Report Vegetation

Vegetation of Amchitka
Island, Aleutian Islands,

Alaska
GEOLOGICAL SURVEY PROFESSIONAL PAPER 648
Vegetation of Amchitka
Island, Aleutian Islands,
Alaska
By HANSFORD T. SHACKLETTE and others
L. W. DURRELL HILDUR KROG
JAMES A. ERDMAN HERMAN PERSSON
JOHN R. KEITH H. SKUJA
WILLIAM M. KLEIN WILLIAM A. WEBER
GEOLOGICAL SURVEY PROFESSIONAL PAPER 648
An account of the plant species and plant

communities that occur in an oceanic
island environment
UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1969

UNITED STATES DEPARTMENT OF THE INTERIOR
WALTER J. HICKEL, Secretary
GEOLOGICAL SURVEY
William T. Pecora, Director
Library of Congress catalog-card No. 76-602502
For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 - Price $1 (paper cover)
CONTENTS
Page
Abstract.__________________________________________ 1 Plant species of Amchitka Island.____________________ 34
Introduction, by Hansford T. Shacklette_______________ 1 Annotated list of vascular plants, by Hansford T.
Objectives and limitations of these studies.________ 1 Shacklette, James A. Erdman, and John R. Keith. 34
Botanical studies that have been conducted. _______ 3 Annotated list of bryophytes, by Herman Persson and
Nomenclature, and notes on collections____________ 3 Hansford T. Shacklette-_--_--_--_-----_---__- 40
Acknowledgments. _ _ ____________________________ 3 Annotated list of lichens, by William A. Weber,
Description of the island and its vegetation, by Hansford James A. Erdman, and Hildur Krog.___________ 47
T. Shacklette-__--__-_-__-______-_____-______ 4 Some algae and other protists, by H. Skuja___-___- 50
Geography.____________________________________ 4 Selected references-_________________________ 55
Climate.______________________________________ 4 Soil fungi and algae, by L. W. DurrelL.___________ 55
Physiography and geology.______________________ 5 Observations on the flora, by William M. Klein____ 57
Soils.__-__-___-__-_-___--________-_______--__- 5 References cited-____-_____________-_---_-____--__-- 58
Plant habitats and plant communities-____________ 9 Index._________-___-___-_____--_---__-_-_________- 61
ILLUSTRATIONS
[Plates follow index]

PLATE 1. Drawings of alga species.
2. Drawings of algae and other protists. Page
FIGURE 1. Map of Amchitka Island, Alaska, showing locations of vegetation study sites.-______._-__.__._..___.-- ._-- 2
2-35. Photographs:
2. Section of a peat deposit with an upper and two lower ash layers that are marked by a thick growth
of the moss Pogonatum urnigerum,- _______________-___--_--_-_---___---_---_____-_------ 8
3. Effects of soil fertility on the growth of the grass Calamagrostis nutkaensis. _____________________ 8
4. Zonation of Honckenya and Senecio at Makarius Bay._________________________________________ 12
5. Sand dune with cover of Elymus community at the seashore, Rifle Range Point-_________________ 13
6. Sea-cliff habitat on the Pacific Ocean coast at Makarius Bay.__-__________-____-----__--_--___ 14
7. Rock pinnacles ("sea stacks"), Makarius Bay __________________________---_---__-___----___ 14
8. Aerial photograph of the Empetrum heath in an area of many lakes.______-__--_---__-____-_-___ 15
9. Hummocky Empetnim-Carex-lichen community in the vicinity of Cyril Cove____________________ 16
10. A well-developed moss mound in a Cladonia-Carex meadow near Cyril Cove_____________________ 17
11. Ecotone of an Empelrum-Carex-lichQH community and a Cladonia-Carex meadow, near Cyril Cove__ 17
12. Organic bird-perch mound showing characteristic columnar summit composed of Poa arctica subsp.
williamsii, near White House Cove.______________-______--_--______---_---_--____----_- 18
13. Streamside community near Cyril Cove____________________________-__-___--____--__-----__- 19
14. Peat-bank community in the Empetrum heath near Constantine Harbor.________________________ 20
15. Frost scar in the Empetrum heath near Cyril Cove.______-___-_-___-_____-_-__--__-___-_--_-_ 20
16. Spring issuing from the margin of a large soligenous bog in a broad valley near Cyril Cove________ 21
17. Wet sedge-meadow community at the margin of a large lake near Cyril Cove____________________ 22
18. Wet sedge meadow near Cyril Cove.______________________----____-___--_-_---_--___------- 22
19. Ice ridge on the leeward bank of a large lake near Cyril Cove._________________________________ 23
20. Small pool supporting a Juncus-Eriophorum community, surrounded by a Cladonia-Carex meadow,
near Cyril Cove._____________________________________________________________________ 24
in
IV CONTENTS
FIGURES 2-35. Photographs Continued Page

21. Bottom of a dry pool near Constantine Harbor bearing a dense growth of Subularia aquatica. ______ 24
22. Vertical view of a Siphula-Scapania community in a shallow rock-bottomed pool near Crown Re?fer
Point__________________________________________-.___-__.__________ 25
23. Deeply entrenched stream in the Empetrum heath near Chitka Point.__________________________ 25
24. Fragments of a roadside bank that has been severely eroded by wind___________________________ 26
25. Lupinus nootkatensis and Deschampsia caespitosa subsp. orientalis growing on the graveled shoulder
of an unused road near Constantine Harbor.____________________________________________ 26
26. The discontinuous heath habitat, with scattered patches of the Salix-Empetrum community________ 27
27. High-altitude land surfaces that are strongly influenced by solifluction processes, between Chitka
Cove and Andesite Point._____________________________________________________________ 29
28. Rock stripes that have developed on a moderate slope between Chitka Cove and Andesite Point. __ 30
29. Fell-field at the summit of a mountain near Chitka Cove______________________________________ 30
30. Solifluction terraces on the high plateau near Rim Point,_____________________________________ 30
31. Abrupt transition from fell-field to discontinouus heath at Chitka Cove_________________________ 31
32. Low-altitude rock outcrop that is used as a bird perch, near Banjo Point._______________________ 32
33. High-altitude bedrock outcrop near Chitka Cove____________________-___-___-_-_____________- 32
34. Rock outcrops on a mountain summit about 1 mile west of Chitka Cove________________________ 32
35. Sitka spruce (Picea sitchensis) that was planted in front of a club building during World War II___. 33
TABLE
Page
TABLE 1. Fungi and algae cultured from soil samples taken on Amchitka Island, Alaska_______.__-__-_______-_-_ 56
VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA
By HANSFORD T. SHACKLETTE and others
ABSTRACT 25 (no earlier reports known) ; and algae, 173 (no earlier re-
An Empetrum heath blankets the low plateaus of Amehitka ports known).
The following taxa were found for the first time in the Aleu-
Island, except where interrupted by streams, lakes, sand
tian Islands: Vascular plants Diapensia lapponica subm>.
dunes, and man-made disturbances. This heath is composed
obova-ta and Poa alpina; bryophytes Aongstroemia. longipes,
of low shrubs of Empetrum, Vaccinium, Loiseleuria, and vary-
Anastrophyllum sphenoloboides, Anomobryum conoinnatum,
ing amounts of many species of sedges, grasses, forbs, lichens,
Brachytliecium acutum, B. salebrosum, Bryum arctic-urn, B. pol-
and bryophytes. Lakes and pools contain the aquatics Hippuris,
lens, B. salinum, Dicranella subulata, Gymnocolea inftata,
fsoetes, Ranunculus, Limosella, and Subularia, and poorly
Hypnum imponen-s, Lophozia opacifolia, Lophozia incisa, Mie-
drained areas around them support communities composed
lichhofcria macrocarpa, Polytrichum sphaerothecium, Spheno 7-o-
largely of Carex or Sphagnum. Moss mounds are abundant
bus sacoatulus, and Tortula mucronifolia ; fungus Nidi-la
throughout, and bird-perch mounds occur near the coasts. The
Candida.
steep cliffs along the shores support a distinctive flora in which
Of the taxa listed above, the following were also found for
Saxifraga, Potentilla, Drafter, Eitrhynchium, and RamaUna are
the first time in Alaska: Anastrophyllum sphenoloboides, Bra-
predominant. Dense stands of Elymus grow intermixed, at
chythecium acutum, Hypnum imponcns, Lophosia opacifolia,
places, with Anemone and Ligusticum on the margins of the
Nidula Candida, Polytrichum sphaerothecium; and Sphenololus
plateaus near the sea coasts. Beaches of sand and cobbles
saccatulus (first report of its occurrence in North America).
commonly have a band of Honckcnya near the water and popu-
Five new species and a new variety of protists are described
lations of a tall Senecio among the driftwood. Sand dunes are
in this report Chla-mydomonas alasJcensis, C. subangulosa, Os-
vegetated with dense stands of Elymus and other grasses or, if
cillatoria facttis, Pseudanabaena- granulifera, Raphidonema-
pebbly, with colonies of Rhacomitrium, Schistidium, Ulota, and
gracile, and Synechococcus cedrorum var. pallidus.
Andreaea.
At altitudes between 400 and 600 feet the Empetrum mats
INTRODUCTION
lie in strips on unstable pebbly or rocky mineral soil; Salix,
Loiseleuria, Lusula, Cladonia, and Thatnnolia may also be com- By HANSFORD T. SHACKLETTE
mon components of the mats. On the unstable soil between the
strips, a few mosses (Andreaea, Rhacomitrium, and Schist- OBJECTIVES AND LIMITATIONS OF THESE STUDIPS
idium), Koenigia, Juncus, and Saaeifraga grow as scattered
colonies or individual plants. The studies reported in this paper were designed to
Solifluction ridges and frost-moved cobble deposits at alti- inventory land and aquatic plants that grow on Am-
tudes between 600 and 900 feet are sparsely colonized with inter- chitka island and to describe the relationships of plant
mixed Draba, Sibbaldia, Diapensia, Lupinns, Potentate, Con- species and plant communities to climatic and edapHc
ostomum, Luzula, Veronica, and Rhacomitrium. Mountain
streamsides and late-melting snow beds have extensive cover
factors. The fresh-water and soil algae and the fungi
of Ant7ielia, Nardia, and Harsupella, with Saxifraga, Geum, listed in this report were collected incidental to other
Pyrola, Leptarrhcna, Hippuris, Vahlodea, and other vascular studies; therefore, the lists do not represent the total
plants. microflora, The fungus Nidula Candida (Peck) White,
At altitudes between 900 and 1,100 feet most areas are covered collected during these studies, was reported by Keeves
with boulder fields in which only a few heath mats of Empetrum,
Salijc, Rhododendron, Vaccinium, and Loiseleuria occur. On the
(1968) ; other collections of fleshy fungi have not be-m
highest peaks, where winds commonly are very strong, only identified. Marine algae were not included in this study.
Carex circinnata and Agrostis horealis grow among the holders At the time of the investigations, access to the moun-
and outcrops that support Umbilicaria, Ocrolechia, Parmelia, tainous parts of the island was difficult because of poor
and other saxieolous lichens. roads and generally inclement weather; according^,
The following numbers of plant taxa are reported to occur only a short time was spent at some mountain sites.
on the island: Vascular plants, 198; bryophytes, 153, lichens, Many sites on other parts of the island were studied in
84; fungi, 25; and algae, 173; total taxa reported, 633. These
taxa are distributed among the 15 habitats and 41 plant com-
detail in both summer and winter. Plant communities
munities that are described in this report. Numbers of taxa that in all major kinds of environments were examined after
are reported for the first time to occur on Amchitka Island traveling on foot or by truck or helicopter to 40 study
are: Vascular plants, 47; bryophytes, 145; lichens, 75; fungi, sites (fig. 1). Dense fog in summer and very strong ccM
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INTRODUCTION
winds in winter at times prevented effective field stud- NOMENCLATURE, AND NOTES ON COLLECTIONS
ies, and good-quality photographs were difficult to ob- The nonmenclature of the plant species in this report
tain under the generally overcast sky. Quantitative eco-
generally adheres to that of the following sourcer:
logical studies that were begun are incomplete and are
Vascular plants, Hulten (1968); mosses, Crum, Steers,
not discussed in this report.
and Anderson (1965); lichens, Hale and Culberson,
BOTANICAL STUDIES THAT HAVE BEEN CONDUCTED (1966); and liverworts, algae, and fungi, from various
sources as judged authentic by Herman Persson, H. Sku-
The first known botanical investigation of Amchitka ja, and L. W. Durrell. Some species found on Amchitka
Island was made by W. H. Ball of the U.S. Coast Sur- Island are not listed in the generally accepted references
vey on July 30, 1873. His collection of plants is in the given above. For these species the nomenclature fol-
U.S. National Herbarium and the Gray Herbarium of lows the usage of specialists in the particular taxo-
Harvard University (Hulten, 1960, p. 12). No other nomic group.
plant collections were made until June 11-12,1929, when Most plants listed in this report were collected by
Misao Tatewaki and K. Takahashi visited Constantine Shacklette, and they bear his collection numbers. He
Harbor. This area was visited by Yoshio Kobayashi in made approximately 790 numbered collections on Am-
the summer of 1931. The reports of these Japanese bot- chitka Island, each at least in triplicate, and one speci-
anists were given by Akiyama (1933), Tatewaki and men of each is held in the U.S. Geological Survey herb-
Kobayashi (1934), and Satake (1932). Eric Hulten arium in Denver, Colo., for eventual deposit in the U.S.
collected plants on this island July 9,1932, near "Square National Herbarium. The other two specimens of each
Bluff" (Bird Cape) and in the middle of the north collection were distributed as follows: Vascular plants
shore. Reports of these collections, as well as sight rec- and bryophytes to the Herbarium of the Swedish Natur-
ords, are given in his "Flora of the Aleutian Islands" al History Museum, Stockholm, and to the Herbarium
(Hulten, 1960). In 1936 O. J. Murie collected plants of the University of Michigan Museums, Ann Arbor;
on Amchitka Island while studying the avifauna. and lichens to the Herbarium of the Botanical Museum,
Hutchison (1942, p. 783) spent 2 clays at Constantine Oslo, and to the Herbarium of the University of Colo-
Harbor in the late summer of 1936 and collected about rado Museum, Boulder. The algae collections were seg-
50 species of flowering plants. These specimens are in regated from the bryophyte and Isoetes collections, ard
the British Museum (Hutchison, 1937, p. 144-145). On the segregates are either being held at the University of
August 22, 1949, George A. Llano collected lichens Uppsala or being cultured from soil samples at Colorado
and bryophytes on this island. State University.
The first systematic collecting of bryophytes, lichens,
and vascular plants on Amchitka Island was done by ACKNOWLEDGMENTS
H. T. Shacklette, in connection with U.S. Geological
This research was supported by the Advanced Be-
Survey investigations, in November and December 1965,
search Projects Agency (ARPA) of the Department
August 1966, and June and July 1967. E. E. C. Clebsch
of Defense and was monitored by Hansford T. Shack-
collected vascular plants and made ecological observa-
lette under ARPA Order No. 938. Dr. Stanley Ruby
tions on the island in late June 1967, and A. J. Sharp
and Lt. Col. Charles Akard, both with ARPA, were
collected bryophytes in early July of the same year. J.
most helpful in expediting this work, and Lieutenant
A. Erdman, M. A. Heifner, J. R. Keith, W. M. Klein,
Colonel Akard also assisted in the first field study.
and H. T. Shacklette collected plants and conducted
Messrs. W. S. Twenhofel, R. E. Davis, R. H. Morris,
ecological studies in July 1967; these studies were con-
and L. M. Gard, all of the U.S. Geological Survey, as-
tinued in August 1967 by Klein and Heifner, assisted by
sisted with administrative matters and field studies, and
R. T. Ward and W. L. Slauson. In September and Oc-
their cooperation is greatly appreciated. The author
tober 1967 and in January and early February 1968, C.
A. Amundsen and Clebsch continued their ecological thanks Messrs. Ted Toren and Joe Brown of the U.S.
studies on the island. Shacklette and Erdman collected Atomic Energy Commission for logistical support on
plants and studied the phytoecology of the island Feb- Amchitka Island in June, July, and August 1967 and
ruary 5-20, 1968. These studies were continued by in February and March 1968. Mr. Odd Clemmetson
Keith, Heifner, and Slauson during March 19-26, 1968, helped in many ways with housing and transportation
and by R. L. Dix, Kenneth Bierly, Richard Cooley, on the island during all periods of field study. The au-
Heifner, David Putman, Skuson, and Walter Younkin thor also thanks Mrs. Jessie Bowles for her assistarce
for various periods during the summer of 1968. in the preparation of specimens and manuscript.
350-274 O - 69 - 2
The critical studies of Amchitka Island specimens extensive network of roads and three airplane runways
made by Drs. L. W. Durrell, Eric Hulten, Hildur Krog, were built in the general area of Constantine Harbor,
Herman Persson, H. Skuja, and W. A. Weber included and one road was constructed from this ar°»a to the
in this report were supported entirely by their own in- northwest point of the island a distance of about 30
stitutions. The author gratefully acknowledges this miles. Apparently, there have been no permanent resi-
support and the invaluable assistance of these dents or commercial establishments on the island since
specialists. the military occupation of World War II.
From 1946 to 1963 the island was visited from time
DESCRIPTION OF THE ISLAND AND ITS VEGETATION
to time by personnel of the Fish and Wildlife Service,
By HANSFORD T. SHACKXETTE the U.S. Weather Bureau, and the U.S. Geological
GEOGRAPHY
Survey (Coats, 1956, p. 84; Powers and others, 1960,
p. 552). In 1964 work was begun to prepr.re a site
Amchitka Island, the southernmost of the Kat Islands, for nuclear testing, and in October 1965 an 80-kiloton
has an area of 114.1 square miles and is the fifth largest nuclear device, buried at a depth of 2,300 feet, was det-
Aleutian Island (Coats, 1956, p. 86). The island is onated (Army Research and Development Fewsmag-
about 35 miles long, in a northwesterly direction, and azine, 1966).
is 3 to 5 miles wide (Powers and others, 1960, p. 522). No native land mammals are on the island. Norway
The Bering Sea is to the north, and the Pacific Ocean, rats were introduced probably during World War II
to the south; therefore, the island is in a region of tur- and are now abundant on the eastern half of the island,
bulent winds and waters. occurring most commonly along the coasts. Foxes, as
The settlement of the island by the Aleuts occurred well as feral dogs remaining from the military occu-
more than 4,000 years ago, and until Russian occupation pation, were exterminated after 1945 by the Fish and
the island apparently supported a relatively large native Wildlife Service as a measure to encourage birds to nest
population (Powers and others, 1960, p. 552). The loca- on the island. Sea otters, Stellar's sealions, and harbor
tions of 40 villages are indicated on a map by Guggen- seals are common in some coves and on the beaches.
heim (1945, p. 22-23), who reported that many sites There are no ants on the island (Hulten, 1960, p. 29),
had been plundered during the military occupation of mosquitoes (if present) are rare, and earthworms are
World War II. All known sites of Aleut huts (bara- not common, but certain species of Diptera and Hymen-
baras) are along the coasts, commonly on sandy de- optera are numerous. The avifauna was reported by
posits. At least one barabara was still habitable in Feb- Murie (1959), Kenyon (1961), and others; waterfowl,
ruary 1968, but all that remains to indicate the location ocean birds, and eagles are abundant, and ptarmigans
of most huts is a rectangular pattern in the vegetation. are frequently seen on the heath. The only trees on the
The Russian expeditions to the Aleutian Islands in island are several Sitka spruce (Picea sitchensis} , now
the 18th century were for the purpose of obtaining furs about 3 feet high, that were planted in front cf military
of sea otter (Barbeau, 1958, p. 28-32). A Russian vil- buildings during the war. Several species of weedy
lage, inhabited by Aleuts and people of Russian de- herbs were probably brought to the island by man,
scent, was established at Constantine Harbor and ap- but they constitute an insignificant part of the present
parently flourished for many years; however, when flora.
Hutchison (1937, p. 145) visited the island in 1936, it CLIMATE
was entirely uninhabited, although the Russian church
was in a good state of preservation. At present, the only The Amchitka Island climate is maritime, being cool,
remaining evidence of this early habitation is a small stormy, and cloudy throughout the year (Arctic
cemetery at the head of Constantine Harbor in which Weather Central, 1950). Average temperatures range
the most recent tombstone bears a death date of 1930. from 0°C in January and February to 9°C in August
In 1913 the entire Aleutian Island chain was set aside (extreme low, 9.5°C in January; extreme high 18.5°C
as a National Wildlife Refuge under the supervision in July). Partial to complete cloud cover is rearly con-
of the Fish and Wildlife Service, U.S. Department of stant; it ranges from 99.7 percent of the tine in July
the Interior. From January 12,1943, until early Novem- to 96.8 percent of the time in October. During the sum-
ber 1950, Amchitka Island was used as a military base mer, fog occurs more than 50 percent of the time, and
and, at times, was occupied by thousands of troops. complete overcast occurs more than 65 percent of the
Many square miles of the eastern part of the island are time. In the winter, there is less fog and overcast. Aver-
today littered with the ruins of military buildings, most age wind speeds range from 22 knots in January to 14
of which are uninhabitable. During this occupation, an knots in June and July (monthly maximum^ of 52-100
knots). Precipitation averages 35 inches annually, in- from the headwalls of the amphitheater valleys g,re
cluding snowfall of 70 inches. The above summary of benches or remnants of valley floors at altitudes of 700
the climate is based on records made from 1943 to 1948 to 800 feet in most places. Most of the bedrock surface
at the airbase (alt 225 ft); doubtless, the mountainous of the benches is grooved and striated, and tarns s.re
part of the island is colder and windier. present on some benches and valley floors.
In this study, soil temperatures were measured with a High plateaus occur in the segment of the island
recording thermograph for 9-day periods in each of 4 northwest of Windy Island, at altitudes between 700
months. For these periods, averages of hourly temper- and 800 feet. Two separate areas of undissected surface
atures, measured at a depth of 20 cm (centimeters) form the two mesas at the west end of the island. The
below the surface of the organic mat (a depth consid- high-plateau surface is underlain by colluvium of
ered to be within the zone of greatest root activity), angular rock fragments.
follow: June, 5.0°C; July 7.7C C; August, 7.2°C; and The oldest rocks, probably early Tertiary in age, r.re
February, 0°C. volcanic ash, tuff, breccia, and lava flows generally of
Barometric pressures fluctuate frequently and often andesitic composition that apparently were largely
greatly. During the 2-week period February 9-19, 1968, erupted and emplaced under the ocean. These rocks
high pressures of 29.48, 28.70, and 29.82 inches of mer- were deformed, uplifted, and eroded before and during
cury alternated with low pressures of 28.23 and 28.15. deposition of the interbedded conglomerate, basaltic
Abrupt changes in weather generally accompanied breccia, and tuff of a later sequence of Oligocene or pos-
these barometric fluctuations. sibly early Miocene age. Erosion reduced the mass to a
surface of low relief, now at an altitude of about 1,100
PHYSIOGRAPHY AND GEOLOGY feet, that was possibly a submarine shoal. Marine sards
The physiography and geology of Amchitka Island and cobble conglomerate are now exposed at altitudes
were described by Coats (1956) and by Powers, Coats, of up to 600 feet.
land Nelson (1960). The discussion that follows is based During late Tertiary and Quaternary times, uplift of
on these reports. at least 500 feet, and possibly more than 1,100 feet, took
The three landforms on the island are, from east to place differentially and spasmodically. Abandoned pm
west, low plateaus, rugged mountains, and high cliffs and beach deposits are inferred to be structurally
plateaus (fig. 1). This varied topography is thought to dislocated remnants of shoreline features of a late Pleis-
have been developed from plateaus of low relief that tocene interglaciial sea level, judged from the fact tl at
were somewhat disturbed by block faulting and were the fauna of the beach deposit is characteristic of water
modified by dissection to greatly different degrees by warmer than that which now surrounds the island.
marine, stream, and glacial erosion. SOILS
Elevated wave-cut platforms, at altitudes ranging
from 135 to about 500 feet, constitute the lower plateaus The soils of Amchitka Island have been insufficiently
of the island. Several successive marine benches com- studied to be classified by soil series; hence, at present
monly are present; many are poorly drained and have they can be placed only in the great soil groups to which
lakes of different sizes and shapes. On eastern Am- they appear to belong. The following outline of these
chitka Island, glacial boulders occur beneath marine soil groups that are represented by the soils on this
gravels at altitudes of more than 100 feet. island is based on the classification of Alaskan soils by
The mountainous part of the island is between Chitka Kellogg and Nygard (1951, p. 33) and on certain con-
Point and Windy Island and is a sinuous ridge that cepts of tundna soils on Kodiak Mand that were pro-
divides the drainage to the Pacific Ocean and to the Ber- posed by Kieger and Wunderlich (1960, p. 27~32).
ing Sea. This ridge has several summits exceeding 1,100 A. Zonal soils
feet in altitude, and the summits iare separated by four 1. Tundra
passes at altitudes of about 750 feet. Five prominent a. Without permafrost (Ando soils of Rieger rnd
spurs extend from the main ridge and form divides be- Wunderlich)
tween major compound amphitheater valleys that head B. Intrazonal soils
in the main ridge. Parts of the ridge and spur summits 1. Bog
are .gentle slopes eroded across the local rock structure 2. Half Bog
and are remnants of an erosional surface of very low C. Azonal soils
1. Alluvial
relief, but it is not known whether the cut surface has a. From local alluvium
been glaciated or even whether it is subaeriial or sub- 2. Lithosol
marine in origin. At the base of the steep slopes leading 3. Regosol
6 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA
Eieger and Wunderlich (1960, p. 27) classified certain in the order Histosols (Soil Survey Staff, 196C, p. 247-
soils as Ando soils; these same soils were classified 248). However, some of the Half Bog soils may belong
earlier by Kellogg and Nygard (1951) as "Tundra to a different order, and they could be classified further
without permafrost." Inasmuch as microclimates and if sufficient data were at hand.
soils on Kodiak Island are similar in many respects to Alluvial. These soils cannot be correlated with the
those on Amchitka Island, it is possible that some of the new system on the basis of available data.
Tundra without permafrost soils of the latter island Lithosols. The Amchitka Island lithosols are classi-
should be named Ando soils. They are, however, not so fied as follows (Soil Survey Staff, 1960, p. 136-139;
strongly influenced by volcanic ash as are those of 1967, p. 93): Order, Inseptisol; suborder, Aquepts;
Kodiak Island. great group, Cryaquepts; and subgroup, Histic Crya-
In 1960 the Soil Survey Staff, U.S. Department of quepts.
Agriculture, published a new system of soil classifica- Regosols. These soils, at both high and low altitudes
tion and in 1967 they expanded and refined the system on Amchitka Island, are classified as follows (Soil Sur-
in a supplement. Their definition of soil requires that vey Staff, 1960, p. 136-143; 1967, p. 98): O"der, In-
plant life be supported; however, an area covered with septisol ; suborder, Ochrepts; great group, Cryochrepts;
water that supports only floating plants or with rocks and subgroup, Lithic Cryochrepts.
that support only lichens is not considered to have soil. The most recent worldwide classification cf soils is
The lower boundary of soil is considered to be (Soil that of Ball (1967), who devised a system for coding
Survey Staff, 1960, p. 1) "the vague lower limit of soils found in terrestrial biological communities that
common rooting of the dominant native perennial may be proposed for conservation in the International
plants, or the vague lower limit of the genetic horizons, Biological Program. This classification recognizes 13
whichever is the deeper." Within the limits of this categories of soil that are designated by letter symbols
definition, an extensive system of classification and with number subscripts. By eliminating the categories
nomenclature was developed on the premise that soils of soils that do not occur on Amchitka Island, the au-
should be classified by their properties rather than by
thor has prepared a key to the soils of Amchitha Island,
their genesis. (For a review, see Shacklette, 1961.) If
using the system of Ball (1967, p. 123-124), as follows:
the soils of Amchitka Island are to be fully classified
by the system of the Soil Survey Staff, a much more 1. Soil with dominantly organic surface horizon at least
intensive study of their properties than has yet been 50 cm deep. If total soil depth less than 50 cm, then
made will be necessary. This new system classifies soils surface organic horizon directly succeeded by un-
into so many more groups than is possible under the altered rock soil type O.
old system that only examples considered to be typical of Soil without dominantly organic surface horizon or
a Great Soil Group are classified below that is, there with organic surface horizon succeeded by mineral
probably is more than one subgroup in an order on soil at less than 50 cm depth 2.
Amchitka Island. The correlation of the old and the 2. Well drained (that is, no evidence of strong im-
new systems of classification, insofar as it can be done pedance or waterlogging above 40 cm depth) 3.
at present, is given below. Poorly drained (that is, evidence in mottled colors
Tundra without permafrost. If these soils on of strong impedance or waterlogging nearer sur-
Amchitka Island are to be related to Ando soils, the face than 40 cm) Soil type P2.
following classification is suggested (Soil Survey Staff, 3. Immature profile (that is, with weakly developed
1960, p. 136-140; 1967, p. 89-90) : Order, Inceptisols; shallow soil formation, possibly with little bio-
suborder, Andepts; great group, Cryandepts; and sub- logical activity) 4.
group, Lithic Cryandepts. If, however, the small Well-developed horizon sequence with moderate to
amount of ash in their horizons as a whole is considered strong biological activity 5.
to be most characteristic, they are classified as follows 4. Immaturity resulting from climatic factors; for ex-
(Soil Survey Staff, 1960, p. 136-143; 1967, p. 107-108) : ample, very low rainfall or temperature, or both
Order, Inseptisols; suborder, Umbrepts; great group, soil type Ii
Cryumbrepts; and subgroup, Lithic Cryumbrepts. Immaturity resulting from lack of time for soil for-
Bog and Half Bog. Because the new system is not mation to proceed; for example, on recent alluvium,
yet complete, these soils can be classified only as being dune sands, and eroded surfaces soil type I2
5. Shallow or simple profiles of A horizons overlying The surficial material on the high plateaus, owing to
parent material soil type F3 less relief, has greater stability than that on the moun-
A(B) C profiles a soil type F4 tain section, and large areas of sparse vegetation occur
ABC profiles; that is, with a horizon accumulation of on the Lithosol of the rock fields. Interrupted vegeta-
clay or iron oxides, or both soil type F5 tion mats, in which Empetrum, Vaccinium^ and Saliw
predominate, develop a thin organic soil in and under
Because the classification of the Soil Survey Staff
them, but the profile of this soil has little in common
(1960, 1967) cannot at present be fitted to all soils on
with that of the Tundra soils at lower altitudes.
the island, subsequent parts of this report will use the
The author did not observe permafrost in any section
classification based primarily on Kellogg and Nygard's
of Amchitka Island. Kellogg and Nygard (1951, p. 11,
(1951) that was presented earlier, and the system of
135) stated that there is none in the Aleutian Islands.
Ball (1967). However, local areas of perennially frozen ground pos-
In 1946 Kellogg and Nygard (1951, p. 41, 43), of the sibly occur under the late-melting snow beds in the
U.S. Department of Agriculture, obtained soil samples
mountains of Amchitka Island, and Lithosols at the
and data for a profile description of a smooth, fairly
margins of permanent snowfields on some of the higher
well drained slope on Amchitka Island. They classified
Aleutian Islands are probably underlain by frozen
the soil as Tundra without permafrost and gave the
ground.
following description of the soil profile:
All soil profiles of the low plateaus of Amchitka Is-
Aoo and Ao. 5 to 0 inches, dark-reddish-brown peaty mat. land examined in the present study contained an ash
Ai. 0 to 5 inches, nearly black humus-rich silty very fine layer 1-1.5 cm thick in the upper part of the B horizon
sandy loam; the slick moist soil is held together by
many roots.
and two ash layers of about the same thickness in the
B2. 5 to 10 inches, nearly black granular silty very fine lower part of the B horizon. A soil profile on Tanaga
sandy loam. Island, about 115 miles east of Amchitka Island, was
B3. 10 to 15 inches +, dark-grayish-brown silty very reported by Anderson and Bank (1952, p. 84) to have
fine sandy loam. In the lower part, horizontal streaks somewhat similar, but more numerous, ash layers. There
suggest ash layers.
have been no volcanoes on Amchitka Island since the
C. At 30 inches, yellowish-brown silty very fine sandy loam.
present soil was formed, but recently active volcanoes
The low plateaus of Amchitka Island, because of on other islands in this region may have produced this
their varied relief, bear soils that include all the soil ash. Powers (1958, p. 67) reported the activity of Cer-
groups on the island. The soil profile given above is berus Volcano on Semisopochnoi Island, which is about
probably typical of soils of the Empetrwm, heath on the 40 miles north of Amchitka Island, as follows: 1772,
undulating plateau surfaces. Land surfaces of steeper smoke; 1790-92, smoke; 1830, smoke; 1873, active. He
slope may have Half Bog soils and soil intergradations referred to smoke as "a word used in records; it refers
between Half Bog and Bog soils at the base of the slope. to steam or steam-and-ash clouds." Therefore, it is
Small areas of local Alluvial soil are common near possible that the ash deposits on Amchitka Island came
streams, and Bog soils occur at lake margins and on from the Semisopochnoi Volcano or from volcanoes on
nearly flat areas of the plateaus. Sand dunes near the other islands nearby. Powers (1958, p. 67) reported for
coasts and beach sands are considered to be Kegosols. Little Sitkin Island, "1776, active; 1828, smoke;" and
Shallow Lithosols develop on rock outcrops, both along for Segula and Kiska Islands, "steaming intermittent."
the coasts and inland. A trench that was dug through a peaty soil deposit of
The mountain section of Amchitka Island, because of the Empetrum heath on Amchitka Island in 1965 ex-
the predominance of rock outcrops and colluvium, con- posed three ash layers in the profile. This trench was
tains most Lithosols. Bog and Half Bog soils, however, examined again in July 1967; a moss, Pogonatum umi-
have developed along some of the high mountain streams gerum, had colonized only the ash layers. The peat part
and around the tarns of the large amphitheaters. Much of the profile bore no mosses or other vegetation; there-
of the total area of unconsolidated surficial deposits at fore, the growth of the moss on the ash made the ash
high altitudes supports litle or no vegetation because of layers very conspicuous (fig. 2). The "preference" of
instability produced from steepness of slope and frost this moss for a mineral soil was known from other ob-
action; these deposits cannot be considered to be soil. servations, but only rarely is such a specific requirement
for a particular substrate demonstrated so clearly.
1 (B) and B horizons are defined (Ball, 1967, p. 121) as, "Horizons
lying between A horizons and the parent material C horizons, which show- Ash layers in the profile described above contained
weathering and release of iron oxides without their transportation, (B) ; preserved stems of sedge and Empetrum. These stems
or which are horizons of deposition of transported sesquioxides and/or
clay, B." were removed from the peat and ash matrix; radiocar-
on the island can be seen by the vigorous growth of many

plant species on bird-perch mounds, calcareous dunes,
kitchen middens, barabara sites, and recently formed
soil banks (fig. 3). The Amchitka Island soils, if com-
pared with agricultural soils or with the soils of most
temperate-zone plant communities, are low in nutrient
elements, especially calcium, phosphorus, and nitrogen.
Kellogg and Nygard (1951, p. 41) gave a chemical
analysis of a soil sample from Amchitka Island and
wrote, "The data * * * show a high exchange capacity
but a very low base saturation. The magnesium is low,
but very high in relation to calcium. Available phospho-
rus is exceedingly low." They (1951, p. 43,116) reported
the following phosphorus content, in parts per million,
of samples from this island:
Ash of a sedge-grass sample __ ____ 24
Soil samples [presumably not ashed] :
Aw horizon______ 12
An horizon_________ 2
B to Chorizons [4 samples] 0
A similar trend in calcium and potassium content of

soils and plants from Amchitka Island was reported by
FIGURE 2. Section of a peat deposit with an upper (a) and two Kellogg and Nygard. They also reported a carbon-nitro-
lower (;&) ash layers that are marked by a thick growth of the
moss Poffonatum urnigerum. Timber (2 in. X 4 in.) in upper
gen ratio (1951, p. 14) of 15 to 1 in the AI soil horizon,
center provides scale. No other plants grew on the vertical as compared with that of 26 to 1 in the B2 horizon, which
bank, which was formed by excavation approximately 2 years suggests the formation of peat in place accompanied by
before this photograph was made on July 28,1967. a loss of nitrogen by leaching.
bon dates of samples from the top, middle, and lower

ash layers were 750 ±250 years, 1,950±250 years, and
I,740=fc250 years B.P. (before present), respectively.
The middle and lower ash layers were so close together,
and the stems in them were so intermixed, that satisfac-
tory separation of the stems was not possible. The dates
of these stem samples, therefore, were averaged as 1,845
years B.P. (Shacklette and Rubin, 1969).
Although mechanical properties of soils related to en-
gineering operations have 'been studied on Amchitba
Island, results have not been published. Such studies,
because of the broad definition of soil that is used, have
but little relevance to ecological investigations.
An evaluation of the soils of this island as media for
the support of native plants is best made by observing
the vegetation that grows on the soils. From this stand-
point, the soils appear to be fertile, because most land
surfaces especially those of the low plateaus are cov- FIGURE 3. Effects of soil fertility on the growth of the grass
ered with dense mats of vegetation. This dense plant Calamagrostis nutkaensis. Soil banks had been formed in dig-
growth, however, does not indicate that the soils meet ging-in a pyramidal tent; later a latrine was located on this
the requirements for optimum growth of most plant tent site and was closed February 4, 1944. The grass is least
species that are present, nor does it indicate that the soils vigorous on the thin rocky soil in the foreground, more vigor-
ous on the soil banks, and most vigorous on the latrine site,
are fertile, as judged by agricultural standards. The where the grass tussocks are about 1 meter high. Photo-
effects of a soil that is more fertile than the average soil graphed February 16, 1968
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 9
Soil profile samples were collected from an Empet- nonrandom association, although many species, if con-
rum heath on Amchitka Island by Shacklette on Au- sidered individually, appear to be somewhat randomly
gust 13,1965. Descriptions of these samples, which were distributed throughout one or more types of habitat.
taken at 4.5-cm intervals of a soil profile 0.5 m (meter) Groups of plants that are composed largely of vascular
thick; their percentages of ash; and their chemical con- plants, of bryophytes, or of lichens were given equal
tents, expressed as percent of ash, follow: consideration in establishing the communities. Some
communities that are described are much larger than
Cal-
others in a real extent and number of component spe-
Sample description Ash Potas- Phos- SO4
cium sitim phorus radical cies. The size of a community is determined largely by
the areal extent of habitat factors in particular combi-
Living heath plants, mixed __ . _ _ . --- 5.7 7.5 7.2 2.0 3.6£
Dead heath plants (Aoo horizon). .... .--- 6.2 8.2 2.2 1.6 4.1*
3.24
nations. Quantitative studies of the compositional ele-
Peat--. .... - 11.0 3.8 1.7 .8
Mucky peat.-.. -- _ ... _ --- __ --........ 13.7 3.5 1.7 .9 3.93 ments of a community have not been completed; the
Peaty muck.. ___ - ________ -...... 26.7 3.4 1.4 .4 2.14
Muck (including ash layer) ..- ____ ....... 57.2 3.7 1.3 .16 .93 present establishment of the communities is subjective
Muck ---.--_--.-.-- __ -.-- __ _ ......... 21.5 2.2 1.1 .6 2.05
Muck (including ash layer) -------- _ . 44.2 3.3 .9 .3 1.33 to a high degree and is likely to be modified when quan-
Muck (including ash layer) - __ ..--....... . 45.0 2.3 1.1 .2 1.17
Muck..-. _ -. __ _ _ ---- _ .-----....... 31.1 1.7 1.0 .4 1.17 titative data are available.
Organic and mineral soil 46.5 2.1 .6 .3 .97
The vegetation of Amchitka Island can be described
conveniently by presenting the species according to
These samples were analyzed by T. F. Harms and their natural occurence in plant communities, then re-
C. S. Papp, U.S. Geological Survey, Denver, Colo. lating these communities to distinct habitats in which
The concentration of these elements in plants is they occur. Causal factors in the development of plant
clearly shown by these data, and a greatly abbreviated communities may be implied in this method of de-
biogeochemical cycling of these elements is indicated. scribing the organization of the vegetation, but they are
That is to say, although the supply of these essential not stated more positively than this investigation war-
nutrients is deficient in the pedosphere as a whole, the rants. Detailed measurements of physical factors of the
amounts that are present are held where they have the habitats are at present almost entirely lacking; like-
greatest beneficial effects on plants in the upper root wise, the responses of many plant species to these fac-
zones and in the plants themselves. The trends in nu- tors are largely unknown. Within these limitations
trient-element concentration in plants and soil horizons imposed on ecological interpretations, however, the
shown in the samples just described were supported by communities and habitats may be described in such a
those found by analyses of other similar samples. manner that the kinds of substrates and microclimates
essential for the formation of the communities are sug-
PLANT HABITATS AND PLANT COMMUNITIES
gested. At the same time, the occurrence of individual
Plant communities of the Aleutian Islands have been species in only one or in several communities is an indi-
described by Hulten (1933, 1960), Tatewaki and Ko- cation of their ecological amplitude, whatever their
bayashi (1934), Walker (1945), and Bank (1951,1956). specific requirements and tolerances may be.
These authors, however, almost entirely neglected the Some plant communities are separated by ecotones
bryophytes and lichens in the communities or associa- that are as great in extent as the adjacent communities
tions that they established; yet these two groups of themselves; hence, the exact delimitations of each com-
cryptogams are very important and, at places, dominant munity can be made only arbitrarily. An example of
elements of plant communities in the Aleutian Islands. communities that have this relationship are the Empet-
One objective of the present study was to give full con- rum-Carex-lichen and the Careoe-lichen meadow com-
sideration to all these groups of plants in describing the munities. Similarly, the latter community may inter-
vegetation of Amchitka Island; therefore, the ecologi- grade, through a broad ecotone, with a wet-sedge
cal units of the former investigators were not adopted meadow community.
in full, although there is a general correspondence of The vegetation on this island is mainly one layered;
many plant communities described in this paper to those there are but few examples of one plant species regu-
of the earlier writers. larly overtopping another, if a few tall herbs and some
The organization of vegetation that is given in this of the minute liverworts and mosses are excluded. There
paper is intended neither to form a hierarchical system are no native trees, and the low shrubs are intertwined
nor to establish plant communities as formal taxonomic with herbs, mosses, and lichens in a manner that allows
entities in a phytosociological system. It is, on the other one, then another, to lie on top of the intricate mat of
hand, a method of naming groups that are easily rec- vegetation. For example, Empetrum may overgrow, or
ognized in the field to be composed of species that have be overgrown by, certain mosses and lichens. The pros-
trate willows have leafy stems that may temporarily plete only insofar as collected specimens and field notes
suppress mosses and lichens, but eventually these indicate the presence of the species. An enumeration of
stems are completely buried by the moss and lichen all species that occur in some communities would add
mat. In some places, saxicolous mosses and lichens grow new names to the list but perhaps would not contribute
over each other. significantly to a better characterization of the
An adequate description of community dynamics on community.
Amchitka Island must await further study. If phases The land habitats of Amchitka Island and the com-
of succession are believed to be evident, however, they munities of plants that occur in these habitats are out-
are so stated. The roles of some biotic, edaphic, and lined as follows:
microclimatic agents in the establishment and perpetu- 1. iStrand
ation of certain communities are strongly indicated by a. Honckenya-Senecio community
characteristics that can be observed in the field. Like- 2. Dune
wise, the interaction of species in a community can. at ia. Poa-Deschampsia-Festuca community
places, be appraised. The interpretations of community h. Elymus community
c. Rhacomitrium-Schistidium-Ulota community
dynamics presented in this paper are, however, only
3. Sea cliff
provisional. a. Eurhynchium-Pucainellia-Caloplaca community
The names of plant communities have been derived b. Potentillea-Draba-Sawifraga community
from either their characteristic species or their distinc- c. Xanthoria-Ramalina, community
tive site features, according to which terminology is d. Elymus-Ligusticum-^nemone community
4. Empetrum heath
more useful for recognizing the communities in the
a. Empetrum-Ccwese-Iichen community
field. In this paper, usage of the term "characteristic b. Cladonia-Carew meadow community
species" agrees in general with that of Costing (1956, p. c. Moss mound community
74-75) and of other writers who include in this term the d. Organic bird-perch mound community
species with a high degree of fidelity (exclusiveness) e. iStreamside community
to a particular community. An estimate of the quantita- f. 'Peat-bank community
g. Frost-scar community
tive characters abundance and cover, however, was also 6. Bog
used in some of the more complex communities to select a. Sphagnum bog community
species that are considered to be characteristic of a com- b. Philonotis-Pamassia community
munity, even though their fidelity to the community is c. Scapania-Nardia^Marsupella community
low. For example, Elymus armarius subsp. irwllis var. 6. Marsh
a. Wet sedge-meadow community
mollis is considered to be a characteristic species of two b. Caltha-Claytonia community
communities, being dominant in one and codominant 7. Lake
with Ligusticum and Anemone in the other; moreover, a. Hippuris-Ranunculus community
it is listed as occurring in several other communities. b. Isoetes-Ranunculus-Limosella community
Similarly, Empetrum nigrum subsp. nigrum is a charac- 8. .Pool
a. Juncus-Eriophorum community
teristic species in two communities and occurs in nine b. Subularia-CaMitriche community
others. To ignore these species as characteristic of some c. SiphulOrScapama community
communities because their degree of fidelity is low 9. Stream
would not be a realistic use of the word "characteristic" a. Fontinalis-Ranunculus community
in its less restrictive meaning and would not convey the 10. Buderal
a. Calamaffrostis community
importance of the species in the community. If quanti- b. Lupinus-Amica community
tative data were available, the importance of species in c. Equisetum-Sagina-Ceratodon community
a community could be more accurately stated than by 11. Discontinuous heath
using the term "characteristic species" in these eco- a. Salix-Empetrum community
logical discussions. b. Andreaea-Schistidium-Rhaoomitrium community
12. iSolifluction
In contrast to the examples given above, some of the a. Fell-field community
species that grow on sea cliffs and in snow beds occur b. Solifluction terrace community
only in a single type of community. They have, there- 13. Alpine meadow
fore, the highest degree of fidelity and, also, are con- a. <Snow-bed community
sidered to be characteristic species. b. Alpine meadow community
14. Inland bedrock
Other species that occur in ia community also are a. Low-altitude bedrock community
listed. The species list given for each community is com- b. High-altitude bedrock community
15. Miscellaneous habitats, substrates, and communities GROUP 1M2. Seasonal short grass.
a. Carcasses Community: Organic bird-perch mound.
b. Imported wood FORMATION 1M21. Seasonal orthophyll short-grass
c. Plantings meadows.
d. Burned areas Communities: Poa-Deschampsia-Fes-
tuca; wet sedge meadow; Calam-
The most recent classification of vegetation on a agrostis.
worldwide basis is that of Fosberg (1967), who devised FORMATION 1M22. Seasonal orthophyll marsh.
a system for use in coding vegetation of terrestrial bio- Community: Juncus-Eriophorum.
logical communities that may be proposed for conserva- CLASS IN. Broad-leafed herbaceous vegetation (closed vege-
tion in the International Biological Program. This sys- tation predominantly of broad-leafed herbaceous
plants).
tem uses a combination of vegetational features that GROUP 1N2. Seasonal broad-leafed herbaceous vegetation.
may be grouped under "structure" (the arrangement in Community: Honckenya-Senecio.
space of the components of vegetation), and "function" FORMATION 1N21. Seasonal broad-leafed herbaceous
(features that suggest special adaptation to past or vegetation.
present environmental situations), and relies on "com- Communities: Streamside; Caltha-
Claytonia; alpine meadow; low-
position" (the species comprising the vegetation) only altitude bedrock.
for the lowest categories. In this system, vegetation is CLASS 1O. Closed bryoid vegetation.
first classified in one of three "Primary Structural GROUP 1O1. Closed bryophyte vegetation.
Groups" closed, open, and sparse or desert. Communities: Moss mound; peat bank; Sca-
I have constructed an outline of classification, based pania-Nardia-Marsupella.
on Fosberg's system, for the plant communities of FORMATION 1O11. Sphagnum bog.
Community: Sphagnum bog.
Amchitka Island that eliminates the categories of vege- FORMATION 1O12. Moss meadow.
tation that do not occur on this island. Fosberg's sys- Communities: PhUonotis-Parnassia;
tem, as presently organized, does not permit all plant snow bed.
communities of this island to be fully classified to GROUP 1O2. Closed lichen vegetation.
Formation or Subformation. The outline that follows FORMATION 1O21. Lichen bog.
Community: Siphula-Scapania.
classifies these communities only insofar as is possible FORMATION 1O22. Lichen meadow.
without devising new categories. Community: Cladonia-Carex mea-
dow.
PRIMARY STRUCTURAL GROUP 1 CLASS IP. Submerged meadows (vegetation of rooted aquatic
herbs).
[Closed vegetation (crowns or peripheries of plants touching or overlapping)]
GROUP 1P1. Evergreen submerged meadows.
CLASS 1C. Dwarf scrub (closed predominantly woody vegetation Community: Subularia-Callitriche.
less than 0.5 m high). FORMATION 1P11. Evergreen watergrass.
GROUP 1C1. Evergreen dwarf scrub. Communities: Isoetes-Ranunculus-
FORMATION 1C12. Evergreen broad sclerophyll dwarf Limosella; Fontinalis-Ranunculus.
scrub. GROUP 1P2. Seasonal submerged meadows (plants, at least
SUBFORMATION 1C12C. Microphyllous evergreen their shoots, disappearing in winter).
dwarf scrub. FORMATION 1P21. Seasonal watergrass.
Community: Empetrum- Community: Hippuris-Ranunculus.
Carex-lichen.
GROUP 1C2. Deciduous dwarf scrub.
PRIMARY STRUCTURAL GROUP 2
FORMATION 1C21. Deciduous orthophyll dwarf scrub.
SUBFORMATION !C21a. Deciduous orthophyll dwarf [Open vegetation (plants or tufts of plants not touching, but crowns not separated by
scrub. more than their diameters; plants, not substratum, dominating the landscape)]
Community: Salix-Em-
petrum. CLASS 2G. Steppe (open herbaceous vegetation, tufts or plants
CLASS 1L. Tall grass (closed herbaceous vegetation more than discrete, yet close enough to dominate the
1 m high, predominantly graminoid). landscape).
GROUP 1L2. Seasonal tall grass (turning brown in winter). GROUP 2G2. Seasonal steppe.
FORMATION 1L21. Seasonal orthophyll tall grass. Communities: Potentilla-Draba-Saxifraga;
Communities: Elymus-Ligusticum- Equisetum-Sagina-Ceratodon.
Anemone; Elymus. CLASS 2H. Bryoid steppe.
CLASS 1M. Short grass (closed herbaceous vegetation, less than GROUP 2H2. Open lichen vegetation.
1 m high, predominantly graminoid). Community: Xanthorea-Ramalina.
350-274 O - 69 - 3
PRIMARY STRUCTURAL GROUP 3 a. Honckenya-Senecio community (1N2).

Soil: Kegosol; type ?2; pH, 6.8-7.0. This com-
[Sparse vegetation or desert (plants so scattered that substratum dominates
landscape)] munity forms a distinct zone of dense vegetation
CLASS 3B. Desert scrub (scattered shrubs in an otherwise bare
near sea level (fig. 4). The Senecio may be as much
or only ephemerally vegetated landscape). as 15 dm (decimeters) high at places and so closely
GROUP 3B1. Evergreen desert scrub. spaced that passage through the community is
Community: Fell-field. difficult. Stands of Lathyrus occur on sand banks
CLASS 3C. Desert herbaceous vegetation (scattered herbaceous of the beach, and scattered plants of Galium com-
plants only).
GROUP 3Cl. Evergreen desert herbaceous vegetation. monly grow in the piles of driftwood.
Communities: Eurhync hium-Puccinellia-Calo- Characteristic species: Honckenya peploides
placa; Andreaea-Schistidium-Rhacomitrium; subsp. major and Senecio pseudo-arnica.
high-altitude bedrock. Other species: Cerastium fischerianum, Cera-
FORMATION 3C12. Evergreen psammophyte desert. todon purpureus, Claytonia sibirica, Cratoneuron
Community: Rhacomitrium-Schistidi-
um-Ulota. filicinum, Eurhynchiwn praelongum, Galvwm
FORMATION 3C13. Lichen tundra sparse phases. aparine, G. trifidvm subsp. cohimbianwn, Hera-
Community: Solifluction terrace. cl&u/m lanatu/m, Juncus a/rcticus subsp. sitchensis,
Lathyrus maritimus subsp. pubescens, Mertensia
The occurrence of most species in the communities maritima,) Peltigera spuria, Rumex obtusifolius,
was determined by collecting specimens in the field. Sagina occidentalis, and Stellaria calycantha
Some of the easily recognized species were, however, in- subsp. calycantha.
cluded in the community descriptions on the basis of
sight records. In both the community descriptions that 2. DUNE HABITAT
follow and the annotated check lists of species, the com-
This habitat type designates sandy and pebbly de-
munities are designated by letters and the habitats by
posits that occur most commonly on top of seacoast
numbers; therefore, the communities and habitats in
which each species was found can easily be determined. bluffs that are from 25 to 100 feet above sea level, where
The community name is followed by the community the sand is moved by wind.
classification according to the system of Fosberg (1967). a. Poa-Deschampsia-Festuca community (1M21).
The great soil group, or groups, and the soil classifi- Soil: Kegosol; type P2 ; pH, 6.7 6.8. On elevated
cation, according to the system of Ball (1967) of the beach deposits of sand and shells that are 100 to
substrate on which the community occurred, follow the
community classification. The approximate pH value,
or range in values, is given for the substrates of most
communities. The pH of the water in which aquatic or
semiaquatic communities occur also is provided. These
values were determined for soils by making a slurry,
or extract, consisting of two parts, by volume, neutral
water and one part soil or other substrate and by
measuring pH with a potentiometer. The pH of water
was measured directly with the same instrument. These
tests were performed at base camp soon after the
samples were collected.
The photographs of habitats and communities were
made by Shacklette, except as noted.
1. STRAND HABITAT
Occurs around the island wherever a shoreline bench

bears a deposit of sand, pebbles, or cobbles. The pres-
ence of abundant driftwood in this habitat indicates
that this habitat is inundated by sea water from time FIGURE 4. Zonation of Honckenya (nearest the bare beach) and
to time. Senecio at Makarius Bay. Photographed June 17,1967.
200 feet above the present sea level. Calcium de- lanatum are sparsely scattered among the tussocks.
rived from the shells probably favors the develop- The lichens Caloplaca sp. and Peltigera comma, were
ment of this community that is composed almost found on moss-covered driftwood in this com-
entirely of grasses and Equisetum. munity.
Characteristic species: Poa eminens, Deschamp-
c. Rha£omitrium-)Schistidium-Ulota community
sia beringensis, and Festuca rubra subsp. aucta.
(3C12).
Other species: Agrostis exarata, Bromus sitch-
ensis var. aleutensis, Elymus arena/ring subsp. Soil: Regosol; type I2 ; pH, 6.8-6.9. On dunes
mollis, Eguisetum arvense, and Poa lanata. composed of dark-colored sand, pebbles, and cobble.
b. Elymus community (1L21). Rhacomitrium and Ulota grow on the cobbles and
Soil: Kegosol; type Lz pH, 6.6-7.0. On dunes pebbles, and Schistidium grows on the sand. This
composed of dark-colored sand derived from vol- community and its unattached moss polsters of
canic rocks (fig. 5). Schistidium, were described and illustrated by
Characteristic species: Elymus arenaris subsp. Shacklette (1966).
mollis var. mollis. This community is dominated by Characteristic species: Rhacomitrium lanugino-
a dense growth of this grass which at places forms sum, Schistidium apocarpum, and Ulota
tussocks that are as much as 12 dm high and are phyllantha.
separated from each other a distance of 14-30 cm. Other species: Andreaea rupestris, Brachythe-
Inasmuch as the distinct tussocks occur on steep cium albicans , Ceratodon purpureus, Drepano-
slopes, soil slippage probably causes their separa- cladus uncinatus, Lecanora atra, L. aleutica,
tion. This community may extend to the sandy Lecidea macrocarpa, Ligusticum scoticum subsp.
beaches and dominate the old beach terraces, as is hultenii, Pogonatum urnigerum, Rhacomitrium, fas-
shown in figure 5; more commonly, however, it oc- cicidare, R. ericoides, Stereocaulon alpinum, Tor-
cupies dunes at slightly higher altitudes. tula mucronifolia and Umbilicaria proboscidea.
Other species: Bryophytes that are able to grow
3. SEA-CLIFF HABITAT
on the almost bare soil between the grass tussocks
include Antitrichia curtipendula, Bryum pseudo- The island is almost completely bordered with pre-
triquetrum, Cephaloziella arctica, Ceratodon pur- cipitous sea cliffs of andesite, breccia, and other igneous
pweus, Eurhynchium praelongum, Lophocolea rocks. At a few locations on the southeast end of the
cuspidata, Nardia scalaris, Plagiothecium roese- island, the cliffs are composed of sedimentary deposits
anum, PogonaPum umigerum, PoJila cruda, Rhy- that give but little support for plants. The cliffs range
tidiadelphus squarrosus, and Tor-tula mucronifolia. in height from only a few feet to at least 100 feet above
The flowering plants Claytonia sibirica, Geum ma- sea level on the central and southeastern parts of the
crophyllum subsp. macrophyllum, and Heracleum island; commonly, they are higher on the northwestern
shores. Plant communities on these cliffs are subjected
to waves and spray of sea water and have but little pro-
tection from the strong winds (fig. 6). The cliffs are
partly covered with ice from time to time during the
winter months.
a. Eurhynchium-Puccilennia-Caloplaca community
(3C1).
Very little, or no, soil present. Occurs on beach
boulders, rock pinnacles, and on cliff faces to the
cliff summits (fig. 7). The characteristic species of
this community have great tolerance to salt water,
and the moss and the grass are nitrophilous in that
they grow very luxuriant where manured by birds.
Characteristic species: Eurhynchium praelon-
gum, PuGcinellia langeana subsp. alaskana, and
FIGURE 5. Sand dune with cover of Elymus community at the Caloplaca granulosa.
seashore, Rifle Range Point. Grass clumps (at top) are about Other species: Blastenia sp., Bryoerythrophyl-
3 feet high. Photographed February 17,1968. lum recurvirostrum, BueUia punctata, Draba hy-
c. XanthoricL-Ramolina community (2H2).

Soil: Lithosol, or rock surfaces; type I2. Occurs
principally near the summits of high cliffs and on
"sea stacks" offshore. Xanthoria appears to require
fertilization by birds, and Ramalina grows only
where there is free air movement; both require-
ments commonly are met on cliff summits (fig. 7).
Characteristic species: Xanthorea candelarw,
Ramalina scoparia, and R. almquistii.
Other species: Amblystegiwn serpens, Carda-
mine umbellata, Caloplaca granulosa, Cystopteris
fragilis subsp. fragttis, Eurhynchium praelongvm,
Ochrolechia frigida, Parmelia saxatilis, P. sul-
-. vS«***»je " ' i>«*«a.-ii-j»«fcJC££S
FIGTJBE 6. Sea-cliff habitat on the Pacific Ocean coast at Makar-

ius Bay. Dense growths of marine algae cover the rocks of
the intertidal zone. The Elymus-Ligusticum-Anemone commu-
nity covers the upper slopes and summits of the cliffs. Photo-
graphed June 17,1967.
perborea. Lecanora aleutica, L. straminea, Physcia

caesia, PotentiUa villosa, Schistidium maritimum,
Ulota phyllantha, Verrucaria maura, Xanthoria
candelaria, and X. elegans.
b. PotentUla-Draba-/Saxifraga community (2G2).
Soil: Lithosol, in rock crevices; type I2 ; pH, «
7.0. Occurs on the vertical cliff faces and rock pin-
nacles from about 10 feet above the beach, or sea
level, to the tops of the cliffs. The vascular plants
are rooted in rock crevices, and the bryophytes
most commonly occur under shallow shelves of the
cliff faces (figs. 6,7).
Characteristic species: PotentiUa villosa, Drdba
hyperborea, and Saxifraga bracteata.
Other species: Amphidium lappomcum, Bar-
fiida cylindrica, Bartramia ithyphylla, Bryum lap-
ponicum, B. stentotrichum, Cardamine umbellata,
Cephaloziella arctica, Claytonia sibirica. Cochlea/rid
officmalis subsp. oblongifolw, Gratoneuron filici-
num, Cystopteris fragUis subsp. fragilis, Eurhyn-
chium praelongum, Lophocolea cuspidata, Luzvla
tundricola, Marchantia polymorpha^ Mielichho-
feria maorocarpa, Montia fontana subsp. fontana,
Platydictya jtmgermannioides, Plantago maritima FIGUBE 7. Rock pinnacles ("sea stacks"), Makarius Bay.
Xanthorea-Ramalina community is on and near the summit,
subsp. juncoides, Pohlia cruda^ Pottia heimii, the Potentilla-Dralta-Samfraga community is near the middle,
Rhacomitrivm fasciculare, Sagina crassicaulis, and the Eiirhynchium-Puccinellia-Caloplaca community is
Schistidium maritimum, Taraxacum trigonolo- near the base of the higher pinnacle. Grass clumps (fore-
fium, TortuZa nmcrontfolw, and Ulota phyllaritha. ground) are about 6 inches high. Photographed June 17, 1967.
cata, Potentilla villosa, Physcia caesia, P. dubia, EurhyncMwn praelongwn, Festuca rubra, Hiero-
JSchistidiwn maritimum, and Ulota phyllanttia. chloe odorata, Leptoforyum pyriforme, Ochrolechia
frigida, Plantago maritima subsp. juncoides, Ra-
d. Elymus-Ligusticum-Anemone community (1L21).
nimculus occidentalis subsp. in&idaris, Salix arctica
Soil: Lithosol, and tundra without permafrost;
type F4 ; pH, 6.4-6.6. On sloping faces of the less subsp. crassijulis, and Viola langsdorffi.
steep cliffs, and at the brink of vertical cliffs (fig. 4. EMPETRUM HEATH HABITAT
6). The soil generally is highly organic but at
places may be sandy or rocky. This community is The undulated surfaces of the old elevated marine
dominant also on large flat-topped sea stacks platforms that occur on the southeastern third of the
around the island, where Elymus commonly is island are covered with a heath composed largely of
more abundant than either Anemone or Ligusticwm. Empetrwm, with sedges, grasses, lichens, and mosses oc-
Chacteristic species: Elymus arenariiis subsp. curring in various proportions. This habitat occurs also
mollis var. mollis, Ligusticum scoticum subsp. Jvul- in low-lying valleys in the mountainous part of the
tenii, and Anemone narcissiflora subsp. villosissima. island, and near Aleut Point, at altitudes ranging from
Other species: Agrostis alaskana, Amblystegiwm about 100 to 600 feet above sea level. The plant com-
serpens, AtKyrium filiw-femina subsp. cyclosorwm, munities at places are very complex in composition and
Brywn stenotrichum, Calamagrostis nutkaensis, structure, commonly having broad ecotones and many
Conioselinum chinense, Dactylorhiza aristata, interruptions (fig. 8).
FIGURE 8. Aerial photograph of the Empetrum heath in an area of many lakes, with the Empetrurn-Carex-lichen community pre-
dominating. A wet sedge-meadow community surrounds the two small lakes in left center. At least 14 well-developed moss
mounds are shown. The trail of a tracked military vehicle is clearly visible in the upper left, although it was made at least
20 years ago. The area shown here isi on the low plateau about midway between Rifle Range Point and Crown Reefer Point.
Large moss mounds are about 25 feet in diameter. Photographed October 20, 1965, by the U.S. Air Force.
a. Empetrum-Carex-\\ohsx\. community (lC12c). Characteristic species: Empetrum nigrum subsp.

nigrum, Carex pluriflora, C. macrochaeta, Cladonia
Soil: Tundra without permafrost; type O; pH,
pacifica, and Sphaerophorus globosus.
5.9-6.0. This community is characteristic of the
Other species: Alectoria ochroleuca, Anemone
summits and slopes of the low ridges and commonly
narcissiflora subsp. villosissima, Antitrichia curti-
is hummocky and interdigitated with ecotonal
pendula, Arnica unalaschcensis, Aulacomnium
variants and interrupted by pools and lakes. At the
palustre, Brachythecium acutum, B. asperrimum,
upper altitudinal limit, the continuity of the com-
Calamagrostis nutkaensis, Calypogeia sphagnicola,
munity is broken by areas of virtually bare pebbly
Campanula lasiocarpa subsp. lasiocarpa, Carda-
ground. In the more hummocky areas, dense stands
mine umbellata, 'Carex anthoxanthea, C. dioica
of Cladoma pacifica occupy the depressions that
subsp. gynocrates, C. lyngbyaei, C. stylosa, Cas-
surround the hummocks composed of Empetrum,
siope lycopodioides, Chrysanthemum arcticum
Carex, and Sphaerophorus (fig. 9). At other places,
subsp. arcticum, Cladonia l>ellidiflora, C. coccifera,
Empetrum, several species of pleurocarpous
C. gracilis, Coptis trifolia, Comicularia divergens,
mosses, and lichens form a dense mat of vegetation
Cornus suecica, Dicranum elongatum, D. groen-
that is fairly smooth and has few surface depres-
landicum, D. howellii, Elymus arenarius subsp.
sions. Herbaceous flowering plants and Lycopo-
mollis var. mollis, Equisetum arvense, Erigeron
dium are scattered throughout this community,
peregrinus subsp. peregrinus, Eurhynchium pncce-
but they rarely form dense stands. Commonly, the
longum, Festuca l>rachyphylla, F. rubra, F. rubra
substrate is a fibrous peat, ranging in thickness
subsp. aucta, Galium trifidum subsp. columbianum,
from a few centimeters to at least 6 dm, which Geum calthifolium, Hylocomium splendens,
overlies fragmented andesite and breccia bedrock,
Hypnum dieckii, Linnaea borealis subsp. borealis,
or, at places, deposits that resemble till, colluvium, Listera cordata var. nephrophylla, Lobaria linita,
or old beach gravel. A thin (2-6 cm) layer of
Loiseleuria procumbens, Lophozia cf. groenlan-
bluish-gray to grayish-brown mineral soil gen-
dica, Luzula multiflora subsp. multiflora var.
erally occurs on these rock substrates. The organic frigida, Lycopodium annotinum subsp. annotinum,
mat and the underlying peat and muck are nearly L. clavatum subsp. clavatum* L. selago subsp.
saturated with water throughout the year, even
selago, Microlepidozia mdkinoana, Mnium glabres-
during the infrequent days in summer when the cens, Mycoltlastus alpinus, Mylia taylori, Ochro-
surface vegetation is dry and crisp.
lechia frigida, Orthocaulis binsteadii, Pedicularis
chamissonis, Peltigera canina, P. malacea, Phleum
commutatum var. americanum* Plagwthecium un-
dulatum, Platanthera chorisiana* P. convallariae-
folia, P. dilatata, Pleurozium schreberi, Pogona-
tum alpinum, Polygonum viviparum, Primula
cuneifolia subsp. saxifragifolia, Ptilium crista-
castrensis, Pyrola asarifolia var. purpurea,
Rhacomitrium lanuginosum, Rhytidiadelphus
loreus, R. triquetrus, Rubus arcticus subsp. stel-
latus, R. chamaemorus, Salix arctica subsp. crassi-
julis, S. cyclophylla, Saxifraga punctata subsp. in-
sularis, Sphagnum capillaceum, S. subsecundum,
Sphenolobus saccatulus, Thamnolia vermicularis,
Tofieldia coccinea, Trientalis europaea subsp. arc-
tica, Vaccinium vitis-idaea, Vahlodea atropur-
purea subsp. latifolia, Veronica stelleri, and Viola
langsdorffii.
b. Cladonia-Carese meadow community (1O22).
FIGURE 9. Hummocky Empetrum-Carex-lichen community in Soil: Tundra without permafrost; type O; pH,
the vicinity of Cyril Cove. The light snow cover emphasizes 5.1-6.5. This community generally occurs adjacent
the irregularity of the surface. Photographed December 2 to the Empetrum-Caresc-lichen. community, and the
1965. two communities merge in a narrow to wide eco-
tone. The chief distinctions of the Cladonia-Careso Other species: Brachythecium cisperrimum, Cal-
meadow community from the adjacent community amagrostis nutkaensis, Calypogeia trichomanis,
are its location on the more level areas or lower Cardamine wribellata, Carew antTuxwmthea, Cera-
parts of gentle slopes, the less prominent role of todon purpweus, Claytonia sibirica, Deschampsia
Empetrum, and greater abundance of Carese lyng- beringensis, Diplophyllum albicans, Drepanocla-
byaei. In general, forbs are more plentiful, and dus revolvens, Empetrum nigrum subsp. nigrwm,
the vegetation mat is wetter in this community than Eriophorwn russeolum subsp. rufescens, Festuca
in the Empetrum-Carese-lichen community (fig. 10, brachyphylla, Gymffiocolea, inflata, Jimcus trigtu-
11). mis subsp. albescens, Lophozia cf. groenlandica,
Characteristic species: Cladonia pacifica, Careso Lycopodium ann^tinum subsp. annatinum, Mylia
lyngbyaei, and C. macrochaeta. taylori, Oncophorus wahlenbergii, Orthocaulis
binsteadii, Petasites frigidus, Pinguicida vulgaris
subsp. macroceras, Platanthera convallariaefolia,
P. dilatata, Pogonatwn alpinum, Scapania irrigua,
S. pcdudosa,) Trichophorum caespitosum, and Viola
langsdorffii.
c. Moss mound community (1O1).
Conical mounds covered with vegetation are
characteristic of many of the central and western
Aleutian Islands. These mounds were first reported
by Hulten (1933, p. 17-18) ; he considered them to
be bird-perch mounds caused by the deposition of
bird manure. This explanation of their origin was
accepted by the following subsequent investigators:
Hutchison (1937); Walker (1945); Bank (1951),
1956); and Fraser and Barnett (1959). These
mounds were described in more detail in a later
FIGURE 10. Well-developed moss mound in a Cladonia-Carese publication by Hulten (1960, p. 29-30).
meadow near Cyril Cove. Mound is about 5 feet high. Photo-
graphed June 10,1967. There are two basic types of conical mounds on
Amchitka Island moss mounds (fig. 10) and rock
mounds (fig. 33). If either of these mounds is used
extensively by birds for perches, the heavy manur-
ing by the birds causes a change in the vegetation
of the mound. Many of the original plant species are
killed by the manure, and coprophilous species
that can thrive in the enriched substrate occupy the
mound. Bird-perch mounds are characterized by a
dense turf of grass at the summit (figs. 12, 32).
On the low plateaus the thousands of moss
mounds are the most conspicuous features of the
landscape (figs. 8, 10). The mounds range in size
from single polsters of moss 15 to 20 cm high to con-
ical plant communities as much as 2 m high and 10
m in diameter. They are organic throughout and
originate from single polsters of vertical-growing
FIGURE 11. Ecotone of an Empetrum-Carex-llchen community
mosses. The manuring by birds plays no part in
(irregular patches of snow-covered vegetation) and a Cla- their early development. The plant community on
doniarCarex meadow, near Cyril Cove. Contrast between the the mounds is largely composed of the same species
two communities was enhanced by a light snowfall, the that grow on the adjacent heath. The substrate, 15
snow having melted on the wetter, Cladonia-Carex meadow.
Strip of snow-covered vegetation (center foreground) is about cm below the mound apex, ranges in pH from 5.1
18 inches wide. Photographed December 7,1965. to 6.5.
A well-developed moss mound has a zonal ar-

rangement of plant species that surrounds the apex
because certain combinations of species follow in
succession up the sides of the mound. The invading
plants eventually suppress the vertical-growing
mosses at the apex, thus destroying the impetus to-
ward development of the mound, and the mound
finally begins to break apart. The zonal arrange-
ment of species in a well-developed moss mound is
given below.
Apex of mound: One of the following species
ordinarily forms the dominant vegetation Rhaco-
mitrium lanuginosum, Dicrcmum elongatum, D. FIGURE 12. Organic bird-perch mound showing characteristic
groenlandicum, D. howellii, D. fuscescens, and On- columnar summit composed of Poo, arctica subsp. williamsii,
cophorus wcMenbergii. Intermixed with these near White House Cove. The grass on the sides of the mound
mosses are a few strands of leafy liverworts, includ- is mostly Calcrniagrostis nutkaensis, but a few plants of Ely-
ing Anastrophyllum sphenoloboides, Bazzania tri- mus arenarius subsp. mollis var. mollis are present. Columnar
summit is about 8 inches in diameter. Photographed June 11,
crenata, Calypogeia sphagnicola, Cephalozia bicus-
1967.
pidata, C. leucantha, Microlepidozia maJcwuxma,
Od&ntoschisma elongatum, Orthocaulis binsteadii,
Riccardia latifrons, and Sphenolobus minutus. Usu- tually the same species, if the mounds are equally
ally, a spreading mat of Antitrichia curtipendula manured. Organic bird-perch mounds are located
eventually grows over the vertical-growing moss of mostly along the coasts, but because the island is
the apex. This moss gives the summit of the mound narrow, sea birds at some inland locations have
a conspicuous yellow color and tends to restrict caused these mounds to develop. These mounds
further vertical development of the mound. originate as moss mounds that are described in 4c;
Sides of mound: Calamagrostis nutkaensis, Carex when the mounds reach a certain height (probably
macrochaeta, C. pluriflora, Cladonia amaurocraea, about 6 dm) birds may begin to use them as look-
C. arbuscula subsp. beringiana, C. bellidiflora, C. out perches. Bird manure favors the growth of
pacifica, C. scabrmscula, Coptis trifolia, Comicul- coprophilous plants and, at the same time, kills the
aria divergens, Cornus suecica, Drepanocladus un- noncoprophilous species that formerly covered the
cinatus, Eurhynchium praelongum, Gymnocolea apex and sides of the moss mound. The vertical-
inflata, Hylocomium splendens, Loiseleuria pro- growing mosses and the characteristic Antitrichia
cumbens, Mycoblastus alpinus, Mylia taylori, Par- at the summit are replaced by 'a dense cylindrical
melia omphalodes, Peltigera aphthosa, Pleurozium turf composed of a grass species. On mounds where
schreberi, Pogonatum alpinum, Polytrichum com- Dicranum elongatum forms the apex, however, the
mune, P. strictum, Ptilidium ciliare, Rhytidiadel- moss summit may persist for an unknown length of
phus loreus, R. triquetrus, Rubus arcticus subsp. time, but it becomes abnormally green.
stellatus, R. chamaemorus, Scdix arctica subsp. In time, the influence of the manure extends
crassijulis, Sphaerophorus globosus, Tofieldia coc- down the aides of the mound, killing most species
cinea, Vaccinium vitis-ida&a subsp. minus, and of mosses and lichens; these plants are replaced
Vahlodea atropurpurea subsp. latifolia. by Calamagrostis and several species of forbs.
Base of mound: Empetrum nigrum subsp. Eventually, the sides of the mound collapse, leav-
nigrum, Calamagrostis nutkaensis, and Carex ing a cylinder of grass turf 6 to 9 dm high protrud-
lyngbyaei. ing from an irregular, hummocky base (fig. 12).
d. Organic bird-perch mound community (1M2). On neighboring Rat Island, foxes were observed to
have caused virtually the same transformation of
Substrate pH, 6.4-6.7. The mounds that support
moss mounds, but the fox-mound lookouts do not
this community (fig. 12) are in ihv Empetrum heath
have cylindrical summits.
habitat and are organic throughout, in contrast to
Listed below are the species of plants found on
the bird-perch mounds formed over boulders and
these mounds.
the conical projections of bedrock that occur at
higher altitudes on Amchitka Island. The vegeta- Apex of mounds: Festuca brachyphylla, Poa-
tion of the two types of mounds is composed of vir- arctica subs. wiUiamsiip Puccinellia langeana
subsp. alasTcana, Rinodina turfacea, Cladonia
alpestris, Eurhynchium praelongum, and Dicra-
num elongatum.
Sides of mound: AntitricTiia curtipendula, Bra-
chythecium alhicans, Calamagrostis nutTcaensis,
Cladonia scabriuscula, Diplophyllum albicans,
Drepanocladus uncinatus, Elymus arenarius subsp.
mollis var. mollis, Eurhynchium praelongum,
Heracleum lanatum, Macrodiplophyllum plicatum,
Peltigera canina, Ranunculus occidentalis subsp.
insularis, Rhacomitrium lanuginosum, Rinodina
turfacea, Sagina crassicaulis, Sphaerophorus glo-
bosus, Ulota phyllantha, Xanthoria candelaria, and
some species listed in 4c that have not yet suc-
cumbed to the effects of the manure.
Base of mound: Calamagrostis nutTcaensis,
Caresc lyngbyaei, Empetrum nigrum subsp. ni-
grum, IcmadophUa ericetorum, Polypodium vul-
FIGUBE 13. iStreamside community near Cyril Cove. The stream
gare subsp. occidentals, and Psoroma hypnorum.
has been dammed by the aquatic moss FontinaMs; the small
e. Streamside oonimunity (1N21). pool above the dam is about 45 cm deep. Streptopus amplexi-
Soil: Tundra without permafrost and alluvial; folius is just beginning to grow on the top of the dam. The
vegetation on the streambanks includes Carex lyngbyaci, Her-
type O; pH, 5.0-6.5. The narrow, deeply entrenched acleum lanatum, Ranuculus occidentalis subsp. insularis, and
streams of gentle slopes and valleys support a dis- Athyrium filix-femina subsp. cyclosorum. Photographed
tinctive plant community that is especially con- June 14, 1967.
spicuous in June, when the bright green of the
streamside plants contrasts strongly with the brown
tones of the adjacent Cladonia-Carese meadows. f. Peat-bank community (lOl).
The vegetation is composed principally of tall forbs Soil: Tundra without permafrost; type O; pH,
and sedges. An understory of lesser forbs, bryo- 5.4-6.1. Numerous peat banks were formed on this
phytes, and lichens that are adapted to reduced island by excavations related to military operations
light grows on the steep stream banks (fig. 13). during World War II. The bare banks, commonly
Characteristic species: Athyrium filix-femina 1 to 2 m high, were opened to primary invasion by
var. cyclosorum, Carex lyngbyaei, Heracleum lana- plants that were adapated to a saturated organic
tum, Geum macrophyttum subsp. macrophyllum, substrate, strong winds, and full exposure to light.
and Dryopteris dttatata subsp. americana. Apparently, only bryophytes and a few lichens are
Other species: Bracliythecium asperrimum, early invaders, for after 20 years vascular plants
Bryum pollens, B. pseudotriquetrum, Calliergon have become established in only a few locations.
sarmentosum, Calypogeia trichomanis, Campylium Areas of seepage on the vertical or overhanging
stellatum, Cardamine umbellata, Cephalozia fiicus- surfaces may support conical or curtainlike growths
pidata, C. media, CTiiloscyplius pallescens, Claopo- of bryophytes (principally Cephalozia bicuspi-
dium crispifolium, Climacium dendroides, Coni- data) at the drip points (fig. 14). All the naturally
oselium chinense, ConocepJialum conicum, Cystop- formed peat banks observed had passed the pri-
teris fragilis subsp. fragilis, Dicranum howellii, mary stages of colonization, and their plant com-
Diplophyllum albicans, Eurhynchium praelongum, munities resembled those of the Empetrum heath.
Fritillaria camschatcensis, Lecanora castanea, Lu- The flowering plant Majanthemum dilatatum,
zula parviflora subsp. parviflora, Marsupella emar- however, appears to grow only on old well-vege-
ginata, Mnium glabrescens, M. pseudopunctatum, tated peat banks, never on the more nearly level
Nardia scalaris, Peltigera aphthosa, P. canina, areas of the heath.
Petasites frigidus, Platydictya jungermannioides, Characteristic species: Pogonatum alpinum,
Ranunculus occidental-is subsp. insularis, Riccardia Polytrichum commune, Cephalozia bicuspidata,
pinguis, Saxifraga punctata subsp. insularis, Sca- and Dicranella heteromalla.
pania paktdosa, 8. undulata, Sphagnum girgen- Other species: Agrostis alaskana, Anomobryum
sohnii, S. teres, and Streptopus amplexifolius. concinnatum, Brachythecium albicans, Bryum arc-
350-274 O -
m-ine umbellata, Deschampsia beringensis, and On-

cophorus woMenbergii.
Other species: Cephaloziella arctica, Chiloscy-
phus pallescens, Dicranella palustris, Dicranum
fuscescens, Diplophyllum albicans, Epilobium
glandulosum, Geum calthifolium, lonaspis epuloti-
ca, Lecidea cf. vernalis, Leptobryum cf. pyriforme,
Luzula multiflora subsp. multiflora var. frigida,
Microlepidozia mdkinoana, Mnium insigne, M.
pseudopunctatum, Nardia scalaris, Philonotis
fontana^ Rhytidiadelphus loreus, Riccardia sinuata,
Rubus arcticus subsp. stellatus, Scapania paludosa,
/Sibbaldia procumbens, Stellaria calycantha subsp.
calycantha, 8. sitchana var. bongardiana, and Ver-
onica serpyllifolia subsp. humifusa.
FIGURE 14. Peat-bank community in the Empetrum heath near
Constantine Harbor. The elongated bryophyte colonies, com-
posed principally of Ccphalozia Mcmpidata, form at drip
points on the saturated bank that is about 2.5 m high. The
colony of large mosses, shown near the upper left, is com-
posed of Polytrichum sphaerothecium. Photographed June 5,
1967.
ticum, Blasia pusilla, Bryoerythrophyllum recur-

virostrum, Calypogeia mriilleriana, Cephalozia am-
bigua, G. pleniceps, Cephaloziella arctica, Cladonia
degenerans, G. gracilis, Dichodontium pellueidum,
Dicranetta heteromalla, D. subulata, Diplophyllum
albicans, Lophozia alpestris, L. cf. groenlandica,
L. indsa, Majanthemum dilatatum, Mnium gla-
brescens, M. insigne, Nardia scalaris, Orthocaulis
FIGURE 15. Frost scar in the Empetrum heath near Cyril Cove.
floerkei, Pannaria pezizoides, Plagiothecium cf. The light snow cover had fallen before the soil had frozen;
undulatum, Plectocolea cf. obovata^ Pogonatum subsequent frost heaving is indicated by the cracks (black
contortum, P. umigerum, Pohlia annotina, P. cf. lines) in the snow. Field notebook gives indication of scale.
cruda, P. nutans, Polytrichum formosum, P. Photographed November 29,1965.
sphaerothecium, Rhacomitrium fasciculare, Sca-
pania scandica, Sphagnum, compactum, S. papillo- 5. BOG HABITAT
sum, and Toninia Idbulata. Most bogs on this island are soligenous and occur on
g. Frost-scar community (3C2). gentle to moderately steep slopes. The borders of lakes
and pools do not support bogs, probably because the
Soil: Tundra without permafrost, or lithosol;
strong winds agitate the water too violently for Sphag-
type ?2; pH 6.0-6.3 Frost scars as much as 1 m or
num, and other bog plants to grow; therefore, there are
more in diameter occur throughout the Empetrum
heath from which the organic mat has been re- no floating bogs on the island. At a few locations the
moved, and mineral soil is exposed. Frost heaving bogs have an unstable surface and may be termed "quak-
is active during the winter and is intensified by ing bogs." Bogs are more common and of larger size on
frequent cycles of freezing and thawing of the soil the low plateaus of the eastern part of the island than
in the mountains, where they are ordinarily restricted
(fig. 15). This unstable substrate of soil, gravel,
and cobbles, or at places angular bedrock frag- to small areas in drainage ways.
ments, provides a suitable niche for species unable a. /Sphagnum, bog community (1O11).
to compete with the vegetation of the unbroken Soil: Bog; type O; pH, 6.1-6.2. Water pH,
heath. « 5.9. These bogs are neither abundant nor large,
Characteristic species: Antennaria dioica, Carda- and at places they intergrade with wet sedge
meadows. Isolated colonies of Sphagnum, com- near Makarius Bay5 where an almost unbroken mat
monly S. capillaceum and /S. subsecundum, occur in of Philonotis covers an area of a hectare or more.
small water-filled depressions in the Empetrum Characteristic species: Philonotis americana and
heath. Typically, this community is best developed Parnassia kotzebuei.
in seepage channels or broader areas of gentle slope Other species: Amblystegium serpens, Carex kel-
(fig. 16), where stands composed largely of several loggii, C. lyngbyaei, Cerastium fontanum subsp.
species of Sphagnum limit the invasion of other triviale, Dicranella heteromalla, D. subulata, Epilo-
plants. bium sertulatum, Juncus triglumis subsp. albescens,
Jungermannia sphaerocarpa, Limosella aquatica,
Lophozia cf. groenlandica, Lupinus nootkatensis,
Platanthera convallariaefolia, P. dttatata, Pogo-
natum umigerum, Pottia heimii, Saxifraga punc-
tata subsp. insularis, and Veronica serpyllifolia
subsp. humifusa.
c. Scapania-Nardia-Marsupella community (1O1).
Soil: Bog; type P,; pH, 6.1-6.8. Although small
colonies of each of the characteristic species occur
in wet areas of the Empetrum heath, these three liv-
erworts commonly form a distinctive community
that occupies drainage channels and wet areas only
at altitudes above 400 feet. These species also grow
in the snow-bed community, but there they have
different vascular-plant associates. The brownish
red to bright red of the Scapania and Marsupella
and the bright green of the Nardia make the com-
FIGURE 16. Spring issuing from the margin of a large soligenous munity conspicuous, even at a distance. The satu-
bog in a broad valley near Cyril Cove. A wet sedge-meadow rated organic mat formed by this community is as
community is in the foreground, and a Sphagnum bog com- much as 20 cm thick.
munity is in the center and background of the picture. At
the time this photograph was made, water temperature in the
Characteristic species: Scapania paludosa, Nar-
spring was 3.3°C5 and under the ice of nearby pools, 0.7°C; dia scalaris, and Marsupella emarginata.
air temperature was -2.3 °C. Photographed February 10, 1968. Other species: Anthelia julacea, Aulacomnium
palustre, Brachythecium asperrimum, Dicranella
palustris, Drepanocladus revolvens, Lycopodium
Characteristic species: Sphagnum teres, IS. squar-
annotinum subsp. annotinum, L. sabinaefolium
ros-um, 8. magellanicum, IS. compactum, S. papillo-
var. sitchense, Oncophorus ^vahlenberg^^ Philo-
sum, and /S. girgensohnii.
notis fontana, and Saxifraga punctata subsp.
Other species: Aulacomnium palustre, Brachy-
insularis.
thecium asperrimum, Calypogeia sphagnicola, 6. MARSH HABITAT
Careso anthoxanthea, C. lyngbyaei, C. pluriflora,
Dicranum angustum, Drepanocladus revolvens, A marsh on this island can be distinguished from a
Eriophorum russeolum subsp. rufescens, Mnium bog during any season of the year by its abundance of
pseudopunctatum, Philonotis americana, Pingui- sedges and forbs, whereas a bog has a preponderance
cula vulgaris subsp. macroceras, Plantago mari- of bryophytes. If the substrate of a marsh is stirred
time subsp. juncoides, Riccardia latifrons, up, even in winter, it gives off a strong odor of hydrogen
Selaginella selaginoides, and Siphula ceratites. sulfide. The substrate of a bog however, is virtually
odorless. Probably chemical compounds in the bryo-
b. Philonotis-Parnassia community (1O12).
phytes of bogs inhibit the production of hydrogen
Soil: Bog; type P2 ; pH, 6.5-6.8. Water pH, 6.3-
sulfide by bacteria.
7.0. Requirement for the development of this com-
munity appears to be a saturated inorganic sub- a. Wet sedge-meadow community (1M21).
strate of very gentle slope through which water Soil: Bog and Half Bog; type O; pH, 5.9-6.2.
moves. The water is less acid than that of Water pH, «=* 6.3. This is the most abundant com-
Sphagnum bogs. The largest observed community munity in the marsh habitat and occurs typically
of this type is on sandy soil below the large spring at the borders of lakes (figs. 8, 17), but it also
FIGURE 18. Wet sedge meadow near Cyril Cove. This area has
a dense stand of Car ex lyng~byaei forming a wide border by
the stream. A Fontinalis-Ranunculus community is in the
stream. Photographed November 23,1965.
setwn, Rcmunculus occidentalis subsp. nelsoni,

Rhinanthus minor subsp. borealis, Rhytidiadel-
phus loreus. Sphagnum girgensohnii, S. papillo-
sum, /S. squarrosum, S. teres, and Viola langsdorffii.
b. Caltha-Claytonia community (1N21).
FIGURE 17. Wet sedge-meadow community at the margin of a
large lake near Cyril Cove. The projecting mats of Carex Soil: Half Bog; type P2 ; pH, 6.1-6.9. Water pH,
are as much as 2 m from base to tip and 30-60 cm wide 6.2-6.9. This community occurs most commonly on
but are being undercut and are thus destroyed by wave action. a wet inorganic substrate that has sufficient move-
A line of new sedge growth is in the center of the picture.
ment of water through it to prevent freezing in
Numerous projections of sedge mats encircle some large lakes
and give a fimbriate appearance to the shoreline, as seen in winter. An area about 2 hectares in extent is
aerial photographs. Photographed June 13, 1967. occupied by this community near Constantine
Harbor, and many smaller areas support this com-
munity where water upwells through the Empetrwm
develops on very wet slopes and beside streams in heath mat.
th&Empetru/mhesith (fig. 18). Characteristic species: CcHiha, palustris subsp.
Characteristic species: Carex lyng~byaei, O. pluri- asarifolia and Claytonia sibirica.
flora, C. anthoxanthea, C. nwcrochaeta, Erio- Other species: Careoc lyngbyaei, Epilofiium
phorum russeolum subsp. rufescens, and Juncus glandulosvm, E. sertulatum, Petasites frigidus,
triglumis subsp. albescens. RammculMs occidentalis subsp. insularis, and
Other species: Aongstroemia longipes, Aula- Rumex obtusifolius.
comniwn palustre, Claytonia sibirica, Drepano-
7. LAKE HABITAT
cladus fluitans, D. revolvens, Epilobium glandulo-
smn, Festuca rubra, Galium trifidvm, subsp. In this report, lakes are distinguished from pools
columbianum, Geum macrophyllum subsp. macro- by their being sufficiently large for wind to strongly
phyllwn, Lophozia alpestris, Mnium glabrescens, agitate the surface and waves to pound the shore, thus
Nardia scalaris, Pedicularis chamissonis, Pellia influencing the kind of community that can develop
neesiana, Philonotis americana, Plantago mcucro- in and around them (fig. 17). The ice on some large
carpa, Pleurozium schreberi, Polytrichum longi- wind-swept lakes affects the formation of the lake banks.
DE SCRIPTION OF THE ISLAND AND ITS VEGETATION 23
If a sudden lowering of the air temperature to a point plants, which are emergent or immersed bottom-rooted
below freezing occurs when gale-velocity winds are aquatics.
blowing, water is lifted from the lake surface, frozen in
a. Hippuris-Ranunculus community (1P21).
midair, and dropped on the leeward side of the lake.
A covering of rough ice then develops from the leeward Soil: Type P2 ; pH, 5.8-6.3. Water pH, 5.1-6.7. The
to the windward side of the lake until the entire lake Hippuris grows in water as much as 1 m deep; in
surface finally is frozen, with thicker ice on the leeward summer the upper part of the plant emerges from
side.The thick ice appears to exert pressure on the the water. Dense stands of this plant occur in some
vegetation mat of the bank such that this mat (largely lakes. Commonly, the Ranunculus is intertwined
unfrozen) is pushed out and up to form the elevated with the Hippuris.
rims characteristic of many lakes (fig. 19). Moreover, Characteristic species: Hippuris vulgaris and Rwn-
the suspended inorganic materials in the windblown unculus trichophyUus var. trichophyllus.
water are deposited on the banks. This process, espe- Other species: None, although a band of sedges
cially active in the more shallow lakes, tends to deepen (principally Carex lyngbyaei} may surround the
the lakes and to elevate the lake banks. lake and extend into the water.
Hamelin and Cook (1967, p. 99) evoked a different b. Isoetes-Ranunculus-LimoseUa community (1P11).
process in explaining the formation of lake ramparts:
"A rise in temperature causes the ice to expand, induces Soil: Type P2 ; pH, 6.4-6.6. Water pH,« 6.6. The
considerable stresses and crowds the edge of the shore, plants of this community commonly grow in lake
the material [gravel and cobbles, in their illustration] water 45 to 60 cm deep. Isoetes grows completely
is pushed, overriding the low shore, for many feet submerged in the deeper water, but the other char-
where it is deposited upon contraction or melting of the acteristic species grow in shallow water near the
ice." This process may also be active on Amchitka shore and may be emergent. All are tenaciously
Island. rooted in mud of the lake bottom and grow through-
Two kinds of lakes are on the low plateaus of the out the year.
island those that have a geologic origin, are relatively Characteristic species: Isoetes muricata subsp. mari-
deep, and have a bedrock or cobble bottom, and those tima, Ranunculus reptans, and Limosella aquatica.
that develop on the peneplaned area of the heath Other species: Commonly none; however, at one lake
habitat, are shallow, and have a muck or soil bottom Scapania subalpina and Sphagnum squarrosum
(Powers and others, 1960, p. 526). With but few grew as submerged aquatics. At some lakes Ranun-
exceptions, only the latter type of lake contains vascular culus hyperboreus subsp. hyperboreus grows on
mud along the shore.
8. FOOL HABITAT
Pools usually are formed in depressions of the vege-

tation and peat mantle on the Empetrum heath, al-
though some are in depressions of bedrock or colluvium,
and some are formed by bryophyte dams (fig. 13). Be-
cause the pools have a periodic fluctuation of water
level, emergent aquatic plants most commonly grow in
them and in the mud along the pool banks.
;f *V
%;#)
.*-4£f.f-
> '^.
" jfe "V
*. Small pools appear to become enlarged by the
process of "swamping," and when they reach a diameter
j .««:
t^, i
^ of about 3 m (if in an exposed location) wind becomes
*. active in "digging" them deeper. The agitated water
holds much silt or humus from the pool bottom in
FIGURE 19. Ice ridge on the leeward bank of a large lake near suspension, and the wind blows this water from the pool
Cyril Cove. The water near the distant shore is not frozen. to the adjacent heath. Deposits of silt on the leeward
Strong winds lift this water, which is frozen in midair and side of pools was observed to be sufficiently thick to
then dropped on the leeward bank and the lake surface.
Top of the tundra bank is about 6 feet above water level. obscure the green color of the Empetrum mat at a dis-
Photographed December 3,1965. tance of 3 m from the pool.
a. Juncus-Eriophorum community (1M22).

Soil: Tundra without permafrost and Alluvial;
type P2 ; pH, 5.0-6.4. Water pH, 4.8-6.3. This
community commonly occurs in shallow pools on
the Empetrum heath (fig. 20) and less commonly
in roadside pools and very wet areas of stream
alluvium. The vegetation along the pool shores is
included here because it is generally inundated
after heavy rains or snow melt.
Characteristic species: Juncus triglumis subsp.
aTbescens and Eriophorum rus s eolum subsp.
rufescens.
Other species: Alopecurus aequalis, Aulacom-
nium palustre, Campylium stellatum, Oarex kel-
loggii, Chiloscyphus pdttescens, Deschampsia
caespitosa subsp. orientalis, Diplophyllum albi-
cans, Drepanocladus revolvens, Juncus ensifolius,
Marchantia polymorpha, MarsupeUa emarginata, FIGURE 21. Bottom of a dry pool near Constantine Harbor
Mnium gldbrescens, Nardia scalaris, Parnassia bearing a dense growth of Subularia aquatica that was
kotzebuei, Pinguicula vulgaris subsp. macroceras, flowering abundantly. Callitriche anceps grows in similar
shallow rock-bottomed pools. Utility pole (background) pro-
Pogonatum alpinum, Ranunculus tricTiophyllus vides scale. Photographed June 17, 1967.
var. trichophyllus, Rhacomitrium aciculare, Ric-
cardia pinguis, /Scapania paludosa, &. undulata,
Selaginella selaginoides, /Sparyanium hyper- Characteristic species: /Subularia aquatica and
fioreum. Veronica americana, and V. serpyllifolia Callitriche anoeps.
subsp. humifusa. Other species: Alopecurus aequalis, Bryum sp.,
Carex kelloggii, Cladonia bettidiflora, and Ran-
b. /Subularia-Gallitriche community (1P1).
unculus reptans.
Soil: Lithosol; type P2 ; pH, ^ 6.6. Water pH,
6.9-7.2. This community occurs in shallow seasonal c. Siphula-Scapania community (1O21).
pools that have bedrock bottoms. The two char- Soil: Tundra without permafrost and Lithosol;
acteristic plants can grow as submersed or emer- type P2 ; pH 6.0-6.4. Water pH, 6.3-6.6. Common
gent aquatics or can persist on damp soil if the in small (60-120 cm in diameter) pools on the Em>-
pools dry up (fig. 21). petrum heath, but most extensive in shallow bedrock
pools of larger size. /Scapania forms a, dense dark
mat on the pool bottom, generally in 5 to 15 cm of
water, and Siphula grows as an emergent aquatic
for much of the year (fig. 22). Both plants can
thrive, however, if the pool dries up. Siphula is the
most hydric lichen on the island and is the only one
observed to grow completely submersed.
Characteristics species: Siphula ceratites and
Scapania paludosa.
Other species: Anthelia julacea, Cladonia
amaurocraea, Isoetes sp., Lophozia incisa, and
Sphagnum compactum.
9. STREAM HABITAT
Most streams are narrow, swift, and short; and the

water, generally clear and colorless, has percolated
FIGURE 20. Small pool supporting a Junctts-Eriophorum com-
munity, surrounded by a Cladonia-Carex meadow, near Cyril through the vegetation mat. The streams flow through-
Cove. Sedges in the pool extend about 1 foot above water sur- out the year at a, fairly steady rate, even under an ice
face. Photographed June 17, 1967. covering, except for periods of greatly increased flow
filled with snowdrifts in winter and are a hazard to foot

travelers. The streams appear to have been entrenched
by the growth and deposition of vegetation at their mar-
gins ; thus they differ from "haggs" of the British moors,
defined by Pearsall (1950, p. 249) as erosion channels in
peat. Streams draining broad valleys generally have al-
luvial fans at their mouths that are bordered by sedge
marshes (fig. 18), whereas those draining the more
nearly level plateaus descend over the sea cliffs in a
series of rapids.
a. Fontinalis-Ranunculus community (1P11).
Soil: Type I2 ; pH, 6.5-6.9. Water pH, 6.8-7.2.
Occurs commonly in rapid, entrenched streams
where the light intensity is low. Fontinalis grows
attached to bedrock or stones in water as much as
1 m deep and develops fronds as much as 60 cm
long. Brachythecium and Hygrohypnum also grow
FIGURE 22. Vertical view of a Siphula-Scapania community in attached to rocks, but they form dense mats in shal-
a shallow rock-bottomed pool near Crown Reefer Point. Dark low water, especially in cascades. These mosses are
colonies of Scapania can be seen on pebbles in the water and shown forming a dam in figure 13. The Ranunculus
at the bases of the Siphula thalli. White thalli are about 2
inches long. Photographed February 18, 1968. is rooted in soil and produces underwater flowers
and fruits throughout the winter.
Characteristic species: Fontinalis neomexicana
following heavy rainfall and snow melt. They com- and Ranunculus trichophyllus var. trichophyllus.
monly are deeply entrenched in the vegetation and peat Other species: Brachythecium asperrmwm, Bry-
mantle (fig. 23) and have a bedrock, gravel, or colluvium oerythrophyllum recurvirostnmi^ Chiloscyphus
bottom. At places the vegetation mat has closed over the pallescens, Hygrohypnum bestii, H. ochraceum,
stream trenches and f ormed "bridges" that may not sup- Scapania irrigua, and S. undulata. Montia fontana
port the weight of a man. The trenches are periodically subsp. fontana grows in the shallow water of cas-
cades but not in deeply entrenched streams. Shallow
parts of the stream margins, if not shaded by high
banks, support the emergent aquatics Mimulus gut-
tatus, Veronica americana, and Carex lyngbyaei.
10. RUDEEAL HABITAT
Soils of this habitat are classified only by type. Many

substrates have the surface qualities of a lithosol or
regosol, and some substrates are not soil. The extensive
disturbance of the ground surfaces by military opera-
tions and by recent construction on the island has opened
many new areas for colonization by plants that are
adapted to a mineral soil and that can withstand the
effects of microclimatic extremes. The species of the
ruderal communities are, with very few exceptions, na-
tive plants that also grow in other habitats on the island
but that commonly grow more vigorously in the areas
which have been disturbed by man. For example, the
largest plants of Lupinus nootkatensis on the island
FIGURE 23. Deeply entrenched stream in the Empetrum heath
grow along roadsides, although smaller plants occur in
near Chitka Point. The narrow stream flows on a bedrock bot- many habitats. Loiseleuria procumbens commonly is a
tom between nearly vertical banks, 1 to 2 m high, of peat and depauperate plant in the Empetrum-Carex-lichen com-
vegetation. Photographed November 22, 1965. munity ; yet a plant that grew in the center of an unused
gravel road had, in about 20 years, formed a dense pros- Hylocomium splendens, Lusula multiflora subsp.
trate shrub 30 cm in diameter. multi-flora var. frigida, L. parmflora subsp. parvi-
The communities of this habitat described below are flora, Nardia scalaris, Pogonatum umigerum,
not so clearly .defined and discrete at all places as are PoMia nutans, Primula cuneifolia subsp. saxifragi-
some of the more "natural" communities on the island and Rhacomitrium ericoides.
probably because many of the component species of the
b. Lupinus- Arnica community (3C2).
ruderal communities have a wide ecological amplitude.
There is, nevertheless, a tendency toward grouping of Soil: Type I2 ; pH, 6.7-7.0. The community is
certain species in niches that may be only slightly sep- common along many roadsides and on disturbed
arated spatially. areas; it is very showy in midsummer because of
the large size of the plants and the profusion of
a. Calamagrostis community (1M21). flowers on the two characteristic species (fig. 25).
Soil: Type I2 ; pH, «5.9. In the construction of
roads during World War II, the organic mantle and
some of the mineral soil below it were pushed aside
to form banks 1/2-'L m high that paralleled the roads.
These banks are covered with an almost unbroken
turf of tall grass intermixed with very few forbs.
After about 20 years of dominance by the grass,
there is now some invasion on the banks of the low
plateaus by Empetrum, and at a few places the
banks are somewhat similar in appearance to the
Empetrum-Caresc-lichen community. At altitudes
of about 400 to 600 feet, wind has severely eroded
the soil banks so that only fragments of the banks
remain. The fragments are held in place only so
long 'as the turf is sufficiently strong to resist wind
erosion (fig. 24) .
Characteristic species: Calamagrostis nutkaensis.
Other species: Antennaria dioica, Bryum FIGTJEE 25. Lupinus nootJcatensis and Deschampsia caespitosa
stenotrichum, Caresc macrochaeta, Cephalozia bi- subsp. orientalis growing on the graveled shoulder of an un-
cuspidata, Ceratodon purpureus, Dicranella subu- used road near Constantine Harbor. Lupine flower spikes are
lata, Ditrichum heteromallum, Gentiana amarella about 8 inches high. Photographed June 15, 1967.
subsp. acuta var. plebeja, Heracleum lanatum.
These plants grow in the loose gravel and mineral
soil of road shoulders, relatively dry ditches, and
similar sites. Other species occupy the wetter ditches
and lower parts of roadside banks. The substrate
is not greatly compacted and is subjected to strong
frost heaving in winter. Heavy rains severely erode
the surfaces; therefore, bryophytes and lichens are
rare, but large colonies of the moss Philonotis occur
at some places in the wet roadside ditches.
Characteristic species : Lupinus nootkatensis and
Arnica unalaschcensis.
Other species : Achillea borealis, Anaphalis mar-
garitacea, Angelica lucida, Antennaria dioica^
Cardamine umbellata, Cerastium fontanum subsp.
triviale, Deschampsia caespitosa subsp. orientalis^
FIGURE 24 Fragments of a roadside bank severely eroded by Epilobium sertulatum, Gentiana aleutica^ Mar-
wind. This remnant of a bank, held in place by a dense turf
of Calamagrostis nutkaensis, is near Sea Otter Point. chantia polymorpha, Mnium glabrescens, Dactylo-
Bank, including turf, is about 4 feet high. Photographed rhiza aristata, Philonotis americana, Polygonum
November 22, 1965. viviparum, and Potentilla egedii subsp. grandis.
c. Equisetum-Sagina-Ceratodon community (2G2).

Soil: Type I2 ; pH, 5.8-6.5. The characteristic
substrate of this community has a hard, very com-
pact, often stony surface that resists frost heaving.
This community occurs on unused well-compacted
roads, graveled parking lots and service areas, frac-
tured concrete slabs, and cracked macadam run-
ways. Some species are rooted in fine-gravel ac-
cumulations 2 cm or less thick or unbroken concrete
or macadam surfaces. Many old gravel-road sur-
faces have an almost unbroken growth of Equise-
tum. Sagina occidentalis forms dense rosettes
rooted in pavement cracks or very compact road
centers. Rhinanthus, Luzula, Hieracium, and Ly-
copodium occur most commonly on well-packed
stony animal trails. FIGURE 26. Discontinuous heath habitat, with scattered
patches of the Saliso-Empetrum community. Large mound
Characteristic species: Equisetum arvense, (center) is a densely vegetated bedrock outcrop. The area is
Sagina occidentalis, and Ceratodon purpureus. between Sea Otter Point and Chitka Point at an altitude of
Other species: Antennaria dioica, Braehythec- about 400 feet. Bedrock outcrop is about 15 feet wide. Photo-
ium acutum, B. albicans, B. asperrimum, Bryum graphed November 22,1965.
stenotrichum, Cladonia pyseidata, Drepanocladus
uncinatus, Empetrum nigrum subsp. nigrum, Eu- Island, the boundary between these two communities
rhynchium praelongum, Hieracium gratiile var. is determined more by the degree of drainage than by
alaskanum, Lecidea macrocarpa, Loiseleuria pro- wind action. The heath at many places extends almost
cumbens, Luzula multi-flora subsp. kobayasii, L. to the lowest points of sheltered valleys, if drainage is
parviflora subsp. parviflora, Lycopodium selago adequate. Conversely, throughout the highest levels of
subsp. selago, Nardia scalaris, Ochrolechia frigida, the low plateaus, sedge meadows occur everywhere that
Placopsis gelida, Platanthera chorisiana, Poa al- drainage is poor. The streamside community and the
pina, P. turneri, Polytrichum formosum, Rhacomi- alpine meadow community are the ones that most
trium ericoides, Rhinanthus minor subsp. borealis, clearly require shelter from wind.
Sagina intermedia, Saxifraga punctata subsp. The striking boundary on Amchitka Island that is re-
insularis, Sibbaldia procumbens, Stellaria caly- lated to wind is the ecotone between the Empetrum
cantha subsp. calycantha, Stereocaulon inter- heath and the alpine region; it is about 400 to 600 feet
medium, S. tomentosum var. alpestre, and Veronica above sea level. At this altitude, wind erosion probably
serpyllifolia subsp. humifusa. is more active than frost action in determining which
plant communities survive (fig. 24).
11. DISCONTINUOUS HEATH HABITAT The strips of vegetation ordinarily are eroded on the
windward side until the roots and subterranean stems
A distinct change in habitats occurs at an altitude of of the dwarf shrubs are exposed and finally die, but on
about 400 feet, where the extensive Empetrum heath the leeward side, new plant growth slowly extends the
of the low plateaus becomes fragmented into small mat outward. Two lichen species, Ochrolechia frigida
strips and patches that alternate with almost bare and Mycoblastus alpinus, are associated with the dead
ground (fig. 26). This disruption of the heath mat margins of the strips; however, it was not determined
probably is caused mostly by wind erosion, but frost whether the lichen growth causes, or is the result of,
heaving doubtless accelerates the process, once the vege- death of the vegetation at the edge of the strips. The
tation cover is broken. Hulten (1960, p. 40-41) wrote, vegetation mats, therefore, appear to "move," in time,
"It is quite clear that the mosaic of plant communities with the wind, but this supposition has not been proved
in the Aleutians is to a large extent regulated by the by actual measurements. The concept of patches of veg-
wind." He noted a very sharp change in vegetation at etation "moving" in this manner was described by Watt
levels where the full force of the wind was felt, which (1947) and by Pearsall (1950, p. 83) as observed in
he reported to be the boundary that separates the British moorlands, and the floristic composition and
meadows from the heath. He referred to this boundary successional changes in these mats were given by Met-
as the "wind plane." On the low plateaus of Amchitka calfe(1950,p.53).
350-274 O - 69 - 5
There is no one direction of prevailing winds on era aphthosa, Polygonum viviparum, Rhacomi-
Amchitka Island as a whole. In the mountains and high trium fasciculare, R. lanuginosum, Sphaerophorus
plateaus, topography exerts a controlling effect on pre- globosus, and Thamnolia vermicularis.
vailing wind directions at specific locations. On slopes,
b. Andreaea-Schistidium-Rhacomitrium community
the strongest winds commonly are upslope, regardless of
(3C1).
compass direction, and the vegetation mats tend to
erode on the lower side and advance on the upper side. Soil: Lithosol; type I; pH, 5.8-6.0. Although
On more nearly level areas the local topographic fea- the area between the heath mats that is covered
tures appear to determine the direction of this "move- with rock fragments and soil may appear to be
ment." More careful observations than have yet been barren, small moss polsters grow on many rocks
made and measurements throughout the year are neces- and in the mineral soil (fig. 26). Andreaea gen-
sary if these processes are to be defined more accurately. erally is attached to small rocks, but at places it
The substrate between and under the vegetation grows on the soil, and the polsters are undercut
patches is composed of volcanic rock fragments in a by wind erosion (Shacklette, 1966, p. 349).
matrix of fine mineral "soil" and is subjected to moder- Schistidium and Rhacomitrium ordinarily grow on
ate frost heaving, but patterned ground generally is not rock fragments, and frost heaving and thawing
formed. This substrate between vegetation patches is only naise and lower the rocks without disrupting
largely devoid of vascular plants, although some scat- the moss polsters.
tered bryophyte colonies occur. Bedrock outcrops in this Characteristic species: Andreaea rupestris, Schis-
area are discussed separately in this report. tidium apocarpum, and Rhacomitrium, lanu-
ginosum.
a. Salisc-Empetrum community (lC21a). Other species: A few colonies of Brachythecium
Soil: Tundra without permafrost and Lithosol; albicans, Conostomum tetragonum, and Drepano-
type F3 ; pH, 6.0-6.1. The fabric of the vegetation cladus uncinatus and rosettes of Draba aleutica and
patches is composed largely of interwoven stems Cerastium aleuticum, which characteristically oc-
and branches of Salim and Empetrum, which hold cur on solifluction terraces, are widely scattered
in place a thin organic soil that supports a few through this community.
herbs, lichens, and mosses. Some of the woody
stems of Salix arctica subsp. crassijulis are almost 12. SOLIFLUCTION HABITAT
1 cm in diameter and appear to be many years old.
The shrubs form roots throughout the length of At altitudes between 600 and 1,100 feet, the effects of
their buried stems; therefore, they may continue to strong frost action are conspicuous in all but the most
grow at the stem tips while dying at the basal end sheltered locations (figs. 27, 28). The land surface is
of the stems. composed largely of scree slopes and colluvial boulder
In the more (sheltered locations of the discontin- fields in which the surface rocks are arranged in ter-
ous heath, areas between the heath patches develop races, rock stripes, <and weakly developed boulder poly-
a conspicuous growth of lichens. Species of Clado- gons. Solifluction terraces and rock stripes occur in
nia grow on the summits and sides of the small pebbly ground and are presently active in reshaping the
hummocks, and dense stands of Thamnolia are ground surface, but observations of the weak boulder
present in the depressed areas between the polygons were inconclusive in determining whether the
hummocks. process of rearrangement was currently active, or
. Characteristic species: Salix arctica subsp. crassi- whether the polygons are relics of frost iaction of former
julis, 8. rotundifolia, and Empetrum nigrum subsp. times. High-altitude rock outcrops and sheltered valleys
nigrum. are discussed separately in this report.
Other species: Alectoria cf. irvingii, A. ochroleu- a. Fell-field community (3B1).
ca, Campanula lasiocarpa subsp. lasiocarpa, Cassi-
ope lycopodioides, Cerastium fischerianum, Clado- Soil: Lithosol; type I; pH, 6.0-7.0. This com-
nia alpestris, C. pacifica, Dicranum howellii, munity is most extensive on the high plateaus of
Epilobium behringianum, Gymnamitrion corallioi- the western part of Amchitka Island (fig. 1), but
des, Hieracium triste, Loiseleuria procumbens, smaller areas occur throughout the mountains.
Luzula arcuata subsp. unalascheensis, L. tundri- Lithosol development is very slight, and many rock
cola, Lycopodium alpinum, L. selago subsp. selago, surfaces have only a partial covering of lichens
Mycoblastus alpinus, Ochrolechm frigida, Peltig- (fig. 29).
* **.
-*: .*
Ste
FIGURE 27. High-altitude land surfaces that are strongly influenced by solifluction processes, between Chitka Cove and Andesite
Point. A light snow cover emphasizes the surface features. Slopes are covered with an almost continuous vegetation mat;
they develop a microrelief in which the ridges and furrows parallel the direction of slope. (Compare with fig. 28.) Photo-
graphed February 7,1968.
Small patches and fragments of a heath community sphagnicola, Campanula lasiocarpa subsp. lasio-
are scattered through some boulder fields; these carpa, Carese circinnata, Cephalozia cf. bicuspidata,
are more common on the steep slopes, where the C. pleniceps, Cetraria nigricans, Conostomum tetra-
rare Rhododendron camtsckaticwin subsp. camt- gonwn, Diapensia lapponica subsp. obovata, Dicra-
schaticwn is a component of the vegetation patches. num JioweUii) Diplophyllum albicans, D. taxifoli-
On the nearly level high plateaus, Lupinus is the um, Drepanocladus revolvens, D. uncinatus, Hypo-
principal initiator of vegetation mats. Single plants gymnia subobscura, Luzula tundricola, Ochrolechia
become established between the boulders on the frigida, Parmelia omphalodes, Pohlia nutans, Rha-
almost barren plateaus, and mosses and lichens grow cowdtrium brevipes, R. ericoides, R. lamiginosum,
in the loose debris and soil that accumulates in the Rhododendron camtschaticum subsp. camtschati-
shelter of these plants. Vegetation mats, as much cum, Salix arctica subsp. crassijuZis, Scapania
as 10 dm in diameter, that contain a significant scandica, Sibbaldia procumbens, /Sphaerophorus
number of heath plants were observed to have de- globosus, Vmbilicaria proboscidea, Vaccinium
veloped around large and obviously very old lupine uliginosum subsp. mwropTiyllum, and Veronica
plants. grandiflora.
Characteristic species: Salioo rotundifolia, S. cy-
b. Solifluction terrace community (3C13).
clophylla, Cassiope lycopodioides, Empetrum ni-
grum subsp. nigrum, and Lupinus nootkatensis. Soil: Lithosol; type Ii; pH, 6.6-6.7. The'ground
Other species: Alectoria ochroleuca, A. pubescens, surface occupied by this community is composed of
Andreaea rupestris, AntitricMa curtipendula, Arc- pebbles, generally not more than 3 cm in diameter,
toa fulvella, Bryum stenotrichum, Calypogeia in a matrix of very loose wet mineral soil. Frost
FIGURE 30. Solifluction terraces on the high plateau near Rim

Point. The terraces support depauperate heath mats, and the
stony ground between the terraces bears a scattered cover of
FIGURE 28. Rock stripes that developed on a moderate slope Lusula tundricola and a few other plants. An animal trail
between Ohitka Cove and Andesite Point. A light drifted snow leads from the foreground to the helicopter. Photographed
cover emphasizes the surface features. Distance between the June 11, 1967.
summits of adjacent ridges is about 30 cm. Ridges are perpen-
dicular to the direction of slope. (Compare with fig. 27.)
Photographed February 7,1968. Characteristic species: Potentilla hyparctica,
Koenigia islandica, Cerastium aleuticum, Draba
aleutica, Juncus biglumis, and Conostomum
tetragonum.
Other species: Andreaea rupestris, Antitrichia
curtipendula, Brachythecium aTbicans, Ca/rdamine
bellidifolia, Cassiope lycopodioides, Cladonia al-
pestris, Claytonia arctica, Dicranum howellii,
Diapensia lapponica subsp. obovata, Diplophyl-
htm albicans, Drepanocladus uncinatus, Herberta
adunca, Hypnum plicatulm, Jungermannia atro-
mrenSj Lu&ula tundricola, Marsupella emarginata,
Poa lanata var. vivipara, Ranunculus occidentalis
subsp. insularis, Rhacomitrium fasciculare, R. lanu-
ginosum, Sagina intermedia, Salix cyclophylla,
Saxifraga punctata subsp. insularis, /S. foliolosa
var. foliolosa^ Sphagnum compactum, Stellaria
FIGURE 29. Fell-field at the summit of a mountain near Ohitka ruscifolia subsp. aleutica, Stereocaulon vesuvia-
Cove. Very little soil forms at this site; vegetation is sparse, num, and Tritomaria quinquedentata.
and strong winds remove most of the fine rock particles.
Photographed February 7, 1968.
13. ALPINE MEADOW HABITAT
action has arranged these materials into rock stripes In the narrow valleys in the mountains, wind velocity
(fig. 28) and terraces (fig. 30) that are perpendicu- is greatly reduced, and in some places snow accumula-
lar to the direction of slope. The brinks of the low tions may persist until mid-June. The valleys afford
terraces may have a weakly developed strip of heath protection from strong frost action, desiccating winds,
plants, but the pebbly ground of rock stripes sup- and severely cold weather and thereby permit growth
ports only single plants or polsters spaced widely of species that are not found elsewhere on the island.
apart. Lichens can scarcely become established on The wettest parts, of the valleys, where snow melt water
the most unstable surfaces, and only one moss, saturates the ground, and the margins of small streams
Conostomum tetragonwn, is well adapted to this that drain the slopes and valleys (fig. 31) support snow-
substrate. bed communities. The more mesic valley walls, slopes,
pella emarginata, Mylia anomala, Nardia com-
pressa, N. scalaris, Oligotrichum hercynicum, O.
parallelum, Osoyria digyna, Pellia neesiana, Pri-
mula cuneifolia subsp. saxifragifolia, Radula poly-
clada, Rhytidiadelphus sguarrosus, Riccardia pin-
guis, Sparganium hyperboreum, Sphagnum com-
pactum, S. papillosum, S. squarrosum, Takakia
ceratophylla, and Veronica serpyllifolia subsp.
humifusa.
b. Alpine meadow community (1JST21).
Soil: Tundra without permafrost or Half Bog;
FIGURE 31. Abrupt transition from fell-field (lower right) to type F5 ; pH, 6.2-6.4. Although all vascular plants
discontinuous heath at Chitka Cove. The streams (fore- of this community (except Pyrola minor} were also
ground) are bordered with snow-bed communities; a large found in the Empetrum heath, the distinctive fea-
wet sedge meadow surrounds the lake; and a moss mound
marks the narrow Empetrum heath above the sea cliffs. tures of this community are in the predominance of
Photographed August 16,1966. large-flowered forbs, the sparsity of Empetrum
and other shrubs, and the lesser importance of
and streamside terraces bear a profusion of showy lichens, sedges, grasses, and bryophytes in the
flowering plants, ferns, and mosses that characterize structure of the community. This community is
the alpine meadows. Lichens are not abundant in this limited to the higher altitudes, generally above 600
habitat. The ecotone between the alpine meadow and feet, in well-sheltered valleys; at lower altitudes
the solifluction terraces, or boulder fields, is at places the Empetrum, heath predominates in the valleys.
very narrow; for example, the distance (measured up Characteristic species: Achillea borealis, Arnica
the steep valley sides) from the stream bed, through unalaschcensis, Chrysanthemum arcticum subsp.
the alpine meadow, to the almost bare terraces and arcticum, Claytonia sibirica, Geum calthifolium,
rock fields may be as little as 10 m. and Ranunculus occidentalis.
Other species: Carex pluriflora, Cephalozia am-
a. Snow-bed community (1O12). bigua, C. bicuspidata, Cetraria ericetorum, Cla-
Soil: Type P2 ; pH, 5.8-6.2. Water pH, 5.4-6.6. donia rangiferina, Cornus suecica, Drepanocladus
The substrate of this community is saturated with revolvens, D. uncinatus, Empetrum nigrum subsp.
water, and large patches of the gray liverwort nigrum, Erigeron peregrinus, Listera cordata var.
Anthelia make the community conspicuous. This nephrophylla, Lophozia alpestris, L. opacifolia,
liverwort is characteristic of snow-bed communi- Lupinus nootkatensis, Nardia geoscyphus, N. scal-
ties in many parts of the northern hemisphere. The aris, Plagiothecium, undulatum, Plantago maritima
yellow-flowered Sataifraga hirculus grows in the subsp. juncoides, Platanthera convallariaefolia, P.
liverwort mats with the diminutive Hippuris mon- dilatata, Pleurozium schreberi, Polygonum vim-
tana and the rare shrub Geum pentapetalum. Col- parum, Pyrola minor, Rhytidiadelphus loreus,
onies of Sphagnum and other hydric bryophytes Sphaerophorus globosus, Sphagnum lindbergii,
complete the mosaic of the snow-bed community, S. magellanicum, Thelypteris limbosperma, T.
which extends along some of the small streams that phegopteris, and Vahlodea atropurpurea subsp.
flow through the discontinuous heath habitat (fig. latifolia.
31). Many species listed below were found only in
14. INLAND BEDROCK HABITAT
this community.
Characteristic species: Anthelia julacea, Sca- Bedrock outcrops at inland locations throughout
pania paludosa, Saxifraga hirculus, and Leptar- the island support plant communities different from
rhena pyrolifolia. those of the sea cliffs. The inland outcrops are remnants
Other species: Andreaea rupestris, Carex bige- of an earlier landform and are resistant to erosion. At
lowii, C. physocarpa, Cephalozia ambigua, C. bicus- places they project only slightly above the general
pidata, Dicranella palustris, Diplophyllum ground level; elsewhere they form irregular masses or
albicans, Drepanocladus revolvens, Geum penta- symmetrical cones that are conspicuous landmarks
petalum, Gymnocolea inflata, Hippuris montana, (figs. 32, 33). The plant communities of the outcrops
Lophocolea cuspidata, Lophozia alpestris, Marsu- are low growing and do not completely cover the rock
Other species: Agrostis borealis, Alectoria ochro-

leuca, A. pubescens, Andreaea rupestris, Angelica
lucida, Antennaria dioica, Antitrichia curtipen-
dula, Bacidea sp., Campamda chamissonis, C.
lasiocarpa subsp. lasiocarpa, Oaresa macrochaeta,
Oeratodon purpureug, Cetraria cucullata, Chrtf-
santhemu/m arcticum subsp. arcticum, Oornicularia
divergens, Dicranum kowellii, Drepanocladus un-
cinatus, Empetrum nigrum subsp. nigrwn, Festuca
bracJiypJiylla^ Gentiana aleutica, Heracleum lana-
tum, lonaspis epulotica var. arctica, Letidea
aleutica, L. macrocarpa, Loiseleuria procumbens,
FIGURE 32. Low-altitude rock outcrop that is used as a bird Lophozia cf. wenzelii, Luzula arcuata subsp. un-
perch, near Banjo Point. The thick turf at the summit is
composed of Festuca firachyphylla. Photographed August 17,
alaschcensis, Lycopodiwn sabinaefolium var.
1966. sitchense, Lycopodium selago subsp. selago, Mnium
glabrescens, Nardia scalaris, Ochrolechia frigida,
Pamnelia saxatilis, P. sulcata, Peltigera aphthosa,
Pertusaria coriacea, Placopsis gelida, Platanthera
chorisiana, Poa arctica subsp. williamsii, Pogo-
natum urnigerwn, Polytrichum junipervrwm,
Primula cu/neifolia subsp. saxifragifolia, Ramalina
almguistii, Rhacomitrium fasciculare^ R. lanugino-
sum, Rhizocarpon Tiochstetteri, JSalios arctica subsp.
crassijulis, S. cyclopJiylla, Thamnolia vermicularis,
Umbilicaria cylindrica, U. proboscidea. Viola
langsdorffii, and XantJioria candelaria.
b. High-altitude bedrock community (301).
Soil: Weakly developed Lithosol; type I; pH,
FIGURE 33. High-altitude bedrock outcrop near Chitka Cove. 6.4^6.6. This community contains few vascular
The fell-field (foreground) has patches of heath plants on plants but many cryptogams; a large part of the
the unstable rocky substrate. Although much of this conical rock surfaces, however, is not vegetated (fig. 34).
outcrop is covered with talus, the loose rocks are not sub-
jected to strong frost action because of the underlying bed- Mosses and lichens can colonize the relatively stable
rock. Outcrop is about 40 feet wide at the base. Photographed rocks of outcrops, whereas the rocks of adjacent
August 16,1966.
surfaces, unless fertilized by birds. The species compo-

sition of the communities appears to be controlled
largely by factors associated with altitude; accord-
ingly, two communities are recognized.
a. Low-altitude bedrock communities (1N21).
Soil: Weakly developed Lithosol; type I2 ; pH,
6.0-6.6. Sites of these communities range in alti-
tude from about 150 to 600 feet above sea level; out-
crops at lower altitudes are on the seacoasts. Some
conical outcrops are used by birds as perches and
are recognizable by the thick turf of grass at the
summits and sides of the mounds (fig. 32). FIGURE 34. Rock outcrops on a mountain summit about 1 mile
west of Chitka Cove. Strong winds and almost constant fog
Characteristic species: Veronica stelleri, Cas- characterize the summits. Vegetation consists of one species
siope lycopodioides, Tofieldia coccmea, and Salix of Carex, one of Affrostis, and several species of lichens and
rotv/ndifolia. mosses. Photographed July 29,1967, by J. A. Erdman.
solifluction terraces and fell-fields are so unstable, man or by ocean currents. Military and other oper-
due to frost action, that fewer of these plants can ations on Amchitka Island have resulted in the im-
grow on them. The conical outcrops at high alti- portation of much wood for buildings, utility poles,
tudes (fig. 33) are used very little as perches by and other uses. The plants which grow on this
birds. At high altitudes, sea birds and ptarmigans wood are as follows: Baeomyces placophyllus,
do not congregate, and raptorial and passerine birds Brachyihecium acutmn,, Ceratodon pwpureus,
are not abundant; therefore, coprophilous species Cetraria ciliaris, Drepanocladus wtcinatus,
of plants are not common on high-altitude outcrops. Eurhynchium praelongum, Hypogynvrda entero-
Characteristic species: Carex drcinnata, Um- morpha, H. pkysodes, Lecanora attophana, Lecidea
bilicaria proboscidea, and Agrostis borealis. cf. symmicta, L. vernaLis (on tar paper over a
Other species: Alectoria cf. irvingii, A. nigricans, wooden roof), Mnium glabrescens, M. pseudopunc-
Andreaea rupestris, Anthelia julacea, Cephdlo&ia tatum, Parmelia saxatilis, P. sidcata, Parmeliop-
leuoantha^ Cerastium aleuticum, Cetraria erice- sis ambigiM, and Pertusaria sp.
torum, Cladonia bettidiflora, Conostomum tetra-
c. Plantings.
gonum, Cornicularia divergens, Diplophyllum
albicans, Empetrum nigrum subsp. nigrwn, Fes- Several Picea sitchensis trees planted in front of
tuca brachyphylla, Gymnomitrion corallioides, buildings during World War II are still living, are
Haematomma ventosum subsp. lapponicum, Hyp- about 1 m high, and had about 3 cm of new growth
num cupressiforme, Lecidea armeniaca,, L. elegan- in 1967 (fig. 35). Similar plantings of this species
twr, L. flavocaerulescens, L. macrocarpa, Lophozia were observed to have been made on Shemya Is-
cf. groenlandica, L. incisa, Luzula tundricola, My- land, and four of the trees were still alive in 1967.
coblastus alpinus, Ochrolechia frigida, Orthocaulis On Amchitka Island the window boxes of a mili-
binsteadii, Parmelia alpicola, P. omphalodes, Pertu- tary building were planted with Anemone narcis-
saria hultenii, Platysmatia lacwwsa, Pogonatuan siflora subsp. villosissima, and these plants are still
alpinum, Pohlia nutans, Rhacomitrium fasciculare, living. No other evidence of plant cultivation was
R. lanuginosum, Rhizocarpon atroalbescens, R. seen on this island.
geographicwn, Sphaerophorus globosus, Sphenolo- d. Burned areas.
Irus minutus, Thelypteris limbosperma, and
Umbilicaria proboscidea. An area of Empetrum heath approximately 1
square kilometer in extent was burned during
15. MISCELLANEOUS HABITATS, SUBSTRATES, AND World War II. Apparently, only the aerial parts
COMMUNITIES
a. Carcasses.
Nitrophilous mosses of the family Splachnaceae
most commonly grow on dung of herbivorous mam-
mals. These animals do not occur on Amchitka
Island, however, and the only splachnaceous moss
that is present, Tetraplodon mnioides, grows on de-
composed carcasses of birds and rats. This moss can
be found on any part of the island (except possibly
at high altitudes) where carcasses happen to lie.
Tetraplodon colonies are conspicuous because of the
density and reddish-brown color of the sporophytes
and the abundance of greenish-yellow spores that
are produced. Byrum stenotrichum was found
growing on a rat carcass, and Pleuro&nt/m schreberi,
on a bird carcass, but these mosses do not commonly
grow on this type of substrate. No other plants were
found on carcasses.
b. Imported wood.
FIGURE 35. Sitka spruce (Picea sitchensis) planted in front of
No native trees grow in the Aleutian Islands; a club building during World War II. Tree is about 3 feet
therefore, all wood on the islands was brought in by high. Photographed February 19,1968.
of plants were burned, whereas the saturated sub- CAPRIFOLIACEAE

strate protected the crowns and roots. After 20 years Linnaea borealis subsp. toorealis Hult. 7298, 4a; 7500, 4a;
the burned area still contrasts strongly to adjacent Erdman 564, 4a. Occurs commonly on banks in a mat of
unburned areas because of the abundance of Cala- Empetrum and mosses; flowers in mid-August.
magrostis nutkaensis and other grasses that revege- CARYOPHYLLACEAE
tated the area after the burn. The grass disclimax Cerastium aleuticum Hult. 7573, 12b; 8226, 14b; Erdman
that develops after a heath fire was described by 563, lib. Found only on unstable soil at high altitudes:
Pearsall (1950, p. 145, 155). Lichens, including flowers in mid-August.
Cladonia cocci/era, C. degenerans, C. gracilis, C. Cerastium êeringianum var. teeringianum Hult. (Hulten.
1960, p. 178).
pacifica, C, pyxidata, and Empetrum nigrum subsp. Cerastium fischerianum Ser. 7496, la; 7995, lla. This plant
nigrum are beginning to revegetate the burned area. has slender light-red stems that are very brittle.
Probably many more years will be required for this * Cerastium f&ntanum subsp. triviale (Link) Jalas. 7470,
area to recover completely from the fire damage. lOb; 8040, 5b. Common on roadsides and sand dunes.
SoncJcenya peploides subsp. major (Hook.) Hult. 7490, la.
Found only on sandy or rocky beaches; forms a zone of
PLANT SPECIES OF AMCHITKA ISLAND
dense vegetation near the high-tide level.
ANNOTATED LIST OF VASCULAR PLANTS Saffina crassicaulis S. Wats. 76.18, 4d; 7619, 4d. Found only
in wet crevices of sea cliffs.
By HANSFOKD T. SHACKLETTE, JAMES A. EKDMAN, and JOHN R. Sagina intermedia Fenzl. 7575, 12b; 8037, lOc; Erdman 578,
KEITH 12b. Occurs on salifluction slopes and rocky roadsides:
forms very dense rosettes on hard soil or in pavement
Duplicates of specimens cited in this list have been cracks.
identified or their names verified, with but few excep- *Sagina occidentalis S. Wats. 7471, lOc; 7654, la. Found on
tions, by Dr. Eric Hulten (written commun., 1967), and upper beach terraces and on old gravel roads.
the nomenclature follows that of his new flora (1968). Stellaria calycantha subsp. calycantha Hult. 7443, 4g; 7633,
Species from Amchitka Island that were reported by la ; 8034, lOc. On disturbed soil at low altitudes.
Stellaria media (L.) Vill. (See Hulten, 1960, p. 176.)
Hulten (1960) but were not found during this study *Stellaria ruscifolia subsp. aleutica Hult. 7580, 12b; 8223,
are cited, but without habitat notes. Collection numbers, 12b. A rare plant of solifluction slopes. Leaves rigid,
in italics, are those of Shacklette unless otherwise speci- glaucous, and sharp pointed. Flowers very sparingly, in
fied; they follow the species name. The habitat and mid-August.
plant community from which a collection was taken, Stellaria sitchana var. tongarMana (Fern.) Hult. 75^6, 4g.
On frost scars and other sites of disturbed soil in the
designated by numbers and letters as set forth in the Empetrum heath.
section of this report on the description of the island
and its vegetation, follow the collection number. One COMPOSITAE
asterisk before the species name indicates that this is a Achillea torealis Bong. 7^39, lOb. Common on roadsides and
report of the first collection of this species on Amchitka on disturbed soil in the Empetrum heath.
Anaphalis margaritacea (L.) Benth. & Hook. f. 7423, lOb.
Island; two asterisks indicate a first collection of the Grows in dense colonies on compacted road shoulders.
species in the Aleutian Islands. Probably an adventive species.
Antennaria dioica (L.) Gaertn. 7255, 14a; 7395, lOc; 7440,
BORAGINACEAE
lOb; 7613, 4g; 7871, lOa. Characteristic of frost scars and
Mertensia maritima subsp. maritime Hult. 8196, la. This other areas of disturbed soil, but was not found above an
plant occurs only on sandy beaches and was found at altitude of 800 feet
only a few locations on the island. Antennaria pallida E. Nels. Cited by Tatewaki and Kobay-
CALLITRICHACEAE ashi (1934, p. 84) as A. alpina Gaertn.; Hulten (1960,
p. 346) considered this citation to refer to A. pallida but
*Callitriche anceps Fern. 7631, 8b. Grows in shallow road- did not cite a collection from Amchitka Island.
side pools; rare. Arnica unalaschcensis Less. 7^79, lOb. Abundant on road-
Callitriche hermaphroditica L. (Hulten, 1960, p. 257). sides and disturbed soil of the Empetrum heath. Begins
CAMPANULACEAE to flower in early August.
Chrysanthemum arcticum subsp. arcticum Hult. 7625, 4a.
Campanula chanvissonis Federov. 75^9, 14a; 7550, 14a. Common throughout the island except at high altitudes.
Found only at altitudes above 300 feet. The common form Begins to flower in late July.
has violet corollas, but at places colonies of many plants Erigeron peregrinus subsp. perearinus Hult. 7469, 4a. The
with lavender to pink corollas occur. Flowers in August. large pink flowers are showy and resemble some cultivated
Campanula, lasiocarpa subsp. lasiocarpa Hult. 7397, 4a; asters. Abundant and widespread; flowering from late
7653, 14a; Erdman 580, 12a. Most abundant in the Empe- July to mid-August.
trum heath, but it occurs also throughout the mountains *Hieracium gracile var. alasJcanum Zahn. 7433, lOc. Not com-
except on the highest peaks. Flowers in August. mon ; found only at altitudes of no more than 200 feet.
PLANT SPECIES OF AMCHITKA ISLAND 35
Hieracium triste Willd. 7561, lla. Not common; found only Carex kclloggii W. Boott. 7485, 8a; 8031, 8b. Commonly oc-
at altitudes above 300 feet. curs in wet sandy soil at low altitudes, where it forms
*Petasites frigidus (L.) Franch. 7624, 4e; 7845, 4b. Not com- dense stools.
mon. Flowers the first of June, before the leaves are well Carex lyngbyaei Hornem. 7451, 6a; 7^67, 4a; 7473, 4b;
developed. 7537, 4c; 7658, 6a; 8199, 5b. Occurs in many different
Scnecio pseudo-arnica Less. 7503, la. Abundant on all sandy habitats at low altitudes and ranges in height from about
or pebbly beaches; at places forms dense stands of tall 10 cm at unfavorable locations to nearly 2 m where g~ow-
(as much as 1.3 m) succulent plants; never found inland. ing in streamside alluvium.
Flowers in mid-August. Carex macrochaeta C. A. Mey. 7478, lOa; 7610, 14a. Wide-
Taraxacum trigonolotitm Dahlstedt. 7491, 3b. Usually grows spread and abundant on the low plateaus.
in crevices of sea cliffs, but occasionally occurs on road- Carex nesophila Holm. Questioned as occurring on Am-
sides. Tatewaki and Kobayashi (1934, p. 89-90) described chitka Island by Hulten (1960, p. 122-123) on the 1 <?sis
a new species, T. aleiiticum Tatewaki & Kitamura, from of the report of C. macrochaeta var. subrigida Kuek.
Amchitka Island, but Hulten (1960, p. 366) considered it apiid Macoun by Tatewaki and Kobayashi (1934, p. 307).
to be T. trigonoloftum. Carex pluri flora Hult. 7536, 4c; 7656, 6a; 8221, 13b. Com-
mon in the Empetrum heath.
CORNACEAE
Carex saxatilis subsp. laxa (Trautv.) Kalela. 7586, 13a. Ap-
Cornus eatwdcnsis L. x succica L. (See Hulten, 1960, p. parently rare; found only in a snow-bed community.
274-275.) No specimens of Cornus that matched Hulten's Carex stylosa C. A. Mey. 8185, 4a. Found only in the Empe-
description of this hybrid were found. trum heath on the low plateaus.
Cornus suecica L. 7404, 4a. Abundant in the moss and lichen Erioplwrum russeolum subsp. rufescens (Anders.) Hyl.
mats at medium to low altitudes. Flowers in mid-June to 7428, 6a; 7534, 8a; 7655, 6a; 8026, 6a. The heads range
early July and produces bright-red fruits abundantly. in color from white to slightly tawny. The rhizomer are
deeply buried in a tough layer of fibrous peat.
CRUCIFERAE
Eriophorum russeolum var. ma jus Sommier. Collected on
Aphragnnts esclisclwltzianus Andrz. (See Hulten, I960, p. Amchitka Island by Hutchison (1937, p. 229) who wrote,
206.) "A very handsome species of bog-cotton with large pale
*Cardamine bellidifolia L. Erdman 567, 12b. Rare; appar- golden tufts." This collection was identified by Hulten
ently restricted to unstable, nearly bare, rocky soil at high but was not recorded from Amchitka Island in his fora.
altitudes. The species was reported from Amchitka Island by Tate-
Cardamine umbellata Greene. 7474, lOb; 7846, 4b; 7903, 3c; waki and Kobayashi (1934, p. 110). It was not found
7997, 4e. Abundant in a wide variety of habitats at alti- during the present study, although especially sought.
tudes below 400 feet. *Triclwphorum caespitosum (L.) Hartm. 8283, 4b. Found
Cochlearia offlcinalis subsp. oftlongifolia (DC.) Hult. 7614- only in moss and lichen mats on the lower terrace at
3b. Occurs typically on the thin wet soil cover of coastal Chitka Cove.
cliffs.
*Draba aleutica Ekman. 7571, 12b. Rare; apparently re- DIAPENSIACEAE
stricted to solifluction slopes, where it forms dense **Diapcnsia lapponica subsp. obovata (F. Schm.) Hult. 7570,
rosettes. 12b; Klein 2829, 12a. This collection provides an iirwr-
Drafter hyperborea (L.) Desv. 7615, 3b; 8022, 3b. A charac- tant range extension for the species. Found growing
teristic species of crevices in sea cliffs. Flowers in June among andesite pebbles of a wet solifluction slope ard in
and July; the large petals are cream colored.
a fell-field. The flowering season was past in mid-August.
*Siil>ularia aquatica L. 8028, 8b. Found flowering the middle
of June in a shallow rock-bottomed pool. Petals of the EMPETRACEAE
minute stellate corolla are white on the upper side and
Empetrum nigrum subsp. nigrum Hult. 7295, 4a; 7535, 4c;
light blue on the lower side.
8054. 14a, Erdman 547, 4a. The most abundant and char-
CYPERACEAE acteristic plant on the island. It flowers in early May,
Carex antlioxantliea Presl. 7465, 4b; 7657, 6a. Grows on and on June 5 the green fruits were almost full size.
ridges in the Empctrum heath and in wet sedge meadows; EQUISETACEAE
nonstooling; never forms large colonies.
*Carcx bigeloicii Torr. 7587, 13a. Apparently very rare, oc- Equisetum arrense L. 7^34, lOc; 7^95, 2a; 7626, 4a; 7914,
curring only in snow-bed communities. Tatewaki and lOc. Abundant and widespread in many habitats, but best
Kobayashi (1934, p. 108) reported C. rigida Good, from developed as colonies on sand dunes and disturbed mir^ral
Amchitka Island; this report probably should be referred soil. The sporangia mature in early June and are pro-
to C. bigelowii. duced in abundance.
Carex circinnata C. A. Mey. 7606, 14b; 7891, 12a ;8228, 14b.
ERICACEAE
This is one of two vascular plant species that can endure
the most severe weather of mountain summits, and al- Cassiopc lycopodioides (Pall.) D. Don. 7254, 14a; 7567,12a;
though it fruits very sparingly, at places it forms large 7852, 12b; 8053, 14a. Common in the Empctrum heath, but
colonies. usually depauperate. At high altitudes it is a major com-
*Carex dioica subsp. gynocratcs (Wormsk.) Hult. Klein 2710, ponent of the discontinuous heath patches. Flowei*s in
4a. Found on a hummock in the center of a pond. June and early July.
Loiseleuria procumbens (L.) Desv. 7264, 14a; 7867, lOc. although single plants are widespread at low altitudes.
Common and well developed in the Empetrum heath; At some locations it forms many dense cylindrical stools,
flowers profusely in mid-June. as much as 0.5 m high, that are separated by narrow
*R1iododendron camtschaticum subsp. camtschaticum Hult. passages between the stools. The development of this
8216, 12a. Apparently occurs only in small patches of unusual type of growth pattern was observed only on
heath near the mountain tops. Flowers in late July. slopes and probably is caused by soil creep.
Vaccinium uliginosum subsp. micropliyllum Lange. 8217, Festuca brachyphylla Schult. 7612, 14a; 7999, 14b. A copro-
I2a. Found only with Rhododendron; neither flowering philous species on this island, found only on the summits
nor fruiting. of bird-perch mounds, where it forms a very dense turf.
Vaccinium vitis-idaea subsp. minus (Lodd.) Hult. 7296, 4c. Considered by Klein, in his section on plant species in this
Most commonly occurs in moss mats of the Empetrum report, to be F. ovina var. braohyphylla (Schult.) Piper.
heath and on moss mounds. Flowers very sparingly and *Festuea nibra L. 7297, 6a; 7408, 4a; 7518, 3d. Found in
none was observed to bear fruits. Some leaves turn bril- several habitats on the low plateaus.
liant red and persist through the winter. Festuca rubra subsp. aucta (Krecz. & Bobr.) Hult. 7408, 4a ;
7463, 4a; 8189, 2a. Vegetative plants are abundart in the
GENTIANACEAE Empetrum heath; fruiting plants are not common in the
*Cfentia-na amarella subsp. acuta var. plebeja ( Cham. & heath, but are abundant at some sites of distur*]^! soil.
Schleoht.) Hult. 7400, lOa. Found only on a disturbed Klein, in his section on plant species in this re'wrt, in-
soil bank. The corolla is dull blue to lavender and is pro- cludes this form in the species listed above.
duced in early August. Hierochloe odorata (L.) Wahlenb. 7849, 3d. Occurs as in-
*Gentiana aleutica Cham. & Schlecht. 7862, lOb; 7683, lOb: dividual plants or small colonies in the Empetrum heath.
8191, 14a. Two color forms occur together one with Phleum oommutatum var. americanum (Fourn.) Hult. 7466,
white, the other with dull purple, corollas. Flowers in 4a. Common 'in the Empetrum heath and on disturbed soil
late July and early August. at low altitudes.
**Poa alpina L. 8032, lOc. This collection represents an impor-
GERANIACEAE tant range extension for this species. The specimens grew
Geranium erianthum DC. (Hulten, I960, p. 256, listed as a in the rocky soil of an old trail, and Hulten (written com-
sight record only.) This very conspicuous plant was mun., 1968) remarked that they were "The smallest speci-
sought during the present study without success. If it mens I have seen." Klein, in his section on plant species
occurs on this island now, it is rare. It was not found on in this report, placed Poa hispidula, P. komarovii, P.
Rat Island, but was common on Shemya Island. lanata, and P. turneri and their varieties in the P. alpina
species complex.
GRAMINEAE *Poa aretica subsp. idlliamsii (Nash) Hult. 7609, 14a. Ap-
Agrostis alaskana Hult 7515, 4f. Found only on a peat bank parently a coprophilous plant. Found only on the summit
near Constantine Harbor. Considered by Klein, in his of a bird-perch mound.
discussion of plant species in this report, to be A. borealis Poa eminens Presl. 7342, 2a; 7493, 2a; 8190, 2a. The short
Hartm. blades are thick and rigid, and the entire plant light pur-
*Agrostis borealis Hartm. 8230, 14b. Found only at altitudes ple. Found only on the deposit of sand and shells at South
above 900 feet. Bight and on sand dunes near Crown Reefer Point.
* Agrostis eorarata Trin. 7438, 2a; 7794, 2a. The culms of this Poa lifepidula var. aleuttoa, Hult. (See Hulten, 1960, p.
plant spread horizontally and are closely appressed to the 94-95.)
sandy deposits on which they grow. Poa Jiispidula var. wmpara, Hult. (See Hulten, 1960, p. 95.)
Alopecurus aequalis Sobol. 7460, 8b; Erdnmn 555, 8a. Found Poa, malacantha Kom. (P. komarovii Roshev., Hulten, 1960,
only in shallow pools and other very wet locations. p. 96).
Bromus sitehensis var. aleutensis (Trin.) Hult. 7492, 2a. *Poa lanata Scribn. & Merr. 8187, 2a. Found only on the de-
On sand and shell deposits at South Bight. posit of sand and shells at South Bight.
Galamagrostis nutkaensis (Presl) Steud. 7213, 4c; 7291, 4c: *Poa lanata var. viin-para Hult. 7566, 12b. Found only in
7294, 4c; 7383, lOa; 7519, 3d; 75,38, 4c. The most common gravelly soil of a roadside ditch at an altitude of about
and widespread grass on the island. It forms almost pure 650 feet.
stands on roadside banks, but is not abundant at high Poa turneri Scribn. 8186, lOc; Erdman 548, lOc. Fonnd only
altitudes. on rock rubble on an old road near Constantine Harbor.
Descharnp&ia alpina (L.) Roeni. & Schult. Cited from Am- Puccinellia langeana subsp. alaskana (Scribn. £ Merr.)
chitka Island by Tatewaki and Kobayashi (1934, p. 97). S0rens, 7504. 3a; 7617, 4d. Apparently a coprophilous spe-
Hulten (1960, p. 85), however, doubted that this species cies. Found only in drain channels of bird perches on the
occurs in Alaska. i coastal cliffs and on the summits of bird-perch mounds.
Deschampsia beringensis Hult. 7475, 4b; 8188, 2a. Common Trisetum spicatum subsp. alaskanum (Nash) Hult. (See
in open areas of the Empetrum heath. The spikelets are Hulten, 1960, p. 90.) Tatewaki and Kobayashi (1934) rec-
purplish red. Considered by Klein, in the section on plant ognized both this taxon and the variety molle Scribn. &
species in this report, to be D. caespitosa (L.) Beauv. Merr. in the Amchitka Island flora.
Deschampsia caespitosa subsp. orientalis Hult. 7412, 8a Vahlodea atropurpurea subsp. latifolia (Hook.) Pors. 7464,
Occurs at the margins of shallow pools, as well as on com- 4a; 8220, 13b. Occurs in the Empetrum heath and in al-
pacted roads. pine meadows. Considered by Klein, in his section on
Elymus arenarius subsp. mollis var. mollis Hult. 7289, 2b; plant species in this report, to be Deschampsia atropur-
7520, 3d. Occurs in large stands only near the coasts. purea (Wahl.) Sheele.
Vahlodea atropurpurea subsp. paramushirensis (Kudo) Luzula wahleribergii subsp. piperi (Oov.) Hult. Amchit>a
Hult. (See Hulten, 1968, p. 115; Amchitka Island occur- Island occurrence shown on the distribution map by Hul-
rence indicated by a dot on the distribution map.) ten (1968, p. 297). Listed from this island as L. divaricata
Watson by Tatewaki and and Kobayashi (1934, p. 112).
HALORAGACEAE
*Hippuris montana Ledeb. 8227, 13a; 8261, 13a. This plant LEGUMINOSAE
has the appearance of a Oalium or a Poffonatum; the Lathyrus maritimus subsp. pubescens (Hartm.) C. Reg^l.
stems are about 10 to 15 cm long, but only the leafy part, 7489, la. Common on beaches and sand banks near the
about 1 to 4 cm long, protrudes from the mud. This plant coasts; flowers and fruits are produced abundantly.
was also found in a Sphagnum mat of a small mountain Lathyrus palustris subsp. pilosus (Cham.) Hult. Hulten
stream that issued from a spring. (1960, p. 255) wrote, "Amchitka, according to Tatew. &
Hippuris vulgaris L. 7627, 7a. Common in lakes and pools Kobay." This report was verified by Hulten (1968, p. 67f).
at low altitudes; the leafy stems commonly emerge from Not found during the present study, although often
the water, which can be as much as 1 m deep. sought.
Hyriophyllum spicatum L. (See Hulten, 1960, p. 268.) Not Lttpinus nootkatensis Donn. 7^86, lOb; 8044, 5b; 8228, 13b.
found during the present study, although extensively An abundant and very showy plant, especially common
sought. Its superficial resemblance to the common Ra- along roadsides; occurs at all altitudes. Two colonies
nunculus trichophyllus may have caused it to be of plants found had pale-pink to nearly white flowers;
overlooked. these may be referred to f. leucanthus Lepage.
*Trifolium repens L. One clone of this adventive plant was
IRIDACEAE found on a disturbed site by W. L. Slauson and M. A.
Iris setosa subsp. setosa Hult. Reported to occur at Chitka Heifner.
'Cove (sight record, by W. M. Klein).
LENTIBULARIACEAE
ISOETACEAE Pinffuicula vulffaris subsp. macroceras (Link) Calder &
Isoetes muricata subsp. maritima (Underw.) Hult. 7373, 7b; Taylor. 7414, 5a ; Erdman 753, 4b. Occurs only at low alti-
7396, 7b; Erdman 575, 7b. Found only in lakes, growing tudes at very wet sites in the Empetrum heath.
in water about 0.5 m deep. The plants are robust, with
conns as much as 3 cm in diameter; many sporelings also LILIACEAE
were found. Fritillaria camschatcensis (L.) Ker-Gawl. Klein 2718, 4e.
Isoetes sp. 8192, 8c. A slender plant with only slight corm de- Abundant in streamside communities throughout the low
velopment; occurs in shallow rock-bottomed pools. Its plateaus.
aspect and habitat suggest that it may be a different taxon *Majanthemum dilatatum (How.) Nels. & Macbr. 7866, 4f.
from that listed above. Found only on peat banks and is only locally abundant.
Streptopus amplexifolius (L.) DC. 7543, 4e. Found only in
JUNCACEAE streamside communities, but was not abundant at any
Juncus arcticus subsp. sitctiensis Engelm. 7634, la - A com- location.
mon species at wet sites; occurs at low to medium-high Tofleldia coccinea Richards. 7262, 14a; 7410, 4a. Common
altitudes. in a wide variety of habitats throughout the island.
* Juncus biglumis L. 7576, 12b; Erdman 579, 12b. These col-
lections are from unstable soil at high altitudes. The LYCOPODIACEAE
plants are from 1 to 8 cm high. Lycopodium alpinum L. Keith 108, lla. Found only in moss
*Juncus ensifolius Wikstr. 7685, 8a; 7882, 8a. Found only at and liverwort mats in a small stream.
pool margins at low altitudes. Even when sterile, plant Lycopodium annotinum subsp. annotinum Hult. 7236, 5c;
can be recognized by its ensiform leaves. 7284, 4a; 7680, 4a ; Keith 107, 4a. Locally abundant in the
*Juncus trialiimis subsp. albescens (Lange) Hult. 7484, 8a; heath of the low plateaus.
8210, 8a. A densely caespitose species that grows in shal- Lycopodium olavatum subsp. clavatum Hult. 7283, 4a;
low pools at low altitudes. The culms are about 15 cm 7681, 4a; Keith 109, 4a. Grows with the species listed
high. above.
Luzula arcuata subsp. unalaschcensis (Buchenau) Hult. Lycopodium sabinaefolium var. sitchense (Rupr.) Fe~n.
7563, lla; 7608, 14a; 8194, 14a. Found only at altitudes 7237, 5c; 7263, 14a. Found commonly on low-a'ltitrde
above 600 feet. bedrock outcrops.
Luzula multi flora subsp. kobayasii (Satake) Hult. 7461, lOc. Lycopodium selago subsp. selago Hult. 7253, 14a; 7432, lOc;
Found on disturbed mineral soil near Constantine Harbor. 7870, 4a. Widely distributed, although not abundant,
Luzula multiflora subsp. multi flora var. frigida (Buchenau) throughout the island.
'Sam. 7419, lOa; 7659, 4a. Specimens of the multifiora
complex are abundant on disturbed sites on the low ONAGRAGEAE
plateaus. Epilobium behringianum Haussk. 7560, lla. Apparently com-
Lusula parviflora subsp. parviflora Hult. 7472, lOa; 7686, mon on the island; however, because of confusion in nom-
4e; 8030, lOc. Found only at low altitudes on banks and enclature with the following species, its abundance is
roadsides; not common. not known with certainty.
Luzula tundricola Gorodk. 7883, 12a; 7907, 12b; 7994, Ha. Epilobium glandulosum Lehm. 7448, 6a; 7547, 4g. A robust
Found only at altitudes above 1,100 feet in exposed fell- plant as much as 40 cm high if growing in wet sedge mea-
fields and on solifluction terraces. dows ; much smaller if growing in the Empetrum heath.
Klein, in his section on plant species in this report, refers POLYPODIACEAE

this species to E. boreale Haussk. Athyrium filix-femina subsp. cyclosorum (Rupr.) Christens.
Epilobiwm sertulatum Haussk. 7436, lOb; 7437, lOb; S039, 7517, 3d; 76S7, 4e. The common tall fern of s c.reamside
5b. A low-growing plant occurring along roadsides and on communities; it also occurs widely scattered through the
disturbed soil. Plants at a site may either have green Empetrum heath.
leaves and pale-pink flowers or be dark red throughout. Cystopteris fragilis subsp. fragilis Hult. 7690, 4e; 7904, 3c.
ORCHIDACEAE Most common in crevices of the sea cliffs, but also occurs
on shaded peat banks.
Dactylorliiza aristata (Fisch.) Soo. 7417, 3d; 8027, lOb. Dryopteris dilatata subsp. americana (Fisch.) Fult 7688,
A showy species that is locally common at low altitudes, 4e. A tall fern that occurs sparingly in streamside
especially on compacted mineral soil. The lip is pale communities.
magenta spotted with purple; other petals are deep laven- Gymnocarpium dryopteris (L.) Newm. (See Hul^n, 1960,
der. One white-flowered specimen was found (Klein 1744)- p. 50, as Dryopteris linneana C. Chr.)
Listera cordata var. nephrophylla (Rydb.) Hult. 7416, 4a; ^Polypodiutn vulgare subsp. occidentale (Hook.) Hult. 8268,
7679, 4a. This very small orchid occurs as scattered in- 4d. Found only at the base, of a bird-perch mound above
dividuals in the moss mat of banks in the Empetrum heath the Pacific Ocean cliffs near the entrance to South Bight.
and in alpine meadows. Plants with either green or pur- On February 8 the leaves were green and b^re many
plish-brown flowers grow intermixed. sori, although the sporangia were empty.
* 'Platanthera chorisiana (Oham.) Rchb. 741S, 4a; 7497, Thelypteris limbosperma (All.) Fuchs. 8244, 14b; 8245, 13b.
14a ; 8210, lOc. Found only on lithosol or on sparsely vege- Found only at high altitudes; in alpine meadows or rock
tated mineral soil. The petals and sepals at anthesis are slides.
greenish yellow with narrow whitish margins; they soon Thelypteris phegopteris (L.) Slosson. 8243, 13b. Found only
'become brown tipped or brown dotted and streaked. in alpine meadows.
Platanthera convallariaefolia (Fisch.) Lindl. 7426, 4b. This
and P. dilatata are common in wet places at low altitudes; PORTULACACEAE
usually both species occur intermixed. The flowers are *Claytonia arctica Adams. Klein 2820, 12b. Found only in
green. a fell-field community; occurs as scattered individuals.
Platanthera, dilatata (Pursh) Lindl. 7468, 4a. Very similar The corolla is white.
to P. convallariaefolia, but has pure-white flowers. The Claytonia sibirica L. 7406, 4b. Very abundant, usually in
flowers of Amchitka Island specimens are not fragrant, wet locations; on low plateaus and in alpine meadows.
contrary to the report of Hutchison (1937, p. 231). Plants with white or pink petals occur together. The
Platanthera tipuloides var. behringiana (Rydb.) Hult. (See white-flowering form was named var. albiflorc, by Tate-
Hulten, 1968, p. 323.) waki and Kobayashi (1934, p. 34) and was reported from
PAPAVERACEAJE Amchitka Island.
*Montia fontana subsp. fontana Hult. 7632, 3b; 7684, 9a;
Papaver alaskanum Hult. Collected in the mountains of 8198, 3b. Found only in cascades and on dripping sea cliffs.
Amchitka Island; the specimen is preserved in the
Colorado State University herbarium. PRIMULACEAE
PLANTAGINACEAE Primula cuneifolia subsp. saxifragifolia (Lehm.) Sm. & For-
rest. 7421, lOa ; 7430, 4a ; 7583, 13a ; 7848, lOa ; 7869, 4a ;
*Plantago macrocarpa Chain. & Schlecht. 7427, 6a. Occurs Erdman 551, 14a. Occurs at all altitudes on the island and
in wet sedge meadows on the low plateaus. is most abundant on disturbed soil. Flowers in late May
Plantaao maritima subsp. juncoides (Lam.) Hult. 7660, 5a; and early June on the low plateaus and in late July and
797^, 3b. Occurs in bogs and on soil of coastal cliffs. early August at high altitudes. The variety albiflora
Koidzumi was reported for Amchitka Island by Tatewaki
POLYGONACEAE
and Kobayashi (1934, p. 72).
*Koenigia islandica L. 7574, 12b; Erdman 562, 12b; Erdman Trientalis europaea subsp. arctica (Fisch.) Hult. 7501, 4a.
577, 12b. Found only on wet mineral soil of solifluction Common in the Empetrum heath at low altitudes; flowers
slopes at high altitudes. in mid-August. Tatewaki and Kobayashi (19^?4, p. 73-
*Oxyria digyna (L.) Hill. 7582, 13a. Rare; found only at 74) described the variety alentica and designated the Am-
high altitudes. chitka Island specimen as the type. Hulten (1960, p.
298) considered this variety to be "a commo^ form of
Polygonum viviparum L. 7435, lOb; 7562, lla; 8218, 13b.
the plant."
Widely distributed but nowhere abundant on the island.
Rumex fenestratus Greene. (See Hulte'n, 1960, p. 167-170.) PYROLACEAE
Hutchison (1937, p. 232) reported this species to be *Pyrola asarifolia var. purpurea (Bunge) Fern. 7611, 4a.
"very luxuriant and fairly common on Amchitka." Only one plant was found, although the species vas sought
* Rumex obtusifolius L. 7544. la. This European and Asiatic extensively. Flowers in mid-August.
species is adventive in the Aleutian Islands (Hulten, 1960, *Pyrola minor L. 8219, 13b. Found only along streams in the
p. 168). It occurs as isolated individuals on wet disturbed mountains; locally abundant. Flowers the latter part
soil, usually in areas of human habitation. of July.
PLANT SPECIES OF AMCHTTKA ISLAND 39
BANUNCULACEAE BUBIACEAE
Anemone narcissiflora subsp. villosissima (DC.) Hult. 7413, *Galium aparinc L. 8195, la. Found only in ruderal habitats,
3d; 7844, 4a. One of the earliest flowering herbs, and usually in driftwood deposits.
very abundant and conspicuous throughout the low Galium trifldum subsp. columbianum (Rydb.) Hult. 7652,
plateaus. la; 7678, 4a; 8045, 6a. Found only at low altitudes; most
CaltJia palustris subsp. asarifolia (DC.) Hult. 7200, 6b; common in deposits of driftwood or in wet sedge meadows
1629, 6b. Produces new leaves under water in November near the ocean.
and December that emerge in early spring; fully developed
leaves and flowers are present in early June. SALICACEAE
Copt is trifolia (L.) Salisb. 7222, 4a; 7628, 4c; 8052, 4a. The Salix arctica subsp. crassijulis (Trautv.) Skvortz. 7420, 3d;
firm shiny leaves persist through the winter, some becom- 7551, 14a; 79S8, 12a; 7996, lla; Erdman 549, 4a. The
ing red, and are replaced by new leaves at the time of largest native woody plant on the island. The prostrate
flowering in late June. stems are as much as 0.5 m long and 1 cm in diameter.
*R<munculus Jiyperboretis subsp. hyperboreiis Hult. 7429, Widely distributed, but most abundant at sites near the
7b. Found only by a small pool, growing on mud; flowers ocean.
the first half of August. Salix cyclophylla Bydb. 7548, 14a; 1851, 12b; 7884, ITa;
Ranunculus occidentalis subsp. insiilaris Hult. 7909, 12b. 8209, 4a. Widely distributed, but most abundant in
Common throughout the island, except at high altitudes. patches of discontinuous heath at high 'altitudes.
Ranunculus occidentalis subsp. nelsoni (DC.) Hult. 7450, Salix polaris subsp. pseudopolaris (Flod.) Hult. Collected
6a. Found only in wet sedge meadows. on this island by Tatewaki and Takahashi, according to
Ranunculus reptans L. 7598, 7b; 7399, 7b; Erdman 553, Hultfin (I960, p. 161).
8b. Grows as a submersed or emergent aquatic, or on Salix rotundifolia Trautv. 7558, 14a; 7S85, 12a; 7586, l£a;
muddy lake shores. The leaf shape ranges from linear 7998, lla; Erdman 560, lla. Found only at altitudes of
to spatulate, and is very similar to that of Limosella 800 feet or more; commonly grows with S. cyclophylla.
aquatica.
Ranunculus trichophyllus var. trichophyllus Hult. 7206, 8a; SAXIFBAGACEAE
7444, 9a; 7674, 9a; 8266, 9a. Grows submersed in rapidly *Leptarrhena pyrolifolia (D. Don) Ser. 7584, 13a; 8222, 13a.
flowing streams, in pools of wet sedge meadows, and in Bestricted to high-altitude snow beds; locally common.
lakes. Flowers in mid-July.
Parnassia kotzebuei Cham. & Schlecht. 7477, 8a;8042, 5b.
ROSAOEAE Found only on wet mineral soil at low altitudes. Flowers
Geum calthifolium Menzies. 7462, 4a. Abundant on the Em- from mid-July to mid-August.
petrutn heath and in alpine meadows; flowers from late Saxifraga bracteata D. Don. 74S8, 3b; 7616, 3b; 7881, f b;
July to mid-August. 7912, 3b. A common and characteristic plant of coastal
Geutn macrophyllum subsp. macrophyllum Hult. 7449, 6a. cliffs.
One of the few tall forbs on the island, reaching a height *Saxifraga foliolosa var. foliolosa Hult. 7579, 12b. Rare;
of 1 m at places. It may be an adventive species (Hulten, found only on wet solifluction slopes. The plant bears
1960, p. 247). many bulbils in the branches of the inflorescence.
Geum pentapetaltim (L.) Makino. 8215, 13a. This low plant *Saxifraga liirculus L. 8225, 13a. Found only in moss and
occurs only at high altitudes in wet snow-bed communi-
liverwort mats of snow beds. This is the only report of
ties. The white flowers appear in mid-July.
this species from the central and western Aleutian
Potentilla hyparctica Malte. 7572, 12b; 7605, 12b; 7993,
12b. This species is restricted to unstable soil at high Islands.
altitudes. Saxifraga pimctata subsp. insularis Hult. 7445, 4e; 7577,
Potentilla egedii subsp. grandis (Torr. & Gray) Hult. 7630, 12b; 7578, 12b;7868, lOc; 7985, 4a; 8043, 5b; 8824, 12b;
lOb. Found only on gravelly soil of a lake shore. Flowers Erdman 554, 5b. Common and widespread, usually oc-
in mid-August. curs in sheltered locations.
Potentilla villosa Pall. 7487, 3c; 7913, 3c; 8021, Be. A very
SCBOPHULARIACEAE
conspicuous plant with silvery pubescent leaves and stems;
the yellow flowers are produced from early June to mid- *Castitteja unalaschcensis (Cham. & Schlecht.) Malte. 8lc,u-
August. Found only on sea cliffs, where it is abundant. son, 15c. Found growing with the introduced Pieea
Rul)us chamaemorus L. 7405, 4a. Abundant in the Empe- sitcliensis that is shown in figure 35.
trum mat; fruits freely, the fruits not yet ripe at the end *Limosella aquatica L. 7221, 7b; 7394, 7b; 8200, 5b. Was not
of August. found flowering. The vegetative organs are similar to
Ruftnx arctlcus subsp. stellatus (Sm.) Boiv. emend. Hult. those of Subularia aquatica and to submersed forms of
7405, 4a; 7476, 4g. The red flowers are produced the first Ranunculus reptans.
week of August, but none that produced fruit was Mimulus guttatus DC. 7622, 9a. Found flowering in late
observed. July to mid-August in shallow streams of the Empetr'wi
Sibbatdia procumbens L. 7411, 4g; 7987, 12a; 8029, lOc. Oc- heath and in cascades at the coastal cliffs.
curs throughout the island; usually grows on disturbed Pedicularis chamissonls Stev. 7422, 4a; Erdman 552, 6a.
mineral soil. The large spikes of reddish-purple flowers, produced in
late July to early August, are conspicuous in the mixtures of several to many species; the same collection
Empetrum heath. number is used for all members of each species that
*Rliinanthus minor subsp. ~borealis (Sterneck) Love. 7431,
lOc; Erdman 574, 6a. Grows in disturbed mineral soil at
were segregated from the mixture.
low altitudes; flowers in late July and early August. In the list that follows, the collection numbers, in
Veronica americana Schwein. 1239, 9a; 7446, 9a; 1516, 8a. italics, following the species name are those of Shack-
Common along margins of swift streams. Flowers from lette, except as noted. The habitat and plant community
late July until December, the latest flowers being pro- in which a species was found, designated by numbers
duced entirely under water.
Veronica grandiflora Gaertn. 79861), 12a. A rare plant of
and letters as set forth in the section on the island and
high-altitude unstable lithosol. The dark-violet flowers its vegetation, follow the collection numbers. The habi-
are borne on very small plants in mid-June. tat notes and community assignments were provided by
Veronica serpyllifolia subsp. humifusa (Dickson) Syme. Shacklette. Species listed as newly reported for the
7403, 8a; 75^5, 4g; 7581, 13a; 8033, lOe; 8041, 5b. Abun- Aleutian Islands (Persson, 1968) are preceded by one
dant on mineral soil at low altitudes; flowers all summer.
Veronica stelleri Pall. 7.^99, 14a; Erdman 550, 14a; Erdman
asterisk; those newly reported for Alaska are indicated
561, 12b. A small pubescent plant that grows on dis- by two asterisks.
turbed soil and lithosol. Bears lavender campanulate I (Persson) am indebted to Dr. T. Amakawa, Shyu-
flowers from late July to mid-August. kan High School, Fukuoka, Japan, for the determina-
SELAGINELLACEAE tion of a Plectocolea that was in poor condition for
*Selaginella selaginoides (L.) Link. 7415, 5a; 8191, 8a.
study; to Dr. Eustace W. Jones, Oxford University,
Found only as scattered plants on the low plateaus. Oxford, England, for confirmation of my determina-
tions of two Anthelia specimens; to Dr. N. Kitagawa,
SPABGANIACEAE
Nara University of Education, Nara, Japan, for con-
Sparganium Jiyperboreum Laest. 7585, 13a; 8193, 8a. The firmation of my determination of a Cephaloziella speci-
plants are very slender and depauperate and grow on the
men and for determination or confirmation of some
mud at snow-bed sites and in shallow pools. Flowers
sparingly in late July. critical and poorly represented Lophozia specimens of
the section Ventricosae; to Dr. Elsa Nyholm, Natur-
UMBELLIFEBAE
historiska Riksmuseum, Stockholm, Sweden, for deter-
Angelica lucida L. 7.J80, lOb. A common plant of the Em- mination of a Sphagnum specimen; to Dr. E. R. B.
petrum heath; flowers in early to mid-August.
Little, Kingston Upon Hull, England, for naming two
Conioselinum chinense (L.) BSP. 7623, 4e; 8197, 3d. Occurs
as scattered plants on banks and streamsides; flowers Riccardia latifrons specimens and confirming the deter-
sparingly in late July and early August. mination of another; and to Dr. M. Mizutani, Hattori
Heracleum lanatum Michx. 7635, la. An abundant coarse Botanical Laboratory, Obi-Nichinan-shi, Japan, for his
plant of many habitats on the low plateaus. critical study of a Riccardia specimen.
Ligusticum scoticum subsp. hultetiii (Fern.) Calder & Identification of the Amchitkan hepatic specimens
Taylor. 7502, 2c. Found only on soil banks at the top of has been rather difficult because these plants cften are
sea cliffs; locally abundant.
represented only by single stems that creep through
VIOLACEAE tufts of other bryophytes; this condition of growth is
Viola langsdorffli Fisch. 7442, 6a; 7^98, 14a; 7677, 4a; 78^7, common in alpine and similar regions. In addition, the
4b. Common to locally abundant in the Empetrum heath sterility of the bryophytes, particularly the hepatics,
and alpine meadows; flowers from early June to mid- is especially pronounced in these specimens and doubt-
August.
less is caused by the severe climate that is charrcteristic
ANNOTATED LIST OF BRYOPHYTES of the Aleutian Islands.
By HERMAN PEBSsoN 2 and HANSFOBD T. SHACKLETTE HEPATICAE

Order TAKAKIALES
Duplicate packets of specimens cited in this list have
TAKAKIACEAE
been examined, and the species have been identified by
me (Persson), except as noted. I (Persson, 1968) have Takakia ccratophylla (Mitt.) Gro. [Lepidosia ce~atophylla
Mitt.]. 7591, 13a. Collected August 16, 1966, and first
also examined the entire collection of many specimens, segregated from a packet of Andreaea rupestris by Dr.
if critical specimens were found in the duplicated Howard Crum. This plant grew in a drain of a snow-bed
packets, and have published an account of the new or community at an altitude of about 650 feet and was
otherwise noteworthy bryophytes from Amchitka associated with large colonies of Anthelia julacea, 8ca-
Island. Many collections, especially of liverworts, are pania paltidosa, Sphagnum compaction, and other snow-
bed bryophytes. Reported for Amchitka Island by Sharp
2 Paleobotaniska avdelningen, Naturhistoriska Riksmuseum, Stock- and Hattori (1967), Hattori, Sharp, Mizutani, and Iwat-
holm, Sweden. suki (1968), and Persson (1968).
Order JUNX3KERMANNTALES HEBBERTACEAB
ADELANTHAOBAE Herberta adunca (Dicks.) Gray [H. hutchinsiae (Gottsche
& Rabenh.) Evans]. 7567'a, 12b. Found only at the base
Odontoschisma elongatum (Lindto.) Evans. 7338, 4c. Found of a boulder on a solifluction terrace, with Tritomaria
only on the summit of a small moss mound that develop- quinquedentata, Marsupella ema-rginata, Diplophyllum
ed on wet frost-heaved soil. albicans, and Jungermannia atrovirens.
ANTHELIACEAE JUNGERMANNIACEAE
Anthelia julacea (L.) Dum. 7235, 5c; 7564, 13a; 7592, 13a; Jungermannia atrovirens Dum. 7567a, 12b. Found only on a
7593, 13a; 8011, 14b; 8019, 14b. A characteristic species of wet solifluction terrace, growing with Herberta adunca
drains from snow-beds at high altitudes and very con- and its associated species.
spicuous because of the large gray colonies that it forms. Jungermannia sphaerocarpa Hook. 8208, 5b. Found only on
Dr. E. W. Jones has kindly confirmed the determinations ridges in wet sandy soil, mixed with Pogonatum urnige-
of Nos. 7592 and 7593. rum, LopJiozia cf. groenlandica, and Dicranella subvlata.
CALYPOGEIAGEAE Mylia anotnala (Hook.) Gray. 7564- 13a. Found only in a
drain channel at an altitude of 650 feet near Chitka Foint,
Calypogeia miilleriana (Schiffn.) K. Mull. 7261«, 4f. On a
intermixed with SpJMgnum papillosum, 8. squarroswn,
shaded peat bank above a small stream in the Empetrum
Anthelia julacea, CepJialozia ambiffita, Pellia neesiana,
heath, with Cephalozia media, Mniutn glabrescens, and
Oymnocolea inflata, Lophozia alpestris, and Cephalozia
Peltigera canina.
Calypogeia sphagnioola (Arn. & Perss.) Warnst. & Loeske. bicuspidata.
7209, 5a; 7214, 4a; 7258, 4c; 7341, 4c; 7455, 5a; 8281, 12a. Mylia taylori (Hook.) Gray. 7335, 4c; 7338, 4c; 7424, 4b.
In bogs at low altitudes, intermixed with Sphagnum Found only on moss mounds in the Empetrum heatt and
capillaceitm, S. magellanicum, and Riceardia latifrons in wet areas of the heath, where it formed tufts with
and on small moss mounds with Dicranum elongatum. Lophozia cf. groenlandica and Orthocaulis binsteadii.
Calypogeia triohomanis (L). Corda. 7540, 4b; 7862, 4e. In Nardia compressa (Hook.) Gray. 8282, 13a. Found only on
the Empetrum heath, intermixed with Oncophorux wah- the bottom of a snow-bed pool, where it formed a dense
lenbergii, Diplophyllum albicam, and Oymnocolea polster.
inflata. Nardia geoscyphus (DeNot.) Lindb. 8232, 13b; 8235, 13b.
Found inly in alpine meadows; grows with Lophozia
CEPHALOZIACEAE opacifolia.
Cephalozia ambigua Massal, 7524, 4f; 7564, 13a; 8232, 13b. Nardia scalaris (Schrad.) Gray. 7265, 4f; 7300, 4g; 7359,
On a peat bank in the Empetrum heath, intermixed with 2b; 7375, 14a; 7387, lOa ; 7389, 8a; 7523% 4f; 7524, 4f;
Mnium glabrescens, Lophozia cf. groenlandica, Dicranella 7526, 4f; 7593, 13a ; 7667 4f; 7668, 4f; 7841, 4f; 7842, 4f;
Jieteromalla, Nardia scalaris, and Scapania sca-ndica, and 7877, 4f; 7879a, 4f; 8036, lOc; 8050, 6a; 8232, 4e; 8233,
at altitudes of about 500 feet with Lophozia alpestris and 4e; 8358, 4f. Abundant in many habitats.
Nardia geoscyphus. Plectocolea cf. obovata (Nees) Mitt. 7525, 4f. Found only
Cephalosia biscuspidata (L.) Dum. 7258, 4c; 7265, 4f; 7300, in a peat bank community. I (Persson) had, with
lOa; 7340, 4e; 7386, lOa; 7522, 4f; 7526, 4f; 7528, 4f; hesitation, identified this specimen as P. obovata. This
7564, 13a; 7688, 4f; 7670, 4f; 7981 (cf.), 12a; 7.978, 4e; specimen was examined by Doctor Amakawa (written
8233, 4e; 8235, 13b; 8262, 4f. One of the most common commun., 1968), who expressed his opinion that is must
leafy liverworts on the island; grows in a wide variety belong to this species.
of habitats. It is the principal component of the pendant
bryophyte colonies of a peat bank shown in figure 14. LEPIDOZIACEAE
Cephalozia teucantha Spruce. 7242, 4c; 7258, 4c; 8281, 14b. Bazzania tricrenata (Wahlenb.) Trevis. 7227, 4c. 7341, 4c.
Found on the summits of moss mounds, intermixed with Found only on moss mounds in the Empetrum heath.
Dicranum elongatum, D. groenlandic-um, Microlepidozia Microlepidozia maJcinoana (St.) Hatt. [Lepidozia t~icho-
makinoana, Cephalozia bicuspidata, Anastrophyllum oladus K. Mull. Mierolepidozia silvatica (Evans) Jnrg.l.
sphenoloboides, and Sphenolobus minutus; also found on 7242, 4c; 7261a, 4g; 734^, 4c; 7343, 4a. Found only in the
bedrock outcrops.
Empetrum heath.
Cephalosia media Lindb. 7261a, 4c; 8087, 4e; 8091, 4e; 8275,
4e. Found on shaded peat banks, intermixed with Mnium LOPHOCOLEACEAE
glabrescens, Calypogeia miilJeriana, and Peltigera canina.
CepJialozia pleniceps (Aust.) Lindb. 7874, 4f; 7899, 12a; Chiloscyphus pallescens (Ehrh.) Dum. 7177, 9a; 717?, 9a;
8264, 4f. Found on peat banks and in a mixture of leafy 7202, 4g; 728S, 9a; 7391, 8a; 7857, 4e. Forms mats in
liverworts from a fell-field community. shallow rapid streams with Brachythecium asperriwum;
also grows on frost scars with Scapania paludosa, and
CEPHALOZIELLIACEAE along pool banks in the Empetrum heath.
Cephalosiella arctica Bryhn & Douin. 7300, 4g; 7345, 2b; Lophocolea cuspidata (Nees) Limpr. 7352, 2b; 7353, 2b;
7640, 3b ; 7641, 3b ; 7642, 3b ; 7643, 3b ; 7644, 3b ; 7673, 4f. 7639, 3b; 7644, 3b; 7863, 13a. Grows as pure colonies or
Grows as dense poMers on sandy banks above the sea- intermixed with Plagiotheeium roeseanum on sandy
shore and in wet crevices of breccia sea cliffs, intermixed banks above the seashore; also in crevices of wet breccia
with Bryum lapponicum, Schtetidium maritimum, Barbu- sea cliffs, and with Rhytidiadelphus squarrosus and
lacylindrica, and Bartramia itJiyphylla. Radula polyclada in a stream margin at Bird
LOPHOZIACEAE G. acutiloba, and G. marginata, all of which range from the

**Anastrophyllum sphenoloboides Sehust. (See Sdiuster, 1969. arctic to temperate regions in the Northern Hemisphere.
They are all polymorphic and related so closely to each
p. 741.) 7242, 4c; 7243, 4c; 7258, c. per., 4c. Found only
other that G. acutiloba and G. marginata could be regarded
at the summits of small moss mounds, intermixed with
as varieties or subspecies of G. inflata." He added (1966,
Dicranum groenlandicum, Cephalozia leucantha, and
p. 113), "However, most forms including the type specimens
Microlepidozia makinoana. This species was represented
of G. marginata and G. montana are rather definitely dif-
in the entire material by only a few stems scattered
ferent from G. inflata in the following features: T/te plants
through tufts of other bryophytes. I (Persson), with
are less flaccid, the leaves are broader and more or less
hesitation, identified it as Lophozia porphyroleuca
concave (sometimes even canaliculate), and the leaves are
(Nees) Schiffn.; the leaves were similar to those of the
rather regularly arranged (leaves on a shoot spread at
latter species, and the trigons of the leaf cells were very
similar angles) and dorsally secund. These differences seem
prominent. Later, I found a fertile stem with a perianth
to be sufficient to make G. marginata specifically distinct
in one of the specimens (No. 7258). Dr. N. Kitagawa
from G. inflata."
(written commun., 1968) examined this specimen and
wrote, "Shacklette No. 7258, which was identified by you The Amchitkan Gymnocolea (the first report of tMs genus
as Lophozia porphyroleuca, seems to be very different in the Aleutian Islands) has the broad, more or less concave
from the latter species in the brownish coloration of the leaves of the Japanese form. With the form that has been
plants, the rounded leaves (not longer than wide), and called G. acutiloba, it shares the only important character
especially in the paroicous inflorescence. I have never be- of this taxon the fairly acute leaf lobes. It is noteworthy
fore seen such a species. I think that it is a new species that Hattori (1958) described a variety acuta of G. montana
related to Isopaches bicrenatus, from which it differs which Kitagawa (1966) placed as a synonym of G. mar-
chiefly in the female bract and the perianth mouth. How- ginata but otherwise did not describe. The Aleutian form
ever, the material is too scanty to describe a new differs from both G. acutiloba, and G. marginata in the dis-
species." tinctly stronger walls, especially at the corners, of its leaf
Several weeks later, Dr. Kitagawa, while examining cells. The broad leaves, as found in the Amchitkan material,
specimens in Dr. Sinske Hattori's herbarium, discovered are occasionally found also in European specurens; the
that the Amchitkan hepatic was a species of Anastrophyl- only leaf of Gymnocolea inflata which Macvicar (1960)
lum which Dr. Rudolph M. Schuster had discovered illustrated in his well-known hepatic flora of Great Britain
earlier in northern Greenland. Dr. Schuster (written could as well have been taken from a flgxire of the Japanese
commun., 1968) examined the fertile specimen (Shack- Gymnocolea. The floras also commonly note that the leaves
lette No. 7258) and confirmed Dr. Kitagawa's identifica- can be concave and even canaliculate. Because th^ habitat
tion of it. of G. inflata ranges from xerophytic to very hytôphytic,
*Gymnocolea inflata (Huds.) Dum. forma. 7888, 4c; 75^0, the species is extremely variable. Apart from the more
4b ; 7564, 13a. On wet areas in the Empetrum heath, inter- commonly used handbooks and floras, the work of Jb'rgensen
mixed with Oncophorus wahlenbergii, Calypogeia tri- (1934) on the hepatics of Norway is recommended, for Ms
chomanes, and Diplophyllum albican-s, and in drain chan- treatment of the ecology and taxonomy of G. inflata. Un-
nels at an altitude of 650 feet. fortunately, this outstanding study of northern hepatics is
In my first report of these collections from Amchitka written in Norwegian and, therefore, is too littT e known
Island (Persson, 1968), I referred them to Gymnocolea outside Scandinavia.
acutiloba (Kaal.) K.M. After a renewed study of the ex- Gymnocolea inflata is widely distributed in the Northern
tremely polymorphous Gymnocolea inflata complex, I have Hemisphere; it is found, among other places, in Siberia,
formed the opinion that G. acutiloba represents only a Greenland, and Iceland. In North America it occurs through-
modification of G. inflata; this opinion is shared with out the continent; its northernmost known legality in
several other taxonomists. Alaska is Mount McKinley National Park, and it is also
The Amchitkan form seems to resemble even more reported from Yukon Territory.
closely the form that Kitagawa (1966), in his monograph In summarizing, I consider the Gymnocolea inflata com-
of Japanese Lophoziaceae, called G. marginata (St.) Hatt, plex to be extremely plastic, and I believe that o^her taxa
under which he placed the entire material of Japanese in the genus probably will prove to be environmental modi-
Gymnocolea and stated that this taxon is endemic to Japan fications. Experiments in the growth of these taxa on sub-
(Hokkaido, Honshu, and Kyushu). The synonyms that he strates of different pH, possibly in water culture, would be
gives are of interest: Sphenolobus marginatus St., 1917; of great interest, especially because G. acutilola is gen-
Cephalozia montana Horik., 1932; Gymnocolea, montana erally regarded as one of the so-called copper mouses.
(Horik.) Hatt, 1948; Gymnocolea inflata (Huds.) Dum. Lophozia alpestris ( Schleich.) Evans, 7524, 4f; 7564, 13a ;
subsp. montana (Horik.) Hatt., 1958; and Gymnocolea 7593, 13a; 7665, 6a; 8282, 13b; 8288, 13b; 8*34, 13b;
montana (Horik.) Hatt. var. acuta Hatt. According to 8235, 13b. Commonly occurs in alpine meadows or snow-
Kitakawa (1966, p. 113), the species grows "on muddy bed communities, but it was also found in a wet sedge
soil and mires at high altitudes." meadow with Pellia neesian-a and on a peat bank.
I have studied good material of the Japanese Gymnocolea Lophozia cf. groenlandica (Nees) Bryhn. 7424, 4b; 7425,
and, with Kitagawa, have observed that it is possible to 4b; 752^, 4f; 7525, 4f; 7528, 4f; 8018, 14b ; 8202, 5b. Most
find forms in some specimens which, according to Kitagawa common on the low plateaus but also found en a high-
(1966, p. 113). ". . . are quite similar to G. inflata, another altitude bedrock outcrop. No. 7525 was named by Dr.
polymorphic species." Kitagawa (1966, p. 111-112) also Kitagawa, who also studied No. 7528 and agreed with
stated, "Gymnocolea contains only three species, G. inflata, me (Persson) on its identification. Kitagawa (written
commun., 1968) wrote that the specimens "agree well 7978, 4e; 8016, 14b; 8019, 4d; 8280, 14b. Grows in a wide
with [R.M.] Schuster's description of the species, except variety of habitats, usually as single stems internixed
for the somewhat larger leaf cells." with other bryophytes.
*Lophosia incisa (Schrad.) Dum. (See Persson, 1968.) 7879, Diplophyllum taxifolium (Wahlenb.) Dum, 7853,12a. Found
4f; 8018, 14b; 8019, 14b; 8285, 8c. Found on peat banks only in a volcanic rubble fell-field.
and andesite rock outcrops at both high and low altitudes. Macrodiplophyllum plicatum (Lindb.) H. Perss. (See Pers-
**Lophozia opacifolia Culmann. (See Persson, 1968.) 8233, son, 1968.) 8010, 4d; 8016, 4d; 8017, 4d; 8020, 4d. Found
13b; 8234, 13b; 8235, 13b. Found only on wet soil by a on an andesite bird-perch mound at an altitude o* 950
stream in an alpine meadow, intermixed with Lophozia, feet.
alpestris, Nardia geoscyphus, and Cephalozia T)icuspidata. Scapania irrigua (Nees) Dum. 7482, 4b; 8264, 9a. Grows on
Lophozia cf. icenzelii (Nees) Steph. 7556, 14a; 7568, 12a. wet peat banks and in small drain channels in peat.
(Specimen 7568 was identified by Dr. Kitigawa.) Found Scapania paludosa K. Mull. 7202, 4g; 7217, 13a; 721?, 5c;
in patches of turf on a solifiuction terrace and on a high- 7225, 4b; 7226, 4b ; 7459, 8c; 7589, 13a; 7864, 4e; 8212, 8a.
altitude bedrock outcrop. Occurs at wet locations on the Empetrum heatl ; at
Orthooaulis Mnsteadii (Kaal.) Buch. 7338, 4c; 7340, 4c; places forms mats in small pools that support Siphula
7341, 4c; 7424, 4b; 7425, 4b; 8281, 14b. Found in the ceratites; also in snow-bed communities.
Empetrum heath, usually intermixed with many other Scapania scandica (Arn. & Buch) Macvic. 7265, 4f; 7524,
bryophytes on moss mounds, and on a high-altitude bed- 4f; 7525, c. per., 4f; 7879, 4f; 7999, 12a. Found in a peat-
rock outcrop. bank community, associated with Mnium glabrescens,
OrthocauUs floerkei (Web. & Mohr) Buch. 8262, 4f. Found Cephalozia atnbigua, and Lophozia cf. groenlandica-; on
only on a peat bank above a small spring at Chitka Cove. an almost bare peat and soil bank with Pogonatum
SphenoloMs mimttus (Crantz) Steph. 1241, 4c; 7242, 4c; urnigerum, Cephalozia bicuspidata, and Nardia scalaris;
7244, 4c; 1341, 4c; 8281, I4b. Found on moss* mounds in and in a fell-field community.
the Empetrum heath and on high-altitude bedrock out- Scapania suoalpina (Nees) Dum. 8293, 7b. Found orly in
crops. a rock-bottomed pond; grows submersed with Limosella
**SpJieno]o1)us saccatulus (Lindb.) K.M. 1906-14 [Junoerman- aguatica.
nia minuta var. grandis Gottsche Mss.; J. riaida Lindb. Scapania undulata (L.) Dum. 7287, 9a; 7288, 9a; 7390. 8a;
j8 grandis Lindb. 1879; J. saccatula Lindb. 1883; Cephalo- 7856, 4e. Found in water of small pools with Nardia
ziopsis saccatula Schffn. 1893; Sphenolobus minutus var. scalaris and Chiloscyplius pallescens, and in streams with
grandis S. Arn. & O. Martensson 1959]. 7424, 4a. Found Brachj/thecium asperrimum and Chiloscyphus pallescens.
in the Empetrum heath in tufts' of Pogonatum alpinum,
intermixed with Lophosia cf. oroenlandica, Mylia taylori, Order METZGERIALES
and Orthocaulis olnsteadii. This is the first report of the
occurrence of this species in North America. ANEURAOEAE
Tritomaria quinquedentata (Huds.) Buch. 7569, 12b; 7567a, Riccardia latifrons Lindb. 7338, 4c; 7455, 5a. On a small
12b. Occurs as small round polsters on loose unstable moss mound, with Ptilidlum ciliare, Mylia taylori,
soil among the pebbles of solifiuction slopes. Odontoschisma elongatum, and Orthocaiilis binsteadii;
also found in a bog with Sphagnum magellanicum, 8.
MARSUPELLACEAE
squarrosum, and S. teres. Dr. E. R. B. Little (written
Gymnomitrion corallioides Nees. 7554, lla; 7594, 14b; Erd- commun., 1968) wrote, "No. 7338 is good Riccwdia
man 612, lla. Found as single polsters on almost bare, latifrons, and I am almost sure that No. 7455 is the same
rocky soil of the discontinuous heath habitat and on high- species."
altitude rock mounds. Riccardia pinguis (L.) Gray. 7185, 8a; 7199, 4e; 7593 13a.
Marsupella emarginata (Ehrh.) Dumort. 7220, 8a; 7234, Occurs in very wet locations at pool and stream margins
5c; 7567a, 12b; 7569, 12b; 7854, 4e. Found as extensive and in drains of snow beds.
colonies in a Philonotis-Parnassia bog, and common in Riccardia simiata (Dum.) Trevis. 7201, 4g. Found on wet
smaller quantities at many other sites. mineral soil of a frost scar in the Empetrum heath. The
determination of this specimen was confirmed by
PTILIDIACEAE
Dr. E. R. B. Little and also, with some hesitation by
PtiUdi-um ciliare (L.) Nees. 7338, 4c. Found at only one lo- Dr. M. Mizutani.
cation on a moss mound. Surprisingly, this conspicuous
liverwort was not found at other locations; apparently, BLASIACEAE
it is actually rare on the island. Blasia pusilla L. 7667, 4f. Found only in a peat-bank com-
munity with Nardia scalaris, Dicranella suoulata, and
RADULACEAE
Cephalozia, Mcuspidata.
Radula polyclada Evans. 7863, 13a; 7864, 13a. Found only
at the margin of a small stream at Bird Cape, with DILAENAOEAE
Rhytidiadelphus squarrosus and Lophocolea cuspidata. Pellia neesiana (Gottsche) Limpr. 7564, 13a; 7665, 6a.
Found in wet sedge meadows with Lophozia alpestris and
SCAPANIACEAE in snow-bed communities with Sphagnum papillosum, S.
Diplophyllum albicans (L.) Dum. 7241, 4f; 7334, 4g; 7392, squarrosum, Anthelia julacea, Mylia anomala, and other
8a ; 7540, 4b ; 7567, 12a ; 7569, 12b; 7593, 13a; 7900, 12a ; bryophytes.
Order MARCHANTIALES Oalliergon sarmentosum (Wahlenb.) Kindb. 8047, 4e. Found

on a concrete spillway at a large spring near Makarius
CONOGEPHALACEAE Bay.
ConocepJwilum conicum (L.) Dum. 7198, 4e. Found only in Campylium stellatum (Hedw.) C. Jens. 7975, 4e; 8212, 8a.
streamside communities on sheltered banks in the Empe- Submerged in a small pool in the Empetrum heath, grow-
triim heath; easily identified in the field by its spicy odor ing with Drepanocladus revolvens and Scapania pnludosa;
after it has been crushed. also found at the margin of a small stream in tl > heath.
MAROHANTIACEAE Cratoneuron fllicinum (Hedw.) Spruce. 7636, la; ??44, 3b;
7645, 3b; 7647, 3b; 7983, 3b. Found on dripping s«a cliffs,
Marchantia polymorpha L. 7285, 3b; 7402, 8a; 7604, lOb. commonly with Vlota phyllantha, Bryum lapponteum,
Not common; found only on the mud bottom of a drained Schistidium maritimum, and Platydictya jungermannio-
pool, in an open steel barrel, and on thin soil of coastal ides; also found at other sites on the seashore.
cliffs. Drepanocladus fiuitans (Hedw.) Warnst. 8284, 6?. Found
SPHAGNOBRYA only in wet moss-lichen mats with Polytrichum tongise-
SPHAGNACEAE tum, on the lower terrace at Cyril Oove.
Drepanocladus revolvens (Sw.) Warnst. 7219, IFa; 7226,
Sphagnum capillaceum (Weiss) Schrank. 7214. 4a. Found 5c; 7567a, 12a; 7975, 13b; 8212, 8a. Occurs at rool mar-
only near the base of a moss mound in the Empetrtim gins, in streams from snow beds and alpine rreadows,
heath. and in fell-fields on the mountain summits.
Sphagnum compaotum Lam. & DC. 7590,13a ; 7676, 8c; 7872, Drepanocladus uncinatus (Hedw.) Warnst. 7188, lib; 7189,
4f; 7970, 5a ; 7992, 12b; 8236, 13a. Found in the drain of a lib ; 7249, 14a ; 7327, 4c ; 7358, 2c; 7380, lOc; 75<>7a, 12b;
snow bed, and on wet soil in the Empetrum heath. 7887, 12a; 7973, 15b; 8006, 4d; 8007, 4d. Abundant in
Sphagnum girgensohnii Buss. 7662, 6a; 7858, 4e. A principal many different habitats and distributed throughout the
component of bogs on the low plateaus. island.
Sphagnum lindoergii Schp. 8237, 13b; 8238, 13b. Found only
Hygrohypnum oestii (Ren. & Bryhn ex Ren.) Holz. ex
in small streams of the alpine meadows.
Broth. 7178, 9a; 7621, 9a; 8048, 9a. Found only at the
Sphagnum magellanicum Brid. 7453, 5a; 7455, 5a; 8236,13b.
margins of, or submersed in, swift streams on the low
A component of bogs on the low plateaus; also found in
plateaus, commonly growing with Hygrohypnum ochra-
small streams of alpine meadows.
ceum, Brachytheciwn asperrimum, and Chiloscyphus
Sphagnum papillosum Lindb. 7564, 13a; 7661, 6a; 7663, 5a;
pallescens.
7672, 4f; 7969, 5a. Most common at high altitudes in
drains from snow beds, but also found on wet peat banks Hygrohypnum ochraceum (Turn, ex Wils.) Loes-ke. 7178,
and wet sedge meadows of the low plateaus. Dr. Elsa 9a; 8269, 9a. Found associated with the spec'es listed
Nyholm kindly determined specimen No. 7969. above; also found with Bryoerythrophyllum recurviros-
Sphagnum squarrosum Sw. ex Crome. 7452, 5a; 7564, 13a; trum on a limonite deposit in a small stream.
7661, 6a; 8293, 7b. A principal component of bogs on the Platydictya jungermannioides (Brid.) Crum. [Amolyste-
low plateaus; also found in wet sedge meadows and in gietta sprucei (Bruch) Loeske]. 7180, 3b; 7182, 3b; 7646,
snow-bed communities. 3b; 8275, 4e. Occurs in crevices and under small rock
Sphagnum subsecundum Nees ex Sturm. 7215, 4a. Found shelves of breccia sea cliffs, at two locations growing with
only near the base of a moss mound in the Empetrum Mielichhoferia macrocarpa, (Shacklette, 1967) ; also found
heath. on soil at the margin of a small stream in the heath.
Sphagnum teres (Schimp.) Angstr. ex C. Hartm. 7454, 5a; AULACOMNIACEAE
7663, 6a; 8260, 4e. Abundant in bogs and wet sedge mea-
dows of the low plateaus; usually bright green in color. Aulacomnium palustre (Hedw.) Schwaegr. 7231, 5c; 7233,
5c; 7402, 8a; 7482, 6a ; 7663, 6a; 7850, 6a. A common moss
ANDREAEOBRYA in Sphagnum bogs, wet sedge meadows, and liverwort
mats in streams.
ANDREACEAE
BARTRAMIAOEAE
Andreaea rupestris Hedw. 7192, lib; 7256, 14a; 7363, 2c;
7591, 13a; 7893, 12b; 8280, 14b. Found at all altitudes; Bartramia ithyphylla Brid. 7641, 31). Found only on thin soil
usually grows on rocks, but at places in the mountains it over breccia sea cliffs.
forms mats in streams of alpine meadows and in snow-bed Conostomum tetragonum (Hedw.) Lindb. 7594, 14b; 8231,
drains (Shacklette, 1966). 12a. Found as small polsters on unstable rocky soil and
bedrock mounds at high altitudes.
EITBRYA Philonotis americana Dism. 7441, lOb, 7457, 5a; 7482, 6a;
8201, 5b. Found in a wet sandy borrow pit, near Makarius
AMBLYSTEGIACEAE Bay, where it formed a dense mat over a large area; also
AmUystegium serpens (Hedw.) B.S.G. 8203, 5b; 8204, 3d. found on wet soil of a roadside ditch and in a wet sedge
Found on a turf bank above the beach near Makarius Bay, meadow.
growing with Leptobryum pyrifonne and Bryum steno- Philonotis fontana (Hedw.) Brid. 7202, 4g; 7203, 4g; 7217,
trichum; also found in a wet sandy 'borrow pit with Phi- 5c. On wet disturbed mineral soil, commonly forming
lonotis americana. large mats.
PLANT SPECIES OF AMCHITKA ISLAND 4-5
BRACHYTHECIACEAE first collected in Alaska on Sitkalidak Island by Eyerdam
(Persson, 1968) not on Amchitka Island, as erroneously
**Brachythecium acutum (Mitt.) Sull. (See Persson, 1968.)
7380, lOc; 7850, 4a; 7373, 15b. Found in Empetrum mats stated by Shacklette (1967).
Polilia annotina (Hedw.) Loeske. 8258, 4f. Found only near
near Constantine Harbor, on broken macadam runways,
and on wet boards in a World War II building.
Constantine Harbor, on a peat and soil bank.
BrackytJiecium albicans (Hedw.) B.S.G. 7-Z89, lib; 7358, 2c;
PoMia cruda (Hedw.) Lindb. 7354, 2b; 1651, 3b; 8264, cf.,
1381, lOc; 1981, 4d; 8002, 12b; 8291, 4f. Commonly found
4f. Found near the seacoast on sandy soil, breccia clifs,
on sparsely vegetated rocky or sandy soil, with Rhacomit-
and peat banks.
rlum lanuginosum, R. ericoides, Drepanocladus uncinatus,
Polilia nutans (Hedw.) Lindb. 7300, lOa; 7672, 4f; 7899,
12a; 7901, 12a; 8018, 14b. Found on soil and peat banks at
and Ceratodon purpureus, but may also be found in other
habitats.
low altitudes, commonly growing with Pogonatum
urnigerum and Ditrichum heteromallum.
Brachythecium asperrimutn (Mitt.) Sull. 7-Z77, 9a ; 1224, 5c;
7287, 9a ; 7343, 4a ; 7380, lOc; 7456, 5a ; 7691, 9a ; 7860, 9a ; OLIMACIACEAE
7983, 9a. Occurs in a wide variety of habitats on the low
Climaciiim dendroides (Hedw.) Web. & Mohr. 7194, 4^;
plateaus.
7195, 4e. Found at only one location on wet soil by a
Eurhynchium praelongum (Hedw.) B.S.G. 1191, 4e; 1216,
small stream in the Empetrum heath. Apparently scare";
3a ; 7258, 4c; 7269, 3a ; 7270, 3a; 1211, 3a; 7343, 4a ; 7350,
this distinctive moss is very easy to recognize in the fieM.
2b ; 1351, 2b ; 7355, 2b ; 7382, lOc ; 7649, 3b ; 7902, 4e ; 7973,
15b; 7977, 4e. This species is widespread at low altitudes DICRANACEAE
and is usually abundant on sea cliffs. At places on the
AmpMdium lapponicum (Hedw.) Schimp. 1183, 3b. Fou^d
Pacific Ocean coast cliffs, it is the only bryophyte found.
only at Cyril Cove, growing below a breccia shelf with
BRYACEAE Pottia lieimii, a few feet above sea level.
*Aongstroemia longipes (Sommerf.) B.S.G. 8051, 6a. Fourd
*Anomo1)ryum concinnatum (Spruce) Lindb. [A. ftliformis
only in a wet sedge meadow near Makarius Bay.
(Dicks.) Solms var. concinnatum (Spruce) Loeske].
Arctoa fulvella (Dicks.) B.S.G. 7853, c. fr., 12a. In a high-
(See Persson, 1968.) 7670, 4f. Found only on a wet peat altitude fell-field, growing with Diplophyllum taxifoliv.m
bank near Constantine Harbor, with CepJialozia bicuspi-
and Rhacomitriutn brevipes.
data, Bryoerytliropliyllum recurvirostrum, Polilia nutans, Diohodontium pellucidum (Hedw.) Schimp. 1521, 4f. On a
Pogonatum urnigerum, and Bryum arcticum.
wet peat bank at Constantine Harbor, growing with
*Bryum arcticum (B. Br.) B.S.G. (See Persson, 1968.)
Cephalozia bicuspidata, Dicranella heteromttlla, and
7673, 4f. Found only in the community listed immediately
other bryophytes.
above.
Dicranella heteromalla (Hedw.) Schimp. 7524, 4f; 7526, 4f;
*Bryiim pattens Sw. sens. lat. 7855, c. fr., 4e. Found only
7528, 4f; 7877, 4f; 8050, c. fr., 5b. Commonly found on wet
in a streamside community about 100 feet above sea level
peat banks; also found on wet sandy soil near Cleveng-er
at Bird Cape, growing with Mnium glabrescens, Plecto-
Lake.
colea cf. obovata, Polilia nutans, and Scapania undulata.
Dicranella palustris (Dicks.) Crundw. ex E. F. Warb.
Bryum pseiidotriqtietrum (Hedw.) Gaertn., Meyers &
[Anisothecium squarrosum (Stark.) Lindb.]. 1201, 4<?r;
Scherb. 1199, 4e; 73^8, 2b. Found at the margin of a small
7205, 4g; 1218, 5c ; 7588, 13a ; 7859, 5c; 7984, 5c. Found in
stream in the heath, growing with Climaciiim dendroides
a wide variety of habitats that range in altitude from the
and Riccardia pinguis; also found on a sandy bank at the
low plateaus to snow beds.
seacoast.
*Dicran-ella subulata (Hedw.) Schimp. (See Persson, 196P )
*Bryum lapponicum Hag Kaur. [B. salinum Hag.] (See Pers-
7385, lOa; 7386, lOa; 7668, 4f; 8201, 5b; 8203, 5b. Tl is
son, 1968.) 7542, 3b; 7640, 3b; 7650, 3b. Apparently re- moss is fairly common on the island, and it is surprising
stricted to sea cliffs that are subjected to salt spray or that it has not been found earlier in the Aleutian Islands.
waves.
Dicranum angustum Lindb. 7453, 5a. Found only in a lo^v-
Bryum, stenotrich urn C. Mull. [B. inclinatum (Brid.) altitude bog, with Sphagnum magellanicum, S. squar-
Bland.]. 7281, 12a ; 7483, lOa ; 7565, 15a; 7648, 3b; 8035, rosum, and 8. teres.
lOc; 8206, 3d. This moss, with Tetraplodon mnioides and Dicranum etongatum Schleich. ex Schwaegr. 7258, 4c;
Pleurosium schreberi, grew on decomposed animal car- 7339, 4c; 7340, 4c; 8246, 4d. A common species at the
casses ; it grows also on mineral and organic soil on summits of moss mounds. It was found on one orgaric
Amchitka Island. bird-perch mound, where it was abnormally dark gre-^n
Leptobryum pyriforme (Hedw.) Wils. 8205, 3d; 8286, cf., and very robust.
4g. Found on a soil-covered sea cliff at Makarius Bay, Dicranum fuscescens Turn. 1316, 4c; 7333, 4ff- Found on
with Amblystegiuni serpens and Bryum stenotrichum; moss mounds, although it is not a common component of
also found on mineral soil of a frost scar. the mound flora, and on frost scars.
*Mielichlioferia macrocarpa (Hook, ex Drumni.) Bruch & Dicranum groenlandicurn Brid. 7242, 4c. The dominant moss
Schinip, ex Jaeg. & Sauerb. (See Persson, 1968.) 1181, on the summits of some small moss mounds.
3b; 1182, 3b; 1541, 3b; 8208, 3b. Shacklette (1967) dis- Dicranum hoicellii Ren. & Lesq. 7233, 4a; 7252, 14a; 7325,
cussed this "copper moss" as it occurs on Amchitka 4c; 7326, 4c; 7977, 4e; 8003, 12b. Very common in a wide
Island, where it is found on sea cliffs, growing only on variety of habitats. According to Persson (1968), prob-
narrow rock strata that have a greater concentration of ably all Aleutian collections named D. scoparium belong
metals than that found in adjacent strata. This moss was to this species.
Onoophorus wahleribergii Brid. 7223, 5c; 1/333, 4g; 75^0, 4b. HYLOCOMIACBAB
This moss forms hard, black conical polsters in wet Hylocomium splendens (Hedw.) B.S.G. 7208, 4c; 7228, 4a;
heath and in sedge meadows that can develop into small 7307, lOa; 73.72, lOa; 7329, 4c; 7331, 4c; 7649, 4c. Found
moss mounds 0.5 m in diameter and as much as 40 cm only on the low plateaus, usually mixed with ti e common
high. These polsters are saturated with water and freeze mosses of the heath, or on moss mounds; at places it is the
into hard icy lumps that one often stumbles over while dominant moss on the collapsed summits of old mounds.
walking on the otherwise resilient vegetation mat.
HYPNACBAB
DITBICHACBAB
Hypnum cupressiforme Hedw. (See Persson, 1P38.) 8007,
Ceratodon purpureus (Hedw.) Brid. 7280, lOc; 7348, 2b; 14b. Found only on an andesite bird-perch mound near
7359, 2c ; 7376, 14a; 7378, lOc ; 7402, 4b ; 7483, lOa ; 8289, Buoy Point, growing with Drepanoctodus uno'natus and
la. Common on compacted soil and on concrete and ma- Pogonatum alpinum.
cadam walks and roads ; also found on peat deposits and **Hypnum dieckii Ren. & Card. (See Persson, 1968.) 7850, 4a.
other organic substrates. Found only in the Empetrum heath near Consta ntine Har-
Ditrichum heteromallum (Hedw.) Brid. 7300, lOa; 8050, bor, growing with Aitlacomnium palustre, Brachythecium
c. fr., 5b. Found in a dense growth of Calamagrostis acutum, and Pogonatum alpinum.
nutkaensis on a roadside bank of mineral soil, and on Hypnum plicatulum (Lindb.) Jaeg. 8002, 12b. Found only
wet sandy soil in a PMlonotte-Parnassia community. at the margin of a vegetation mat on a solifluction terrace.
ENTODONTACEAE Ptilium crista-castrensis (Hedw.) DeNot. 7230, 4a; 7306,
4a. Usually sparse, but at several locations on the Empe-
Pleurozium schreberi (Brid.) Mitt. 7309, 4a ; 73-77, 4c ; 7328, trum heath it was observed to form almost pure colonies
4c; 7514, 15a; 7663, 6a; 7664, 6a. Very abundant as a 5 sq m or more in size.
component of the Empetrum heath mat.
LBUCODONTACEAB
FONTINALACEAE
Antitrichia curtipendula (Hedw.) Brid. 7209, 4c; 7210, 4d;
Fontinalis neomexicana Sull. & Lesq. 7176, 9a; 7240, 9a; 7248, 14a ; 7250, 14a; 7313, 4c ; 7318, 4c; 732?, 4c; 7330,
7447, 9a; 8207, 9a. Grows submersed in swift clear 4c ; 7331, 4c; 73/A 2b ; 7890, 12b ; 8002,12b; 8008, 4d. Com-
streams of the Empetrum heath. The "fronds" are as mon, and at places very abundant, on the Empetrum
much as 60 cm long, and some of them fruit abundantly. heath ; also found on bird-perch mounds and on solifluction
GBIMMIACBAB terraces. Its yellow color makes it conspicuous as the
summit moss on many "mature" moss mourns, visible
Rhacomitrium aciculare (Hedw.) Brid. 7186, 8a. Only one even on aerial photographs (fig. 8). W. A. Weber (oral
polster was found; it was growing with Riccardia pinguis commun., 1967) suggested that in the treeless Aleutian
on the bottom of a small pool that had been drained. Islands moss mounds substitute for trees in providing the
Rhacomitrium brevipes Kindb. ex Mac. 7853, 12a; 8267, 12a. elevated habitat characteristic of this moss throughout
One of the two polsters found had formed a dense cylin-
its range.
drical cushion about 8 cm high in a fell-field community,
where it had entrapped much windblown silt. MNIACBAE
Rhacomitrium ericoides Brid. [R. canescen-s (Hedw.) Brid. Mnium glabrescens Kindb. (See Persson, 1968.) 72616, 4e;
var. ericoides (Brid.) B.S.G.] 7357, 2c; 736-7, 2c; 7279, 7388, lOb; 7402, 8a; 7483, 4a; 7524, 4f; 758(7, 4f; 7528
lOc; 7666, lOa; 7890, 12a. Not common; found on sand 4f; 7862, 4e; 8264, 4e; 8284, 6a. A common species on many
dunes near the coast (Shacklette, 1966), on hard soil different substrates throughout the island.
of a gravel road, and in a fell-field community. Mnium insigne Mitt. 7261o, 4g; 8288, 4f. Foun-l on stony
Rhacomitrium fasciculare (Hedw.) Brid. 7373, 14a; 7508, soil of a frost scar hi the Empetrum heath and on a shaded
3b; 7556, lla; 7873, 4f; 800.7, 12b; 8880, 14b. Found on peat bank.
sea cliffs, low- and high-altitude rock mounds, and soli- Mnium pseudopunwtatum Bruch & Schimp. (See Persson,
fluction terraces. 1968.) 7196, 4e; 7203, 4g; 7260, 4e; 7458, 5a. F<nnd only at
Rhacomitrium lanuginosum (Hedw.) Brid. 7.790, lla; 7.79-7, low altitudes, usually on very wet soil and in bogs.
lib; 7207, 4c; 7241, 14a; 725.7, 14a; 73.74, 4c; 73.75, 4c;
7362, 2c; 7372, 14a ; 7568, 12b; 7889, 12a; 7892, 12a ; 7901, ORTHOTBICHACEAB
12a; 7990, 12a; 8020, 4d. Very abundant throughout the Ulota phyttantha Brid. 7271, 3a; 7273, 3c; 7364, 2c; 7507,
island. Probably initiates mound formation more com- 3h; 7638, 3b; 7911, 3b; 8024, 4d. Found on rocks near
monly than any other moss on the island. the sea, and, at one location, on an organic bird-perch
Schistidium apooarpum (Hedw.) B.S.G. [Grimmia apocarpa mound. A facultative halophyte.
Hedw.]. 7366, 2c. Unattached polsters of a spheroidal
shape that were composed of this species were described PLAGIOTHBCIACEAE
from Amchitka Island (Shacklette, 1966), growing on Plagiothecium roeseanum B.S.G. 7353, 2b. Found only on
a pebbly sand dune near Cyril Cove. sandy banks above the beach at Cyril Cove, growing with
Sohistidium maritimiim B.S.G. [Grimmia maritima Turn.]. Eurhynchium praelongum, Lophocolea cusfMata, and
7267, 3a ; 7272, 3a ; 7274, 3c; 7507, 3c; 7649, 3b; 7650, 3b; Pohlia cruda.
8025, 3a. Apparently a strict halophyte; grows only on Plagiothecium undulatnm (Hedw.) B.S.G. 7227, 4a; 8239,
rocks that receive sea spray or waves. 13b; 8273, cf., 4f. Occurs sparingly as single strands in
the moss mat of the Empetrum heath and in alpine *Tortula mucronifolia Schwaegr. (See Persson, 1968.) 75SV,
meadows. 2b; 7531, 2b; 7532, 2b; 8247, 3b. In loose sand of dun?s
near the coast and on breccia sea cliffs.
POLYTRICHACEAE
Oligotrichum Jiercynicum (Hedw.) Lam. & DC. 8240, 13a. RHYTIDIACEAE
Found on mineral soil in the bed of a stream that flowed Rhytidiadelphus loreus (Hedw.) Warnst. 7204, 4g; 7228, 4a;
from a snow field. 7232, 4a; 7308, 4a; 7327, 4c; 7664, 6a. Abundant, especially
Oligotrichum parattelum (Mitt.) Kindb. 823%, 13a. Found in wet locations in the Empetrum heath.
only on soil by a stream that flowed from a snow bed at Rhytidiadelphus squarrosus (Hedw.) Warnst. 7344, 2Ii;
Chitka Cove. 7863, 13a. On sandy soil near the seacoast, growing among
Poffonatum alpinum (Hedw.) Rohl. 1229, 4a: 7252, 14b; dense stools of Elymus, and in a snow-bed community.
1311, 4a; 7384, cf., 4f; 7402, 8a ; 7425, 4b ; 7523, 4f; 7841, Rhytidiadelphus triquetrus (Hedw.) Warnst. 7309, 4a. A
4f; 7850, cf., 4a; 8006, cf., 14b; 8257, 4f. Found on low common species in the moss mats on the low plateaus.
terraces of the Empetrum heath, usually growing in the
SPLAOHNACEAE
thick moss mat, and on high-altitude bedrock outcrops.
At one location it grew with Marchantia polymorpha on Tetraplodon mnioides (Hedw.) B.S.G. 75^4, 15a; 7880, 15a;
the peat bottom of a drained pond. 7968, 15a; 8242, 15a. Entirely restricted to decomposed
Pogonatum contortum (Brid.) Loeske. (See Persson, 1968.) carcasses of birds and rats; it forms dense polsters tint
7206, 4f. Found only on the vertical walls of a narrow fruit profusely.
deep trench that had been dug in peat during World II.
THUIDIAOEAE
Poffonatum urnigerum (Hedw.) P. Beauv. 7261, 14a; 7265,
4f; 7300, lOa; 7355, 2b; 7356, 2c; 7385, lOa; 7481, lOa; Claopodium crispifolium (Hook.) Ren. & Card. 8274, ê-
7672, 4f; 8201, 5b; 8202, 5b; 8213, lOa; 8214, lOa. This Found only on soil at the margin of a stream in the heath
moss commonly grows on disturbed soil and peat banks near Cyril Cove.
(flg.2).
Polytrichum commune Hedw. 7332, 4c; 7377, 4f. On moss ANNOTATED LIST OF LICHENS
mounds and peat banks of the low plateaus.
Polytrichum formosum Hedw. 7241, 4f; 7299, 4f; 7381, cf., By WILLIAM A. WEBER,' JAMES A. EBDMAN, and HILDUR KEO?*
lOc. On peat banks and broken macadam pavement. Shown below is the only published list of lichens that grow on
Polytrichum juniperinum Hedw. 7374,14a. Apparently rare; Amchitka Island, and all species on the list are new records for
found only on an andesite outcrop at an altitude of about this island, unless otherwise noted. To our knowledge, the only
400 feet. former Amchitkan collections of significance are those made
Polytrichum longisetum Brid. [P. yracile Sm.]. 8284, 6a. by Eric Hulten on July 9, 1932, and those of George A. Llano
Found only in wet moss-lichen mats on the lower terrace made on August 22, 1947. Inventories of lichens collected frcm
at Cyril Cove, with Drepanocladus flultans. the Aleutian Island chain were published by Hedrick (1936) and
**Polytrichum sphaerothecium (Besch.) Broth. [Pogonatum Degelius (1937).
sphaerothecium Besch.]. (See Persson, 1968.) 7529, 4f. Duplicates of specimens cited in this list that were collect?d
This is the first collection of this moss in Alaska, and by Hansford T. Shacklette in November and December 1965
apparently the second collection in North America. Persson were identified by Krog. All subsequent collections were identi-
(1968) made the observation that this moss, in its world- fied by Weber, except those of Stereocaulon determined by I.
wide distribution, is often associated with Bryoxiphium Mackenzie Lamb, Harvard University, Cambridge, Mass.; sev-
norvegicum and that these mosses are found only in non- eral packets of Umbilicaria sent to George A. Llano, National
glaciated regions of the world. Science Foundation, Washington, D.C.; and Parmeliopsis am-
Polytrichum strictum Menz. ex Brid. 7335, 4c. Found only Mgua identified by Mason E. Hale, Jr., Smithsonian Institution,
on a small moss mound. Washington, D.O. Identification of the rocks on which the
POTTIACEAE saxicolous lichens grew was made by W. D. Quinlivan, U.S.
Geological Survey.
Barbula cylindrica (Tayl. ex Mack.) Schimp. ex Boul. 764^, Nomenclature follows that of Hale and Culberson (1966) in-
3b;-7642, 3b; 7644, 3b. Found only on breccia cliffs of the sofar as possible. Collection numbers, in italics, are those of
seacoast. Shacklette, unless otherwise specified; they follow the species
Bryoerythrophyllum recurvirostrum (Hedw.) Chen. 7671, name. The habitat and plant community from which a species
4f; 8270, 9a; 8276, 3a. Found growing with Anomobryum was taken, designated by numbers and letters, as set forth
concinnatum, Pohlia nutans, Pogonatum urnigerum, and in the section "Description of the Island and Its Vegetation" of
Dicranella subulata on a peat bank at Constantine Harbor this report, follow the collection number.
and in a spring on a deposit of limonite, where it was
heavily incrusted with iron compounds. CLADONIACEAE
Pottict heimii (Hedw.) Fiirur. ex Hampe sens, lat, 7184, Baeomyces placophyllus Ach. Brdma/n 646, 15b. On a wood°n
3b; 8203a., 5b; 8204, 3b. Found growing with Amphidium sill of a World War II hut above Constantine Harbor.
lapponicum on breccia sea cliffs at Cyril Cove and on Cladonia alpestris (L.) Rabenh. Hutchison s.n.; 8023, 4d;
sandy soil of a borrow pit near Makarius Bay, associated Erdman 613, 12b. Hutchison (1937, p. 144-145) reported:
with Philonotis americana. It also grew on turf banks at
Makarius Bay, associated with Aniblystegium serpens, 3 University of Colorado Museum, Boulder, Colo.
Leptobryum pyriforme, and Brytim stenotrichum. * Botanical Museum, Oslo, Norway.
''The yellow-grey Cladonia alpestris, one of the reindeer from a crustose to semi-fruticose growth due to sand
mosses, formed the groundwork of the moors [of Am- abrasion and deposition."
chitka], into which the foot crunched as if walking on Lecanora allophana (Ach.) Rohl. [L. subfusca (L.) Ach.,
snow. It was mixed with empetrum * * *." She un- in part]. Erdman 632, 15b. On a small plank in the margin
doubtedly was referring to the dominant Cladonia pa- of a wet sedge meadow.
cifica, recently described by Ahti (1961) in his treatment Lecanora atra (Hnds.) Ach. Erdman 641, 2c. C ~\ a small
of the reindeer-lichen complex. Cladonia alpestris occurs 'boulder in a sand-dune blowout.
widely scattered over the heath mat, often in patches, but Lecanora oasta<nea (Hepp) Th. Fr. Erdman 67f, 4e. With
it is less abundant than C. pacifica. bryophytes in a streamside community.
Cladonia amaurocraea (F16rke) Schaer. Erdman 653, 4c; Lecanora straminea (Wahlenb.) Ach. Erdman 630-B, 3g.
Erdman 666, Sc. Uncommon ; found on a few moss mounds On andesite beach rocks.
and at the edge of a tundra pool. OchrolecMa friffida (Sw.) Lynge. Hulten s.n. (Degelius,
Cladonia arbiiscula subsp. Tjeringiana Ahti [C. sylvatica 1937, p 126) ; 7277, 3d; 7555, 14a; 7597, 14b; 7895, 12a;
(L.) Hoffm.]. St evens s.n.; 7539, 4c. On the summit of a 7898, 12a; Erdman 565, 4a; Erdman 566, 4a; Erdman
tundra mound above Cyril Cove. Merrill (1929, p. 42) 610, lla; Erdman 663, 14b; Erdman 673, lOc. Occurs in
stated that "D. H. Stevens found the species [C. sylvatica a wide range of habitats, at places overgrowing the heath
subsp. sylvestris Oed.] at Amehitka and Ilac Islands, mat. Degelius (1937, p. 126) stated that it "seems to be a
and it has come to hand from Tanana crossing." No habi- very common species on these [Aleutian] islands." This
tat was given. taxon is highly polymorphic.
Cladonia bellidiflora (Aeh.) Schaer. 7245, 4c ; 7332, 4c; 1513, Placopsis gelida (L.) Linds. [Lecanora gelida (L.) Ach.].
4a; 7596, cf., 14b; 7602, 14b; 8012, 4a; Erdman 595, 8b. Hulten s.n. (Degelius, 1937, p. 125) ; 7£97, 14a; Erdman
Widespread but not abundant; found in a variety of 596, lOc. Common, especially on loose cobb^s of old
habitats. roadbeds.
Cladonia coccifera (L.) Willd. 7301, 15d; 7510, 4a; 7511, LECIDEACEAE
4a; 7512, 4a; Erdman 620, 15d. Fairly uncommon; how-
ever, it is more abundant and conspicuous in burned Bacidea sp. Erdman 643, 14a. On andesitic tuff breccia rocks.
areas than elsewhere. Lecidea cf. aleutica Degel. Erdman 629, in part, 14a. On
Cladonia degenerans (Fl6rke) Spreng. [C. cerasphora andesite sill or flow boulders.
Vain.]. 7303, 15d; Erdman 608, in part, 4f. Uncommon Lecidea armeniaca (DC.) Fr. Erdman 592, 14b. Common
throughout the Empetrum- heath. on high-altitude basaltic rocks above Chitka Cove.
Lecidea elegantior H. Magn. Erdman 589, 14b. On high-
Cladonia gracilis (L.) Willd. 7302, 15d; 7505, 4a; 7669, 4f;
altitude basaltic rocks above Chitka Cove. Not listed by
Erdman 608, in part, 4f; Erdman 609, in part, 4f; Erdman
Hale and Culberson (1966) ; first reported for North
617, 15d ; Erdnian 618, 15d. Occurs in a variety of habitats
America by Weber and Viereck (1967).
on the low plateaus, but is common only in burned areas.
Lecidea flavocaerulescens Hornein. Erdman 587, 14b. Com-
Cladonia pacifica Ahti [C, impexa Harm.]. 7210, 4c ; 7323,
mon on high-altitude basaltic rocks above Chitka Cove.
4c ; Erdman 568, 4a. The dominant lichen in the Empetrum
Lecidea macrocarpa (DC.) 'Steud. [L. platycarpa Ach.;
heath; forms pure clumps in surface depressions, thus
L. steriza (Ach.) Vain.]. Erdman 590, 14b; Erdman 597,
giving a dotted appearance to the ground surface as seen
lOc ; Erdman 639, in part, 14a; Erdman 642, 2c. On breccia
from the air (fig. 8).
rocks, extending from sea level to high-altitud? fell-fields.
Cladonia puxidata (L.) Hoffm. 7290, lOc; Erdman 619-A, Lecidea cf. symmicta (Ach.) Ach. Erd-man 604, 15b. Found
15d. Sparse, except in burned areas, where it is abundant. on the wind-eroded wooden frame of a World War II
Cladonia rangiferina (L.) G. H. Web. ex Wigg. Erdman
hut.
582, 13b. Observed only on a thick Empetrum mat in a Lecidea vernalis (L.) Ach. Erdman 603, 15b ; cf. Erdman 667,
small valley above Chitka Cove; apparently uncommon. 4g. On tar-paper roofing of a hut at high altitude and in
Cladonia scabriuscula (Del. ex Duby) Leight. 7211, 4c; 7324, a frost scar of the Empetrum heath.
4c; Erdman 624, 4d. A component species of moss mounds. Mycoblastus alpinus (Fr.) Kernst. [M. sanguinarius (L.)
LECANORACEAE Norm., in part]. 7597, 15b; Erdman 566, 4a; Erdman 611,
lla; Erdman 652, 4c. Occasionally found intermixed with
HaemaJomma ventosum subsp. lapponicum (Has.) Laund. OchrolecMa frigida.
[H. lapponicum Has.]. Erdman 586, 14b. On a high-alti- Rhisocarpon atroalbescens (Nyl.) Zahlbr. Erdman 591, 14b.
tude rock outcrop. Common on high-altitude basaltic rocks ab^ve Chitka
Icmadopliila, ericetorum (L.) Zahlbr. Erdman 633, in part, Cove.
4d. At the base of an organic bird-perch mound. RMzocarpon geographicum (L.) DC. Hulten «.«. (Degelius,
lonaspis epulotioa var. arctica (Lynge) H. Magn. Erdman 1937, p. 117) ; Erdman 661, 14b. On an andesite outcrop of
639, in part, 14a; Erdman 645, 4g. On a latite boulder a scree ridge above Chitka Cove.
above Constantine Harbor and on basalt andesite scoria RMzocarpon hochstetteri (Korb.) Vain. Hulten s.n. (Degel-
rocks in a frost scar. ius, 1937, p. 117) ; Erdman 629, in part, 14a; Erdman 639,
Lecanora aleutica. H. Magn. ex Hedrick. 7365, 2c; Erdman 14a. Apparently common on rock outcrops throughout the
630-D, 3a. Common on boulders, primarily on those along island.
the beaches. I. Mackenzie Lamb examined specimen 7365. Toninia lobulata (Somin.) Lynge. Erdman 640, 41 This
which grew on a sand dune, and wrote (written commun., lichen was found covering the peat banks of an old gun
1968), "Apparently a Lecanora species that has gone emplacement.
PANNARIACEAB Pertusaria hultenii Erichs. Erdman 588, 14b. Saxicolous;
Pannaria pezizoides (G. Web.) Trev. 7843, 4f; 7865, 4f; on basaltic outcrop above Chitka Cove.
Erdman 601, 4f. Collected from soil or peat banks at both Pertusaria sp. Erdman 571, 15b. Corticolous; on a utility
ends of the island. pole.
Psoroma hypnorum (Vahl) S. Gray. Erdman 633, in part, PHYSCIACEAE
4d. At the base of an organic bird-perch mound.
Buellia punotata (Hoff.) Mass. [B. pullata Tuck.], Erdman
PARMELIACEAE 630-C, in part, 3a. On andesite beach boulders.
Cetraria ciliaris Ach. [C. orbata (Nyl.) Fink]. Erdman 572, Physcia caesia (Hoffm.) Hampe. 7276, 3c; Erdman 630-C,
15b. On a felled utility pole; possibly an adventive species in part, 3a. On sea cliffs and beach boulders.
introduced during World War II occupation. Physcia dubia (Hoffm.) Lett. 7276, 3c. Found growing with
Cetraria cucullata (Bell.) Ach. Erdman 656, 14a. Found P. caesia.
near the summit of a large rock mound. Rinodina turfacea (Wahlenb.) Korb. 7982, 4d; Erdman 671,
Cetraria ericetorum Opiz [C. crispa (Ach.) Nyl.; C. island- 4d. A nitrophilous lichen found on a few organic bird-
ica var. crispa (Ach.) Nyl.]. Erdman 581, 13b; Erdman perch mounds.
615, 14b. Observed and collected only above Chitka Cove; SPHAEROPHORAGEAE
apparently uncommon.
SphaeropJiorus globosus (Huds.) Vain. Hulten s.n. (lege-
Cetraria niaricans (Retz.) Nyl. 7897, 12a. In the rubble of a
lius, 1937, p. 107) ; 7246, 4c ; 7320, 4c; 7595,14b ; 7599, 14b ;
high-altitude rock field.
7897, 12a ; 7991, 14b; 8014, 4d ; 8015, 4d; Erdman 569, 4a;
Hypogymnia enter omorplia (Ach.) Nyl. 7282, 15b. On a
Erdman 583, 13b. Degelius (1937, p. 107) described this
bridge timber near Cyril Cove; possibly introduced with
species as very common throughout the Aleutian Islands.
construction materials.
It is the dominant lichen on the summits and ridge^ of
Hypogymnia pJiysodes (L.) Nyl. [Parmelia physodes (L.)
the hummocks in the Empetrum heath.
Ach.; P. duplicata var. doughlasiocla Gyeln.]. Erdman
570, 15b. On a felled utility pole. S TEREOCAULAOEAE
Hypogymnia subobscura (Vain.) Poelt [Parmelia sub- Stereocaulon alpimim Laur. 7360, 2c. On a sand dune above
obscura Vain.]. 7897, 12a. In the rubble of a high-altitude Cyril Cove.
rock field, mixed with several fruticose lichens. Stereocaulon intermedium (Sav.) H. Magn. Erdman 599,
Parmelia alpicola Th. Fr. Erdman. 601, 14b. On boulders lOc. Common on a gravel roadbed south of Gonstartine
at high altitudes. Harbor. I. Mackenzie Lamb (written conimun., 1968)
Parmelia omphalodes (L.) Ach. 7603, 14b; 7888, 12a; Erd- considered it to be "a stunted, subcrustose state of 8.
man 655, 4c. On rocks at high altitudes. intermedium."
Parmelia saxatilis (L.) Ach. 7277«, 3c; Erdman 606, 15b; Stereocaulon tomentosum var. alpestre Flot. [8. tom-entosum
Erdman 621, 14a. On rock outcrops above or on the beach, Fr.]. Erdman 600, lOc. Associated with 8. intermedium
and on wooden supports of a World War II shed. on a gravel roadbed south of Constantine Harbor.
Parmelia sulcata Tayl, 7275, 3c; Erdman 605, 15b; Erdman Stereocaulon vcsuvianum Pers. [8. denudatum Plk.-; 8.
625, 14a. On breccia outcrops above or on the beach, vesuvianum var. denudatum (Florke) Lamb]. Erdman
and on wooden supports of a World War II shed. 612, 12b. On upland slopes above the Pacific Ocean, op-
Parmeliopsis ambigua (Wulf.) Nyl. Erdman 631, 15b. On posite Chitka Cove.
a small plank at the edge of a wet sedge meadow.
Platysmatia lacunosa (Ach.) Culb. & Culb. [Cetraria lacu- STICTACEAE
nosa, Ach.]. 7601, I4b. On a rock mound above Chitka Lobaria, linita (Ach.) Rabenh. [Sticta pulmonaria var.
Oove. linita, (Ach.) Tuck.]. Hulten s.n. (Degelius, 1937, p. 108),
PELTIGERACEAE Erdman 636, 4a. At the base of a heath bank above Cyril
Cove.
Peltigera apJitJwsa (L.) Willd. 7259, 4e; 7319, 4c; 7321, 4c;
Erdman 638, 14a ; Erdman 651, lla. A fairly common ter- TELOSCHI STAGE AE
ricolous lichen on shaded, often vertical, substrate sur- Blastenia sp. Erdman 630-C, in part, 3a. On andesite beach
faces. boulders.
Peltiaera canina (L.) Willd. [P. oanina var. membranacea Caloplaca aranulosa (Mull. Arg.) Jatta. Erdman 594 : 3a.
Ach.; P. membranacea (Ach.) Nyl.]. Hulten s.n. (Degelius A conspicuous saxicolous crustose lichen; common on
1937, p. 110) ; 7261c, 4e; Erdman 623, 4d; Erdman 634, supralittoral beach boulders and sea stacks.
4a; Erdman 669, 2b. A common terricolous lichen on Caloplaca sp. (Ferruginea group). Erdman 668, 2b. Or the
shaded, often vertical, substrate surfaces. end of a large driftwood log.
Peltiaera malacea (Ach.) Funck. Erdman 635, 4a. At the XantJwria, candelaria (L.) Th. Fr. [TeloscMstes ca*tdel-
base of a heath bank above Cyril Cove. ariits (L.) Fink]. 7369, 4d; Erdman 627, 14a; Erf man
Peltiaera, spuria (Ach.) DC. [P. canina var. spuria (Ach.) 630-C, in part, 3a; Erdman 649, 14a. On rock outcrops
Schaer.]. 7637, la. Found on the soil of a second beach fertilized by birds.
terrace at Constantine Harbor. Xanthoria elegans (Link) Th. Fr. [Caloplaca. elegans
(Link) Th. Fr.]. Erdman 637, 3a. Occurs on beach boul-
PERTUSARIAOEAE ders and sea cliffs. Tatewaki and Kobayashi (1934, p. 6)
Pertusaria coriacea (Th. Fr.) Th. Fr. Erdman 2, 14a. On stated that "This saxicolous association is developed on
a flow breccia outcrop. such precipitous cliffs that the vascular plants cannot
grow. The beautiful red colour of Caloplaca elegans maura, the calcivorous [sic] lichen, is distribrted over
adorns the monotonous gloomy rock surface." the supralittoral rocks, giving a dark aspect to this belt."
UMBILICABIACBAE SOME ALGAE AND OTHER PROTISTS

Umbilicaria cylindrioa (L.) Del. [Gyrophora cylindrica
(L.) Ach.]. 7367, 14a; Erdman 628, 14a. On an andesite By H. SKUJA S
boulder pile near Cyril Cove. Although Llano (1950) cited
no collections from Amchitka Island, he did state (p. Collections of algae from three types of habitat on
11) that, in the Aleutian Islands, U. cylindrico, is "the Amchitka Island were sent to me for identificat:on. The
dominating Umbilicaria on all exposed lava rocks, in habitat descriptions and the annotated list, including
association with U. proboscidea, U. hyperborea, and U. descriptions of new taxa, follow.
torrefacta * * *." The following taxa, found in a liverwort polster
Umbilicaria proboscidea (L.) Schrad. 73670, 14a; 7506, 2c;
7607, 14b; 7894, 12a; Erdman 585, 12a; Erdman 602, (mostly Marsupella emarginata) , were collected (No.
14b; Erdman 650, 14b; Erdman 660, 14a; Erdman 664, 7234} by H. T. Shacklette, November 22, 1965, from a
14a. Common on rocks in a wide variety of habitats. small stream near Chitka Point:
USNEACBAE CYANOPHYCEAE
Alectoria cf. irvingii Llano. 8013, 14b- Found on an ande- Chroococcus varius A. Br.
site bird perch on a high peak north of Buoy Point. We Gloeocapsa magma ( Breb.) Kiitz.
have not compared this collection with the type from ScMzothria caloicola (Ag.) Gom.
Anaktuvuk Pass, but it appears to agree well with the ScMzothriae lardacea (Ces.) Gom.
type description that is, it resembles Comicularia di- Scytonema mirabile (Dillw.) Born.
vergens, medulla Pd-, with long fissural pseudocyphellae. Stigonema minutum (Ag.) Hass.
Alectoria nigricans (Ach.) Nyl. 7596, 14b. Mixed with Cla- Stigonema ocellatum Lyngb.
donia bellidiflora at the base of an andesite rock mound
south of Chitka Cove. DESMIDIACEAE
Alectoria oehroleuca (Hoffm.) Mass. 7193, 14a; 7368, 14a; Actinotaenium cucurbita (Breb.) Telling. Cells28/u-30/t long,
7553, 14a; 7553, 14a; 7886, 12a; Erdman 567, 4a. A con- 13p-15p broad. (PI. 1, fig. 1.)
spicuous fruticose lichen that apparently is of wide eco- Cosmarium cyclicum Lund. var. arcticum Nordst.
logical amplitude, for it occurs as scattered clumps on Cosmarium decedens (Reinsch) Racib. Cells 45^-48/t long,
breaks in the Empetrum heath, as well as on rocks ait the 24/4-26/t broad, and 17/X-19/H thick; breadth of isthmus
highest altitudes on the island. 18/X-20/*. (PI. 1, fig. 7.)
Alectoria pubescens (L.) R. H. Howe [Parmelia lanata Cosmarium obliquum Nordst. Cells 20^-23/11 long, 17/i-18/t
(L.) Wallr.]. 7371, 14a; 7857, 12a; Erdman 657, 14a. On broad, and 13^-14/t thick; breadth of isthmur* 14/t-16jtt.
rocks in outcrops or rubble fields, in many places mixed (PI. 1, figs. 8-9.)
with other fruticose lichens. Cosmarium tatricum Racib. Cells 4lM-52p lonf, 23/i-27j«.
Comicularia divergens Ach. [Alcctoria divergens (Ach.) broad, and 17/*-20/i thick; breadth of isthmus 18/t-21/t.
Nyl.]. 7247a 14a; 7598, 14b; Erdman 614, 14b; Erdman (PI. 1, figs. 4-6.)
647, 4a; Erdman 654, 4c; Erdman 659, 14a. Widespread, Cosmarium variolatum Lund. Cells 45^-52/1 long, 25/i-27/u
but most common on rock mounds and outcrops. broad, and 17/*-20/i thick; breadth of isthmus 12/t-15/i.
Ramalina almquistii Vain. 7370, 14a; 7906, 3c; Erdman 626, (PI. 1, figs. 2, 3.)
14a. On breccia cliffs and inland outcrops. Cylindrocystis brebissonii Menegh.
Ramalina scoparia Vain. 7078, 3c; 7505, 3c. Commonly Cylindrocystis brebissonii var. minor W. et G. S West.
occurs with R. almquistii. Cylindrocystis crassa De Bary.
Siphula ceratites (Wahlenb.) FT. 7266, 8c; 7675, 8c.; Erd- Mesotaenium micrococcum Kiitz.
man 665, 8c. Forms dense colonies along the edges of Staurastrum punctulatum Breb. var. pygmaeum (Breb.)
small pools. W. et G. S. West. A quadrangular form. Length 36/t-40M,
Thamnolia vermicularis (Sw.) Ach. ex Schaer. IT. sub- breadth 34/t-40/t; isthmus 18/i-20ju broad; cell wall uni-
uliformis (Bhrh.) W. Culb.]. 7257, 14a; 7559, 14a; Erd- formly granulate, granules minute and acute.
man 672, 4a. Common throughout the island in a wide
variety of habitats but is best developed in the uplands, DIATOMACEAE
where it forms pure stands in the eroded Empetrum Achnanthes spp.
heath. All material fluoresces under ultraviolet light. Caloneis silicula (Ehrnb.) Cl.
Epithemia turgida (Ehrnb.) Ktitz.
VERRUCARIACEAE
Eunotia arcus Ehrnb.
Verrucaria maura Wahlenb. ex. Ach. Erdman 630-A, 3a. Eunotia gracilis (Ehrnb.) Rbh.
A dominant but inconspicuous lichen on the rocks of the Eunotia lunaris (Ehrnb.) Grun. var. capitata Gr^n. Length
beaches. In describing the plant communities of the 50/i-86/*, breadth 4/t.
Aleutian Islands, Tatewaki and Kobayashi (1934) placed Eunotia praerupta Ehrnb. var. muscicola Boye-Pe4:.
Verrucaria in the supralittoral belt, which is just above
the mean high-water level, where the plant is washed by 5 Uppsala Universitets Institution fOr Systematisk Botanik, Uppsala,
occasional high tides. They stated (p. 6), "Verrucaria Sweden.
Eunotia praerupta var. bidens (W. Sm.) Grun. Cells 15/4- slightly convex, rarely conspicuously depressed. (PI. 1,
75/4 long, 5/4-10/4 wide; striae 10-12 in 10/4. fig. 12.)
Eunotia robusta Ralfs var. diadema (Bhrnb.) Ralfs. Cosmarium nitidulum De Not.
Frustulia rhomboides (Bhrnb.) D.T. var. saxonica (Rbh.) Cosmarium subcrenatum Hantzsch forma. Cells 30/4-33/4
D.T. long, 25/4-26/4 broad, and 17.5/4-18.5/4 thick; isthmus 11/4-
Gomphonema angustatum (Kiitz.) Rbh. var. prod<uctum 12/i broad; semicells subtrapezo-rectangular, with flat-
Grun. Length 38/4-45/4, breadth lO/i-12/4. tened 4-crenate ends and 4-5 crenae at each side; the low
Melosira distans (Bhrnb.) Kiitz. var. alpigena Grun. frontal swelling above the isthmus with generally 5,
Breadth 7/4-9/4, height of the whole cell 10/4-12/4; striae rarely 4 or 6, strong rows of 3-5 angular granules.
obsolete, valvae conspicuously dotted. (PI. 1, fig. 11.)
Pinnularia gibba Bhrnb. Cosmarium subpachydermum Schmidle forma. Cells 38/«^-41/4
Pinnularia lata (Br6b.) W. Sm. Length 85/4-103/4, breadth long, 28/4-32/4 broad, and 20/4-22/4 thick; isthmus 13/4-14/4
28/4-30/4; striae 4 in 10/t. broad; cell wall scrobiculate, but without a refractive
Pinnularia microstauron (Bhrnb.) Cl. thickening in the center of each semicelL (PL 1, fig. 14.)
Pinnularia viridis (Nitzsch) Bhrnb. var. rupestris Compare also C. pseudonitidulum Nordst.
(Hantzsch) Cl. Cosmarium subundulatum Wille. Cells 60/4-65/4 long, 42ju--45/i
broad, and 28/i thick; isthmus 20ju-22/i broad; semicells
Tabellaria flocculosa (Roth.) Kiitz.
high semicircular with a 10-undulate margin; membrane
DINOPHYCBAB distinctly punctato-scrobiculate with a large but low
lenticular inflation in the center of each semicell. (PI. 1,
Gloeodinium montanum Klebs. Diameter of cells with in-
tegument 41/4-58/4, without integument 25/4-38/4; integu- fig. 10.)
ment or cell wall thick and lamellose, colorless or yellow-
Cylindrocystis brebissonii Menegh.
ish or brownish; cell content brown; 2-celled aggregates Cylindrocystis crassa De Bary.
Euastrum ansatum Ehrnb. Cells 78ju-87ju long, 39/4-44/4 wide,
with integument 57/4-60/4X60/4-85/4. (PI. 1, figs. 20-23.)
and 27/t-29ju thick; apex 20ju-22/4, isthmus 13/4-15ju broad.
Peridinium cinctum (O.F.M.) Ehrnb. Euastrum Udentatum Nag. Cells 53ju-61/t long, 24/4-26/t
The following taxa were found in a shallow pool on broad, and about 20ju thick; apex 22ft, isthmus 10/4-12ju
andesite bedrock at a quarry near Banjo Point, with broad.
Euastrum denticulatum (Kirchn.) Gay. Cells 33/4-35/t long,
Isoetes sp. (44 chromosomes). Collected by H. T. Shack- 22ju-26/i broad, and 15ju-17ju thick; apex 16ju-20/4, isthmus
lette, July 28,1967. 5/4-6/4 broad.
Euastrum denticulatum var. quadrifarium Krieger. C^lls
OYANOPHYCBAE
about 20/t long, 160 broad, and 10/t thick; apex 13/4, isthmus
Aphanothece castagnei (Br6b.) Rbh. 4/t broad.
Oscillatoria amoena (Kiitz.) Gom. Trichomes straight, with Euastrum didelta Ralfs. Cells 125/i-135/t long, 64/t-66/t bread,
more or less attenuated and frequently curved ends, 4/4-6/4 and 37ju-39ju thick; apex 30ju-31ju, isthmus 20/t-21/4 broad.
broad, not constricted or only slightly constricted at the Euastrum elegans (Br6b.) Kiitz. A typical form; c^lls
crosswalls, and grayish blue green or olivaceous and 30ju-32ju long, 20/1-21/4 broad, and 12/t thick; isthmus 5/t-6/t
pale violet; cells nearly quadrate or slightly shorter, or broad.
longer than broad, granulated at dissepiments; end cells Euastrum oblongum (Grev.) Ralfs.
capitate, with broadly conical apex. (PI. 2, fig. 25a-d.) Euastrum pectinatum Br4b. Cells 74/t-76/t long, 46/t-48/t
Oscillatoria- amphibia Ag. Trichomes nearly straight or broad, and 34/4-35/t thick; width of the polar lobe 27/4-?9ji4,
coiled, 2/1-2.5/4 broad, ends nonattenuated and not capitate, width of the isthmus 19/i-21/t.
not constricted at the joints, and pale blue green; cells Hormidium flaccidum A. Br. Filaments «=> 4/4-5/t broad, not
4/4-7.5/4 long, with two granules at the septa; end cells constricted at the crosswalls; cells 2-3% times longer
rounded. than wide; chloroplast a unilateral plate with a pyrenoid.
Synechococcus maior Schrot. Cells oblong or ellipsoidal with Mougeotia sp. ster.
rounded apices, 19/4-21/t broad, and 24/4-34/4 long; contents Oocystis parva W. et G. S. West. Solitary or in families of
homogeneous and blue green. 2-4 cells, inclosed by the enlarged mother cell wall; c^lls
ellipsoidal or fusiform with pointed poles, 3/i-7/t bread,
CHLOROPHYCBAB s.l. 7/4-10/t long; chloroplasts 1-2 parietal discs, usually with-
Chlorella elUpsoidea Gerneck. out pyrenoids. (PI. 1, fig. 15.)
Chlorella vulgaris Beyerinck. Pediastrum braunii Wartm. Coenobia 7- to 10-celled, cells
Cosmarium humile (Gay) Nordst. 10/4-16/t in diameter.
Cosmarium laeve Rbh. forma. Cells 24/t-26/4 long, 17/4-18/4 Scenedesmus acutus (Meyen) Chod.
broad, and 1V-12/4 thick, with 4/4-5/t broad isthmus; cer- Scenedesmus ecornis (Ralfs) Chod.
tain specimens have one semicell of the type, the other Scenedesmus spinosus Chod.
one of var. septentrionale Wille. (PI. 1, fig. 13.) Scotiella nivalis (Shuttlew.) Fritsch. Cells solitary,
Cosmarium margaritiferum Menegh. ellipsoidal, 12/4-14/4 broad and 20/4-24/4 long; cell vail
Cosmarium meneghini Br6b. var. concinnum Rbh. Cells. colorless, with 6-8 longitudinal ribbons or low wirgs;
25/4-27/t long, 19/4-20/t broad, and 13/4-14/4 thick; apex single parietal chloroplast with one pyrenoid.
13/4-14/4, isthmus 4/4-5/4 broad; cell ends truncate or often Zygogonium ericetorum Kiitz.
DIATOMACEAE Pinnularia appendiculata (Ag.) Cl. Cells 28/4-33^ long and

Caloneis silicula var. longissima Schirschow. Cells linear, 4/1-6/1 broad; striae about 16 in 10/i.
with an evident widening in the middle, 80/i-85/j long and Pinnularia brevicostata Cl. Cells 83/i-92/i long and1 13^-15^
lO/i-14/i broad; striae about 10 in 10/i. broad; striae 8-9 in 10/j.
Caloneis silicula var. alpina Cl. Cells oblong, with con- Pinnularia divergens W. Sm. Cells 58/i-66(U long anc1 15/i-17/i
spicuously triundulate sides, 32^-37/j long and 12/j-15,u broad; striae 10-12 in 10/*.
broad. Pinnularia gentilis (Donk.) Cl. Cells 200ju-205/i long, 34/i-
Cymbella aequalis W. Sm. Cells 17^-25^ long and 36/i broad; striae 6-7 in 10/i.
broad; striae 13-15 in lO/i. Pinnularia gibba Ehrnb. Cells 65/I-90/I long, lO/i-lSa broad;
Cymbella, angustata (W. Sm.) 01. striae about 10 in 10/i.
Cymbella angustata var. hybrida (Grun.) R. Ross. Pinnularia interrupta W. Sm.
Cymbella cuspidata Kiitz. Cells 50^-62^ long and 15 Pinnularia lata (Breb.) W. Sm. Cells 80fj.-85fi long and 30/i-
23/j broad; striae 10-13 in 10/j. 33/* broad; striae 3 4 in lOû.
Cymbella delicatula Kiitz. Cells 25/^-36^ long and Pinnulariamaior (Kiitz.) Cl.
broad. Pinnularia microstauron (Ehrnb.) Cl. Cells 34^-72/i long
Cymbella gracilis 01. Cells 56/i-60/i long and 8ju-10,u broad; and Sju-ISM broad; striae 10-12 in 10/i.
striae 12-13 in 10,u. Pinnularia microstauron forma biundulata O. MiHl. Cells
Cymbella hebridica (Greg.) Grun. Cells 29/t-37/i long and S-ifi-72/j. long and 8/*-13/i broad; striae 10-12 in IQu.
7{j.-8/ji broad; striae 10-12 in 10/i. Pinnularia viridis (Nitzsch) Ehrnb.
Cymbella naviculiformis Anersw. Cells 38^-50^ long and Pinnularia viridis var. sudetica (Hilse) Hust.
12ju-16,u, broad; striae 14-18 in 10/i. Stauroneis anceps Ehrnb. Cells 48/*-57/* long and 10/*-13)U
Cymbella perpusilla A. 01. Cells 16^-20^ long and broad; striae about 20-23 in 10/i.
broad. Stauroneis anceps forma linearis (Ehrnb.) Cl.
Cymbella pusilla Grun.
Cymbella tumidula Grun. Cells 28/J-32/J long and CHRYSOPHYCEAE
broad. (Only cysts were found.)
Cymbella ventricosa Kiitz. The following taxa were found in a tundra lake about
Eunotia diodon Ehrnb. Cells 52,u-70/j long and 10^-15^ wide ; 50 cm deep near Cyril Bay, with Isoetes sp. (66 chromo-
striae 12-14 in 10p.
Eunotia pectinalis (Kiitz.) Rbh. var. minor (Kiitz.) Rbh.
somes) . Collected by H. T. Shacklette, July 28,1967.
Cells 10|U-17/j long and about 4/t wide ; striae 15-18 in 10/i. CYANOPHYCEAE
Eunotia praerupta var. bidens (W. Sm., Grun. Cells 15/J-75/J
Lyngbya rivulariarum Gom.
long and S/i-lO/i wide; striae 10-12 in 10/t.
Merismopedia elegans A. Br.
Eunotia robusta Ralfs var. diadema (Ehrnb.) Ralfs. Cells
Merismopedia glauca (Ehrb.) Nag.
47,u-50ju long and 20^-21/1 broad, with 6-undulate dorsal
Nostoc microscopicum Carm.
margin; striae about 10 in 10/t.
Oscillatoria amoena (Kiitz.) Gom.
Eunotia septentrionalis Ostr. Cells 200-23,11 long, 4^-5/1 Oscillatoria amphibia Ag.
broad; striae 17-18 in 10/i. Oscillatoria facilis Skuja, sp. nov.
Fragilaria construens (Ehrnb.) Grun. var. binodis (Ehrnb.) Trichomata solitaria vel in strata expansa, tenuia,
Grun. arachnoidea, olivacea vel sordide fusco-viridia agglomerata,
Fragilaria construens var. subsalina Hust. aut plusminusve recta aut subflexuosa et elongata, ad geni-
Fragilaria nitssohioides Grun. cula leniter constricta, apice breve attenuata e4; paullo
Frustulia rJiomboides var. saxonica (Rbh.) D.T. Cells 50^- curvata, l/u-1.3/u crassa; cellulis diametro trichomatis 2-4
105/t long and 13,u-200 broad. plo longioribus, protoplasto homogeneo sed cum cl romato-
Gomphonema angmtatum var. productum Grun. plasmate a centroplasmate sat limitato, pallide griseo-
Hantzschia amp7iioxys (Ehrnb.) Grun. aerugineo vel olivaceo et subfusco; cellula apicali elongate
Melosira distans var. alpigena Grun. Breadth 6î-10(H, height conoidea, superne parum obtusata. (PL 2, fig. 26a-e.)
of the whole cell 50-17,u; striae 16-18 in 100; valvae Trichomes single or forming a flat arachnoid, ol'vaceous
conspicuously dotted. or brownish-green thallus, more or less straight or slightly
Navicula contenta Grun. Cells 80-140 long and 20-30 broad. flexuous, slightly constricted at the cross-walls, l/j-1.3/*
Navicula cryptocephala Kiitz. Cells 250-400 long and 50-80 broad, and short at the ends but clearly attenuated and
broad; striae 16-18 in 10/i. curved; cells 2-4 times as long as wide, with a homogeneous,
Navicula minima Grun. Cells 10^-16^ long and 3/i-4^ broad. pale-grayish-blue-green or olivaceous and brownish proto-
Navicula minima var. atomoides (Grun.) 01. Cells lOî-16/i plast ; end cell elongate, conoid with slightly obtusa te apex,
long and 3/t 4/t broad. not capitate and without a calyptra. (PI. 2, flg. 26«-e.)
Neidium afflne (Ehrnb.) 01. var. minus Cl. 38^-45^ long and
This species resembles O. neglecta Lemm. but 1 as both
curved and attenuated apices, longer cells, and a different
8^-10^1 broad; striae 20-23 in 10,u.
color of thallus and trlchomes; compare also O. exit's Skuja
Neidium iridis (Ehrnb.) 01. var. amphigomphus (Ehrnb.)
(Skuja, 1964, p. 51, table 7, figs. 1-3), which is considerably
V.H. Cells 7(V-130/* long and 28^-33^ broad; striae 16-18 thicker, more constricted at the crosswalls, and of a different
in WfjL. color. Also different are O. deflesea W. et G. S. West, O. sub-
Nitsscliiapalea (Kiitz) W. Sm. tilissima Kiitz., O. limnetica Lemm. with var. acicularis
Nitsschia stagnorum Rbh. Nygaard, and others.
Oscillatoria grossegranulata Skuja. brana tenuissima, perspicua; contentu cellularum pallidis-

Trichomes solitary among other filamentous algae, elon- sime aerugineo, fere achroo, homogeneo vel granulis subtilis,
gate, nearly straight or slightly flexuous, not attenuated or sparsis ornato; multiplicatio fit divisione transversal!.
curved at the ends, and distinctly constricted at the cross- (PL 2, fig. 24.)
walls, thus torulose, 5.5/J-6/* broad; cells cylindrical, 1-1% A forma genuina cellulis paullo minoribus, colore prl-
times longer than wide, 6/^-12^, rarely 16/*, long with clear- lidissimo primum differt.
and pale-olivaceous or light-grayish-blue-green coarsely Cells solitary, mobile, single or two together after division,
granulated protoplast, and at times with granulated septa. short cylindrical or ellipsoidal with rounded apices, 2.5;*-
(PI. 2, fig. 28.) 3.5/i broad and 3.'5ju-6/i long; membrane delicate and hyaline;
Oscillatoria splendida Grev. contents very pale blue green, almost colorless, homogeneous
A very characteristic species, with straight or flexuous or with some minute granules; multiplication by transverse
trichomes 1.7/*-2/j broad which usually are not constricted at division. (PL 2, fig. 24.)
the crosswalls but are rather short and abruptly attenuated This variety is well characterized by its smaller size and
at the more or less curved ends; cells 2^1 times longer than the very pale protoplast.
broad, with often granulated septa and a clear blue-green Compare also S. notatus Skuja forma (Skuja, 1964, p. 24,
homogeneous protoplast; end cell very elongated and table 1, figs. 39-40), which is considerably larger, entirely
pointed, more or less bent, capitate at the apex, without colorless, and with a differently constituted protoplast.
calyptra. (PI. 2, fig. 27.) Synechococcus maior Schrot.
The thallus of this alga also has a characteristic intense
odor which resembles that of fertile soil. CHLOROPHYCEAE s.l.
Oscillatoria, sp. Ankistrod&smus braunii (Nag.) Collins var.
Trichomes nearly straight or slightly bent, unconstricted Cells may be fusiform but usually slightly curved or
at the joints and not attenuated or capitate at the ends, lunate with more or less acutely pointed ends, solitary or
0.5/4-0.7/* broad; cells mostly 2-4 times, rarely 5 times, as after division (forming of autospores) in pairs or in forrs
long as wide, with a homogeneous pale-blue-green protoplast contacting each other, 7/i-14/i long, 1/^-2^ broad; cell wall
but with 2 granules at the dissepiments; end cell simply thin, smooth, and colorless; chloroplast single, parietal,
rounded. usually unilateral and with a small rounded medial excava-
Only a few trichomes seen; probably a small form of tion, without a pyrenoid. (PL 2, fig. 49.)
0. amphibia. (PL 2, fig. 29.) Only a few cells were seen. Considerably smaller than the
Phormidium frigidiim F. E. Fritsch. typical A. braunii, but otherwise similar; compare w'th
Pseudanabaena catenata Lauterb. Vischer's (1920) description. Perhaps also it is a relative of
Pseudanabaena granulifera Skuja, sp. nov. some Koliella species (Hindak, 1963).
Trichomata solitaria, inter algas benthonicas sparsa, sat Ankistrodesnuts spiralis (Turner) Lemm.
brevia, plerumque 6-20-cellularia, 15/^-50^ longa, fere recta Chaetoplaca nom. nov. genericum. Syn. Chaetopedia Skuja
vel leviter arcuata, 1.5ju-1.8/i crassa, apices versus non 1948 non Pascher 1939. Fain. Chaetoplaeaceae nom. nov.
attenuata, ad dissepimenta hyalina sat crassa evidenter fam.
constricta, motu proprio praedita; cellulis 1-1% plo raro Chaetoplaca crassiseta Skuja var. puclla Skuja (Skuja,
2% plo longioribus quam latis, breve cylindricis, lateribus 1964, p. 110, table 16, figs. 29-33). Has characteristic 4-
baud concavis; protoplasto pallide olivaceo vel dilute and 8-celled colonies.
viridi-aerugineo, granulato, granulis sat grossis praecipue Chlamydomionas alaskensis Skuja, sp. nov.
peripheriter locatis; cellula apicali superne late conoidea. Cellulae rotundato ellipsoidales et ovoideae vel interdum
(PL 2, figs. 30-32.) fere globosae, 12^-18^-21(1 longae, 10/j-15/*-l9/i latae,
Trichomes solitary among other benthonic algae, short, fiagellis binis aut cellula e aequilongis aut paullo brevioril 'is
usually 6- to 20-celled and 15/j-SO/j long, mobile, nearly praed-tae; membrana modice crassa, plerumque distin"te
straight or slightly curved, 1.5ju-1.8/i broad, not attenuated porosa raro fere laevi, achroa, papilla antica parva n^n-
at the ends, distinctly constricted at the joints, dissepiments nunquam vix evoluta instructa; chromatophoro parietal!
hyaline, rather thick ; cells short-cylindrical l-l%-2% times crasso, bursaeformi, multipartite sed cum partibus inter se
longer than wide, with pale olivaceous or light-greenish-blue dense contiguis, pyrenoide destitutis; stigmate fuscorutro,
coarsely granulated protoplast, granules mostly peripheral ; med-ocri, breve baculiformi in parte anteriore; vacuolis
end cell at apex conoid. (PI. 2, figs. 30-32.) contractilibus binis, parvis in polo antico ad basin flagel-
This species perhaps resembles most nearly P. minuta lorum, nucleo nucleolato media in cellula sito. Multiplies tio
Skuja forma (Skuja, 1956, p. 70, table 6, figs. 27-28), from divisione protoplast! cellulae matricali in duas vel fre-
which it is distinguished by its considerably shorter and quenter quattuor cellulas filiales. (PL 2, figs. 33^11.)
smaller trichomes and differently colored, coarsely granu- Cells rounded ellipsoidal or sometimes almost globose,
lated protoplast. The typical P. minuta (Skuja, 1948, p. 57, 12ju-18/i-2V long and 10^-15/^-19^ wide, with two fiagella
table 5, figs. 6-7) is also closely allied but is broader and which are as long as or slightly shorter than the cell; cell
has more rounded cells usually with homogeneous or only wall moderately thick, distinctly porous or almost smooth,
finely granulated protoplast. Compare further P. galeata colorless, with anterior semicircular small papilla which
Bocher and P. bipes Bocher. sometimes are entirely lacking; chloroplast parietal, thick
Synechococcus cedrontm Sauv. var. pallidus Skuja var. nov. but divided into a number of separate closely joined bod! <?s,
Cellulae solitariae, motu proprio praeditae, singulae vel without a pyrenoid ; eyespot brownish red, small, short sub-
geminatae, breve cylindricae vel ellipsoideae, apicibus linear, in the anterior half of the cell; contractile valuoles
rotundatis, 2.5^-3/1 raro 3.5/i latae et 3.5^-6^ longae; mem- two, small, at the base of the flagella; nucleus more or less
centrally placed. Asexual reproduction by the division of Hortnidium flaccidum A. Br.

the cell protoplast usually into 4 or more, rarely 2, daughter Mougeotia sp. ster. .
cells or zoospores. (PI. 2, figs. 33-41.) Oedogonium sp. ster.
C. alasTcensis is in the subgenus Chloromonas. This species Pedict strum braunii Wartm.
is similar to C. vulgaris Anachin but is considerably smaller Planctococcus sphaerocystiformis Korschik.
and has a porous cell wall, a smaller papilla, and a higher Free-floating spherical colonies, 25ju-30/i in dirmeter; the
placed, shorter eyespot. It is also somewhat like C. jemt- gelantinous nonlamellate and colorless envelope relatively
landica Skuja but differs in being almost twice as large wide, including a central group of 4 globose cells, 9/j-12ju in
and in having a porous cell wall, shorter flagella, and a diameter. Cells have a thin smooth wall and a thin or only
higher placed sublinear eyespot. Should also be compared moderately thick parietal multipartite chloroplast covering
with some other chloromonads. most of the wall; and lack a pyrenoid. (PI. 2, fig. 47.)
Chlamydomonas subangulosa. Skuja sp. nov. Probably not a separate intraspeciflc taxon but only a
Cellulae subanguloso-ovales vel late-ellipsoideae, 13/i-16ju 4-celled form of P. sphaerocystiformis.
longae, 9/i-14/* latae, flagellis binis cellulae circiter aequi- Raphidonema gracile Skuja, sp. nov.
longis vel paullo brevioribus instructae ; membrana delicata, Cellulae solitariae vel post divisionem factern binatae,
achroa levisve, papilla antica mediocri hemisphaerica vel elongatae et cylindraceo-fusiformes, 1.7ju-2.3ju latae, 12^-25/u,
truncato-rotundata praedita ; chromatophoro poculiformi in longae, apicibus subito angustatis, polls subacuto rotundatis,
parte anteriore extus longitudinaliter striato-carinato, parte plerumque plusminusve curvatis; membrana tenui, achroa
postriore valde incrassato et pyrenoide depresso-globoso sat laevisve; chromatophoro parietali, plusminusve unilateral!,
rnagno donato; stigma te fuscorubro mediocri, rotundo vel sine pyrenoide sed cum excavatione ventral! med'a in cellula
elliptico in parte anteriore; vacuolis contractilibus binis in ubi nucleus locatus. Propagatio fit cellularum divisione in
polo antico ad basin flagellorum locatis; nucleo nucleolato duas partes. (PI. 2, fig. 48o-f.)
in excavatione chromatophori sito. Multiplicatio divisione Cells free floating, solitary, or in pairs after division,
protoplast! cellulae maternae plerumque in 4 cellulas flliales. elongate fusiform to somewhat cylindrical, with slightly
(PI. 2, figs. 42-46.) tapering and subacute rounded ends, usually more or less
Cells slightly rectangular or broad-ellipsoidal, 13ju-16ju curved, 1.7ju-2.3ju broad and 12/A-25//. long; cell wall thin,
long and 9/t-14/* wide, with paired flagella which are as smooth, and colorless ; chloroplast single, parietal, and more
long as, or slightly shorter than, the cell; cell wall thin, or less unilateral, without a pyrenoid but commonly with a
smooth and colorless, with anterior semicircular medium- rounded ventral sinus in the middle where the nucleus is
sized papilla; chloroplast cup shaped, striped on the outside located. Reproduction by a transverse division into two
of the anterior part, the posterior portion massive and with daughter cells that remain connected for some time. (PI. 2,
one rather large pyrenoid; eyespot brownish red, small, at flg.48«-/.)
the anterior end of the chloroplast. Asexual reproduction This form perhaps most nearly resembles F.oliella cor-
by division of the cell protoplast to form 4 zoospores or contica Hindak (Hindak, 1963, p. 107, table 2 (20), fig. 1),
daughter cells. (PL 2, figs. 42-46.) although it differs considerably because of its smaller and
This species is in the subgenus Euchlamydomon-as, and less cylindrical cells. I believe now that the rnair differences
is in some respects most similar to C. angulosa Dill but between Koliella Hindak and Raphidonema Lagerheim are
differs in its smaller size, its only medium-sized papilla, not of a general but of an entirely gradual character and,
and the longitudinally-striped chloroplast on the outside, therefore, of only secondary value taxonomically. Compare
punctiforrn stigma, and the not angular but more or less in this respect also the key to both genera by Hindak (1963,
rounded smaller pyrenoid. C. leptobasis Skuja is somewhat p. 99) and the figures in his plates 1-6, figures 19-24).
similar, but it is smaller, more rounded-ovoid, with longer Scenedesmus aoutus (Meyen) Chod.
flagella, and it has a thinner, simple cup-shaped chloroplast Scenedesmus armatus (Chod.) G. M. Smith
with a small globose pyrenoid. C. subangulosa should also Scenedesmus brevispina (G. M. Smith) Chod.
be compared with C. saxonensis Skuja and with some other Scenedesmtis ecornis (Ralfs) Chod.
species of the subgenus Euchlamydomonas. Scenedesmus quadricauda (Turp.) Breb.
Chlorococcum humicola (Nag.) Rbh. Scenedesmus quadricauda var. maximus W. et G. S. West
Typical form. Cells globose, solitary or often gregarious Scenedesmus spinosiis Chod.
in small amorphous clumps, variable in size within the same Sphaerozosma granulatum Roy et Biss.
aggregation namely 7^-20^-25^ in diameter; cell wall thin Staurastrum margaritacetim (Ehrnb.) Menegh. Granulation
or moderately thick, colorless and smooth but in an older of the cell wall more or less reduced.
state sometimes more or less evidently lamellose; chloro-
plast cup shaped, thin, covering the whole wall or nearly EUGLENOPHYCEAE
so, with one pyrenoid in a thickened part of the plastid. Anisonema acinus Duj.
Reproduction by the forming of usually 8-16, rarely as many Euglena pisc-ifonnis Klebs.
as 32, aplanospores in a mother cell. (PI. 1, figs. 16-19.) Euglena viridis Ehrnb. var. olivacea Klebs.
Closterium acutum Breb. Typical form; cells lOO/i-123/i Menoidium incur vum (Fres.) Klebs.
long and 4/*-4.5/t broad.
Euastrum denticulatum (Kirchn.) Gay. CHRYSOPHYCEAE
Euastrum elegans (Br6b.) Kiitz. Bodo minimus Klebs.
Gloeocystis rupestris (Lyngb.) Rbh. Compare also Skuja Honas uniguttata Skuja.
(1964, p. 106, table 15, fig. 11.) Pleuromonas jaculans Perty.
DIATOMACEAE short, budlike protruding branchlets from which subse-
quently the terminal more or less tetraradiate hyaline spore
Achnanthes minutissima Ktitz.
Cyml) lla austriaca Grun. or conidium arises. (PI. 2, figs. 50-53.)
Cynibelld delicatula Kiitz. SELECTED REFERENCES
Cymbellahebridica (Greg.) Grun.
[For references cited, see general listing of references]
Cymbella ventricosa Kiitz.
Fragilaria construens var. Mnodis (Ehrnb.) Grun. Cleve-Euler, Astrid, 1951-55, Die Diatomeen von Schweden uni
Melosira distans var. pfaffiana (Beinsch) Grun. Finnland: Kgl. Svenska Vetenskapsakad., Handl. ser. 4; v. 2,
Neidium duMum (Ehrnb.) Cl. Cells 40/i-48/i long, 10/i-12/t no. 1, 151 p., 1951; v. 3, no. 3, 143 p., 1952; pt. 2, v. 4, no. 1,
broad. 149 p., 1953; pt. 3, v. 4, no. 5, 240 p., 1953; pt. 4, v. 5, no. 4,
Neidium iridis var. ampMffomphus (Ehrnb.) V. H. Cells 217 p., 1955.
80^-120/1 long, 27/i-30/i broad. Croasdale, H. T, 1955, Freshwater algae of Alaska; Pt. 1, Sonre
Nitzschia gracilis Hantzsch. Desmids from the interior: Farlowia, v. 4, no. 4, p. 513-563.
Pinnularia cardinaliculus Cl. A slightly larger form length 1956, Freshwater algae of Alaska; Pt. 1, Some Desmic's
of the cells 100^-115/1*, breadth 21/t-22/i; striae 9-10 in 10/j. from the interior; Pt. 2, Actinotaenium, Micrasterias, and
Pinnular'M dactylus Ehrnb. Length of the cells 230/i-250/tt, Cosmarium: Am. Microscop. Soc. Trans., v. 75, no. 1, p. 1-77.
breadth 46/1-48/u,; striae 5-6 in 10/t. 1957, Freshwater algae of Alaska; Pt. 1, Some Desmids
Pinnularia gentilis (Donk.) Cl. from the interior; Pt. 3, Cosmariae concluded: Am. Micro-
Pinnularia gi~b~ba Ehrnb. scop. Soc. Trans., v. 76, no. 2, p. 116-158.
Pinnularia gibba forma sul)undulata A. Meyer. 1958, Freshwater algae of Alaska; Pt. 2, Some new fonrs
Pinnularia intermpta W. Sm. from the plankton of Karluk Lake: Am. Microscop. So°.
Pinnularia interrupta forma minutissima Hust. Trans., v. 77, no. 1, p. 31-35.
Pinnulariamaior (Kiitz.) Cl. Hirano, M., 1968, Desmids of arctic Alaska: Kyoto Univ. Biol.
Pinnularia microstauron (Ehrnb.) Cl. Lab. Contr. 21, p. 1-53.
Pinnularia microstauron var. ambigua Meist. Krieger, Willi, and Gerloff, Johannes, 1962, Die Gattur*
Pinnularia viridis (Nitzsch) Ehrnb. Cosmarium, Pt. 1: Weinheim, Germany, J. Cramer, 112 p.
Stauroneis alabamae Heiden var. angulata Heid. Length 1965, Die Gattung Cosmarium, Pt. 2: Weinheim, Ger-
75/1-80^, breadth 17/j. According to Cleve-Euler (1953, many, J. Cramer, p. 113-240.
p. 211-212) Heiden's taxon is identical with 8. nol)iUs Nilsson, Sv., 1964, Freshwater Hyphomycetes: Symbolae Bet.
Schumann, but see also Cleve (1894, p. 148). Upsaliensis, v. 18, no. 2,130 p.
Stauroneis anceps Ehrnb. Length of the cells 63/i-71/i, Polunin, Nicholas, compiler and ed., 1947, Botany of the Cana-
breadth 14^-15/i; striae about 25 in 10/t. dian Eastern Arctic; Pt. 2, Thallophyta and Bryophyta:
Stenopterobia intermedia (Lewis) Fricke. Length of the Canada Natl. Mus. Bull. 97, Biol. Ser. 26, 573 p.
cells 200/1-215/tt, breadth ll/i-13/j; costae short, 4-5 in lO/i; Proschkina-Lawrenko, A. I., Zabelina, M. M., Kisselew, T. A.,
transverse striae very delicate, about 20 in 10/tt. and Scheschukowa, V. S., 1951, Diatomovije vodorosli: Mos-
Tal>ellariaflocculosa (Roth.) Kiitz. cow, Opredelenie Presnovodnych Vodoroslei SSSR 4.
Ross, Robert, 1947, Freshwater Diatomeae (Bacillariophyta), p.
CHLOROMONADOPHYCEAE 178-233, in Polunin, Nicholas, compiler and ed., Botany of
Vacularia virescens Cienk. Mostly encysted. the Canadian Eastern Arctic, Pt. 2: Canada Natl. Mus. Bull.
97, Biol. Ser. 26, 573 p.
CRYPYOPHYCEAE Starmach, Karol, 1966, Cyanophyta-Sinice, Glaucophyta-Glauio-
Cryptaulax milgaris Skuja. fity, v. 2 of Flora Slodkowodna Polski: Warszaw, Panstwoê
Cyathomonas truncata (Fres.) Fisch. Wydawnictwo Naukowe, 807 p.
Taylor, W. R., 1928, The alpine algal vegetation of the moun-
DINOPHYCEAE tains of British Columbia: Acad. Nat. Sci. Philadelphia
Peridinium cunningtonii Lemm. tab. contactum Lef. Proc., v. 80, p. 45-114.
1934, The fresh-water algae of Newfoundland, Pt. 1:
RHIZOPODA Michigan Acad. Sci., Arts, and Letters Papers, v. 19, p.
Paulinella chromatopJiora Lauterb. The ovoid test with a 217-280.
low collar around the aperture, 29/t-31/u. long and 25/t*-26/j 1935, The fresh-water algae of Newfoundland, Pt. 2:
wide; cyanells 2, about 25/t long and 4.5/i-5/x broad. Michigan Acad. Sci., Arts, and Letters Papers, v. 20, p.
185-230.
HYPHOMYCETES Vischer, W., 1920, Sur le polymorphisms de I'Ankistrodesmus
Tetracladium marchalianum De Wildem. Braunii (Naegeli) Collins: Schweizer. Zeitschr. Hydrologie,
Mycelium of this swamp species was found on decaying v. 1, p. 5-50.
leaves of Isoetes sp. The evidently septate hyphae are SOIL FUNGI AND ALGAE
sparsely or moderately branched and irregularly undulate
or are more or less nodulose (constricted always at the By L. W. DuEREix 6
septa), colorless or pale brownish ; older hyphae as much as The fungi and algae were cultured from soils that
6/1 broad, the terminal hyphae about 1.5/tt broad, with cells
that are 3-8 times as long as broad ; sporo- or conidiophores
surrounded the roots of several grasses (Alopecurus
lateral, relatively short and usually unicellular thus with- « Professor Emeritus of Botany and Plant Pathology, Colorado State
out septa, the terminal parts slightly capitate with many University, Fort Collins, Colo.
aequalis, Deschampsia caespitosa, Phleum alpinum, 7, in which six taxa representing as many genera were
and Trisetum spicatum) that had been collected for found, was the richest sample. In some of the genera,
transplanting in a common garden in order to study notably Phoma, Fvxarium, and Gephalosporium, it is
their ecotypic variation. The grasses, together with the difficult if not impossible to make determinations of
upper 3-4 inches of soil that adhered to their roots, were species.
collected on August 7, 1967, from upland sites and were In my studies of Alaskaii soil samples from Peters
placed in plastic bags to keep them moist during ship- Lake in the Brooks Range, Barter Island, the shores of
ment. While collecting the specimens, no special precau- the Arctic Ocean, and Matanuska, I have isolated 28
tions were taken to prevent contamination of the soil species of fungi. Judged from those and hundreds of
by foreign microorganisms; the preliminary inventory other samples from throughout the world, none of the
of species that were found in the soil samples may, taxa found on Amchitka Island can be considered to
nevertheless, be indicative of the microflora at the sites. represent unique distributions all are widespread taxa.
To isolate the fungi from the soil samples, small Much more intensive and careful sampling would be
fragments of soil were planted on Rose-Bengal agar. required to reveal unique elements of the microflora, if
This method of culture gives a more reliable index of they are present.
the fungi that are present than does the old dilution Algae were cultured from 10 of the 21 soil samples by
technique. Table 1 gives a list of the fungi that were placing soil fragments on white quartz sane1 that had
isolated from the 21 samples. Fungi were cultured from been autoclaved in small Erlenmeyer flasks. The sand
all but one of the samples, and two or more taxa were was then wetted with modified Bristol-Roach solution
identified from more than half of the samples. Sample (Bold, 1949) to add some nutrients and to stimulate
TABLE 1. Fungi and algae cultured from soil samples taken on Amchitka Island, Alaska
[X indicates the presence, .... the absence, and ? the questionable presence of the organism in the soil sample)
Soil sample
Organism
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21
Fungi
Alternaria tennis auct. Wiltshire ......... .... y
Aspergittus nidulans (Nidan) Winter..... ......................... . .....
terreus Thorn. ...-..-----......-...... --... ...
Cephalosporium sp... .--.------.....-.-.... --.-._._ . . y y y
Curvularia sp_.... .................................... .
Fusarinm sp _ .......................... .... y X X
X y
Mucor corticolus Hagen. .......... ..... y y
racemosus Fresenius... ........ ..... X
variant Povah.... __ ............... .. .... y
sp......... ............................................................
Penicillium cambertii Thorn.. .................. _ ......
citrinum Thorn-----..-.--. . . .....
frequentans Westling _ ...... ----- y
oxaltimm Thorn. -...-..-.- . .....
Sp... ..-.--..__...._.______.__.._______. _.__.________._._._ . , .. X
Phoma sp. .............................. y y
Pythium sp.---- -.-. ? y X
StachybotTysatTaCor&a...--^- ........... . ..... y
Stemphylium ilicis Tengwall. . . .... X
Trichoderma viride Pers. ex Fries... --.-. y
Verticillium sp. ...................... ...
Algae
Chlorella vulgaris Beyerinck. y X X
Chlorococcum humicola (Naeg.) Rabenhorst..-.- X X X X X
y
Palmella sp...-. .. ?
Phvrmidium antarcticum W. et G. S. West y X
X
X
Stichococcus subtilis (Kuetz.) Klercker.. y X X
Stigonema turfaceum (Berk.) Cooke __ . y
SOIL SAMPLES
1. Black sandy loam, pH 7.0. 8. Black sandy loam, pH 5.9. 15. Gray organic matter and clay, r*I 6.5.
2. Black sandy loam, pH 6.0. 9. Black sandy loam, pH 6.4. 16. Gray sandy loam, pH 6.2.
3. Gray organic clay, pH 6.0. 10. Black sandy loam, pH 6.5. 17. Gray organic matter and clay, pH 6.5.
4. Gray organic clay, pH 5.9. 11. Gray sandy loam, pH 6.5. 18. Black sandy loam, pH 6.2.
5. Gray organic clay, pH 5.5. 12. Gray sandy loam, pH 6.6. 19. Black sandy loam, pH 6.5.
6. Black sandy loam, pH 6.4. 13. Loam (muddy), pH 6.6. 20. Gray organic matter and clay, pH 6.5.
7. Black sandy loam, pH 6.2. 14. Dark sandy loam, pH 6.8. 21. Gray organic matter and clay, pH 6.9.
growth. In about 60 days, algae, if present in the sample, chitka Island have now approached this second phase,
will form a green growth on the sand. As in fungi, sam- as is indicated by the comments on the taxa that folk w.
ple 7 was among the richest in algae of those examined, The third and fourth phases, biosystematic and encyclo-
having four species that represent three genera. The pedic, are mostly in the future, although some work 1 as
absence of algae from 11 of the samples may have been now begun which might appropriately be called
because of limited soil sampling. Growth in a culture biosystematic.
frequently originates from a single propagule; there- The collections of the Gramiiieae were compared with
fore, more extensive sampling at a site would have those in the U.S. National Herbarium. The names tl at
increased the likelihood of including these single re- I suggest are based primarily upon these studies and re-
productive bodies. flect my own views regarding the application of specific
and subspecific categories which may be summarized
OBSERVATIONS ON THE FLORA » as follows:
By WILLIAM M. KLEIN 8 1. When two taxa intergrade over a considerable pT.rt
Since 1965 several Government agencies have sup- of their distribution range there can be, in my
ported studies of the vegetation and ecology of Amchit- opinion, no real justification for recognizing thQ.m
ka Island. Botanists of the U.S. Geological Survey and as distinct species. The occurrence of intermediate
others have collected extensively on the island, with types commonly indicates genetic continuity be-
vascular plants, bryophytes, and lichens having re- tween populations which are best treated as con-
ceived the most attention. I collected vascular plants on specific. A satisfactory treatment, however, vlll
this island during the summer of 1967 to provide tax- come only after the geography and pattern of
onomic support for ecological investigations. These variation are more thoroughly understood. Until
recent studies have resulted in the accumulation of a such time, it seems best to follow a conservative
large number of plant specimens, and the Amchitka course in the application of new names.
Island flora is now probably more thoroughly collected 2. Intraspecific taxa should be recognized where this
than that of any other island in the Near Island and appears to be an appropriate way of dealing with
Rat Island groups. the variation pattern. Recognition of such taxa
After I had critically studied some of these collec- usually requires intensive field and herbarium
tions, it became apparent that the recent collecting ef- studies, and if possible these studies should be
forts had brought the understanding of this flora to correlated with experimental findings. In irost
a new level and that certain binomials could no longer instances where intraspecific taxa are distin-
be applied. The observations in this report indicate the guished, there is geographic and ecological dif?-r-
progress that has been made in these floristic studies entiation, and taxa can be delimited on the basis
and emphasize the need for a clarification of certain of morphology.
vascular plant names as applied to this flora. 3. The occurrence of two or more distinct morphologi-
The floristic studies of the Aleutian Islands have cal forms in the same populations throughout the
followed the usual course of events as described by range of a taxon may indicate the polymorphic
Davis and Hey wood (1963). The first and so-called nature of the population rather than fundamental
pioneering phase in the Aleutian Islands was led by differences. The application of names to the vir-
Hulten (1960). In this stage taxonomic decisions must tually endless number of variants can only burien
be based upon limited material, and judgments regard- the taxonomic system and inevitably obscure the
ing the status of a particular taxon must at times be more important relationships.
arbitrary. Through more comprehensive collections and
herbarium studies knowledge of the flora enters the Determinations of Epilobium were made by Dr. P. A.
second or consolidation phase where many of the var- Munz of the Rancho Santa Ana Botanic Garden, Clare-
iants described as species are found to be variants of mont, Calif. Collections of Salix were sent to Dr. G. W.
previously described taxa, Intensified exploration, also Argus of the W. P. Fraser Herbarium, University of
in this phase, may lead to the discovery of new species Saskatchewan, Saskatoon, Saskatchewan. Collections
and may make possible conclusions regarding phyto- of Ca-rex were sent to Dr. F. J. Hermann of the U.S.
geographic relationships. The floristic studies of Am- Forest Service. The assistance of these specialists is
gratefully acknowledged.
7 This study was supported by U.S. Department of the Interior contract In the vegetation studies and the annotated lis^ of
14-08-0001-10935.
8 Department of Botany and Plant Pathology, Colorado State Uni-
vascular plants of this report, Shacklette, Erdman, and
versity, Fort Collins, Colo. Keith used the names that are given in Hulten's fora
(1968), except for some references to Amchitka Island Poa alpina L. [P. Jiispidula Vasey; P. Jcomarovii Roshew;
taxa that were not collected in the present study. Hul- P. lanata (Scribn.) Merr.; P. turneri Scribn.]. An ex-
tremely complex group; the above-listed taxa cited by
ten's 1968 nomenclature was adopted because of the ad- Hultgn (1969, 1968) for Amchitka Island probably can
vantages gained by following a comprehensive modern all be assigned to the P. alpina complex .All of these taxa
treatment of the Alaskan flora. Results of taxonomic produce viviparous forms that appear to be environ-
studies that I report below are preliminary to more com- mentally induced.
plete morphological and cytological investigations of ONAGRACEAE
certain critical groups of taxa from this island.
Epilotitim boreale Hausskn. [E. Iglandulosum Lehm.
The following list includes only the families on which (Hultgn, 1960, p. 263-264; 1968, p. 690)]. Material deter-
some critical work has been done and the taxa that have mined by P. A. Munz to be E. boreale is very common on
binomials whose proper application was in doubt. Sug- Amchitka Island, although this species is not renorted by
gested names for species of four families in the Amchit- Hultgn for any of the Aleutian Islands. It seems likely
that this plant is being called E. glandulosum, which is
ka Island flora follow.
reported to be common along the entire island chain.
CYPERACEAE
SALICACEAE
Hermann's determinations of species in this family agreed
with those of Hultdn for all but one species, Carex kelloggii Salix arctica Pall. [8. crassijulis Trautv. (Hult6n, 1960,
Boott. Two collections of this species, 7485 and 8031, were p. 158-160) ; 8. arctica Pall, subsp. crassijulis (Trautv.)
made by Shacklette. Collection 7485 was determined as C. Skvortz. (Hulten, 1968, p. 340)]. Salix arctici, accord-
Jiindsii C. P. Clark and collection 8031 as possibly C. ing to G. W. Argus (written commun., 1968), is a highly
aquatills Wahl., although it was noted that the material of variable species and it is impossible at this point to
the latter was too immature for a critical identification to be distinguish intraspecific taxa.
made.
REFERENCES CITED
GRAMINEAE
Agrostis borettlis Hartm. [A. ala-skana Hult. (Hult£n, 1960, Ahti, T., 1961, Taxonomic studies on reindeer lichens (Cla-
p. 78; 1968, p. 99) ]. Agrostis alaskana is noted by Hultfin donia, subgenus Cladina) : Bot. Soc. Zool. Eot. Fenn.
(1960, p. 77) to be the most common species of the genus "Vanamo" Annales, v. 32, no. 1, p. 1-160.
in the Aleutian Islands. It intergrades with A. oorealis Akiyama, S., 1933, Carices of the Aleutian Islands collected by
and should be placed in that complex. Mr. Y. Kobayashi: Tokyo, Bot. Mag., v. 47, p. 67-69.
Descliampsia atropurpurea (Wahl.) Sheele [Vahlodea Anderson, S. T., and Bank, T. P., 2d, 1952, Pollen and radio-
atropurpurea (Wahl.) Fr. (Hulten, 1960, p. 88; 1968, carbon studies of Aleutian soil profiles: Science, v. 116, p.
p. 115)]. Vahlodea is treated as a segregate of 84-86.
Deschampsia in the U.S. National Herbarium and in most Arctic Weather Central, 1950, Climate, weather, and flying
American works. conditions of Alaska and Eastern Siberia: Alasl a, Elmen-
DescHiampsia caespitosa (L.) Beauv. [D. beringensix Hult. dorf Air Force Base, llth Weather Squadron, 1 p.
(Hulten, 1960, p. 85-87; 1968, p. 114)]. Deschampsia Army Research and Development Newsmagazine, 19"6, ARPA
leringensls is cited by Hult6n (1960, p. 86) as "one of reports on "Longshot" nuclear sounding test: February, p.
the most common grasses on the Aleutians." This species 29 [anonymous].
cannot be distinguished from D. caespitosa, and Kawano Ball, D. F., 1967, Classification of soils, in Peterken, G. F., com-
(1963) treats it in that complex. Hulten (1968) noted that piler, Guide to the Check Sheet for IBP areas: Oxford
many specimens show "hybrid influence from D. and Edinburgh, Blackwell Sci. Pubs., p. 121-12".
caespitosa." Bank, T. P., 2d, 1951, Botanical and ethnobotanical studies in
Festuca ovina var. ~brachypliylla (Schult.) Piper [F. ~bracliy- the Aleutian Islands; Pt. 1, Aleutian vegetation and Aleut
phylla Schult. (Hulten, 1960, p. 103-104; 1968, p. 168)]. culture: Michigan Acad. Sci., Arts, and Letters Papers, v.
Festuca frrachypliylla intergrades widely with F. ovina 37, p. 13-30.
and affinities are best shown by maintaining it in this 1956, Birthplace of the winds: New York, Thomas Y.
complex, which may also include F. ovina subsp. afosfcew- Crowell Co., 274 p.
sis Holmen (Hulten, 1968). Barbeau, Marius, 1958, Pathfinders in the North Pacific: Oald-
Festuca rulra L. [F. rubra subsp. acuta (Krecz & Bobr.) well, Idaho, The Caxton Printers, Ltd., 235 p.
Hult. (Hulten, 1960, p. 104; 1968, p. 171)]. Festuca rulra Bold, H. C., 1949, The morphology of Chlamydomonas chlamy-
is a highly variable species and some of the variants that dogama sp. nov.: Torrey Bot. Club Bull., v. 76, no 2, p. 101-
have been described appear to represent expressions of 108.
polymorphic systems and should not be accorded formal Cleve, P. T., 1894, Synopsis of the naviculoid diatoms, Pt. 1:
taxonomic recognition. Kgl. Svenska Vetenskapsakad., Handl., v. 26, no. 2, 194 p.
Plileum alpinum L. [P. alpinum var. americanum Fourn. Cleve-Euler, Astrid, 1953, Die Diatomeen von Schveden und
(Hulten, 1960, p. 74-75) ; P. commutatum Gandoger var. Finnland; Pt. 3, Monoraphideae, Biraphideae 1: Kg1!.
americanum (Fourn.) Hult. (Hult6n, 1968, p. 88)]. The Svenska Vetenskapsakad., Handl., ser. 4, v- 4, no. 5, 240 p.
recognition of infraspecific taxa and the application here Coats, R. R., 1956, Reconnaissance geology of soire western
of another specific epithet does not appear to be justified Aleutian Islands, Alaska: U.S. Geol. Survey Bull. 1028-E,
on the basis of existing evidence. p. 83-100.
REFERENCES CITED 59
Crum, Howard, Steere, W. C., and Anderson, L. E., 1965, A Llano, G. A., 1950, A monograph of the lichen family Umfr'li-
list of the mosses of North America: The Bryologist, v. cariaceae in the Western Hemisphere: U.S. Office Naval
68, no. 4, p. 377-432. Research Navexos P-831, 281 p.
Davis, P. H., and Heywood, V. H., 1963, Principles of angio- Macvicar, S. M., 1960. The student's handbook of British hep"t-
sperm taxonomy: Princeton, D. Van Nostrand Co., 558 p. ics: Codicote, Herts, Wheldon, & Wesley, Ltd., 464 p.
Degelius, Gunnar, 1937, Lichens from southern Alaska and Merrill, G. K., 1929, A new list of Alaskan lichens in the gerus
the Aleutian Islands, collected by Dr. E. Hult£n: Medd. Caladonia: The Bryologist, v. 32, no. 3, p. 41-50.
Goteborgs Bot. Tradgard, v. 12, p. 105-144. Metcalfe, G., 1950, The ecology of the Cairngorms; Pt. 2, Tê
Fosberg, F. B., 1967, Classification of vegetation for general mountain Callunetum: Jour. Ecology, v. 38, no. 1, p. 46-74.
purposes, in Peterken, G. F., compiler, Guide to the Check Murie, O. J., 1959, Fauna of the Aleutian Islands and Alaskan
Sheet for IBP areas: Oxford and Edinburgh, Blackwell Peninsula, with notes on invertebrates and fishes collected
Set Pubs., p. 73-120. in the Aleutians, 1936-38, by V. B. Scheffer: U.S. Dent.
Fraser, G. D., and Barnett, H. F., 1959, Geology of the Delarof Interior, Fish and Wildlife Service, no. 61, 406 p.
and westernmost Andreanof Islands, Aleutian Islands, Costing, H. J., 1956, The study of plant communities: San Fran-
Alaska: U.S. Geol. Survey Bull. 1028-1, p. 211-248. cisco, W. H. Freeman & Co., 440 p.
Guggenheim, Paul, 1945, An anthropological campaign on Pear sail, W. H, 1950, Mountains and moorlands: London, Col-
Amchitka: Sci. Monthly, v. 61, p. 21-32. lins Clear-Type Press, 312 p.
Hale, M. E., Jr., and Culberson, W. L., 1966, A third check- Persson, Herman, 1968, Bryophytes from the Aleutian Islards,
list of the lichens of the continental United States and Alaska, collected mainly by Hansford T. Shacklette: Svensk
Canada: The Bryologist, v. 69, no. 2, p. 141-182. Bot. Tidskr., v. 62, no. 2, p. 369-387.
Hamelin, L.-E., and Cook, F. A., 1967, Le p6riglaciaire par Powers, H. A., 1958, Alaska Peninsula-Aleutian Islands, in Wil-
1'image Illustrated glossary of periglacial phenomena: liams, Howel, ed., Landscapes of Alaska their geologic
Quebec, Les Presses de 1'Universite Laval, 237 p. evolution : California Univ. Press, p. 61-75.
Hattori, Sinske, 1958, The Hepaiticae of Ontake Mountain, Mid- Powers, H. A., Coats, B. R., and Nelson, W. H., 1960, Geology rnd
dle Japan: Hattori Bot. Lab. Jour., v. 20, p. 33-53. submarine physiography of Amchitka Island, Alaska: U.S.
Hattori, Sinske, Sharp, A. J., Mizutani, M., and Iwatsuki, Z., Geol. Survey Bull. 1028-P, p. 521-554.
1968, TaTcakia ceratophylla and T. lepidozioides of Pacific Beeves, Fontaine, Jr., 1968, A perennial Nidula from Alaska:
North America and a short history of the genus : Misc. Bryol. Am. Midland Naturalist, v. 80, no. 1, p. 272-273.
et Lichenol., v. 4, no. 9, p. 137-149. Rieger, Samuel, and Wunderlich, B. E., 1960, Soil survey and
Hedrick, Joyce, 1936. Lichens from the Aleutian Islands and vegetation of northeastern Kodiak Island area, Alaska: U.S.
the Alaska Peninsula: Michigan Acad. Sci., Arts, and Let- Dept. Agriculture, Soil Conservation Service, Soil Survey
ters Papers, v. 21, p. 75-80. Ser. 1956, no. 17, 46 p.
Hindak, Fr., 1963, Systematik der Gattungen Koliella gen. nov. Satake, Y., 1932, Juncaceae of the Aleutian Islands collected by
und Raphidonema Lagerh.: Nova Hedwigia, v. 6, nos. 1, 2, Mr. Y. Kobayashi in 1931: Tokyo, Bot. Mag., v. 46, p. 185-
p. 95-125. 187.
Hulten, Eric, 1933, Aleutiska Oarna, en Geografisk och Schuster, B. M., 1969, The Hepaticae and Anthoceratae of North
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1968, Flora of Alaska and neighboring territories, a 1966, Unattached moss polsters on Amchitka Island, Alas-
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1,008 p. 1967, Copper mosses as indicators of metal concentra-
Hutchison, I. W., 1937, Stepping stones from Alaska to Asia: tions: U.S. Geol. Survey Bull. 1198-G, 18 p.
London, Blackie & Son, Ltd., 246 p. Shacklette, H. T., and Bubin, Meyer, 1969, Badiocarbon dating
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Jorgensen, E., 1934, Norges Levermoser: Bergens Mus. Skr., no.
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principal soil groups of Alaska: U.S. Dept of Agriculture 399 p.
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60 VEGETATION OF AMCHTTKA ISLAND, ALEUTIAN ISLANDS, ALASKA
Soil Survey Staff, 1960, Soil classification, a comprehensive sys- Walker, E. H., 1945, Plants of the Aleutian Islands, in Collins,
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Vischer, W., 1920, Sur le polymorphisme de I'Ankistrodesmus National Park, Alaska: The Bryologist, v. 70, no. 2, p.
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v. 1, p. 5-50.
INDEX
[Italic page numbers indicate major references]
A Page B Page Pagi

Achittea borealis............................ 26,31,34 Bacidea sp._...._.___..______... 32,48 Corel____._._-_.____ ....... 57
Achnanthes minutissima...................... 55 Baeomyces placophyllus...... .......-.---.. . 33,47 anthoxanthea..................... 16,17,21,22,35
spp -.._- __.-- __ __ 50 Barbula cylindrica......................... 14,41,47 aquatilis........................... .. ... 58
Acknowledgments..........._.__--.... 3,40,57 Bartramiaithyphytta....................... 14,41,44 bigelowii.................................. 31,35
Actinotaenium cucurbita.................... 50;pl.l Bartramiaceae- ...____..._.... ..... 44 circinnata-.-......................... 29,33,35
Adelanthaceae................................ 41 Bazzaniatncrenata.----...................... 18,41 dioica gynocrates.......................... 16,35
Age of ash layers, radiocarbon dating. -..._.__ 8 Blasia pusitta .............................. 20,43 hindsii.................................... 58
Agrostis alaskana....................... 15,19,36,58 Blasiaceae.................................... 43 kdloggii............................. 21,24,35,58
borealis............................. 32,33,36,58 Blastenia sp.___....__...______.... 13,49 lyngbyaei............ 16,17,18,19,21,22,23,25,35
exarata................................... 13,36 Bodo minimus................................ 54 macrochaeta................ 16,17,18,22,26,32,35
Alectoria divergent............................ 50 Bog habitat-.....-.. -.---..--- ------ 10,20 subrigida.............................. 35
irvingii................................ 28,33,50 Bog soil......_...........__-.____-..- 6 nesoptiila........................ ... 35
nigricans................................. 33,50 Boraginaceae...- -..-------..---..-----.. - 34 physocarpa.... _ - - 31
ochroleuca....................... 16,28,29,32,50 Brachytheciaceae_.-...-..---.-.-..----.-.-- 45 pluriflora........................ 16,18,21,22,31
pubescent ............................ 29,32,50 Brachythecium....... .........__........... 25 saiatUislaxa....................... ....... 35
Algae_.___.__._....__._-.___.. 60 acutum... ....................... 16,27,33,45,46 ttylosa.................................... 16,35
cultured from, soil samples---.-..--------- 56 albicans....................... 13,19,27,28,30,45 Carex-Cladonia meadow community---------- 10, 16
selected references........................ 55 asperrimum...... 16,17,19,21,25,27,41,43,44,45 Carer-lichen community, ecotones around.... 9
soil.----...---------------.-------..-....- 66 Bryaceae...-........ ............ - ... 45 Carex-lichen-Empetrum community------- 10, 16, 17
Alluvial soil...._.__..................... 6 Bryum sp.-.. .......-.. .. ... - 24 Caryophyllaceae ......... 34
Alopecurusaequalis........................ 24,36,55 Bromus sitchensis aleutensis....-....----.....- 13,36 Cassiope lycopodioides............. 16,28,29,30,32,35
Alpine meadow community. ----.-------._.__ 10, 31 Bryoerythrophyllum recurvirostrum............ 13, CastilUja unalaschcensis.............. . 39
Alpine meadow habitat...................... 10,30 20,25,44,45,47 Cephalosporium sp . .. .- .. ... 56
Alternaria tennis.............................. 56 Bryophytes, annotated list....... ...... 40 Cephalozia ambigua..................... 20,31,41,43
Am blystegiaceae....---.-...-.....--.-----.-. 44 Bryoxiphium norvegicum...................... 47 bicuspidata ........... 18,19,26,29,31,41,43,45
Amblystegiella sprucei......................... 44 Bryum arcticum............................. -19,45 Uucantha........................... 18,33,41,42
Amblystegium serpens............. 14,15,21,44,45,47 inclinatum- .........___..... . ... 45 media..................................... 19,41
Amphidium lappmicum................... 14,45,47 lapponicum... -...---....-..-..-..-- 14,41,44,45 montana................. - - 42
Anabaena variabilis........................... 56 pollens.................................... 19,45 pUniceps.............................. 20,29,41
Anaphalis margaritacea....................... 26,34 pseudotriquetrum ......-.....-.-.---.. 13,19,45 Cephaloziaceae...----. ------- -- ------ 41
Anastrophyllum.............................. 42 *o/in«n»--.----- --..---.-- -------- 45 Cephalozietta............------------ ---- 40
sphendtoboides..--...-................. 18,41,42 stenotrichum......... 14,15,26,27,29,33,44,45,47 arctica..............- ........... 13,14,20,41
Andreaceae................................... 44 Buellia pullata................................ 49 Cephalozielliaceae.--------------------------- 41
Andreaea rupestris....... 13,28,29,30,31,32,33,40,44 punctata......................-...---..... 13,49 Cephaloziopsis saccatula....................... 43
Andreaea-Schistidium-Rhacomitrium commu- Burned areas, plant communities...--------- 33 Cerastium aleuticum.................... 28,30,33,34
nity--.---.------.-------.... 10,25 beeringianum beeringianum................ 34
Anemone..................................... 10 fischerianum........................... 12,28,34
narcissiflora villosissima................ 15,16,39 Odamagrottis................................. 18 fontanumtwale.--.................... 21,26,34
Anemone-Elymus-Ligusticum community_.. 10, 16 community_ - - -- . - 10, SO Ceratodon purpureus..... 12,13,17,26,27,32,33,45,46
Aneuraceae .........................._... 43 nutkaensis........... 15,16,17,18,19,26,34,36,46 Ceratodon-Equisetum-Sagina community . - 10,27
Angelica lucida............................ 26,32,40 CdlUergonsarmentosum......................- 19,44 Cetraria ciliaris............................... 49
Anisonema acinus............................ 54 Callitrichaceae..___.__...__........ 34 crispa................................. - 49
Anisothecium squarrosum..................... 45 Callitricheanceps...---....................... 24,34 cacMflote - - - " 32,49
Ankistrodesmusbraunii............. ....... 53; pi. 2 hermaphroditica........................... 34 erfcetoram----------------------------- 31,33,49
spir<dis................................... 53 Cattitriche-Subularia community. ___ - _ - - - 10,24 islandica crispa....................-...... 49
Anomobryum condnnatum................. 19,45,47 Caloneis silicula. ............................ 50 lacunosa..................... ------------- 49
filiformis.................................. 45 silicula alpina .......................... 52 nigricans..................... ----- - 29,49
Antennaria dioica.................... 20,26,27,32,34 longissima........................---- 52 orbata....................-...-............ 49
pallida ....... ___..-..___--...... 34 Caloplaca elegant..... ___.__....... - 49 CAoeiopedJo.-.-------------------------------- 53
Antheba...................................... 40 gramdosa.............................. 13,14,49 Chaetoplaca -- - 53
julacea..................... 21,24,31,33,40,41,43 sp........................................ 13,49 crassiseta puella..............-...-...... 53
Antheliaceae....__................___. 41 Caloplaca-Eurhynchium-Puccinetta community- 10, 13 Chttoscyphus pallescens........ 19,20,24,25,41,43,44
Antitrichia.. ........._......___._..... 18 Caltha palustris asarifolia..................... 22,39 CMamydomonas alaskensis............... 53,54; pi. 2
curtipendula............ 13,16,18,19,29,30,32,46 Caltha-Claytonia community._.._.... ... 10,22 angulosa.. ---------- ------------------- 54
Aongstroemialongipes..-. _____......_... 22,45 Calypogeia mulleriana........................ 20,41 jemtlandica.....................-------... 54
Aphanothece castagnei......................... 51 sphagmcola...................... 16,18,21,29,41 leptobasis.........-----.. --- ------- 54
Aphragmus eschscholtzianus................... 35 trictiomanis-........................ 17,19,41,42 saxonensis................... ... - --- 54
Arctoafulvetta.----........................... 29,45 Calypogeiaceae..-------.-......---.. .--.--- 41 subangulosa. ..----- -------- - 54; pi. 2
Arnica unalaschcensis................... 16,26,31,34 Ca mpanula chamissonis... _ - 32,34 vulgaris. ------------ --------- ------- 54
Arnica-Lupinus community.................. 10,26 lasiocarpa lasiocarpa............. 16,28,29,32,34 Cldorella ellipsoidea..-------.-.------- -- ------ 51
Ash layers-.................................. 7 Campanulaceae---.- _ _ . _. 34 vulgaris...... ._ . . ---- --- 51,56
Aspergillus nidulans.......................... 56 Campyliumstellatum---..._............... 19,24,44 Chlorococcum tiumicola... -_ - 54,56; pi. 1
terreus.................................... 56 Caprifoliaceae..-------..-. ----------- 34 Chloromonadophyceae.------- - ------ 55
Athyrium felix-femina cydosorum........... 15,19,38 Carcasses, plant communities ------------ S3 Chloromonas.......----......----------------- 54
Aulacomniaceae.---.__..................... 44 Cardaminebdlidifolia......................... 30,35 Chlorophyceae.---------------- -- ----- 51,53
Aulacomnmm palustre .......... 16,21,22,24,44,46 umbellata.................. 14,16,17,19,20,26,35 Chroococcus varius............................ 50
61
62 INDEX
Page Page Page

Chrysanthemum arcticum arcticum...... 16,31,32,34 Conocephalum conicum....................... 19,44 Ditrichumheteromallum ................. 26,45,46
Chrysophyceae.------------------------------ 52,54 Conostomum tetragonum............. 28,29,30,33,44 Drabaaleutica............................. 28,30,35
Citraria ciharis.................................. 33 Coptis trifolia.............................. 16,18,39 hyperborea.---......................... 13,14,35
Cladoma alpestris.................... 19,28,30,47,48 Cornaceae...............--...-...-.--.--....- 35 Draba-Saiifraga-Potentilla communr' "_ __... 10, 14
amaurocraea........................... 18,24,48 Cornicularia divergent................ 16,18,32,33,50 Drepanocladus fluitans_____________ 22,44
arbuscula beringiana...................... 18,48 Cornus canadensis x suecica................... 35 reealvens................... 17,21,22,24,29,31,44
belhdiflora.................... 16,18,23,24,48,50 suecica ........................... 16,18,31,35 uncinatus................................. 13,
cerasphora................................... 48 Cosmarium cydicum arcticum................. 50 18,19,27,28,29,30,31,32,33,44,45,46
coccifera............................... 16,34,48 decedens............................... 50; pi. 1 Dryopteris dilatata amencana.................. 19,38
degenerans............................. 20,34,48 humile.................................... 51 linneana.................................. 38
gratilis............................. 16,20,34,48 laeve................................... 51; pi. 1 Dune habitat __._____________ 10,/2
impexa...................................... 48 septentrionale-...--................... 51
paciflca....................... 16,17,18,28,34,48 margaritiferum............................ 51 E
pyxidata............................... 27,34,48 meneghini concinnum.................. 51; pi. 1 Elymus...................................... 47
rangiferina................................ 31,48 nitidulum................................. 51 arenariusmollis........................... 13
Salix-Empetrum community---.---..-------- 28 obliquum.............................. 50; pi. 1 mottis mollis.............. 1% 13,15,16,19,36
scabriuscula .......................... 18,19,48 pseudonitidulum.......................... 51 community------______..._____ 10, IS
sylvatica..................................... 48 subcrenatum........................... 51; pi. 1 sea stacks....------------_..-_.__- 15
sylvestris................................. 48 subpachydermum....................... 51; pi. 1 Elymus-Ligusticum-Anemone comnrinity..--- 10, IB
Cladonia-Carex meadow community---------- 10, 16 subundulatum......................... 51; pi. 1 Empetraceae -----________.___..- 35
Cladoniaceae..---------..--___...__._._--_ 47 tatricum............................... 50; pi. 1 Ecotones, alpine region..--------------------- 27
Cladi>p\ora...................................... 56 variolatum............................. 50; pi. 1 between plant communities.----------_ 9
Claopodium crisp'folium...................... 19,47 Cratoneuronfllicinum...................... 12,14,44 Empetrum nigrum nigrum.................... 10,
Claytonia arctica.............................. 30,38 Cruciferae.---.--..---_.-.-__..--.-_...- 35 16,17,18,19,27,28,2^, 31,32,33,34,36
sibinca.................... 12,13,14,17,22,31,38 Cryptaulazvulgaris----....................... 55 Empetrum heath, alpine community..... _.... .31
albiflora................................. 38 Crypyophyceae.-.-..-....-......-...--.__. 55 discontinuous heath habitat- - __... __ 27
Claytonia-Caltha community.-...-___.__ 10,22 Curvularia sp..---------__.---__.. T -----_.- 56 frost-scar community.--------------------.-..20
Climaciaceae..-... .....-.-..----.-..-.....--- 45 Cyanophyceae__.................__. 50,51,52 pool habitat_._____.___._.-.-. 23
Climacium dendroides......................... 19,45 Cyathomonas truncata......................... 55 Siphula-Scapania community....-----._- 24
Climate of area..........-..-.--------.- ..- . 4 Cylindrocystis brebissonii...................... 50,51 soil profile_.-._...__..._____... 7
Closteriumacutum............................... 54 brebissonii minor.......................... 50
Cochlearia offlcinalis oblongifdia............... 14,35 crassa..................... __..--.____ 50,51 Sphagnum ........................... 21
Communities, alpine meadow.--..-_____-_-_- 10, 31 Cymbetta aequalis............................. 52 vascular plants........... ..-.----.-..- S4
Andreaea-Schistidium-Rhacomitrium....... 10,28 angustata................................. 52 wet sedge-meadow community........... 22
Clamagrostis.............................. 10,26 hybrida............................... 52 Empetrum heath habitat... ----- ... 10,15
Caltha-Claytonia.......................... 10,22 austriaca.................................. 55 organic bird-perch mounds.... .... 18
Cladonia-Carex meadow......._...._. 10,16 cuspidala.............. ------- --------- 52 Empetrum-Carex-lichen community .... 10, /6,17
Elymus.................................. 10, IS delicatula................................. 52,55 ecotones-..--___.-...-..--....-.-....- 9
Ely'mus-Ligusticum-Anemone............. 10, IB graalis.................................... 52 Loiseleuria procumbens................... 25
Empetrum-Carez-lich&i ................. lo,16 hebridica.................................. 52,55 ruderal habitat-___.._.. ----- 25
Equisetum-Sagina-Ceratodon.............. 10,27 naviculiformis............................. 52 Empetrum-Salix community..---...--......- 10,28
Eurhynchium-Puccinellia-Caloplaca....... 10, IS perpusilla .............................. 52 Entodontaceae ...__.__..-.. - - 46
fell-field.. -....-- ...-.-..-.--....... 10,28 pusitta.................................... 52 Epilobtum.................................... 57
Fontinalis-Ranuncidus................... 10,25 tumidula.................................. 52 behringianum............................. 28,37
frost-scar.-.--.---_,....--.....-..-._. 10,20 eentricosa................................. 52,55 boreale.................................... 38,58
high-altitude bedrock................_, 10,32 Cyperaceae..-_._____-----._-----------.-- 35,58 glandulosum........................ 20,22,37,58
Hippuris-Ranunculus.................... 10,2S Cystopteris fragilis fragilis.................. 14,19,38 sertulatum ....................... 21,22,26,38
Honckenya-Senecio........................ 10, .72 Epithemia turgida............................. 50
Isoetes-Ranunculus-Limosella ..... _..... 10,2S D Equisetaceae.-..._....... --------------- 35
Juncus-Eriophorum...................... 10,24 Dactylmhiza aristata....................... 15,26,38 Eyuisetum.................................... 13
low-altitude bedrock.....__.__..___. 10,32 Deschamsia alpina............................ 36 arvense............................. 13,16,27,35
Lupinus-Arnica.......................... 10,26 atropurpurea. -------- . . __- 36,58 Equisetum-Sagina- Ceratodon community.... - - 10, 87
miscellaneous plant. ........._........ S3 beringensU....................... 13,17,20,36,58 Ericaceae -..-- ___-.--.---_-----._.--- 35
moss mound .._.-------..._.....-.- 10,77 caespttosa.............................. 36,56,58 Erigeronperegrinus-.......................... 31
organic bird-perch mound.. _____. __ 10, /8 orientalis-......................... 24,26,36 peregrinusperegrinus..... . ....... 16,34
peat-bank..__._.______...._. 10, 19 Deschampsia-Festuca-Poa community -_.. 10, /2 Eriphorum russeolum majus.................. 35
Philonotis-Parnassia................... 10,2^,46 Description of island....--.-----.----- ... 4 russeolum rufescens.............. 17,21,22,24,35
plant...____..______._.-._. 9 Desmidiaceae.. -.----.--...-----------.. 50 Eriophorum-Juncus community-------------- 10,24
burned areas...._-..---.._---...- S3 Diapensia lapponica obovata................ 29,30,35 Euastrum ansatum........................... 51
carcasses .._............._..... S3 Diapensiaceae__-...---,.-_-... ..... 36 bidentatum................................ 51
imported wood-.---------..--.-....-- S3 Diatomaceae....--_......_.. --..... 52,55 denticulatum.............................. 51,54
plantings..-..__.-..- .-._...._- SS DichodonliumpeUucidum.............. ...--.. 20,45 quadrifarium.......................... 51
Poa-Deschampsia-Festuca................. 10,12 Dicranaceae...-.__--------_----...-.---- 45 didelta. ................................ 51
PotentiUa-Draba-Sax:fraga................. 10, 14 Dicranella heteromalla................ 19,20,21,41,45 elegans.... .--.--.... .-...-...-.. ...- 51,54
Rhacomitrium-Schistidium- Ulota.......... 10,13 palustris-.......................... 20,21,31,45 oblongum. ------------- . ----------- 51
Salix-Empetrum.......................... 10,28 subulata................... 20,21,26,41,43,45,47 pectinatum --.____-..------_----------- 51
Scapania-Nardia-Marsupella ............ 10,21 Dicranumangustum.......................... 21,45 Euchlamydomonas............................ 54
Siphula-Scapania......................... 10,24 elongatum....................... 16,18,19,41,45 Euglenapisciformis........................... 54
snow-bed...-.--___..._______... 10, SI fuscescens.............................. 18,20,45 viridis olieacea............................- 54
solifluction terrace._________-...- 10,29 groenlandicum.--------...... 16,18,41,42,45 Euglenophyceae..--..-.-.- .-.-.- .- 54
Sphagnum bog.._.__._____._ 10,20 howettii................. 16,18,19,28,29,30,32,45 Eunotia arcus .............................. 50
streamside. --------_.-.______ 10,19 scoparium................................ 45 diodon ............ - 52
Subularia-Callitriche...................... 10,24 Dilaenaceae....-...---------..---.----------- 43 gracilis.............................------- 50
wet sedge-meadow...___...__.___ 10,£/ Dinophyceae....--.---------------- ------- 51,55 lunaris capitata..........................- 50
Xanthoria-Ramalina...................... 10,14 Diplophyllum albicans........................ 17, pectinalis...................-...---------- 52
Compositae_..----..____.__...___ 34 19,20,24,29,30,31,33,41,42,43 praeruptabidens.......................... 51,52
Conical mounds, basic types._.............. 17 taiifolium............................. 29,43,45 muscicola....................--------- 50
Conioselinum chinense..................... 15,19,40 Discontinuous heath habitat-. . -._.- 10,27 robusta diadema........................... 51,52
Conocephalaceae...____.___...__ 44 Ditrichaceae____________.___. 46 septentrionalU.... -------------------...-- 52
INDEX 63
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Eurhynchium praelongum..................... 12, Habitats Continued Lathyrus maritimus pubescent................. 12,37
13,14,15,16,19,27,33,45,46 pool...__..........__--.....--..__... #3 palustris pilosus.......................... 37
Eurhynchium-PuccineUia-Caloplaca commu- ruderal-.............._.____...._... 85 Lecanora aleutica........... ______... 13,14,48
nity..---.----....--...-...-... 10, 13 sea-cliff--.-..---...-.--.- -----.-.. IS allophana................................. 33,48
solifluction...____________..... 88 atra........ _____ ..______.__ 13,48
strand.-.-....- --. . ... ...-- 1% castanea.................................. 19,48
Fell-field community.----.--.-.._-------- W,28 stream ---.------...--.-.-._.-...-..- 84 gelida..................................... 48
Festuca brachyphytta........... 16,17,18,32,33,36,58 Haematomma lapponicum..................... 48 straminea................................. 14,48
ocina..................................... 58 ventosumlapponicum..... ____....... 33,48 subfusca..... -------------------------__ 48
alaskensis............................. 58 Half Bog soil...-.....-...-....-........- 6 Lecanoraceae.-------___- -____......... 48
brachyphytta.......................... 36,58 Haloragaceae.------.-........._........... 37 Lecidea aleutica.......... ____ .. ...__ 32,48
rubra............................ 15,16,22,36,58 Hantzschia amphioxys......................... 52 armeniaca................................ 33,48
aucta........................... 13,16,36,58 Heracleum lanatum.............. 12,13,19,26,32,40 elegantior................................. 33,48
Festuca-Poa-Deschampsia community. --_. 10, 1% Herberta adunca....................-.-------- 30,41 flavocaerulescens.......................... 33,48
Flora, observations.---.-------.----- .------ 57 hutchinsiae.... ...._._.____....... 41 macrocarpa...........-.......... 13,27,32,33,48
Fontinalaceae...... _....................... 46 Herbertaceae--...... ....................... 41 platycarpa................................ 48
Fontinalis neomexicana........................ 25,46 Hieracium.................................... 27 symmicta.......................... ._... 33,48
Fontinalis-Ranunculus community. -. _...-- 10. S5 gracilealaskanum................---...... 27,34 vernaKt................................ 20,33,48
Fragilaria construens binodis.................. 52.55 Mate...................................... 28,35 Lecideaceae..............___..____._. 48
construens subsalina....................... 52 Hierochloe odorata............................. 15,36 Leguminosae.....-----.-------..-........-- 37
nitzschioides............................... 52 High-altitude bedrock community --------- 10, 38 Lentibulariaceae. - 37
Fritillaria camschatcensis...................... 19,37 Hippuris montana............................ 31,37 Lepidoiiaceratoptiylla .--.................... 40
Frost action, solifiuction habitat.. ........ 28 vulgaris................................... 23,37 trichodadus............................... 41
Frost-scar community... ----.--..---------- 10,50 Hippuris-Ranunculus community. ----------- 10, S3 Lepidoziaceae.. __ __ __ __ _._.. .. ._._ 41
Frustulia rhomboides saxonica................. 51,52 Honckenya peploides major.................... 12,34 Leptarrhena pyrolifolia........................ 31,39
Fungi, cultured from soil samples . _... 56 Honckenya-Senecio community --------- 10, 1% Leptobryum pyriforme................ 15,20,44,45,47
Nidula Candida........................... 1 Hormidiumflaccidum......................... 51,54 Leucodontaceae....----.._-.._- _ .____ 46
soil.._ - - - _..____ SB Hygrohypnum................................ 25 ~Lichen-Empetrum-Carex community..... - 10, 16, 17
Fusarium sp ._- - . ..... 56 buM...................................... 25,44 Lichens, annotated list. . __......... 1ft
ochraceum................................ 25,44 Ligusticum................................... 10
O Hylocomiaceae.--.---..._...._----------- 46 scoticum hultenii....................... 13,15,40
Galium....................................... 37 Hylocomiumsplendens.................. 16,18,26,46 Ligusticum-Anemone-Elymus community. 10, 15
aparine................................... 12,39 Hyphomycetes. - . 55 LDiaceae ......__...... ___--------- 37
trifidum columbianum............... 12,16,22,39 Hypnaceae.---. ----._---.-...---.---------. 46 Limosettaaquatica...................... 21,23,39,43
Gentiana aleutica........................... 26,32,36 Hypnum cupressiforme........................ 33,46 Limosella-Isoetes-Ranunculus community... -. 10, S3
amarella acuta plebeja..................... 26,36 dieckii.................................... 16,46 Linnaea borealis borealis....................... 16,34
Qentianaceae-... ..................._.-... 36 plicatulum................................ 30,46 Listera cordata nephrophylla. ....... _. __ 16,31,38
Geography of area.......-__...__.. .-.-. 4 Hypogymnia enteromorpha.................... 33,49 Lithosols, classification...-.--.-.-------- -.- 6
Geology of area... - 5 physodes.................................. 33,49 Lobarialinita................................. 16,49
Qeraniaceae...........-..----..........---... 36 subobscura................-...----.-.--.-- 29,49 Loiseleuria procumbens........ 16,18,25,27,28,32,36
Geranium erianthum.......................... 36 Lophocolea cuspidata......... ._- 13,14,31,41,43,46
Qeum calthifolium....................... 16,20,31,39 Lophocoleaceae ...._......______... 41
macrophyllum macrophyllum........ 13,19,22,39 Icmadophilaericetorum................ ........ 19,48 Lophozia..................................... 40
pentapetalum............................. 31,39 Inland bedrock habitat............-.....--- W,S1 alpestris...................... 20,22,31,41,42,43
Oloeocapsa magma............................ 50 lonaspisepulotica.-..-------.................. 20 groenlandka............ 16,17,20,21,33,41,42,43
Oloeocystis ampla............................. 56 epuloticaarctica..-........................ 32,48 ineisa............................... 20,24,33,43
rupestris.................................. 54 Iridaceae...--..--.---- -_..---..-----.---- 37 opacifolia.............................. 31,41,43
Gloeodiniummontanum.-...-.-............ 51; pi. 1 Iris setosa setosa............................... 37 porphyroleuca........ -__.__.___. 42
Gomphonema angustatum productum.......... 51,52 Isoe'taceae..-------------------- ---------- 37 wenzelii.-................................ 32,43
Qramineae...__._..................... 36,58 Isoetes muricata maritima..................... 23,37 Lophoziaceae...-....-_._................. 42
collections..------- ----._.----.-..---. 57 sp... _ .... .-. -. 24,37,52 Low-altitude bedrock communities..... 10,3^
Grimmia apocarpa.........----.-....---.--.-- 46 Isoltes-Ranunculus-Limosella community.. -. - 10, S3 Lupinus...................................... 29
maritima................................. 46 Isopaches bicrenatus........................... 42 nootkatensis-..................... 21,25,26,29,37
Lupinus-Arnica community...__...__.... 10, 88
Grimmiaceae.. ......_-----.--.----. 46
Luzula....................................... 27
Gymnocarpium dryopteris..................... 38 arcuate unalaschcensis................. 28,32r 37
Juncaceae.-.---.- -..------..----- .. . 37
Gymnocolea acutttoba.......................... 42 Juncus arcticus sitchensis...................... 12,37 multiflora kobayasii....................... 27,37
inflata........................... 17,18,31,41,42 biglumis.................................. 30,37 multiflorafrigida................ 16,20,26, 37
montana ............................ 42 ensifolius................................. 24,37 parviflora parviflora................. 19,26,27,37
marginata................................. 42 triglumis albescens............... 17,21,22,24,37 tundricola..................... 14,28,29,30,33, 37
montana......-.-.--.....-... ............. 42 Juncus-Eriophorum community-. . ... 10, 84 wahlenbergiipiperi........................ 37
acuta....... - _ . 42 Lycopodiaceae - .. 37
Jungermanniaatrovirens...................... 30,41
Gymnomitrion corattioides... -------_..... 28,33,43 minuta grandis...------------------------- 43 Lycopodium.................................. 16 27
Gyrophora cylindrica...-.--.............--.... 50 rigida /8 grandis.........-.......--.----..-. 43 alpinum.................................. 28 37
saccatula..................... ------------- 43 annotinum annotinum.............. 16,17,21.37
H sphaerocarpa.............................. 21,41 clavatum clavalum........................ 16 37
Jungermanniaceae ._..... - 41 sabinaefoliumsitchense................. 21,32,37
Habitats, alpine meadow. so selagoselago.. ................. 16,27,28,32,37
bog-....--.-----.--- 80 Lyngbya rivulariarum......................... 52
discontinuous health. 87
K
Koenigia islandica............................. 30,38 M
Empetrum heath .__......._....-.. IB Kolieaa....................................... 53
inland bedrock. _ - . . 37 corcontica.......................--- ------- 54 Macrodiplophyllumplicatum.-.----------..... 19 43
lake...__....____.....____....... 88 Majanthemum dilatatum................... 19,20 37
land, associated plant communities, out- Marchantiapolymorpha.............. 14,24,26,44 47
lined-. ..-__. --------....... 10 Lake habitat- - -- - 10, *» Marchantiaceae.-----.------ --------------- 44
miscellaneous .... .... _ -------- 11,33 Land habitats, associated plant communities, Marsh habitat-.. 10 91
marsh..-__--.--.- .--.-....._......... 81 outlined.-.. - ----- -------- 10 Marsupetta emarginata..... 19,21,24,30,31,41,43 50
plant..._. _ _ ___._..........--.-. 9 miscellaneous_ ._ - --- - - 11,33 Marsupetta-Scapania-Nardia community... - - 10 81
64 INDEX
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Marsupellaceae ..___...__..__.__- 43 Oscillatoria Continued Pinnularia Continued
Melosira distans dlpigena.........--...-.-..... 51,52 splendida............................... 53; pi. 2 lata---................................... 51,52
distansp)'affiana................ _. ... 55 subtilissima............................... 52 motor..................................... 52,55
Menoidium incunum......................... 54 sp........................................ 53 microstauron................ .._.__ 51,52.55
Merismopedia elegans......................... 52 Oxyria digyna................................. 31,38 ambigua......... ..................... 55
glauca.................................... 52 biundulata............................ 52
Mertemia mantima........................... 12 P piridis ---------------------------- 52,55
maritima maritima........................ 34 Palmetto, sp................................... 56 rupestris........--....--.......-...-.. 51
Mesotaenlum micrococcum.---------.......... 50 Pannaria pezizoides........................... 20,49 sudetica.-...-.-..-.--................. 52
Microlepidozia makinoana............ 16,18,20,41,42 Pannariaceae..........._..........._.-... 49 Placopsisgelida............................ 27,32,48
sllvatica................................... 41 Papaver alaskanum........................... 38 Plagiotheciaceae ..-.-.-------.--..-..-----._ 46
Mielichhoferia macrocarpa ................ 14,44,45 Papaveraceae.....-...-.---.__..-..--.-_ 38 Plagiothecium roeseanum................... 13,41,46
Mimulusguttatus............................. 25,39 Parmeliaalpicola............................. 33,49 undulatum ....................... 16,20,31,46
Mniaceae. .................................... 46 duplicata douglasicola..................... 49 Planctococcus sphaerocystiformis............ 54; pi. 2
Mnium glabrescens........................... 16, lanata.................................... 50 Plant communities.-.__.-.-...-.__....... 9
19,20,22,24,26,32,33, 41,43,45,46 omphalodes. ........................ 18,29,33,49 associated land habitats, outlined ...... 10
insigne.................................... 20,46 physodes.................................. 49 classifications...--....._..........-.--.- 11
pseudopunctatum................ 19,20,21,33,46 saiatilis .......................... 14,32,33,49 miscellaneous ._..__.......__.... 11, SS
Monas uniguttata............................. 54 subobscura................................ 49 Plant habitats---..---------....------------- 9
Montia fontana fontana.................... 14,25,38 sulcata. -------------------------- 14,32,33,49 Plant species-.....--. .... ........... 34
Montospora brevis............................. 56 Parmeliaceae.-.-------------.----.._...--.- 49 nomenclature-_........__ .__...... 3
Mortierella vinacea............................ 56 Parmeliopsis ambigua--.... ................... 33,49 Plantaginaceae.---...-_-.----.-----.--.---- 38
Moss mound community........_.___... 10, 17 Parnassia kctzebuei........................ 21,24,39 Plantago macrocarpa...... ... ... ----- - 22,38
Mougeotia.................................... 51,54 Parnassia-Philonotis community... ___ - 10, tl, 46 maritima juncoides...... ......... 14,15,21,31,38
Mucor corticolus.............................. 56 Paroicous inflorescence ._............... 42 Platanthera chorisiana ................ 16,27,32,38
racemosus................................. 56 Paulinella chromatophora..................... 55 convallariaefolia.................. 16,17,21,31,38
varians................................... 56 Peat-bank community. .---...--.---------. 10,19 dilatata.......................... 16,17,21,31,38
sp...-._ - - __.........___. 56 Pediastrum braunii........................... 51,54 tipuloides behringiana..------------------- 38
Mycoblastus alpinus.............. 16,18,27,28,33,48 Pediculans chamissonis.................... 16,22,39 Platydictya jungerman nioides ............ 14,19,44
sanguinarius.............................. 48 Pdlianeesiana ................. 22,31,41,42,43 Platysmatialacunosa ......-...-...--......- 33,49
Mylia anomala............................ 31,41,43 Peltigera aphtltosa-................... 18,19,28,32,49 Plectocolea .........__ . . - 40
taylori........................... 16,17,18,41,43 canina..--......... .......__... 13,16,19,41,49 obovata .............................. 20,41,45
Myriophyllum spicatum....................... 37 canina membranacea............------..-- 49 PUuromonasjaculans. ...................... 54
spuria.-..--------.------.-.---------- 49 Pleurozium schreberi. .......... 16,18,22,31,33,45,46
N malacea... ......................._--..-. 16,49 Poa alpina .............................. 27,36,58
spuria.... ...._. . .....__........ 12,49 arcticawilliamsii.-.................... 18,32,36
Names, flora, specific and subspecific catego- Peltigeraceae................................. 49 eminens. .......... ---- --.- 13,36
ries............................... 57 Penicillium cambertii ........................ 56 hispidula ............................... 36,58
Nardia co mp ressa............................ .31,41 citrinum.................................. 56 aleutica............................... 36
geoscyphus............................. 31,41,43 freguentans.....................-...--.--- 56 vivipara ~..-.......--------..---.-..-. 36
scalaris-... 13,19,20,21, 22,24,26,27,31,32,41,43 oxalticum.---------.-.-----------------.-. 56 komarovii. ---_.---. --------------------- 36,58
Nardia-Marsupella-Scapania community.-... 10, HI sp................. _-- .-- -- .-- 56 lanata.................................. 13,36,58
Navicula contenta................ ............. 52 Peridinium cinctum........................... 51 vivipara...-----.....--.-.-..-...-----. 30,36
cryptocephala....--....--................. 52 cunningtonii contactum....-..-..--.....-.- 55 malacantha................................ 36
minima................................... 52 Pertusaria coriacea..-.-......--.-----.-------- 32,49 turneri-. ........................... 27,36,58
atomoides............................. 52 hultenii................................... 33,49 Poa-Veschampsia-Festuca community 10, H
Neidium affine minus......................... 52 sp---.... -. ...- 33,49 Pogonatum...-........... . . .. 37
dubium................................... 55 Pertusariaceae..-----.-...__-----.---.------ 49 alpinum............. 16,17,18,19,24,33,43,46,47
iridis amphigomphus...................... 52,55 Petasitesfrigidus -................... 17,19,22,35 contortum.......----.-.-......-...---..... 20,47
Nidula Candida.. ............................ 1 Philonotis..................................... 26 sphaerothetium...... .... ----- - -- 47
Nitzschia gracilis.............................. 55 americana....................... 21,22,26,44,47 urnigerum.--.-..- 7,13,20,21,2P, 32,41,43,45,47
palea..................................... 52 fontana......-...--..--...-.-....---... 20,21,44 Pohlia annotina~~........................... 20,45
stagnorum................................ 52 PMlonotis-Parnassia bog-..........--------... 43 cruda .. . - - -- 13,14,20,45
Nonmenclature of plant species ............. 3 community_.._._____.-.-.-.-. 10,27,46 naten* - 20 26,29,33,45,47
Nostoc microscopicum......................... 52 Phleumalpinum.............................. 56,58 Polygonaceae - ----------- ...-- ... 38
paludosum................................ 56 alpinum americanum..................... 58 Palygonum viviparum................ 16,26,28,31,38
O
commutatum americanum.............. 16,36,58 Polypodiaceae . .. . ... - ------- 38
Phomasp -----------------.-----..---...... 56 Polypodium vulgare octidentale................ 19,38
Ochrolechiafrigida........ 14,15,16,27,28,29,32,33,48 Phormidium antarcticum ...........-..-.-.-. 56 Polytrichaceae- -. . - ....... 47
Odontoschisma elongatum................... 18,41,43 crouani...... .........__ . . 56 Polytrichum commune..........-------. - 18,19,47
Oedogonium.................................. 54 frigidum.---.---.-----------------.....-.. 53 formosum.......-.----.-------......--- 20,27,47
Oligotrichum hercynicum.-_..-.-............ .. 31,47 motte......--...----....--.......-----.... 56 gracile............................ ------ 47
parallelum............................... 31,47 subcapitatum..................... ----- 56 juniperinum.....-----------.--- ---------- 32,47
Onagraceae................................... 37,58 Phystia caesia.... ..-.--------..-----. ... 14,15,49 longisetum ........................... 22,44,47
Oncophorus wahlenbergii-...... 17,18,20,21,41,42,46 dubia..................................... 15,49 sphaerothecium... ------------------------ 20,47
Oocystispana.............................. 51; pi. 1 Physciaceae.----.----._.._. _......... 49 strictum......--.----------------- -------- 18,47
Orchidaceae..........._...................... 38 Physiography of area.--------.-.--..---..-.-- 5 Pool habitat------------------------- 10,23
Organic bird-perch mound community.. - _. 10,18 Picea sitchensis............................. 4,33.39 Portulacaceae.---------------- ------ 38
Orthocaulis binsteadii-............ 16,17,18,33,41,43 Pinguicula vulgaris macroceras- ......... 17,21,24,37 Potentillaegedtigrandis...-------- --------- 26,39
floerkei ................................. 20,43 Pinnularia appendiculata... .................. 52 hyparctica----- --------------- ---------- 30,39
Orthotrichaceae......................._.... 46 brevicostata .............................. 52 viUosa.. . - .-- 14,15,39
Oscillatoria amoena...................... 51,52; pi. 2 cardinalicidus.......-.....--.-.-.-------.- 55 Potentilla-Draba-Sa t ifraga community-------- 10, 14
amphibia.... ...-.....-.. ..... 51,52,53; pi. 2 dactylus- ............ .......-...-.. .-. 55 Portia ftdww------------------------ 14,21,45,47
defleia.................................... 52 divergens.................................. 52 Pottiaceae ----------- -------------- 47
exilis ................................... 52 gentilis ---------------------------------- 52,55 Primula cuneifolia saxifragifolia...... 16,26,31,32,38
facilis.................................. 52; pi. 2 gibba- .............................. 51,52,55 cuneifolia saiifragifolia albiflora............ 38
grossegranulata......................... 53; pi. 2 subundulata.------..---......--..-... 55 Primulaceae - --.----- ------------ 38
limnetica acicularis........................ 52 interrupta..... --------------_.......---- 52,55 Protists.... -..--__..----- ------------ B0
neglecta.............................. ..... 52 mtnufiwtma...................... - 55 selected references.. - - --- BB
INDEX 65
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Pseudanabaena bipes.......................... 53 Sagina-Ceratodon-Equisetum community...... 10,27 Sphenolobus marginalus....................... 42
catenata................................... 53 Salicaceae..-.__.... .... ._.__..... 39,58 minutus............................ 18,33,41,43
galeata.................................... 53 Salii......................................... 7,57 grandis......................... ...... 43
granulifera............................. 53; pi. 2 arctica.................................... 58 saccatulus................................ I"),43
minuta................................... 53 crassijulis... ........ 15,16,18,28,29,32,39,58 Splachnaceae.-. - - _ _ 47
Psoroma hypnorum........................... 19,49 crassijulis .............................. 58 Siachybotrys atra........ _................... 56
Ptilidiaceae_......--...-.-.._........... 43 cyclophyUa................. ...... 16,29,30,32,39 Staurastrum margaritaceum................... 54
Ptilidium ciliare.............................. 18,43 polaris pseudopolaris...................... 39 punctulatum pyjmaeum- ............... 50
Ptilium crista-castrensis....................... 16,46 rotundifolia......................... 28,29,32,39 Stauroneis alabamaeangulata... --------------- 55
Puccinellia langeana alaskana.............. 13,18,36 Salix-Empetrum community.......... _.... - 10,28 anceps.................................... 5"?.55
Puccinettia-Caloplaca-Eurhynchium commu- Samples, soil-profile, chemical contents....... 9 linearis.................---........... 52
nity-.. ........................... 10,13 Saiifraga bracteata............................ 14,39 nobilis.---..-----.--..--.................. 55
Pyrola asarifolia purpurea..................... 16,38 foliolosafoliolosa.......................... 30,39 Stellaria calycantha calycantha.... ....... 12,20,27,34
minor..................................... 31,38 hirculus ................................ 31,39 media..................................... 34
Pyrolaceae................................... 38 punctata insularis .......... 16,19,21,27,30,39 ruscifolia aleutica.......................... 30,34
Pythiumsp................................... 56 Saxifraga-Potentilla-Draba community........ 10,.74 sitchana bongardiana...................... 20,34
Saxifragaceae. ................................ 39 Stemphylium ilicis............................ 56
R Scapania irrigua........................... 17,25,43 Stenopterobia intermedia....................... 55
paludosa............. 17,19,20,21,24,31,40,41,43 Stereocaulaceae...----.._--- -_-- -- 49
Radiocarbon dating, ash layers...------------ 7,8 scandica............................ 20,29,41,43 Stereocaulon alpinum......................... 13,49
RaduJapolyclada........................... 31,41,43 subalpina................................. 23,43 denudatu m................................ 49
Radulaceae ..............._......._... 43 undulata ...................... 19,24,25,43,45 inter medium..............................27,49
Ramalina almquistii....................... 14,32,50 Scapania-Nardia-Marsupella community..... 10,2.7 tomentosum............................... 49
on cliff summits-.-----..--------------.-- 14 Scapania-Siphula community...__...._.-._ 10,24 alpestre............................... 27,49
scoparia.................................. 14,50 Scapaniaceae..-...----.--..----.-.-...-.-.-.- 43 vesuvianum..................--.--.--..... 30,49
Ramalina-Xanthoria community ----------- 10,14 Scenedesmusacutus........................... 51,54 denudatum............................ 49
Ranunculaceae.-.-._-------.-.-_---------.-.. 39 armatus.................................. 54 Stichococcus subtilis..------------------------- 56
Ranunculus hyperboreus hyperboreus.......... 23,39 brevispina................................. 54 Stictapulmonarialinita-.._... ................ 49
occidentalis............................... 31 ecornis.................................... 51,54 Stictaceae.-....-----.----.----.------------- 49
insularis..................... 15,19,22,30,39 quadricauda............................... 54 Stigonema minutum.. ..-.--------.-........ 50
nelsoni............................... 22,39 maximus ........................... 54 ocellatum.....-.-.--..-..---.--.........-- 50
reptans................................ 23,24,39 spinosus.................................. 51,54 turfaceum... ..-- .--- . .-....-.--.. 56
trichophyllus.............................. 37 Schistidium apocarpum -------------------- 13,28,46 Strand habitat ---------------------------- 10.72
trichophyllus.................... 23,24,25,39 maritimum...................... 14,15,41,44,46 Stream habitat. - - - -- -- 17,*4
Ranunculus-Fontinalis community........... 10,25 Schistidium-Rhacomitrium-Andreaea communi- Streamside community. -- -- - 17, 19
Ranunculus-Hippuris community............ 10,23 ty.-.--.---------.-.--------.-. 10,28 Streptopus amplexifolius...................... 19,37
Ranunculus-Limosella-Isoetes community.---- 10,23 Schistidium-Ulota-Rhacomitrium community.. 10,73 Studies, floristic- --.--.--------------------- 57
Raphidonema................................. 54 Schizthrix calcicola........................... 50 Substrates, miscellaneous.~ --. . 33
gracile................................. 54; pi. 2 lardacea...... ............_..-..--.--.... 50 Subulariaaquatica......................... 24,?5,39
References, cited, general listing-.______ . 58 Scotiella nitalis............................... 51 Subularia-Callitriche community-..-...-. 10,24
selected, algae and protists............... 55 Scrophulariaceae........ _ _--.-.-------_---- 39 Synechococcus cedrorum pallidus.....--..... 53; pi. 2
Regosols ................................... 6 Scytonena mirabile.... - - - 50 moi'or ----------------------------------- £1,53
Rhacomitrium aaciulare....................... 24,46 Sea-cliff habitat.............................. 10,73 notatus....-....---..---.-------.-------.- 53
breeipes................................ 29,45,46 Selaginellaselaginoides.---..--....... -... 21,24,40
canescens......................... ........ 46 Selaginellaceae--.........._....... ----- 40
ericoides...................... 13,26,27,29,45,46 Selected references on algae and other protists-. 55 Tabellaria flocculosa .....--...---.-...---.-. 51,55
fasciculare.............. 13,14,20,28,30,32,33,46 Senecio pseudo-arnica.....-..----..... ...-.. 12,3.5 Takakiaceratophylla.------................... F1,40
lanuginosum... 13,16,18,19,28,29,30,32,33,45,46 Senecio-Honckenya community - -..-..-..-- 10, 1% Takakiaceae-_.... . .--. .-.. . ... 40
Rhacomitrium-Andreaea-Schistidium commu- Sibbaldiaprocumbens--.--.............. 20,27,29,39 Taraxacum aleuticum.---..-------..---.-...-- 35
nity. ---.- ... - . 10,28 Siphulaceratites........................ 21,24,43,50 trigonolobum.....-------........--.....--. 14,35
Rhacomitrium-Schisiditum- Ulota community.. 10, / ? Siphula-Scapania community................. 10,24 Teloschistaceae..-----.- _-.----...--.-.--.. 49
Rhinanthux.--................................ 27 Snow-bed community.-------.--------------- 10,37 Teloschistes candelarius. .....---.---....----- 49
minor borealis.......................... 22,27,40 Soils, algae...---.-,._--.-----.------------.-- 55 Tetracladium marchalianum................ 55; pi. 2
Rhizocarpon atroalbescens..................... 33,48 chemical composition _______.. --._ 8,9 Tetraplodon mnioides...................... 33,45,47
geographicum............................. 33,48 classification .....___........ ..... 5 Thamnolia ..-.--. ............... ... ----- 28
hochstetteri................................ 32,48 fungi.........-.._..----...--_------------ 55 s ubidiformis ----------------------------- 50
Rhizopoda................................... 55 vermicularis ...................... 16,28,32,50
Solifluction habitat...--.---...----------.- 10,28
Rhododendron camtschaticum camtschaticum... 29,36 Thelypterislimbosperma ................. 31,33,38
Rhytidiaceae.-._....................__... 47 Solifluction terrace community.....-...-....- 10,29 phegopteris . ................ . 31,38
Rhytidiadelphus loreus............ 16,18,20,22,31,47 Sparganiaceae...--......--......--.---.-.-.-. 40 Thuidiaceae....-----..-.--------------------- 47
squarrosus....................... 13,31,41,43,47 Sparganium hyperboreum...--.---.--.--... 24,31,40 Tofieldia coccinea.. ---------------- 16,18,32,37
triquetrus.............................. 16,18,47 Sphaerophoraceae.......--..-....--....-..--. 49 Toninialobulata.............................. 20,48
Riccardia..................................... 40 Sphaerophorus globosus...... 16,18,19,28,29,31,33,49 Tortula mucroni folia....................... 13,14,47
latifrons. ........................ 18,21,40,41,43 Sphaerozosma granulatum..... ......--.-...--- 54 Trichoderma viride ---------------------------- 56
pinguis....................... 19,24,31,43,45,46 Sphagnaceae .............-....---.-...--.. 44 Trichophorum caespitosum. ------------------- 17,35
sin uata................................... 20,43 Trientalis europaea arctica-............. ----- 16,38
Rinodina turfacea............................. 19,49 Sphagnum... ................................. 40
europaea arctica aleutica-.. ... --_ - 38
Rosaceae..-.. - -- __ _ ._ ___ 39 bog community.----.-.--------------- 10,20,44 Trifolium repens.............................. 37
Rubiaceae.................................... 39 capillaceum--....................... 16,21,41,44 Trisetum spicatum..... ----------------------- 56
Rubus arcticus stellatus................. 16,18,20,39 compactum................ 20,21,24,30,31,40,44 spicatum alaskanum...................... 36
chamaemorus.......................... 16,18,39 Empetrum heath habitat...---.. ------ 21 alaskanum molle...................... 36
Ruderal habitat.....-..-..----..-,.......... 10,25 girgensohnii- ... -_____--..___.._.- 19,21,22,44 Tritomaria quinqwdentata------... ------- 30,41,43
Rumei fenestratus............................ 38 lindbergii..- -.-.............--- .--.-.... 31,44 Tundra without permafrost.---------------- 6
obtusifohux............................ 12, 22,38 Tundra without permafrost, profile ___ 7
magellanicum................. 21,31,41,43,44,45
papillosum................... 20,21,22,31,41,43 U
Sagina crassicaulis......................... 14,19,34 squarrosum. ............ 21,22,23,31,41,43,44,45 Ulota phyllantha.................. 13,14,15,19,44,46
intermedia........ ..................... 27,30,34 subsecundum. -...--... .. ....-.-.-. 16,21,44 Ulota-Rhacomitrium-Schistidium community.- 10, IS
ocddentalis............................. 12,27,34 teres.......................... 19,21,22,43,44,45 Umbelliferae. __..-_ .......... - 40
66 INDEX
Page Page Page

Umbilicaria eylindnca,......................... 32,50 Vahlodea atropurpurea........................ 58 Viola langsdorffii.... .............. 15,1% 17,22,32,40
hyperborea................................ 50 atropurpurea latifolia ......_----- 16,18,31,36 Violaceae..._.. .--..-.....--. -..-..._. 40
proboscidea .................... 13,29,32,33,50 paramushirensis...................... 37
torr«/octo ----------------------- 50 Vascular plants, annotated list..... 34 W, X, Z
UmbUiranaoeae 50 Vegetation on island . ._. - 4 Wet sedge-meadow community............... 10,*/
sneaceae _-_. ........ .. ------- .... Veronica, americana........................ 24,25,40 Wood, imported, plant communities ...-. 33
v grandiflora................................ 29,40 Xanthona, candelaria.................... 14,19,32,49
Vaccinium-..- ............................ 7 serpylUfolia humifusa......... 20,21,24,27,31,40 elegans.................................... 14,49
uliginosummicrophyllum................. 29,36 stellen................................. ib,dA4U onseastacks 14
«>«« iitnen
mtiS'idaea................................. ifi
ib Tlerrwcorio maura............................. 14,50 " community.
Xanthona-Ramalina -----------------------
.... 10, U
minus................................ 18,36 Verrucariaceae --.. -.. .. -. .- _ 50
Vacularia vireseens............... _ - _ 55 Verticillium sp .-.-.__ .. -- 56 Zygogonium ericetorum........................ 51
PLATES
PLATE 1
FIGURE 1. Actinotaenium cucurbita (Br6b.) Telling.
2, 3. Cosmarium variolatum Lund.
4-6. C. tatricum Racib.
7. C. decedens (Reinsch) Racib.
8, 9. C. obliquum Nordst.
10. C. subundulatum Willie.
11. C. subcrenatum Hantzsch forma.
12. C. meneghini Br6b. var concinnum Rbh.
13. C. laeve Rbh. forma.
14. C. subpachydermum Schmidle forma.
15. Oocystis parva W. et G. S. West.
16-19. Chlorococcum humicola (Nag.) Rbh.
20-23. Gloeodinium montanum Klebs.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 648 PLATE 1
DRAWINGS OF ALGA SPECIES

PLATE 2
FIGURE 24. Synechococcus cedrorum Sauv. var. pallidus, var. nov.
25. Oscillatoria amoena (Kutz.) Gom.
26. 0. facilis, sp. nov.
27. 0. splendida Grev.
28. 0. grossegranulata Skuja.
29. 0. sp. cf. amphibia Ag. forma.
30-32. Pseudanabaena granulifera, sp. nov.
33-41. Chlamydomonas alaskensis, sp. nov.
42-46. C. subangulosa, sp. nov.
47. Planctococcus sphaerocystiformis Korschik.
48. Raphidonema gracile, sp. nov.
49. Ankislrodesmus braunii (Nag.) Collins var.
50-53. Tetracladium marchalianum De Wildem.
U.S. GOVERNMENT PRINTING OFFICE: 1969 O 350-274

GEOLOGICAL SURVEY PROFESSIONAL PAPER 648 PLATE 2
DRAWINGS OF ALGAE AND OTHER PROTISTS

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Vegetation of Amchitka

Island, Aleutian Islands,

GEOLOGICAL SURVEY PROFESSIONAL PAPER 648

An account of the plant species and plant

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1969

Library of Congress catalog-card No. 76-602502

For sale by the Superintendent of Documents, U.S. Government Printing Office

[Plates follow index]

FIGURES 2-35. Photographs Continued Page

By HANSFORD T. SHACKLETTE and others

on the island can be seen by the vigorous growth of many

A similar trend in calcium and potassium content of

bon dates of samples from the top, middle, and lower

PRIMARY STRUCTURAL GROUP 3 a. Honckenya-Senecio community (1N2).

Occurs around the island wherever a shoreline bench

c. XanthoricL-Ramolina community (2H2).

-. vS«***»je " ' i>«*«a.-ii-j»«fcJC££S

FIGTJBE 6. Sea-cliff habitat on the Pacific Ocean coast at Makar-

perborea. Lecanora aleutica, L. straminea, Physcia

a. Empetrum-Carex-\\ohsx\. community (lC12c). Characteristic species: Empetrum nigrum subsp.

A well-developed moss mound has a zonal ar-

m-ine umbellata, Deschampsia beringensis, and On-

ticum, Blasia pusilla, Bryoerythrophyllum recur-

setwn, Rcmunculus occidentalis subsp. nelsoni,

Pools usually are formed in depressions of the vege-

a. Juncus-Eriophorum community (1M22).

Most streams are narrow, swift, and short; and the

filled with snowdrifts in winter and are a hazard to foot

Soils of this habitat are classified only by type. Many

c. Equisetum-Sagina-Ceratodon community (2G2).

FIGURE 30. Solifluction terraces on the high plateau near Rim

Other species: Agrostis borealis, Alectoria ochro-

surfaces, unless fertilized by birds. The species compo-

of plants were burned, whereas the saturated sub- CAPRIFOLIACEAE

Klein, in his section on plant species in this report, refers POLYPODIACEAE

By HERMAN PEBSsoN 2 and HANSFOBD T. SHACKLETTE HEPATICAE

LOPHOZIACEAE G. acutiloba, and G. marginata, all of which range from the

Order MARCHANTIALES Oalliergon sarmentosum (Wahlenb.) Kindb. 8047, 4e. Found

UMBILICABIACBAE SOME ALGAE AND OTHER PROTISTS

DIATOMACEAE Pinnularia appendiculata (Ag.) Cl. Cells 28/4-33^ long and

Oscillatoria grossegranulata Skuja. brana tenuissima, perspicua; contentu cellularum pallidis-

centrally placed. Asexual reproduction by the division of Hortnidium flaccidum A. Br.

[Italic page numbers indicate major references]

A Page B Page Pagi

Page Page Page

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DRAWINGS OF ALGA SPECIES

U.S. GOVERNMENT PRINTING OFFICE: 1969 O 350-274

DRAWINGS OF ALGAE AND OTHER PROTISTS

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