetrachloride will relieve the irritation.
To understand the operation of the bloodsucking mechanism of the
various offending insects one should consult the previous chapter on
mouth parts and the other chapters appropriate to this subject. The
VENOMOUS AND URTICARIAL AHTHROPODS 573
student will profit much by a careful study of "Zoonosen der Haut in
warmeren Landern" by Martini (loc. cit.). In general the reactions are
the result of both mechanical and chemical processes.
Some individuals apparently suffer no ill effect from bedbug bites,
not even a swelling at the site of the bite; others react violently to even
one bite. These differences in tolerance to a given species are not fully
understood. Martini remarks that doubtless allergic processes play a
role in these reactions.
Insects that inflict a very painful bite, such as the stable fly, Stomoxys
calcitrans (Linn.), and most salt-marsh mosquitoes, e.g., Aedes dorsalis
(Meig. ), are apparently seldom vectors of disease, while species with
benign bites, such as Anopheles maculipennis (Meig.), are commonly
potent vectors. May it not be that in order to become a successful vector
of disease, the arthropod must first modify the severity of its bite!
Conenoses or kissing bugs, belonging to the family Reduviidae (see
Chapter VIn), commonly inflict painful "bites:' Their mouth parts (see
Chapter VI) are well adapted for piercing the skin of the host. Reduviids
are essentially predaceous, attacking many species of insects, particularly
soft-bodied forms from which they suck the body fluids. Attack upon
humans is made principally, if not wholly, in self-defense. Persons pick-
ing up boards, sticks, or stones, etc., may accidentally pick up one of
these insects; likewise in plucking a leaf or flower from a tree or other
plant the fingers may close upon the insect as well, with the result that
a very painful bite maybe suffered.
The principal offenders are about 18 to 20 mm in length, and all bear
a general resemblance to the illustrations, Figs. 40 and 41. Among the
species inflicting painful bites are the following: Reduvius personatus
(Linn. ), known as the "kissing bug"; Triatoma sanguisuga (Lec.), the
"bloodsucking conenose" or "big bedbug"; Triatoma protracta (Uhler),
the "China bedbug"; and Rasahus biguttatus (Say), the "two-spotted
corsair."
The symptoms produced· by Triatoma protracta (Uhler), the usual
offender in California, are described as follows: "In a few minutes after
a bite the patient develops nausea, flushed face, palpitation of the heart,
rapid breathing, rapid pulse, followed by profuse urticaria all over the
body. The symptoms vary with individuals in their intensity."
The symptoms described for Rasahus biguttatus (Say) are as follows:
"Next day the injured part shows a local cellutitis with a central spot;
around this spot there frequently appears a bulbous vesicle about the
size of a ten-cent piece and filled with a dark grumous fluid; a smaller
ulcer forms underneath the vesicle, the necrotic area being generally
limited to the central part, while the surrounding tissues are more or
less swollen and somewhat painful."
Biting water bugs. The order Hempitera (see Chapter VIII) contains
a number of families of aquatic forms several of which included biting
species; among these are the families Belostomatidae and Notonectidae.
The giant water bugs Lethocerus, Belostoma, and Benacus, belonging
to the family Belostomatidae, are among the largest of the water bugs,
measuring 2~ to 4~ inches in length and possessing formidable beaks.
They feed on other aquatic insects, also young frogs, fish, etc., and since
they are winged and readily attracted to lights, they are commonly
known as electric light bugs. They have been known to attack birds.
Ewing 9 describes the effect of the bite as follows:
. . . at 9:30 A.M. a giant water bug Benacus griseus (Say) was allowed to
bite the back of the right index finger. The beak was left inserted for a few
seconds. Immediately a burning sensation followed. Two minutes later the
same bug was allowed to puncture the back of the left index finger for several
seconds. A burning sensation was produced. Soon some swelling was noted,
and a reddened area developed about the point of the puncture. Pain con-
tinued but diminished during the forenoon and by noon the reddened area had
become reduced. By 1:30 P.M. a small red spot was all that was left at the
puncture. . . . When Benacus griseus bites it emits a milky fluid from the tip
of the beak, and the beak adheres to the skin after penetration, so that the
skin is pulled up when the beak is withdrawn.
Back swimmers belonging to the family Notonectidae may also in-
flict a painful bite. These predaceous bugs swim on their backs, hence
the common name "back swimmers." The bite is nearly as severe as a
bee sting.
Bloodsucking phytophagous bugs. Numerous instances of bloodsucking
among phytophagous Hemiptera have been reported. Much information
concerning these cases has been assembled by Usinger (1934, lac. cit.)
and Myers 10 (1929).
Among the species exhibiting this fortuitous bloodsucking behavior
are members of the following families: Membracidae (treehoppers)
such as Ceresa bubalus (Fabr.); Cicadellidae (leafhoppers and sharp-
shooters) such as Eutettix tenellus (Bak.) and Erythroneura comes
(Say); Miridae, such as Jrbisia solani (Heid.) and SOpidae marginata
Uhler; Corizidae, Leptocoris trivittatus (Say).
Usinger remarks that the change from the sucking of plant juices to
bloodsucking at first appears to be very great. "However, upon compari-
son of the chemical constituents, it is found that in general the same
elements are found in plants as in blood and often in very similar com-
binations although in very different proportions."
Thrips biting man. Thrips (Order Thysanoptera) are minute plant-
feeding (sapsucking) insects (see Chapter VI for description of mouth
parts); however, there have been numerous reports of their attacking
man and of their ability to suck blood. Baileyll states that while working
on experimental plots he experienced bites from the onion thrips, Thrips
tabaci Lind. He felt slight pricks on the arms, face, and neck, both when
perspiring and when not. He observed that the larvae (second instar)
were more prone to bite than the adults and that the alimentary canal
took on a reddish-brown appearance after feeding. Small pinkish dots
appeared on the skin and disappeared in one to two days. No swelling
occurred but there was a slight itching sensation. He had similar ex-
periences with the pear thrips, Taeniothrips inconsequens (U zel).
Several other species of thrips have been reported in a similar con-
nection, e.g., Heliothrips indicus Bagnall, a cotton pest of the Sudan;
Thrips imaginis Bagnall, reported for Australia; Limothrips cerealium
Haliday, for Germany; Gynaitkothrips uzeli Zimmerman, for Algiers;
and other species. It would appear that many species of thrips are thus
involved and that this behavior is not restricted to anyone species.
Urticarial hairs. The caterpillars of many species of Lepidoptera, (at
least eight families) possess urticating hairs. Among the families which
have urticarial larvae is the Satumiidae of which the genus H emi-
leuca is especially offensive. H emileuca oliviae Cockerell, the range
caterpillar, is reported to be a menace to cattlemen in New MexicoY
A rash known as the "brown-tail rash" is traceable to the caterpillar of the
brown-tail moth (Nygmia phaeorrhoea Don.), a common and very de-
structive shade tree pest in Europe and in America, espeCially in New
England. When the caterpillars of this species molt, myriads of tiny
barbed hairs are shed with the skin. The cocoons of the pupated cater-
pillars as well as the adult moths possess these hairs. These hairs are
blown about by the wind and coming in contact with the skin of the
neck, face, hands, or other exposed parts of the body produce a very
severe dermatitis. The hairs are hollow, and it has been shown by
Tyzzer13 that they contain a definite poisonous principle which is in-
jected into the circulation by the sharp-pointed hair in contact with the
skin, thus producing the rash. Ingestion of the hairs by swallowing or
inhaling in breathing may cause serious internal disturbances.
Bishopp14 describes the symptoms produced by contact with the
"puss" caterpillar (Megalopyge opercularis S. & A.) as follows:
Almost immediately after any portion of the body comes in contact with
one of these caterpillars an intense burning pain is felt, described by some as
similar to a severe nettle sting. This usually becomes worse accompanied by
itching for several minutes and persists from 1 to 12 hours and sometimes
longer. Almost immediately after a sting the area touched by the caterpillar
shows minute raised whitish spots or papules which soon become red, followed
by spreading of the inflammatory area for several inches and often accom-
panied by general swelling of the portion of the body stung. Stings on the wrist
have been followed by a swelling of the entire arm to almost double its normal
size. A feeling of numbness which almost assumes the characteristics of paraly-
sis accompanies the swelling. This is usually confined to the member attacked
but may be generalized. Apparently stings on the neck are even worse, as the
writer has one record of a man who was stung severely on the neck and com-
pletely incapacitated, being confIned to the hospital for 6 days. These paralytic
symptoms are often accompanied by nausea and sometimes by vomiting. The
stings are especially severe among young children, who often develop consider-
able fever and nervous symptoms. These sometimes last for a day or two and
are accompanied by nausea, especially during the first few hours. Usually with-
in two or three hours after a sting, the reddened pimple-like swellings at the
site assume the appearances of small vesicles or blisters. These usually persist
for a few hours and then apparently harden through absorption, leaving a
roughened area. In some instances the discoloration of the skin surrounding the
point of attack is rather marked, varying from a deep red to almost black. The
paralytic symptoms usually subside with the pain, but the local lesions often
persist for several days.
Students concerned with the subject of urticarial hairs will need to
consult Weidner's'5 (1936) work which includes a comprehensive bibli-
ography on poisonous insect hairs. Weidner lists the following families
of Lepidoptera which include caterpillars with hairs causing skin irrita-
tion; namely, Morphidae, Morpho hercules Dalm.; Arctiidae, Lithosia
caniola Rbn., L. griseola Rbn.; Lymantriidae, Nygmia phaeorrhoea Don.,
[= Euproctis chrysorrhoea (L.)], Porthesia similis Fuessly, and several
others; Thaumetopoeidae, Thaumetopoea pinivora Tr., Anaphe infracta
Wlsgh., and others; Lasiocampidae, Macrothylacia rubi (L.), Dendroli-
mus pini (L.), Lasiocampa quercus (L.), and others; Noctuidae, a few
species occasionally cause irritation; Nymphalidae, larval hairs may
pierce the skin, e.g., Vanessa io (L.) and Hamadryas antiopa (L.)
(mourning cloak); Saturniidae, Automeris io (Fabr.) (10 moth), Hem-
ileuca maia Drury (buck moth), and others; Megalopygidae, Lagoa
crispata Pack. (white flannel moth), and others; Limacodidae, Sibine
stimulea Clem. (the saddle-back caterpillar), and others.
Blister beetles. Blister beetles belong to the family Meloidae (Can-
tharidae) Order Coleoptera and are so deSignated because of their vesi-
cating properties, i.e., the application of the pulverized bodies or even
simple contact with the insect may produce a blistering of the skin.
The Meloidae (Cantharidae) are described by Comstock: "The blis-
ter beetles are of medium or large size. The body is comparatively soft;
the head is broad, vertical and abruptly narrowed into a neck; the pro-
thorax is narrower than the wing covers, which are soft and flexible; the
legs are long and slender; the hind tarsi are four-jointed, and the fore
and middle tarsi are five-jointed."
The blister beetles deposit their eggs on the ground; the larvae are
active and feed, it is said, in some species on the eggs of locusts and
solitary bees; others are predaceous. They undergo a number of changes
not usual to insects, their 'development being termed "hypermetamor-
phoSiS." The adults are vegetable-feeding.
Spanish fly. The Spanish fly, Lytta vesicatoria (Linn.), is a Euro-
pean species of beetle found most abundantly during the early summer
in Spain, southern France, and other parts of Europe (Fig. 182). It is
golden green or bluish in color, and ranges from one-half to three-quar-
ters of an inch in length. It makes its appearance quite suddenly in early
summer, when it may be collected by the hundreds, clinging prinCipally
to such vegetation as ash, privet, and lilac. The peculiar hypermetamor-
A B
FIG. 182. The Spanish fly, Lytta vesicatoria; (A) female , (B) male.
phosis of these insects and the subterranean predaceous larval habits give
to them some obscurity during their early development; the sudden ap-
pearance and equally sudden disappearance, owing to short adult life,
give rise to the belief that they are migrating forms .
Cantharidin, a crystalline prinCiple, the anhydrid of cantharidic acid,
was isolated by Robiquet in 1812 from the Spanish fly, Lytta vesicatol'ia
(Linn. ). Cantharidin penetrates the epidermis quite readily and pro-
duces, even in very small quantity (7i 0 mg) violent superficial irritation,
resulting in vesication (blistering) in a few hours. Even when applied
to the skin, cantharidin irritates the kidneys so that "fly blisters" are
contraindicated in nephritis. It was formerly used as an aphordisiac, but
its effects may be dangerous to life; hence its use for this purpose has
been largely discontinued.
The collection and preparation of the beetles for medicinal purposes
provides an occupation for many persons for a brief period. Collecting
and preparing the insects requires special precautions owing to their
vesicating properties. The best quality of cantharidin produced from the
pulverized beetles is the result of special care in the drying, which must
be gradual.
Other blister beetles, causing severe vesicular dermatitis in Africa,
are Mylabris nubica de Marseul, Epicauta tomentosa Maeklin, and
Epicauta sapphirina Maeklin, according to Chalmers and King. 16 Also
Paederus crebripunctatus Epp. is reported to be a severe vesicating
beetle of East Africa 17 affecting Europeans and Africans similarly, al-
though less severely on habitually exposed parts of natives. The term
"Nairobi eye" applies to the conjunctivitis caused when the juices of
crushed beetles are rubbed into the eye. The active principle is can-
tharidin. Roberts and Tonking17 (loc. cit.) recommend a cold compress
of saturated solution of magnesium sulfate.
At least two species, Sessinia collaris (Sharp) and Sessinia decolar
Fairm., belonging to the family Oedomeridae, cause severe blistering
on some of the mid-Pacific Islands where they are called coconut
beetles. Is These beetles fairly swarm about the newly opened male
flowers of the coconut where they feed on pollen. They are readily at-
tracted by light. Corning in contact with one of these beetles causes a
sharp momentary pain, like a burn from hot oil, but the large blister
which forms in a few hours causes little pain.
SPIDERS
Class Arachnida-Order Araneida (Araneae)
General characteristics. Spiders are arachnids in which the prosoma
is uniform, bearing not more than eight eyes, and joined to the opistho-
soma by a pedicle. The opisthosoma is usually unsegmented and bears
not more than four, usually three, pairs of spinnerets. There is no telson.
The chelicerae are two-segmented, moderately large and unchelate, and
contain a poison gland. The pedipalps are six-segmented, leg-like and
tactile in function. The legs consist of seven segments; the tarsi have
two or three claws. Respiration is by book lungs or trachea or, normally,
both. The pedipalps of the male are modified as intromittent organs
(Savory, lac. cit. p. 59).
Though spiders are universally feared, no doubt because of the
knowledge that they are able to kill insects and other small animals by
introducing a venom with the bite, it is nevertheless true that out of the
more than two thousand genera in more than thirty families only a very
few species are actually dangerous to man.
Tarantulas. The term tarantula was first applied to a European spe-
cies, Lycosa tarantula (Linn.), a member of the family Lycosidae (wolf
spiders). In Italy in the vicinity of Taranto there occurred a spider scare
during the seventeenth century which gave rise to a condition known as
"tarantism" resulting from the bite of this species of spider. To rid the
body of the venom those bitten engaged in a frenzied dance known as
the "taranteIla."19
The following account of tarantism is from the Cambridge Natural
History, Vol. 4, p. 361:
The bite of the spider was supposed to induce a species of madness which
found its expression-and its cure-in frantic and extravagant contortions of
the body. If the dance was not sufficiently frenzied, death ensued. In the case
of survivors, the symptoms were said to recur on the anniversary of the bite.
Particular descriptions of music were supposed to incite the patient to the
excessive exertion necessary for his relief; hence the name "Tarantella."
In the middle ages epidemics of "tarantism" were of frequent occurrence
and spread with alarming rapidity. They were seizures of an hysterical char-
acter, analogous to the ancient Bacchic dances, and quite unconnected with the
venom of the spider from which they took their name. The condition of exalta-
tion and frenzy was contagious, and would run through whole districts, with
its subsequent relapse to a state of utter prostration and exhaustion. The evil
reputation of the Tarantula appears to have exceedingly little basis in fact.
In the southwestern United States the term "tarantula" is applied to
the very large spiders belonging to the family Aviculariidae, also known
as "bird spiders." Many of these spiders measure about five inches in
spread of legs. Among these is Eurypelma californica Ausserer; Baerg 20
. reports that this species has been credited with prodigious jumping
power (10 to 25 feet) and is greatly feared on account of its alleged
poisonous nature.
Many tests have been made with the poison of this tarantula. On white
rats and guinea pigs both the bite and injections have been employed. The
injections were made by grinding up both poison glands in distilled water, and
also in physiological salt solution. On guinea pigs no serious effects have ever
been observed. On white rats, one to two months old, four deaths have been
observed. The bite and the injection proved equally fatal. As a rule the bite
of this tarantula is not fatal to rats, but nevertheless it produces fairly definite
symptoms. At first the rat runs about excitedly, and in a jumping and jerky
manner. Then it becomes more quiet and appears to have considerable pain
in the wounded leg. For much of the time the eyes are closed. In about four or
five hours the rat shows evidence of recovery and in another hour it is normal.
On myself I tried the bite of this tarantula twice, and subsequently I have
been bitten by accident. The relatively dull fangs produce a pain that may be
compared to that made by a pin prick. It lasts for only 15 to 30 minutes and
is not accompanied by any inflammation or swelling.
The author has also been bitten by this species with similar benign
effects.
Sericopelma communis Cambro is a largc- black species of "tarantula"
common in the Panama Canal Zone, where it is generally feared. Baerg
(loc. cit.) allowed a spider of this species to bite him on the finger. He
allowed only one fang to puncture the skill. The finger felt numb in a
few minutes, and in 10 minutes the pain was quite severe. There followed
considerable swelling of the finger, hand, and wrist. After two hours
Baerg put the hand in hot water for 30 Ininutes, when the pain and
swelling subsided. A lame feeling in the small and third fingers remained
for several days. Baerg concludes that altllOugh decidedly painful, the
bite of this tarantula is probably not dangerouS.
Atrax formidabilis Cambridge and A. rubustus Cambridge are cited
by Thorp and Woodson '9 (1945) as dangerous spiders in Australia. The
bite may cause severe symptoms, and death may result. They are funnel-
web spiders of fairly large size, up to 1 Y2 iIlches in length excluding the
legs. The fore part of the body is glossy ebony black, the black abdomen
is covered with a velvety pile, while the under surface bears tufts and
bruShes or rea naIr.
DANGEROUS Latrodectus SPIDERS

Genus Latrodectus. Dangerous spiders belonging to the genus Latro-
dectus are found throughout the world. The genus belongs to the family
Theridiidae, the comb-footed spiders. They have eight eyes and three
tarsal claws; the hind pair of legs is comP-footed. They spin irregular
webs in which the female spiders hang DeIly upward. In the United
States at least two species are recognized; namely, Latrodectus mactans
(Fabr.) and L. geometricus C. Koch; in H;;lwaii, L. mactans is common;
in Mexico Latrodectus mactans is known as arana capulina, the "cherry
spider," also as chintatlahue; in Central ArJlerica it is known as pallu or
cassampulga; in the West Indies L. geometricus and L. curacaviensis
(Muller) are known as cul rouge; in South America L. mactans prevails
in many parts to Argentina as does L. geometricus; in Europe there is
L. guttatus, the malmignatte, also L. congoblatus Koch, the falangio in
Italy, and L. tredecimguttatus (Rossi) (with 13 triangular or half-moon-
shaped spots on the underside); in Russia, L. lugubris Motsch is known
as the karakurt (the black wolf); in Africa L. geometricus, L. concinnus
Cambridge, and L. indistinctus Cambridge are known as knoppie spin-
nekop (shoe-button spider); in Asia L. lugtt bris is widely distributed and
is known as the karakurt (black wolf) among the Tartars; in Australia,
L. hasseltii Thorell is common.
The black widow spider, LactrodectUS mactans (Fabr.), was first
described from America by Fabricius in 1775, under the name Aranea
mactans. Like many other species the specific name of this spider has
many synonyms, among them the following: Lathrodectus malmignathus
var. tropica van Haaselt, Latrodectus perfidus Walck., L. insularis Dahl.,

L. datatus C. Koch, L. apicalis Butler. Many common names are also
applied to this spider, among them in addition to "black widow" are
"hourglass spider," "shoe-button spider." "Pokomoo," a name used by
the California Indians, who referred to this species as "a small black
spider with a red spot under his belly."
FIG. 183. The black widow spider,

Latrodectus mac tans; (upper left)
mature female with egg sac, (upper
right) egg and first instar spiders,
(lower left) second and third instar
spiders, (lower right) fourth instar
spiders.
The adult female is glossy black to sepia and densely clothed with
short, almost microscopic hairs which give it a naked appearance. An
irregular white stripe, a remnant of nymphal pattern, is sometimes pres-
ent on the dorsal anterior margin of the abdomen. The characteristic
crimson hourglass marking on the underside of the abdomen (Fig. 183),
rarely altogether absent, varies among individuals from the distinct
hourglass marking to a design comprising two or more distinct triangles
or occasionally only an irregular longitudinal area. An occasional speci-
men has a crimson spot at the posterior end of abdomen on the dorsal
side, just above the spinerets. The abdomen is globose and often likened
to a shoe button. The average width of the abdomen is 6 millimeters, or
~ inch; and the length over all (legs extended) is about 40 millimeter;
that is, about 1 Yz inches. The abdomen of gravid females often measures
9 by 13 millimeters ( % by Yz inch).
The color pattern of the adult male (Fig. 184), while exhibiting con-
siderable variation, approaches that of the immature female spider. Oc-
casional mature males are almost black but retain some of the abdominal
markings of the immature form. The terminal segment of each palpus is
shaped like a knob (black) at the front of the head and contains the
ejaculatory sexual apparatus, a portion of which resembles a coiled watch
spring. The abdomen measures about 3 millimeters, or VB inch, in diame-
ter, and the length over all is about 30 millimeters (about 1 VB inches).
Distribution and habitat. This speCies, like most of the members of
the genus, favors warmer climates, although it is abundant in some of
the northern states of the United States. It has been reported in nearly
every state in the United States. The distribution of this spider is re-
FIG . 184. Male black widow spider,

Latrodectus mactans. Dorsal view
(left), ventral view (right).
ported as ranging from Canada in North America to Tierra del Fuego

in South America, including Mexico, Central America, Hawaii, and the
West Indies; it has been t~ken at an altitude of 8,000 feet in Colorado.21
In its natural habitat 'the black widow spider is found with her web
and egg sacs in protected darkened locations, such as vacated rodent
burrows, under stones, logs, and long grass; in hollow stumps, and in
brush piles. Convenient abode is found in darker corners of barns,
stables, privies, pump houses, garages, fruit drying sheds, piles of boxes
and crates, wood piles, stone piles, etc.
As a rule the females are not aggressive unless agitated or hungry.
When guarding the egg sac the female is particularly prone to bite.
Mature female spiders are present throughout the year but are more
abundant in the late summer and early fall. Many females have reached
maturity by that time, while a few are carried over from the brood of
the previous year. The mature males live but a few weeks. The observa-
tions coincide with the incidence of poisonous bites by months, i.e., the
majority of the spider-bite cases are recorded during July to October,
inclusive. .
Once a web is established in a suitable location, the female spends
the rest of her life feeding on the prey ensnared in this crude but ef-
fective web and guarding such egg sacs as she may deposit.
Feeding habits. Whether the prey be a nocturnal moth, cricket, or
domestic fly, the technique of capturing, killing, and finally sucking the
fluids from the victim is very consistent. The spider depends largely upon
vibrations of the web as an indication of a trespasser, or prospective
meal. The coarse, permanent web is not particularly viscid in nature, but
inadvertent insect visitors become temporarily entangled and in strug-
gling to free themselves inform the owner of the presence. The spider
always approaches the victim backwards, extending a freshly spun
strand of viscid silk with either one or both hind legs, and attempts
to tie down the thrashing appendages. If the captured prey appears
particularly obstreperous, the spider ejects from the spinnerets large
viscous droplets which dry quickly after the manner of rubber cement,
and if the victim becomes entangled by these jets, escape is impOSSible.
At about this point a lethal bite is usually administered. After being bitten
the victim struggles violently, and in the course of a few minutes of
progressively weaker tremors, dies. The body fluids are sucked from
the trussed up victim at the leisure of the captor. After the meal is fin-
ished, all points of attachment between the remains of the prey and the
web are cut loose, allowing it to drop from the web.
The diet of the black widow consists largely of insects of the locality,
small spiders, and even centipedes and sow bugs. It is surprising how
many insects' an individual spider consumes during its lifetime. Accurate
records kept of the food of isolated specimens have shown a total, in the
life of an individual spider, as high as 250 houseflies, 33 vinegar flies
( Drosophila), two crickets, and one small specimen of Latrodectus
mactans (Fabr.). In considering the economic status of this spider its
large diet of prevailing pests is a matter which should not be put aside
lightly. It is interesting to note in connection with the diet that one
individual (a male) was reared on a diet of its own species exclusively.
Mating habits. After molting the last time the male leaves its web
and seeks a mate. In this active, wandering state the male makes no
attempt to capture prey but will occasionally suck up a small amount
of water or liquid food if the opportunity is offered. If fortunate in find-
ing a likely mate, the male vibrates his abdomen rapidly, causing the
entire web to vibrate; the female may produce reciprocating movements.
Cautiously the male approaches and strokes the female with his forelegs.
It is a dangerous game the suitor plays, for if the female is not ready for
his advances, death may result. On the other hand, if the. female accepts
his advances, the wooing begins. If agreeable, the female remains quiet
and allows herself to be spun up in a delicate web. Once the web is suc-
cessfully spun the male effects coitus by applying the spring-like appa-
ratus of either pal pus to the female genital opening. Occasionally this
is repeated. After coitus the female easily frees herself and in many
instances ensnares and feeds upon her mate. The infrequent observance
and recognition of the male of Latrodectus mactans (Fabr.), together
with the mariticidal habit of the female, has given rise to the name of
"black widow." However, if other food is immediately available, often
the male is not killed and in due course of time dies a natural death. In
the laboratory the males will readily mate a second time, but the females
do not show such a tendency.
Comstock reports that in some male spiders the seminal fluid is trans-
mitted from the sexual organs (in the abdomen), which lack any ejacula-
tory apparatus, to the palpi and is there stored for some time previous to
mating. A delicate web appears to be spun upon which the fluid is emit-
ted and then collected by the palpi. The process has been observed by
the author in L. mactans (Fabr.).
Life history. The life history of the black widow spider from egg to
maturity requires about four months under laboratory conditions with
ample food. 22 The gravid female, when ready to deposit her eggs, forms
a loosely woven cup of silk which hangs downward and while clinging
inverted to its rim emits the eggs singly with rapid but regular upward
flexures of the abdomen. The eggs, which appear to be forced into an ex-
panding, gelatine-like film, gradually fill and adhere to the silken cup.
The open end of the cup is then covered with loose strands of silk and
the whole enclosed in a tough, watertight covering of silk The entire
-process consumes from one to three hours. Shortly after the egg sac is
completed, the film surrounding the eggs seems to evaporate, and the
eggs are free to roll about within their envelope. Egg-laying usually takes
place during the night.
In California these white or buff-colored egg sacs have been found
suspended in webs out of doors from March to October, inclusive. Egg-
laying may take place in the laboratory where food is plentiful through-
out the year. The egg sacs (Fig. 183) measure from 12 to 15 millimeters
(lh to % inch) in diameter, are usually oval in shape, and may contain
from 25 to 917 (Lawson,23 1933) spherical eggs, each of which is about
1 millimeter (1/32 inch) in diameter. Females have been observed to
spin from one to nine egg sacs a season. One spider under the writer's
observation spun seven egg sacs; the eggs in the last one did not hatch.
We have never observed egg deposition on the part of mature virgin
females. Fertile, mature females in isolation have produced egg sacs in
the fall, and s';Lrviving the winter, have produced additional egg sacs
the following spring, both groups of eggs being fertile.
The time between the deposition of successive groups of eggs varies
from about one week to about four months. The incubation period de-
pends on the temperature, and at normal summer temperatures requires
about 20 days in the interior of California, the observed extreme range
being from 14 to 30 days. At a sustained temperature of 27 0 -+- 1 0 C the
incubation period was about 30 days. The majority of the eggs usually
hatch but not simultaneously. In the case of several egg sacs, each being
formed by one female, the later lots appear to contain many sterile eggs.
The spiderlings after hatching spend some time-varying from four
days in the summer to about one month in cooler weather-within the
egg sac before emerging from one or more small holes which they make
in the tightly woven envelope. The first molt, previous to which the
spider cannot feed, occurs from one to two weeks after hatching. Usually
the entire first instar (and sometimes the second) is spent within the egg
sac, and at emergence the molted skins are left behind together with the
egg remnants. There is a tendency on the part of the spiderlings to cluster
for a few days after emerging from the egg sac, and cannibalism rules
during this time. The spinnerets appear to be capable of functioning at
the time of emergence, but the extremely delicate web is capable of hold-
ing only the smallest of prey, such as gnats, mosquitoes, and other tiny
spiders. The mother if confined with her young will not feed upon them
even though extremely hungry.
Shortly after emerging and after a brief period of clustering, the
nymphal spiders disperse by means of nearly invisible strands of silk.
For several weeks they move about in the vicinity of their birthplace
and suffer a high mortality from other predaceous spiders, and as already
stated, from their own species. When about one-third grown the female
spiders establish themselves in some protected niche, construct small,
loosely woven webs of their own, lacking in specific design, or, rarely,
take possession of an abandoned funnel, sheet, or irregular web. Once
settled they remain in the chosen lair, capturing progressively larger prey
and extending the web as they approach maturity.
The number of molts that the black widow has varies, and the length
of the intervening periods is even more inconstant, seemingly conditioned
by the season and the amount of food assimilated. The average number
of skins cast by the male is five. At optimum temperatures and with
plenty of food this number is often reduced to three; under less favorable
conditions, resulting in slower growth, a series of six skins may be shed.
The sexes may be distinguished by the palpi, or feelers, which in the
male are swollen or knob-like (Fig. 184), while in the female they are
slender. Subsequent to acquiring this secondary sexual character the
male molts once (sometimes twice) before attaining maturity, at which
time the web is abandoned, and his search for a mate begins.
The female takes longer to mature and has an average of seven molts,
with a range from six to eight. When preparing to molt, she eats nothing
for several days. The old skin splits around the margin of the carapace,
slips off the abdomen, and the spider then gradually pulls its legs free
from its old sheaths, leaving the "ghost" of itself on or near the web. The
entire process requires about an hour. The newly molted spider is rather
delicate and usually remains at rest for a day or so after molting. Indi-
viduals occasionally die during the molting process.
The immature stages. First instar. The abdomen of the newly hatched
spiderling is opalescent white with no markings. The cephalothorax and ap-
pendages are white to pale yellow; short hair covers the body, becoming dark
at the tips of the legs. The tarsi, or last segment of the legs, have two notched
claws and a supplementary third between. Eight simple eyes are on the ante-
rior margin of the cephalothorax in two rows of four, and the anterior medians
are comparatively dark.
Second instar. All eyes become darker and a black band extends down the
center and around the margin of the carapace. Also the mouth parts and the
appendages become darker as well as the margin of the sternum of the cephalo-
thorax. A double row of black dots extends down the mid region of the dorsum
of the abdomen which remains whitish. On the underside of the abdomen the
white area takes on a broad hourglass design outlined by a dark brown border.
Third instar. From this stage to maturity a wide variation in color pattern
occurs. Distinct lateral stripes begin to appear on the dorsum of the abdomen,
in the region of the dots of the second instar. Intervening areas take on a pale
greenish-yellow cast, and the legs acquire four black bands, and at each end of
the patella, one near the center of the tibia, and one at the junction of the tibia
and the metatarsus. The longitudinal white area on the underside of the ab-
domen becomes tinged with crimson.
Fourth instar. Dark stripes or bands become distinct and faintly bordered
with buff. The spinnerets take on a mottled appearance. Black bands at the
leg joints become more distinct.
Fifth instar. The central dorsal white stripe on the abdomen tends to be
constricted at intervals and acquires a reddish tinge near the tip. All white
areas become lightly colored with brown. Males usually mature at this point.
Sixth and seventh instars. Usually only the females go through these two
stages. All coloration is much darker, and the more variable remaining white
areas become more and more restricted. Often a series of reddish spots are
formed along the middorsal region of the abdomen.
Eighth instar. Only the females pass through this stage, which is often
difficult to distinguish from the mature form. They are usually all black or
sepia w.ith the exception of the characteristic crimson markings and an occa-
sional white band on the anterior margin of abdomen.
Longevity. The length of life of individual spiders, as one might

expect depends on such factors as food supply, natural enemies, includ-
ing man, etc. Under optimum conditions of food, temperature, humidity,
protection, etc., the complete life cycle from egg to maturity requires at
least four months. Spiderlings emerging from eggs laid in July and
hatched in August will, of course, pass the winter in an immature stage
which thus materially extends the length of time required to complete
the life cycle. Activity on the part of both the spiders and the insect
prey is greatly reduced during the winter months and thus largely ac-
counts for the retardation in development. When a brood emerges in
late spring or early summer, the females generally reach maturity before
cold weather sets in, but egg laying is held over until the following
spring, and hence the life cycle is extended over a complete year. Ma-
ture males have not been found overwintering.
Under laboratory conditions a few females have lived through the
second and third summers, giving a life span of nearly two years.
Arachnidism (spider bite). While spiders in general have been con-
sidered poisonous, though largely erroneously, for centuries, the group
to which the black widow belongs in particular has been classed as
poisonous for little more than a century. Many of the early reports of
spider bite traceable to the black widow came from the southern states
and from 1889 to 1894 were frequently mentioned in Insect Life (Riley
and Howard, 1889-94). After the rapid increase in the population of
California during the latter part of the nineteenth century, reports of
poisonous spider bites began to be received from this state. In 1932
Bogen 24 listed a total of 380 cases from 18 states, of which 250 were
from California.
The chain of symptoms resulting from the bite of the black widow
spider is so striking that once recognized there is little danger of confus-
ing it with that of other venomous forms or with an acute abdominal
condition. Cases of arachnidism, or spider-bite poisoning, suggest numer-
ous acute abdominal conditions such as a ruptured peptic ulcer, acute
appendicitis, renal colic, enteritis, food poisoning, etc.
The bite itself (similar to a pinprick) is not always felt, and often
there is but little evidence of a lesion. However, a slight local swelling
and two tiny red spots may occur, and local redness is usually in evi-
dence at the point of attack.
Pain, usually in the region of the bite, is felt almost immediately and
increases in intensity, reaching its maximum in one to three hours and
generally continuing for 12 to 48 hours, gradually subsiding. Rigidity and
spasm of most of the larger muscle groups of the body (particularly
those of the abdomen) are constant symptoms. The abdominal muscles
become "board-like," but local tenderness as in appendicitis is almost
always absent. There is a slight rise in body temperature, increased blood
pressure, a definite leucocytosis, and usually an increase in the pressure
of the spinal fluid There is usually profuse perspiration and often a
tendency to nausea. The degree with which these symptoms are present
varies in individual cases, and other symptoms such as chills, urinary
retention, constipation, hyperactive reflexes, priapism, and a burning sen-
sation of the skin are frequently reported.
Baerg,25 who permitted himself to be bitten (basal joint of the third
finger of the left hand) by the black widow spider, when reporting on the
effect of the bite, states:
Referring briefly to some of the general effects of the case, I would say that
the sharp pain in the finger, or rather in the left hand, was the most prominent
feature. Very nearly as unpleasant was the aching pain which was most violent
in the thick m1}scles of the lower part of the back, and present in almost all the
muscles of the shoulders, chest, and legs. There was no marked tendency
towards profuse perspiration. I sweated heavily only when I first went to bed,
and later after each one of the hot baths. I covered up well after these baths in
order to bring about sweating, and I believe that it aided in recovery. There
was no evidence of constipation. One dose of magnesium citrate brought fairly
prompt results. On the day I left the hospital I took a second dose in order to
facilitate recovery as much as pOSSible.
Baerg's physician, Dr. E. F. Ellis, added the following note:
The subjective symptoms in Mr. Baerg's case have been very graphically
described by him. The objective symptoms would indicate, as observed by
me, that there is a very marked phagocytosis locally around the area of the
spider bite. The toxicity of the bite was such that the phagocytes very shortly
offered no resistance to the systemic invasion of the poison. The poison in
my opinion was partly transmitted through the blood stream and partly
through the nerve trunk which in this case was the median nerve. Strange
to say in this particular instance the patient had a marked vasomotor disturb-
ance on the flexor side of the forearm, as was evidenced by a narrow strip
something like an inch in width, extending up almost to the elbow in which
there was very marked diaphoresis. This was present during the first 24 hours
after the bite. The toxicity was also manifested by vasomotor changes in the
lumbar muscles and muscles of the extremities, and in all the large joints of the
body, as was shown by intermittent pains and symptoms similar to intermit-
tent claudication. There seems also to be a disposition, on his part, to unload
very slowly, by elimination, the products of poison. More so than is the case
with bites of any of the snakes including the rattler that I have observed."
Clinical case records. The following case records from the Woodland
( California) Clinic are typical of arachnidism:
CASE No.1
Age: 41 Sex: Male Nationality: Dutch
Occupation: laborer .. Date admitted: 9-3-31. .Date dismissed: 9-8-31
9-3-31. About 8:30 P.M. on September 3 patient was taken with a sharp
pain in the glans penis as he was sitting on the stool of an open privy. In about
VENO¥OUS AND URTICARIAL ARTHROPODS 589
15 to 20 minutes the pain spread to the abdomen, hips, and thighs, the pain
being steady, deep, and dull, and so severe that the patient doubled up and
rolled on the floor. His abdomen got hard with the onset of the pain. In about
two hours pain had developed in the back. He was given two hypodermics
(morphine) on entry which relieved him some but stated that he slept only
one-half hour during night. Heat was applied to legs and back. Around 4 this
morning (9-4-31) the pain seemed to shift to legs (below knees) and feet. In
addition to type of pain described in the abdomen, he had an intense burning
sensation of skin, worse on soles of the feet. Patient has slight pain in chest,
no appetite, and feels weak and shaky all over. Patient unable to urinate.
9-4-31. Patient now has pain in back and legs. Sweating profusely. Pulse,
80. Abdomen absolutely rigid-"board-like." Legs flaccid, neck not stiff. Ex-
amination of penis reveals old scars, no recent lesion seen. Impression-
arachnidism.
9-5-31. Pain practically gone. Voiding and eating well.
9-7-31. Free of pain. Except for weakness, feels well.
9-8-31. Free of pain. Feels well. Dismissed.
Chart shows patient's temperature Ructuated but not abnormally, pulse
somewhat above normal, and respiration normal.
CASE No.2
Age: 24 Sex: Male Nationality: American
Occupation: laborer.. Date admitted: 6-17-32 .. Date dismissed: 6-31-32
6-17-32. About 9 P.M. last evening patient retired and after getting in bed
something dropped on his right shoulder. After turning on the light he dis-
covered a large jet-black spider which he killed. About midnight he noted a
tightness in his abdomen and a rather severe lumbar backache. Went to a
local physician who gave him a hypodermic about 2 A.M. Abdominal tenseness
and pain increased and patient began to note leg pain.
6-17-32. Patient admitted at 3:30 A.M. and examined. Eight hundred
cubic centimeters of 5 per cent glucose in normal saline given intravenously
after which pulse, which was thready, became fuller and stronger. At 4:45 A.M.
considerable back pain and given morphine sulphate (14 grain).
6-18-32. Abdomen softer. Considerable pain in feet. Profuse perspiration-
now apparently out of danger.
6-20-32. Abdomen soft, eyelid edema gone. Considerable discomfort in
both feet. Still perspiring but reduced.
6-31-32. Patient discharged. Fingernails look blue.
Chart shows patient's temperature to be slightly above normal, pulse
fluctuating, and respiration normal.
Of 37 persons with cases of arachnidism treated at the Woodland

(California) Clinic during a period of 10 years, about one-half were
bitten on the genitalia while using a privy.26 Four of the patients were
females and the remainder males, the majority of whom were laborers
and farmers between the ages of twenty and fifty. Five school children
were among those treated. The temperature and pulse of the patients
were usually slightly above normal, and the respiration normal. Mor-
phine sulphate (]i4 grain) was employed in 25 cases and other drugs
such as codeine, amidopyrine, atropine, hyoSCine, etc., in the remainder.
Although spider-bite cases were admitted to this hospital every month in
the year except December and January, the greatest number were ad-
mitted during June, July, and August. There was no correlation between
the time of day and the bites. The spider was seen in 11 cases. None of
the patients reported any after effects on the follow-up inquiry.
Only two deaths were reported in these records, one (6/ 25/ 24) of a
male 78 years of age, with heart and syphilitic complications, and the
other (7/ 13/ 25) also of a male (an itinerant) aged 44 years who died
four days after being bitten. Post-mortem examination of the latter pa-
tient showed a ballooned intestine, and he was recorded as dying of a
paralytic ileus resulting from a black widow spider bite.
Venom apparatus. Spiders are equipped with a poison apparatus for
pnrposes of killing their prey. The venom apparatus of Latrodectus
FIG . 185 (A) chelicerae and
'] venom glands of female black widow

spider, Latrodectus mactalls; (B)
separated chelicera; and (C) a freed
gland. (Photograph by Charles Lad-
enheim.)
ABC
mactans (Fabr.) consists of two poison sacs each with an attached duct
leading into the heavily chitinized chelicerae (horny fangs) from which
the poison is expelled at the time of biting (Fig. 185) . The venom ap-
paratus is present in both the male and the female spider. In the male
the venom is primarily of use in the immature stages. The mature male
does not attack prey, and the poison apparatus appears to become in-
active with maturity and remains small in size. In the case of the female
spider the venom apparatus increases in size and strength with maturity.
The large quantity of the venom present in the poison glands of the
female black widow spider makes her presence perilous. The poison
glands of the mature female average about 0.40 mm in diameter and
2.70 mm in length, and in the mature male they average 0.16 mm in
diameter and 0.66 mm in length.
The poison glands are located in the cephalothorax extending pos-
teriorly about two-thirds of the way to the junction of the cephalothorax
with the abdomen. The long narrow glands curve outwardly around the
fovea media, that is, the indentation in the center of the cephalothorax
for muscular attachment; this is due to the position of the sucking
stomach directly below. If the glands were over the sucking stomach
they would be squeezed together every time the sucking stomach IDled
with food and cause a loss of venom. In the female (all stages) the
glands are opaque aI).d filled with venom, while in the mature male
they are rather translucent but according to experimental evidence con-
tain sufficient venom to produce symptoms of arachnidism in white mice.
Nature of the venom. There is good evidence showing that the so-
called poison glands are not glandular in nature, but function as ab-
sorptive organs which take up the poisonous constituents from the body
fluid of the spider. Sachs (1902) and Kobert in 1901 and 1906, according
to Bogen, isolated from the spider body a specific poisonous principle
named "arachnolysin" which they claim has a hemolytic effect on the
blood of various animals [see also Hall and Vogelsang 21 (1932)]. Ex-
perimental evidence indicates that the venom of L. mactans acts pri-
marily as a neurotoxin. 28 Spider poison is not limited to the poison glands
but is also carried in the body fluids as pointed out by Sachs and by
Kobert.
Blyth and Blyth29 also state that the poison of Latrodectus lugubris,
. is not only found in special glands, but is also diffused through the body.
. it is a generic type of poison of spiders; the active principle is neither
a glucoside, acid, nor an alkaloid. It does not dialyse, and drying destroys
its activity; it has the characters of a toxalbumin, and has much similarity
to the action of ricin and abrin. The "Kara-Kurt" poison dissolves the color-
ing matter of the red blood corpuscles even with a dilution of 1: 127,000;
it has a paralyzing effect on the heart, either due to action on the motor
ganglia, or possibly a direct action on the muscle itself. The blood pressure
sinks, the walls of the smallest arteries and capillaries become so changed
as to allow the transudation of the blood and serum, producing punctiform
hemorrhages and edema. This is best seen in the lungs. . . . The poison also
has a paralyzing action on the central nervous system, but it is not clear
whether this action is primary, or whether it depends on the circulation
troubles.
The fatal dose of the poison, injected subcutaneously or intravenously,
is extremely small. Cats are killed by quantities equal to 0.2 to 0.35 milligram
per kilogram body weight. Repeated injections of nonfatal doses confer im-
munity.
Kellogg 3o (1915) states that high temperatures destroy the action of

extracted black widow spider venom. It can withstand a temperature
of 56° C (132° F) for 40 minutes, but it is wholly destroyed if heated
for 45 minutes at 70° C (158° F). The venom cannot be crystallized,
since it is destroyed by desiccation. The best means of preserving the
extracted venom is in glycerin, where it can be kept for several months.
Eating, black widow spiders caused no harm to white mice, guinea pigs,
nor a monkey; neither did canned spinach cooked with black widow
spider in it. 31
Treatment. When bitten by the black widow spider, the patient
should be treated with local antiseptics, such as tincture o~ iodine, at
the pOint of injury to prevent secondary infection, and should be kept
as quiet as possible; a phYSician summoned at once. Since, among other
properties, the venom appears to be neurotoxic and its effect little short
of instantaneous, first-aid measures for snake bite are of little value.
Medical records, according to Bogen (1926, 1932), list more than 75
different remedies and, of all these, three seem to be outstanding as
palliatives; namely, spinal puncture, intravenous injections of magnesium
sulphate, and intramuscular administration of convalescent serum when
given within eight hours.
Gilbert and Stewart32 (1935) point out that, as "the toxin directly
stimulates the myoneural junctions or that it acts on the nerve endings,
to find a type of therapy which would have a direct depressant effect
upon these structures would be ideal." Because calcium apparently de-
presses the neuromuscular junctions, Gilbert and Stewart selected cal-
cium salts for the treatment of black widow spider bite. They report on
this treatment as follows, "We found that intravenous injections of 10
per cent calcium chloride gave instantaneous and prolonged relief of
the pain and at the same time produced immediate relaxation of the
muscle spasm so commonly seen in these patients. However, calcium
chloride is not given thus without considerable danger. Its necrotic
action on tissue outside a vein is only too well known. This danger is
greatly magnified when its use is attempted in the treatment of children.
Therefore, calcium gluconate (10 cc. of 10 per cent solution, intrave-
nously ), which does not have this objectionable feature, was used and
found to produce equally as spectacular results as the calcium chloride.
The intramuscular route, advisable for children, gave relief within a
minute's time. Calcium lactate orally was ineffective as far as determined,
probably because of its incomplete and slow absorption." Halter and
Kuze1l 26 also report similar immediate relief, usually in about one minute,
follOWing intravenous treatment of adult males with both calcium chlo-
ride and calcium gluconate (10 cc of 10 per cent). In some cases, these
authors suggest, a repeat may be necessary at intervals of one to two
hours for four or five times.
As Bogen (loc. cit., 1932) states, in part, " . . . despite its severe
symptoms arachnidism is in the majority of cases a self-limited condition,
and generally clears up spontaneously within a few days."
Control. The black widow spider is frequently found in garages,
basements, in living quarters, in old outbuildings (particularly privies,
barns, pump houses, stables), and woodpiles. Frequent disturbance of
spider webs with a broom or brush is suggested, and the spiders should
be killed in any convenient way. Workers in dried fruit industries find
numerous black widow spiders under and in the drying trays when
handling these. The use of gloves is suggested in this instance. Out-of-
doors the spider may be found in vacant lots and open fields and on hill-
sides, building their webs in rock piles, heaps of rubbish, under logs,
under projecting banks, in deserted squirrel or rabbit holes, under low
wooden or concrete bridges, culverts, etc.
Since the egg sacs are conspicuous and are not carried about by the
spider, they may be readily collected and destroyed. Great care should
be exercised when collecting egg sacs because the female spider guards
the sac closely and is particularly vicious at that time. The public should
be encouraged to collect and burn or otherwise destroy the egg sacs of
the black widow spider. Where accessible, the adult spider can be
brushed from its web with a broom or stick and stamped upon, or a
suitable fly spray may be used to knock the spider to the floor where
it should then be crushed. The use of a blow torch when no hazard is
involved is effective in the destruction of eggs and spiders as well. Be-
cause of the danger from spider bite when using privies in rural sections,
it is suggested that the undersides of the seats and corners of the box
be painted well with creosote. Privies should be properly constructed to
exclude spiders and notably flies on which they feed. Thorough and
liberal spraying of all parts particularly under the seats with DDT fly
spray, benzene hexachloride, chlordane, or the like is strongly urged.
The adult female black widow spider is extremely difficult to kill with
the usual fly sprays which act for the most part as moderate repellents
and at best only render the spider temporarily paralyzed. It is of interest
to note that a good kill of black widow spiders has been achieved in-
cidental to the use of DDT spray in fly control operations in stables and
dairies. Chlordane and benzene hexachloride sprays are reported to be
fairly effective. Pyrethrum fly spray fortified with an organic thiocyanate
(Lethane 384) is a useful contact spray in the household. 33 Corners in
garages, outbuildings, basements, etc., may be sprayed with a penetrating
creosote which has some repellent effect against spiders.
Natural enemies. Under natural conditions the black widow spider
is held at least moderately in check by its natural enemies. Among the
various spider-hunting wasps and mud daubers there appear to be cer-
tain species which completely ignore this spider, as for example the
yellow-marked mud dauber, Sceliphron caementarius Drury, while other
species of mud daubers store the black widow spider in their nests; thus
Irving and Hinman 34 discovered that the blue mud dauber, Chalybion
cyaneum (Klug), captures great numbers of this spider. They found 285
black widow spiders had been stored in 15 nests, an average of 19 per
nest. The large San Diegan alligator lizard, Gerrhonotus multicarinatus
webii (Baird), in southern California, has been suggested by Cowles 35
as probably an important factor in cutting down the incidence of this
spider on his property. Cowles points out that the elimination of cats is
essential because they will eat this lizard. Egg predators, such as the
scelionid wasp, Baeus latrodecti Dozier, and the chloropid fly, Pseudo-
gaurax signata (Loew) (Gaurax araneae Coq.), play an important role.
The chloropid fly, Pseudogaurax signata (Loew), can be reared suc-
c<f§sfully in captivity as proved by George Elwood Jenks, who has in-
dicated his success to the author by correspondence and has beautifully
illustrated the work of this interesting fly in the August, 1936, number
of Popular Science Monthly. The larvae of this fly lie free in the egg sac
and completely consume the eggs of the spider. The fly deposits its
glistening white eggs on the outside of the spider's egg sac. The larvae
hatch in five or six days and gain entrance into the sac by pushing their
way through the fibers. The length of the larval stage is eight to nine
days, according to Kaston and Jenks,36 and the pupal stage requires 11
to 12 days.
With protection afforded by man-made structures which exclude the
natural enemies, the spider thrives and multiplies rapidly.
Other spiders. Instead of being objects of admiration because of their
beauty, many of the beautiful garden spiders, orb weavers, are greatly
feared. These spiders are harmless and what is more, they are beneficial
because they feed on insects which may be harmful to the garden. Two
common garden spiders are Miranda aurantia (Lucas), the golden orb
weaver, and Argiope argentata (Fabr.), the silvered orb weaver; both
of these construct beautiful geometrical webs. 37 They belong to the
family Argiopidae.
The remarkable trap-door spiders represented by the California spe-
cies, Bothriocyrtum californicum Cambridge, and the eastern trap-door
spider, Pacylomerus audouini Lucas, belong to the family Aviculariidae
already mentioned as including the tarantulas. The trap-door spiders
are perfectly harmless. (Passmore. 38)
The burrowing spider, Brachythele longitarsus Simon, commonly
causes consternation in the home when winter rains and cold weather
drive it undercover. Like the trap-door spider, which it resembles, it is
quite harmless.
Steatoda borealis (Hentz), belonging to the family Theridiidae, re-
sembles the black widow spider rather closely but is slightly smaller;
the color is sepia and it bears no red markings. The egg sac is small
and loosely woven. It is a harmless species.
The banana spider, Heteropoda venatoria Koch, belongs to the family
Heteropodidae and is frequently brought into northern fruit markets in
VENOMOUS AND URTICARIAL ,ARTHROPODS 595
bunches of bananas or other produce from the tropics. These rather large
(spread of 3Y2 inches), long-legged, dark-brown spiders are commonly
mistaken for tarantulas. The bite of this species is painful and at times
may cause severe envenomization.
In the house as well as out of doors one commonly encounters the
small jumping spiders, belonging to the family Attidae. Members of the
genus Phidippus may be seen stalking houseflies on the windows of the
dining room or kitchen. These spiders have been known to bite, but so
far as the author is aware, the effects are ben.ign. ' ..
The "pruning spider," Gliptocranium ga/J1eracanthoides Nicolet, is
reported by Escomeps to be a particularly -venomous species in Peru.
SCORPIONS
Class Arachnida, Order Scorpionida
Genera) cnaraclerlstks. -::'corpions are easiiy Teeogn'neo. 'oy Ynei:r more
or less crab-like appearance but particularly by the long fleshy five-
segmented tail-like postabdomen terminating in a bulbous sac and promi-
nent sting (Fig. 186). The pedipalps are greatly enlarged, and the last
two segments form strong chelae or pincers. The true jaws, chelicerae,
are small and partly concealed from above by the front edge of the
carapace. There are four pairs of terminally clawed legs.
The cephalothorax bears a pair of conspicuous eyes near the mid-
dorsal line (median eyes), and several smaller ocelli in groups of from
two to five, on the lateral margins (lateral eyes). Some species are eye-
less. Scorpions breathe by means of lung booles. They are ovoviviparous,
and when the young are born they are carried attached by their pincers
to the body of the mother. Although the sexes are very similar in appear-
ance, the males have a longer cauda and broader chelae. An excellent
account of the morphology as well as characters used in classification of
scorpions is given by Moreno 4o in "Scorpiologia Cubana."
Scorpions are found most commonly in warmer climates. They are
nocturnal, remaining hidden during the day beneath loose stones, loose
bark of fallen trees, boards, piles of lumber, floors of outbuildings, and
debris; some bury themselves in sand or loose earth. They feed upon
large insects and spiders, which they seize with their chelae and sting
with their powerful sting, which is thrust forward over the scorpion's
back.
Scorpion sting. The "aculeus" or sting of the scorpion is situated
terminally on the final bulbous segment. This segment qpntains a'pair of
venom glands, which are separated by a muscular septum. From the
glands are given off fine efferent ducts opening at the apex of the sting
(Pawlowsky41 ). The sting curves downward when the "tail" is extended
but upward and forward when the scorpion poises for attack or defense,
the entire tail-like postabdomen being curved dorsally and forward. The
victim is struck quickly and repeatedly, the thrust being made quite
close to the front of the carapace.
The venom is a transparent liquid, acid in reaction. The toxic princi-
ple is said to be a neurotoxin. "It also has a lecithide which haemolyzes
FIG. 186. A scorpion, Hadrll1'lIs

hirslltus. X .6.
nucleated as well as nonnucleated blood corpuscles. . . . Although the

sting of a scorpion is very painful, the poison as a rule does not produce
general symptom.> in adults, but in children under five years of age the
sting frequently causes death," according to Waterman!2 whose notes
on the subject of scorpion poisoning should be consulted by physicians
concerned with this matter. Waterman states that the "diagnosis is gen-
erally easy if a history of the sting is obtained, a slow full pulse easily
compressible, with rapid respirations, a pulse respiration ratio of 3: 1,
2: 1, or 1: 1; salivation, vomiting, glycosuria and epigastric pain and
tendemess-a characteristic picture of scorpion sting." The puncture
made by the aculeus rnay be visible.
The symptoms caused by the sting of the Durango scorpion, Cen-
truroides sufJusus Pocock [Centrurus gracilis (Latreille)]' are described
by Baerg (lac. cit., 1939) as follows:
Immediately following the sharp pain produced by the sting is a feeling

of numbness or drowsiness, then there is an itching sensation in the nose,
mouth and throat that makes the victim distort the face, rub nose and mouth,
and sneeze. There is at first an excessive production of saliva; this and a
curious feeling that is described as the sensation of a ball of hair in the throat,
induce the victim to swallow as rapidly as possible. The tongue is sluggish, so
that communication is often by signs. The muscles of the lower jaw are con-
tracted so that it is difficult, or impossible, to give medicine through the mouth.
There is a disorder of movements in arms and legs. The temperature rises
rapidly to 104 0 or 104.8 0 F., the salivary secretion now diminishes and there is
a scarcity of urine. The senses of touch and sight are affected, objects appear
large on touching them, hair feels rigid, face feels bulky, a veil seems to be
interposed between the eyes and various objects, strong light is unpleasant to
patients. Luminous objects, such as a candle, are surrounded by a red circle.
Frequently there is a pronounced strabismus. There may be a hemorrhage of
the stomach, intestine and lungs. The convulsions come in waves and increase
in severity for an hour and a half to two hours, or in severe cases until a fatal
result. When the case ends in death, respiration stops a full minute before the
pulse ceases to beat. When the patient survives for three hours he is usually
considered out of danger; yet death may occur six to eight hours after the
patient was stung. It is then probably due to nervous exhaustion following
the long periods of convulsions.
Even though stunt by a presumably innocuous scorpion the person

bitten should be taken to a physician if the bite has been inflicted on the
face, the back of the neck or the genitalia. Ordinary scorpion stings may
be given first-aid treatment according to Stahnke,43 as follows, "A piece
of ice should be applied as soon as possible against the site of the sting;
then more ice should be finely crushed and placed in water, using more
ice than water. With a piece of ice against the site of the sting place
the hand in the ice mixture. . . . Where a large dose of venom has
been received the hand should be kept in this mixture for about two
hours . . . a large ice pack of finely crushed ice in a thin cloth may be
placed over the site of the sting." The function of the ice is to decrease
absorption or to localize the venom; it has no effect on the venom itself.
The order Scorpionida is divided into six or seven families, depending
upon the author. These are Scorpionidae, Buthidae, Vaejovidae, Chac-
tidae, Ischnuridae, Bothriuridae, and Chaerilidae. According to Ewing,44

four of these families occur in the United States and northern Mexico,
and are separated as follows:
A.' Sternum subpentagonal, with sides almost parallel.

B.' Membrane at base of last tarsal segment of most of the legs with a
single spur; postabdomen frequently reduced. .. Scorpionidae
B.' Membrane at base of last tarsal segment of most of the legs with
two spurs.
C.' Only two ocelli on each lateral margin of carapace .Chactidae
C: With three to five ocelli on each lateral margin of carapace
Vaejovidae
A.' Sternum triangular, the sides being strongly convergent anteriorly; mem-
brane at the base of last tarsal segment of most of the legs with two
unbranched spurs; fixed arm of chelicerae without ventral tooth
Buthidae
Over 300 species of scorpions are known. Of the 40 species found

in the United States, according to Stahnke (lac. cit.) only two are dan-
gerously virulent to man (see below).
Family Buthidae. This is the most widely distributed of the families
of scorpions and is distinguished from the other American families by the
shape of the sternum which is triangular, the sides being strongly con-
vergent anteriorly (Ewing). To this family belong Centruroides sufJusus
Pocock, the so-called Durango scorpion, the common scorpion of the
State of Durango, as well as of other neighboring states in Mexico and
portions of Arizona. During the period from 1890 to 1926, according to
Baerg (1929, lac. cit.), " . . . a period of 36 years, there have been
1,608 deaths" from scorpion stings in Durango. The majority of victims
are children from one to seven years. The sympt<~ms caused by the sting
of this scorpion are described above.
Centruroides sculpturatus Ewing, said by Stahnke (lac. cit.) to be
confined at present to Arizona, is a small species, about 2Yt inches in
length; it is generally of a solid yellow-straw color. It is a dangerously
virulent species and is reported to be abundant.
Centruroides gertschi Stahnke, so far as is known, according to
Stahnke (lac. cit.), is also confined to Arizona but is much less abundant
than C. sculpturatus. It is about the same size as the latter but has two
irregular black stripes down its entire back; its basic color is yellow.
This is also a dangerous species.
Centruroides vittatus (Say) [C. carolinianus (Wood)] is the common
striped scorpion of the United States. It is widely distributed, having
been reported from Georgia, Florida, Kansas, Texas, Arkansas, Louisiana,
New Mexico, and South Carolina. Concerning the sting of this species,
Ewing (1928,. loco cit.) writes, "The writer has induced this species to
sting him and has observed the effects of its sting on others. At the time
of the stinging there is a sharp pain, but this soon subsides. A small
swollen area, or wheal, usually develops about the puncture point. This
soon disappears. There are no permanent effects of the sting reported
for the species as far as known to the writer:' Stahnke (loc. cit.) also
regards this species as relatively harmless.
Centruroides nigrescens (Pocock) is known as the black scorpion.
It is reported from Texas and Mexico. It measures 10 cm in length and
is a dark chestnut-brown to jet-black color.
Centruroides californicus (Girard) is the widely distributed striped
scorpion of California. It resembles C. vittatus (Say) very closely.
Buthus quinquestriatus Hemprich and Ehrenberg is a common
Egyptian and North African species, more especially in upper Egypt,
according to Wilson/ 5 who states that it is of a sandy-yellow color tend-
ing to brown and measures about 10 cm in length. He also states that
it is undoubtedly the commonest species in thilt region and is generally
believed to be the most dangerous; it is frequently found in houses and
is the species in all probability giving rise to the numerous cases of
scorpion sting said to be most commonly fatal in upper Egypt. Shulov46
reports having traced four fatal cases in children between the ages of
6 and 13 in Palestine.
Family Scorpionidae. According to Ewing this family is well repre-
sented in Central America and the tropical I(~gions of the Old W orId,
but is poorly represented in the United States.
Diplocentrus whitei (Gerv.) is a very dark reddish-brown scorpion
from 5 to 7 cm in length. This species has been taken on the Mojave
Desert, California. Ewing writes that he has been inclined to associate
serious scorpion stings with this species, be(~ause of the descriptions
given by persons living near the Mexican botllldary who have had ex-
perience with cases of severe scorpion stings. On the other hand, Baerg
(loc. cit.) states that several punctures from two scorpions failed to
produce any appreciable effect on him except the slight pain of the
puncture.
Family Chactidae. This family differs from all other North American
scorpion families in having only two occeli on each side of the carapace.
Broteas alleni (Wood) is a small dark-brown species measuring about
3 cm in length. It has been taken in lower California and at Fort Tejon,
California.
Family Vaejovidae. This is the best represented family in North and
Central America. It contains a number of very large species.
Hadrurus hirsutus (Wood) (Fig. 186) is the giant hairy scorpion, our
largest species, measuring 11 to 12 em in length. The body is dark yel-
lowish and hairy. It is found in the southwestern United States and
northern Mexico. The writer has taken this species in the Imperial Val-
ley, California.
Hadrurus aztecus Pocock is the Mexican hairy scorpion and closely
resembles H. hirsutus (Wood).
Vaejovis spinigerus Wood is the stripe-tailed scorpion. It has four
longitudinal dark stripes on the underside of the "tail." It measures from
5 to 8 cm in length. It is a typical desert species, occurring in rocky
waste places of Texas, New Mexico, and California, where it is common.
Baerg (loc. cit.) reports that its sting caused only a slight pain which
disappeared in less than half an hour; its poison had no appreciable
effect on white rats.
Vae;ovis boreus (Girard) is the northern scorpion occurring in North
and South Dakota, Idaho, Wyoming, Nebraska, and Montana. It is a
dark, yellowish-brown, unmarked species measuring from 3.5 to 5 cm in
length. Its sting, though painful, is benign in effect.
Uroctonus mordax Thorell is the mordant scorpion, a dark-brown
medium-sized Pacific coast species. It is the commonest species in the
San Francisco Bay region. Its sting is about as painful as that of a yellow
jacket, but causes as a rule less swelling; the effects soon disappear. It
occurs under loose rocks, beneath bark of fallen trees, under rubbish,
tent floors, and the like.
Scorpion control. The scorpion hazard on premises may be largely
reduced by the elimination of favorable hiding places, such as boards,
loose rocks, rubbish, platforms, and the like. Creosote sprays have a
substantial repellent effect, but unless a clear solution can be obtained,
may not be desirable about the yard. In some localities where scorpions
are abundant and dangerous a bounty has been paid. Thus Baerg (loc.
cit.) reports that in 1928 (May 1 to July 31) 12,941 scorpions were
collected in and near the city of Durango by scorpion collectors (alacra-
neros ), and a bounty of two and one-half cents for the females and two
cents for the males was paid. Baerg also states that Dr. Brachetti ad-
vises that the "powder of chrysanthemum will drive scorpions away, or
even kill them, and so does creoline-a dilute solution of creoline to be
sprinkled on the floors and on the flower beds about the house."
WHIP SCORPIONS
Class Arachnida, Order Pedipalpida
Characteristics of Pedipalpida. The Pedipalpida are tropical and sub-
tropical arachnids although very unevenly distributed. They are said to
be absent from Europe and North Africa (Savory). The term "whip
scorpion" is applied to the Family Thelyphonidae because the terminal
end of the abdomen is provided with a long, slender, many-segmented

appendage (Fig. 187).
The giant whip scorpion, Mastigoproctus giganteus (Lucas), occurs
in Florida where, according to Ewing, (1928 lac. cit.,) it is found on the
ground under various litter and under logs, boards, and lumber lying
on moist ground. It feeds on almost all kinds of larger insects and other
FIG. 187. Whip scorpion (Pedipalpida) Masti-

goproctus gigantetls. X .8.
arthropods if not too hard or too active. A closely related species, if not
identical, occurs in southern California, mainly in sandy desert places
where it burrows in sand under debris. It is commonly regarded as
poisonous, although it cannot sting but may bite. The writer has found
that many persons living in the arid parts of California fear this creature
very much, but knows of no evidence to justify this fear . Ewing states
that on no occasion was there more than a trivial mechanical effect from
the bite, similar to that of a slight pinprick. He states that when handled
it gives off a repellent fluid which has the odor of vinegar. This fluid may
possibly produce some irritation to persons with a tender skin.
SOLPUGIDS
Class Arachnida, Order Solpugida
Characteristics of the Solpugida. The solpugids (Fig. 188), commonly
known as "sun spiders" and "wind scorpions," are in general appearance
spider-like, although there is no pedicle; they are very hairy, largely
nocturnal, occurring mainly in desert, tropical, and subtropical regions.
FIG. 188. (Left) a soIpugid, commonly called "sun spider"; (right) eggs.
They are common in many parts of California and have been reported
as far north as Nebraska. The chelicerae are large and powerful and
are two-segmented. The second segment is movable and articulates in
such fashion as to work in a more or less vertical plane. Food is crushed
to a pulp, the fluid is swallowed, and the hard parts are ejected. The
first pair of legs are used as tactile organs. Respiration is tracheate. They
are commonly but erroneously regarded as exceedingly venomous. The
writer has been told that the presence of one of these animals in a water-
ing trough would result in the death of any animal drinking from the
same. There is evidently not the slightest foundation for this belief. Al-
though these animals are able to inflict a painful bite by means of their
powerful jaws, the effect is fleeting. Poison glands are absent.
There are said to be only twelve species in the United States, all but
one belonging to the two genera, Eremobates, e.g., Eremobates formi,
carius (Koch) and E. formidabilis (Simon); and Ammotrecha, e.g.,
Ammotrecha limbata (Lucas).
VENOMOUS TICKS
Class Arachnida, Order Acarina
Ticks belonging to both familes, Ixodidae and Argasidae, may cause
local as well as systemic disturbances by their bite (see Chapter XX).
Ordinarily little or no injury results from the mere bite of an ixodine
tick; the writer has known of Dermacentor occidentalis Neumann and
Dermacentor variabilis (Say) to remain attached to a person for days
without causing great inconvenience and occaSionally quite unobserved
by the host. However, Nuttall (1911, loco cit.) records a number of cases
cited by other authors in which the bite of Ixodes ricinus (Linn.) has
caused serious consequences, notably a case described by Johannessen
of a "boy where the tick's body was removed but the capitulum re-
mained embedded in the skin at the back of the head. Swelling followed
at the point of injury, accompanied by headaches, stiffening and cramps
in the muscles of one side, partial loss of memory and polyuria; the pupils
became dilated, etc. The boy made a slow recovery." The bite of Ixodes
pacificus commonly results in more or less marked systemic disturbances
as observed by the author in California.
Quite a number of species belonging to the family Argasidae are
known to cause more or less serious consequences by their bites alone,
notably Ornithodoros moubata (Murray), O. coriaceus Koch, O. talate
(G.-M.), and O. turicata (Duges).
Ornithodoros moubata (Murray) has been reported repeatedly as
causing marked disturbances by its bite. Wellman, as quoted by Nuttall
and Warburton (1908, loco cit., p. 98), "states that the bite is very pain-
ful, the swelling and irritation (especially in Europeans) not subSiding
for days. The wheals are hard, raised and swell most disagreeably if
scratched, and this even a week after being bitten. The bite of young
ticks (nymphae) is said by the natives to be more severe than that of
the adults."
Ornithodoros coriaceus Koch. This species (Fig. 189) occurs com-
monly in the more mountainous coastal counties of California, having
been first described from Mexico, where it is known as the "tlalaja." In
California it is known as the "pajaroello." The writer has collected it on
Mount Hamilton where it flourishes in the deer beds among the low
scrub oaks (Quercus dumosa). The following description of the species
is a translation by Nuttall from the original:
Shaped like the sole of a shoe, thick margined, roughly shagreened, yellow-
ish-earthly color, spotted rusty-red, legs toothed dorsally. Length 9.3 mm. Body
about twice as long as wide, width fairly uniform, indented on the sides,
. pointed above the mouth parts, rounded posteriorly, a thick turned-up border
all around; the whole surface above and below thickly granulated like fish skin
(shagreen), the granules flat above, consequently, the whole leathery, on the
back unequal folds and grooves. Beneath in the front of the body a deep groove
FIG. 189. Showing Ornithodoros coriaceus just backing away from her
eggs recently deposited in the sand. Note protective coloration of the tick.
x5.
running to the stigmata and on the inner protrusion the rather large round
quite clearly marked eyes. The coxae gradually thicken toward the distal ex-
tremity and are somewhat bent; the other articles somewhat compressed and
clearly notched or round toothed. The whole surface, above and below, dirty
yellowish-earthy color, rusty-red spots irregularly distributed throughout.
Capitulum and palps light yellow. Legs gray-brown. Female.
The "pajaroello" is more feared than the rattlesnake by persons living
where it exists, and many harrowing tales are told regarding the loss of
an arm or leg, or even death resulting from its bite. Much of this is, of
course, gross exaggeration. However, the following is an account of two
bites which a former student suffered some years ago in July. He ex-
perienced a sharp pain on the left arm and upon rolling up his sleeve
discovered a large tick, partly engorged, attached to the upper arm in
front. He dislodged the tick and sucked the lesion. The lesion when
first discovered showed a small dark purple ring surrounding a bright
red spot, the point of attachment. The discoloration disappeared in a
short time, but the arm was "highly irritable for two or three days and
at the point of attachment a minute clear scab formed." The tick proved
to be a "pajaroello."
The second bite took place two weeks later while he was seated in
a thicket of willows (the first bite had occurred while he was riding over
FIG. 190. Egg of OrnithodO'ros

coriaceus, larvae in the act of emerg-
ing, and two fully emerged in-
dividuals. X 14.
a brush-grown hill), and in this case the sharp pain involved the left
leg. An almost fully engorged tick (a~ain a pajaroello) measuring about
three-quarters of an inch in length and about one-half inch in width
was removed from just above the shin. Once more a bright red spot was
visible at the point of attachment, surrounded by an irregular purple
ring about three-quarters of an inch in diameter. In about an hour the
leg began to swell in the vicinity of the lesion, and in about three hours
the entire lower leg was tremendously swollen. The coloration about
the point of attachment had widened conSiderably, was puffy, and a
clear lymph exuded freely from the lesion. The young man lanced the
wound, causing the blood to flow freely, and treated it with crystals of
potassium permanganate, binding the leg with cotton and gauze. During
the following night he reports experiencing a generally disagreeable
feeling, the entire lower leg "irritable and numb." On the following day
the bite on the arm became "irritable" again and was treated as had
been the leg as he feared bad results. For several weeks both lesions
exuded a clear lymph from beneath an "oily-looking, transparent, red
mottled scab" which remained in evidence for two or three months.
Life history of Ornithodoros coriaceus Koch. The "pajaroello" de-
posits large plum-colored spherical eggs (Fig. 190). In the laboratory
these are deposited on sand in slight depressions. There are commonly
four to seven layings at intervals of from several days to several weeks
during the months of May to July, inclusive (as early as February under
laboratory conditions), and the female is known to deposit eggs for at
least two successive seasons. The greatest number of eggs observed at
one laying was 802, with a total of 1,158 for one season. The incubation
period at a maintained temperature of from 24° to 26° C was from 19
to 29 days, with an average of about 22 days.
The larvae (Fig. 190) are very active, scattering quickly and attach-
ing readily to a host, particularly rabbits in the laboratory. Experimen-
tally the human may also serve as a larval host. The ear of a rabbit ap-
parently affords a most satisfactory point for attachment. The larva re-
mains attached to the host for a period of about seven days, becoming
quite globular and much enlarged. Under favorable conditions the larval
stage lasts 10 to 12 days.
Sexually differentiated ticks appeared after the fourth molt. Others
did not become sexually differentiated with five molts. There may be six
or seven nymphal stages. Ordinarily the tick molts once for each en-
gorgement, but there may be two molts between feedings. Smith47 re-
ports that reared females began ovipositing four to six months after the
first feeding. He records the adult longevity of reared, fed ticks as rang-
ing from about eight months to more than three years and seven months
for males and from 13 months to more than five years for a female.
CENTIPEDES ~ND MILLIPEDES

Class Myriapoda
Myriapoda. The Myriapoda are worm-like animals with separate
head, possessing antennae, and many fairly similar segments, each
possessing one or two pairs of segmented appendages. Like the insects
they are tracheated and for the most part terrestrial.
The class Myriapoda is divided into four or five orders, among which
are Chilopoda, the centipedes, with only one pair of appendages to each
segment; and the Diplopoda, the millipedes, with two pairs of append-
ages to each segment. They inhabit moist situations and feed principally
on decaying vegetable matter. An example is Julus nemorensis Koch, a
so-called "thousand-legged worm" (Fig. 8b).
Centipedes. The Chilopoda have only one pair of appendages to each
segment which are widely separated at the bases, the antennae are
many-jointed, the genital pore is located on the terminal body segment.
The larger species, at least, are predaceous, feeding mainly on insects.
Notwithstanding the confusing abundance of walking appendages, the
centipedes crawl very rapidly.
The centipedes are provided with powerful poison claws located
immediately ventral to the mouth and connected by means of a hollow
tube with large poison glands. The first pair of legs which terminate
in these claws are bent in position to form maxillipeds.
The larger centipedes (Fig. 191) are commonly regarded as veno-
mous and are generally much feared. Large insects are quickly killed
when the poison claws close upon them. It is true that the larger species
belonging to the genus Scolopendra and even the smaller house centi-
pede, Cermatia (= Scutigera) forceps (Raf.), are able to pierce the skin
FIG. 191. A venomous centi-

pede, Scolopendra heros. X .66.
with their poison claws and cause severe pain with same swelling at the
site of the bite. Cases are reported showing a double reddish streak on
the skin when the centipede had crawled. This condition may be caused
by the dragging sharp tips of the terminal pair of legs. Among the several
formidable-appearing species of the genus Scolopendra are S. heros
Gir. of the southern United States, S. polymorpha Wood of the south-
western states and Mexico, S. viridis Say, a Mexican species, and S.
sumichrasti Sauss. from the Canal Zone. All of these species may reach
a length of from five to six inches. Many of the centipedes are markedly
phosphorescent, notably Geophilus electricus (Linn.) and G. phos-
phoreus Gervais.
Millipedes. The DipJopoda are commonly separated into two groups
depending upon the presence or absence of repugnatorial glands. In the
Chilognatha these glands are present and are capable of producing
irritating effects (Burt' S ). Certain large tropical species, such as Rhino-
cricus spp., are able to squirt irritating fluids a distance of 18 to 33 inches.
Halstad and Ryckman 49 (1949) report a conclusively proven case of
vesicular dermatitis in Montemorelos, Nuevo Leon, Mexico, caused by
a diplopod of the genus Orthoporus.
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INDEX
._.
Bold face figures indicate bold face headings in text
Aarons, T., 276, 293 Aedes ( Continued)

Abalos, J. W., 107, 154 pionips Dyar, 192
Acalyptratae, 147 pseudo$cutellarias Theobald, 248
Acanthiidae, 95 punctor (Kirby), 191, 192
Acanthocephala, 43 $Capularn (Rondani), 240, 250
Acanthocheilonema perstam (Manson), scutellarias (Walker), 241
162, 249 seoulensis Yamada, 189
Acariasis, 35, 39, 525 simpsoni, Theobald, 189, 239
chorioptic, 533 sollicitans (Walker), 181, 188,243
intestinal, 556 squamiger (Coquillett), 181, 188
psoroptic, 525, 534 stimulans (Walker), 250
pulmonary, 545 stokesi Evans, 240
sarcoptic, 525, 526 taeniorhynchus, (Wiedemann), 188,
Acarina, 57, 58, 464, 525 243
Acarus Linn., 466 togoi (Theobald), 249
Acarus siro (see Tyroglyphus siro) tormentor Dyar and Knab, 249
Achorion schoenleini Lebert, 119 triseriatus (Say), 189,240,249
Ackert, J. E., 89, 338 varipalpus (Coquillett), 189,261
Addis, C. J., 160 ventrovittis Dyar, 191
Adler, S., 159 vexans (Meigen), 181, 182, 188, 264,
Aedes Meigen, 179, 182, 209, 246 265
boreal, 190 vittatus (Bigot), 238
flood-water, 188 Aedini, 176, 179, 182, 187
salt-marsh, 187 characteristics, 187
tree-bole, 189 Aerosols, 281
Aedes aegypti (Linn.), 3, 4, 5, 7, 176, Agramonte, A., 3, 233
192, 235, 237, 241, 242, 249, 250, Aitken, T. H. G., 207, 208, 311, 503
293 AZces americana americana Jardine, 476
africanus Theobald, 239 AlderHies, 55
albopictus (Skuse), 240, 241 Aldrich, J. M., 167,313,327,389
aldrichi Dyar and Knab, 181, 182 Alessandrini, G.,·225
argenteus (Poiret) (see Aedes aegypti) Aleurobius farinae (see Tyroglyphus
atropalpus (Coquillett), 249 farinae)
calopus (Meigen) (see Aedes aegypti) Alicata, J. E., 437
cantator (Coquillett), 243 Allergy, 39
cataphylla Dyar, 191 Allodermanyssus sanguineus (Hirst), 545
communis (DeGeer), 191, 192 Allolobophora chlorotica Sav., 327
communis tahoemis Dyar, 190, 191 Amara (Carabidae), 89
dorsalis (Meigen), 27, 187, 188, 189, Amblycera, 130
242, 277, 573 Amblyomma Koch, 479
flavescens (M uell. ), 192 Amblyomma americanum (Linn.), 479,
fluviatilis (Lutz), 240 489, 490, 495
hexodontus Dyar, 191 cajennense (Fabr.), 107,480,490
increpitus, Dyar, 191 hebraeum Koch, 498
leucocelaenus ( Dyar and Shannon) , variegatum (Fabr.), 498
239 Ammotrecha Banks, 603
luteocephalus Newstead, 189,240 Ammotrecha limbata (Lucas), 603
nearcticus Dyar, 191 Amoebiasis, 39
nigromaculis (Ludlow), 243 Anacimas Enderlein, 312
palustrn Dyar, 191 Anaphe infracta Walsingham, 576
611
Anaplasma marginale Theiler, 498 Anopheles (Continued)
marginale centrale Theiler, 498 pseudopunctipennis Theobald, 182,
Anaplasmosis, bovine, 56, 498 197, 207
Anarsia lineatella ZeI!., 554 pseudopunctipennis franciscanus Mc-
Anasa tristis (DeGeer), 95 Cracken, 197, 203, 206, 207, 224,
Ancona, H. L., 133 265, 276
Ancylostoma canium (Ercolani), 336 punctipennis (Say), 182, 193, 197,
duodenale (Dubini), 29 203, 205, 207, 224, 226, 249, 265,
Anderson, J. F., 121, 365 276
Andre, C., 335 quadrimaculatus Say, 182, 201, 203,
Angina pestosa, 439 205, 207, 224, 226, 227, 249, 261,
Anigstein, L., 495 264, 265, 272
Anisomorpha buprestoides (Stoll), 38 sacharovi Favr., 197, 198, 265
Annelida, 31 sargenti Theobald, 203
Anomiopsyllus nudatus Baker, 435 stephensi Liston, 226
Anopheles Meigen, 196, 208, 216, 246 subpictus Grassi, 225
biting habits, 205 urnbTosus Theobald, 202
breeding habits, 201 walkeri Theobald, 208, 249
egg characters, 198 Anophelines, hibernating, 226
life history, 203 key to North American, 208
mating, 197 malaria-carrying, 3
Anopheles albimanus Wiedemann, 17, North American, 206
203,205,208, 265 vectors of malaria, 227
atropos Dyar and Knab, 208 Anophelini, 176, 179, 195
aztecus Hoffman, 206 "Anophelism without malaria," 3, 225
barberi Coquillett, 189, 203, 208 Anoplura, 26, 55, 75, 109, U3, 127
bellator Dyar and Knab, 203, 274 Antarcophthirus trichechi (Boh.), 113
bradleyi King, 208 Anthocoridae, 110
crucians Wiedemann, 208 Anthonomus grandis Boh., 554
culicifacies Giles, 15, 196, 273 Anthomyidae, 147, 316
eluteus Edwards (see Anopheles sac- Anthrax, 2, 36, 40, 98, 302
harovi) Anthrenus museorum Linn., 86
farauti Laveran, 248, 280 scrophulariae (Linn.), 87
freeborni Aitken, 196, 197, 198, 201, Ant lions, 313
203, 206, 249, 261, 273 Antigen, flea, 563
gambiae Giles, 12, 249, 293 Antficola, 501, 518
georgianus King, 208 Antricola coprophilus (McIntosh), 518
grabhamii Theobald, 196 marginatus Banks, 518
hYTcanus sinensis Wiedemann, 196, Ants, 56, 570
245, 248 agricultural, 570
maculatus Theobald, 202 California fire, 570
maculipennis Meigen, 3, 27, 32, 176, California harvester, 570
198,201,224,227,242,573 larvae, 53
Mexican fire, 570
maculipennis var. atroparvus van Thiel, velvet, 571
198, 225 white, 55
var. elutus Edwards, 198 woolly, 571
var. labranchiae Falleroni, 198 Anto, 453
vaT. melanoi5n Hackett, 198 Antunes, W. A., 237
vat. messeae Falleroni, 198 Apatolestes Williston, 298, 311
vat. subalpinus Hackett and Lewis, Aphididae, 94
198 Aphid lion, 24
var. typicus Meigen, 198 Aphids, 55
maculipennis freeborni Aitken, 182 corn-root, 22
minimus Theobald, 203 Aphis foot, 533
minimus fiaviTostris Ludlow, 205, 264 Aphis maidiradicis Forbes, 22
occidentalis Dyar and Knab, 206, 224 Aphodius, 88
plumbeus Stephens, 203 Aptera, 54, 75
INDEX 613
Apterygota, 48, 49 Auchmeronmyia luteo14 (Fabr.), 385
Arachnida, 32,46, 56,464 Austen, Major E. E., 326, 327, 340, 352,
anatomy, external, 56 369, 385
internal, 58 Automeris io (Fabr.), 576
characteristics, 525, 578 Aviculariidae, 579, 594
claSSification, 58
development, 57 Babcock, O. G., 130, 387
mouth parts, 75 Babesia argentina Lignieres, 486
orders, key to, 59 bigemina (Smith and Kilbourne), 2,
Arachnidism, 587 40,41,466,483,485,486
Arachnolysin, 591 caballi (Nuttall), 497
Aragao, H. B., 250 canis (Piana and Galli-Valerio), 497
Aranea mactans Fabr. (see Latrodectus Babesiosis (see Piroplasmosis, canine)
mactans) Bacillus anthracis Cohn, 2, 40, 302, 331
Araneida (Araneae), 58, 578 botulinum (see Clostridium bOtfJli-
Arctiidae, 576 num)
Arctomys sibirica Schreber, 424,432 dysenteriae (see Shigel14 dysenteriae)
Argas, 501, 515 icteroides, 236
Argas americanus Packard (see Argas paratyphosus A, 333
persicus) pestis, 13
brumpti Neumann, 518 Back swimmers, 109, 574
rnianensis Brumpt, 518 Bacot, A. W., 419, 421, 430, 431
miniatus Koch (see Argas persicu8) Bacterium tu14rensis (see Pasteurel14
persicus ( Oken), 5, 467, 468, 470, tu14rensis) •
515, 517 Bader, M. N., 495
reflexus (Fabr.), 518 Badger, L. F., 437, 490
vespertilionis (Latreille), 518 Baer, W. S., 406
Argasidae, 467, 468, 470, 501,603 Baerg, W. ]., 579, 580, 588, 597, 598,
Argasinae, 466 599
Argiope argentata (Fabr.), 594 Baeus 14trodecti Dozier, 594
Argiopidae, 594 Bagdad boil (see Oriental sore)
Argyritarsis Robineau-Desvoidy, 196 Bailey, S. F., 575
ArUus cristatus (Linn.), 103, 108 Balfour, A., 517
Armillifer Sambon, 557 Bancroft, E., 1,2,245
Armillifer moniliformis (Diesing), 558 Bancroft's filaria ( see Wuchereria ban-
Arnold, F. T., Jr., 282 crofti)
Arribalzagia Theobald, 196 Bang, F., 546
Arthropoda, 31 Banks, N., 505, 525, 547
Arthropods, and disease, 35 Barber, M. A., 83, 279
intermediate hosts of helminths, 41 Barium carbonate, 453
pathological conditions caused by, 37 Bark lice, 55
venomous, 563 Barraud, P. J., 198
Ascaris Linn., 42 Bartonel14 bacilliformis Noguchi, 156
Ascaris equorum Goeze, 336 Bartonellosis, 156
lumbricoides Linn., 29 Bass, C. C., 217
Ashburn, P. M., 5, 240 Bastianelli, G., 3, 216
Ashcraft, J. B., 388 Basu, B. C., 305
Asopinae, 110 Bates, L. B., 503
Astacus spp., 42 Bats, relation to mosquitoes, 290
Atheta occidentalis, 90 Battistini, 157
Atrax formidabilis Cambridge, 580 Bauer, J. H., 4, 13, 237
robustus Cambridge, 580 Bdellonyssus Fonseca, 539
Atta (Formicidae), 22 Bdellonyssus bacoti (Hirst), 539
Attagenus pellio Linn., 86 brasiliensis (Fonseca), 539
Attaphi14 fungicola Wheeler, 22 bursa (Berlese), 540, 541, 547
Attidae, 595 nagayoi (Yamada), 539
Atylotus Osten Sacken, 312 Beauperthy, L. D., 2
Aubert, 5 Beck, M. D., 438, 505, 511
Bedbugs, 5,32, 35, 39, 95 Benacus griseus (Say), 574
big (see Triatoma sanguisuga) Bengston, I. A., 388
bites, 97 Bennett, B. L., 243
China (see Triatoma protracta) Bennett, D. H., 387
common, 96 Benzene hexachloride, 107, 593
control, 98 Bequaert, J. C., 152, 352, 359, 413, 414
disease transmission, 97, 107 Bequaertomyia Brennan, 311
Indian, 95 Berbillinae, 432
methods of distribution, 97 Berry, C. E., 459
tropical, 95 Berry, G. P., 243
Bedford, G. V., 409 Bertram, D. S., 539, 540
Bee sting, 564 Bethylidae, 572
morphology, 564 Betts, M. C., 451
operation of, 567 Beyer, G. E., 224
reaction to, 563 Big head, rabbit (see Myxomatosis)
treatment, 568 Bignami, A. E., 3, 14, 216
Bees, 53, 56 Bironella Theobald, 196
Beetles, 56, 85 Bishop, E. L., 263, 546
beaver, 92 Bishopp, F. C., 128, 133, 134, 136, 139.
blister, 89, 91, 576 364, 366, 394, 395, 413, 476, 488,
burying, 92 500,539,540,543,575
carpet, 38, 87 Biting lice (see Louse, biting)
carrion, 87, 92 Black death (see Plague)
cellar, 84 Blacklock, B., 385
characteristics, 86 Blacklock, D. B., 152,385
churchyard, 89 Black muzzle (sheep mange), 531
cockchafer, 88, 91 Black widow spider, 28, 33, 580
coconut, 90, 578 bite, 587
darkling ground, 91 control, 592
dung, 88 distribution and habitat, 582
fiat, 92 feeding habits, 583
grain, 90, 92 life history, 584
hide, 87 longevity, 586
intermediate hosts, 43, 87 mating habits, 583
June, 91 treatment of bite, 592
ladybird, 24 venom, nature of, 591
larder, 87 venom apparatus, 590
larvae, 53 Blaizot, L., 119
leaf, 91 Blake, F. G., 551
leather, 92 Blaps mortisaga Linn., 89
mandibles, 64 Blatella germanica (Linn.), 79, 81, 82
May, 88, 91 83
meal, 84 Blatta orientalis Linn., 78, 79, 81, 83
parasitic, 91 Blattaria, 55, 75
red-legged ham, 90 Blattidae, 43, 78
rodent, 91, 92 Blattner, R. J., 244, 546
rose chafer, 90 Blocking, 159
rove, 38,86 Blood worm (Chiromidae), 164
scavanger, 86 Blowflies, 25, 39, 146, 316, 327
skin, 92 control, 387
stink, 90 wool-maggot, 386
tule, 90 Blowfly strike (see Myiasis, cutaneous)
vesicating, 89 Blue, R., 428
Bell, E. J., 551 Bluebottle flies, 39
Belostoma, 574 Blyth, A. W., 591
Belostomatidae, 109, 574 Blyth, M. W., 591
Belschner, H. G., 387 Bogen, E., 587, 591, 592
Benacus. 574 Bolbodimyia Bigot, 312
INDEX 615
Bollinger, 0., 303 Bubo virginianus pacificus Cassin, 414
Book louse, 28, 55 Bugs, 94
Boophilus Curtice, 478, 480 ambush,110
Boophilus annulatus (Say), 2, 12, 33, 40, assassin, 11 0
41, 464, 466, 468, 469, 480, 485, back swimmers (see Back swimmers)
498, 499 bat, 109, 110
annulatus microplus Canestrini, 480, bedbug (see Bedbugs)
485, 498 blood-sucking, 574
australis Fuller (see Boophilus annu- chinch, III
latus microplus) conenose (see Conenose bugs)
bovis Riley (see Boophilus annulatus) copra, 90
decoloratus (Koch), 479, 485, 498 damsel, 111
BoOpiidae, 130 flower, 110
Booponus intonsus Aid., 388 giant water (see Water bugs)
Borell, A. E., 109 gnat (see Gnats)
Borrelia anserina, 467 leaf, 110
gallinarum Blanchard, 517 marsh treaders, 110
hermsi Davis, 512 poultry, 95
parkeri Davis, 512 shore, 110
recurrentis Lebert, 119, 120, ..510, 511 squash, 95, 111
Botflies, 25, 35, 146 stinkbug, 95
horse, 39, 147, 389 swallow, 95
robust, 146 toad, 109
sheep, 38 Burgess, R. W., 169, 293
Bothriocyrtum californicum Cambridge, Burke, A. W., 238
594 Burrill, A. C., 164
Bots, 389 Burroughs, A. L., 433, 436
pathogenesis, 393 Burt, E., 607
rodent, 405 Busvine, J. R., 127
sheep, 400 Buthidae, 598
treatment, 394 Buthus afer Leach, 564
Boutonneuse fever, 490 quinquestriatus Hemprich and Ehren-
Bovicola bovis (Linn.), 131, 132, 136, berg, 599
139 ButterHies, 50, 53, 56
caprae (Gurlt), 138 mouth parts, 74
Boyd, M. F., 204, 505 Buxton, P. A., 16, 116, 127, 203, 352,
Boynton, W. R., 499 356, 360
Brachetti, F., 600 Byam, W., 122
Brachycera, 144, 145
Brachythele longitarsus Simon, 594 Caddis Hies, 56
Bradley, G. R., 285 Cahn, A. R., 476
Braun, M., 373, 537 Calabar swellings, 307
Brennan, J. M., 311, 547 Calero, C., 383
Brenton, 488 Calliphora Robineau-Desvoidy, 327
Briggs, L. R., 504 Calliphora erythrocephala (Meigen) (see
Brill's disease, 120, 437 Calliphora vicina)
Bristletails, 54 vicina Robineau-Desvoidy, 327, 331,
Brody, A. L., 250 375
Brookman, B., 244, 546 vomitoria (Linn.), 327, 331, 374
Broteas alieni ( Wood), 599 Calliphoridae, 316, 327, 334, 372
Brown, H. W., 246, 247, 539 Callitroga americana Cushing and Pat-
Brown-tail rash, 575 ton, 39, 380, 382, 383, 384, 387
Bruce, D., 2, 355, 356, 357 hominivorax (Coquillett), 383
Bruce, W. G., 384 macellaria (Fabr.), 384
Bruce, W. N., 420 Calmette, A., 564
Brues, C. T., 157 Calyptratae, 146
Brug's filaria (see Wuchereria malayi) Cambournac, F. J. c., 273
Brumpt, E., 105, 497 Cameron, A. E., 530
Cameron, D., 541 Cercopithecus patas, 114
Cancer, Gongylonema pulchrum as agent Ceresa bubalus (Fabr.), 574
of, 83 C ermatia forceps (Raf.), 607
Canthariasis, 89 Cestoda, 29, 42 (see Tapeworm)
Cantharidae (Coleoptera), 576 C etonia aurata (Linn.), 88
Cantharidin, 577, 578 Chactidae, 598, 599
Carahidae (Coleoptera), 89, 91 Chaetopoda, 31
Carbon bisulfide, in rodent control, 457 Chaetotaxy, 144
Cardosa, E., 238, 239 Cbagas, C., 6, 104
Carpenter, S. J., 293 Chagas' disease, 6, 37, 41, 44, lO4
Carri6n's disease, 156 control, 107
Carroll, J., 193, 233 transmission of infection, 105
Carroll, J. H., 122 Chagasia Cruz, 195, 196
Carter, H. R., 19, 205, 233, 237, 257 Chalmers, A. J., 578
Case, A. A., 89 Chalybion coeruleum (Linn.), 570
Caseflies, 56 cyaneum (Klug), 593
Caseworms, 42 Chandler, A. C., 240, 241
Castaneda, M. R., 437 Chaoboridae, 145, 165
Castellani, A., 1, 333, 355, 356 Chaoborus astictopus Dyer and Shannon,
Caterpillars, 53 166
with irritating hairs, 576 lacustris Freeborn, 166
"puss" (see Megalopyge opercularis) Chapin, R. M., 499
range (see H emileuca oliviae) Cheese skippers, 376
saddle-back (see Sibine stimulea) Chelonethida, 58
Cattani, J., 333 Cheshire, F. R., 565, 567
Cattle grubs, 394 Chevril, R., 378
economic losses, 317 Cheyletidae, 557
injury, 397 Chigger mites, 547
life history, 395 control, 550
treatment, 398 dermatitis, 550
Cattle tick, southern (see Boophilus an- Chilognatha, 607
nulatus) Chilomastix mesnili (Wenyon), 336
Causey, O. R., 374 Chilopoda, 31, 606
Cavia asperea Pallas, 433 Chironomidae, 146, 160, 164
Ceder, E. T., 437 Chironomus plumosus Burrill, 165
Centipedes, 31, 606 Chitwood, B. G., 84
Centruroides californicus (Girard), 599 Chlordane, 85, 107, 281, 289, 593
carolinianus (see Centruroides vittatus) Chloropidae, 147, 166
gertschi Stahnke, 598 Chlorotabanus Lutz, 312
nigrescens ( Pocock) , 599 Choanotaenia infundibulum (Bloch), 42,
sculpturatus EWing, 598 338
sutfusus Pocock, 597, 598 Cholera, 39, 333
Centrurus gracilis (Latreille), 597 Choriomeningitis, lymphatic, 496
C ephalomyia ovis (Linn.), 405 Chorioptes Gervais and van Beneden, 553
Cephenemyia ;ellisoni Townsend, 402 Chorioptes bovis (Gerlach), 533
phobiger Clark, 402 equi (Gerlach) (see Chorioptes sym-
pratti Hunter, 402 biotes)
rufibarbus Meigen, 402 ovis (Railliet), 534
stimulator Clark, 402 symbiotes Verheyen, 533
trompe Linn., 402 Chowning, W. M., 488
ulrichii Brauer, 402 Christophers, S. R., 180, 198, 221, 230
Ceratophyllus acutus (see Diamanus Chrysomelidae, 91
montanus) Chrysomya albiceps (Wiedemann), 387
gallinae Schrank, 421, 431, 445 bezziana (Villen.), 387
idahoensis (see Oropsylla idahoensis) dux ( Esch. ) , 387
niger Fox, 441, 445 flaviceps (Walker), 387
Ceratopogon stellifer Coquillett, 161 megacephala (Fahr.), 374, 380
Ceratopogonidae, 146, 160 rufifacies (Macq.), 387
INDEX 617
Chrysopidae, 24 Cockroaches (Continued)
Chrysops Meigen, 6, 297, 311 American, 80
Chrysops americana Osten Sacken, 310 Australian, 80
callida Osten Sacken, 310 control, 84
celer Osten Sacken, S10 Cuban, 81
dimidiata v. d. Wulp, 249, 306, 310 digestive system, 51
discalis Williston, 6, :306, 310 dusty-tail, 81
hyalina Shannon, 311 feeding habits, 78
silacea Austen, 306 German, 79
Chrysozoma Meigen, 298, 311 hosts of threadworms, 43
Chun, J. W. H., 423, 440 intermediate hosts of nematode para-
Chung, Huei-Lan, 504 sites, 83
Cicadas, 55, 94 life history, 81
Cicadellidae, 94, 574 mouth parts, 62
Cicadidae, 94 Oriental, 79
Cimex adiunctus, 100 species of sanitary importance, 79
boueti J oyeux, 95 Surinam, 81
columbarius J enyns, 95 tropical, 80
hemipterus (Fabr.), 95, 98, 100 vectors, 82
lectularis Linn., 32, 95, 96, 98, 100, Coenurus cerebra lis (Batsch) (see Multi-
306 ceps multiceps)
pilosellus (Horv.), 96, 100 Colchicum autumnale, 29
pipistrelli Jenyns, 95 Cole, M. M., 474
rotundatus (Sign.), 95, 98 Coleoptera, 56, 75, 85
Cimexopsis nyctalis List, 100 families, key to, 91
Cimiddae, 95, 110 larvae, 53
key to North Americ~n, 99 Collembola, 54, 75
Cirripedes, 25 Colobus caudatus, 114
Citellophilus tesquorum (Wagner), 433 Columbicola columbae (Linn.), 131, 134
Citellus Oken, 434, 454 Commensalism, 22
Citellus beecheyi beecheyi (Richardson), Compound 1080, 453, 457
431, 435 Comstock, J. H., 48, 49, 148, 576, 584
beecheyi douglasii (1lichardson), 454 Comte, C., 6, 121
beecheyi fisheri (Merriam), 454 Conenose bugs, 6, 28, 37, 41, 100, 250
beldingi ~ Merriam), 4'54 ni'tes, \W~, '5'1~
columbianus (Ord), 434 bloodsucking, 104
oregonus (Merriam), 454 China bedbug, 103
richardsoni (Sabine), 243 cross bug, 103
Clark, P. W., 452 kissing bug, 102
Clostr.idium botulinum (Van Ermen- life history, 101
gem), 388 Mexican bedbug, 104
Cnemidocoptes laevis vllr. gallinae (Rail- spotted corsair, 104
liet) , 533 wheel bug, 103
mutans (Robin and Lanquentin), 532 Conjunctivitis, 167
Cnephia minus (Dyat and Shannon), habronemic, 337
150 Connor, M. E., 292
pecuarum (Riley), 150 Conorhinus megistus (Burm.), 105
Coccidiida, 217 Conseil, E., 6, 119, 121
Coccinellidae, 24 Cook, S. S., 285
Cochliomyia americana Cushing and Cooley, R. A., 468, 473, 474, 475, 480,
Patton, 39, 380 492, 501, 503, 513
hominivorax (Co(juillett), 383 Cope, O. B., 289
macellaria (Fabr.), :380, 384 Copeman, S. M., 323, 340
Cochran, J. R., 205 Coprobionts, 372
Cockchafers, 88, 91 Coquillettidia Dyar, 209
Cockerell, T. D. A., 78 Cordylobia anthropophaga (E. Blanch-
Cockroaches, 28, 55, 78 ard), 385
agents of contamination, 36 Cordyluridae, 147
Coreidae, 111 Culicoides (Continued)
Corizidae, 574 dovei Hall, 161
C orizus hyalinus (Fabr.), 38 fmens Poey, 163
Cornell, V. H., 243, 505 grahami Austen, 163
Corrodentia, 55, 75 gutipennis Coquillett, 161
Corsair, two-spotted (see Rasahus bigut- melleus Coquillett, 161
tatus ) Culiseta incidens (Thomson), 180, 186,
Cotton-boll weevil (see Anthonomus 276
grandis) inornata (Williston), 187
Courtet, M., 224 morsitans Theobald, 187
Covell, G., 230 Cumming, R G., 352
Cowles, R B., 594 Curran, C. H., 143, 145, 146, 316, 327,
Cox, T. A., 153 390
C oxie1la burneti (Derrick), 495, 496 Cushing, E. C., 380, 384
Crab louse (see Louse, crab) Cuterebra Clark, 405
Craig, C. F., 5, 240 Cuterebra beameri Hall, 405
Crayfish, 31, 42 buccata Fabr., 405
Cremastogastor (Formicidae), 181 emasculator Fitch, 405
Creophilus (Staphylinidae), 87 horripilam Clark, 405
Cresylic acid, 274, 275, 278 peromysci Delmat, 405
Crickets, 54 tenebrosa Coquillett, 405
Crithidia Leger, 7 Cuterebridae, 146, 389, 402, 40.5
Croton bug, 79 Cyclops, 43
Crouch, W. E., 451 Cyclorrhapha, 144, 145, 146, 299
Crustacea, 31, 43 Cynomys, 434
Cryptocerata, 95, 109 Cyprinodon variegatus, 292
Ctenocephalides canis (Curtis), 42, 140, Cytoleichidae, 557
421, 439, 446, 563 Cytoleichus nudus (Vizioli), 557
felis (Bouche), 420, 421, 439, 446
Ctenopsyllus segnis (SchOnch), 421, 435, Da Fonseca, F., 538, 541
443 Dalmat, H. T., 405
Cuclutogaster heterographus Nitzsch, 133 Dampf, A., 152, 163
Cucujidae (Coleoptera), 90, 92 Damsel Hies (Odonata), 55
Culex Linn., 209, 246 Darling, S. T., 224, 336
Culex annulirostris Skuse, 245 Da Rocha-Lima, H., 6, 122
apicalis Adams, 181 Das Gupta, B. M., 20
erraticus Dyar and Knab, 272 Davidson, A., 104
fasciatus Fabr. (see Aedes aegypti) Davis, G. E., 98, 475, 493, 495, 502, 509
fatigans Wiedemann, 2, 185 Davis, N. C., 237
pipiens Linn., 176, 185,274,276,546 DayHies, 55
pipiens molestus ForskiU, 185 DDT, 7, 85
pipiens pallens Coquillett, 185, 245, bedbug control, 98, 107
248 Hea control, 447
quinquefasciatus Say, 248, 274 Hy control, 322, 335, 342, 345, 346,
quinquefasciatus Wiedemann, 241, 246 360
tarsalis Coquillett, 182, 186, 244, 273, gnat control, 154, 160, 168, 166, 170
276 lousicide, 125, 130
thalassius Theobald, 240 mite control, 540, 551
tritaeniorhynchus Giles, 245 mosquito control, 269, 274, 289
Culicidae, 145, 175 spider control, 593
key, 208 tick control, 500, 501, 512
Culicini, 176, 182, 209 Deay, H. 0., 81
characters, 185 Deer Hies, 6, 146, 297, 306, 412
Culicoides Latreille, 161 Deinocerites Theobald, 176, 209
Culicoides austeni Carter, Ingram, and Delhaye, R, 374
Macfie, 162, 249 Delhi boil (see Oriental sore)
canithorax' Hoffman, 161 Delousing, 123
diabolicus Hoffman, 161 Demodex bovis Stiles. 537
INDEX 619
Demodex (Continued) Diamanus montanus (Baker), 13, 421,
cati Megnin, 537 431,433,435,436,441
canis Leydig, 537 Dicladocera Lutz, 312
equi Railliet, 537 Dichloro-diphenyl-trichloroethane (see
folliculorum Simon, 39, 537 DDT)
phylloides Csokor, 537 Dimethylphthalate, 154, 160, 554
Demodicidae, 537 Dipetalonema perstans (Manson), 162
Dendrolimus pini (Linn.), 576 Diphyllobothrium latum (Linn.), 336
Dengue fever, 5, 192, 240 Diplocentrus whitei (Gerv.), 599
Mediterranean (see Sand fly fever) Diplococcus pemphigi contagiosi Wherry,
mosquito transmission of, 240 119
Dennis, E. W., 486 Diploglossata, 55, 75
Dermacentor, 473 Diplopoda, 31, 606
key to genus, 477 Dips, arsenical, 499
Dermacentor albipictus (Packard), 464, benzene hexachloride, 130, 500
476,477,489,498,500 DDT emulsion, 130, 500
andersoni Stiles, 5, 6, 27, 41, 243, 306, sulfur-rotenone, 130, 499
464, 465, 466, 469, 470, 474, 477, Diptera, 48, 51, 56, 75, 143
488, 489, 493, 494, 495, 496, 498, classification, 144
499 larvae, 53
halli McIntosh, 477 mandibles, 64
hunteri Bishopp, 477 Dipylidium caninum (Linn.), 5, 42, 140,
nitens Neumann, 472 336, 447
occidentalis Marx, 477 Dirofilaria immitis (Leidy), 249
occidentalis Neumann, 5, 468, 489, Disease control, cost of, 19
490, 493, 498, 603 Disease transmission, 39
parumapertus Neumann, 468, 477 by bedbugs, 97
parumapertus marginatus Banks, 489, control, 16
490 cyclico-developmental, 40
reticulatus (Fabr.), 497 cyclico-propagative, 40
variabilis (Say), 474, 477, 488, 489, fecal, 40
490,493,495,498,500,603 hereditary, 41
venustus (see Dermacentor andersoni) propagative, 40
Dermacentroxenus rickettsi Walbach, 5, transovarian, 41
488 Diseases, gastrointestinal, 332
Dermanyssidae, 538, 542 Dixidae, 145, 165
Dermanyssus Duges, 542 Dobsonflies, 55
Dermanyssus americanus Ewing, 545 Docophorus cygni Denny, 133
gallinae (DeGeer), 33, 542, 546 icterodes Nitzsch, 133
Dermaptera, 55, 75 Dodge, H. R., 546
Dermatitis, chigger, 550 Doerr, R., 5
epidemics, 555 Doetschman, 548
Dermatobia Brauer, 403 Dolichoderinae, 570
Dermatobia hominis (Linn. ) 39, 402, Dolichopsyllidae, 438, 441
405 Dolve, R. M., 339
Dermatophilus penetrans (Linn.), 442 Donat, F., 6, 107
Dermatosis, 38 (see Acariasis) Dorman, S. C., 328
Dermestes lardarius Linn., 87 Dormitator latifrons, 292
vulpinus Fabr., 86, 87 Dorylinae, 570
Dermestidae, 86, 87, 92 Doty, A. H., 257
Derris, 127, 136, 398 Doty, R. E., 450
DesoH, P., 374 Douglas, J. R., 436, 459, 476
de Souza, G. S., 1 Dove, W. E., 161, 287, 539
Devil's darning needles, 55 Downs, W. G., 283
Dewevre, L., 5, 118, 119 Dracunculus medinensis (Linn.), 43
D'Herelle, F., 225 Dragonflies, 42, 55, 290 .
Diachlorus Osten Sacken, 312 Drosophila, 166
Diamanus Jordan. 441 Duck clubs, as mosquito hazards, 288
Duke, H. L., 27 Eumops perotis californicus (Merriam) ,
Dumdum fever (see Kala-azar) 109
Dunn, L. H., 319, 403, 434, 503 Euphorbia, 29
Dust louse, 55 Euphorbiaceae, 7
Dutton, J. E., 5, 355, 356, 357, 467, 503 Euproctis chrysorrhea (Linn.), 564, 576
Dyar, H. G., 150, 165, 179, 190 Eurypelma californica Ausserer, 579
Dyer, R. E., 437, 490, 495 Eusimulium minus Dyar and Shannon,
150
Earwigs, 55 pecuarum (Riley), 150
Echidnophaga gallinacea ( Westwood), Eutamias quadrimaculatus, 511
437, 442, 445, 459 Eutettix tenellus (Bak.), 574
Echinophthiriidae, 113 Eutrombicula Ewing, 548
Echinophthirus phocae (Lucas), 114 Eutrombicula alfreddugesi (Oudemans),
Ecology, importance of, 14 39,548,549
Ectoparasites, 24, 41 batatas (Linn.), 548, 549
Edwards, F. W., 175, 176, 184, 185, 196 masoni (Ewing), 548, 549
EI debab, 307 Evans, A. M., 151
Elephantiasis, 245, 247 Evans, F. C., 459, 546
Ellenberger, W. P., 499 Ewing, H. E., 438, 525, 542, 543, 545,
Ellis, E. F., '588 547,549,550,574,598,599,601
Elmendorf, J. E., Jr., 230 Exopterygota, 48
Elson, J. A., 10l Eyles, D. E., 182, 249
Embioptera, 55, 75 Eypris, 571
Encephalitis, 186 Eypris californicus (Ashmead), 572
Japanese "B" virus, 244
St. Louis, 546 Faichnie, N., 333
Encephalomyelitis, equine, 7, 242, 546 Falleroni, D., 3, 198, 225
Endamoeba coli (Grassi), 336 Fallis, A. M., 401
histolytica (Schaudinn), 29, 333 Fannia Robineau-Desvoidy, 324
typhosa, 336 Fannia canicularis (Linn.), 324
Enderlein, G., 299 incisurata (Zett.), 325
Endoparasites, 24 manicata (Meigen), 325
Endopterygota, 48 scalaris (Fabr.), 325, 331, 378
Enicocephalidae, III Fantham, H. B., 5
Enterobius Leach, 42 Fasciola hepatica Linn., 31
Enterobius vermicularis (Linn.), 39, 336 Faust, E. C., 14, 444, 558
Entomophobia, 37 Favus, 118
Envenomization, 38 F elicola subrostratus (Nitzsch), 139
Eomenacanthus stramineus Nitzsch, 132, Feng, Lan-Chou, 190, 504
133 Fernald, H. T., 18
Ephemerida, 42, 55, 75 Ferris, G. F., 91, 109, 113, 114, 137, 414, ...
Epicauta hirticornis (Haag-Rutenberg), 415
89 Fever, breakbone (see Dengue fever)
Epizootics, 43 Bullis, 495
Eremobates Banks, 602 Colorado tick, 492
Eremobates formicarius (Koch), 602 deer-fly (see Tularemia)
formidabilis (Simon), 603 dengue (see Dengue fever)
Eretmapodites chrysogaster Graham, 240 dumdum (see Kala-azar)
Eristalis Latreille, 378 East coast, 496
Eristalis tenax (Linn.), 376, 377 estivo-autumnal, 214, 217
Erythroneura comes (Say), 574 five-day (see Fever, trench)
Escomel, E., 595 jail (see Typhus fever)
Esenbeckia Rondani, 311 Japanese flood (see Tsutsugamushi
Eskey, C. R., 431, 436, 441 disease)
Espundia (see Leishmaniasis, American Kedani (see Tsutsugamushi disease)
mucocutaneous) araya, 156
Essig, E. 0., 54, 572 phlebotamus (see Pappataci fever)
Esten, W. N., 83, 330 Q (see Q fever)
INDEX 621
Fever (Continued) Flies (C ontinued)
quartan, 214. 218 caribou warble, 400
quotidian, 214 cheese, 376
relapsing (see Relapsing fever) chin, 392
seven-day (see Sand fly fever) crane, 145
tertian, 214, 217 deer (see Deer flies)
three-day (see Sand fly fever) drone, 377
trench, 122 . ear, 6
van der Scheer (see Sand fly fever) eye, 166
yellow ( see Yellow fever) flesh (see Flesh flies)
Fibiger, J., 83 flower, 146
Figley, K. D., 39 frit, 147, 166
Filaria bancrofti ( Cobbold) (see gad, 6,302
Wuchereria bancrofti) Glossina (see Tsetse flies)
loa (Cobbold), 6 greenbottle (see Greenbottle Hies)
FilariasiS, 2, 245 greenhead (see Horseflies)
asymptomatic, 247 head-maggot (see Head-maggot flies)
Bancroft's, 247 heel, 394
heartworm, 249 Hippelates (see Hippelates flies)
inflammatory, 247 horn (see Horn flies)
malayi, 248 horse (see Horseflies)
mosqUito species involved in, 248 house (see Houseflies)
obstructive, 247 louse (see Louse flies)
vectors, 248 mango, 306
Filariidae, 43 moth, 145, 154
Finlay, C., 3, 233 nose, 389, 392
Flea control, 447 oestrid, 389
Flea repellents, 448 ox warble, 402
Fleas, 39, 56 Phlebotomus, 154, 155
chigoe, 56, 443 pigeon, 413
commoner species, 438 pomace, 166
control, 447 Psychoda, 154
digestive tract, 419 sand (see Sand flies)
ground-squirrel, 13 screwworm (see Screwworm flies)
hosts, 421 snipe, 146, 313
as hosts of cestodes, 447 soldier, 376
household, 445 stable (see Stomoxys flies)
human, 29 tachina, 146
larvae, 53 tsetse (see Tsetse Hies)
life history, 419 tumbu,385
longeVity, 421 warble, 39, 400, 402
mandibles, 64 wool-maggot (see Flesh Hies, Blow-
mouth parts, 73, 416 Hies)
plague infected, 435 Florence, L., 128
in plague transmission, 36 Florio, L., 493
rat, 36, 442 Flugge, C., 5
sticktight, 29, 445 Fluke (Trematoda), 25, 31, 42
structural characteristics, 416 liver ( see Fasciola hepatica)
western hen, 445 lung, 42
Fleshflies, 25, 316, 327, 379 poultry, 42
Fletcher, J., 368 Fly control, 338
Flies, 1, 56, 316 community, 344
bat, 414 DDT, 345
black, 146, 147, 151 garbage cans, 346
blow (see Blowflies) privies, 348
bluebottle (see Bluebottle flies) septic tanks, 347
bot (see Botflies) sewage treatment plants, 348
buffalo, 369 fiytraps. 349
Fly control (Continued) Gasterophilus (Continued)
rural, 338 intestinalis (DeGeer), 39, 389, 390
DDT, 342 nasalis (Linn.), 389, 392
dairy barns, 341 pecorum (Fabr.), 393
manure management, 339 veterinus (Clark), 392
Fly larvae, bots, 389 Gasterophilus Leach, 146, 389
cattle grubs, 394 Gastiaburu, J. C., 157
head bots, 389 Gaurax araneae Coquillett, 594
warbles, 389 Gelastocoridae, 109
Folger, A. R., 364 Geophilus electricus (Linn.), 607
Forde, R. M., 5, 355 phosphoreus Gervais, 607
Formica rofa Linn., 22 Gerrhonotus multicarinatus webii
Formicidae, 570 ( Baird), 594
Formicinae, 570 Gerridae, 110
Foster, M. R., 124 Gerroidea, 110
Fowl-pox, 250 Giardia lamblia Stiles, 336
Fox, C., 425, 443 Gibson, N. R. E., 165
Fox, I., 438 Gigantorhynchus gigas (Bloch), 43, 88
Framboesia tropica (see Yaws) Gilbert, E. W., 592
Franca, M., 239 Gilbert, Q. 0., 375
Francis, E., 6, 98, 245, 305, 306 Gill, C. A., 333
Franz., K., ;) Gillette, H. 1'. S., 274
Freeborn, S. B., 183, 189, 200, 364 Ginsburg, J. M., 279, 280, 285
Frogatt, W. W., 386, 533 Gjullin, C. M., 181
Frost, F. M., 198, 201 Glasgow, R. D., 154
Frost, W. R., 365 Gliptocranium gasteracanthoides Nicolet,
Fumigation, for bedbugs, 99 595
calcium cyanide, 454 Gliricola porcelli (Linn.), 130
carbon bisulfide, 457 Glossina Wiedemann, 352
carbon monoxide, 454 Glossina austeni Newstead, 358, 359
clothing, 123 brevipalpus N ewstead, 358
hydrocyanic acid, 99 caliginea Austen, 359
lice (body), 123 fusca ( Walker), 358
methyl bromide, 124, 454, 458 fusca var. congolensis Newstead and
rats (burrows), 454 Evans, 358
squirrels (ground), 457 fuscipleuris Austen, 358
sulfur dioxide, 454 honingtoni Newstead and Evans, 358
Fundulus diaphanus, 292 longipalpus (Wiedemann), 358, 359
dispar, 292 longipennis Corti, 358
heterocliteus, 292 medicorum Austen, 358
maialis, 292 moristans Westwood, 2, 5, 355, 358,
similis, 292 359
Furman, D. P., 130, 139, 499, 546, 548 moristans var. pallida Shire., 359
moristans var. paradoxa Shire., 359
Gahan, J. B., 283 nigrofusca N ewstead, 359
Gajardo-Tobar, R., 383 pallicera Bigot, 359
Galeb, 0., 83 pollidipes Austen, 358, 359
Gallinippers, American, 175 polpalis (Robineau-Desvoidy), 5, 352,
Galli-Valerio, B., 29 356,357,358,359
Gallus domesticus, 540, 541 polpalis var. maculata Newstead, 359
Gambusia afJinis (Baird and Girard), 291 polpalis var. wellmani Austen, 359
Gammexane, 154 schwetzi Newstead and Evans, 359
Garman, R., 302 severini Newstead, 359
Gasterophilidae, 147, 389 submoristans Newstead, 359
as human parasites, 404 swynnertoni Austen, 358, 359
Gasterophilus equi (Clark), 390 tabaniformis West., 359
haemOf'rhoidalis (Linn.), 389, 392 Gnats, 56, 143
inermis (Brauer), 389, 392 bite, 151
INDEX 623
Gnats (Continued) Gupta, P. C. Sen, 158
bloodsucking, 6 GutberIet, J. E., 338
Bodega black, 161 Gymnocerata, 95, 109
buffalo, 146, 147, 148, 151 Gynaitkothrips uzeli Zimmerman, 575
chaoborid, 165, 175 Gyropidae, 130
characteristics, 147 Gyropus ovalis Nitzsch, 130
cholera, 151
Columbacz, 151 Haas, V. H., 431, 436
control, 153, 161 Habronemiasis, 337
eye, 166 Habronema Diesing, 337
life history, 148 Habronema muscae (Carter), 837
relation to disease, 151 Hackett, L. W., 3, 198, 225
Goldberger, J., 121, 151, 555 Hadrurus aztecus Pocock, 600
Goldman, L., 159 Hadwen, S., 395, 494
Golgi, C., 185, 215, 219 Haeckel, E. H., 15
Gongylonema pulchrum Molin, 43, 83, Haemagogus capricornii (Lutz), 239
88 equinus Theobald, 240
Gongylonemiasis, 43, 88 spegazzini Brethes, 239
Goniocotes bidentatus Scopoli, 134 splendens Williston, 240
chrysocephalus Gieb., 134 Haemaphysalis Koch, 477
hologastes Nitzsch, 133 Haemaphysalis cinnabarina Koch, 493
rectangulatus Nitzsch, 134 humerosa Warburton and Nuttall, 495
Goniodes colchicus Denny, 134 leachi (Audoin), 478, 497
damicornis Nitzsch, 134 leporis-palustris (Packard), 477, 489,
dissimilis Nitzsch, 133 493
falcicornis Nitzsch, 134 Haematobia irritans Robineau-Desvoidy
gigas Tasch., 133 (see H aematobia stimulans)
numidianus Denny, 134 serrata Robineau-Desvoidy, 242,366
stylifer Nitzsch, 131, 133 stimulans Meigen, 366
Goniops Aldrich, 298, 311 Haematomyzidae, 114
Good, N. E., 438 Haematomyzus elephantis Piaget, 1I4
Gordon, R. M., 539, 540 Haematopinidae, 114, 127
Gorgas, W. C., 257 Haematopinoididae, 114
Gorrie, R. H., 546 Haematopinus asini (Linn.), 129
Gouck, H. K., 474, 501 eurysternus Nitzsch, 128, 130
Gould, G. E., 81 macrocephalus (Burm.), 129
Graham, G. M., 505 quadripertusus Fahr., 128
Graham, H., 5, 241 mis (Linn.), 114, 127
Graham-Smith, G. S., 166, 331 tuberculatus (Burm.), 128
Granette, P., 287 urius Nitzsch, 127
Grasshopper, 49, 51, 54 Haematosiphon inodora (Duges), 95, 99
Grassi, B., 3, 14, 216, 225, 338 Haemodipsus ventricosus (Denny), 129,
Gray, H. F., 257, 265, 283, 285 306
Gray, J. A., 546 Haemoproteus columbae Celli and San
Graybill, H. W., 470, 480 Felice, 414
Greenberg, M., 545 lophortyx O'Roke, 414
Greenbottle Hies, 39, 379 Haemosporiidea, 217
Greenwald, M., 205 Hairs, urticarial, 575
Griffitts, T. H. D., 205 Halarachne zalophi Oudemans, 546
Grinnell, M. E., 114 Hale, W. C., 328
Grocer's itch, 556 Hall, D. G., 161, 169,327,328,334
Grubs, 53 (see also Cattle grubs) Hall, M. C., 41, 42, 83, 103, 129, 376,
Griinberg, K., 353 394
Grundmann, A. W., 243 Hall, W. W., 591
Grylloblattodea, 55, 75 Halstad, B. W., 607
Guberlet, J. E., 389 Halter, B. L., 592
Guinea worm, 43 Hamadryas, 114
Gunderson, 342 Hamadryas antiopa (Linn.), 576
Hamlyn-Harris, R., 292 HesS, A. D., 290
Hammer, 0.,366 H eterandria formosa, 292
Hammon, W. M., 206, 243, 244, 493, Heterodoxus armiferus Paine (see He-
546, 547 terodoxus longitarsus)
Hampton, B. C., 424 longitarsus Pia get, 130, 139
Hardman, N. H., 224 Heterometabola, 50
Hare, J. E., 412 Heteropoda venatoria Koch, 594
Haring, C. M., 242 Heteropodidae, 594
H armolita grandis (Riley), 554 Hewitt, C. G., 285, 316
Harpactocorinae, 100, 108 He){apoda, 31, 46
Harpagomyia de Meijere, 181 Heymons, R., 557
Harpalinae, 89 Heys, F. M., 244, .546
Harrison, L., 130 Hill, H. R., 559
Harvestmen, 58 Hill, R. B., 273
Harwood, P. D., 376 Hindle, E., 158, 192, 224, 503
Hasseltine, H. E., 443 Hine, J. S., 299, 309
Hawes, I. L., 114 Hinman, E. H., 41, 161, 246, 249, 290,
Hayashi, N., 551 593
Hayne, J. B., 279 Hippelates, 39
Hazato, H., 551 classification, 169
Head-maggot fly, 38, 400 Hippelates bicolor Coquillett, 170
1ife history, 400 bishoppi Sabrosky, 170
Hearle, E., 399, 400, 476 collusor Townsend, 170
Heartwater, 498 convexus Loew, 170
Heartworm, of dogs, 249 dissidens (Tucker), 170
Hebridae, 1l0, III dorsalis Loew, 170
Hectopsyllidae, 438, 442 {lavipes Loew, 167, 170
Hegh, E., 292, 352 hermsi Sabrosky, 170
Heim, F., 86 microcentrus Coquillet, 170
Heliothrips indicus Bagnall, 575 montanus Sabrosky, 170
Helminths, 27, 39, 83, 84 nobilis Loew, 170
intermediate hosts of, 41 pallipes Loew, 168, 170
Hemileuca maia Drury, 576 particeps Becker, 170
oliviae Cockerell, 575 plebqus Loew, 170
Hemimerids, 55 proboscideus Will., 170
Hemiptera, 48, 55, 75, 94
pusio Loew, 167, 168, 170
mandibles, 64, 66 robertsoni Sabrosky, 170
mouth parts, 65
Hippelates flies, 166, 336
Hemiptera-Heteroptera, 48, 94
key to, 109 classification, 169
Hemiptera-Homoptera, 48, 94 control, 170
Henning, M. W., 493, 496 Hippobosca Linn., 413
Herman, C. M., 231, 413, 414 Hippobosca camelina Leach, 413
Hermetia illucens (Linn.), 376 capensis v. Olfers (see Hippobosca
Herms, H. P., 242 longipennis)
Herms, W. B., 11, 12, 17, 19, 26, 37, 82, equina Linn., 413
91, 129, 138, 167, 198, 200, 201, tulva Austen, 413
227, 242, 257, 265, 28.5, 293, 303, longipennis Fabr., 413
307, 321, 344, 362, 375, 403, 414, maculata (see Hippobosca variegata)
498,502,505,507,514,579,603 martinaglia Bedford, 413
Herpetomonas davidi Lafont, 7 rufipes v. Olfers, 413
pyrrhocoris Zotta et Galli-Valerio, 29 struthionis O. E. Janeson, 413
Herrick, G. W., 236 variegata von Miihlfeld, 413
Hertig, M., 155, 156, 157, 158, 160 Hippoboscidae, 26,32, 109,414
Hesperocimex coloradensis List, 100 characteristics, 4 H)
Hesperoctenes eumops Ferris and Hippocrates, 215
Usinger, 109 Hirschfelder, A. D., 124
hermsi Ferris and Usinger, 109 Hirst, S., 537, 539, 541
INDEX 625
Hirudinea, 31 Houseflies (Continued)
Hirudo medicinalis Linn., 31 lesser housefly, 324
Hisette, J., 152 life history, 319
Hobmaier, A., 558 influence of temperature, 319
Hobmaier, M., 558 preferred breeding places, 320
Hodes, H. L., 245 longevity, 324
Hodge, C. F., 321 metamorphosis, 50
Hoeppli, R., 347, 378 mouth parts, 60, 61, 70
Hoffman, W. A., 161, 196 range of flight, 323
Hollis, M. D., 263 Howard, L. 0., 2, 3, 102, 104, 179, 193,
Holocopops kerteszi Kieff., 161 256, 257, 291, 307, 317, 334, 587
Holsendorf, B. E., 452 Howe, L., 334
H omalomyia ( see Fannia) Howell, D. K, 498, 499
Homoptera, 75 Howitt, B., 242, 546
Honorato, A., 383 Howlett, F. M., 323
Hooke, R., 179 Hu, S. M. K., 247
Hooker, W. E., 464 Hudson, K H., 44
Hookworms, 27 (see Ancylostoma duo- Hudson, N. P., 13, 237
denale) Huebner, R. J., 496, 545
Hopkins, J. P., 282 Huff, C. G., 44, 219, 223, 414
Hoplopsyllus anomalus Baker, 431, 435 Hughes, L. E., 539
Horn flies, 366 Huie, D., 331, 333
characteristics, 367 Hull, J. B., 161
control, 368 Hunter, G. W., 3rd, 224
damage, 368 Hunter, W. D., 18, 464, 476, 488
life history, 367 Hurlbut, H. S., 265
Hornets, 568 Hyalomma Koch, 479
Horntails, 56 Hyalomma aegyptium (Linn.), 479
Horseflies, 6, 35, 146, 242, 297 lusitanicum (Koch), 498
anthrax (relation to), 37, 302 Hydrocyanic acid gas, 98, 443
bite, 301 Hydrometridae, 110
black, 308 Hylemyia nidicola Aldrich, 388
black-and-white, 308 Hymenolepis carioca (Magal.), 338
breeding history, 299 diminuta (Rudolphi), 336, 447
control, 307 nana (v. Siebold), 336
greenhead, 308 Hymenoptera, 56, 75
life history, 299 larvae, 53
mouth parts, 68 mandibles, 64
surra (relation to), 303 Hypoderma Latreille, 394, 403
Horvath, G., 95 Hypoderma aegagri Brauer, 396
Hoskins, W. M., 130, 328 bovis (DeGeer), 39, 394, 396
Hosoi, T., 245 lineata (de Villers), 394, 399, 402
Hotson, H. H., 389 Hypodermatidae, 389, 394
Houseflies, 1, 316 Hystrichopsyllidae, 438, 442
abundance, 318
agents of contamination, 36, 39 Ichthyol, 538
carriers of intestinal protozoa, 336 Imamura, A., 551
control (see Fly control) Imes, M., 137, 514~ 535
DDT-resistant, 346 Impetigo, 5, 118
dispensers of parasitic worm eggs, 335 Indalone, 154
germ carriers, 329 Infantile paralysis, 364
for cholera, 333 Infection, chain of, 36
for gastroinestinal diseases, 332 mechanism of, 36
for ophthalmia, 334 Infection carriers, mechanical, 39
for poliomyelitis, 334 Insecticides, 18
for tuberculosis, 335 Insects, 31, 46
for yaws, 333 anatomy, external, 36
intermediate hosts for Cestoda, 42 internal, 51
Insects (Continued) Katydids, 54
biting, 572 Keay, G., 550
bloodsucking, 574 Ked (see Tick, sheep)
classification, 54 Keener, G. G., Jr., 290
larvae, 53 Keilin, D., 26, 327
orders, 54, 75 Kellog, V. L., 137, 591
parasitoid, 24 KeIser, R. A., 242, 243
stinging, 564 Kessel, E. L., 418, 420
wings, 46 Kilbourne, F. L., 2, 41, 484, 485
Irbisia solani (Heid.), 574 King, A. F. A., 215
Iris, R. C., 283 King, H. H., 576
Irving, W. G., 593 King, W. V., 224,262,285
Ischnocera, 130 Kinghorn, A., 5
Ischnopsyllidae, 438 Kirshner, A., 158
Ischnuridae, 598 Kishman, K. V., 158
Isoptera, 55, 75 Kissing bug, 102, 573
Itch (see Scabies) Kitaoka, M., 245
Ivy, A. C., 13, 14 Kitasato, S., 424
Ixodes, 466, 472 Kitselman, C. H., 243
Ixodes californicus Banks, 473 Klebsiella paralytica, 476
holocyclus Neumann, 494, 495 Kleine, F. K., 357
howelli Cootey and Kohls, 473 Knipe, F. W., 273
paci/icus Cooley and Kohls, 473, 493, Knipling, E. F., 287, 389, 391, 392, 395
495, 603 Knowles, R., 14, 158
ricinus (Linn.), 466, 470, 473, 497, Knowlton, G. F., 243
498, 603 Kobert, R., 591
Ixodidae, 467, 468, 470, 472, 603 Koch, C. L., 466
Ixodinae, 466 Koch, R., 3, 467
Ixodiphagus caucurtei du Buysson, 492 Kofoid, C. A., 107
Ixodoidea, 464, 466 Kohls, G. M., 464, 473, 480, 490, 493,
classification, 470 501, 513, 551, 553
Izumi, E. A., 546 Komp, W. H. W., 179
Izumi, E. M., 244, 547 Koszalka, M. F., 563, 568
Kriggsman, B. J., 369
Jachowski, L. A., Jr., 191 Krober, 0., 299
Jahnes, W. G., Jr., 224 Kumm, H. W., 168
James, M. T., 372 Kurtpinar, H., 396, 399
I anthinosoma lutzii (Theobald), 403 Kuzell, W. C., 592
I atropa basiacantha, 44
Jaundice, infectious, 237 Laake, E. W., 130, 136, 380, 382, 384,
Jellison, W. L., 438, 459, 464, 496, 545 395, 398
Jenkins, D. W., 192, 548, 549 Lacaillade, C. W., Jr., 243
Jenks, G. E., 594 Lacewings, 55
Jepson, F. P., 332, 333 Lachnosterna arcuata Smith, 88
Jo, K., 245 Laemmert, H. W., 239
Jobling, B., 73, 502 Lafont, A., 7
Johannsen, O. A., 143, 165 Lagoa crispata Pack., 576
Johns, F. M., 217 Lake, G. C., 98
Johnson, E. P., 153 Lal, R. B., 333
Jordan, E. 0., 332 Laminosioptes cysticola (Vizioli), 557
Jordan, Karl, 441 Lamson, G. H., Jr., 90, 129, 139
Jorge, R., 432 Lancisi, 256
Joyeux, C., 95 Larkspur, tincture of, 126
Iulus nemorensis Koch, 606 Larrousse, F., 492, 497
Larvae, phantom, ~66
Kadner, C. G., 414 Lasiocampa quercus (Linn.), 576
Kala-azar, 98, 158 Lasiocampidae, 576
Kaston, B. J., 594 Lasius (Formicidae), 22
INDEX 627
Lathrodectus malmignathus var. tropica Leptocoris trivittatus (Say), 574
van Haaselt ( see Latrodectus Leptomonas davidi Lafont, 7, 29
mactam) Leptopsylla musculi Duges, 443
'-atrodectus. 580 Leptospira icteroides Noguchi, 236
Latrodectus apicalis Butler (see Latro- Lepus bairdi Hayden, 434
dectus mactam) irritam Lucas, 548
concinnus Cambridj!;e, 580 Lethocerus, 574
conglobatus Koch, 580 Lethocerus americanus (Leidy), 95
curacaviensis (M liller ), 580 Leucocytozoon Danilewsky, 153
datatus C. Koch ( see Latrodectus Leucocytozoon amatis Wickware, 6
mactam) Leucophaea simondi Mathis and Leger,
geometricus C. Koch, 580 153
guttatus, 580 smithi Volkmar, 153
hasseltii Thorell, 580 surinamensis (Linn.), 81
indistinctus Cambridge, 580 Leucosticte tephrocotis dawsoni Grinnell,
insularis Dahl (see Latrodectus mac- 473
tam) Leucotabanus Lutz, 312
lugubris Motsch, 580 Lewis, D. G., 198
mactam (Fabr.), 28, 33, 580 Lewis, D. J., 354
per/idus Walck ( see Latrodectus Lewis, J. L., Jr., 158
mactam) Libellula quadrimaculata Linn., 42
tredecimguttatus (Rossi), 580 Lice (see Louse)
Liiuger, P., 7 Lillie, R. D., 306
Laurent, A., 493 Lima, S., 237
Laveran, A., 2, 355 Limacodidae, 576
Laveran, C. L. A., 215 Limberneck, 388
Law, J., 531 Limnophora, 403
Lazear, J. W., 3, 233 Limothrips cerealium Haliday, 575
Leaf insects, 55 Lindquist, A. W., 166,346,448
Leafhoppers, 55, 94 Linguatula Frolich, 557
Le Conte, J. L., 102 Linguatula rhinaria Moniez (see Ling_
Leeches, 27, 31 uatula serrata)
Leeson, H. S., 117 serrata Frolich, 557
Leidy, J., 332 Linguatulidae, 58, 557
Leiognathus bacoti Hirst, 539 Linne, C. von, 466
sylviarum ( Canestrini et Fanzogo), Linognathus pedalis (Osborn), 129
541, 546 piliferus (Burm.), 129
Leiper, R R, 6 stenopsis (Burm.), 129
Leiperia gracilis (Diesing), 559 vituli (Linn.), 114, 128, 130
Leishmania Ross, 7, 158 Linsley, E. G., 91
Leishmania brasiliensis Vianna, 159 Lipeurus baculus Nitzsch; 131, 134
donovani (Laveran et Mesnil), 158 caponis (Linn.), 131, 133
Leishmaniasis, 158 heterographus Nitzsch, 133, 134
American mucocutaneous, 159 numidae Denny, 134
Lemann, J., 224 poly trapezius Nitzsch, 133
Leonard, M. D., 313 squalidus Nitzsch, 133
Lepidoptera, 51, 56, 75, 575, 576 variabilis Nitzsch, 133
larvae, 53 Liponissidae, 538
mouth parts, 74 Liponyssus bacoti ( Hirst) (see Bdel-
Le Prince, J. A., 12, 205, 257, 278 lonyssus bacoti)
Leprosy, 98 setosus Kolenati, 538
Leptidae, 146 sylviarum (Canestrini and Fanzago),
Leptinidae, 91, 92 244, 541
Leptinillus aplodontiae Ferris, 91 Lipoptena cervi (Linn.), 412
validus (Horn), 91 depressa (Say), 412
Leptinus testaceus Muller, 91 ferris Bequaert (see Lipoptena subu-
Lcptoconopinae, 162 lata)
Leptoconops torrens Tow~end, 161 mazamae Rondani, 412
628 MEDICAL ENTO!\lOI,OGY
Lipoptena (Continued) Lucilia (Continued)

subulata Coquillett, 412 sericata ( Meigen) (see Phaenicia
List, G. M., 95 sericata)
Liston, W. G., 4, 426 sylvarum Meigen, 328
Lithosia caniola Hbn., 576 Lumbricus terrestris Linn., 31
griseola Hbn., 576 Lumsden, L. L., 349
Litosomoides carinii (Trav.), 539 Lundy, H. W., 546
Livingston, S. K, 406 Lycosa tarantula (Linn.), 578
Livingstone, David, 352, 467, 503 Lycosidae, 578
Lizard, San Diegan alligator, 594 Lygaeidae, III
Loa loa (Cobbold), 6, 249, 306 Lymantriidae, 576
Locusts, 54 Lynchia americana (Leach), 414
Loiasis, 306 fusca (Macquart), 414
Long, J. D., 458 hirsuta Ferris, 414
Longfellow, R. C., 83 lividicolor Bigot ( see Pseudolynchia
Lophortyx californica californica Shaw, canariensis)
414 maura Bigot ( see Psuedolynchia
Lounsbury, C. P., 496, 497 canariensis)
Louse, 5, 35,39,55,113 Lyperosia exigua (de Meij.), 369
bird, 55 irritans Linn., 366
biting, 55, 130 Lytta vesicatoria (Linn.), 89, 564, 577
classification, 130
control, 139 Macaca, 114
injury, 131 Macacus, 114
life history, 131 McCalla, L. P., 488
cattle, 128, 132 MacCallum, W. C., 3, 14, 181, 215
crab, 115, 116, 127 McCaw, W. D., 233
dog, 5, 41 McClure, H. E., 244, 546
domesticated mammals, 136 McCoy, G. W., 13, 422, 432, 434
hen, 28, 29, 132 McDowell, M. M., 495
hog, 128 McIntosh, A., 478
human body, 116, 118, 126 Mackie, D. B., 459
human head, 114 Mackie, F. P., 5, 119
Macracanthorhynchlls hi fll din a c eus
mouse, 306
poultry, 132, 134 (Pallas), 43, 88
Macrodactylus subspinosus (Fab.), 90
pubiC, 115
Macronyssidae, 538
rabbit, 30
Macronyssus Kolenati, 538
sucking, 55
Macropsyllidae, 438
characteristics, 113 Macrothylacia rubi (Linn.), 576
classification, 113 Madden, A. H., 448
control, 129 Madsen, D. E., 243
mouth parts, 68 Maggotts, bird, 388
of mammals, 127 Congo foot, 385
teniasis (relation to), 140 fly, 53
typhus transmission, 121 foot, 388
Louse flies, 109, 410 head, of deer, 402
birds, 413 head, of horse, 402
deer, 412 head, of sheep, 38, 39, 400, 401
mouth parts, 72 identification, 372
Louisicides, 124 surgical, 405
Low, G., 2, 3, 216 wool, 386
Lowe, H., 162 Malaraeus telchinum (Roth.), 435
Lucania para, 292 Malaria, 1, 2, 3, 14, 213, 466
Lucanidae, 64 amphibian, 232
Lucilia caesar (Linn.), 329, 331, 388,
c annopheline vectors, 227
406 avian, 231
iIlfistris (Meigen), 329 cause, 36
INDEX 629
'..lalaria (Continued) Mastigophora, 355
Ceylon epidemic, 14 Mastigoproctus giganteus (Lucas), 601
estivo-autumnal, 217 ' Mastitis, bovine, 168, 336
falciparum, 23 Mastomys coucha (A. Smith), 433
history, 215 Matheson, R, 194, 504
human, 213 Matthysee, J. G., 129, 130, 132
human vectors of, 227 Maurer's dots, 218
losses due to, 19 Maver, M. B., 489, 490
plasmodia, 217 Maxcy, K. F., 551
quartan, 214, 218 Maxwell, J. A., 444
saurian, 232 Mayflies, 39, 42, 55
simian, 232 Mayne, M. B. (see Mitzmain, M. B.)
surveys, 230 Mayo, N. S., 528, 529
synonyms for, 213 Mazzotti, L., 152
Maldonado, A., 44 Meador, C. N., 504
Mallis, A., 155, 571 Meal worms, 43, 88, 89
Mallophaga, 26, 28, 29, 55, 75, U3, 130 Mease, J., 483
Mangabeira, 0., 160 Mecoptera, 56, 75
Mange, 56 Megaloptera, 55, 75
bovine, 531 Megalopyge opercularis S. and A., 575
canine, 531 Megalopygidae, 576
equine, 529, 530 Megarhinini, 176, 181, 184,209
follicular, 537 characteristics, 184
foot, 534 Megarhinus Robineau-Desvoidy, 209
notoedric, 531 Megarhinus inornatus Walker, 176, 185
psoroptic, 536 mtilus Coquillett, 185
swine, 528 septentrionalis Dyar and Knab, 185
tail, 533 Melander, 570
Manson, P., 2, 3, 216, 245, 246 Melania libertina, 25
Manson, P. T., 216 Melanolestes picipes (Herrich-Schaeffer),
Mansonella ozzardi (Manson), 163, 249 101, 108
Mansonia Blanchard, 194,209,246 Meleney, H. E., 176,376
Mansonia africana (Theobald), 240 Melilotus altissima, 225
annulifera Theobald, 248 MeIIanby, K., 440, 527
perturbans (Walker), 181, 194, 249 Melnikoff, R., 5
Mansonioides Theobald, 209 Meloidae, 91, 576
Mantodea, 55, 75 Melolontha melolontha (Linn.), 88
Manure, in relation to houseflies, 339 Melophagus ovinus Linn., 73, 410, 412
chemical treatment, 341 Membracidae, 94, 574
close packing, 340 Menopon biseriatum Piaget, 29
composting pits, 340 fulcomaculatum Denny, 134
disposal, 339 gallinae Linn., 28, 29, 130, 132
on lawns, 347 numidae Gieb., 134
wastage, 339 pallidum Nitzsch, 28, 29, 130, 132
Manwell, R. D., 215 phaeostomum Nitzsch, 134
Marchand, W., 300 stramineum Nitzsch, 133
Marchoux, E., 5, 467, 517 titan Piaget, 29
Marett, P. J., 160 Menoponidae, 130
Margaropus, 478 Mercurialis, 1
Margaropus winthemi Rorsch, 478 Merrill, M. R., 243
Marmoto /Wviventer f/aviventer (Aud. Merriman, G., 823
and Bach.), 434 Mesembrinella, 327
Martin, C. J., 430 Mesnil, F., 355
Martin, E., 3, 16, 198, 225, 568, 573 Mesoveliidae, 110
Martin, E. D., 557 Mestor megistus (Burm.), 6, 101
Martin, R., 7 Metamorphosis, 49
Martin, M., 131 Metcalf, R L., 217
Mason, C. J., 83, 330 Metcalf, Z. P., 35
Methyl bromide, 124, 454, 458 Moe, L. H., 499
Metopiidae, 146, 816, 827 Mohler, J. R, 484
Meyer, K. F., 16, 242, 483, 435, 505 Mole crickets, 54
Michener, C. D., 549, 550 Moniezi expansa (Rudolphi), 42
Michener, M. H., 550 M onopsyllus vision (Baker), 434
Mickel, C. E., 571 Montfils, A. J., 2
Microfilaria diurna Manson, 806 Moore, J. A., 224
Microlynchia pusilla (Speiser), 414 Moore, J. J., 489
Microtus montebelli (Sado), 551, 553 Moore, W., 118, 124
Midges, 146, 164 Mooser, H., 437
biting, 146, 160 Morbus errorum, 118
Dixa, 145, 165, 175 Moreno, A., 595
owl, 145 Morgan, T. H., 12, 13
punkies, 161 Morishita, K., 107
sand flies, 161 Morphidae, 576
Miller, A., 293, 546 Morpho hercules DaIm., 576
Millipedes, 81, 607 Mosquito abatement, 256
Milne, A. D., 5, 467, 503 aircraft in, 285
Milzer, A., 496 drainage, 265
Miranda aurantia (Lucas), 594 ditch maintenance, 267
Miridae, 1l0, 574 ditches, 266
Missiroli, A., 3, 198, 225 salt-marsh drainage, 268
Mitamura, T., 245, 551 education of public, 259
Mites, 56 essentials, 261
air-sac, 557 history, 256
auricular, 536 inspection for breeding places, 276
characteristics, 525 larvicides, 278
chigger (see Chigger mites) natural enemies, 290
depluming, 533 organization for, 258
dermanyssid, 542 personnel, 258
encephalitis (relation to), 546 repellents, 287
follicle, 39, 537 surveys for, 260
itch, 26, 39, 526 water management in, 261
characteristics, 526 controlled reflooding, 264
life history, 527 flushing, 264
liponyssid, 538 impounded water, 263
mange (see Mites, itch) irrigation, 262
meal, 556 in rice fields, 272
northern fowl, 541 salinification, 265
oribatid, 42 water-level management, 264
poultry, 33, 244, 525 Mosquito hawks, 55
quill, 557 Mosquitoes, 1, 35, 39, 56, 145, 175
red chicken, 542 anatomy, 184
control, 543 biting habits (Anophelini), 205
habits, 543 breeding habits (Anophelini), 201
prevention, 544 dengue fever transmission, 240
sarcoptic, 526 egg characters, 198
scab, 39, 534 filariasis (relation to), 248
life history, 535 flight habits, 181
sheep, 38 food habits, 181
treatment, 536 importance, 175
scaly leg, 532 larvae, 58
tropical fowl, 541 life history, 179
tropical rat, 539 Anopheline, 203
Mitzmain, M. B., 6, 182, 205, 226, 303, malaria transmission, 36
304,306,309,362,420,421 mating, 197
Miyagawa. Y., 551 mouth parts, 68
Miyajima, M., 551, 553 salt-marsh, 187
INDEX 631
Mosquitoes (Continued) Myiasis (Continued)
snow, 190 ophthalmomyiasis, 404
surveys, 260 semi-obligate, 380
terminalia, 176 urinary, 378
tree-hole, 18, 189 MYL insecticide powder, 124
yellow fever, 41, 192 Mylabris cichorii (Linn.), 89
Most, H., 249 nubica de Marseul, 578
Moths, 53, 56 obtectus (Say), 554
mouth parts, 74 quadrimaculatus (Fabr.), 554
Mouth parts, 60 Myobia Tatti Skidmore, 531
classification, 60 Myriapoda. 606
importance, 60 Myrientomata berlese, 54
morphology, 62 Myrmeleon, 318
types, 61 Myrmicinae, 570
anopluran, 61, 68, 75 MyxomatOSiS, rabbit, 250
dipteron, 61, 68, 75 Myzomyia Blanchard, 196
hemipteron, 61, 65, 75 Myzorhynchus, Blanchard, 196
hymenopteron, 62, 74, 75
lepidopteron, 62, 74, 75 Nabarro, D., 5, 355
orthopteron, 61, 62, 75 Nabidae, 110, 111
siphonapteron, 62, 73, 75 Nagahata, K., 245
Mud daubers, 570, 593 Nagana, 2, 358
Muir, J. T., 376 Nagaya, M., 551
Muller, P., 7 Nairobi eye, 576
Multiceps multiceps (Leske), 401 Najera, A. D., 147
Mumford, E. P., 378 Napier, L. E., 158
Mumu (see Filariasis, inflammatory) Nash, T. A. M., 354
Muridae, 432 Naturalistic control, 17
Murinae, 433 Naucoridae, 109
Murray, A., 114, 467 Necator americanus (Stiles), 336
Murray, W. D., 247 Necrobia rufipes Fabr., 90
Murrina, 336 Necrobionts, 39, 372
Mus musculus (Linn.), 438, 449, 545 Necrophorus, 87
Musca autumnalis DeGeer, 326 Needham, J. G., 48, 49
corvina ( see Musca autumnalis) Neff, J. A., 288
domestica Linn., 42, 61, 168, 316, 326, Nelson, Y., 82
329, 331, 333, 334, 336, 388, 378, Nemathelminthes, 27
496 Nematocera, 144, 145
nebulo Wiedemann, 324 Nematoda, 48, 83, 87
sorbens Wiedemann, 324 Neochrysops, 311
vestustissima Walker, 324 Neocid, 125
vicina Macquart, 324 Neomyzomyia Theobald, 196
Muscidae, 146, 147, 316, 352, 372 Neopsylla inopina Roth., 434
Muscina stabulans (Fallen), 326 setosa Wagner, 433
Muscoidea, 146, 316 Neotoma, 103,434,435
Museum pests, 87 Neotoma c. cinerea (Ord), 434
Mutillidae, 571 Nepidae, 109
Mutualism, 22 Neumann, L. G., 466
Myers, J. G., 574 Neuroptera, 55, 75
Myiasis, 35, 39, 372 Newstead, R., 352
accidental, 372 Newton, M. V. B., 409
cutaneous, 383 Nicoll, W., 335
dermal, creeping, 402 Nicolle, C. N., 6, 119, 121
dermal, traumatic, 379 Nicotine sulfate (Black Leaf 40),136
epidemic, 383 Nits, 115
gastric, 376 Noctuidae, 576
intestinal, 374, 876 Noguchi, H., 157
obligate, 389 Nosopsyllus Jordan, 441
Nosopsyllus fasciatus (Bosc), 355, 420, Oriental sore, 159
421, 430, 431, 437, 441, 443, 447 Ornithobius bucephalus Pia get, 134
Notoedres minor var. cati (Hering), 531 Ornithoctona erythrocephala ( Leach) ,
minor var. cuniculi Gerlach, 531 414
muris (Meigen), 531 Ornithodoros, 501
Notonectidae, 109, 574 Ornithodoros asperus Warburton, 509
Nott, J. C., 1 coriaceus Koch, 470, 502, 603, 606
Nuttall, G. H. F., 4, 22, 23, 26, 86, 97, erraticus Lucas, 503, 509
11~ 117, 11~ 121, 18~ 303, 332, hermsi Wheeler, 468, 470, 505
333, 466, 470, 472, 494, 503, 517, megnini Duges, 470
603 moubata (Murray), 5, 107, 467, 502,
N uttaWa equi (La veran ), 497 509, 603
Nycteribiidae, 410, 414 normandi Larrousse, 503
Nygamia phaeorrhyea Don., 564, 575, papillipes Birula, 503
576 parkeri Cooley, 503
Nymphalidae, 576 savignyi (Audouin), 107,509
Nysius euphorbiae Horvath, 7, 29 talaie (Guerin-Meneville), 503, 505,
509, 603
Obermeier, O. H. F., 119 tartakovskyi Olenev, 509
Octocentor Colley, 472 tholozani (Laboulbene et Megnin),
Odocoileus hemi;nus columbianus (Rich- 503, 509
ardson), 402, 499 turicata (Duges), 107,509,603
hemionus hemionus (Rafinesque), 402, venezuelensis Brumpt, 503, 509
499 verricosus Olenev, Sassuchin, and
virginianus, 402 Fenik, 509
Odonata, 42, 55, 75 O'Roke, E. C., 153, 414
Oeciacus hirundinus (J enyns ), 95 Oropsy lla idahoensis (Baker), 434
vicarius Horv., 95, 100 silantiewi (Wagner), 432
Oedemagena tarand; (Linn.), 400 Oroya fever, 156
Oedemeridae, 90, 578 Orthopodomyia Theobald, 194, 209
Oestridae, 146, 147, 389, 400 Orthopodomyia signifera ( Coquillett),
Oestrius ovis Linn., .'38, 400, 402, 405 194
Ogata, M., 425 Orthoporus, 607
Orthoptera, 54, 75
Oil, bamber, 287
Orthopterygium huancui, 44
citronella, 287
Orthorrhapha, 144, 299
drips, 281
Oryzoephilus surinamensis (Linn.), 90
linseed, raw, 139
Osborn, H., 129, 137, 400
petroleum, 278
Oscinidae, 147
Okumura, T., 551, 553
Oscinis collusor Townsend, 167
Olfersia maura Bigot (see Pseudolynchia Osten Sacken, C. R., 299
canariensis) Otitis, parasitic (see Otoacariasis)
Onchocerca caecutiens Brumpt (see Otoacariasis, 536
Onchocerca volvulus) Otobius, 501, 513
gibsoni Cleland and Johnston, 153 Otobius lagophilus Cooley and Kohls, 513
gutterosa Neumann, 1.53 megnini (Duges), 38, 513
volvulus (Leuckart), 6, 43, 152, 249 Otodectes cynotis (Hering), 536
Onchocerciasis, 6, 151 Oudemans, A. C., 525
bovine, 152
Onychophora (Protracheata), 31 Packchanian, A., 107
Ophthalmia, 334 Pacylomerus audouini Lucas, 594
Ophthalmomyiasis, 404 Paederus cerbripunctatus, Epp., 578
Opifex Hutton, 176 ragasa, 110
Opiliones, 58 Pahvant Valley plague (see Tularemia)
Opisocrostis tuberculatus (Baker), 434 Paine, J. H., 139
Orchopeas sexdentatus (Baker), 435 Pajaroello (see Ornithodoros coriaceus)
Orenstein, A. J., 20 Palacios, A. M., 147
Organ of Berlese, 95, 99 Panchlora cubensis Sauss., 81
INDEX 633
Panchloridae, 81 Pediculus humanus Linn., 5, 6, 26, 114,
Pangonia Rondani, 298 116, 121
Pangoniinae, 298, 311 humanis capitis DeGeer, 26, 114
Pamtrongylus geniculatus (Latr.), 106 humanis corporis DeGeer, 26, 116, 121
megistus (Burm.), 6 m;obergi Ferris, 114
Pantopoda, 58 schaffi Fahrenholz, 114
Pappataci fever, 5, 41 (see Sand fly vestimenti Nitzsch, 114, 116
fever) Pedipalpida, 58, 600
Paragonimus westermani (Kerbert), 25, Pelecorhynchidae, 311
42 Pellagra, 151
Paraplasma flavigenum Seidelin, 236 Pellitteri, O. J., 545
Paraponera clavata (Fabr.), 570 Pelopaeus californicus Sauss., 570
Parasimulium furcatum Malloch, 150 Pence, R. J., 155
Parasites, 22 Penna, H., 238, 239
classes, 24 Pentastomiasis, 557
Parasitism, 22 Pentastomida, 58, 557
effects, 25 Pentatomidae, 110
forms, 24 Peripatus, 31
origin, 28 Periplaneta americana (Linn.), 80, 81,
parasitoid, 24 83
social, 24 australasiae (Fabr.), 80
Paris green, 273, 278, 279 Perlids, 55
Parish, H. E., 380, 384 Phaenicia sericata (Meigen), 328, 334,
Parker, G. H., 375 374,387,388,406
Parker, R. R., 324, 434, 475, 486, 488, Phalangida (Opilones), 58
489,490,491,492,493,495 Phanurus emersoni (See Telenomus
Parman, D. C., 308, 445 emersoni)
Passer domesticus, 389, 540 Phasmida, 55, 75
Passmore, L., 594 Phidippus (Attidae), 595
Pasteur, L., 2 Philaematomyia Austen, 369
Pasteurella pestis (Lehman and N eu- Philaematomyia imignis Austen, 369
mann),35,40,424,429 Philip, C. B., 250, 302, 311, 475, 496,
tularemis (McCoy and Chapin), 6, 539,551,552,553
41, 98, 305, 493 Philopteridae, 131
Patino-Camargo, L., 156 Phlebotominae, 154
Patton, W. S., 98, 151, 158, 380 Phlebotomus anthophorus Addis, 160
Paul, J. R., 334 argentipes Annandale and Brunetti,
Pauropoda, 31 158
Pawlowsky, E. N., 568, 595 chinemis Patton and Hindle, 158
Peacock, A. D., 118 diabolicu8 Hall, 159
Pearse, A. S., 15, 435 intermedius Lutz and Neiva, 159
Peat, A. A., 168 limai Fonseca, 159
Peck, J. L., 245 longipalpus Lutz and Neiva, 159
Pedicinus Gervais, 114 minutus Rond, 158
Pedicinus albidus (Rudow), 114 noguchii Shannon, 157
ancoratus Ferris, 114 papatasii Scopoli, 5,41, 157, 158, 159
eurygaster (Burm.), 114 perniciosus N ewstead, 158
hamadryas Mjoberg, 114 pessoai Coutinho, 159
longiceps Piaget, 114 sergenti Parrot, 158, 159
obtusus (Rudow), 114 stewarti Mangabeira and Galindo, 160
pafas (Fahrenholtz), 114 texanus Dampf, 160
pictus Ferris, 114 verrucarum Townsend, 156, 157
Pediculidae, 114 vexator Coquillett, 159
Pediculoides ventricosus (Newport), 525, Phormia metallica Townsend, 388
554 regina (Meigen), 329, 334, 379, 387,
Pediculosis, 118 406
pubic, 116 Phthiriasis, 115
Pediculus Linn., 114 Phthirus Nuttall, 114
Phthirus inguinalis Redi, 115 Plasmodium (Continued)
pubis (Linn.), 114, 115 DOOle Stevens, 219
Phyllodromiidae, 81 praecox Blanchard, 217
Phymatidae, 110 relictum (Grassi and Feletti), 231
Phytoparasites, 24 vivax (Grassi and Feletti), 14,27,218,
Pierce, W. D., 117 221, 226
Pieris rapae (Linn.), 74 Plath, O. E., 388, 389
Pinkeye, 39, 167 Platyhelminthes, 29
Pinworms, 39 Platynus maculicolis (Dej.), 90
Piper, S. E., 456 Platypsyllidae, 91, 92
Piophile casei (Linn.), 376 Platypsyllus castoris Ritsema, 91
Piophilidae, 376 Plecoptera, 42, 55, 75
Piratinae, 108 Plica palonica, 115
Piroplasmosis, canine, 497 Pneumonyssus Haan and Grijns, 546
equine, 497 Pneumonyssus simicola Banks, 546
Piroplasma bigeminum ( see Babesia Pogonomyrmex barbatus (F. Smith), 570
bigeminal californicus (Buckly), 570
Pithecus, 114 Poliomyelitis, 334, 364
Pithecus rhesus (Desmarest), 121, 239 Polistes bellicosus Cresson, 570
Pityriasis, 5 Pollenia rudis (Fabr.), 327
Plague, 1, 423 Pollitzer, R., 423
bubonic, 35, 425 Polyctenidae, 109, III
Commission, Indian, 426 Polyp lax serratus (Burro.), 306
in field rodents, 431 spinulosus (Burm.), 355
Heas as vectors, 425 Pomerantz, C., 545
historical, 12, 423 Ponerinae, 570
in man, 424 Porchinsky, J., 307
pneumonic, 425 Porocephalidae, 557
septicemic, 425 Porocephalus Humboldt, 557
sylvatic, 4, 454 Porocephalus crocodili (Wheeler), 558
transmission by Heas, 429 Porter, G. S., 505, 511
Plant lice, 94 Porthesia similis Fuessly, 576
Plasmodia, 217 Potheir, O. L., 224
Plasmodium Marchiafava and Celli, 217, Praying mantids, 50
219 Primicimex cavernis Barber, 99
asexual cycle, 219 Prince, L. H., 406
effect of temperature on, 226 Privies, 348
life cycle, 219 Pmsimulium fulvum Coquillett, 150
sexual cycle, 221 hirtipes (Fries), 150
Plasmodium agamae Wenyon, 232 Prosthogonimus pellucidus (v. Linstow),
brasilianum Gonder and Berenberg- 42
Gossler, 232 Protocalliphora azurea (Fallen), 388
bufonis Fantham, Porter, and Richard- hesperia S. and D., 389
son, 232 hiruda S. and D., 389
catesbianca Fantham, Porter, and splendida Macquart, 389
Richardson, 232 Protomonadida, 355
cathemerium Hartman, 231 Protoroa, 29
cynomolgi Mayer, 232 intestinal, 336
diploglossi Aragao and Neiva, 232 Protura, 54, 75
elongatum Huff, 219, 231 Proust, A., 86
falciparum (Welch), 215, 217 Prurigo, 5
/loridense Thompson and Huff, 232 Pseudogaurax signata ( Loew ), 594
gallinaceum Brumpt, 219, 231 Pseudolynchia brunnea (Latreille), 414
giganteum Theiler, 232 canariensis (Macquart), 73, 413
inui Halberstadler and Prowazek, 232 Pseudoscorpionida (Chelonethida), 58
knowlesi Sinton and Mulligan, 14, 232 Psilopa petrolei Coquillett, 143
kochi (Laveran), 232 Psocidae, 28
malarlae (Laveran), 2, 25, 218, 226 Psocids, 55
INDEX 635
Psocoptera, 28 Rattus (Continued)
Psorophora Robineau-Desvoidy, 194, 209 rattus norvegicus (Erxleben), 83, 438,
Psorophora ciliata (Fabr.), 175 443, 449, 539
celumbiae (Dyar and Knab), 194 rattus rattus (Linn.), 83, 438, 442, 449
confinnis (L.-A.), 272 Rau, P., 82
discolor (Coquillett), 272 Readio, P. A., 101
ferox Humboldt, 249 Rebrassier, R. E., 557
lutzii (Theobald), 403 "Red spiders," 556
Psoroptes Gervais, 534 Redtenbacher, J., 49
Psoroptes communis var. bovis (Ger- Reduviidae, 6, 24, 28, 37, 100, 1l0, 573
lach), 536 key to Reduviidae of medical im-
communis var. equi Gerlach, 536 portance, 108
communis var. ovis (Hering), 33, 39, Reduviinae, 100, 108
525, 534 Reduvius personatus (Linn.), 102, 108,
Psychoda alternata Say, 154 573
pacifica Kincaid, 154 Reed, A. C., 214
Psychodidae, 145, 154 Reed, W., 193, 233
Psychodinae, 154 Reeves, W. C., 182, 206, 243, 244, 293,
Pterostichus, 89 546
Pterygota, 48 Regan, W. M., 364
Ptinus, 86 Reid, W. M., 89, 338
Pulex dugessi Baker, 439 Relapsing fever, 5, 6, 59, 98
irritans Linn., 29, 140, 420, 421, 429, louse-borne, Asiatic, 119
439, 443, 563 epidemic, 119
Pulicidae, 438, 439 European, 119, 504
Punkies (see Midges) spirochetes, 509
Pupipara, 410 tick-borne, African, 5, 502, 503, 509
Puri, I., 180 CalifOrnian, 16, 41, 44, 506, 511
Pycnogonida (Pantopoda), 58 Central American, 503, 509
Pyrethrum, 163, 279, 414, 593 endemic, 504, 511
Pyrosoma bigeminum T. Smith, 486 North American, 511
South American, 503, 509
Q fever, 495 Texan (see North American)
Quayle, H. J., 257 United States, 504
Quercus agrifolia, 189 vectors, 509
lobata, 189 Reservoir animals, 43
Reynolds, F. H. K., 241, 243
Radcliffe, L., 292 Rhagionidae, 146,313
Radford, C. D., 525 Rhagovelia, 110
Raimbert, A., 2 Rhiginia, 110
Raillietina cesticillus (Molin), 89, 338 Rhinocricus, 607
Ransom, B. H., 83, 338 Rhinoestrus purpureus (Brauer), 402
Rao, B. A., 273 Rhipicentor, 478
Raphidiodea, 56, 75 Rhipicentor bicornis Nuttall and War-
Rasahus biguttatus (Say), 104, 108, 573 burton, 478
thoracicus Stal, 104, 108 Rhipicephalus, 478
Rats, 449 Rhipicephalus appendiculatus Neumann,
biology of, 450 . 496, 497
as carriers of fleas, 442 bursa Canestrini et Fanzago, 498
control, 451 capensis Koch, 496, 497
pOisons, 452 evertsi Neumann, 469, 496, 497
Rattus concolor browni, 553 sanguineus (Latreiile), 107, 478, 497,
/lavipectus yunanensis, 553 498
hawaiiensis Stone, 441, 449, 450 simus Koch, 497, 498
rattus alexandrinus (Geoffroy-Saint Rhodnius prolixus Stal, 105, 106, 108,
Hilaire and Audouin), 438, 442, 250
449 Rhopalopsyllus cavicola (Wayenberg),
rattus frugivorus Raflnesque, 449 433
Rhynchophthirina, 114 Russell, P. F., 176, 203, 215, 227, 273
Rhynchotaenia Bnlthes, 209 Rutgers 612, 154, 287
Rice, L., 240, 241 Ryckman, R. E., 607
Richards, C. S., 249
Richards, J. T., 495 Sabin, A. B., 334
Ricketts, H. T., 5, 6, 121, 488, 490, 492 Sabrosky, C. W., 170, 182
Rickettsia akari, 540, 545 Sacculina carcini (Rathke), 25
burneti (see Coxiella burneti) Sachs, H., 591
diaporica Cox, 495 Sadusk, J. F., 551
mooseri Montiero, 122, 437 Saint John, J. H., 241, 503
orientalis Nagayo, 551 Saldidae, 110
prowazeki da Rocha-Lima, 6, 120 Salimbeni, A., 5, 467, 517
prowazeki mooseri, 122 Salmon, D. E., 466, 483
quintana da Rocha-Lima, 122 Salmon flies, 55
rickettsi (Wolbach), 5, 41, 488 Salmonella paratyphi (Kayser), 333
ruminatium (Cowdry), 498 Sambon, L. W., 3, 43, 84, 151, 216, 557
tsutsugamushi (H yashi ), 551 Sanarelli, G., 236
Rickettsialpox, 545 Sand flies, 44, 145, 154
Riley, C. V., 587 control, 160
Roberts, A., 433 life history, 155
Roberts, H. R, 178 species, 159
Roberts, J. 1., 578 Sand fly fever, 157
Robertson, M., 357 Sanders, D. A., 168, 336, 497
Robiquet, 577 Sao Paulo fever (see Spotted fever)
ROcky Mountain spotted fever, 5, 41, 43, Sarcobionts, 39
44, 486 Sarcophaga carnaria (Linn.), 329, 331
control, 492 hacmorrhoidalis (Fall.), 329, 383
infection in nature, 489 Sarcophagidae, 146,316,327,372
mechanism of infection, 490 Sarcopsylla penetrans (see Dermatophilus
reactivation, 491 pf!netrans)
tick transmission of, 488 Sarcoptes Latreille, 526
symptoms, 486 Sarcoptes minor var. felis Gerlach, 531
Rodent control, 448 scabiei (Linn.), 26, 39, 526
field rodents, 454 scabiei Var' auckeniae Raulliet, 532
methods, 451 scabiei var. bovis Robin, 531
rats, 449 scabiei var. canis Gerlach, 531
Rodhain, J., 219 scabiei var. caprae FUrstenberg, 532
Romafia, C., 105, 107, 154 scabiei var. equi Gerlach, 526
Rook, A. F., 155 scabiei var. hominis (Hering), 525
Root, F. M., 24, 161, 336 scabiei var. ovis Megnin, 531
Rose chafer, 90 scabiei var. suis Gerlach, 526
Rosen, L., 293 Sarcoptidae, 526, 534, 536
Rosenau, M. J., 364 Sata, K, 551
Ross, E. S., 178 Saturniidae, 575, 576
Ross, I. C., 494 Savory, T. H., 58, 578
Ross, P. H., 5, 467, 503 Sawdust, in mosquito control, 284
Ross, R., 3, 13, 215, 217, 256 Sawflies, 56
Rotenone, 130, 398 Sawyer, W. A., 365
Roubaud, E., 225, 336, 385 Scabies, bovine, 536
Roundworm of man (see Ascaris lumbri- chorioptic, 533
coides) human, 56, 526
Rovelli, G., 338 ovlne, 534
Rowe, J. A., 243, 345 Scale insects, 55
Roy, D. N., 167 Scarabaeidae, 43, 88, 89, 91
Rozeboom, L. E., 176, 201, 204, 227 Scatophagidae, 147
Rucker, W. C., 421 Sceliphron caementarius Drury, 593
Rumreich, A., 437, 490 Schamberg, J. F., 555
Russell, J. C., 182, 196 Scharff, D. K, 205
INDEX
Schizotrypanum cruzi Chagas, 6, 104 Sialids, 55
Schliffner's dots, 218, 219 Sibine stimulea Clem., 576
Schultz, C., 191 Sigmodon hispidus, 539
Schwardt, H. A., 301, 302 Sign of Romafia, 105
Schwarz, E. A., 167 Silpha, 87
Sciuridae, 433, 454 Silphidae, 87, 92
Sciurus douglasii albolimbatus Allen, 505 Silver, J., 451
hudsonicus richardsoni Bachman, 434 Silverfish, 49, 54
Scolopendra Linn., 607 Silvius Meigen, 298, 31J
Scolopendra heros Gir., 607 Simmons, J. S., 241, 243
polymorpha Wood, 607 Simmons, P., 376
sumichrasti Sauss., 607 Simond, P. L., 4, 426
viridis Say, 607 Simpson, R. W., 155
Scorpionflies, 56 Simuliidae, 6, 146, 147
Scorpionida, 58, 595 characteristics, 147
Scorpionidae, 598, 599 classification, 150
Scorpions, 56, 58, 595 life history, 148
characteristics, 595 species, 150
control, 600 Simulium callidum Dyar and Shannon,:
Durango, 597, 599 152
venom, 563, 596 columbaschenis Fabr., 151
whip (see Whip scorpions) damnosum Theobald, 6, 152, 250
wind (see Solpugida) meridionale Riley, 150
Scott, O. K., 249 metallicum Bellardi, 152
Screwworm flies, 379, 380 neavei Roubaud, 152
control, 384 nigroparvum (Twinn), 153
life history, 384 occidentale Townsend, 150, 153
Screwworms, 39 ochraceum Walker, 152, 250
Scrub typhus (see Tsutsugamushi dis- ornatum Meigen, 148, 153
ease) pictipes Hagen, 150
Scutigera forceps (Rafinesque) (see Cer- venustum Say, 6, 151, 153
matia forceps) vittata Zetterstedt, 150
Seale, A., 291 Sinea diadema (Fabr.), 108
Seaman, E. A., 249 Sinton, J. A., 230 '
Sea-spiders, 58 Siphonaptera, 56, 64, 75, 416, 4!)6
Seddon, H. R., 517 key to families, 438
Seifert, 0., 374 mouth parts, 73
Seki, 0., 245 Siphunculata (Anoplura), U3
Sellards, A. W., 243 Siphunculina funicola de Meyere, 16'1'
Semnopithicus maurus, 114 Siroptera, 83
Sense organs, accidental injury to, 38 Sitotroga cerealella (Oliv.), 554
Serafin, J., Jr., 238, 239 Skidmore, L. V., 153,531
Sericopelma communis Cambridge, 580 Sleeping sickness, 356
Sessinia collaris (Sharp), 90, 578 Commission, 358
decolor Fairm., 90, 578 Gambian, 5, 357
Shannon, R. C., 150, 157, 165, 239 historical, 5, 357
Sharp, N. A. D., 162 Rhodesian, 5, 357
Sharpshooters, 94 transmission, 357
Shaughnessy, H. J., 496 Smart, J., 147, 148, 150
Sheeley, W. J., 384 Smith, C. N., 474, 500, 501
Shelmire, B., 539 Smith, J. B., 188, 257
Shigella dysenteriae (Shiga), 333 Smith, L. M., 162
Shimizu, M., 245 Smith, M. B., 244
Shipley, A. E., 184 Smith, M. G., 546
Shiraki, T., 81 Smith, R. O. A., 158
Shope, R. E., 250 Smith, T., 2, 28, 41, 153, 466, 483, 484,
Shortt, H. E., 158 485, 486
Shrimp, 31 Snails, 25
Snakedoctors, 55 Spotted fever (Continued)
Snakeflies, 56 mechanism of infection, 490
Snodgrass, R. E., 56, 60, 62, 74, 75, 77, transmission by ticks; 488
416, 525 SpringtaiIs, 49, 54
SnowHeas, 54 Squill, 452
Sodium fluoride, 85, 134, 139 Squirrel, ground, 454
Solerwpotes capillatus Enderl., 128 Beechey (see CiteUm b. beecheyi)
Solenopsis xyloni var. maniosa Wheeler, breeding, 455
570 Chickaree (see Sciurus douglasii al-
Solifugae, 58 bolimbatus )
Solpiguda, 58, 602 control, 455
Soothsayers, 55 Douglas ground (see Citellus beecheyi
Soper, F. L., 4, 12, 237, 238, 239, 293 douglasii)
Sopidae marginata Uhler, 574 estivation and hibernation, 455
Southwell, T., 158 ground, 454
Sowbug, 31 habits, 455
Spanish fly, 89, 577 picket-pin (see Citellus beldingi)
Species sanitation, 17 plague transmitted by, 13
Spencer, R. R., 493 Tamarack (see Sciurus douglasii al-
Speotyto cunicularia, 459 bolimbatus )
Spermophilus refuscens, 433 Stable flies (see Stomoxys flies)
Sphaerophthalma occidentalis ( Linn. ) , Stage, H. H., 181, 182
571 Stahnke, H. L., 597, 598
Sphex luctuosa Smith, 570 Stallybrass, C. 0., 43
vio!aceipennis Lepeletier, 570 Staphylinidae, 38, 86, 90, 92
Spicknall, C. G., 158 Staphylococcus aureus Rosenbach, 330
Spiders, 56, 576 Steatoda borealis (Hentz), 594
banana, 594 Stegomyia fasciata (Fabr.) (see Aedes
bite, 587 aegypti)
. black widow ( see Black widow Steinhaus, E. A., 490, 495, 551
spider) Stenotabanus Lutz, 312
burrowing, 594 Stephanostoma haemorrhoidalis (Fall.),
garden, 594 383
narvest, 58 Stephens, J. W. W., 5, 221
Latrodectus, 580 Sternostomum rhirwlethrum Trouessart,
sun, 58 546
tarantula (see Tarantulas) Stevens, G., 505
trap-door, 594 Stevens, I. M., 505, 511
Spirochaeta carteri Manson, 6 Steward, J. S., 152, 153
duttoni Novy and Knapp, 5, 509 Stewart, C. M., 592
gallinarum Blanchard, 5, 467 Stewart, M., 493, 499
hispanica Sadi de Buen, 510 Stewart, M. A., 38, 376, 406, 435, 459
kochi Novy, 510 Stilbometopa impressa (Bigot), 414
marchouxi Nuttall, 517 podopostyla Speiser, 414
neotropicalis Bates and St. John, 510 Stiles, C. W., 349, 486
obermeieri, 119 Stiles, G. W., Jr., 466, 498, 499
persica Dschunkowsky, 510 Sting, bee, 564
recurrentis Lebert, 5, 6, 41, 119, 505 operation, 567
turicatae Brumpt, 509 reaction to, 568
venezuelense Brumpt, 509 treatment, 568
Spirochetosis, fowl, 5, 467, 517 Stokes, A., 4, 13, 237
Spirura Tytipleurites (Deslgch.), 83 Stomoxys calcitrans (Linn.), 61, 305,
Spiruridae, 43, 87 322,338,360,365,573
Splenomegaly, tropical (see Kala-azar) irwrnata Grunberg, 369
Spotted fever, 56, 121, 490 (see also nigra Macquart, 369
Rocky Mountain Spotted fever) omega Newstead, 369
control and prevention, 492 Stomoxys Hies, 60, 61, 360
infection in nature, 489 breeding habits, 362
INDEX 639
Stomoxys flies (Continued) Tabanus atl'atus Fabr., 68, 308
as cattle pest, 364 costalis Wiedemann (see Tabanus vi-
characteristics, 360 carius )
control, 366 dorsovittata Walker, 357
habits, 360 gilatus Townsend, 300
life history, 362 lineola Fabr., 301, 309
longevity, 363 nemoralis Meigen, 307
mouth parts, 69 phaenops Osten Sacken, 302
poliomyelitis (relation to), 364 punctifur Osten Sacken, 242, 807, 30B
Stone, Alan, 176, 227, 299, 301, 310, 311 striatus Fabr., 6, 303, 304, 305, 309
Stoneflies, 42, 55 stygius Say, 308
Stonemyia Brennan, 311 sulcifro1l$ Macquart, 309
Storer, T. I., 388, 451, 454, 455, 459 tomentosus Macquart, 307
Stratiomyidae, 376 vicarius (Walker), 308
Straw itch, 554 Tabardillo, 120, 121
Streblidae, 410, 414 Tachnidae, 146
Strepsiptera, 56, 75 Tadarida macrotis (Gray), 109
Strong, R. P., 15, 152, 157 Taenia hydatigena Pallas, 336
Stubbs, T. H., 224 pisiformis (Bloch), 336
Stunkard, H. W., 23, 24, 42 saginata Goez, 31
Stygeromyia Austen, 369 solium Linn., 29, 336
Stygeromyia maculosa Austen, 369 Taeniasis, fowl, 338
sanguinaria Austen, 369 Taeniothrips inconsequens (Uzel), 575
Sucking lice (see Louse, sucking) Tamiya, T., 551
Sulkin, S. E., 244, 543, 546, 547 Tampi, M. K., 245
Sullivan, E. C., 443 Tapeworms, 2, 5, 25, 41, 336
Supella supellectilium Serv., 80, 81 beef (see Taenia saginata)
Surcouf, J., 311 dog (see Dipylidium caninum)
Surra, 6, 303 fowl, 42, 89, 338
Surveys, mosquito, 260 intermediate hosts for, 42
rat-flea, 443 pork (see Taenia solium)
Swaminath, C. S., 158 sheep, 42
Sweet, W. C., 273 transmission by fleas, 447
Swellengrebel, N. H., 23, 430 Tarantulas, 578
SWingle, L. D., 411 Tardigrada, 58
Swynnerton, C. F. M., 352, 360 Tarsonemidae, 554
Sylvilagus nuttali nuttali (Bachman), 434 Tate, H. D., 480
Symbiosis, 22 Tatem lobengulae De Wint, 433
Symphoromyia atripes Bigot, 313 Taussig, S., 5
hirta Johns, 314 Taylor, R. M., 239
kincaidi Aldrich, 313 TDE, 154, 166, 281, 290
pachyceras Williston, 313 Telenomus emersoni (Girault), 308
Symphoromyia Frauenfeld, 313 Telford, H. S., 136
Symphyla, 31 Telini fly, 89
Synxenoderus comosus List, 99 Temple, J. U., 493
Syringophilus bipectinatus Heller, 557 Ten Broeck, C., 243
columbae Hirst, 557 Tenebrio molitor, Linn., 88, 89
Syrphidae, 146, 378 Tenebrionidae, 43, 84, 88, 89, 91
Syvertan, J. T., 243 Tenjui, S., 245
Termites, 55
Tabanidae, 6, 146, 297 Terrapane carolina (Linn.), 549
breeding history, 299 Tetranychidae, 556
control, 307 Tetranychus bimaculatus Harvey, 556
key to genera, 311 telarius (Linn.), 556
life history, 299 Texas cattle fever, 2, 12, 40, 41, 56, 466,
species, 308 480,482
Tabaninae, 297, 311 Texas fever tick, 480
Tabanus Linne, 298, 313 life history, 481
Texas fly (see Horn fly) Ticks (Continued)
Thallium sulfate, 453 venemous, 603
Thaumetopoea pinivora Tr., 576 wood, 483, 488
Thaumetopoeidae, 576 Tictin, J., 5
Thayer, W. S., 224 Tilden, E. B., 157
Theiler, M., 13,497,498 Tillyard, R. J., 48, 49
Theileria parva (Theiler), 496 Ti"~:JUlidae, 145
Thelyphonidae, 600 Ti;1:zoni, G., 333
Theobaldia Neveu-Lemaire, 209 Tobia petechial fever (see Spotted fever)
Theodor, 0., 159 Todd, F. M., 12
Theridiidae, 580, 594 Todd, J. L., 5, 357, 467, 503
Thomas, L. J., 476 Toepfer, H., 122
Thompson, M. G., 385 Tongue worms, 58, 557
Thrassis acamantis (Roth.), 435 Tanking, H. D., 578
Threadworms, 43 Torti, F., 215
Threlkeld, W. L., 153 Townsend, C. H. T., 155, 156, 151
Thrips, 55, 574 Toxascaris leonina (v. Linstow), S36
mouth parts, 64 Trask, J. D., 334
Thrips imaginis Bagnall, 575 Treehoppers, 55, 94
tubuc\ U",d, 515 Trematoda, 31, 42
Thysanoptera, 55, 75, 574 Treponema pertenue Castellani, 1, 168,
Thysanura, 54, 75 333
Tick control, 499
recurrentis (see Borrelia recurrentis)
on dogs, 500 Triatoma ambigua Neiva, 106
on live stock, 499 barberi USinger, 106
on vegetation, 500 delpontei Romafia y Abalos, 106
dimidiata (Latr.), 106
Tick fever, 461
:/favida (Neiva), 106
Colorado, 492
gerstaeckeri (Still), 106
Ticks, 56, 464
hegneri Mazzotti, 106
American dog, 474
heidemanni Neiva, 106, 107
cattle, 41
jntestaTlS (Klug), 250
Cayenne, 490
longipes Barber, 101, 106
characteristics, 467 megista (Burm.), 6, 105
control (see Tick control)
phyllosoma (Burm.), 106
diseases carried by, 482 protracta (Uhler), 6, 27, 32, 95, 101,
dog, 478, 490 106, 107, 108, 250, 573
ear, 38, 513 protfacta woodi Usinger, 106
fowl, 515 rubida (Uhler), 106
control, 518 rubida uhleri Neiva, 106
damage, 517 rubrofasciata (DeGeer), 101, 106, 107
life history, 515 sordida (StilI), 106
high vector potential, 465 1'riatominae, 100
historical, 466 Trichinella spira lis (Owen) 29
horse, 476 Trichocephalus trichiurus (Linn.), 336
hosts, 465 frichodectes bovis (Linn.), 131
life history, 468 breviceps Rudow, 139
"lone star," 479 canis DeGeer, 5, 42, 138, 140, 447
longevity, 470 climax (see Bovicola caprae)
mouth parts, 468 crassipes Rudow, 138
paralysis, 493 equi (Linn.), 137
pigeon (see Argas reflexus) hermsi Kellogg and Nakayama, 138
Rocky Mountain wood, 474 latus Nitzsch (see Trichodectes canis)
sheep, 410 limbatus Gerv., 138
spotted fever, transmission by, 488 ma/or Piaget, 138
Texas cattle fever (see Texas fever avis Linn., 137
tick) parumpilosu; Pia get (see Trichodectes
tularemia, transmission by, 493 equi)
INDEX 641
Trichodectes (Continued) Typhus fever, 5, 44, 120
penicillatus Piaget, 138 endemic, 122
pilosus Giebel, 137 epidemics, 125
scalaris Nitzsch (see Bovicola bovis) Mexican, 120, 121
sphaerocephalus Nitzsch (see Tricho- murine (flea-borne), 122, 437
dectes avis) transmission by lice, 121
subrostratus Nitzsch, 139 Tyroglyphidae, 556
tibialis Piaget, 139 Tyroglyphus americanus Banks, 556
Trichodectidae, 131 farinae (DeGeer), 556
Trichoptera, 42, 56, 75 longior Gerv., 556
Trinoton luridum Nitzsch, 130, 133 siro (Linn.), 556
Troctes divinatoria (Miill.), 28 Tyzzer, E. E., 187, 243, 575
Trombicula Berlese, 548
Trombicula akamushi (Brumpt), 551, Umbellaria californica, 189
553 Underhill, G. W., 153
autumnalis (Shaw), 548 Unsworth, K., 539, 540
deliensis Walch, 553 Uranotaenia Lynch-Arribalzaga, 209
my otis Ewing, 554 Uroctonus mordax Thorell, 600
thalzahuatl (Murray) (see Eutrom- USinger, R. L., 94, 95, 99, 106, 108, 109,
bicula alfreddugesi) 574
Trornbibiidae, 547 Uta (see Leishmaniasis, American mu-
Trombiculinae, 547 cocutaneous )
Trypanosoma Gruby, 355 Uvarov, B. P., 16
Trypanosoma berberum Edmond and Et.
Sergent, 307 Vaejovidae, 598, 599
brucei Plimm. and Bradf., 27, 355, 357 Vaejovis boreus (Girard), 600
conorhini Donovan, 107 spinigerus Wood, 600
cruzi Chagas, 6, 104, 107 Vagabond's disease, 118
evansi Steel, 303, 305, 355 Valentin, 355
gambiense Dutton, 5, 27,29, 355, 356, Valley black gnat, 161
357 Van Allen, A., 438
hippicum Darling, 60, 336 Van Leeuwenhoek, A., 2
lewisi (Kent), 355 Vanessa io (Linn.), 576
rhodesiense Stephens and Fantham, 15, Vanillism, 556
27,356,357 Vannote, R. L., 279
Trypanosomiasis, 354 Vargas, L., 147, 152, 154
Algerian, 307 Vector control, 17
American, 104
Veeder, M. A., 1,332
Brazilian, 6, 41, 104
Veliidae, 110
Panamanian, 336
Trypanosomidae, 355 Venoms, insect, 563
Tsetse flies, 2, 5, 352 introduction of, 564
characteristics, 352 Verjbitsky, D. T., 4, 426
control, 359 Vermileo, 313
life history, 353 Verruga peruana, 156
sleeping sickness (relation to), 356 Vespa crabro, Linn, 568
Tsutsugamushi disease, 44, 56, 551 germanica Fabr., 568
Tuberculosis, 335 maculata (Linn.) (see Vespula macu-
Tularemia, 6, 41, 98, 305, 454 lata)
tick transmission of" 493 vulgaris Linn., 568
Tunga penetrans (Linn.), 29, 442, 443, Vespidae, 568
445 Vespula diaholica (Sauss.), 568
Tupaia belangeri versurae, 553 maculata (Linn.), 568
Turk, F. A., 546 pennsylvanica (Sauss.), 568
Turner, T. B., 168 Viruola pestosa, 439
Turner, W. E., 464 Vitzthum, H., 525, 538
Tyler, J. R., 157 Vogelsang, W. A., 591
Typhoid fever, 28, 39 Von Geldern, C. E., 571
Walking stick, 38, 55 Wilson, H. G., 346

Wallace, G. 0., 476 Wilson, H. M., 262
Walton, W. R., 144 Wilson, L. B., 4BB
Warble (see Grubs, cattle) Windred, G. L., 371
Warburton, C., 395, 470, 472, 5OS.527, Wing venation, 48
603 Winterbottom's sign, 356
Ward, R., 3S4 Wisseman, C. L., 54S
Warren, G., 216 W ohlfartia magnifica (Schin.), 387,
Wasps, 53, 56, 570 nuba, 406
mutillid,571 vigil (Walker), 383
Watanabe, S., 245 Woke, P. A., 185, 193
Water moths, 56 Wolbach, S. B., 5, 486
Water-bears, 58 Wood, F. D., 107
Water bugs, biting, 574 Wood, H. P., 133, 134, 543
giant, 95 Wood, General Leonard, 234
velvet, III Wood, S. F., 106, 101
Water creepers, 109 Wood, S. W., 104
scorpions, 109 Woodland, J. C., 495
striders, 110 Woodruff, L. C., 82
treaders, no Woods, C. M., 4~~
Waterhouse, A. T., 109 Woodward, T. E., 464
Waterman, J. A., 596 Woodworth, C. W., 257
Watson, M., 256 Woodworth, H. E., 388
Watt, J. Y. C., 374, 378 Workman, W. G., 437
Watts, C. N., 448 Worm lion, 313
Weathersbee, A. A., 282 Worms, African eye, 306
Webb, J. L., 302 filarial, 249
Weber, N. A., 570 meal (see Meal worms)
Webster, F. M., 556 monoxenous, 42
Weevils, 56 parasitic, 336
Wehr, E. E., 391
Weidner, H., 576
'l,lle{l''S dh-ease, 2S{l
Weller, B., 506
Weller, T. B., 224
Wellman, C., 603
Wells, K. W., 389, $91
/
/J thorn-headed, 43, 88
thread (see Threadworms)
Wrigglers, 5<3
Wu, C. Y., 423, 440
Wu, L., 423, 440
Wu, Y. F., 148
Wuchereria bancrofti (Cobbold), 2, 40,
Wells, R. W., 136, $02, 395, § 43, 186, 245, 307
West, L. S., 215 Bora Bora strain, 249
Westwood, J. 0., 109. life cycle, 246
Wharton, G. W., 548, 553 Wuchereria malayi Brug, 248
Wheeler, C. M., 107, 125, 242, 436, Wyeomyia Theobald, 209
459,506,507
Wheeler, W. M., 2;2 Xanthocephalus xanthocephalus ( Bon-
Whip scorpion, 58, 600 aparte), 244
characteristics, 600 Xenodiagnostic, 105
White, R., 225 Xenodusa cava (Lec.), 22
Whitman, L., 239 montana (Csy.), 22
Whitneyomyia Bequaert, 312 Xenopsylla astia (Rothschild), 440
Whittingham, H. E., 41, 155 brasiliensis (Baker), 440, 443
Wilcox, A., 217 cheopis (Rothschild), 421, 426, 429,
Wilder, R. M., 6, 121 431, 436, 437, 439, 443, 447
Williams, C. L., 442, 443 hawaiiensis Jordan, 441
Williams, D. H., 404
WtlIiams, L. L., Jr" 202, 204, 285 Yao, H. Y., 331, 333
Williams, R. W., 5S9, 546, 550 Yates, W. W., 181
Williston, S. W., 410 Yaws, 1,39, 168,333
Wilson, D. B., 12, 237, 293 Yellow fever, 1, 41, 232
INDEX 643
Yellow fever (Continued) Yorke, Wo, 5
Commission, 233 YOUl1g, Co W o, 158
discoveries, recent, 237 Young, Mo Do, 224
epidemics, 233 Yuan, 10 Co, 331, 333
etiology, 236 Yust, H. R., 280
historical, 3, 4, 12, 233 Yutuc, 303, 304
jungle, 238
mo~quito, 1, 192 Zeidler, 00, 7
vectors, 239 Zelus exsanguis (Stal), 108
virus transmission, 237 socius Uhler, 108
Yellow jackets (see Hornets) Zinsser, Ho, 437
Yen, Co, 247 ZoOparasites, 24
Yersin, Ao, 424 Zoraptera, 55, 75

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MEDICAL ENTOMOLOGY

MACMILLAN AND CO., LIMITED

THE MAC)ULLAN COMPANY

BY vVILLIAM B. HERMS_, SC.D.

THE MACMILLAN COMPANY: ~EW YORK

All rights reserved-no part of this book may be

SECOND PRINTING, 1953

niques of chemical control (use of insecticides) and the neglect of knowl-

MUCH of the matter contained in the following pages was prepared

PREFACE to the first edition ix

lUSTORY; CLASSIFICATION; COMMON SPECIES; THE BITE; RELATION TO

LINE CONTROL IN BROMELIADS; SEWER INLETS AND CATCH BASINS;

HORSE BOTFLIES; CATTLE GRUBS; LIFE HISTORY AND HABITS; HEAD

Historical. In the King James version of the Old Testament (Exodus

fever. In 1854 Beauperthuy/ a French physician in the West Indies,

SCOPE AND METHOD

Scope. The preceding brief outline of the historical development of

A ground squirrel was inoculated subcutaneously with a broth culture

Although human malaria was not known to be transmissible to labo-

2. ---, 1943. "Preparation for a career as a medical entomologist,"

Symbiosis and parasitism. Biologists are not agreed as to the definition

a. Protozoa: unicellular animals; e.g., Endamoeba histolytica (Schau-

FIG. 2. Examples of parasitic flat-

FIG. 6. Examples of the

thorax; two pairs of wings on thorax (may be reduced or

FIG. 9. Examples of the Class Insecta. (A) a reduviid (conenose) , Xl;

Texas fever tick; Dermanyssus gallinae (DeGeer) , the poultry

by bites or by skin invasions. Some of these irritations could be appro-

infections are notable for their wild-animal reservoirs: Rocky Mountain

STRUCTURE, DEVELOPMENT, AND

The Insecta (Hexapodaj constitute the largest class in numbers of

A PRIMITIVE WING TILLYARD B TABANUS

change is found in the Apterygota (silverfish and springtails), which

nymph, (3) imago or sexually mature adult. This type of metamorphosis

FIG . 13. Illustrating primitive meta-

2. A more complicated situation exists in the case of the Anopheles

FIG. 16. Showing digestive h'act of cockroach, Order Orthoptera (top);

Insect classification. The medical entomologist must be equipped with

wings , nerve-wiugecl insects; biliug mouth parls; complex metam(Jr -

FIG. 17. Showing external morphology-of a spider. C, coxa; Cit., chelicera:

o Some authors include among the Arachnida the Pycnogonida ( Pantopoda )

Key to the Terrestrial Orders of the Class Arachnidaf

t Adapted after various authors.

INSECT AND ARACHNID MOUTH PARTS

Importance of mouth parts. No doubt all insects possessing mouth

1. Orthopteron type: generalized mouth parts consisting of opposable jaws

MORPHOLOGY OF MOUTH PARTS

~IDE VIEW FRONT VIEW

FIG. 19. Head and mouth parts of a cockroach. Orthopteron (mandibulate)

FIG. 20. Head and mouth parts

cal labium forms a prominent beak-like proboscis which is usually three-

/OORSAL STYLET \ VENTRAL STYLET

RIGHT HALF OF SUCKING TUBE

RIGHT HALF OF SAC TUBE

LABRUM LEFT MANDIBLE LEFT MAX ILLA LABELLA

covering to the concaved hypopharynx, thus giving rise to a food tube.

SIDE VIEW VENTRAL VIEW

~~L ABIAL PALP

ovinus Linn., is nearly as long as the combined haustellum and labellum

FIG. 29. Head and mouth