DARWIN'S GHOST

The Origin ofSpecies Updated

STEVF JONES

R A N D OM H O USE NEW YO R K
 Copyright © 1999,2000 by Steve Jones

All rights reserved under International and Pan-American Copyright Conventions.

Published in the United States by Random House,Inc.,New York.

RANDOM HOUSE and colophon are registered trademarks of Random House,Inc.

This work was originally published in slightly different form in Great Britain
by Doubleday,London,in 1999.

Library of Congress Cataloging-in-Publication Data

Jones,Steve
[Almost like a whale]
Darwin's ghost: The origin of species updated I Steve Jones.
p. cm.
Includes bibliographical references (p. )
ISBN 0-375-50103-7 (alk. paper)
1. Natural selection. 2. Evolution (Biology) 1. Title.
QH375.J66 1999
576.8¢2-dc21 99-53246

Random House website address: www.atrandom.com

Printed in the United States of America on acid free paper

2 4 6 8 9 7 5 3

First Edition
 To Dick Lewontin,
who showed me what evolution
can and cannot explain
L
 CONTENTS

The Origin ofSpecies: Facsimile Title Page and List of Contents IX

An Historical Sketch of the Progress of Opinion

on the Origin of Species XVll

Introduction 1
I Variation Under Domestication 21
II Variation Under Nature 40
III Struggle for Existence 55
IV Natural Selection 69
V Laws of Variation 1 02
VI Difficulties on Theory 1 19
VII Instinct 1 44
VIII Hybridism 1 69
IX On the Imperfection of the Geological Record 1 90
X On the Geological Succession of Organic Beings 213
XI Geographical Distribution 235
XII Geographical Distribution-continued 257
XIII Mutual Affinities of Organic Beings; Morphology;
Embryology; Rudimentary Organs 275
Interlude: Almost Like a Whale? 309
XIV Recapitulation and Conclusion 33 1
Further Reading 35 1
Index 36 1
But with regard to the material world, we can at least go so
far as this-we can perceive that events are brought about
not by insulated interpositions of Divine power, exerted in
each particular case, but by the establishment of general
laws.

-W. WHEWELL, Bridgewater Treatise

To conclude, therefore, let no man out of a weak conceit of

sobriety, or an ill-applied moderation, think. or maintain,
that a man can search too far or be too well studied in the
book of God's word, or in the book of God's works; divinity
or philosophy; but rather let men endeavour an endless
progress or proficience in both.

-BACON, Advancement ofLearning

 ON

THE ORIGIN OF SPECIES

BY MEANS OF NATURAL SELECTION,

OR THB

PRESERVATION OF FAVOURED RACES IN THE STRUGGLE

FOR LIFE.

By CHARLES DARWIN, M.A.,

FBLLOW OF THE ROYAL, GF..or.oGICAI., LtNNJEAN, ETC., SOCIETJES j
'
AUTHOR OF 'JOURNAL 01' RESEARCHES DURING H. M. 8. BEAGLE S VOYAGE
ROUNt> THE WORLD.'

LONDON:
JOHN MURRAY, ALBEMARLE STREET.

1859.
 CONTENTS.

INTRODUOTION .. Page 1

OHAPTER I.

V AJUATION trNDBB DoDBTIOATION.

Causes of Variability - Effects of Habit - Correlation of Growth­

Inheritance - Character of Domestio Varieties - Difficulty of
distinguishing between Varieties and Species-Origin ofDomestio
Varieties from one or more Species - Domestio Pigeons, their
Differences and Origin - Prinoiple of Selection anciently followed,
its Effects-Methodical and Unconscious Seleotion-Unknown
Origin of our Domestio Productions - (''iroumstanoeB favourable
to Man's power of Selection .. 7-43

OHAPTER II.

VARIA.TION UNDO NAT'O'BB.

Variability - Individual ditr�06B - Doubtful species - Wide

ranging, muoh diffused, and common species vary DlOIIt-Spe­
cies of the larger genera in any country vary more than the species
of the smaller genera-Many of the species of the larger genera
resemble varieties in being very olosely, but unequally, related
to each other, and in having restricted ranges •• 44-59
 CHAPTER III.

STRUGGLE FOR EXISTENCE.

lleal'tl on naturnl selection-The term used in a wide sense-Geo­
mctrical powers of increase - Rapid increase of naturnlised
animals and plants-Nature of the checks to increase-Compe­
tition universal - Effects of climate - Protection from the
number of individuals-Complex relations of all animals and
plants throughout nature-Struggle for life most severe between
individuals and varieties of the same species; often severe be··
tween species of the same genus-The l'elation of organism to
organism the most important of all relations Page 60-79

CHAPTER IV.

NATURAl. SEI.ECTION.
Natural Selection - its power compared with man's selection - itA
power on chal'acters of trifling importance-its power at all ages
and on both sexes -Sexual Selection -On the generality of intel'­
crosses between individuals of the same species - Circumstances
favourable and unfavourable to Natural Selection, namely,
intercrossing, isOlation, numller of individuals -Slow action­
Extinction caused by Natural Selection-Divergence of Cha­
racter, related to the diversity of inhabitants of any small area,
and to naturalisation-Action of Natural Selection, through
Divergence of Character and Extinction, on the descendants
from a common parent -Explains the Grouping of all organic
beings 80-130

CHAPTER V.

LAWS OF VARIATION.

Effects of external conditions - Use and disuse, combined with

natural selection; organs of flight and of vision - Acclimatisa­
tion-Correlation of growth - Compensation and economy of
growth - False correlations-Multiple, rudimentary, and lowly
organised structures variable - Parts developed in an unusual
manner are highly variable: specific characters more variable
than generic: secondary sexual characters variable -Species of
the same genus vary in an analogous manne r - Reversions to
long-lost characters - Summary .. .. 131-170
 CHAPTER VI.

DIFFICULTIES ON THEORY.

Difficulties on the theory of descent with modification-Transitions­

Absence or rarity of transitional varieties-Transition s in h ab its
of life-Diversified habits in the same species-Species with
habits widely different from those of their allies-Organs o f
extreme perfection-Means of transition-Cases of difficulty­
Natura non facit saltum-Organs of small importance-Organll
not in all cases absolutelyperfect-The law of Unity of Type
and of the Conditions of Existence embraced by the theory of
Natural Selection Page 171-206

CHAPTER VII.

I NSTIN C T.

Instincts comparable with habits, but different in their origin­

Instincts graduated - Aphides and ants:"- Instincts variable­
Domestic inst.incts, their origin - Natural instincts of the cuckoo,
ostrich, and parasitic bees - Slave-making ants - Hive-bee, its
cell-making instinct - Difficulties on the theory of the Natural
Selection of instincts - Neuter or sterile insects - Summary
207-244

CHAPTER VIII.

HYBRIDISM.
Distinction between the sterility of first crosses and of hybrids­
Sterility various in degree, not universal, affected b y close inter­
breeding, removed by domestication-Laws goveming the sterility
of hybrids - Sterility not a special endowment, but incidental
on other differences - Causes of the sterility of first crosses and
of hyblids - Parallelism between the effects of changed con­
ditions of life and crossing - Fertility of varieties when c roSBed
(\11(1 of their mongrel oft'spring not universal - Hy brid s and
mongrels compared independently of their fertility - Summary
245-278
 ·CHAP'l'ER IX.

ON THE IMPERFEOTION OF THE GEOLOGICAL RECORD.

On the absence of intermediate varieties at the present day - On

the nature of extinct intermediate varieties; on their number­
On the vast lapse of time, as inferred from the rate of depOsi­
tion and of denudation - On the poorness of our pa.lreontological
oollections - On the intermittence of geological formations­
On the absence of intermediate varieties in anyone formation
- On the sudden appearance of groups of species - On their
sudden appearance in the lowest known fossiliferous strata
Page 279-311

CHAPTER X.

ON THE GEOLOGIOAL SUCCESSION OF ORGANIO BEINGS.

On the slow and successive appearance of new species - On their

different rates of change - Species once lost do not reappear­
Groups of species follow the same general mles in their appear­
ance and disappearance as do single species - On Extinction­

On simultaneous changes in the forms of life throughout the

world- On· the affinities of extinct species to each other and to
living species - un the state of development of ancient forms -
On the succession of the same types within the same areas­
Summary of preceding and present cbapters . . 312-345

CHAPTER XI.

GEOGRAPHICAL DISTRIBUTION.

Present distribution cannot be a.ccounted for by differences in phy­

sical oonditions - Importance of l>arriers - Affinityof the pro­
ductions of the same continent - Centres of creation - Means
of dispersal, by changes of climat6 and of the level of the land,
and by occasional means - Dispersal dming the Gla.cial period
co-extensive with the world . . •• 346-382
 CHAPTER XII.

GEOGRAPHIOAL DISTRIBUTION-continued.

Distribution of fresh-water productions - On the inhabitants of

oceanic islands -Absence of Batrachians and of terrestrial Mam­
mals - On the relation of the inhabitants of islands to those of
the nearest mainland- On colonisation from the nearest source
with subsequent modification-Summary of the last and pre­
sent chapters Page 383-410

CHAPTER XIII.

MUTUAL AFFINITIES OF ORGANIO BEINGS: MORPHOLOGY:

EMBRYOLOGY: RUDnlENTARY ORGANS.

CLASSIFIOATION, groups subordinate to groups-Naturai system­

Rules and difficulties in classification, explained on the theory of
descent with modification-Classification of varieties - Descent
always used in classification-Analogical or adaptive characters
- Affinities, general, complex and radiating - Extinction se­
parates and defines groups - MORPHOLOGY, between members
of the same class, between parts of the same individual­
EMBRYOLOGY, laws of, explained by variations not supervening
at an early age, and being inherited at a corresponding age
- RUDnlENTARY ORGANS; their origin explained - Summary

411-458

CHAPTER X I V.

REOAPITULATION AND CoNOLUSION.

Recapitulation of the difficulties on the theory of Natural Selection

-Recapitulation of the general and special circumstances in its
favour-Causes of the general belief in the immutability of
species -How far the theory of natural selection may be
extended - Effects of its adoption on the study of Natural
history - Conclu.ding remarks 459-490

INDEX •• •• 491
How extremely stupid not to have thought of that!

-T. H. HUXLEY, on reading The Origin ofSpecies

If a man will begin with certainties, he shall end in doubts;

but if he will be content to begin with doubts, he shall end
in certainties.

-BACON, Advancement ofLearning

 AN H I S T O R I CA L S K E T C H O F T H E P R O G R E S S O F

O P INI O N O N T H E ORI G IN OF S P EC I E S

Two of the worst of all lines of English poetry, written in 1 799 by John
Hookham Frere:

The feather'd race with pinions skim the air­

Not so the mackerel, and still less the bear!

However poor that verse, it has a moral. The lines come from the poet's
somewhat neglected work "The Progress of Man; Poetry of the Anti­
Jacobin." Birds, bears and fish carry a political message. T hings are as
they are and it is folly to change them. The French Revolution disturbed
the God-given order; to proclaim the rights of man was as absurd as to
suggest that mackerel-or even bears-might fly.
A pair of quotations from The Origin ofSpecies, published sixty y ears
later:

It is conceivable that flying-fish, which now glide far through the

air, slightly turning and rising by the aid of their fluttering fins,
might have been modified into perfectly winged animals. If this had
been effected, who would have ever imagined that in an early tran­
sitional state they had been inhabitants of the open ocean, and had
used their incipient organs of flight exclusively, as far as we know,
to escape being devoured by other fish?
XVlll An Historical Sketch

In North America the black bear was seen by Hearne swimming

for hours with widely open mouth, thus catching, almost like a
whale, insects in the water . . . we might expect, on my theory,
that such individuals would occasionally have given rise to new
species, having anomalous habits, and with their structure either
slightly or considerably modified from that of their proper type.

Thus Charles Darwin on what evolution might do. Although it had

not yet made a whale-bear with a taste for aquatic insects, it could. After
all, evolution had already produced such improbable creatures as a fish
that skimmed the air. The idea oflife as fixed in a divine mold was dead.
Instead, all was change.
Before Darwin, the great majority of naturalists believed that species
were immutable productions, and had been separately created. Today, his
theory that they undergo modification and are the descendants of pre­
existing forms is accepted by everyone (or by everyone not determined
to disbelieve it) . Most people would, if asked, find it hard to explain why.
We all know that men and chimps are relatives and that plants, animals
and everything else descend from a common ancestor. The struggle for
existence, the survival of the fittest and the origin of species are wisdom
of the most conventional kind. Evolution happened; and, in 1 996, even
the Pope agreed (although he would admit only that "new knowledge
leads us to recognize in the theory of evolution more than a hypothesis").
All this is rather like Galileo and the earth's journey around the sun.
We know he was right, but what was his evidence? Why, in his dispute
with the Vatican, did he (as some say) mutter "eppur si muove'' "but still
-

it moves"? In the absence of high technology his proof was subtle. It in­
volved the movement of "wandering stars"-planets-against a back­
ground of fixed bodies deep in space. The pattern made sense only if the
Earth itself was a planet and not a stationary object around which the sky
turned. Such evidence, persuasive though it might be, is unknown to
most of those who believe his ideas.
Evolution is much the same. Although the notion is as simple as that
of the solar system, Darwinism is not the obvious explanation of how the
world works. Common sense tells us that life-like the Sun-revolves
around ourselves. The idea has but one fault: it is wrong.
Satellites make it impossible to deny the structure of the universe. In
 An Historical Sketch XIX

the same way, genes are a triumphant proof of the fact of evolution. Dar­
win's theory of common descent does for biology what Galileo did for
the planets. It was laid out in a book written for the general reader, the
only bestseller to change man's conception of himsel£ An idea put for­
ward in 1 859 is still the cement that binds the marvelous discoveries of
today. The Origin of Species is, without doubt, the book of the millen­
nIum.
Nowhere else is the case for evolution better put. Darwin called his
work "one long argument." Modern biologists can but agree. To read his
four hundred pages is to be amazed by how well their reasoning accom­
modates each new finding as it appears. The Origin's logic is as powerful
today as when it was written.
Evolution is inevitable. It depends on mistakes in reproduction. De­
scent always involves modification, because any copy, be it of a picture
or a gene, must be less than exact. Information cannot be transmitted
without loss, and a duplicate of a copy is, in its turn, less perfect than
what went before. To reproduce in succession an original again and again
is to make-to evolve-something new. What went in emerges trans­
formed by errors of descent, the raw material of biological change.
That ore is turned into precious metal by natural selection, the furnace
within which diversity is tried. Life is a struggle. As more individuals are
born than can possibly survive, a grain in the balance will determine
which individual shall live and which shall die. The slightest advantage
in any one being, at any age or during any season, over those with which
it comes into competition, will turn the balance. Natural selection is sim­
ple. It picks up inherited differences in the capacity to reproduce. If one
version multiplies itself better than others it will take over and, in the
end, a new form of life-a new species-will emerge.
Errors of descent are the stuff of evolution. Variation in the ability to
copy them-natural selection-gives it a direction. Nature does not
favor beauty, or strength, or ferocity; all it can do is to advance those best
able to multiply themselves. Although its products include the most
beautiful and most repulsive of beings there is no mystery to Darwin's
machine: it is no more than genetics plus time.
His evidence, however, is that of a century and a half ago and leaves
many gaps before his case can be considered proven. All-or nearly all­
have now been filled. My own book brings Darwin up-to-date. It is, as
xx An Historical Sketch

far as is possible, an attempt to rewrite The Origin of Species. I use its

plan, developing as it does from farms to fossils, from beehives to islands,
as a framework, but my own Grand Facts (another phrase beloved of
Darwin) are set firmly at the beginning of the twenty-first century. Dar­
wins Ghost tries to read Charles Darwin's mind with the benefit of scien­
tific hindsight. Its title hints at my aspiration: to be a ghostwriter is the
most that I can claim (and I have to disappoint those who assume that
this is a work channeled from beyond the grave by its great originator,
helpful though that would have been).
The theory of evolution unites biology as his millennium comes to an
end. Evolution has become more than a science, as its ideas are used, wit­
tingly or otherwise, in economics, politics, history, art and more. No ed­
ucated person can afford to ignore them; and they have no excuse for
doing so, as parts of the Darwinian story are so well told in the many ex­
cellent books that describe aspects of his theory for a general audience.
No book, though, tells it all; there is no modern and nontechnical treat­
ment of evolutionary biology, in all its depth and vast diversity. To
rewrite The Origin ofSpecies is more than most biologists would dare, but
daunting (and in some ways hopeless) though that task might be, I have
attempted it. If an apology is called for, I make it here. For my own book
I can make but one claim: it is the least original of its decade. So sturdy
is the model upon which it is based, however, that in this case to be de­
rivative is a strength and not a weakness.
The main difficulty in achieving my goal has been to know what to
leave out. The Origin marks the foundation of modern biology, and of
large parts of geology as well. I have tried to reflect its breadth and have,
no doubt, failed. My own volume tries to use Darwin's logic to illumi­
nate the discoveries of today. It is not a history of evolutionary ideas, or
of life; nor a biography of Darwin, or of animals and plants, but an ar­
gument that will, I trust, persuade my readers of the truth of evolution.
To keep it within bounds (and it is the same length as the original, with
much the same division among chapters) I have had to pick and choose,
and to omit certain topics in order to allow others a chance.
My standard for inclusion is a mention in Darwin's great work. I allow
a marginal exception. That volume contains but one substantial sentence
on humans ("Light will be thrown on the origin of man and his his­
tory"), but so much is now known about our past that I devote my final
 An Historical Sketch XXI

chapter to the subject. Darwin himself rules out several topics. His book
has nothing to do with the origin of the primary mental powers, any
more than with life itself. The beginnings oflife and of consciousness are
equally absent from my own pages. They also lack any discussion of the
Darwinian basis of society. Such topics, if they are part of science at all,
are not what most evolutionists see as evolution and are notable by their
absence from its technical literature. They must defer, if only on grounds
of space, to the objective truths of a century and a half of discovery in ge­
ology, genetics and all the rest.
It would be presumptuous to present this essay as more than a shadow
of its original, in content or in form. The Origin is the high point of the
literature of fact. Darwin wrote well because he read well. In a single
summer, his diary records, he enjoyed Hamlet, Othello, Mansfield Park,
Sense and Sensibility, Boswell's Tour ofthe Hebrides, The Arabian Nights
and Robinson Crusoe. His own prose is like a Victorian country house. It
radiates confidence from whatever direction it is viewed; as literature, as
autobiography or as brilliant science.
Compare Darwin's account of his first sight of the Galapagos with that
of Herman Melville, whose The Encantadas (another name for the is­
lands) was published in 1 854. Darwin is vivid and direct: "In the morn­
ing ( 1 7th) we landed on Chatham Island, which, like the others, rises
with a tame and rounded oudine, broken here and there by scattered
hillocks, the remains of former craters. Nothing could be less inviting·
than the first appearance. A broken field of black basaltic lava, thrown
into the most rugged waves, and crossed by great fissures, is everywhere
covered by stunted, sun-burnt brushwood, which shows litde signs of
life. The dry and parched surface, being heated by the noon-day sun,
gave to the air a close and sultry feeling, like that from a stove: we fan­
cied even that the bushes smelt unpleasandy."
Melville is, in contrast, feeble: "But the special curse, as one may call
it, of the Encantadas, that which exalts them in desolation above Idumea
and the Pole, is that to them change never comes; neither the change of
seasons nor of sorrows. Cut by the Equator, they know not autumn, and
they know not spring; while, already reduced to the lees of fire, ruin it­
self can work litde more upon them. The showers refresh the deserts, but
in these isles rain never falls. Like split Syrian gourds left withering in the
sun, they are cracked by an everlasting drought beneath a torrid sky."
XXll An Historical Sketch

Nobody could copy Darwin's language. I have not tried to do so (al­

though I have filched a few of his sentences in the hope of improving the
tone of my own work). As a hint at what is to be gained from reading
The Origin in the original, I include (where they exist, from the fourth
chapter onwards) his summaries, his own list of chapter contents and the
book's final "Recapitulation and Conclusion."
It is possible to read each sentence of The Origin ofSpecies unchanged
in a modern context in the hope of revealing meanings unknown to the
nineteenth century. I do not propose to do so. Although the structure of
its argument is intact and the order of chapters the same, I use Darwin's
masterpiece as a scaffold rather than a straitjacket. This is a postmod­
ernist treatment of evolution, with the strengths and weaknesses so im­
plied. Its architecture is of an earlier age, but it is constructed with today's
materials. The Origin is also, it must be said, a work of high Victorian se­
riousness, with no concession to any desire to be entertained. In these
more flippant times I yield to the temptation to leaven a scientific narra­
tive with tales from the curious history of evolution and those who
study it.
Students of evolution face another danger. Boris Vian's mystical novel
Froth on the Day dream has a character who dedicates his existence to the
petrified vomit of a thinly disguised Jean-Paul Sartre. Biologists (like
Marxists) share that inelegant habit. What, they ask, did the patriarch
mean? Could he be wrong, or is it forbidden even to suggest such a
heretical idea? To interpret sacred texts has a fascination of its own. I have
tried to avoid it.
Darwin is the best biographized of all scientists, and among that dull
and sometimes arrogant race stands out as both attractive and modest.
Even his family has an allure. His grandfather Erasmus published a the­
ory of evolution in heroic couplets and appears in Frankenstein: "The
event on which this fiction is founded has been supposed, by Dr. Dar­
win, and some of the physiological writers of Germany, as not of impos­
sible occurrence." He was a Lunatick, a member of a group that included
Joseph Priestley and Josiah Wedgwood. When not engaged in the dis­
covery of oxygen or the introduction of industry into England they de­
signed a machine "capable of pronouncing the Lord's Prayer, the Creed
and the Ten Commandments in the Vulgar Tongue." Robert Darwin,
Charles's father, was a physician who mixed with (and lent money to) the
aristocracy, and Charles's own grandson Bernard played golf for England.
 An Historical Sketch xxiii

This book is about Darwin's science, the heart of biology. Its roots are
in the past, but it is the key to the present. Its subjects include the AIDS
virus and the blue whale, dog shows and the garbage that floats in the Pa­
cific. Milton, some say, was the last man to know everything (or to know
enough about most things to discuss them with authority) . Darwin was
the last biologist who could claim that. His mind was, he said, "a ma­
chine for grinding general laws out of a large collection of facts." The
hundreds of books and papers referred to in the manuscript of which The
Origin was to be a "sketch" include The Cottage Gardener and Country
Gentleman's Companion, the India Sporting Review and the Philosophical
Transactions of the Royal Society of London. Charles Darwin wrote to
scores of people, expert and amateur, in search of information, and wove
their lore into his case.
Nobody could do that now. So great is today's knowledge that there
are no Miltons even of biology, no one who has sufficient command of
the field to debate it with any colleague, from whatever sphere. To un­
derstand evolution involves interests so disparate that it is impossible to
embrace them all. That is the joy-and the tragedy-of modern science.
Because we now know so much about how life works, evolution has
become the science of the exceptions. It would be tedious to consider the
feuds about details that consume the subject (although not to do so guar­
antees that every biologist will find something to dislike in this book) .
Even so, and bitter as the disputes are, no scientist denies the central
truth of The Origin, the idea of descent with modification.
Darwin did not have that comfort. He had to persuade an audience
unused even to the notion that life could change that it all shared a pedi­
gree. Against much opposition-the Daily Telegraph, no less, urged its
readers to vote against an election candidate who had given him a favor­
able review-he succeeded. There had been ideas about evolution before
Darwin's time, but he was the first to provide not just a mechanism but
the proof that it worked.
In spite of his twenty-year search for evidence, Darwin was so con­
scious of the gaps in his thesis that he might never have made it public.
His book is full of apologies: "To treat this subject at all properly, a long
catalogue of dry facts should be given; but these I shall reserve for my fu­
ture work . . . It is hopeless to attempt to convince any one of the truth
of this proposition without giving the long array of facts which I have
collected, and which cannot possibly be here introduced . . . I must here
XXlV An Historical Sketch

treat the subject with extreme brevity, though I have the materials pre­
pared for an ample discussion." Today's readers may feel a certain relief
that the promised book never appeared. By happy chance, Darwin was
stung into publication of a summary of his ideas by an unexpected letter
from Alfred Russel Wallace containing the same notion.
Powerful though his logic may have been, Darwin's great weakness was
his failure to understand what now seems simple. In 1 859, ignorance
about inheritance was as general as it had been for a thousand years. In
1 726, Mary Toft, of Godalming, saw a rabbit while she was pregnant.
Then, she said, she gave birth to one rabbit after another. After the first
dozen, George I sent his court anatomist to examine her. In his Short
Narrative ofan Extraordinary Delivery ofRabbets he attested the truth of
the tale and suggested that the animals had jumped down her fallopian
tubes. Mary Toft was soon exposed as a fraud (and became the subject of
a ballad by Pope and a sketch by Hogarth). In spite of a noble attempt to
transcend such tales, Darwin remained confused. In 1 866, Gregor
Mendel at last got things right, and his work, in its clarity and elegance,
even gained a mention in the Encyclopaedia Britannica. Darwin never
saw it.
Its importance was not appreciated until the first years of the twenti­
eth century. Darwin had a messy scheme of his own, based in part on the
mixing of substances present in the blood. He soon saw that it was
Hawed. A heredity based on dilution leads any useful character to be
thinned as the generations succeed. It would blur any emerging division
among species and evolution would come to a stop. Although Darwin
agonized about the problem he had no reason to be anxious. DNA
speaks a digital rather than an analog language and inheritance is based
not on liquids but on particles-genes-that can be recovered un­
changed at any time. Even a slight advantage can be summed over the
years. Genetics saved The Origin and its central question-how varieties
are transformed into species-can now be answered in Mendelian terms.
To modern biologists, species are republics of genes, separated from
their neighbors by sexual barriers. Any favorable change in the DNA­
an ability to manage on half the food, or to have twice the number of off­
spring-will spread to fill the state, but will never get into another one.
To define species by genes does not always work, because some are
caught in the act of promotion toward an identity of their own and be-
 An Historical Sketch xxv

cause distance can put a stop to sex even within the same one. Even so,
to interpret their origin in genetic terms is the keystone of the arch that
bridges the ancient gap between the study of inheritance and that of life's
diversity.
Darwin's thesis was that the world's variety arose, not from forgotten
disasters, but through processes visible today. For him, the present was
the key to the past. Evolution was driven by the simple, slow and potent
mechanism of natural selection. As this acts solely by accumulating
slight, successive, favorable variations, it can produce no great or sudden
modification; it can act only by very short and slow steps. What limit, he
asked, can be put to this power, acting during long ages and rigidly scru­
tinizing the whole constitution, structure, and habits of each creature­
favoring the good and rejecting the bad? With such a machine at its
disposal, nature had no reason to make leaps.
Geology persuaded Darwin that there was no need to calion ancient
cataclysms, be they biblical floods or gigantic earthquakes, to shape the
earth. A tiny stream, given long enough, could carve a giant valley and a
shallow sea make, as it dried, a plain a thousand miles across. Could not
life be formed in the same way? If landscapes could be transfigured by
slow change, so, surely, could flesh. The idea of a universe preserved since
the Creation was dead.
That belief made biology into a system of knowledge rather than a set
of random facts. Any theory with such ambitions was bound to attract
criticism. It did, and, ever since 1 859, it has continued to do so.
The Origin is two things: a bold statement of the idea of evolution,
and a work of persuasion as to how it took place. It contains a mass of
evidence that makes a compelling case for evolutionary change. In his
old age, faced with a wave of inconvenient discoveries, Darwin began to
complicate his ideas. His famous description of a swimming bear, quoted
on the first page of my book, conceals within itself an irony. The phrase
"almost like a whale" comes from the sixth edition of The Origin, pub­
lished in 1 872. In 1 859, Darwin was more confident. His Leviathan was
unrestrained: "I can see no difficulty in a race of bears being rendered, by
natural selection, more aquatic in their structure and habits, with larger
and larger mouths, till a creature was produced as monstrous as a whale. "
I base my own book on the clarity of his first edition, but that apologetic
"almost" is at the heart of biology.
XXVi An Historical Sketch

The fact of evolution has survived. It is a dance to the music of time

that unites all who join in. Its theme is more elaborate than once it
seemed. Darwinism is often rendered as a dignified waltz to the melody
of natural selection. That is less than fair to its author. Darwin himself
saw that descent with modification could happen in many ways. Acci­
dent might dictate what plants or animals arrive on an empty island, and
some variants might increase or be lost by chance. He was, however, con­
vinced that natural selection has been the main means of modification.
Species, to him, were but varieties writ large, a single step in the process
of slow change that unites biology.
Darwin's great idea-of life as a series of successful mistakes-is sim­
ple, so simple, indeed, that it seems almost impossible that it could make
such complicated things. Its enemies still maintain-without cease­
that evolution is so blind that it could never build an eye and that to un­
derstand the mystery of life (or, at least, of humankind) must demand
mysterious forces from outside the world of science. Such claims are easy
to dismiss, but they cast their venerable shadow wide. Although all biol­
ogists accept the truth of evolution, some have almost a compulsion to
complicate its ideas.
Darwin's theory has been much revised, but rarely to its advantage.
The idea of natural selection Cl$. evolution's sole agent and of all change as
gradual are each, perhaps, too simple. Faced with the facts of the 1 850s,
.

Darwin complained about "difficulties SC> grave that to this day I can
never reflect on them without being staggered." He has more to be con­
cerned about today. Charles Darwin's feet, like those of any idol, have
been much inspected for signs of clay and a few traces have been found.
However, those who hope to replace his ideas often fail to notice quite
how well his icon has lasted in the face of a century and a half's explo­
sion of knowledge.
Every evolutionist has, by definition, the benefit of hindsight. Some of
Darwin's problems have been solved, others restated in modern terms. To
read today's biology is, quite often, to relive the argument of The Origin
in modified language. There have, of course, been many shifts in opin­
ion, and an evolutionary "theory of everything" of the kind proclaimed
by physics is still far away. Even so, and in spite of the many wonderful
facts that illuminate the science, there have been rather few new ideas
since his time. As a result, this book emerges (somewhat to my surprise)
 An Historical Sketch XXVll

as a work of refreshing conventionality. Darwin's thesis is perfectly able

to support a century and more of scientific advance.
I have never met a biology undergraduate who has read The Origin of
Species. Even scientists, familiar as they are with its contents (or what
they believe them to be), tend to honor it in the breach rather than the
observance. It is, though, much studied by students of the humanities as
an element of a philosophy or an English literature course. There is noth­
ing wrong with that. It was, after all, quick to enter the literary canon:
Anna Karenina's last thought before she decides on suicide is, "Yes, the
struggle for existence and hatred is all that holds men together." Unfor­
tunately, the subject's bible is often presented for what it is not, a work
of metaphysics rather than of science. Darwin and Melville both say a lot
about whales, but to rely on The Origin in a philosophy class is like using
Moby Dick as a zoology text. One is fact, the other metaphor. The Col­
lege of Liberal Arts often finds it hard to tell the difference.
Evolution is to the social sciences as statues are to birds: a convenient
platform upon which to deposit badly digested ideas. It has been debased
since it began by those who use the idea to support their own creed. Dar­
winism was not the first (and will probably not be the last) science to be
abused for political ends.
Bishop Berkeley saw sociology as a branch of physics. In 1 7 1 3-soon
after Newton's Principi�he came up with the view of society as a par­
allel case of the universe. It was ruled by gravity, by a Law of Moral Force,
a "principle of attraction in the Spirits or Minds of men" that draws them
into "families, friendships, and all the various species of society." People,
like the planets, find distant objects less attractive than those close at
hand. If men are governed by the Earth's attraction (and to jump off a
cliff shows that they are) , why should society not be so? The Founding
Fathers saw in the United States Constitution "how by the attraction of
gravitation the various parts are held in their orbits, and represent in
Congress, the judiciary and the President a sort of imitation of the solar
system." Other thinkers preferred a "political anatomy," derived from
William Harvey (who discovered the circulation of the blood) . Two leg­
islative houses were needed, a stronger and a weaker, because the heart
had two ventricles, different in size.
Anyone who came up with a planetary or cardiac philosophy of life
would today be laughed out of court. Evolution has joined those sciences
XXVlll An Historical Sketch

in the dock. Walter Bagehot, in his Thoughts on the Application of the

Principles ofNatural Selection and Inheritance to Political Society, proposed
that "what was put forward for mere animal history may, with a change
of form, but an identical essence, be applied to human history." Most of
what his many successors claim about the same thing does no more than
restate the obvious in biological terms. The rest-whatever it might be­
is not science.
The Olympian vagueness of their notions is illustrated by the writings
ofTeilhard de Chardin. He linked biology to the Spirit of Christmas in
a gaseous envelope called the noosphere: "Life physically culminates in
man, just as energy culminates in life . . . The phenomenon of Man was
essentially pre-ordained from the beginning." The Origin does not have
much sarcasm, but the "Historical Sketch" that begins its later editions
mentions a Dr. Freke who had, in a paper of wonderful obscurity,
claimed precedence for its ideas. As Darwin says: ''As Dr. Freke has now
(1 86 1 ) published his Essay . . . the difficult attempt to give any idea of
his views would be superfluous on my part." That does for Teilhard and
his heirs too.
I once spent thirty years studying the evolutionary genetics of snails.
Although my research decorated the margins of the subject (and is so in­
cidental to this book as scarcely to appear in it), I still have no real idea
what makes them tick. Society is, it seems, easier to explain. Charles Dar­
win saw where the importance of his theory lay ("Species are not-it is
like confessing a murder-immutable") and was opposed to the naive
use of his ideas in human affairs. This book has nothing on the various
attempts, more or less infantile, to apply Darwinism to civilization.
Darwin and Wallace presented their ideas to the Linnean Society of
London in 1 858. They had little impact. Thomas Bell, a dentist with an
interest in reptiles and then president of the society, claimed in his review
of the year that it had not "been marked by any of those striking discov­
eries which at once revolutionise, so to speak, the department of science
on which they bear; it is only at remote intervals that we can reasonably
expect any sudden and brilliant innovation which shall produce a
marked and permanent impression on the character of any brand of
knowledge, or confer a lasting and important service on mankind."
Bell's lack of judgment is reflected in his own book, Kaiygonomia, or
the Laws of Female Beauty (with plates bound as a separate volume to
 An Historical Sketch xxix

allow them to be locked away from inquisitive eyes) . It listed "defects in

the Intellectual system of Women (4); Defects in the Mechanical system
ofWomen (17) and Defects in the Vital system of Women (9) . " The rep­
tilian dentist had failed to notice the alibi that his society's paper was to
provide for students of society. He dismissed the talk that was to change
biology for ever, and went on to note that "a Bacon or a Newton, a Davy
or a Daguerre, is an occasional phenomenon, whose existence and career
seem to be specially appointed by Providence." Darwin and Wallace did
not count.
The sad truth is that the idea of descent with modification did not
need Providence, or Darwin, at all. Sooner or later, like any discovery, it
would have appeared in another guise. In science, revolutions are bound
to happen. Nowadays, no biologist could work without Darwin's theory.
Evolution is the grammar of their science. It accepts his painfully recog­
nized fact that life, like the English language, works to rules that, even if
filled with exceptions, make sense.
In spite of the Thomas Bells of this world, science (unlike the arts) can
be detached from those who do it. For that reason, I refer to no living sci­
entist by name. lowe a debt of gratitude to the many friends and col­
leagues who have commented on (and often disagreed with) my
manuscript. They include Douda Bensasson, Sam Berry, John Brook­
field, Bryan Clarke, Michael Coates, Jerry Coyne, Andrew Leigh-Brown,
Adrian Lister, Ursula Mackenzie, James Mallet, John McCririck,
Michael Morgan, David Parkin, Norma Percy, Mark Ridley and Kay
Taylor. All have been gracious about my use of their time and, too often,
my reluctance to accept their criticisms.
Darwin's ability to rule over his disciples from beyond the grave is such
that I hope that he will be forgiven an occasional appearance in this vol­
ume. His spirit is on every page.
\
 ON

THE ORIGIN OF SPECIES

INTRODUCTION

According to a 1991 opinion poll, a hundred million Americans believe

that "God created man pretty much in his present form at one time dur­
ing the last ten thousand years." A large majority saw no reason to op­
pose the teaching of creationism in schools. They followed in a long
tradition. A text of 1923, Hell and the High Schools, claimed that: "The
Germans who poisoned the wells and springs of northern France and
Belgium and fed little children poisoned candy were angels compared to
the text-book writers and publishers who are poisoning the books used
in our schools . . . Next to the fall of Adam and Eve, Evolution and the
teaching of Evolution in tax-supported schools is the greatest curse that
ever fell upon this earth."
Fifty pieces of legislation tried to put a stop to the subject. All failed.
Undeterred, Alabama called for a note to be pasted into textbooks: "This
book may discuss evolution, a controversial theory some scientists give as
a scientific explanation for the origin of living things, such as plants, an­
imals and humans . . . No one was present when life first appeared on
earth. Therefore, any statement about life's origins should be considered
as theory, not fact." In 1999 the Kansas Board of Education voted to re­
move evolution from the school curriculum and no doubt other states
will try similar tricks.
Such intolerance is new. At the end of the last century few clerics op­
posed the idea of evolution. In spite of polemic against a "genealogical
table which begins in the mud, has a monkey in the middle and an infi-
2 Introduction

del at the tail" most were ready to accept a compromise between The
Origin and the Bible. A Day of Creation might be millions of years long,
or might represent six real days that marked the origin of a spiritual Man
after the long ages it took all else to evolve. Real bigotry had to wait for
modern times.
The creationist movement is part of a triumphal New Ignorance that
rules in many places, the United States more than most. In fact, the ma­
jority of those determined to tell lies to children believe in Darwin's the­
ory and understand how it works, without noticing. Evolution is
embedded in the American consciousness for a simple and terrible rea­
son. For the past two decades the nation has lived through an episode
that has, with extraordinary speed, laid bare the argument of The Origin
ofSpecies. The organism involved was unknown in the nineteenth cen­
tury, but is now familiar. It is the AIDS virus.
Creationists find it easy to accept the science of AIDS. Its arrival so
close to the millennium and the Last Judgment is a useful illustration of
God's wrath. Homosexuals, they claim, have declared war on nature, and
nature has exacted an awful retribution. Fundamentalists admit the evo­
lution of a virus as nature's revenge but will not concede that the same
process acts upon life as a whole.
Even to anti-evolutionists, AIDS is proof of descent with modification
because they can see it happening. Its agent has changed in its brief his­
tory and has adapted to overcome the many challenges with which it is
faced. As death approaches, a patient may be the home of creatures--de­
scendants of those that infected him-as different as are humans and
apes. Every continent, with its own sexual habits, has its own exquisitely
adjusted set of viruses; and AIDS has relatives in animals quite different
from ourselves. Darwin would have been delighted to see the workings
of his machine so starkly exposed.
Science makes patterns from ideas. If AIDS can evolve, so can any­
thing else. The Origin uses freshwater bears and flying fish to make a case
that applies to all forms of life. For its opponents, in contrast, what is true
for viruses cannot be true of birds or fish, let alone a man. The existence
of an animal as unlikely as a whale is, for them, proof that evolution does
not work.
The other view of the origin of whales, men or viruses is simple. As
many more individuals of each species are born than can possibly survive
 Introduction 3

and as, consequently, there is a frequently recurring struggle for exis­

tence, it follows that any being, if it vary however slightly in any manner
profitable to itself, under the complex and sometimes varying conditions
of life, will have a better chance of surviving, and thus be naturally se­
lected. From the strong principle of inheritance, any selected variety will
tend to propagate its new and modified form.
Every part of Darwin's thesis is open to test. The clues-from fossils,
genes or geography-differ in each case, but from all of them comes the
conclusion that the whole of life is kin. That is no mere assertion, but a
chain of deduction with every link complete. The biography of the AIDS
virus, one of Nature's newest and tiniest products, is almost complete
and that of whales-the largest animals ever seen-is fragmentary, but
they are cousins under the skin. The AIDS virus is change seen under the
microscope, and the whale the same process viewed, in glimpses and over
long ages, through a biological telescope. Evolution at the extremes of
size is an apt prelude to the great drama that is Darwinism.

Creationists often deny the possibility of an intermediate between two

species. Take whales and land animals. What use are flippers on solid
ground, or feet in the sea? "There are simply no transitional forms in the
fossil record between the marine mammals and their supposed land
mammal ancestors . . . It is quite entertaining to attempt to visualize
what the intermediates may have looked like. Starting with a cow, one
could even imagine one line of descent which prematurely became ex­
tinct, due to what might be called an udder failure." The complaint (and
the leaden humor) is not new. A London newspaper of 1859 said of Dar­
win's "whale" passage that "With such a range and plasticity . . . we know
not where to stop-centaurs, dryads and hamadryads and (perhaps)
mermaids once filled our seas."
Nobody has ever seen a mermaid, or even a dinosaur. Evolution is,
most of the time, an attempt to reconstruct a history whose pace is far
slower than that of those who study it. �DS is unique because genes and
time come together on a human scale. Darwin himself saw disease as a
model of change. Almost the first recorded hint of his theory is in a note
made on the Beagle. He was told by the surgeon on a whaling ship that
lice from Sandwich Islanders will not survive on Europeans. How, he

�
4 Introduction

asked, could this be-unless each had diverged from the same ancestor?
Why should a Creator, if parasites were needed, not make a universal
louse for all mankind?
AIDS came to notice in 1 98 1 with a report of a sudden increase in a
certain form of pneumonia. As the sober language of the Morbidity and
Mortality �ek(y Report of the United States Centers for Disease Control
put it: "The fact that these patients were all homosexuals suggests an as­
sociation between some aspect of homosexual life-style or disease ac­
quired through sexual contact and Pneumocystis pneumonia in this
population." The illness became notorious with the death of the actor
Rock Hudson in 1 985. By then, more than twelve thousand Americans
were dead or dying. Within a decade, half a million had perished. No­
body guessed that such a rare disease would become a pandemic. Camus,
in The Plague, has it that: ''A pestilence isn't a thing made to man's mea­
sure; therefore we tell ourselves that pestilence is a mere bogey of the
mind, a bad dream that will pass away. But it doesn't always pass away,
and from one bad dream to another it is men who pass away." They did
and, more and more, they will.
AIDS, like the Great Pox of the fifteenth century, is spread by sex. The
ground was well prepared before its seeds were planted. In the 1 970s, five
thousand gay men moved to San Francisco each year. By 1 980, venereal
disease was widespread-and four out of every five of the patients were
homosexual men. A typical AIDS victim admitted to sex with eleven
hundred people in his lifetime, while some claimed as many as twenty
thousand partners. Most of the city's homosexual males had the viral ill­
ness known as hepatitis B, and many suffered from gay bowel syndrome,
multiple gut infections acquired from the curious sexual habits of part of
their community. Casual sex in bathhouses-the Cornhole, the Boom
Boom Room, the Toilet Bowl-helped the diseases to spread. AIDS,
though, was new.
It was greeted with hysteria. Some claimed that the virus had been
placed in Tutankhamen's tomb to punish those who defiled his grave and
had come to America with an exhibition of his treasures. An analyst stud­
ied what he called its psycho-incubation. AIDS victims, he said, had suf­
fered an emotional emergency in childhood that made them feel
abandoned and later led to illness. The editor of Burkes Peerage went fur­
ther. To preserve the purity of the human race his publication would not
 Introduction 5

list any family in which a member was known to have the disease: "We
are .worried that AIDS may not be a simple infection, even if conveyed
in an unusual way, but an indication of a genetic defect."
Although some dissenters tried to associate its symptoms with the use
of capsules of amyl nitrate to enhance erotic pleasure, the real cause was
soon found. The culprit is a virus, the human immunodeficiency virus,
or HIY.
Like a whale, the virus is built on an inherited plan coded by genes,
each one liable to accident every time it is copied. HN is unusual even
among viruses. As a retrovirus, its genes are based not on DNA, but on
its relative RNA (a molecule used in most creatures to translate, rather
than to transmit, the genetic message). All retroviruses-and they come
in many forms-contain about ten thousand RNA units, or "bases." The
AIDS virus subverts its host's cells. It forces them to make replicas of it­
self with an enzyme whose job is to copy information from the invader's
RNA into human DNA. Each new particle hides itself in a cloak of cell
membrane into which it inserts a protein. This is the key to the infection
as it fits into matched molecules on the surface of blood cells and opens
the door to their interior.
The lock that turns to an enemy's key is most abundant on certain
cells of the immune system. These multiply in response to infection, but
cannot cope with the challenge. Billions of new particles are made each
day, and although most are at once destroyed, they soon prevail. Soon
after the virus arrives, the number of protective cells falls, only to rise as
the body's fight back begins. Then, the immune system begins to col­
lapse. The first sign of illness is a malaise no worse than influenza. This
clears up, but HIV stays at work. As the defenders are driven back, other
diseases gain a hold. For most people, the transition from inf<E.ction to
overt illness takes from six to ten years.
As AIDS advances there may be pneumonia, �gaI r /Cft-infections,
:!�� diar­
rhea, weight loss and a viral form of blindness. }i cancer called Kaposi's
sarcoma, otherwise found among aged Jewish men, quite often appears.
Its first sign is purple marks on the skin, but as it progresses it kills. Ka­
posi's sarcoma is caused by a herpes-like virus common in the homosex­
ual community. It gains entry because the body's defenses have been
undermined. If the patient does not first surrender to a fungus, bac­
terium or cancer, he wastes away.
6 Introduction

The history of AIDS, over days, years and centuries, is simple. It in­
volves descent, accompanied by modification. Each virus divides once a
day. Mutation is followed by natural selection that allows the invader to
adapt to the body's defenses, to the drugs used to treat it and to the sex­
ual habits of the society in which it lives. Some changes are, it seems, un­
heeded by selection and build up at random as the generations pass. In
time (and it does not take long) new forms of virus emerge.
The genes tell the story. They link a patient with the person who in­
fected him, with others long dead, and with the viruses of apes, cats and
whales. Except in its details, and the trivial matter of size, the evolution
of the AIDS virus is that of every other being.
The disease is not very contagious. The chance of infection for a
woman each time she has sex with an HIV-positive man is one in several
hundred; and a stab with a contaminated needle is more likely to pass on
hepatitis than to transmit the virus. Nevertheless, by 1 992 it was the
main cause of death for young American men. By the end of that decade,
one in a hundred of the world's sexually active population carried it,
most as yet unaware of their plight. Ignorance played a part: many Tan­
zanians, for instance, believed that insecticide sprays protected against
contagion. The epidemic has moved on from the bathhouses. Most cases
are in the tropics, with a new outbreak in the states of the former Soviet
Union. There, drug injection is rife and the incidence of syphilis (a sign
of an AIDS outbreak on the way) has shot up by a hundred times.
The long incubation period means that no Third World country has
yet faced the full truth of what is to come. In London, one person in six
died in the last great epidemic, the Great Plague of the 1 660s, which
then faded away. By 1 998 half the adult population of some Mrican
cities was HIV-positive. AIDS is already Mrica's greatest killer and the
continent's death rate may increase by five times in the next decade. In
Botswana, children born today have a life expectancy of twenty years.
Without AIDS, the figure would be nearer seventy. The virus has entered
the public domain. It will be almost impossible to drive it out.
AIDS is an illness of social change; of travel and of promiscuity. In the
United States most infections are passed on by homosexuals, by drug in­
jectors and by those unfortunate enough to receive a transfusion of con­
taminated blood. In Mrica, India and Asia almost all cases come from sex
between men and women or by transfer from mothers to babies. Like all
 Introduction 7

diseases-or flowers or songbirds-the virus is delicately adjusted to the

challenges it must face.
HN is bad at making exact copies of itself, which is one reason why
it does so well. The error rate of its copying enzyme gives it a mutation
rate a million times higher than that of its host. The virus is, like Mr. Mi­
cawber, always waiting for something to turn up, and-most of the
time-it does. Each new particle has, on average, a single change in its
RNA. As a result, any patient soon contains a vast diversity of intruders.
Most are defective and do not survive, let alone infect anyone else. Oth­
ers withstand all that the body or human ingenuity can do.
A patient is an island to which its invaders must adapt or die. Natural
selection is hard at work from the moment they arrive. The immune sys­
tem is good at its job and billions of HN particles succumb to it. How­
ever, a few emerge unharmed because genetic accidents alter their signals
of identity. They are missed by the body's defenses, multiply and prevail.
Selection also explains why drugs failed for so long. There has been no
lack of research. Fifty times as much has been spent per AIDS death in
the United States as on each American who dies from stroke. The main
conclusion from years of work and millions of dollars is that evolution is
a relentless foe.
The first drugs attacked the copying enzyme in the hope that if the
virus could not translate its information it would fail. Others used chem­
icals that look enough like components of DNA to fit into the chain as
it grows and to stop it dead. Some treatments used "anti-sense" nucleic
acids, mirror images of genetic information that can bind to viral genes
and block their action. Much has been spent, to little effect, on the search
for a vaccine.
As years went by with no success, desperate patients turned to quacks,
who stimulated their immune systems with ozone blown into the rectum
or with expensive potions made of ground-up tortoise shells. Now, at
last, there is hope. The survival of Americans with the disease has dou­
bled in the past decade and in Europe the death rate is a fifth of what it
was. Soon, doctors dared to ask whether the virus itself could be wiped
out. In some people a combination of drugs reduced it to undetectable
levels and-perhaps-Ied to a cure. The optimism did not last. The
viruses hide in the blood cells and reappear when the medication is
stopped.
8 Introduction

Now, for those able to withstand (and, at ten thousand dollars a year,
afford) the treatment, the illness can at least be kept under control. In
1 997 the gay newspaper The Bay Area Reporter was, for the first time in
two decades, able to use the headline "No Obituaries." The therapy re­
quires dozens of pills a day, and to stop even for a short time allows the
virus to rebound. Many abandon the treatment as it is so exacting, but
the days when diagnosis meant death are over.
Every AIDS patient is a monument to the theory of evolution. Nat­
ural selection alters the identity of the virus as the disease progresses.
Drugs, too, lead to evolutionary change.
The best evidence for a theory comes when two experiments give the
same result. The drug Ritonavir was introduced in the mid- 1 990s. At
first, there was success, with-in most patients-the number of viruses
reduced by a hundred times. Within months, all suffered an ominous
climb back to the original levels and, as the treatment lost its power, the
disease continued its course. Every virus, from London to San Francisco,
gained its resistance with the same mix of four mutations at different
points in its genome. What is more, and although the genetic accidents
occurred at random, evolution utilized them in the same order each
time. It must wait first for a change in a site to which Ritonavir attaches.
That causes only a slight improvement in the virus' ability to cope, but
is at once seized upon by natural selection. The three subsequent alter­
ations, too, are picked up in the same sequence in every patient. In com­
bination they increase resistance twentyfold.
Each point in the viral RNA has an error rate of around one in ten
thousand. The chance of the four changes happening at once is that fig­
ure, multiplied by itself four times-one in ten million billion, a total
greater than the number of particles made in the entire course of an ill­
ness. It could never be reached by the accidents of mutation in a single
individual, let alone within the hundreds who have evolved resistance.
Evolution triumphs because it turns to natural selection, the plodding
accumulation of error.
AIDS is Darwinism unadorned: a faulty copying machine that alters
faster than its opponents. The shadow of evolution also lies over the epi­
demic as a whole. For HN, in its dawn, with San Francisco in a new
Summer of Love, it was bliss to be alive. Any agent of infection was guar­
anteed a welcome. The pressure was on, and a gene that helped a virus to
attack a new victim was favored, whatever harm it did to its host. From
 Introduction 9

the viral point of view, the death of a patient (and his HIV particles)
meant little, as long as he had already infected dozens more.
In those early days, virulence was all. AIDS became more lethal as the
epidemic grew. Americans infected in the plague's first few months sur­
vived for longer than those who caught it later. By then, selection had
brought to the fore the variants quickest to copy themselves even if they
killed their carriers, because they spread faster than the others. Soon, ho­
mosexuals became more cautious. For the virus, the change in behavior
was bad news because it made it more difficult to reach a new host. The
new erotic environment forced the disease to be kinder. Variants slow to
kill were favored because their bearers lasted long enough to pass the
Virus on.
Where did AIDS come from? The records reveal a fragment of its his­
tory, but the genes say much more. They reveal a shared past of descent,
accompanied by modification. The early epidemic in Europe infected
not just homosexuals but African immigrants who belonged to quite a
different sexual community. Why should it attack these disparate
groups?
Nucleic acids gave the answer. The "signature site," a length of three
hundred or so RNA bases, changes at great speed. It was a hint that AIDS
is more than one disease. When any group of organisms-viruses, mice,
butterflies or whales-is studied in enough detail, what seems at first
sight an entity often turns out to be a set of distinct but related organ­
isms; in other words, different species. HIV is no exception.
Some humor to relieve a grim story. A man goes into a Szechuan
restaurant in Aberystwyth (a town on the western edge of Wales) and is
served by a Chinese waiter who speaks perfect Welsh. Beckoning the
boss, the customer asks where he found this prodigy. The answer: "Keep
your voice down, he thinks he's learned English!"
In other words, from a Chinese speaker's point of view, Welsh and
English are mere dialects of each other, each just as easy or hard to un­
derstand. They are members of the Indo-European group of languages,
descended from a common ancestor. Although anglophones find the
Welsh tongue impenetrable, the only way to test how distinct it might
be is to put it into context, with Chinese as a separate group with which
Welsh and English can be compared. The difference between the two is
then seen, in global context, to be tiny.
There is a hidden structure in the languages of the world. The various
10 Introduction

degrees of difference of those from the same stock can be expressed by

groups subordinate to groups, and the proper or even only arrangement
would be genealogical. This would be strictly natural as it would connect
"
together ali languages, extinct and modern, and would give the filiation
and origin of each tongue. British and American English are closer than
English and French, and those two Indo-European dialects are more dis­
tant relatives of Greek. Greek, in its turn, is not much like Urdu, but
each retains enough of its past to hint at a common ancestry. The ances­
tral language can itself be reconstructed from the hints held in its much
diverged descendants.
Information is transferred by genes in much the same way as it is by
words. For each, evolution is inevitable. Literary fragments-fossil
speech-and some daring guesses about the rate of change reveal the
family ties. French and English split not long ago (to Dumas, after all,
English was just French badly pronounced); and, strange though it
sounds, there is also nothing special about Welsh. It broke away from its
sisters long after Chinese separated from the ancestor of them all . Lan­
guage is the key to history. If we all spoke the same one-Chinese, Welsh
or English-and there were no records of the past, it would be impossi­
ble to tell where our shared tongue came from. Of course, we do not. In­
stead, descent with modification uncovers the biography hidden in every
sentence and reconstructs the past of those who speak them.
For the human immunodeficiency virus, changes within a patient-a
mere shift in genetic accent-or in the particles that pass from one vic­
tim to the next track its evolution over months or years. If (and the as­
sumption is a bold one) the great limbs of the AIDS tree follow the same
rules as those of its twigs, the epidemics of today, it should be possible to
work out the past growth of any branch, however ancient. To compare
the human immunodeficiency virus with its distant relatives-the Chi­
nese speakers of the viral world-hints at the origin of the plague itself
Genes reveal two separate human immunodeficiency viruses. They
differ in almost half the letters in the signature site. The virus involved in
the homosexual outbreak is known as HIV- l , that mainly responsible for
the illness in African heterosexuals as HIV-2. Each is divided into sec­
ondary clusters. HIV- l (like the Indo-European languages, with their
eastern and western offshoots) has two great divisions. The first is uni­
versal, while the other, much rarer, has its home in Cameroon, Gabon
 Introduction 11

and Equatorial Guinea. A third distinct type was found in 1 998 in just
two people in Cameroon. The main cluster contains ten or so subtypes
(HIV- 1A, - l B and so on), each of which diverges by up to a quarter in
the genetic autograph. Minor variants in signature disclose the presence
of distinct strains of each subtype.
The virus has a structure of relatedness that traces its roots further and
further into the past. Its hint at a grand natural system proclaims quite
plainly a shared descent from common parents. As the modified descen­
dants proceeding from a single progenitor become broken up into
groups within groups, the growth rate of the young shoots of the tree of
illness can reveal its distant past.
A species can originate but once. Few biologists are lucky enough to
see it happen, even at the giddy pace of evolution among viruses. How is
it possible to trace the origin of a disease most of whose victims are dead?
Nucleic acids tell the story. They reveal a recent eruption of diversity. In
a family tree of the agent of AIDS drawn for the United States, differ­
ences explode, like a firework, from a common source-the mark of the
disease's sudden arrival and rapid spread. The pattern of global related­
ness, too, looks more like a shrub than an oak. For HIV- 1 , all the main
branches join in a common node. The HIV-2 pedigree looks much the
same. For each, the point where the shoots all meet marks the origin of
an epidemic.
A chronometer is hidden within all words, and in each length of nu­
cleic acid. It depends on a simple and regular mechanism, the buildup of
errors with time. If the moment when its hands were set is known and
the machinery ticks smoothly, its rate can be measured. This molecular
clock, as it is known, can then be used to estimate when any member of
the HIV family diverged from any other. The timer's rate varies, in part
because patients who die soon after infection leave no time for it to tick,
while those who struggle on for years undergo more evolution. Even
worse, it seems to move at different speeds in different subtypes. Never­
theless, the clock in the genes makes sense, and HIV- l samples collected
in the 1 990s have moved on from those taken ten years earlier. To turn
back the hands to zero puts the start of the global outbreak at some time
in the 1 940s. HIV-2, the genes show, began its international career at
much the same moment.
The earliest known specimen of the human immunodeficiency virus
12 Introduction

was found long after the death of its victim. It came from a fossil, the pre­
served remains of an anonymous African inhabitant of the Congo city of
Leopoldville (now called Kinshasa and the capital of the Democratic Re­
public. More than a thousand blood samples left over from the first years
of Congolese independence were tested, but although several were HIV­
positive, just one, from a patient who died in 1 959, retained any viral
genes. It was quite similar to the common ancestor of today's HIV- l
viruses as reconstructed from the genetic signatures o f their descendants.
AIDS, it seems, started within an African (a discovery greeted by the
Ghanaian Times as "a shameful and vulgar attempt to push this latest
white man's burden onto the door of the black man") . The similarity of
the reconstructed and the actual ancestor suggests that the global epi­
demic began soon after the Second World War, perhaps from a single
virus particle. Startling as that seems, it is no more than what happens
every few years as new waves of Asian flu sweep across the world.
The virus did not take long to escape from its native land. Only one
of its routes out of Africa has been explored. In 1 976, a young Norwe­
gian sailor died of a mysterious illness. A sample of his tissue was, many
years later, found to contain traces of the Cameroon branch of HIV- l .
He was infected on a trip to West Africa, soon after his fifteenth birth­
day in 1 960. The voyage gave him plenty of chances for sex. His records
show that, somewhere on the journey, he caught gonorrhea. The doctor
missed his other, fatal, illness.
Fourteen years after the young Norwegian brought home his viral
cargo, a bisexual German musician who liked to hire prostitutes to take
part in orgies became the first European to be diagnosed with AIDS. The
sailor was by then a truck driver. He often visited Cologne, the orgiast's
home town. The next victim was a French barmaid from Reims. Reims,
too, was on his regular route. The history of a dead Scandinavian, pre­
served in a bottle, uncovers one viral pathway into Europe. Most of the
others will never be found.
The genes show that the invasion force involved j ust a few intrepid
voyagers. All the subtypes of HIV- I are found in its native continent, but
other places have only a few. The earliest samples taken in North Amer­
ica and Europe all belong to subtype 1 B, proof that a small number of
travelers brought the virus from Africa to the developed world.
Once in a newfound land the migrants must evolve to cope. HIV- l ,
 Introduction 13

the killer of the Western world, has remained an exclusive beast. Each of
the eight or so modern subtypes sticks to its own community and has
adapted to fit its sexual tastes. In Thailand, HN- 1 B was the main form
at the start of the 1 990s. It had become something of a specialist at travel
by the anal route but is, in its new nation of sex tourists, in retreat before
HN- 1 E, a virus that prefers conventional sex. In Russia, drug users have
four subtypes of HIV- 1 , while female prostitutes and homosexuals each
lay claim to their own.
AIDS' ability to cope with human vice is helped by its own sex life.
Sex, in all its guises, is no more than a way to mix up genes. If someone
is infected with two HN subtypes, the viral enzymes can reshuffie their
RNA to yield a generation of viruses with new combinations of genetic
material. About one HN- 1 subtype in ten is a blend of the RNA of
other, older, forms. HN- 1 E-today's Thai form-is a hybrid between
the African HIV- 1A (which has not itself reached Asia) and an undis­
covered donor of the gene that makes the protein coat of the Thai virus.
Some strains are a patchwork of genes from four or more sources.
One definition of what species are (and it can be hard to tell) turns on
sex. If two individuals-viruses or whales-can blend their genes to
make young with elements from each, they belong to the same species.
If they cannot, they are distinct. People who pick up both HIV- 1 and
HN-2 (and such unfortunates do exist) never produce hybrids. In con­
trast, those inflicted with, say, HIV-IA and HN- 1 B often generate new
mixtures. In the world of traditional biology, HN- 1 and HN-2 would
be defined as distinct species, 1A and 1 B as varieties of the same one. A
mutation for drug resistance within HIV- 1 might spread to every sub­
type of that virus, but would never enter HN-2. Descent with modifi­
cation has gone so far that what was once an entity has been divided into
two; and a virus has completed its journey along the Darwinian road to
an identity of its own.
Much of the story of AIDS is evolution on a human scale, the tale of
an opportunist in the modern world. Its genes also hint at its remote an­
cestors.
Primate genes show that chimps, humans and gorillas are close kin,
while orangs are more distant and monkeys further still from that
almost-human trio. The monkeys themselves have a deep split between
the Old World and the New. However, the family trees of host and the
14 Introduction

AIDS-like viruses found in most African primates are not at all alike. The
disease and its victims have, it seems, trodden different paths, and AIDS
must have entered humans from another animal.
An accidental experiment shows how a change of scene can lead to dis­
aster. A harmless African monkey virus was passed in error to Asian mon­
keys. They all died within a few months. When blood from the first
Asian monkey was then injected into a second victim and from him in
rapid series to a third, fourth or fifth, the virus became more lethal at
each step. Later generations of monkeys died in weeks. As with the gay
men of San Francisco, rapid transfer favored the viruses fastest to copy
themselves, whatever harm they did. In one zoo, macaques picked up an
AIDS-like illness from the talapoin monkey. In the talapoin the virus is
harmless, but in macaques it kills-and moves from animal to animal by
simple contact, with no need for sex. Humans have, so far, been spared
such a fate.
Genes show that HIV- 1 and HIV-2 are not much related and come
from different sources. HN- 1 resembles a virus found in West African
populations of chimpanzees, while HIV-2 is closer to another from the
sooty mangabey. They entered humans more than once. The three main
groups of HIV- 1 each resemble different lineages of chimp virus, and
HN-2 has half a dozen distinct types. Other primates have their own
viruses that do little harm to those who bear them.
Why should a primate virus attack humans? Plenty of hunters have
been bitten by monkeys, and Victorian travelers ate them with more or
less enjoyment (although one found "something extremely disgusting in
the idea of eating what appears, when skinned and dressed, so like a
child") . The great French hunter Paul Du Chaillu recorded that in 1 86 1
the Fang people o f Cameroon gave up the purchase of corpses as food in
favor of gorillas. Those animals are now protected, but the Fang still talk
of gorilla tongue as a delicacy and, with today's opening of the forests by
timber companies, there is a new call for "bush meat" to feed their work­
ers. Ape meat has also become a popular restaurant dish in parts of
Africa.
Many-perhaps most-diseases come from animals: rabies from
dogs, anthrax from cattle and Lyme disease from deer. To have a pet is a
good way to become ill. Parrots give psittacosis to their owners, and pet
snakes often pass on salmonella. The god Thoth, inventor of speech, was
 Introduction 15

worshipped in the form of a baboon. In the Pithecussae, the Monkey

Cities, those animals, it was said, waited at table and no doubt bit their
customers from time to time. Pets, though, may not be to blame. Some
claim that AIDS is an epidemic sparked off by scientists who used mon­
keys to develop vaccines, or by doctors who used unclean needles in the
fury of vaccination in Mrica after World War II.
However it began, the human immunodeficiency virus must be far
older than any of its extant strains. The fossil virus from Kinshasa is more
similar to modern HN- 1 than to any chimp virus, so that the transfer to
humans happened long before the young Congolese met his fate. Per­
haps, over the centuries, there were local outbreaks of AIDS in Mrica,
each the result of a separate infection from an ape. They did not spread
in an age without travel, transfusion or frantic promiscuity. Not until the
1 9 50s could the illness expand to fill the world. Genes also hint at its pre­
history. Many animals have viral diseases of the immune system, and a
virus quite similar to those of primates causes a similar affiiction in cats.
In the 1 980s such an epidemic led to the death of thousands of Mediter­
ranean dolphins. Its cause was a distant relative of the agent of AIDS.

The human immunodeficiency virus contains in its brief history the en­
tire argument of The Origin ofSpecies: variation, a struggle for existence,
and natural selection that in time leads to new forms of life. Geography
tells part of its story, as do fossils, and its genes are a link to distant rela­
tives with which it shared an ancestor long ago. They reveal a hierarchy
of order as evidence of descent from a common source pushed further
and further into the past.
Our lives are too short to understand the evolution of other beings in
such detail. Take the aquatic bear for which Darwin suffered such mock­
ery. Could it ever have made its way toward the sea? "What does it take to
become a whale, to live at the other end of the scale from a virus? Now,
we know.
"Whales, like all mammals, breathe air and give milk. "When did they
take to the water and what were they before they made the move? Their
new life involved more than a change of medium. They grew, to a hun­
dred and fifty tons in the case of the blue whale (which is to humans as
we are to mice) . The skull and neck became shorter and the nose moved
16 Introduction

backward. The ear closed and sound now passes through a layer of fat.
Legs evolved into fins, with extra bones in the back to match. Beneath
the skin were other changes. The deepest diver can make it to four hun­
dred feet without artificial aids and holds his breath for a few minutes to
do so. The sperm whale dives to a mile and more and can stay under for
two hours. The change is in its chemistry rather than its lungs. Whale
muscles contain large amounts of myoglobin, a protein that pulls oxygen
from the blood. Their oxygen is kept not as a gas, but as a chemical com­
pound. This in turn allows the lungs to collapse at depth as a defense
against the bends (nitrogen bubbles in the blood) as the animal comes to
the surface.
The remains of extinct creatures mark each step in the move from land
to sea. The fossil evidence is confirmed by the record of the genes. If
whales survive, their history will soon be as well understood as is that of
the AIDS virus. The technology is much the same, and the evolutionary
logic is identical.
Most of the fossils suggest that the distant ancestors of whales were
hyena-like beasts called mesonychids, scavengers for carrion and hunters
of fish. They underwent a radical change of habit. The Simla Hills of
Northern India, with their cool mountain climate, were a holiday haven
for the British rulers of the Raj. There, several thousand feet above sea
level, was found a fifty-three-million-year-old jawbone from Himalaya­
cetus, the first known ancestor of today's whales. The fossil came from a
beast that seems to have spent time both in fresh water and in a long-lost
sea.
A fifty-million-year-old skull discovered in the Kala Chitta Hills of
Pakistan came from an animal further on the way to whaledom. Paki­
cetus, as it was called, lacks the fatty earplugs of its descendants. Its days
were passed between land and water, with an inner ear midway between
those of whales and of land animals, allowing it to hear both in the air
and beneath the surface. Those oldest whales lived in a vanished ocean,
the Tethys Sea, now replaced by India and Pakistan, its floor thrust into
the skies to build the highest peaks in the world. Only later did their de­
scendants escape to fill the seven seas.
A younger version, found in Egypt, was christened Ambulocetus natans
(the swimming walking-whale) after its large back legs, with seven-inch
toes. Ambulocetus was about the size of a sea lion, with a long tail quite
 Introduction 17

different from a whale's flukes. Another relative, from three million years
later, has its limbs reduced by a third. Yet another version, Basilosaurus
(whose name reflects its first designation as a "king lizard," regal indeed
at seventy feet) lived about forty-five million years ago. It had small but
perfectly formed rear limbs projecting a few inches from its body. With
these fossils, almost all the steps from land animal to leviathan have been
found.
The mammals of today hint at how the first whales leapt into the
waves. Many can, with more or less embarrassment, make their way in
water. Dogs paddle, humans do the breaststroke, seals swim better still.
Modern whales do the job so well that they cannot walk on land. The
feet of Ambulocetus put it between wind and water, in the otter league,
with back feet bigger than those in front. It swam better than any dog
and may have been as good as a sea otter, as it moved not just with kicks
of its rear legs but by flexing its spine. This was a large step toward the
whales, which do the same with the help of an evolutionary novelty, a
pair of giant flukes.
Although old bones are quite persuasive about what is needed to make
a whale, they are, like all ancient remnants, above all rare. Nobody will
ever reconstruct the biography of whales from the fragments of their an­
cestors. Fortunately, whales-as much as viruses-are living fossils. The
vestiges of limb bones show that, once upon a time, they had legs (and
now and again a modern whale is born with small hind legs of its own)
but the genes say more about whence they came.
The chronicle of the DNA is most obvious when it goes wrong. One
whale suffered from "a peculiar snow-white wrinkled forehead . . . The
rest of his body was so streaked, and spotted, and marbled with the same
shrouded hue that, in the end, he had gained his distinctive appellation
of the white Whale. "
Moby Dick was not quite a fiction, for a white sperm whale (Mocha
Dick, named after the island off the coast of Chile where he made his on­
slaughts) attacked whaleboats in the 1 830s. A snow-white example was
once caught by the Japanese. The beluga whale's name in Old Norse
means "corpse whale" because of its resemblance to a drowned body
blanched by weeks in the water. It shares (with white mice and albino
people) an inherited defect in the ability to make the dark pigment called
melanin. The human disease Chediak-Higashi syndrome arises from a
18 Introduction

mutation that causes silvery hair. In mice the same gene is called, from
its effects on coat color, the beige mutation. Patients (or mice) who in­
herit it get cancer and die young, because they lack a class ofwhite blood
cell whose job is to destroy tumors. Chediak-Higashi is connected to
AIDS, because the HN virus does some of its damage when it attacks
such cells. The altered gene has also been found in mink, cattle-and a
single killer whale.
Genes can track down the Moby Dicks of this world as well as they
can its viruses. One whale sampled in the North Atlantic in 1 964 found
its way three decades later to the meat counter of a department store in
Osaka after having been captured, supposedly as part of a scientific sur­
vey. Whole populations can be tracked in this way. The humpbacks of
Hawaii almost all carry the same pattern of DNA, with virtually no vari­
ation among them. The Hawaiians had no names for whales and the old
whalers never found them around the islands. Perhaps, like AIDS on its
first forays into Europe, they are recent immigrants, descendants of a few
founders and a limited pool of genes.
DNA links whales to the other mammals. They are not, alas, rdated
to bears (who sit firmly among the dogs) . Instead their molecular her­
itage shows them to be close to the hoofed mammals, the ungulates.
Within that group (which includes deer, horses, pigs, giraffes, hippos
and dephants) whales are nearer to those with an even number of toes­
pigs, deer or hippos-than to ungulates with one, three or five toes, such
as horses and rhinoceri.
The real key to their past lies, by chance, among some rdatives of the
AIDS virus. They have been hidden in the DNA of those great beasts
since long before the first whale took to the oceans.
For some viruses, the war between host and parasite gives way to truce.
They become integrated into the chromosomes of their carriers and are
copied each time the proprietor's cell divides. Because such hangers-on
do no harm, they are transmitted for millions of generations, dormant in
the same place in the DNA. Much of our own inherited material is made
of such decayed retroviruses, some shared with other primates, others
with more distant beasts. Their arrival may have been marked by an epi­
demic rather like AIDS. If it was, all hint of bad blood between vehicle
and passenger has long disappeared. Perhaps, in the distant future, the
sole evidence ofAIDS itself will be a few silent sections of DNA scattered
among the genes of our remote descendants.
 Introduction 19

Such biochemical hitchhikers can be used to search for kinship, not of

viruses, but of those who carry them. They reveal the history of animals
billions of times their size as, if two species have the same retrovirus in­
serted into the same position in their genes, they must descend from a
common ancestor.
The retroviruses ofwhales hint at an unexpected past. Whales, hippos,
deer and giraffes have three shared elements of this kind, each in the
same place in the DNA. The crucial trio of passengers is absent from all
other even-toed ungulates, pigs and camels included. They show that a
deer is more related to a whale than it is to a pig. Had Darwin speculated
about an aquatic and open-mouthed stag (or even giraffe) he would not
have been far from the truth.
Other viral wanderers narrow the search for the whales' immediate
kin. Whales do not, it must be said, look much like any other animal.
Fossils may tie them to the ungulates, but they seem quite distinct from
any extant member of the group. However, a further ten shared elements
show whales to be closely related to hippopotami. Deer, pigs and camels
are, in turn, more and more distant relatives of those unlikely sibs.
The molecular tree of hoofed mammals sprang from a common root
sixty-five million (rather than sixty) years ago. Like that of immunodefi­
ciency viruses it is bushy, which makes it hard to sort out where the
branches leading to horses, rhinos and the others split off. Nevertheless,
the marriage of hippopotamus with whale is clear. The molecular clock,
its rate set by the bones that fill the rocks, points at an ancestor common
to whales and hippos some fifty-five million years ago, just before Hi­
malayacetus, the earliest known fossil whale.
Whales and hippos may not much resemble each other nowadays, but
retain some hints of kinship. Hippos spend their time in the water and
on land; and some early whales did the same. Young hippos swim before
they can walk, and hippos and whales each nurse their young underwa­
ter as they squeak and squeal through river or sea. Both are hairless, nei­
ther can sweat and their males each keep their testicles inside the body
rather than in a convenient bag.
Although genes hint that the animal most like a whale is a hip­
popotamus, there is no manifest link between that ponderous river
dweller and those swift predators, the mesonychids. New fossils, though,
show the mesonychids to be descendants of an ancestor shared with
whales, rather than on the direct whale line. Although-as for AIDS
20 Introduction

viruses-much remains to be learned about the ancient history of

leviathans, the case for their evolution is impossible to deny.

The virus and the whale each tell a story of how descent with modifica­
tion leads, in time, to new forms of life. Each, in its disproportionate
way, affirms the truth of evolution. Although much is obscure, it is im­
possible to entertain any doubt, after a century of the most deliberate
study and dispassionate judgment, that the view which most naturalists
once entertained-namely that each species has been independently cre­
ated-is erroneous.
To deny the truth on grounds of faith alone debases both science and
religion. The point was made by Galileo himself Summoned to explain
his views and their conflict with Scripture, he argued that the church had
no choice but to agree with the discoveries of science. It would, he said,
be "a terrible detriment for the souls if people found themselves con­
vinced by proof of something that it was made a sin to believe." Cre­
ationists have not yet faced that fact.
No biologist can work without the theory of evolution. Like Galileo's
notion of a solar system with the sun at its center, Darwin's long argu­
ment makes sense of their subject. Ideas of origin were once, like Moby
Dick, allegories. They helped to comprehend not the structure but the
meaning of the universe. Some still hope to find symbolic significance in
Darwinism. They will not: but his work turned the study of life into a
science rather than a collection of unrelated anecdotes.
 C H A P T E R I

VARIAT I O N UNDER D O M ES T I CATI O N

Character o f Domestic Varieties - Relation between Man and his Do­

mestics - Origin of Domestic Varieties from one or more Species -
Principles of Selection, anciently followed - Domestic dogs, their Dif­
ferences and Origin - Methodical and Unconscious Selection - Breed
and Identity - Evolution on the Farm - Zoological Gardens: the Call
of the Tame - Loss of Variety under Domestication - The Wolf be­
neath the Skin - Difficulty of distinguishing between Varieties and
Species

Man has a strange relationship with his domestic animals. The Victorian
explorer William Burchell found himself unable to eat zebra when he was
near starvation in Mrica, because of its resemblance to his favorite mare.
The French government, alarmed by the waste of good protein, had
managed in the 1 860s to persuade its citizens to feed on horse, but in
London, the Society for the Propagation of Horse Flesh as an Article of
Food failed in the endeavor, in spite of a launch banquet of Salmon with
Racehorse Sauce, Filet of Pegasus, and, to follow, a Gateau Veterinaire.
Even so, at about that time, the Live Stock Journal and Fancier's Gazette
complained that "in some parts of England cats are not wholly despised
as an article of diet" and that a notorious gang of cat eaters in West
Bromwich meant that fanciers "cannot keep a favourite a week."
Animals, as they become domestic, enter an uncertain domain be­
tween the real and the artificial. They persuade man to accept the living
world as part of himself, promoted from food to member of the family.
22 D A RW I N ' S G H O S T

In the Middle Ages pigs were tried and hanged for murder, and only
forty years ago a female rhinoceros was elected, by a large majority, to the
Sao Paulo City Council. In an equivalent confusion today, a third of all
dog owners are happy to identifY their pet as closer to their heart than is
anyone else in the household.
As the wilderness creeps into the home, boundaries that were once dis­
tinct become blurred. Greeks, Egyptians and Icelanders each had sacred
dogs-Cerberus, Anubis and Garm-to guard the entrance to the next
world. In them, and in William Burchell's mare, the wild undergoes a
spiritual transformation beyond the reach of science.
It is just one step further than a change that affirms the central truth
of evolution: that variation within existing forms can, with human help,
bring forth new kinds of creatures quite different from their ancestors.
Evolution on the farm is a small-scale version of that in Nature. A wild
beast does not at once become tame, or a new breed arise in an instant.
Its passage from one way of life to another obeys the rules that apply to
the real world. Much can be learned about the course of the great stream
of evolution from domestication, its minor tributary. It shows that
species are not set in stone, but are always in flux.
Many people turn to The Origin in a search for a philosophy of life.
Most are disappointed to find that the first chapter is mainly about pi­
geons. It shows how breeders have produced birds as distinct as the
pouter, the runt, the turbit and the laugher from that drab source, the
rock pigeon, as proof of how animals can change. Darwin himself joined
the Philoperisteron, the smartest club of the London Fancy. He visited
breeders in their gin palaces and became a considerable expert on the
birds. His own became a pleasure: "the greatest treat . . . that can be of­
fered to any human being." Darwin did his job too well. One reader of
his manuscript suggested that it needed much more on pigeon breeding:
"Everyone is interested in pigeons . . . The book would be reviewed in
every journal in the kingdom, and would soon be on every library table."
Variation under domestication-in fields, in zoos and in living
rooms-is still powerful evidence for his theory. When we look to the in­
dividuals of the same variety or sub-variety of our older cultivated plants
and animals, one of the first points which strikes us is that they generally
differ much more from one another than do the individuals of any one
species or variety in a state of nature. Pigeons alone have dozens of
 Variation Under Domestication 23

breeds. The transformation of what was once a plain and unambiguous

bird-and its equivalents in the garden or around the fireside-tells an
evolutionary story.
Pigeons have been succeeded in Cockney affections by dogs (as is
manifest to anyone who walks in London's streets or parks) . The city's
dogs are beasts very different from a timber wolf, its municipal roses
quite distinct from their wild relatives. All are a product of selection.
Some wolves or roses were, for some reason, better able to cope with man
and, unlike their savage kin, survived and multiplied. Their progeny in
turn were winnowed and, in time, the wild became domestic. Those who
chose servitude changed to do so. Lord Somerville, quoted in The Ori­
gin, had it of breeders that: "It would seem as if they had chalked out
upon a wall a form perfect in itself, and then had given it existence." As
a result, the best place to see evolution is on the farm.
Most people see humans as at the top of a pyramid whose foundations
were laid four billion years ago. Mankind is, it seems, the master of the
fate of all other beings, most of all those used for food or work.
But who is at the center of the worldwide web of exploitation? The an­
swer is not humans, but pigeons, dogs or roses, for their lives depend on
man's readiness to be used. More than nine tenths of all the flesh in
Britain-except that of man himself.-is of domestic animals. Mice, deer
and all these islands' aboriginal inhabitants are left far behind. Although
the servants repay their debt with milk or bread, affection or golf courses,
their victory is clear. Fifty million dogs live in the United States, while
ten thousand wolves remain. Wolves are hunted down because they dare
to kill a few of man's billion sheep, themselves the relatives of the last
mouflons. To be born free was, for wolves or wild sheep, a dead end.
The grand enslavement began in the Middle East, with rye, thirteen
thousand years ago. Those who grew it were hunters, and long remained
so. The hunter-gatherers of the Yangtze cultivated rice twelve thousand
years ago, and the first domestic squash was grown in clearings in South
American jungle a thousand years later. Until the modern age there were
no domestic plants with an origin in Australia or Southern Mrica. It is
not that these countries, so rich in species, do not by strange chance pos­
sess the aboriginal stocks of any useful plants, but that the native plants
have not been improved by continued selection up to a standard of per­
fection comparable with that given to the plants in countries anciently
24 D A RW I N ' S G H O S T

civilized. This is itself evidence for evolution, as not until variation was
noticed by man did he create new animals and plants. Cows or wheat are
special only because they were exposed to human choice.
Wheat descends from three grasses that hybridized on the Anatolian
steppes. One, einkorn, still grows there. The DNA of wild and cultivated
einkorn shows that just the plants from the Karas;adag Mountains, be­
tween the Tigris and the Euphrates, gave genes to wheat. There the mod­
ern world was born, as those hills, with the lowlands to the south, were
also the birthplace of grapes, olives, chickpeas and bitter vetch. These
crops were, in time, enriched by plants from other places-rice from
China, maize, potatoes and more from South America-and by the do­
mestic animals of today (among whom pigs came first, followed by goats,
sheep and, much later, cattle) . Their descendants are evidence of how na­
ture has been molded to human ends.

On the Breeds ofthe Domestic Dog. Nowhere has domestication gone fur­
ther than in the household. Dogs bear witness to its power. The first dog
show in England was held in Newcastle in the year of publication of The
Origin. It featured familiar breeds such as pointers, setters, and spaniels.
The Americans took up the sport a few years later with a display of
Queen Victoria's own deerhounds, a dog called Nellie who walked on her
hind legs because the front pair was missing, and the prize of a pearl­
handled revolver for the overall winner.
Now, the United States has eleven thousand shows a year and seven
billion dollars is spent on veterinary fees alone. The forms generated in
the brief search for perfection are remarkable. The champion of the 1 998
Westminster Kennel Club Dog Show, America's premier event, was Fair­
wood Frolic, a Norwich terrier. She was selected from the best dogs in
groups that included a malamute, a toy poodle, a long-haired dachshund
and an Old English sheepdog. Previous winners have come from breeds
as extreme as the pug (whose eyes have sometimes furtively to be pressed
back into their sockets by their owners) . To survey the arena at Madison
Square Garden or at the National Exhibition Centre in Birmingham
(where Crufts, the English championship, is held) is to see how plastic
flesh can be. A dog show is evolution chalked out for all to behold.
All these breeds descend from some wild ancestor. There are thirty-five
named species of wild dog, which gives plenty of candidates to choose
from. Some are solitary like the fox, others social like the jackal; some as
 Variation Under Domestication 25

small as the crab-eating zorro of South America, a few as large as the Arc­
tic wolf The bones of an even bigger animal, the dire wolf, are preserved
in the La Brea Tar Pits in Los Angeles. It died out ten thousand years ago.
Just one dog (given the Latin name Canis flmiliaris by the great clas­
sifier Linnaeus) has been domesticated. Linnaeus defined that familiar
beast as distinct from all others on the basis of its upturned tail, found in
none of its kin. Other characteristic features were that domestic dogs lick
their wounds, are often infected by gonorrhea, and urinate when they
hear certain musical notes.
The variation within the only dog to place its future in the hands of
man transcends that in all its relatives put together. Linnaeus com­
mented on the domestic dog's diversity and mentions the Naked Dog,
the Fat Alco and the Techichi (a barkless New World dog with a wild and
melancholy air) as variants of his Canis flmiliaris. Chihuahuas stand six
inches high at the shoulder, Irish wolfhounds three feet. A St. Bernard
weighs fifty times as much as a Pomeranian, and the range of color, shape
and temper of pet dogs is almost beyond imagination. Certainly, if they
were not our familiar domestics, at least a score of dog breeds might be
chosen, which, if shown to a mammalologist, would be ranked by him
as well-defined species. If the history of dogs as a product of man's whim
were not so familiar, the numbers given distinct labels by science would
rocket. Only because we see them as mere breeds are they confined
within a single Latin name.
The sole alternative to admitting the variation of dogs as evidence for
evolution is to believe that each breed descends from a separate wild an­
cestor, now extinct with no token of its passing. The doctrine of the ori­
gin of domestic races from several aboriginal stocks was once carried to
absurd extremes. As Nature was thought to be static and its products un­
able to change, breeders claimed that there must have existed at least a
score of species of wild cattle, as many sheep, and several goats in Europe
alone, and several even within Great Britain.
Even now, the idea of separate ancestry for dog breeds is not dead. It
is, however, wrong. But what was the cur whose descendants burst into
such variety? So different are the domestic kinds that it seems impossible
that they could descend with modification from a single source. Now,
the genes make it plain. The dog's sole ancestor is one celebrated animal,
the wolf
Wolf bones are found near those of humans as long ago as the time of
26 D A RW I N ' S G H O S T

Boxgrove Man, who lived in southern England four hundred thousand

years before the present. Nobody knows the nature of the relationship.
Perhaps men and wolves fed in the same place, or wolves were pests who
raided the hunters' camps. The people of Boxgrove may sometimes have
picked up a cub to be tamed, fattened and eaten later. Whether neigh­
bor, camp follower or convenient snack, the wolves changed little as their
owners were transformed. A mere ten thousand years since, when hu­
mans were more or less what they are today, the skulls of wolves recov­
ered from a drowned camp of the nomads who crossed from Siberia to
Alaska were almost the same as those of their Boxgrove ancestors. They
had shorter faces and their teeth were more crowded, but in all appear­
ance they were wild animals.
Then, hunters began to use arrows rather than spears. Wolves--dogs,
as they soon became-became more useful as they could chase and pull
down wounded prey. Such a creature took at once a large step toward the
fireside. They deserved honor, and Darwin noted with disapproval how
the people of Tierra del Fuego would devour their old women rather
than their dogs in times of shortage. At Ein Mallah in Israel, in a grave
of the earliest farmers, is the skeleton of a puppy buried next to a child.
A wild animal had become a member of the family. Soon, its muzzle
shrank, its teeth became smaller, its eye grew large and round, and the
modern dog had arrived.
The singing dog of New Guinea is a relic of that primitive beast. The
dingo, too, is not much changed since those days. Dogs got to most of
the -world (except Mrica, which was dog-free until the Iron Age of A.D.
500) as soon as humans did but, worldwide, changed little for thousands
of years.
One set of genes, in dog or man, is special. Mitochondrial DNA is
found in certain cell organelles whose job is to generate energy. It passes
(like the mirror image of a human surname) through females alone; from
mothers to daughters and sons, but daughters alone transmit it to the
next generation. It contains within itself the history of bitches. Like all
genes or surnames it accumulates changes as the generations pass. How
distinct such DNA might be in any pair of lineages hints at how long it
has been since they last shared a mother.
A survey of dozens of breeds of dogs, and of wolves, coyotes and foxes,
shows that wolves and dogs are indeed the closest kin. Dog "surnames"
 Variation Under Domestication 27

fall into two distinct groups, evidence that the animals were tamed twice
in different places. As their ancestors were in contact with humans for
thousands of years, a mere two domestications suggests that to become a
servant is not as easy as it seems.
The story of dogs is written both in bones and in genes; and, unfor­
tunately, they disagree. Fossils prove that wolf and dog split some ten
thousand years ago and that the paths of wolf and coyote diverged a mil­
lion years earlier. Mitochondrial DNA, though, hints at an earlier do­
mestication. To put the mitochondria of wolf and dog into coyote
context puts their division at a hundred thousand years before the pres­
ent, which-given that the first modern humans appeared at about that
time-makes little sense. Fossils may be rare, but they are not ambigu­
ous; and the tale plainly told by dog bones is more convincing than that
hinted at by their genes.
Whenever it happened, what did it take for the dog to abandon the
wild? Biology and culture worked together. Nobody claims ownership of
a wolf, but a dog without a home is a pariah. Its existence depends on its
relationship with man. Dogs choose their masters well, with a close fit
between family income and dog ownership. Sometimes, though, they are
ejected. Italy has a million feral dogs, driven back some way toward na­
ture. They live, like their ancestors, in packs of a dozen or so. Their exis­
tence is miserable by comparison with that of their domestic cousins. For
food they roam from rubbish tip to rubbish tip. They have a parasitic
rather than an affectionate relationship with ourselves. Servitude has
even destroyed their native ability to raise young. As a result, only one
feral pup in twenty lives through its first year, and were it not for a steady
influx of other outcasts the packs would soon die out. For our favorite
pet, to become housebroken led to a dramatic increase in the quality of
life. It was, however, a journey down a one-way street.
Dogs have paid a price for easy living. To become domestic stifles the
world of the senses. Wolves are fierce, fearful and filled with stress; dogs
calm, docile and, most of the time, carefree. Pets are by their nature a
parody of a wild animal. What made the wolf an emblem of dread has
been much diluted. Its ears, once pricked, are floppy, and the sounds of
the world are dulled. Its sharp eyes are blurred by a fringe of hair and can
no longer stare an opponent into submission. The lupine tail, an expres­
sion of rage or delight, is in many breeds so curled as to bear no message
28 D A RW I N ' S G H O S T

at all. Most pets cannot even raise their hackles in anger as their hair is
too long. All this comes from an unconscious preference by man for an
animal that knows its place.
What was once done without thought has been echoed by science. In
1 950s Russia, silver foxes were farmed for fur. They were savage, suspi­
cious and liable to die from anxiety. On a certain collective, in an attempt
to improve matters, only those willing to accept human company were
chosen as parents. Within twenty years and a mere ten thousand foxes,
the farmers saw a great shift in their charges. The ranch was filled with
well-behaved animals that looked more like dogs than foxes, with a low­
ered tail and drooping ears. Many had piebald coats, quite unlike their
unrestrained kin, and the females reproduced-like dogs-twice rather
than once each year. To breed for tameness was enough to make the
change. The other characteristics followed.
Many of the qualities of today's dogs arise from the simple human
taste for young over adults. For most of the time, a pet's job is to be a sur­
rogate child. Any animal that looked like a pup-a short muzzle with
small teeth, round and attractive eyes, a friendly nature-was bred from
at the expense of his more brutish sibs. Familiarity alone prevents our
seeing how universally and largely the minds of our domestic animals
have been modified by domestication.
The end result of evolution through man's desire was to produce a sex­
ually mature wolf pup. A wolf-sized dog, a Labrador, say, has a brain a
fifth smaller than that of its wild relative; the size, indeed, of the brain of
a three-month-old wol£ It is scarcely possible to doubt that the love of
man has become instinctive in the dog. Any pet behaves like a j uvenile
version of its ancestor. It sits around in the hope of a meal and licks the
hand that feeds it, as wolf cubs lick their mother's face to persuade her to
disgorge meat. Most wild dogs cooperate to raise their young, either in
packs or (as in jackals) with each pair helped by a "maiden aunt." The
success of their tamed heir turns on its ability to persuade men to act as
aunts.
How far any dog is allowed to progress into maturity depends on what
its owners need: mere affection, or much more. The key is man's power
of accumulative selection: Nature gives successive variations; man adds
them up in certain directions useful to him.
The hounds that protect sheep in the mountains of southern Eu-
 Variation Under Domestication 29

rope-the Pyrenean mountain dog or the Anatolian shepherd-look

fierce, but are in fact enormous infants. They are brought up with the
flock and come to regard them as members of their family. An Anatolian
dog is quite unable to hunt. Even if starved it will not eat a carcass un­
less it has already been cut open-a job done by its mother in the years
before domestication. If an enemy attacks the flock, the guardian scares
it off with clumsy friendship. It alarms the nervous predator with hos­
pitable barks as it wags its tail and runs up in welcome. Such canine Peter
Pans even have low levels of the brain chemicals used in nerve transmis­
SlOn.
Herd dogs have not regressed quite so far. They are granted an adoles­
cence. A good collie enters into the spirit of the hunt, up to a point. It
fixes a sheep with its eye and stalks it, but does no more. Under no cir­
cumstance will it bite its prey. Corgis-a favorite of the Royal Family­
take the next step. Because cattle, their usual target, are so large, to move
them they must eye, stalk and nip at the animals' heels. Catahoula leop­
ard cowhog dogs go still further toward the adult wol£ They form packs
to harry cattle across the range until the herd runs to where it should.
Distinct breeds have been around for a long time. An image of a
pharaoh hound is on the tomb ofTutankhamen. The Pekingese appeared
in second-century China as a pet bred to look like the spirit-lion of Bud­
dha. So sacred were they that, when the British occupied Peking, the Em­
press ordered the execution of her favorites to prevent their capture by
the white devils.
A new form, with its own appearance and personality, can arise with
great speed. Although the Inuit used dogs to haul sledges across the Arc­
tic long before the arrival of the white man, the tandem harness, with its
many animals linked together, was introduced by Europeans. Today's
sled dogs are, over more than ten miles, the fastest mammals on land.
Their marathon, the Iditarod, extends over a thousand miles of ice and
snow from Anchorage to Nome. The winners do it in ten days. Teams at
the turn of the century were a mixed and disreputable bunch but within
a few years the victors began to converge on a common type. Soon, they
looked like the familiar beasts of today, and now all the champions share
the same thick coat, long lope and ability to work as a team. Even their
feet have changed, with a specialized pad that does not pick up snow.
Some of the diversity in size or in temperament of various breeds de-
30 D A RW I N ' S G H O S T

scends from the ancestral wolves. Some-like that in "sporting plants, "
by which term gardeners mean a single bud o r offset, which suddenly as­
sumes a new and sometimes very different character-has arisen since
the animals became part of our household arrangements. Such changes
happen because the genetic material of dogs, like that of viruses, is not
copied to the letter. Any mammal, with about a hundred thousand
genes, suffers from a constant influx of mutations. Most are harmless or
are hidden by unaltered versions of the same gene. Some, though, cause
changes that can be selected by an alert breeder.
Wherever the variation comes from, it is used by fanciers to make new
forms that, sooner or later, attract labels of their own. More than three
hundred distinct dog breeds are recognized. Quite what that means is a
matter of taste, because a breed does not exist until it has been named.
The American Kennel Club has as its mission "the maintenance of the
purity of thoroughbred dogs." Their self-imposed task is impossible.
Each stock, pure though it may seem, is always evolving as it responds to
genetic change and to its masters' desires.
Some of those are open, but most are not. There is a kind of selection
which may be called unconscious, and which results from everyone try­
ing to possess and breed from the best individual animals, with no wish
or expectation of permanently altering the breed.
The bulldog was once a savage animal, "unequalled for high courage
and stoutness of heart," whose speciality was to fly at the face of its
quarry and to use its massive jaws to bite and hold on to the animal's
nose. Its own set-back nostrils helped the dog to breathe as it did so. The
sport was outlawed in 1 835. By 1 900 the bulldog had-with no con­
scious attempt to change it-become "a ladies' dog as its kindliness of
disposition admirably fits it." The purists were far from happy, particu­
larly when furtive crosses with pugs further tamed the breed. Compared
to its fierce original, "the disgusting abortions exhibited at the shows are
deformities from foot to muzzle . . . and totally incapable of coping with
a veteran bull."
All breeds have altered, on purpose or by accident. A 1 570 book rec­
ognized just seventeen: Terar, Harier, Bloudhound, Gasehunde, Tumber,
Stealer, Setter, Wappe, Turnspit, and others. By 1 850, one writer recog­
nized forty breeds, and the Crufts show of 1 890 had two hundred and
twenty classes (although, admittedly, the last included "stuffed dogs, or
dogs made of wood, china, etc.") .
 Variation Under Domestication 31

Slow and insensible changes could never be recognized unless actual

measurements or careful drawings of the breeds in question had been
made long ago, which might serve for comparison. For one lineage that
is possible. The King Charles spaniel, much incorporated in portraits of
Charles II, was in the seventeenth century a black-and-tan animal. It has
been much modified since his time. Now, it has four varieties: the origi­
nal, another with white marks, yet another reddish-brown, and the last
reddish-brown with white spots. The decision as to what to accept as a
distinct class is quite arbitrary. The American Kennel Club is conserva­
tive and recognizes a mere hundred and fifty or so breeds (of which the
King Charles spaniel is-or was-just one). A certain nationalism takes
hold. Canada, for instance, insists on a separate name for the Nova Sco­
tia duck-tolling retriever, accepted nowhere else. The Kennel Club of
England is happy to subdivide its pets. The brown variety of the royal
spaniel has its own identity as the Blenheim King Charles spaniel, and a
form that looks like the original but has a long face is graced with the title
of the cavalier King Charles. In 1 997, a new American dog was born.
The cavalier King Charles was at last acknowledged by the American
Kennel Club and gained its own category in their shows.
Try the patience of owners as it might, such uncertainty is testament
to evolution. A breed, like the dialect of a language, can hardly be said to
have had a definite origin. It is hard to delimit any new kind of dog be­
cause when a system of its nature in flux is forced into fixed categories,
boundary disputes are inevitable.
Whether accepted by purists or not, breeds must, because they look
distinct, vary in their genes. Such differences, from duck-tolling retriever
to cavalier King Charles, are quite small. They involve only the attributes
selected by man. He cannot select any deviation of structure excepting
such as is externally visible; and indeed he rarely cares for what is inter­
nal. As a result, variants safe from human scrutiny have changed less than
have size, shape or behavior. Much of the hidden diversity of the ancient
camp followers remains, silent and unaltered, within the stocks of today.
Distinct as a terrier and a greyhound might appear, they are alike under
the skin. Breeds are pure only in the eyes of their owners. So much vari­
ability remains within each canine stock that some cities plan to use
DNA fingerprints on feces on the street to identify serial offenders.
As fanciers select their preferred types, the stream of ancient genes for
behavior, shape and size splits into a series of rivulets that narrow as se-
32 D A RW I N ' S G H O S T

lection goes on. Once a lineage is established, the patricians of the canine
world have their mates chosen from among their relatives. As a result,
they become inbred; they share a heritage because of descent from a com­
mon ancestor. A noble dog bonds to noble people and Irish wolfhounds
have, their owners say, an inborn ability to recognize those who bear the
blood of Irish kings.
Some canine genes are (like many of those for human disease) harm­
ful when present in double dose. The deformations that emerge from
such genetic mishaps are seized on by fanciers as fuel for their explo­
rations of the wilder shores of taste. In the kennel, after all, we see adap­
tation, not to the animal's own good, but to man's use or fancy. The
malign effects suffered by inbred animals show how evolution can ex­
ploit hidden diversity.
Time magazine claimed in 1 994 that a quarter of all pedigree dogs suf­
fer from a genetic disorder. The estimate may be too high, but without
question many have been damaged through the efforts of their owners.
If man goes on selecting, and thus augmenting, any peculiarity, he will
almost certainly unconsciously modify other parts of the structure. Some
canine difficulties arise because a change bred for in a particular charac­
ter alters the development of a different one. Almost all stocks have their
own problems. Chows are almost blind because their turned-in eyelids
give them a quizzical look, while bloodhounds, selected for a droop in
the lower eyelid and a mourhful face, suffer from inflamed eyes. In the
same way, Doberman pinschers suck their flanks and there are problems
of paranoia among basset hounds. The three hundred and fifty canine
diseases known to be inherited include some-cataracts, retinal cancers,
pathological falling asleep (a trait, like flank sucking, common in
Dobermans)-close to those of humans. The dog may repay part of its
debt to its owners as it helps them to understand their illness.
An Italian botanist, Odoardo Beccari, once speculated that our pets
had missed their chance because they evolved after an epoch when life
was plastic: "Had man been associated with the dog during the plasma­
tive epoch, I believe that to the expression of our face and to the sound
of our voice there would have been aroused in the dog, owing to the at­
tention with which he listens to us and observes us, analogous move­
ments in its vocal organs, which, instead of expressing themselves by
inarticulate sounds, would have enabled it to talk and to learn a lan-
 Variation Under Domestication 33

guage." Speechless though they remain, dogs are, because of the evolu­
tionary pressures imposed by their owners, silent witnesses to the mal­
leability of nature.

Selection. Many creatures have been subdivided by man in his search for
the excellent. They show evolution hard at work, but nowadays, too
often, in reverse: the problem is the loss of breeds as they fall from favor,
rather than their constant emergence. Some three thousand named types
of ass, cow, goat, horse, pig, sheep and water buffalo are known, a third
of them at risk of extinction. Their history is in their DNA. The genes
show that horses (like dogs) were tamed more than once, from Europe's
huge herds. Cattle, too, were domesticated twice. Those from Mrica and
Europe are on their own branch, those from India on the other. Mrica
has two varieties, a small form found in the west, and zebu, large animals
with humps, common throughout the continent. Zebu are susceptible to
a fatal blood parasite and can be helped with genes for resistance bred in
from their kin.
Farmers have much simplified their plants and animals. As they relive
the process of evolution by selecting from the finest, farmers lose the sec­
ond best-and their genes. Like dog fanciers, they have begun to realize
that diversity has been eroded at a fearful rate by unnatural selection. The
problem is a real one: because of the growth of the human population
the world will have to produce as much food during the next century as
it has since the beginning of agriculture. Farmers try, by careful crossing,
to retain variety. Even so, their domestics have become a much reduced
sample of the wild. The entire United States soybean crop-sixty million
tons of it-descends from a dozen strains collected in northeastern
China. In the 1 970s most of the country's maize harvest was lost as all
the billions of identical plants, superior as they were, succumbed to the
same fungus. Animals also suffer from man's obsession with the excel­
lent. Sunny Boy, a Dutch bull of superb quality, died in 1 997 after siring
his two-millionth calf. Half a dozen other bulls have each bestowed more
than a million successful sperm upon the world's cows, which means that
the genes of untold others have been lost.
Nature is now pillaged for what remains. Animals are preserved as
frozen embryos or as sperm freeze-dried to a powder and brought back
34 D A RW I N ' S G H O S T

to life. The world has seven hundred seed banks, each a reservoir of di­
versity. Botanists search for genes in the ancestors of today's crops, and
for new sources to exploit. The anti-cancer drug Taxol came, after all,
from an American yew; and when Leningrad was besieged people starved
rather than eat the seeds of rare wild grains. The Third World today com­
plains about collectors who hunt for wealth among the local plants.
The first attempt to rescue Nature's diversity began in 1 828. The goal
of the new Zoological Society of London, as set forth in its Prospectus,
was to introduce new kinds of animals into England "for domestication
or for stocking our farm-yards, woods, pleasure-grounds and wastes."
The society had not a single success. It triumphed in a different way, as
a display cabinet of curious animals for Victorian London.
Now, London Zoo, like all others, has changed its image. Its aim is no
longer to pillage the wild for man's use, but to protect what remains
against domestication. Zookeepers see themselves as the last hope for
many of the products of evolution, but-like the first farmers and the
members of the American Kennel Club-they are testimony to its inex­
orable force.
To some extent zoos have succeeded. Without them we would have no
Arabian oryx or European bison at all. They face two problems. First, a
zoo can sample just a tiny part of the diversity present in the wild. An an­
imal that once ranged over thousands of miles is forced to migrate to a
tiny island. As in European AIDS viruses or the humpback whales of
Hawaii, just a small proportion of the ancestral genes can make it to the
new home. Keepers must breed from what little they possess-which it­
self means evolution. They must also, perforce, choose as parents those
animals best able to cope with their new environment. As a result, their
charges begin to change, as cows and dogs have changed.
A zoological garden bears the unwelcome message that, because of
man's inadvertent selection, any animal taken from the wild becomes do­
mestic, a travesty of its natural self Evolution is as hard at work on caged
animals as on those born free. In time they will emerge as beings quite
different from what they were. Those who conserve animals in the hope
of returning their descendants to Nature may be disappointed by what
they let loose. Their failure shows how descent with modification is im­
possible to avoid.
In an attempt to save the wild, more and more has been tamed. Pes-
 Variation Under Domestication 35

simists claim that by the middle of the next century the world's zoos­
an area no larger than metropolitan Denver-will be the last stronghold
of all the hundred and sixty kinds of primate, sixty out of seventy-two
wild cats, and most of the world's wild dogs. Two thousand vertebrates
have their future behind bars. Already, many are reduced to a wild pop­
ulation of fewer than a thousand. The prospect for most of them is bleak.
The first zoo was founded by King Shulgia of Mesopotamia in 2000
B . C . He kept lions in his park near the divine city of Nippur. He was fol­

lowed by the Chinese Emperor Wen Wang with his "Garden of Intelli­
gence" and by Henry VI of England, who caged a lion in the Tower of
London (and gave free entry to those who brought a dog or cat for its
lunch) . Caged beasts testified to wealth; and Aristotle and Theodore
Roosevelt each had zoos of their own. The liberation of the inhabitants
of the Paris Zoo at the time of the siege in 1 870 had practical as well as
symbolic significance. Most of the emancipated creatures were eaten.
At first, the inhabitants of such cabinets of curiosity died soon after ar­
rival, because nobody knew how to look after them. In one, the gorillas
were given sausages and a pint of beer for breakfast, followed later by
cheese sandwiches, boiled potatoes and mutton, and more beer. Few sur­
vived (the visitors did not help; when Philadelphia Zoo opened in 1 874,
its sloth was poked to death by umbrellas within a week) and those that
did were reluctant to breed in public.
Akbar, the great sixteenth-century Mogul of India, is mentioned in
The Origin as the proprietor of twenty thousand pigeons. He crossed his
stocks, "which method was never practiced before," and "improved them
astonishingly." He also owned a thousand cheetahs, but these undomes­
tic beasts were contrary when it came to sex. The whole menagerie pro­
duced just a single litter. The next cheetah to be born in captivity was in
Philadelphia in 1 956. Nothing is more easy than to tame an animal, and
few things more difficult than to get it to breed freely under confinement
(in the cheetah, the female's need for a sexual chase by several males be­
fore she can ovulate is part of the problem). That may be why so few an­
imals have been domesticated and why dogs, horses and cows were
housebroken just once or twice.
For wild animals, from wolves to cheetahs, to accept human hospital­
ity reveals variation usually hidden and provides selection with new and
often unwelcome opportunities. Zoos must cope with a legacy of dam-
36 D A R W I N 'S G H OST

aged genes. Although of no apparent value in themselves, they hint at

how much diversity is concealed beneath the face of nature. If faulty
genes are rare and the animal common, then few will be unlucky in the
biological lottery, as most of the defective versions are masked by a nor­
mal one. Sometimes, though, two damaged pieces of DNA get together
and the carrier pays the price. This happens most often when relatives
mate, as each descends from a shared ancestor who might himself have
carried a single faulty replica of the gene. When brother and sister white­
footed mice are crossed in the comfort of a cage, their progeny released
into the wild die at four times the rate of those of unrelated parents.
For zoos, evolution is a problem harder to solve than is the diet of the
gorilla. A limited pool of parents and a new environment means a sub­
sequent genetic change. White tigers have been known since 1 834 and
were once much pursued by hunters. Now hundreds are in captivity.
Each descends from a single male, Mohan, captured by the Maharajah of
Rewa in the 1 950s. More than a hundred have been bred in Cincinnati
Zoo alone. Albino cats are popular, and the Philadelphia white lion
brought in a million dollars a year in tickets. It appeared as a result of do­
mestication, when close relatives were bred together (with the help of
careful family planning, as its father is also its grandfather and great­
grandfather) . The tamed white cats reveal a genetic secret of the wild:
that life is full of hidden imperfections that are noticed by selection as
soon as they appear.
Zookeepers suffer from other kinds of unwanted evolution. As wild
animals are hard to breed in captivity, their guardians tend to choose
those best able to cope with their new circumstances. That means
change-toward tameness and, as in the foxes kept for fur, toward other
things as well. All Przewalski's horses-reduced to a core of thirty-two
animals in the 1 940s-descend from a dozen wild ancestors caught in
Mongolia, and from a single female Mongolian pony. In their early years,
inadvertent preference for the animals most ready to accept captivity in­
creased the contribution of their tame progenitor until some herds traced
a fifth of their ancestry to that single pony. Although keepers have now
set out to purge the stock of their ancestor's insistent genes, the instant
response to selection for good behavior has a lesson for all other inhabi­
tants of zoos. Although the horse's saviors hoped to keep a wild animal
safe, what they have done is to evolve a new one.
 Variation Under Domestication 37

Today's conservationists are well aware of such dangers. Their philos­

ophy is quite opposed to that of farmers, who try as hard as they can to
select the best. Hundreds of "species survival plans" have been made in
the hope of combating the power of natural selection. They involve the
exchange of animals among zoos to keep up diversity and reduce local
adaptation and, at the same time, strenuous attempts to ensure that all
families have the same number of young in order to minimize the chance
of inherited differences in reproductive success. All this is an attempt to
bend the Darwinian rules and will merely delay rather than solve the
problem of evolution behind bars.
Caged and domesticated animals each show how selection can pro­
mote new forms. Cattle, sheep and goats never forget a face (even from
photographs) and recognize not just individuals but whole breeds by the
way they look. Given the choice a cow prefers to associate, and to mate,
with one of its own kind, be it a Friesian or a Hereford. What is more, a
male tends to fall for a female who reminds him of his mother. After all,
she brought him up and belongs-by definition-to his own kind. The
oedipal effect is strong. Young male lambs raised by goats, or young male
kids raised by sheep, are interested only in sex with animals like their fos­
ter mothers; male sheep with female goats, for example. A male's prefer­
ence for sex with a partner like his mother keeps a breed to itself.-a first
step in the promotion from variety to the elevated status of species; and
a further statement of how the domestic can relive the experience of the
wild.

Not all prisoners change their nature. No matter how much an unnatu­
rally tame horse-or a white tiger-alters as its ancestors retreat into the
past, each retains much of its history in its genes. Arabian oryx returned
to Oman after eight generations in captivity could still navigate across
the desert to find water a hundred miles away. No inmate of a zoo has
been so domesticated as to change its scientific name, to lose its natural
identity in favor of one imposed by man. Nobody enters a lion's cage un­
escorted and, in the United States, elephant keepers have the most dan­
gerous job of all-more so than the police, with one keeper in six
hundred killed each year.
Dogs, too, retain a past much older than their alliance with another
38 D A R W I N 'S G H OST

species. Each is a barely evolved wolf, a descendant of a dangerous ani­

mal. That is part of their attraction. Their owners glory in their pet's vic­
arious tie to Nature. Jack London, in his 1 903 novel The Call ofthe Wild,
tells the tale of a pampered domestic animal, a cross between a St.
Bernard and a Border collie, stolen and taken to the Klondike. The ani­
mal learns the truth: "Kill or be killed, eat or be eaten was the law; and
this mandate, down out of the depths of Time, he obeyed. " The book
sold two million copies. London, a social Darwinist of virulent stamp,
had a political agenda but his story has a scientific message. After a series
of bloody adventures, Buck, the canine hero, emerges in Nietzschean
style as the leader of a pack of purebred wolves. The "dominant primor­
dial beast"-the wolf within-had at last come to the fore. Domesticity
was a mere stage in his history, reversed when the wild called with more
insistence than did the fireside.
By owning a dog, any dog, men welcome into the home a beast that
preserves much of its primordial self Overgrown juveniles though they
are, evolution by human choice has not removed the instincts of their an­
cestors. In the United States, a hundred times more people are murdered
than are killed by dogs. Even so, their pets slaughter about twenty peo­
ple a year (although the wolf itself has killed no one within the past cen­
tury) . Like wolves, dogs attack the weak, be they young, old, or drunk.
Packs of feral animals have pulled children from bicycles and eaten them,
and a mere half-dozen beagles, dachshunds and terriers once devoured
an eighty-year-old woman. The homicidal packs relive their past. All do­
mestics retain some of their ancestry and every child licked-or sav­
aged-by its pet is fresh proof of the importance of descent.
Now, the dog has disappeared, victim of evolution. Two centuries after
it gained its scientific name, the International Commission on Zoologi­
cal Nomenclature has stripped it of its identity. Their action accepts the
transition between breed and species .
.As anyone who walks in a park knows, any dog is ready to discuss its
genes with any other, as long as it is in heat. Even so, there are barriers to
full sexual agreement among breeds. No owner of a pedigree bitch is
happy to see it mate with a mongrel, and breeders ensure that most of
their aristocrats never come across an animal not of their own kind. If an
owner's guard slips and male chihuahua meets female Great Dane, me­
chanical constraints to copulation come into play, or (if roles are re-
 Variation Under Domestication 39

versed) the birth of a giant pup to a mother so much smaller than the fa­
ther causes problems. Even when large forms like Great Dane and St.
Bernard are mated, the young are defective, as they inherit so many genes
for abnormal growth.
Linnaeus-a classifier of what he saw as a fixed biological universe­
named the domestic dog as a separate species, Canis familiaris. For him,
it had the same status as the elephant or the tiger: an animal so removed
from its relatives as to demand a label of its own. The International Com­
mission on Zoological Nomenclature defines its species more stringently.
Dogs as an entity were doomed because, once out of its owner's sight, any
dog is happy to have sex with any other-and even, given the chance,
with a wolE The DNA traffic has been in both directions. Dog mito­
chondria hint at an influx of Russian wolf genes and, because of crosses
between wild and tame, most European wolves have accepted genes from
their descendants.
Sex was, for the lexicographers of life, enough to destroy canine inde­
pendence. In 1 993 the Smithsonian Institution's Mammal Species of the
World, the Whos Who of mammals, admitted the domestic dog only as a
subspecies of the wolf Canis lupus. As Canis lupus familiaris it joins the
American wolf Canis lupus occidentalis and the European wolf, Canis
lupus lupus; each so much alike as to be ranked as mere varieties of the
same animal. The Smithsonian's decision is at the center of the theory of
evolution. It shows how arbitrary is the distinction between breed and
subspecies, and how a subspecies may, through sexual choice, gain a per­
sonality of its own. The accumulative action of selection, whether ap­
plied methodically and more quickly, or unconsciously and more slowly,
will always cause life to change. Dogs are still wolves beneath the skin,
and for every creature, domestic or wild, once evolution is at work a
change of identity must follow.
 C H A P T E R I I

VARIATION UNDER NATURE

Variability - Doubtful species - Essence versus individual difference - Hid­

den or cryptic species - Species resemble varieties in being closely, but
unequally, related - Defining life's frontiers - The undiscovered uni­
verse and the overcrowded Ark - Races of mice, in Laboratory and Na­
ture - Hybridism and Identity - The law meets evolution - Genes
and the bird-watcher's dilemma

Bird-watchers and ornithologists are not at all the same. To the latter,
everything about birds is of interest-how they migrate, where they
breed, or what they eat. Bird-watchers have a single concern, which is to
see as many kinds as they can. Once seen, as soon forgotten, or, at least,
ticked off and added to the Life List that is the basis of their self-esteem.
I went through the same phase. After the usual interest in stamps and
an eccentric deviation into cheese labels, I was given a pair of binoculars.
At the age of twelve I eavesdropped on a group of excited amateurs
(twitchers, as in much of the world they call themselves nowadays) as
they peered at some gulls bobbing, on a dim winter day, on the then
filthy waters of the River Mersey. All agreed: one of the birds was not an
ordinary herring gull, but the much scarcer glaucous gull, seldom seen so
far south. My problem was that I could not see any difference. A mem­
ber of the flock was a rarity, but which was it? Did it count? Could I
check the box in my bird book? It was my introduction to the ethics of
science. I admit it: I made the mark, but then I erased it.
Twitchers, like scientists, belong to a fellowship of faith. They play
 Variation Under Nature 41

cards against Nature. It is possible to win every time by faking one's

hand, but to do so removes the point of the game. That is the strength
of science, and its greatest weakness. Without collective trust it could not
work. Instead there would be the dismal apparatus of mutual suspicion
familiar to every accountant.
Birding is refreshingly free from fraud. It has had its scandals, such as
the notorious case of the Hastings Rarities (a set of bizarre sightings on
the South Coast in Edwardian times), the dubious claim of a Dalmatian
pelican in Colchester in the 1 960s, and more. Even so-and whatever
the rivalry amongst the twitching fanatics-most of those involved in
the sport play by the rules.
One problem baffles the most ethical bird-watcher. Stamps (or cheese
labels) are easy. The 1 853 One-Shilling Cape of Good Hope Triangular
(or the 1 954 Vache qui Rit) is, or is not, genuine. Fakery apart, there is
no reason to question the object itself. But what if it is ambiguous? Then
opinion, the enemy of science, creeps in. Is one kind of bird really unlike
another? How different does it have to be to count as distinct? What, in­
deed, is a "species" in the first place? Does it have a scientific definition,
or is it all in the eye of the beholder? The question is at the center of the
theory of evolution.
Most people can tell gulls from terns. Many can separate the herring
gull from the lesser black-backed (look at the back, which is pale in the
first and dark in the other) . My friend the glaucous gull is more subtle­
as large as a greater black-backed, but as pale as a herring gull, without
its black wingtips. A real expert can even sort out the Iceland gull (like a
small glaucous, with longer wings). Birders still argue about the existence
of the yellow-legged gull as a distinct entity, but its "bold, confident
look" is said to be diagnostic.
But how to deal with variation within each bird? Herring gulls from
Estonian bogs in the 1 950s had, some say, yellow legs, but these have
now disappeared in favor of the pink legs found everywhere else. Those
from the Atlantic islands are on occasion blown to Britain. Their backs
are almost as black as those of lesser black-backeds. A "generally stouter
bill" might help, but what use is a generalization when an individual
must be classified? If the despondent twitcher were to travel to Iceland
he would be frustrated by hybrids between the Iceland gull and its com­
monplace relatives. Which box does he check?
42 D A RW I N 'S G H OST

Species and nations have a lot in common. What, for example, is a

German? The tribe has a shared and guttural means of communication
that interrupts intercourse of most kinds, but the attribute is equivocal,
for Austrians speak the same language. Since 1 9 1 3, the country has de­
fined its own citizens by descent, by German blood (whatever that might
be) . It includes within the realm the remnants of the Saxon diaspora
(many of whom-Romanians included--cannot speak German at all) ,
but cuts out children born in Berlin of Turkish parents. One badly be­
haved teenager was deported to Turkey even though he was born in Ger­
many; while at the same time a hundred thousand Romanian-speakers of
approved blood were allowed in. Until the fall of the Berlin Wall, indeed,
a geographical barrier made many of the nation's citizens more alien to
one another than Westerners or Easterners were to the French or the
Poles. A century ago German identity itself meant little, as there were
only Prussians, Bavarians and Rhinelanders, political entities of their
own, each now reduced to variants within some greater Teutonic whole.
The problem of how to define Germans, or any nation, arises because
the question is ambiguous: does it turn on shared appearance and be­
havior, on geography, or on descent? Is a country an historical entity, or
should it be identified only on criteria that apply today? How much can
frontiers be allowed to leak before a nation loses its essence? When will
Germans be seen as Europeans, as Prussians have become German?
Such problems of identity turn on natural variation, the raw material
of evolutionary change. Like a politician, the twitcher has to deal not just
with differences among individuals but with the subtle distinctions that
separate each kind. The difficulty of how to define domestic breeds has
been magnified and transferred to the world as a whole. Twitchers are
asking a question older than Darwinism. How should they deal with
forms that possess in some considerable degree the nature of species but
are not classified as such?

Taxonomy, the science of ordering life, has to worry about that problem.
Needless to say, many animals and plants are easy to tell apart. If they
were not, bird-watching and natural history museums would each go out
of business. One tribe in New Guinea recognizes a hundred and thirty­
six kinds of birds, just one fewer than the number accepted by the ex-
 Variation Under Nature 43

perts. Experts and tribesmen have the same philosophy. Each needs an
archetype, a gold standard, to allow their specimens to be put in the cor­
rect cabinet.
Once, all taxonomists worked in the same way. An animal was killed
and its remains stuffed, pinned or bottled. Then, it was described in the
scientific literature. The cadaver was the "type" against which others
could be checked. In 1 868, in China, the French missionary Pere Ar­
mand David saw the skin of a black-legged white bear. It resembled an­
imals shown in ancient works of art and until then assumed to be polar
bears brought back from the north by hunters. The first specimen of the
mysterious beast was collected in 1 929 by Theodore and Kermit Roo­
sevelt, the sons of the president. They shot a giant panda asleep under a
tree. Its body gave the animal entry to the pantheon of mammals as Ail­
uropoda melanoleuca. It joined the world of science as had all its relatives,
as a corpse.
Now, fewer than a thousand pandas are left. In China, to kill one
means the death penalty. Taxonomists, too, are more careful with their
material than once they were. The essence of a species can now (or so
museum-keepers believe) be preserved not in its bones but in its genes.
The Bulo Burti boubou shrike of Somalia was recognized in 1 99 1 on the
basis of the DNA sequence in a feather shed by a captive bird. The type
specimen-the very substance-of this new form is a set of dark bands
on a photographic plate. The rest of the bird was released.
Not all pandas-or Bulo Burti boubou shrikes-are alike. They may
look the same but are, like whales, dogs, or viruses, full of diversity. Clas­
sifiers hence face a fatal temptation: to split their animals into too many
groups. As in the Kennel Club or the United Nations, quarrels break out
between those who like to subdivide the world and those who hope to
unify it.
A rich nineteenth-century collector, Isaac Leigh, was interested in the
freshwater mussels of North America. He named more than a thousand
kinds on the basis of tiny variations in shell shape and size. Now the
number has been reduced by two thirds. A hundred and two of his types
are classified as one. Isaac Leigh was too enthusiastic about his varieties.
His cherished diversity was no more than that between people with
brown or blue eyes or between the pink- and yellow-legged herring gulls
that once infested Estonian marshes. He had, nevertheless, put his finger
44 D A RW I N 'S G H OST

on a problem that still plagues museums. How should they fix the fron­
tiers between supposed entities when each is filled with variation?
Genetics, the science of differences, has not made their job any easier.
Before it began it had often been asserted-but the assertion was quite
incapable of proof.-that the amount ofvariation under nature is strictly
limited in quantity. Now, the claim can be tested, and it fails.
Most members of most species do not look much different one from
the next. Any fruit fly is much like another, and even their best friends
find it hard to tell mice apart. In spite of some exceptions-the colorful
snails or butterflies that come in dozens of forms and are still studied by
a few outmoded naturalists-to share a Latin name imposes, almost by
definition, a certain uniformity upon those who bear it. That comforts
both creationists and experts on taxonomy. They like to see existence as
a set of neat ideals, each filled with some pure Platonic essence. However,
a great deal is hidden within even the most uniform creature. Genetics
shows that no one-not even the glorified chemists which most biolo­
gists have become-can any longer suppose that all the individuals of the
same species are cast in the very same mold.
Systematists are far from pleased at finding variability in important
characters, and there are not many men who will laboriously examine in­
ternal and important organs, and compare them in many specimens of
the same species. When biologists look for diversity with microscopes or
DNA-sequencing machines, they usually find it. Individuality is every­
where. Some of the variation in shape or size comes from the conditions
in which plants or animals grow. Even so, to move eggs between nests, or
to plant the same seeds in different places, shows that many of the dif­
ferences in such attributes do reflect genetic variation.
Much more variety lies beneath the surface. Most animals, even the
simplest, have inherited cues of identity on their cells; precursors of the
blood groups and the other genes of the immune system found in more
complicated creatures. Chromosomes vary in shape, size and arrange­
ment; and the proteins themselves are filled with difference. Most mole­
cules move when placed in an electrical field. They slow down when
passed through a jelly that filters them by shape, size and electrical
charge, to a degree that is altered by slight changes in their structure. The
technique reveals a mass of variation in certain proteins, the products of
genes. Now, it is much applied to DNA, either to measure the lengths of
 Variation Under Nature 45

the pieces that emerge when the molecule is cut i n particular places, or
to help put its individual units into order. The method was the key to a
universe of variability and to what became known as the "find them and
grind them" era of evolutionary biology.
For a time it seemed that all Darwinian problems could be solved by
a glance at differences in protein structure, cheap, simple and tedious as
the work was. All conceivable plants and animals (and some scarely so)
were subject to the art. Variation goes from zero in a few plants and in
sea elephants to situations in which half the genes tested in a particular
species may be present as alternative forms. Species can, in the new world
of the molecules, no longer be seen as absolutes. They are not units, but
groups of individuals, each with a biological personality of its own. That
poses a question about their very nature. The DNA of two unrelated
mice or deer is separated by a million or more differences. How can tax­
onomy work if its subjects can no longer claim an unblemished identity?
In fact, genetics gives the science a status it once lacked. An animal's
place need no longer depend on the opinion of an expert. Instead, it can
be put into context, its inborn essence shared with neighbors, with other
populations of its own kind and with forms more or less distinct. The
genes reveal a hierarchy of difference, from population to variety to
species and on to life's higher divisions. Sometimes the gradation is clear,
and sometimes less so, but order nested within order is all around. The
pedigree of life can now be drawn in nucleic acids.
At first sight, some animals seem detached from such biological real­
ity. The differences between two populations of the same snail in adja­
cent Pyrenean valleys are greater than those between man and chimp.
Nevertheless, the overall picture is clear. Thousands of plants and ani­
mals reveal a close fit between the divisions revealed by genes and the
groups long used by classifiers. From seaweeds to mammals, separate
populations of the same species share about nine tenths of their protein
diversity, while the variation common to any pair of related species
ranges from a third to three quarters (although birds, it must be said, do
not fit into this neat picture as the differences among them are small).
When mammals are compared with birds, or flies with snails, descent
with modification is undisguised because it has gone on for so long.
Genes also show the actual numbers of distinct kinds of plants or an­
imals to be far higher than experts once thought; in part because so little
46 D A RW I N 'S G H OST

of the world has been explored, but, quite often, because DNA shows
that what was at one time classified as a single form of life is in fact sev­
eral.
Nobody knows how many different species there might be, even in a
taxonomy based on external appearance. It takes a long time to sort out
even simple groups. The first list of British butterflies was made in the
year of the Great Fire of London, and by 1 7 1 0 almost half of those
known today had been discovered. Not until the late nineteenth century,
the era of the great amateur naturalists, was the catalog complete.
The situation is worse for less familiar beings. A million and a half
kinds of plants and animals have their own names, but even among well­
studied groups, such as insects or worms, three times as many may re­
main to be described. To gas the inhabitants of a tropical tree can reveal
a thousand new kinds of beetle. About seventy thousand fungi have been
given a label, but the experts agree that a million are still undiscovered.
The sea is an unknown-and enormous-land; if its water were evenly
spread, the whole globe would be drowned a mile deep. The oceans are
filled with mystery and may hold from half a million sorts of animal to
twenty times as many. A handful of marine mud can hide a hundred dif­
ferent kinds of nematode worm. In 1 995 a whole new phylum (the
largest inclusive group of creatures), one of just thirty-five in the macro­
scopic world, was found on the sea bottom, its members attached to the
lips of lobsters. So much is unexplored-the deep sea, the rainforest,
even the soil in suburban gardens-that the world may contain a hun­
dred million different kinds of plants and animals.
Some are hidden not beneath the oceans, but nearer at hand. As the
Victorians surveyed the globe, five hundred new mammals such as the
mountain gorilla and the terrible mouse (the size of a fox terrier) were
brought to light each year. The number has dropped to an annual hun­
dred or so, but splendid discoveries are still made. In 1 992 strange horns
were seen in the homes of Laotian hunters. Two years later, their owners
were tracked down, scattered over a range of wooded mountains. They
are the saola, a bridge between oxen and antelopes. Eleven of the eighty
known kinds of whales and dolphins were discovered in the present cen­
tury. In the last few years remote places have revealed new deer, new pri­
mates and a whole host of rodents. The world's list of mammals is about
five thousand long, but three thousand more may be waiting in the
 Variation Under Nature 47

wings. Plants, too, hide a multitude of unknown forms. Almost half the
palms of Madagascar have been discovered in the past decade. In Mex­
ico, a new relative of maize was found, a plant of great potential whose
presence had been missed by generations of collectors.
Molecular variation under nature reveals divisions invisible to the
most skilled taxonomist. To journey into the rainforest or the garden will
uncover new kinds of being, but to sequence the DNA of those we know
may reveal many more.
The pipistrelle is Europe's most common bat. What seemed a single
animal is now known to be two, distinct in their genes and with squeaks
of different pitch. When it comes to the leopard frog of North America,
what was once classified as a species is now thought to be at least twenty­
seven, different in their DNA and quite unable to cross.
Genes ask old questions about species in a new way, often to the dis­
comfiture of those who classify the world. Most Mrican elephants roam
in herds through the savanna, but others live a more solitary existence in
forests. They meet at salt licks and live on leaves and fruit. The molecules
show the forest elephants to be distinct from their familiar kin. Whether
each deserves a scientific name of its own is so far undecided. In deter­
mining whether a form should be ranked as a species or a variety, the
opinion of naturalists having sound judgment and wide experience once
seemed the only guide to follow. Today, evidence of separation also lies
in the genes: in whether two animals can mate and have offspring. No­
body has yet dared to try to test the sexual desires of the forest and the
savanna elephant, and in most newfound species the chance to do so
never arises. As a result, biochemists have taken on the naturalists' role
and DNA has become the touchstone of identity.

The issue of who belongs where in the natural world can sometimes be
sidestepped with "varieties," "races" or "subspecies." What these are often
depends on who studies them. If one compares several floras-lists of the
plants in a particular place-drawn up by different botanists, it is possi­
ble to see what a surprising number of forms have been ranked by one as
good species, and by another as mere varieties. The same is true of ani­
mals. Seventeen and a half thousand species of butterfly have been de­
scribed-but they are divided into a hundred thousand subspecies. All
48 D A RW I N ' S G H O S T

this points at the quandary faced by those who make lists. Where do the
boundaries lie?
Noah, the world's second taxonomist (after Adam) had to decide
whom to allow onto his Ark. He took on board seven pairs of each of the
biblical clean animals (ruminants, those who chew the cud and have
cloven feet) and a pair of each of a selection of the unclean beasts (in­
cluding, it seems, all the insects-those that "walk on many feet") . His
Ark is estimated by biblical scholars to have been four hundred and fifty
feet long. Nowadays it would have to be a lot bigger. As well as three
hundred and thirty thousand kinds of beetle, the Ark might-depend­
ing on its captain's views on classification-have to accept the half-dozen
named varieties of tiger and twice as many leopards. Chevrotains-a
group of foot-high deer found in Asia and Africa-have a hundred and
twenty subspecies (or so some experts claim) , each of which would have
to argue itself on board.
There is a rift in the fraternity of museum-keepers. They need to clas­
sify their specimens in order to preserve them. But what is to be done
about those awkward cases in which a bird or a mosquito cannot be
checked off in its own box? The essence of any collection of stamps or of
teapots must be that each specimen belongs in a distinct category. If they
do not, the whole system breaks down. In biology, the urge for order has
to defer to the reality of change. For plants and animals-unlike cheese
labels-even to discuss whether a particular form is a species or a variety
is, quite often, vainly to beat the air.
Familiarity breeds varieties. It is within the best-known countries that
we find the greatest number of forms of doubtful value, because only in
such places is enough known to blur the boundaries between what might
appear at first to be distinct. Seven forms of European wild cat have been
named-and their ranges coincide exactly with national boundaries. In
the same way, many subspecies and races of voles, butterflies and more
have been described from the islands around the British coast. Most at­
tain that status only because Britain's plants and animals have been so
much studied.
If any animal or plant in a state of nature be highly useful to man, or
from any cause closely attract his attention, varieties of it will almost uni­
versally be found recorded. The mouse has always been in the public eye,
although its image has changed from pest to scientific specimen. It re-
 Variation Under Nature 49

veals, as does no other animal, how much varieties and species have the
same general character and how hard it can be to tell one from the other.
Mouse bones are found in association with man six thousand years
ago, at Catal Hiiyiik in Turkey. There they were useful, as the skeletons
suggest that they were skinned (perhaps to make clothes). Apollo
Smintheus, God of Mice, was worshiped in Homeric times, and, some
say, in Crete until the Middle Ages. The ancients knew of their diversity.
Pliny speaks of white mice used in fortune-telling, and, in China, gov­
ernment records list thirty such animals caught between A. D . 307 and
1 64 1 . Mendel himself bred gray and white specimens in his monastery
room, but had to keep his results secret as pets were forbidden. But not
until the present century did the mouse come into its own.
Just before the First World War, Miss Abbie Lathrop, a Massachusetts
schoolteacher and failed poultry farmer, set up a business for the sale of
pet mice. Her first attempt with "waltzers," mice with an inner-ear de­
fect that caused them to dance, did not much amuse the public. Her
business took off when she began to sell animals to laboratories.
From her stocks came many of the mice now used by their millions in
research. As brothers were mated with sisters, forced incest caused the an­
imals within each inbred line to become, in effect, identical twins, all
with the same genes. Those refined beasts were a small sample of the di­
versity of their wild ancestors. Variation no longer circulated through the
population as a whole, but was parceled out among lines, each a reposi­
tory of a part of the total. For mice in Nature, individuals are different
but populations are much the same. In the laboratory the opposite is
true. How much the lines differ is, as a result, a measure of variation in
the wild. Each stock has become, in effect, a new and artificial race.
Mouse lines are distinct in many ways. Some are fat, some thin, some
active, some passive. Some prefer sweet foods, others salty. Some, given
the choice between water and a beverage the strength of gin go, to a
mouse, for the latter, while others are strict teetotalers. A hundred thou­
sand mice are used each year to test what substances might cause cancer.
A certain chemical increases the incidence of cancer in some strains, but
decreases it in others. Size, weight, color, heartbeat, sexual activity-all
vary among lines and all are testimony to how easily a wild beast can be
subdivided.
To read the catalog of strains kept by the Jackson Laboratory in Maine
50 D A RW I N ' S G H O S T

(the mouse's Library of Congress) is to retrace their history. The C57BI

stock, the most used of all, originated with Miss Lathrop in 1 92 1 . It is
now divided into substrains, kept separate in the lab, which since those
days have diverged in how much they rattle their tails, the rate at which
they age and in their liability to cancer.
Wild races of mice, too, can be defined by genetic differences that
seem too great to be accommodated in a single scientific name. Most
come from isolated places: not laboratory cages, but islands and remote
valleys. There they can evolve undisturbed. More than forty races have
been named in Europe alone. The mice of the Welsh island of Skokholm
are a quarter again as large as those on the mainland, while those on the
Isle of May, in the Firth of Forth off Edinburgh, are docile. Some popu­
lations even have different numbers of chromosomes. Most mice have
twenty pairs, but in many places certain chromosomes become fused to­
gether. More than a hundred fusions are known, to give local races in the
Alps, in Greece and on the Orkneys. The Val Tellavina in the Italian Alps
contains, within ten miles, five distinct chromosomal types. Because
such animals find it difficult to cross with others, each race can be de­
fined not just by how it looks, but by how easy it finds it to have sex with
the neighbors.
The biography of mice is much the same as that of gulls, or mussels,
or any other animal studied in enough detail. At first, there seemed to be
just a single kind. Pliny introduced the term musculus or "little mouse"
to separate mice from rats. It was borrowed by Linnaeus as the scientific
name for his familiar Swedish animal, Mus musculus. By 1 940, there were
almost one hundred and fifty species of mice, each with its own Latin
name, but then, in a move toward simplicity, all were lumped into a
worldwide mouse under the Linnaean label (although a dozen or so sub­
species were allowed, to satisfy the splitters) .
That was too bold. At least seven genetically separate units are hidden
within the supposed entity and each now has its own Linnaean name.
Some are widespread like Mus domesticus, common in the Americas,
Western Europe, and Australia, while some are confined (one to Spain
and Portugal, which have a mouse of their own) . Some have odd
habits-the steppe mouse, for instance, makes hillocks in which it stores
grain and spends the winter. Although a few of the various kinds can be
crossed in the lab, most never mate in the wild. Their DNA is quite dis-
 Variation Under Nature 51

tinct. Genetic tests on inbred strains show some to be of hybrid origin.

Indeed, part of the heritage of C57Bl comes from quite another species,
as a hint at a checkered past under Miss Lathrop's care.
The mouse's hierarchy of change is not unbroken. Where the true Mus
musculus (an animal of Scandinavia and northeastern Europe) meets its
western and southern relative Mus domesticus, as it does in a great curve
from Denmark through Germany and across the Balkans to the Black
Sea, they hybridize. Different as they are, their males and females can
mate; but the two do not merge. Just a few miles away from the zone,
their DNA patterns are distinct. Within it, the offspring of mixed pairs
are unhealthy and full of parasites, and their sons are sterile. This is
enough to save European mice from fusion.
Even so, the frontiers are blurred and, over a territory thousands of
miles long, but no more than ten across, have sprung a leak. That reflects
a history of divergence as varieties make the painful transition toward an
identity of their own. The center of genetic diversity of the world's mice
is in Pakistan and India. Mus musculus and Mus domesticus began their
journey there, in a common homeland and with a shared pool of genes.
Their ancestors traveled in man's wake, in separate waves north and
south around the icy Alps as farmers moved west. As they went, they
evolved, until, at last, when the circle was closed and the mice met, each
had changed enough to render them incompatible. As a result, each kind
has been promoted to the status of a species rather than remaining as
mere varieties of the same one.
Some places-the Pyrenees, or the Alps-are full of such hybrid ani­
mals and plants, from newts and toads to hedgehogs and oaks. At the
time of the ice ages their inhabitants were driven into fastnesses in south­
western and southeastern Europe. There they were transformed, and de­
scent with modification created forms that, when they met, were less able
to exchange genes than before. Each, like the mice, is an evolutionary
adolescent, poised on the edge of independence, but not yet ready to cut
itself off from its relatives.
The inescapable conclusion from all this is that species are strongly
marked and permanent varieties; their permanence assured by a block to
the exchange of genes. Hybrid zones show how hard it is to draw a dis­
tinction between the two. Mice, newts, oaks and more are alive and are
always in flux. Because of evolution, all must change as the years or the
52 D A RW I N ' S G H O S T

miles roll by. As descent always involves modification, resemblance de­

creases as a shared ancestor recedes into the past. Just as in families or na­
tions, the decision as to whom to let in is often a matter of taste. What
seem like solid frontiers may soon melt away.
The courts have long been involved in demarcation disputes. The con­
servation movement means that they have no choice but to be dragged
into biology as well. The judicial difficulties that arise when fitting vari­
ation into pigeonholes are testimony to evolution.
The United States Endangered Species Act is all-embracing in its de­
fense of those at risk of extinction: it can protect "any distinct popula­
tion segment of any species of vertebrate fish or wildlife which
interbreeds when mature" or is "an important component in the evolu­
tionary legacy of the species." What, if anything, can that mean and who
is to decide on which population needs protection?
The dusky seaside sparrow-an animal defined by its unusually dark
feathers-once lived in a patch of scrub around Cape Canaveral. In the
1960s it went into decline. The U.S . Fish and Wildlife Service spent five
million dollars to protect its last scrap of habitat, but still the numbers
went down. Five of the last six birds (all males) were captured. The final
hope for their precious genes was to cross them with another kind of
sparrow from nearby. The courts quashed the plans on the grounds that
the progeny would be hybrids, with no rights under the law. The bird
was declared extinct in 1987.
Conservationists were outraged. How could this piece of American
heritage be allowed to die? Although the government refused to become
involved, such is the majesty of a Latin name that the fight goes on. The
attempt to save some version of the bird continues, at Disney World.
The precious sparrow was defined, as are many birds, by a slight dif­
ference in the way it looks. It takes an expert to separate the dusky ver­
sion from its vulgar and unprotected neighbors. The tale has a biological
twist. Dusky seaside DNA is, as far as can be seen, identical to that of
other, commonplace, sparrows nearby. The famous bird is not a separate
entity at all. The genes do reveal a deep split between the seaside spar­
rows of the Atlantic coast of Florida (the dusky form included) and
those-to the naked eye almost the same-on the Gulf of Mexico.
Where does the boundary lie? Which is the species and which the vari­
ety?
 Variation Under Nature 53

In 1 990, the Fish and Wildlife Service gave up the anti-Darwinian

struggle. It no longer placed hybrids outside the law. Texas cougars were
released into Florida in the hope that they would breed with and sustain
the last Florida panthers (already inbred, with most of the males plagued
by undescended testicles) . The federal government is now reduced to the
preservation of "evolutionarily significant units," whatever those might
be. The dignity of the law has met the volatility of nature and has been
forced to retreat.

Whatever species may be (and they are not what birders or governments
hope), they are not fixed. Instead, their boundaries change before our
eyes. What is a mere variety to some is granted its own identity by oth­
ers. Quite often, animals that are similar on the surface differ in their
genes. All this is grist for evolution, for the transition between variation
within a single kind and the origin of a new one.
The biggest difficulty about species is to decide what they are. The
problem is time-to describe in the two dimensions of today something
that evolved in three. For classifiers, what matters most is the future. All
plants and animals are, with hindsight, the same because they all descend
from an ancestor three billion years old. At the present day they may hy­
bridize or stay apart, and biologists can spend useful lives in deciding
where lines should be drawn. To taxonomy, though, their essence lies in
years to come. Museums assume (and it seems fair) that cats and dogs are
separate because there will never be an animal that traces a shared descent
from dogs and cats. That assumption is impossible to test.
As a result, when it comes to the nature of species, a certain pragma­
tism is called for. The word comes from the Latin specere, "to look at";
but simple appearance is not enough. No one definition has yet satisfied
all naturalists; yet every naturalist knows vaguely what he means when he
speaks of a species. The amount of difference considered necessary to
give to two forms the rank of species is quite indefinite. One text on evo­
lution reviews seven statements of what the mythic word might mean
and-as does every attempt to impose order on the chaos of life-fails.
Of course, to an evolutionist, it should.
Most biologists have a guilty secret: they started as bird-watchers.
Lord Rutherford claimed that all science is either physics or stamp col-
54 D A RW I N ' S G H O S T

lecting. In some ways he was right, but biologists have the excuse that
they were the first to see that life is not just an album of specimens. In­
stead, it is fluid, and in its inconstancy reflects the fact of change.
Bird-watching is a more refined pastime than it was in the days when
Liverpool was a port and a twelve-year-old could tick off gulls and at the
same time check funnels for shipping-line symbols. Now the twitchers
have their own association. The UK400 Club is so named because to see
four hundred different kinds of birds in Britain is the mark of a lifetime's
dedication. Twitching is a competitive business. Three of the top five en­
thusiasts for the sport are called Steve {and three of the top hundred are
women} . All have more than five hundred feathers in their caps. For
them, the glaucous gull is dull, given the choice of great black-headed,
Mediterranean, laughing, Franklin's, little, Sabine's, Bonaparte's, black­
headed, slender-billed, ring-billed, common, herring, Thayers', yellow­
legged, Iceland, lesser black-backed, greater black-backed, Ross's and
ivory gulls to be checked off on the British list.
Even for the heroes of the birding world, evolution raises its vexatious
head. Take the yellow-legged gull, admitted as a "tick" after much vacil­
lation about its status {confirmed when it spread to northern Europe in
the 1 970s and lived alongside its cousins but did not breed with them} .
The UK400 guide has it that "this species is distinct from Herring Gull.
In fact, it is more related to the Lesser Black-backed Gull, with which it
sometimes hybridises. The races atlantis, michahellis, cachinnans,
barabensis and mongolicus are included within this complex, whilst Ar­
menian Gull armenicus is considered by some to be a further species.
This isolated form breeds on the Armenian Lakes, Turkey and Iran and
winters in northern Israel. The Arctic races heuglini and taimyrensis are
best treated as Siberian races of Lesser Black-backed Gull, whilst the race
vegae is best lumped with Herring Gull or treated as a separate species."
That statement, in its petulant tone, contains within itself the theory
of evolution: that differences blend into one another in an insensible se­
ries, and a series impresses the mind with the idea of an actual passage.
For twitchers to treat life as a glorified stamp album is entirely to miss its
point.
 C H A P T E R I I I

STRU G G LE F O R EXISTENCE

Geometrical powers o f increase in Man and other creatures - Rapid increase

of naturalized animals and plants, from cows to water weeds - Compe­
tition universal: cycles, stalemate or disaster may each result from a Strug­
gle for Existence - The term used in a wide sense - Nature of the checks
to increase - Effects of climate, and the risks of the marketplace -
Shortage of food - Complex relations of all animals and plants through­
out Nature - The battle for sex - The fisherman and the struggle -
The relation between organisms and the failure of conservation

The U.S. National Park Service is expert at letting in daylight on magic.

Ellis Island was, for forty years after it closed, a magical place. Because it
could not be visited it could be invented. As the gateway to seventeen
million immigrants, it was a metaphor for America itsel£
Now, the island is just another New York tourist attraction, run with
federal efficiency for the descendants of those who passed through it and
no more mystical than Disney World. Instead it is instructive. A helpful
bar chart in the reception hall shows the size of the United States popu­
lation from its foundation, projected into the future. The picture is of
continuous growth to the middle of the next century, when the display
stops with the promise of four hundred million Americans.
The nation now holds two hundred and seventy million people, most
of them descendants of European immigrants. Its history has a message
for evolution: that the existence of any creature is a constant struggle
against relentless forces.
56 D A RW I N ' S G H O S T

America is full of failures. The Irish arrived and died out, as did the
Welsh. The Vikings were around even earlier but abandoned the coun­
try in 1 0 1 3. Some stopped on the way, in Greenland. Their numbers
reached six thousand, but by 1 36 1 , when a priest came to see what had
happened to his flock, all had gone. They had ignored the sea in order to
farm. The record of the ice shows that a series of bad summers had
starved them. The colonists even killed and ate their dogs.
Later attempts by the English to find a toehold in the Americas also
collapsed or fell into terrible difficulty. The "Lost Colony" of Roanoke
was founded on an island off the coast of the present North Carolina by
a hundred and fourteen people, who landed under the direction of Sir
Walter Raleigh in 1 587. Although the first New World child of English
parents was born there, three years later neither she nor anyone else was
left. Another venture in Virginia received seven thousand immigrants in
the two decades after 1 606. In that time, six thousand died, their strug­
gle over. Tree rings show that the worst drought for five hundred years
had struck just before their arrival, and another almost as fierce followed
a few years later. Crop failure and famine killed the settlers.
Most migrants share a history of repeated disaster by many and dra­
matic success by a few. Not many attempts to reintroduce threatened an­
imals into the wild have worked. Millions of dollars have been spent to
airlift the eggs of Kemp's ridley sea turtle to Texas. Not a solitary "head­
started" animal made it back to the beach. Most of the claimed successes
are mitigated failures. The California condor was declared extinct in
1 987. Since then, some have been released from their Los Angeles "con­
dorminium" but the few who survive must be trained to come to food.
They, too, have come to grief in the struggle for life.
All colonists have a natural desire to make their new land more like
home. In the nineteenth century, all over the world, acclimation societies
were founded to do the job. The rabbits let loose in South Australia in
1 859 succeeded, but they followed an earlier antipodean attempt that
did not. In New Zealand, a hundred and thirty kinds of birds were set
free. Just twenty lasted to the 1 980s. What mattered was how many were
released. Eight out of ten species with more than a hundred set loose
prospered, compared to a quarter of those with fewer birds. Four hun­
dred yellowhammers were released in New Zealand, where the bird
flourishes, thirty into Australia, where it sank from sight. Any small
group of animals gambles with limited capital in the great casino of Na-
 Struggle for Existence 57

ture. As a visitor to Las Vegas soon finds out, that means inevitable ruin.
Sooner or later, the money has gone and it is impossible to continue.
Even the Australian rabbit's luck was short-lived. An accidental escape of
a virus from a government laboratory may soon put paid to them over
the entire continent.
In the end, though, the Europeans who went to America made good.
Two decades after its foundation, Virginia was ten thousand strong.
Throughout the eighteenth century its population grew, mainly through
their own sexual efforts rather than by immigration. Patrick Henry
("Give me liberty or give me death" ), first governor of an independent
Virginia, himself had eighteen brothers and sisters. Now the state has
seven million people. A few years after the Revolution, an elderly Rhode
Island woman claimed five hundred descendants, half of them alive at
the time of her death. The population of England and Wales increased
by a quarter in the first half of the eighteenth century, while that of
North America multiplied six times. Benjamin Franklin, in his 1 75 1
essay Observations concerning the Increase ofMankind, peopling of Coun­
tries, etc., concluded that "our people must at least be doubled every
twenty years . . . in another century . . . the greatest number of English­
men will be on this side of the water."
Thomas Malthus, an English cleric, was alarmed by such figures.
American growth could not, he said, be sustained. It was "a rapidity of
increase probably without parallel in history." The figures suggested to
him that populations grew by doubling-from two to four to eight to
more than a thousand in just ten generations-whereas resources in­
creased from two to four to six and so on: "I think I may fairly make two
postulata. First, that food is necessary to the existence of man. Second,
that the passion between the sexes is necessary and will remain nearly in
its present state . . . I say, that the power of population is infinitely
greater than the power in the earth to produce subsistence for man." His
interest was in morals and not in biology. God "ordained that population
should increase much faster than food" and had provided a sense of re­
straint to prevent it from so doing. Marx and Engels also saw the paral­
lel between the economies of Nature and of man (although neither was
fond of Malthus; to Marx he was "a shameless sycophant of the ruling
classes" and his doctrine, to Engels, a "vile, infamous theory, a revolting
blasphemy against nature and mankind").
Tertullian had, long before, seen what such expansion implied: "We
58 D A RW I N ' S G H O S T

have grown burdensome to the world . . . nature no longer provides us

sustenance. In truth, pestilence and famine and wars and earthquakes
must be looked upon as a remedy for nations, a means of pruning the
overgrowth of the human race."
If the burden is reduced by the survival of one set of genes at the ex­
pense of others when faced by plague, war or famine, then a nation (or a
species) has no choice but to evolve. In 1 838, at the age of twenty-nine,
Darwin read (for amusement, as he claimed) Malthus on Population.
Here, he saw, was the basis for a theory of change: that, owing to the
struggle for life, any variation, however slight, if it be profitable to any
individual in its complex relations to Nature, will tend to its preservation
and will be inherited by its offspring. Any species-any country-that
fails to adapt will join the pantheon of the extinct.
A struggle for existence inevitably follows from the high rate at which
all organic beings tend to increase. We have better evidence on this sub­
ject than mere theoretical calculations: namely, the numerous recorded
cases of the astonishingly rapid increase of various animals in a state of
Nature, when circumstances have been favorable to them. Still more
striking is the evidence from our domestic animals of many kinds that
have run wild in several parts of the world.
In the sixteenth century, the Spanish colonists of Hispaniola consid­
ered quitting their colony because of an outbreak of a fearsome stinging
ant. Fortunately, prayers to St. Saturnin ensured that the plague sub­
sided. The Englishmen on Barbados a couple of centuries later were vis­
ited by the same scourge, and their offer of twenty thousand pounds for
a solution worked just as well. On both islands ant numbers decreased,
and the animal is now no more than a harmless member of the local
fauna. Why it burst into such abundance and then declined, nobody
knows, but something in its balance of life had changed.
Cattle were introduced into the Americas in 1 493 by Columbus. The
overgrowth had at once a chance to hint at what it can do. Two centuries
after those first cows, huge herds ran wild in Mexico as a wave of beef
moved into ungrazed lands to the north. The Jesuits abandoned a ranch
with its five thousand cattle in Argentina in 1 638. The animals grew into
a swarm. A traveler saw plantations and orchards whose walls were built
of skulls stacked "seven, eight, or nine deep, placed evenly like stones,
with the horns projecting. " The same happened with wild horses. At the
time of the Gold Rush, thousands were driven to their deaths over the
 Struggle for Existence 59

cliffs of Santa Barbara to make room for cattle. Plants can do even bet­
ter. A pretty aquatic fern, Salvinia, has been introduced into much of the
tropics from Brazil. Everywhere it has become a pest. It blocks rivers and
interferes with water supplies. Salvinia reached Australia in 1 952. It can
double every three days and, by 1 977, Lake Moondarra in northern
Queensland was covered by fifty thousand tons of weed.
Any creature will, given time and resources, increase in this way. Most
do not. Gilbert White published his Natural History ofSelborne in 1 789
as an account of the animals and plants of his Hampshire village. In it he
describes a struggle for existence among its swifts: "I am now confirmed
in the opinion that we have every year the same number of pairs invari­
ably; at least, the result of my enquiry has been exactly the same for a
long time past. The number that I constantly find are eight pairs, about
half of which nest in the church, and the rest in some of the lowest and
meanest thatched cottages. Now, as these eight pairs-allowance being
made for accidents-breed yearly eight pairs more, what becomes annu­
ally of this increase?"
Today, Selborne has doubled in size. The straw-roofed cottages are
gone, or have been reborn as bijou homes whose thatch is held down by
mesh that birds cannot penetrate. The church tower was altered in the
1 950s and swifts can no longer get in. Even so, their numbers have
changed little. In 1 983 the village held twelve pairs, which, given its
growth, is almost the same density as that recorded by Gilbert White.
The birds he saw could, with unrestricted increase, have brought forth
ten thousand billion billion billion descendants in the two hundred years
since he counted them. Something (not, it seems, a shortage of places to
nest) holds them in check. Swifts died in multitudes. If any of their
deaths were influenced by genes, natural selection was at work.
Eggs or very young animals seem generally to suffer most, but this is
not invariably the case. Seedlings, also, are destroyed in vast numbers by
various enemies. In a tract of tropical grassland, a hundred and seventy
thousand palm seeds were scattered. A third became seedlings, a tenth
saplings-but just fifteen reached maturity. Even worse, most young
plants and animals must attempt to force themselves into a place already
more or less full. Thousands of seeds can germinate on a square yard of
bare English ground. A few years later, when a community has become
entrenched, but a single one of those thousands will succeed.
The struggle for existence is a term best used in a large and metaphor-
60 D A RW I N ' S G H O S T

ical sense. It may be of one individual with another of the same species,
or with the individuals of distinct species, or with the physical conditions
of life. It is the doctrine of Malthus applied with manifold force to the
whole animal and vegetable kingdoms. It includes not just survival or
speedy growth, but sexual success; and what is fought over can be food
or sunshine, a place to live or a mate to fertilize.
The battle is a complex affair, with many players. It may involve dis­
tant relatives or close kin. War breaks out between unexpected enemies.
Humpback whales carry a thousand pounds of barnacles. The turbulent
water around an encrusted whale feeds the passengers but slows down
their vehicle. The whale retaliates with skin, grown at a rate three hun­
dred times that of our own as a kind of antifouling to slough off the tiny
travelers in an intimate contest between two implausible foes. Often the
conflict is at its most bitter when relatives are involved. Britain has two
common land snails, much studied by geneticists and hard to distinguish
at a glance. Since the 1960s, one has retreated at the expense of the other
at a rate great enough to render it extinct within a century.
The struggle may last for years. Even the most peaceful place is full of
strife, with any weakness among its inhabitants at once exploited. It can,
as in snails, lead to victory or to defeat; or to an uneasy truce. Sometimes,
the advantage passes from side to side as a reminder that the war of Na­
ture never stops.
From season to season, the number of snow grouse shot in Canada
fluctuates by five times. The rhythm has, say the records of the Hudson's
Bay Company, existed since 1821 (and has in all likelihood gone on for
twenty times as long) . The cause seems simple. Lynx kill thousands of the
birds and as the big cats become common they drive down their prey.
The lynx pay the price and many die of starvation. In the Malthusian
struggle of cat and bird, it seems that first one prevails and then the other.
The truth is more complicated. Grouse are not the lynx's favorite food.
Instead, the cats prefer snowshoe hares, animals sometimes abundant but
in other years almost never seen. The number of hares cycles by forty
times. The lynx, in return, vary seven times over from peak to trough.
What leads to this ten-year rhythm, in perfect time across the whole of
Arctic Canada?
Far more is involved than a simple battle for survival between grouse,
hare and cat. First, the unfortunate hares pay a psychological price for
their predicament. Almost every hare during a population decline is
 Struggle for Existence 61

killed by a predator and the few survivors suffer much stress. This re­
duces their ability to reproduce. The time needed to recover a certain
equilibrium after the predators have gone makes for a lag before the next
hare boom. Other contestants are also involved. Hares eat the fresh twigs
of willow and birch. When these are plentiful, the hares are well fed and
have many young, most of whom survive. Soon, their numbers begin to
shoot up. Parents, children and grandchildren all begin to gnaw back the
twigs. When the food is gone, the population crashes and the hungry re­
mainder are forced into the open, where they become prey for lynx.
The trees can defend themselves. For a couple of years after each
plague their new branches are filled with poisons that make them bitter
and hard to eat. As a result, hare numbers stay low after a crash. In time,
new generations of sweeter twigs appear, the animals return and the cycle
begins again. That of the lynx is a symptom of the hares' own swings in
abundance, which turn as much on food as on predators. Grouse are
dragged in as the hungry lynx turn to them when their main item of diet
disappears. Other reluctant players include squirrels, coyotes and ravens,
all of whom seesaw in synchrony. Red-backed voles dance to the beat of
a different (and as yet unknown) drummer, with cycles of their own. Be­
cause the system has not settled into the usual glum stalemate of Nature,
the complex and unexpected checks and relations between organic be­
ings which have to struggle together in the same country are revealed in
all their intricacy.
Fluctuations of this kind can lead to catastrophe. In 1 980, a thousand
black long-snouted weevils-a grazer on the Salvinia weed in its native
Brazil-were released onto the fern-clogged Lake Moondarra. By April
1 98 1 the lake had a hundred million weevils, but in August came disas­
ter. The fern population collapsed from a peak of fifty thousand tons to
a remnant ofless than a ton. The weevil population crashed in sympathy
and, soon, both participants disappeared.
The ancient and intimate dance of plants and animals on the Cana­
dian prairie stops when settlers move in. The confusion of Nature yields
to the tedium of the farm and, as fields break up forests, the number of
hares steadies. When a patch of forest has been filled, the excess can move
to an empty wood with plenty of young twigs. The circle has been bro­
ken by agriculture and, although the struggle goes on, the evidence is
hidden because the numbers of each player do not change.
Such stability is true of most of Nature-but it does not mean peace.
62 D A RW I N ' S G H O S T

Life, like a silent forest, may seem to be in harmony, but often that is no
more than chaos postponed. The battle may be long deferred, and it may
involve not different kinds-lynx, hare and birch-but the same. In­
deed, the struggle almost invariably will be most severe between the in­
dividuals of the same species, for they frequent the same districts, require
the same food and are exposed to the same dangers.
Chile has huge forests of southern beech, filled with plants that started
their lives at the time of Columbus. For mile after mile, trees of the same
kind and about the same height cover the land. A beech grove is at first
sight as calm a place as any on Earth. Its branches stretch a hundred feet
toward the sun and shade out most of the light. Little can survive be­
neath the canopy-not even beech itsel£ Most places have no young
plants, which is odd in a forest that has lasted for so long. The beeches,
in the absence of recruits to replace plants dying of old age, seem
doomed. Every tree makes innumerable seeds in its five-hundred-year ex­
istence, but for few do any succeed.
Sometimes, though, there is an earthquake, eruption, or storm.
Records from the 1 700s show that a typical stand is wrecked by gales or
fire within three short centuries. Then, life erupts and the survivors have
a chance. The sun floods in, young plants shoot upwards and the strug­
gle starts anew as the winners block light from their inferiors. All the
trees in a beech forest are the same age because they all started to grow at
the same time. Their conflict may last for just a few months in a hundred
years or more but is as vicious as the fight between lynx and hare. What
seems a forest primeval is in truth an interval between catastrophes. Bat­
tle within battle must ever be recurring with varying success; and yet in
the long run the forces are so nicely balanced, that the face of Nature re­
mains uniform for long periods of time.
The forest is a cathedral, but it often burns down. Lightning strikes
the earth two hundred million times a year and sparks off innumerable
fires. The pioneers of North America remarked on the park-like land­
scape of New England. Its timber was scattered not for aesthetic reasons
but because it had been burned by lightning or by Indians. Not before
fire control did Longfellow's "murmuring pines and the hemlocks,
bearded with moss, and in garments green" have a chance to grow. Five
thousand years ago the jungles of South America were reduced by fire to
copses encircled by grassland. Much earlier even Ireland was covered
 Struggle for Existence 63

with tropical forest. Its rocks are filled with ancient charcoal, the rem­
nant of a forgotten firestorm.

Farmers try to second-guess the struggle for existence when they move
plants or animals from their native land to somewhere new. How their
subjects survive and what kills them in their novel abode are clues to
what they face at home.
Penzance, Nice and Alicante are proud of the palms along their
seafronts. All are aliens as, apart from a few dwarf specimens in southern
Spain, Europe has no native palm trees. They fail because their shoots are
damaged by frost. A single cold night in fifty years-a moment of defeat
in Nature's struggle-will stop them.
The denudation of the Cornish Riviera is a trifle compared to the risks
faced by coffee growers. Wild coffee is found nowhere in the world where
the mean temperature in any month drops below that of an English
spring. A millennium and a half ago, farmers got into the act. In the
Ethiopian province of Kaffa, coffee was used to season food. A thousand
years later it was roasted, ground and used as a drink by Arabs. Now mil­
lions of tons are grown each year. Almost half the crop comes from Brazil
and Colombia, half a globe away from its home. For most of the time,
conditions are ideal-but, sometimes, it freezes.
The New York Coffee, Sugar, and Cocoa Exchange gambles against
the weather, with all the uproar of an "open outcry" trading floor. The
growers and roasters deal in futures, a bet on the price in months or years
to come. They hope to reduce their exposure to the risks of cold. Their
opponents count on the profit to be made from the changes that follow
a sudden frost. Fortunes can be won or lost on a guess about bad weather
as the contest between speculators moves back and forth.
The market is evidence of a struggle against the elements. Coffee's re­
cent peaks in price were reached in 1 979, 1 98 1 , 1 986 and 1 995. The
cycle is driven by chance changes in climate rather than, as in the snow­
shoe hare, a shifting balance between implacable foes, but its peaks and
troughs are just as much a response to stress (in this case mitigated by
greed) . A frost year would kill the crop without human intervention; and
most good harvests are followed by a slump brought on by excess pro­
duction in a buoyant market.
64 D A RW I N ' S G H O S T

Coffee has survived, but its insecurity in an alien home puts the econ­
omy in constant danger. From the 1 890s, "coffee presidents" ruled
Brazil. Crop failure, followed by glut and price collapse, led to unem­
ployment and revolution, as a reminder that the natural state of any
ecosystem is of convalescence between frightful battles.
For coffee, the weather is a simple enemy-it kills it. For other plants,
even when climate, for instance extreme cold, acts directly, it will be the
least vigorous, or those which have got least food through the advancing
winter, which will suffer most. Many crops struggle more with starvation
than with hard weather. Farmers fertilize their fields because more food
means better harvests. Often, some of the fertilizer leaks out and its re­
cipients respond . .& the River Po spews filth into the Adriatic it releases
algae from their struggle. They form mats for a hundred miles down the
coast.
Parts of the open ocean are so pure as to support little life. Great quad­
rants of the Pacific, from the Equator to Antarctica, have almost none of
the tiny marine plants-the plankton-that abound elsewhere. The seas
have plenty of the nitrogen and phosphorus needed for growth. What
the plants search for and fail to find are minute amounts of iron. With­
out it, the green machinery that soaks up the sun's energy is starved. A
thousand pounds of iron salts added to a five-mile square of sea near the
Galapagos caused the numbers of plankton to explode. A shortage of a
few parts per million of a simple but crucial item limits life over whole
oceans. The seas were more productive at the time of the ice ages because
of the battle for iron, which, in those cold and dusty times, was blown
far out to sea from the parched land.
Much of the struggle for existence-on land as much as in the sea­
is for a share of the sun's energy. The sunnier a place, the more kinds of
animals and plants it contains; and, within any locality, animals that
keep themselves at a constant temperature are less numerous than those
that do not. In general, the more active a creature, the less common it
is-in Britain, for example, land mammals are a hundred times as abun­
dant as birds of the same size, in part because fliers need more energy to
stay alive.
The fuel for nature's commerce is not well used. On land, about one
part in three hundred is trapped by plants, in the sea even less. The en­
ergy flows upward, but does not go far. Most food chains have four or
 Struggle for Existence 65

five steps from bottom-the plankton in the ocean or the grass of an

African savanna-to top, be it lion, eagle, or killer whale.
When a deer eats a leaf, it gains less than a tenth of its worth as food.
The lion that feeds on the deer does rather better. Even so, more than
half the prey's value is wasted. Too many links in the chain leave too lit­
tle for those at the top. It becomes impossible to add another and to
evolve a beast that preys on blue whales, or on lions. Such an animal
would have to be fast, fierce, and of necessity rare. Most of the islands of
the Outer Hebrides have no eagles as they are not big enough to provide
enough food. A bird that ate eagles would need most of Scotland to sup­
pon itself
Short though life's chains may be, their links are tightly packed. South
Australia was once covered with a spiny scrub called the jarrah. It had to
cope with drought and with the giant kangaroos that browsed upon its
leaves. A ten-yard square may contain a hundred kinds of plants. Intense
competition means that each bush or flower is forced into its own nar­
row niche and that together, they exploit all that sun, soil and rain can
offer. Some are tall, some shon; while some roots penetrate deep and
others spread out.
Now, all have been replaced by a great field containing a small part of
the five hundred trillion wheat plants in the world. Wheat is sheltered
from ecological reality. It grows for half the year and wastes most of the
sun's energy as it passes through its upright stalks. The fields flood in
winter and in summer dry and starve because every plant grows to the
same depth. Entire harvests may be destroyed by pests. The jarrah,
shaped by generations of struggle, was more resilient. Where cultivated
fields have been abandoned, the jarrah creeps back. Wheat, in contrast,
cannot force its way into the native vegetation without human help.
For jarrah, wheat and giant kangaroos the conflict is obvious. Any
plant or animal must avoid its enemies, find a place to live and get
enough to eat in order to have a chance to pass on its genes. However, to
win those engagements is just to set the stage for the greatest contest of
all: the battle for sex.
When it comes to that interesting pastime, most members of most
species come to grief When they fail depends on their habits. For ele­
phants, the majority survive, although not all have offspring, but for or­
chids (which may have a billion seeds) a minute proponion last to have
66 D A RW I N ' S G H O S T

issue of their own. Investment in each stage depends on its chances of

success. An acorn is tiny compared to its parent, but a kiwi lays an egg a
quarter her own weight.
Malthus himself saw that the struggle for existence involved both sur­
vival and sexual success. With his Christian standards, he was a strict ex­
aminer. Far better for the excess to fail at once and die young, rather than
interfere with God's procreative plan. A boy who "cannot get subsistence
from his parents . . . and if society do not want his labour" should face
the facts. ''At Nature's mighty feast there is no vacant cover for him. She
tells him to be gone, and will quickly execute her orders." Any attempt
to rig the test by "unnatural promiscuous concubinage . . . violations of
the marriage bed, and improper acts to conceal the consequences of ir­
regular connexions" was a vice. Human nature was, to Malthus, red in
tooth and claw, and must be controlled (he favored "moral restraint" as
an escape, and lived up to it, with a marriage at thirty-eight and just three
children of his own) . His image of the cruel fate of the surplus popula­
tion still sums up, in the public mind, the struggle for existence.
Tennyson's sanguinary phrase reflects not a rule of life, but despair at
the death of a friend. Nature is often not at all like that. The battle may
be for light rather than land, or for traces of metal instead of terrified
prey. It may be confined to a few moments in a whole lifetime. The war
of nature is not incessant; no fear is felt, death is generally prompt, and
the vigorous, the healthy and the happy survive and multiply.
Except, that is, when politics comes in. Then, the struggle is to the
end. Mark Twain parodied it: "I confined an Irish Catholic from Tip­
perary, and as soon as he seemed tame I added a Scotch Presbyterian
from Aberdeen. Next a Turk from Constantinople . . . a Buddhist from
China; a Brahman from Benares. Finally, a Salvation Army Colonel from
Wapping." A few days later "there was but a chaos of gory odds and ends
of turbans and fezzes and plaids and bones and flesh-not a specimen
left alive. These Reasoning Animals had disagreed on a theological detail
and carried the matter to a Higher Court."
Those animals, reason as they might, have often destroyed their re­
sources, and their lives, as the consequences stare them in the face. Fish­
ermen are closer to the wild than are coffee traders and act-like any
predator-only in their own short-term interests. Politicians try to con­
trol them, but almost always fail. As in the beech forests of South Amer-
 Struggle for Existence 67

ica, lethal skirmishes are separated by intervals of calm. The annual quota
of Alaskan halibut is taken in two days of mad scramble-the Halibut
Olympics. For the rest of the year, millions of dollars of capital lie idle
and, it seems, at peace. So incensed are its owners that the controls are to
be lifted, at whatever cost to the fish.
Such battles go back to the dawn ofAmerica. The Pilgrim Fathers ob­
tained a grant of land in North Virginia (now New England) to set up a
cod fishery. They had reason to be hopeful. John Cabot, a century and a
half earlier, had scooped fish six feet long from the sea in baskets. The Pil­
grims were better at dogma than fishing and had to pillage the Indians
to stay alive. So devoid was the country of their accustomed diet that
they reported with some chagrin that the sole "dish they could presente
their friends with was a lobster. "
In the end, cod triumphed and New England boomed. The court that
burned the witches of Salem had a codfish on its seal, the first coins
minted in Massachusetts carried its image, and a wooden figure of that
noble animal was hung from the roof of the Old State House in Boston.
The fishery spread north and soon Cape Cod, the Grand Banks and the
Gulf of St. Lawrence provided enough fish to feed the United States and
much of Europe.
The stock seemed boundless: after all, a female codfish can live for two
decades and lay nine million eggs a year. As Alexandre Dumas wrote in
his Grande dictionnaire de cuisine: "It has been calculated that if no acci­
dent prevented the hatching of the eggs and each egg reached maturity,
it would take only three years to fill the sea so that you could walk across
the Atlantic dryshod on the backs of cod. "
The steam trawler was invented in England in 1 88 1 . The break­
through was followed by Clarence Birdseye's discovery that cod could be
frozen, even at sea. Before long, the waters off North America were plun­
dered. The steam-powered harpoon appeared in 1 864. The number of
whales it killed rose from thirty in that year to sixty-six thousand in
1 96 1 .
In 1 992, the Canadian government closed the seas off Newfoundland.
The stocks had fallen to twenty thousand tons from more than a million.
Although the number of fishermen in the world has doubled since the
Second World War, most stocks are in decline and a third of the output
of the continental shelf already ends up on the table. Whales, too, were
68 D A RW I N ' S G H O S T

hunted to near extinction before the moratorium of 1 986. In the 1 960s,

the Soviet Union built the Sovetskaya Rossiya, a whaler the size of an air­
craft carrier. They kept the industry alive (and cheated on their quotas)
to pay for it. It was pensioned off as a marine slaughterhouse for Aus­
tralian sheep.
Now, the old Massachusetts whaling port of New Bedford has be­
come, like Ellis Island, a tourist attraction. It celebrates not a triumph
over the struggle for existence, but a catastrophe. The cod-fishing villages
to the north, in Newfoundland, are an even starker memorial. In spite of
a billion dollars in aid, many have disappeared, while others are sunk in
poverty. They are a monument to Malthus. He saw that "the positive
checks to population are extremely various . . . Under this head, there­
fore, may be enumerated all unwholesome occupations, severe labour
and exposure to the seasons, extreme poverty, bad nursing of children,
great towns, excesses of all kinds, the whole train of common diseases
and epidemics, wars, plague and famines." The fishermen of North
America faced most of them; and, like their prey, have failed.
 C H A P T E R I V

NATURAL S ELECTI O N

Natural selection: the factory for the almost impossible - Its power compared
with man's selection - Illustrations of the action of Natural Selection,
from Manchester to the Caribbean - Speed of action - Its power at all
ages and on both sexes - Evolution's second paper: Sexual Selection -
Sex, age and death - Cuckoldry and the balance of the sexes - On the
intercrossing of individuals - Circumstances favorable to Natural Selec­
tion, as illustrated by agents of disease - Extinction caused by Natural
Selection - Divergence of Character related to the diversity of re­
sources - Action of Natural Selection on the descendants from a com­
mon parent - Superiority of the work of Nature to that of Man

I once worked for a year or so, for what seemed good reasons at the time,
as a fitter's mate in a soap factory on the Wirral Peninsula, Liverpool's
Left Bank. It was a formative episode; and was also, by chance, my first
exposure to the theory of evolution.
To make soap powder, a liquid is blown through a nozzle. As it streams
out, the pressure drops and a cloud of particles forms. These fall into a
tank and after some clandestine coloration and perfumery are packaged
and sold. In my day, thirty years ago, the spray came through a simple
pipe that narrowed from one end to the other. It did its job quite well,
but had problems with irregularities in the size of the grains, liquid
spilling through or-worst of all-blockages in t;l'ie tube.
Those problems have been solved. The succes� is in the nozzle. What
used to be a simple pipe has become an intricate duct, longer than be-
70 D A RW I N ' S G H O S T

fore, with many constrictions and chambers. The liquid follows a com­
plex path before it sprays from the hole. Each type of powder has its own
nozzle design, which does the job with great efficiency.
What caused such progress? Soap companies hire plenty of scientists,
who have long studied what happens when a liquid sprays out to become
a powder. The problem is too hard to allow even the finest engineers to
do what they enjoy most, to explore the question with mathematics and
design the best solution. Because that failed, they tried another ap­
proach. It was the key to evolution, design without a designer: the preser­
vation of favorable variations and the rejection of those injurious. It was,
in other words, natural selection.
The engineers used the idea that molds life itself: descent with modi­
fication. Take a nozzle that works quite well and make copies, each
changed at random. Test them for how well they make powder. Then,
impose a struggle for existence by insisting that not all can survive. Many
of the altered devices are no better (or worse) than the parental form.
They are discarded, but the few able to do a superior job are allowed to
reproduce and are copied-but again not perfectly. As generations pass
there emerges, as if by magic, a new and efficient pipe of complex and
unexpected shape.
Natural selection is a machine that makes almost impossible things.
Consider a typical protein such as whale myoglobin. That molecule is
but one of a hundred thousand or so proteins in the animal's body and
contains a hundred and fifty-three units called amino acids. These come
in about twenty forms. The number of possible combinations of amino
acids in a structure the size of myoglobin is hence twenty raised to the
power of a hundred and fifty-three. The figure, ten with about two hun­
dred zeros after it, is beyond imagination and is far more than all the pro­
teins in all the whales, all the animals and all the plants that have ever
lived. Such a molecule could never arise by accident. Instead, a rather or­
dinary device, natural selection, has carved out not just myoglobin but
millions of other proteins and the organisms they build.
Selection is simple, efficient and inexorable. George Bernard Shaw did
not like it: "When its whole significance dawns on you, your heart sinks
into a heap of sand within you. There is a hideous fatalism about it, a
ghastly and damnable reduction of beauty and intelligence, of strength
and purpose, of honour and aspiration." Unfortunately for Shaw, beauty
and intelligence have themselves now turned to that fatal device for help.
 Natural Selection 71

Some artists use a computer to generate altered versions of an original.

Then, by aesthetic preference, they select each generation until a work to
their taste has emerged. Architects do the same, and machines even write
plays (albeit not to Shavian standard) .
Like the tapeworm or the tree kangaroo, the buildings, plots or noz­
zles made by selection may grate upon the senses of a purist. Even so,
they work, and do so without a single blueprint or equation. Instead,
they have evolved through descent with modification. Darwin is loose on
the shop Hoor, and industry has become a branch of the biological sci­
ences. Natural selection has returned the compliment.

Illustrations o/the action o/Natural Selection. In Britain in 1 690 three mil­

lion tons of coal were mined. By 1 950 the figure was eighty times higher.
Evolution has been busy since the invention of the Hying shuttle-the
New Machine for Opening and Dressing Wool-two and a half cen­
turies ago. The Industrial Revolution was a great experiment in biology.
In the eighteenth century, Manchester was described by Daniel Defoe
as the greatest mere village in England-a market town, but little else.
Within a hundred years, its population had multiplied and it had be­
come Cottonopolis . .As de Tocqueville put it: "From this filthy sewer
pure gold Hows. Here, civilisation works its miracles, and civilized man
is turned back into savage." London, too, in its different way, had grown.
The inhabitants of each city and every other had to adapt or die. Many
died-but many more evolved to stay alive.
To stand on a clear day by the monument on Highgate Hill where, five
hundred years ago, Dick Whittington was persuaded to turn again, is,
sometimes, to take in a splendid view. It extends across London to the
Downs, twenty miles to the south. Too often, though, the best to be seen
is the glum towers of the City of London as they peer through a sul­
phurous haze. Smog is a London word, coined to describe the smoky fog
that once killed thousands. Just thirty years ago it let through a third less
sun in the East End than in the more afHuent west of the city.
1 848 was a year of revolutions, brought about by the social changes
that followed the spread of manufacturing. In February, in Paris, the
populace rose and overthrew the emperor. A few months later Austria,
Germany and Hungary were in tumult. The revolts were a sign of the po­
litical storms that were to form the modern world.
In Britain, the main upheaval was among the moths. In that year there
72 D A RW I N ' S G H O S T

was a report from Manchester of a black form (a melanic) in the pep­

pered moth, an animal that had until then been gray. By the time of Dar­
win's death in 1 882, the melanic type was the commonest form in much
of industrial Britain. Collectors were quick to notice: "Near our large
towns where there are factories and where vast quantities of soot are day
by day poured out from countless chimneys, fouling and polluting the
atmosphere with noxious vapours and gasses, this peppered moth has
during the last fifty years undergone a remarkable change. The white has
entirely disappeared, and the wings have become totally black." It was
the nation's response to the Industrial Revolution.
The force of selection was simple. Dark moths could not be seen by
birds against the filthy trees on which they sat and were safer from attack
than the original form (which was camouflaged against the lichens of
rural woodlands) . The dark moths appeared in London by 1 897. By
then, just one Manchester peppered moth in fifty was of the original
type. In rural Britain, blacks were rare or absent (although a few were
found in East Anglia, a place much polluted by drifts of smoke from the
Midlands) . The rest of the world tells the same story, as melanics turned
up in peppered moths in the industrial parts of Europe, and, from 1 906,
in North America too. Japanese peppered moths remain white, but they
live in the countryside and have not gone into towns. All over the world,
natural selection had responded to the new challenge. It caused melanics
to spread so fast after 1 848 that a simple sum shows them to be half again
as able to succeed in cities as was the original form.
With smoke-control, the dark moths began to disappear. As the soap­
works were cleaned up, the frequency of blacks in the Wirral fell from 95
percent in the late 1 950s to 5 percent at the turn of the millennium.
Soon, the new form will be gone, and the latest episode in the moth's
evolutionary adventures will be over.
London still has black mallards, black pigeons and black squirrels
(which became less common in America at the time of its own Industrial
Revolution, because dark fur was in fashion). Life in the city affected sex
as much as survival. Unlike their gray relatives, black pigeons-the coo­
ing majority in Trafalgar Square-breed throughout the year. Their pig­
ment soaks up more of the sun and persuades the calendar in the brain
that winter days are long and that it is time to turn on the testosterone.
Cities, unlike the birds' ancestral cliffs, provide enough food to breed in
cold weather. As a result, melanics, scarce in the countryside, prevail.
 Natural Selection 73

Town dwellers must deal with a world of change, or die. That is noth­
ing new. The countryside is as merciless as the toughest slum. Every plant
or animal descends from ancestors tried by a struggle for existence for
three and a half billion years before cities came on the scene. Man selects
only for his own good, Nature only for that of the being which she tends
(which is why the streets are not full of marauding sheep or tomatoes) .
The Industrial Revolution was a test of the theory of evolution. Its re­
sults were often unexpected, sometimes unwelcome and always un­
planned. So familiar is the testimony of natural selection that we do not
recognize the best proof of its power: the fit of life to where it lives. When
we see leaf-eating insects green and bark-feeders mottled gray; the alpine
ptarmigan white in winter, the red grouse the color of heather and the
black grouse that of peaty earth, we must believe that these tints are of
service to those birds and insects in preserving them from danger.
Evolution, say its critics, is not a real science, because it cannot predict
the future. It has, no doubt, done pretty well at explaining the past. But
how can we know what it will do next? Even Thomas Henry Huxley,
Darwin's Bulldog, felt that his hero "does not so much prove that natural
selection does occur, as that it must occur, but in fact no other sort of
demonstration is attainable." He was wrong. Now, evolution has joined
the scientific mainstream. It designs its own experiments to confirm what
mere observation might suggest.
The Caribbean is full of scores of different kinds of lizards called
anoles, climbers skilled enough to make their way up a vertical sheet of
glass. On each island, the lizards fall into half a dozen physical types, spe­
cialized for the forest crown, for trunks or twigs, or for bushes, grass or
the ground. Bare rocky places have lizards with shorter legs than those
from sites with large trees, because short limbs allow a lizard to be agile
at the cost of being slow. A lizard able to sprint along a thick branch will
fall off a stick, which is not much help when it tries to catch food or es­
cape enemIes.
In 1 977 small groups of lizards were moved from their home on a Ba­
hamian island shrouded in thick vegetation to an islet without lizards of
its own. There, the cover consisted only of thin twigs. Ten years later, the
emigrants had changed. Their limbs were shorter than those of their an­
cestors and had evolved the pattern of stubby legs on thin branches
found throughout the Caribbean. On the new island, natural selection
had been at work, daily and hourly scrutinizing every variation, even the
74 D A RW I N ' S G H O S T

slightest, rejecting that which is bad, preserving and adding up all that is
good; silently and insensibly working at the improvement of every or­
ganic being in relation to its organic and inorganic conditions of life.

Short legs (or black wings) help their owners to stay alive. That is of
course important, but selection, in the end, depends only on the relative
number of offspring. A long, happy and well-adapted existence, without
issue, is of no use at all. What matters is how many genes are passed on.
As no animal can do everything, evolution, like economics, is full of
compromIse.
Any investment-in flesh or in cash-trades off stability against
growth. The highest yields are from a risky bet, while security means a
lower rate of interest. Because (as Malthusians and financiers each know)
compound interest is powerful, any speculation in genetic futures needs
to squeeze in as many rounds of interest, with as high a rate of return, as
possible. The market, however, calls for a decision between a long life
and a fertile one, with success in one vocation paid for by failure at an­
other. Chickens bred for meat lay fewer eggs. Even among the egg layers
those selected to lay young die before their time so that the number of
eggs goes up not at all. A balance of cost and benefit molds what any
business (evolution included) can do.
The ecological market determines when plants and animals should
make the biggest investment of all: to reproduce. Variation in when to
breed, in how much risk to take, and in how many young to produce is
potent fuel for evolution. In their happier days, when some were the size
of a man, North Atlantic cod delayed their sex lives until they were four
or five years old. Now, the giants have gone and the few that escape the
net are tiny. With no chance of old age, evolution favored those that re­
produced as soon as they could. Those reduced animals have struck a bal­
ance between mortality and sex, and now lay eggs at the age of two.
For cod and all other creatures, sex and death are weighed in the scales
of selection. Trinidadian guppies have different lives in different places.
Fish in the lower rivers are much harassed by predators. As a result, most
die young. Because their enemies cannot get above the waterfalls, gup­
pies in mountain streams are free from attack and are guaranteed a death
deferred. As a result, lower and upper rivers evolve their own investment
 Natural Selection 75

strategy. Upstream guppies rejoice i n peace and quiet, while life for their
lowland relatives is shorter and nastier. With so few chances of an erotic
encounter before Nemesis arrives, the downstream fish put all their eggs
into one sexual basket. They mature more rapidly and have more and
smaller young, each with a lower chance of survival and faster growth,
than do fish from higher up. Fish moved from the lower parts to the
headwaters soon evolve local ways. Within ten years they grow larger,
delay sex and have fewer offspring. The males adapt to their new and re­
laxed home by evolving at ten times the rate of their consorts.
As natural selection adjusts a guppy's existence, it illustrates the links
between sex, age and death. They are universal and unavoidable. The
more a fruit fly copulates, the sooner it dies, and the more cones a Dou­
glas fir makes, the slower it can grow. Life is a gamble. Any animal must
decide whether to make a large stake in the hope of a payoff, or to delay
a bet in the hope of better odds. As insurance companies realize, that
logic applies to death as much as to sex. If astrology worked and every­
one knew when they would die, nobody could sell any policies. Clients
told that not much time is left would buy coverage at once, but those
promised many happy years would spend the money elsewhere.
Insurance and evolution must each cope with the fact that risk goes up
with time. As I write this chapter I have, as a fifty-something male, a
chance of around one in a thousand of death before I finish it; and, I have
to say, I am not too worried. The chance is hundreds of times bigger than
that of a lottery win, should I be fool enough to buy a ticket. However,
the two risks are subtly different. Not every gambler wins the lottery, but
all of us (and every fish) will die. A sixteen-year-old faces a one in two
thousand probability of demise before his next birthday, while for cente­
narians the figure drops to one in two. Unlike lottery odds, the yearly
danger of death alters with time-for humans, doubling about every
eight years.
The worth of a young animal, calculated in the currency of how many
young it can have, is, as a result, higher than that of an old one. The force
of selection, like the value of a new insurance policy, weakens with age
because less time is left for the contract to run (which is why, for down­
stream guppies, a short life means a heavy investment in premiums) . A
slight increase in the chance of reproduction when young is worth more
than a sexual triumph long delayed. This means that evolution favors
76 D A RW I N ' S G H O S T

youthful vigor at the expense oflater decline. Why should it worry if the
price of sex is to become a burned-out wreck? Any gene able to help its
carriers to copy their DNA will spread, however evil its effects-if they
are long enough deferred. Selection favors only those who can pass on
the genes and cares not at all for those who cannot. Age is a tax on sex,
levied by natural selection. It is as much a product of the struggle for ex­
istence as are the black wings of the peppered moth.

Sexual Selection. Sex is a marketplace for natural selection. As a merger

between genetic enterprises it is, like any other business with two part­
ners, liable to discord. To find a mate, fight off the opposition, suffer sex­
ual congress and pregnancy, and raise a brood, are all expensive and
dangerous. Each involves subtle differences in the investment strategy of
the parties involved. Sometimes there is a sexual war that may lead to
traits that seem opposed to the interests of those who bear them. They
are a reminder that the biological battle goes much further than the
search for food or for light: there is, in addition, an unavoidable struggle
to pass on genes.
Evolution is an examination with two papers. To succeed demands a
pass in both. The first involves staying alive for long enough to have a
chance to breed, while the mark in the second depends on the number
of progeny. Malthus assumed that sex was so attractive that, given the
chance, all would indulge as much as they could and that birth rates
would always be near their maximum. Death was what counted. He was
wrong. Until the eighteenth century, the English married late-at
twenty-five for women and even older for men. The rapid growth in
numbers at the beginning of the Industrial Revolution came not from a
decline in the power of death, but from an emancipation of the national
libido. The age of marriage dropped by several years so that changes in
fertility, not in survival, caused the population explosion that gave Dar­
win his theory.
Inherited differences in sexual ability fuel a subtle and unexpected
kind of evolution. It gives rise to attributes that seem contrary to the very
idea of selection because they reduce the survival of those who bear them.
In fact, they are evidence of its subtlety and power.
The dropouts from the first part of the ecological exam are easy to
identify as they are, after all, dead. Variation in sexual success is more dif­
ficult to measure, particularly for the male candidates {who tend to dis-
 Natural Selection 77

appear before their papers are ready for marking) . The battle is of male
against male, of female against female, and of either sex against the self­
ish interests of the other.
The successes and failures of two hundred marked female spar­
rowhawks in Eskdale in southern Scotland were followed for a quarter of
a century. Like the swifts in Selborne the number of birds in the valley
stayed about the same. That stability hides great differences in how well
each did in the contest to transmit DNA. Most of the birds are in an evo­
lutionary dead end. Three out of every four females die young, fail to
find a mate, or have no progeny; but some triumph in the sexual strug­
gle. A fifth of the young produced nine tenths of the next generation. A
few live for a decade and have two dozen fledglings, while many more die
before they have any chicks at all.
For sparrowhawks, life is unfair, with no question of equal shares, be
they for food or for sex. Man himself insists on inequality. As a result,
wealth-which leads to happy old age and plenty of sex-soon becomes
concentrated among a few. Half the private land in Scotland is owned by
three hundred and fifty people; and the greatest proprietor of all, at a
quarter of a million acres, is the Duke of Buccleuch. The Duke's lesser ti­
tles include a couple of earldoms, a barony or two-and the lordship of
Eskdale.
How many of the testamentary differences among birds or dukes are
due to good genes, how many to good parents and how much to good
luck nobody knows. When it comes to the nobility, good (or aristocratic)
parents are what count. For the birds of the ducal acres, success may say
more about genes. Whatever the reason, in Eskdale or anywhere else,
most sparrowhawk-most animal-DNA does not make it to the next
generation.
Dukes vary more than duchesses in the ability to make the reproduc­
tive grade, because the struggle for sexual existence nearly always bears
more upon males. They put less effort into each encounter, because
sperm are cheaper than eggs and because those who make them are not
left holding the baby. Males have more chances for sex-but they are
limited by the numbers of partners available. The imbalance means that,
in general, any female, however unattractive, will find a mate, but that
just the best males will win. Males must, as a result, fight for access to fe­
males as much as for any other resource in short supply.
Their struggle is manifest in many ways. Often, the winner must hold
78 D A RW I N ' S G H O S T

a territory against his ardent rivals. Shoot a male red grouse and another
moves into his space, proof of how crowded existence must be. If a bird
is injected with the male hormone testosterone, the size of his patch in­
creases. Like a bodybuilder attacked by steroid rage, he displaces his un­
lucky neighbors from their homes. They may survive, but the battle to
transmit their genes is over.
Many of Nature's most attractive features-flowers, birdsong, man­
drill bottoms-result from rivalry among male sex cells for access to eggs.
Male alligators have been described as fighting, bellowing and whirling
round, like Indians in a war dance, for the possession of the females; male
salmons have been seen fighting all day long; male stag beetles often bear
wounds from the huge mandibles of other males. Many are armored; for
the shield may be as important for victory as the sword or the spear. Even
the giraffe's neck is involved. If access to high leaves is important, why
stretch necks rather than legs-and why do males have longer necks than
females? In fact, they fight (and die) over mates by battering each other
with their heads, and a longer handle makes a more effective club. Be­
cause their brain is ten feet higher than their heart, giraffes have the high­
est blood pressure of any mammal. The sexual struggle is to blame.
Sexual selection molds the public image of biology, but natural selec­
tion is always on the alert to stop it going too far. In Uganda in the
1 930s, almost every male elephant had tusks, structures evolved (at least
in part) as statements of reproductive excellence. Sixty years later, ivory
poachers had much reduced the number of animals, with those with the
largest tusks at greatest risk. Now, a third of adults are tuskless, because
the negative effects of ivory on survival outweighed its role in allure.
Some females are firm in monogamy, but many more go in for mani­
fold males. The successful (and anxious) partners follow their mate and
beat off the competition and its insistent sperm. Dogs have the unsubtle
strategy of copulating for long enough to give their sperm a head start
over the next candidate, while mice and many insects insert a plug to
keep out a later deposit. Those excluded can fight back, with a hooked
penis to unblock the female or a nozzle to siphon out any earlier contri­
bution. The conflict goes on after sex, because a sperm has to compete
with others donated by the previous swain. A typical mammal has a hun­
dred million sperm in each ejaculate, which leaves plenty of room for an
internal struggle. A hidden battle rages inside any female who mates
 Natural Selection 79

more than once. A quick DNA test shows which partner succeeded as a
father. Often, the last male to mate sires all or most of the young. It hence
pays a male who mates with a female for the first time to make a lot of
sperm to flood out an earlier donation. As a result, ejaculates with a new
partner are several times larger than those produced for a familiar mate.
Whoever is on top, the sexual interests of males and females differ. It
pays all males to be lazy, selfish and debauched, while any female is bet­
ter off with an active, helpful and faithful spouse. Such paragons are hard
to find and may be less trustworthy than they seem. Certain flies court
for days before a male can persuade a female to respond, but once he suc­
ceeds, he inserts a cocktail of chemicals. Some force the female to lay eggs
sooner than otherwise she would, and reduce her sexual interest in later
males. Because of the male poisons, a female who mates many times dies
long before one who mates just once.
Plants do not escape the battle of the sexes. The flower is an organ of
allure, evolved to attract the pollinators who move male genes. In flow­
ers that are both male and female, a reduction in size makes little differ­
ence to female success, as just a single bee can deliver the crucial genes.
It has, in contrast, a great effect on the export of pollen. Plenty of polli­
nators means lots of male sex cells on the move. Flowers are silent
screams of masculine passion.
Often, their cries are unheard. In some orchids, a mere one plant in
fifty attracts even a single insect. Pollinators do thirty billion dollars'
worth of work for American farmers each year. In orchards, with millions
of flowers in bloom at once, sexual competition is so intense that grow­
ers hire hives of peripatetic bees to ensure that there are enough to go
round. Insecticides have killed half the honeybees in the United States
over the past decade or so. Many plants now fail to set seed, and some
crops-such as blueberry-have collapsed altogether. The sexual strug­
gle in American plants is more intense than ever.
Males are not always the aggressive sex. The female spotted hyena has
a structure remarkably like a penis: an enlarged and erectile clitoris, as big
as her partner's genital organ. For intercourse, she rolls it back, rather like
a man might roll up the sleeve of his shirt. She gives birth through her
penis-and it must be a painful business, as around a sixth of all females
die when they have their first cubs. When it comes to sex, the female
spotted hyena has an unrelaxed existence that calls for some steroid
80 D A RW I N ' S G H O S T

rage-and the appendage that grows from it. Her bizarre organ appears
because she makes lots of testosterone, a hormone more often associated
with males. As in the grouse, that useful chemical is involved in a sexual
battle. In spotted hyenas, females are in charge. The top bitch passes on
her rank to her daughters, who must battle to keep it. Status pays, be­
cause females at the top get more meat and have twice as many young as
do others. Hyenas have twins, and newborn sisters attack each other
within minutes. One is killed, or lives as a cringing inferior who may
never have cubs of her own. All this stress leads to the production of male
hormones-a great aid to courage-by females. The ladylike penis fol­
lows on.
Because of its well-endowed female, Aristotle believed the spotted
hyena to be a hermaphrodite. He was wrong, but animals in which boy­
girl meets girl-boy are common. They are an experiment in the struggle
of the sexes. A long debate determines who will be unlucky and bear the
cost of eggs rather than sperm. To mate may take hours, with an elabo­
rate nuptial dance. Careful bargaining ensures a fair division of the bill
(although in some slugs the argument goes so far as to involve each part­
ner trying to bite off the other's penis) . In sea slugs the first penis enters,
and then the other, while in hermaphrodite fish a series of packets of eggs
are traded for an equal number of bundles of sperm.
'
Sexual conflict is as bitter as the struggle for food or for light. It can
produce traits that are favored only because they increase sexual success,
even as they reduce the chances of staying alive. Alarmed by the cost of
such a structure, Darwin wrote that "The sight of a feather in a peacock's
tail, whenever I gaze at it, makes me sick." The outlandish organ, and
many like it, reduces its bearers' hopes of survival and as a result might
seem evidence against evolution. However, the birds' gaudy train does
not deny the truth of Darwinism. Instead it is a reminder of the rigors of
its second examination paper. Such bizarre decorations are driven by a
particular force of selection called sexual selection. That depends, not on
a struggle for existence, but on a struggle between the males for posses­
sion of the females; the result is not death to the unsuccessful competi­
tor, but few or no offspring.
In sex, who wins is often a matter of status, determined by fights,
threats and submissions. The risk is real: among baboons, for example,
half of the males who try to take over a group from its owner are seriously
 Natural Selection 81

injured. There can never be enough status to go around, because a top

dog exists only by virtue of his inferiors. Such battles can be reduced with
the use of signals to establish whether a challenge is worthwhile. To sport
a badge of rank is cheaper than to face the risk of violent death. The sym­
bols include massive antlers, loud croaks, or characters invisible to the
human eye. Male and female blue tits look the same to us, but in the ul­
traviolet (seen by birds) the males have a conspicuous patch on the head.
Male insects make complicated scents, or sounds inaudible to ourselves.
Many have ornate genitals, studded with knobs and spikes. Such titilla­
tors (from the Latin titilfare, "to tickle") were once seen as keys to open
a partner's lock and ensure that a female accepted a male of her own kind.
They are, however, found only among insects whose females mate many
times. Those with faithful females have simple penises. A spike may
hence be a statement of excellence, rather than identity, tested by one
party before the other is allowed in.

However bizarre the behavior involved, sex is, in the end, a struggle to
find someone to fuse with. Sperm and egg do not grow or build a home,
but they do unite. All biographies come in two parts. A long and tedious
episode of advance from egg to adult is followed by a brief fight to find
another cell. Selection is hard at work on both legs of the journey from
egg to grave. The opposed priorities of sex and survival explain why the
two sexes look so unalike, not just in trivial characters such as antlers or
tails, but in their very essence.
To meet someone else it pays to be common and to move around a lot.
As a result, sexual selection favors small, active and abundant sex cells. To
get a head start on the way to adulthood needs a large and well-stocked
embryo, and natural selection is all in favor of reproductive cells that are
big, stuffed with food, and reluctant to waste energy in going places. The
balance between the two forces explains the evolution of sperm and
egg-and of the sexes themselves.
Males have small sex cells, females large. The difference (the true def­
inition of what the sexes actually are) results from an ancient conflict.
Long ago (and in some simple organisms today), sex cells were all the
same size and fused to make an embryo, well provided with food. Then
self-interest made an appearance and one partner moved to making
82 D A RW I N ' S G H O S T

smaller but more abundant cells. He (for such was, from that moment,
his gender) might have hungry young, but there were more of them. His
success was limited by the risk of those small cells fusing to make an em­
bryo with too little food to grow at all. Selection favored union with the
large cells of his ally, and so the sexes were born. Males have been frauds
since they began.
All males make more sperm than can succeed. This leads to a struggle
among their sex cells. Often, a male shoots himself--metaphorically­
in the foot as, once an egg has been fertilized, the sperm shut out may
well be his own. If he is the only male in the reproductive arena, there is
not much point in lots of sperm (or in being big enough to make it). As
a result, in most animals (with a few familiar exceptions, such as cows,
deer or humans) males are smaller than females. Sexual selection has in­
sisted on small and active sons, natural selection on large and well-fed
daughters. Some males are tiny. The male anglerfish is a worm-like ap­
pendage attached to his mate. Certain barnacles, indeed, were once
thought to be hermaphrodites, but the male is there, snug inside, busy
with sperm and not much else. In the vastness of the sea each male has
but a small chance of a sexual encounter. The best strategy is to have
small males who stick like glue on the rare occasions when one makes the
grade.
Not until the battle escalates, with males rather than sperm at war, do
there evolve the virile beasts so beloved of television producers. Fish show
how size can matter. Many start as one sex, and change into the other as
they grow. Some are male first, some female. Precedence follows a simple
rule. In those whose sex cells are ejected into the water (with competi­
tion among sperm rather than among males themselves) it pays to be
male first, because any male, small as he is, can generate large amounts
of sperm while an older (and larger) female can make more and better­
fed eggs. Not unless the battle is among males for possession of the fe­
males does it pay to make eggs first and to delay manhood until full size
and aggressive potential is reached.
The sexual equation has two terms. Males compete for females, while
females choose the males that they find, for some reason, attractive. Male
battles are obvious enough, but can the aesthetic tastes of females drive
males to such lengths? It may seem childish to attribute any effect to such
apparently weak means; but if man can in a short time give elegant car-
 Natural Selection 83

riage and beauty to his bantams, according to his standard o f beauty,

there is no good reason to doubt that female birds, by selecting, during
thousands of generations, the most melodious or beautiful males, ac­
cording to their standard of beauty, might produce a marked effect.
Females do not always approve of male ardor, because it wastes time
and cuts down their own chance to choose. As well as sex, females have
to find food and a place to live and lay eggs. They may be loath to accept
a partner and some female insects are decorated with sharp spikes to dis­
courage courtship.
Females, nevertheless, do prefer the frog with the deepest croak, the
bowerbird with the most decorated nest, the plant with the most sym­
metrical Hower, and so--endlessly--on. Why? There are many ideas
about the evolution of such sexual habits, and perhaps almost as many
truths behind them.
The choice may be eased by a meal, or somewhere to live. Many male
insects give their mates a tasty grub before sex. The larger the gift, the
more acceptable the partner. Female fruit Hies digest the ejaculate itself
to help them make eggs; but other insects ask for more. A certain moth
is protected from its enemies by a poison picked up from plants. A male
passes on with his sperm a solid dose of the substance, which in turn
reaches the eggs. His sex scent (used to attract the females) is based on
the poison, and the more scent he can make, the safer his partner and her
eggs and the better mate he will be.
Sexual selection leads to a folie a deux. If the best choose the brightest,
female preference and male trait will evolve together. The system has
volatility built in, and once the sexual fuse is lit, an evolutionary explo­
sion is more or less guaranteed. If a sexy male has sexy sons who are cho­
sen by the most fastidious females, the process will run away with itself
to give characters as strange as the blue tit's crown. How far it can go is
limited by natural selection. Even an ultraviolet coronet is expensive, be­
cause it can be seen by sparrowhawks.
Females often choose mates on the basis of what genes a male might
carry. Because of the dangers of incest, a brother or a cousin is a less at­
tractive partner than is a nonrelative. Whole systems of kin recognition
have evolved to avoid such matings. Mice, for example, excrete scents in
their urine that enable females to avoid their close kin. But why should
the choice involve a showy crown or a pair of luminous buttocks? What
84 D A RW I N ' S G H O S T

does a female gain from such a well-endowed partner? His characters

seem at first sight useless or even perverse. It may be-once more-a
matter of marketing. An extravagant signal could be a general statement
of a male's ability to cope. A peacock able to manage his accessory must,
no doubt, be quite a lusty specimen and could make an excellent father.
As soap makers know, advertisements are expensive. That is part of
their point, for if a company can afford lavish publicity it must be a solid
concern. Because cheats devalue the currency, business goes a long way
to protect its brands. The Distiller's Company in London-makers of
Gordon's Gin and Johnnie Walker Whisky-has a "black museum" of
fakes from around the world (the least plausible a shampoo, Johnny
Washy's Old Sloshy, which a customer once drank by mistake) . The
company always sues those who mimic them, and almost always wins.
France once had a brutal and much-feared band of bank robbers called
the Moustache Gang. Cashiers learned to hand over the money as soon
as they appeared. In a few months there appeared squads of counterfeit
gangs with the trademark but not the violence. Quite soon, the intim­
idatory power of upper-lip hair began to wane. In the same way, a feeble
male sparrow painted with the black bib of sexual dominance is attacked
by other males for his presumption.
If a male can afford an expensive symbol, to pick him as a mate may
help, because his young may be as well endowed. In tree frogs, females
prefer males who make a longer call-and they gain nothing but his
DNA, because he disappears before the tadpoles hatch. When half the
eggs of a female are fertilized by long-call males, and the rest with sperm
from those with short calls, the former grow faster. The genes advertised
by their father in his lengthy croaks do indeed help his offspring.
The work of natural and of sexual selection stands in great contrast.
Natural selection often reaches similar ends with different means. Thus,
wasps or snakes that warn of danger each use black and yellow, while
both grouse and peppered moths have mottled camouflage. Sexual char­
acters are more capricious. Each haughty male has a badge unique to his
own kind. Some bowerbirds are bright and showy, but others are drab
and excite their females with elaborate shelters instead. There is no gen­
eral statement of sexual excellence (apart, perhaps, from size) common to
all the male birds, or to all flowers. Perhaps the characters are arbitrary
and the ancestral female's choice was frivolous. The machinery of sexual
selection did the rest.
 Natural Selection 85

Males can be beguiled into a signal by the workings of the female

mind. A bird brain is quite naive. A female oystercatcher given an egg
several times the normal size will abandon her nest to sit on what seems
a manifest fake. The message that says "egg" overwhelms what little com­
mon sense she has. Female zebra finches prefer sex with males with red
plastic leg rings and even have a penchant for a mate decorated with a
white chef's hat. Eccentric as these quirks seem, they are no more than
enhanced versions of what a male zebra finch already has: a red bill and
a white facial stripe proclaiming him to belong to the correct stock. Fe­
male bias is wired deep into the brain, poised to send her sexual partners
into an evolutionary blind alley.
For sex, as for business, what matters in the end is not advertising but
sales. Feeble players-the discount stores of the sexual world-some­
times do as well as their more impressive opponents. The bluegill sunfish
has a society based on lies. Each fish sets off down a path to one of sev­
eral sexual styles. At the age of two, a male makes a momentous deci­
sion-whether to become adult or to stay young. Some mature while
small, but others delay adulthood for six years or more, by which time
they are several times larger than their precocious sibs. When maturity
comes at last, the slow developers are big enough to defend a home.
They own grand premises, but they have a problem: their cunning
sibs, who lurk nearby. When a landlord has enticed a female onto his
patch, the rivals dash in and emit semen. It lacks an impressive wrapper
and comes without a guarantee, but is cheap enough to undercut their
rival. When the sexual scoundrels grow too big to sneak in, they again
change their style. They begin to resemble females, until they can saunter
unafraid onto a territory, the sole risk that of courtship by its besotted
holder. When a real female appears a transvestite's deception pays off. He
fertilizes her eggs and makes a hurried exit.
A large male courts and inseminates females far better than does a
prowler or cross-dresser. It has, though, taken him years to achieve his
sexual peak-and for most of that time his rivals have, in devious ways,
been passing on genes. To lurk or lie needs rapid maturity. A cheat's fee­
ble frame and ability to exploit the market is as much a product of sex­
ual selection as the beefy bodies of his rivals.
For a sunfish or a supermarket, the best game to play depends on what
everybody else is up to. If most of the competition is territorial, it pays
to cheat, because the water is full of potential cuckolds. If impostors be-
86 D A RW I N ' S G H O S T

come too common, then there will not be enough hosts to go round. It
pays to be different and to do what others do not. If a strategy becomes
rare it gains an advantage-but that is lost as soon as it becomes com­
mon. In time, the system settles down and householders and sneaks pass
on genes with equal efficiency.
That explains why most animals have equal numbers of each sex. After
all, males and females are themselves no more than alternative solutions
to the problem of handing on genes. If a single male can fertilize dozens
of partners, why the spares? Why not a hundred females and one male?
The reason is simple: as in sunfish, the rarer caste is always better off. If
so few males exist that each always finds lots of mates, then it pays a par­
ent to have sons. A shortage of females puts an equivalent premium on
daughters. Soon, things come to a balance, with equal investment into
the transfer of genes through each gender.

Sexual selection helps explain the evolution of characters that at first

sight make no sense. Unfortunately, it can also be a general excuse. It is
unwise to attribute all sexual differences to this agency; for we see pecu­
liarities arising and becoming attached to the male sex in our domestic
animals (the wattle in male carrier pigeons, horn-like protruberances in
the cocks of certain fowls, and so on) which we cannot believe to be ei­
ther useful to the males in battle or attractive to the females.
We see analogous cases under Nature. The penis raises man above the
primates. Our organ is, in comparative terms, huge. The gorilla has a
guilty secret: its one-and-a-quarter-inch member. The chimpanzee, a
copulator of gigantic appetite (a male manages hundreds of sexual en­
counters with dozens of females each year), does little better.
Why do men have such large genitals? Some calI on sexual selection as
the cause of penis expansion. Like the peacock's tail, it simply ran away
with itselE A penis may not cost as much in metabolic terms as does a
showy tail, but think how useful it would be if all that too solid flesh were
to be recycled into brain or muscle. Our unique organ might all be a
matter of female choice. A natural experiment seems to test the idea. In
New Guinea, men of the Ketengban tribe enhance what Nature has pro­
vided with a pointed gourd-a phallocarp. Without this additional foot
or so of reassurance, they say, they feel naked. Perhaps this is what the
 Natural Selection 87

penis would evolve into were it not held back by some contrary force. It
is, above all, a signal of virility.
In most places, of course, the penis is not a signal of social or genetic
excellence at all, because so few get to see it. The organ is not flaunted in
the sexual marketplace, but hidden away until the last possible moment.
Even Don Giovanni did not wear a phallocarp. The Duke of Edinburgh,
on a visit to New Guinea long ago, was given one, but has always refused
to reveal whether he put it on. Most Western males follow in the ducal
path. They are proud of their appendage, but in private. As far as is
known, societies that expose or conceal that useful organ have no differ­
ence in penis size, but as both exist it is fatally easy to appeal to the op­
tion that fits the theory. Of course, things might have been different in
the days before trousers, and Homo erectus may have lived up to his name
in more ways than one. To test the idea that today's male accessory results
from sexual selection long ago is even harder than to establish whether it
is at work now.
The dangerous flexibility of the penis argument warns of the risks of
searching too hard for the hand of selection. It has long been fashionable
among evolutionists to mock the claims of the Reverend William Paley,
whose Natural Theology (and its watch found upon a heath as proof of a
divine watchmaker) multiplied examples of the perfection of the body as
proof of the existence of a Creator. Too often, enthusiasts for evolution
do the same as they hail every quirk among plants or animals as evidence
for selection's power. Sometimes they are right; but to assume that every­
thing must be adapted simply because it evolved is to practice theology
rather than science. The feats of evolution are such that blind faith in its
abilities is not needed.

On the Intercrossing of Individuals. Sexual selection-with all its ex­

pense-depends, of course, on the presence of two sexes in the first place.
Most animals fulfill their destiny with that curious strategy. The division
of reproductive labor has so many drawbacks that it is hard to see why it
has not been quashed. Pious Jewish men in the second century chanted
three times a day: "Praise be to God that He has not created me a
woman!" In fact the real question is why they (or anybody else) were cre­
ated as men. Why should any female bother with such creatures? They
seem almost useless. To give up males could at once double a female's
88 D A RW I N ' S G H O S T

output of daughters who would copy her DNA, and each child would
contain just her own genes, undiluted by those of a stranger.
Even so, when it comes to chastity (or at least to asexual reproduc­
tion), most plants and animals refuse to go the whole hog. Many try to
escape sex, but they usually fail. Some creatures indulge but once a year,
with long periods of abstinence as they make copies of themselves. Oth­
ers are hermaphrodites, while some persist for countless generations
without males at all. For most creatures, however, it seems that a cross
with another individual is occasionally-perhaps at very long intervals­
indispensable. The persistence of sex is one of the puzzles of biology. As
is the case for the tail of the peacock, the theory of sex is well ahead of
the facts needed to support it.
Many hermaphrodites-slugs, for instance-fall into what Woody
Allen called "sex with someone you really love": they mate with them­
selves. As a result, their progeny are liable to inherit two copies of the
same gene and are more alike than average. If the process goes on for long
enough, all the members of a line become, in effect, identical twins. On
the way, genes are exposed that are best kept hidden. Although the sur­
vivors are purged of inherited weakness they have lost their variety.
For British slugs, sex stops at Preston. North of there, the familiar large
slug of gardens and wild places retires from the sexual arena and takes in­
cest to its logical conclusion. The slug of Scotland and Scandinavia is in
effect a single strain of billions of identical animals. In southern parts, the
slugs of Welsh or Cornish mountaintops (and, oddly enough, of parts of
Cambridge) take the same reproductive route. The pattern of less sex in
cold places is found in many creatures. It hints at one reason why sex
might maintain itself: faced with the predictable enemies of frost and
starvation it is better to evolve a single set of hardy genes that are never
broken up by admixture with others. In the steamier parts of the world­
London, or the Amazon Basin-the adversaries come from biology
rather than the weather; they are other animals (parasites included).
These can themselves evolve, and a constant production of new genetic
combinations through sex is essential if their victims are to have a
chance.
Sex is not just an escape from the painful slowness of evolution, but
an ingenious way to make scapegoats who, by their own sacrifice, save
others from having to atone for their genetical sins. Several bad genes
 Natural Selection 89

may come together in one shuffie of the reproductive cards. All are dis­
posed of at the cost of a single bearer's death. In Levitical style, he carries
the faults of many with him as he goes. In genetics, as in life, sex and guilt
are close companions; and each is just as far from a convincing explana­
tion.

Circumstances Favorable to Natural Selection. What helps natural selec­

tion? A large amount of inheritable and diversified variability is favor­
able. A large number of individuals, by giving a better chance for the
appearance within any given period of profitable variations, will com­
pensate for a lesser amount of variability in each individual and is an ex­
tremely important element of success; and a large continental area,
which will exist for long periods in a broken condition, will be the most
favorable for the production of many new forms of life.
In other words, bacteria are bound to win their war against medicine.
Nowhere else does the evolutionary battle take place in an arena where,
in effect, one player holds all the cards. Bacteria show what natural se­
lection needs and what it can do when it gets the chance.
The Murray Collection is a series of reference strains of harmful bac­
teria gathered between 1 9 1 4 and 1 950 and kept in suspended animation
ever since. Every strain is, when reanimated with warmth and food, sus­
ceptible to every one of the dozens of antibiotics used today. They are a
reminder of what a revolution those drugs made. Before the Second
World War, wards were filled with patients close to a horrible death from
infections of the blood. After penicillin, they could be cured with a few
injections. Those glorious days will soon be over, because of evolution.
Antibiotics have become a human right. Penicillin was first used in the
1 940s. Its history is an object lesson in natural selection. The first resis­
tant strain was found within a year of its use and soon spread. Twenty
years ago, the drug could kill the bacterium that causes meningitis. In
many places-the United States and France included-three quarters
can now defy it. The more the drugs are used, the more resistance

spreads. In Norway, where antibiotics are controlled, only one strain in

five hundred of the septicemia bug is resistant to more than one drug,
while in Greece, where such remedies are available over the counter, half
the strains are.
Natural selection, with the help of stupidity, has triumphed over med-
90 D A RW I N ' S G H O S T

icine. Twelve million doses of antibiotic are given each year in the United
States to fight colds or sore throats, against which they do not work. In
the third world, things are even worse, and in Kenya, powerful drugs like
tetracycline and ampicillin are sold on the street. Farmers who add the
chemicals to animal feed pour yet more into the environment. Their
"growth promoters" include drugs that might be needed in medicine. To
put them into food could almost have been designed to speed evolution.
Again, Mrica leads the way. It is easier to add a powder than to clean up
a farm, and Kenyan chicken guts are filled with bacteria resistant to tetra­
cycline. Now, fruit trees are sprayed to cure their diseases, and salmon
farmers use drugs by the sack.
Why have the bugs done so well? First, there are a lot of them about.
Just a tenth of the cells of our bodies are human. Most of the rest belong
to bacteria (although a few fungi, mites and worms leaven the mix) .
When things are good, the inhabitants of British guts double in number
every twenty minutes or so, compared to the fifty years that it takes the
population of these islands, even in expansive times, to do the same.
With a world population of around ten with thirty zeros after it, bacte­
ria are so common that the most improbable events are, in effect, bound
to happen. Mutation is almost guaranteed and one individual among bil­
lions is certain to draw the successful ticket in the genetic lottery. That
explains why many of them rarely indulge in sex. For most bacteria there
is no need to exchange genes with another when, quite soon, the same
one will turn up in your own family.
When necessary, bacteria cheat to ensure a supply of new mutations.
DNA is supported by a mechanism that cuts down the number of mis­
takes each time it is copied. As soon as things get tough our enemies have
a clever stratagem. They circumvent their own repair machinery and, as
a result, increase the mutation rate in a last-ditch attempt to generate
variants that may save them. Some drugs spark off this emergency re­
sponse and help their targets to defend themselves.
Even so, the rate of change per gene is small, at about one in ten mil­
lion. All new variants are at first rare and most disappear, purely by acci­
dent, before selection notices them. There were, no doubt, hundreds of
moths with brand new genes for black wings in nineteenth-century
Manchester. Most of them failed, for reasons unrelated to pollution­
they starved, or drowned in a Mancunian downpour. The chances of suc-
 Natural Selection 91

cess for any new gene are smaller than they seem. The peppered moth
was common enough to cope with the accidental death of most of its
new melanics (although it took twenty years for the gene to succeed) . For
the billions of bacteria in a single decent-sized pustule, the chance loss of
a few mutations is less of a worry.
Geography, too, plays a part in the evolutionary equation. If every­
body lives in the same place and faces the same challenges, then selection
is absolute: a new mutation succeeds, or it fails and the whole population
dies. In a divided group, a few individuals may find a safe refuge in which
to await a favorable change.
New York in the 1980s suffered an outbreak of tuberculosis, concen­
trated among the poor. Black men had a rate of infection fifty times the
national average. A lengthy course of the right medicine can cure the dis­
ease. In New York, though, resistance became impossible to contain.
Most patients failed to complete their treatment: a single dose made
them feel better, there were side-effects, and the lives of many were in
such chaos that they could not manage a course of therapy. In Harlem,
only a few patients took more than a few pills before giving up.
As a result, drug resistance flourished in the poorer boroughs. It soon
spread to the affluent parts of the city, in which the disease had seemed
to be defeated. A reservoir of resistance, maintained by low doses of
drugs, was enough to overcome all efforts to get rid of the disease. Most
European cities are great continents of people, but those of the United
States are divided by social barriers, with success on one of their many is­
lands of humanity nullified by failure in another. In places in which the
population is treated as a whole, tuberculosis has been conquered.
The spread of bacterial genes is helped by sex as much as by politics.
Although some bugs rarely indulge, others enjoy it on demand, and in
many ingenious ways. For them, venereal disease evolved early on. In­
fectious third parties called plasmids, sections of mobile DNA inserted
into the genetic material, are (rather like AIDS viruses in human cells)
multiplied each time their hosts divide. Some can hop from host to host,
carrying resistance genes as they go. In time, a single plasmid may accu­
mulate many such genes and become invaluable to its carriers.
The Murray Collection had plenty of plasmids, but no resistance
genes at all . Their descendants are full of genes that enable them to cope
with several antibiotics at once. Many of the multiple-resistance plas-
92 D A RW I N ' S G H O S T

mids were first seen in hospitals, but are now everywhere. In Madagascar
a single strain of plague bacillus can resist ampicillin, chloramphenicol,
streptomycin, spectinomycin, kanamycin, tetracycline and sulfona­
mides. All seven resistances are carried on a single short length of mobile
DNA. With sixty million air travelers a year, such elements can move at
some speed. A resistant agent of pneumonia, first seen in Spain, was
found within five years in the United States, Korea and South Africa.
Even worse, plasmids can hop between species. Some found in harmless
denizens of guts have entered pathogens (such as the agent of gonorrhea)
and, at a stroke, rendered them safe from several drugs.
Bacteria show how natural selection builds its defenses. Evolution is
happy to pick up and use whatever is at hand. It presses new mutations
into service as they arise and is just as ready to make do with what is al­
ready around. Resistant bacteria may break down an antibiotic, block its
entry, pump it out, store it where it will do no harm, or change the shape
of its target molecule. Sometimes, complete new pieces of biochemical
equipment evolve, but more often workaday genes are pressed into ser­
vice. Tetracycline can be coped with in twenty different ways and peni­
cillin in almost as many. The most effective destroyers of drugs are
ordinary enzymes made in huge amounts by resistant strains. The bugs
pay a price, as their economy is so distorted by the need to fight off the
enemy that they become bacterial drug addicts, unable to survive with­
out the poison given to exterminate them.
The last new class of such drugs was discovered twenty years ago and
no more are on the horizon. Medicine's finest days may soon be over, but
antibiotics, in their brief flowering, have revealed as can nothing else
what evolution needs to do its finest work.

Extinction. The creative force of evolution has a dark side, for life today
was earned at the cost of the death of almost all that went before. The
idea of a past now gone alarms fundamentalists because it casts doubt on
the perfection of God's plan. Thomas Jefferson was so concerned that he
told the explorers Lewis and Clark to keep an eye open for mastodons as
they traveled through their new continent. Charles Lyell, the geologist
whose work formed Darwin's views, also denied the idea ofloss. Instead,
he envisaged a time when "the pterodactyl might flit again through um­
brageous groves of tree-ferns." Extinction, however, is a crucial part of
the evolutionary machine and is as inevitable as is the origin of species.
 Natural Selection 93

Because o f the conservation movement, such catastrophes have be­

come impossible to ignore. Public concern about the fate of the planet
suffers from overkill. Twenty years of lament about the imminent disas­
ter has led to a general view that things cannot be as bad as is painted.
Many who once cared about the environment now share a Voltairean
sentiment that the easiest way out of the crisis might be to strangle the
last panda with the guts of the last blue whale. However, today's cata­
clysm is no different from many others (and is far smaller than some of
the accidents that befell ancient life). Few plants and animals last for
long. The descendants of a very few, transformed by natural selection,
make up the world today. Some survivors manage to remain unchanged
for tens of millions of years, but for most, death soon follows birth.
Why do plants or animals meet their end? If they arise by the slow
process of natural selection, why should they not disappear for the same
reason? If each area is already fully stocked with inhabitants, it follows
that as each selected and favored form increases in number, so will the
less favored forms decrease and become rare. As better adapted successors
emerge, others are forced into a corner. They are driven back in the great
battle for life until, at last, they are gone. Like the American Motors Cor­
poration, they have been driven to ruin by their better-adapted succes­
sors.
The evidence is in the fossil record. Although there have been some
hesitations in the creative process, with more new forms arising at cer­
tain times, the big picture is consistent. Over the past five hundred mil­
lion years, through all its ecological alarms and excursions, new kinds
appeared at an almost constant rate. A survey of tens of thousands of ma­
rine animals over that time gives a rate of four hundred and fifty new
species a year. The world is more or less full. Any new species must push
out a predecessor to have a chance and will, sooner or later, be squeezed
out in its turn.
Sometimes, a few survivors of an earlier age remain as relics of an an­
cient race. Lungfish first appeared about four hundred million years ago.
They were an active and diverse group that adapted themselves to what­
ever they were faced with. A cadet branch of the family played an im­
portant part in the transition of vertebrates from water to land. Then, all
of a sudden, lungfish evolution slowed down, while around them that of
other vertebrates exploded. Now a mere half dozen kinds are left. They
live a dreary existence in the lakes and rivers of Mrica, South America
94 D A RW I N ' S G H O S T

and Australia, shrouded in mud for much of the year. They are living fos­
sils, reminders of a universe now lost.
Any form represented by few individuals will, during fluctuations in
the seasons or in the number of its enemies, run a good chance of utter
extinction. As new species in the course of time are formed through nat­
ural selection, others will become rarer. Then, bad luck begins to play a
part. Although time and the gradual appearance of new and better forms
kills most of them, chance is also important. Bighorn sheep in the Rock­
ies have been studied for almost a century; and in that time all groups of
fifty or fewer animals became extinct, while nearly all those with more
than a hundred survived. When gambling with Nature, it pays to have a
strong hand.
It is hard to be sure about extinction. Nobody, after all, writes to the
newspapers about the last cuckoo of spring. The youngest fossil of the
coelacanth, a fish at one time thought to be important in the origin of
land animals, is eighty million years old, and it was once presumed to be
gone forever. An example of the remarkable beast was caught off the
coast of South Africa in 1 938. For a time coelacanths seemed to have a
population of a few hundred and to teeter on the edge of demise, but,
sixty years on, a specimen on sale in an Indonesian market led to the dis­
covery of many more, thousands of miles away off the coast of Sulawesi.
Coelacanths may be common all around the Indian Ocean.
The same uncertainty applies in other places. About half a million
kinds of beetle are known but, because most were found by gassing trop­
ical trees, many are recorded from just a single location. An absence on a
second visit may not mean that they have gone, but that they are on an­
other tree. Species thought extinct quite often reappear. In California,
where there has been much concern about the loss of Mediterranean
plants, more kinds supposed to have been driven out have been redis­
covered than have in fact disappeared. Even so, when it comes to the de­
struction of what evolution has made, we live in interesting times. They
prove how fast selection can carry out its baleful work as soon as it gets
the chance. All over the world, life has been swept away, as if by some
murderous pestilence. That pestilence is man and his hangers-on.
At any moment, a hundred thousand people are suspended over the
Atlantic. Some smuggle alien plants and animals, but many more have
seeds, insects and more in their cuffs or their baggage. Other creatures
 Natural Selection 95

travel in soil or packing crates, or are introduced for food o r ornament.

The United States Customs Service intercepts three thousand species of
potential pest each year, but many get through. Some make a home in
their new world and drive out the natives. The worst culprits are those­
like men or rats-able to eat many things. If they choose to graze on the
eggs of some rare bird, they can drive it to extinction while they sustain
themselves on other food. On Stephens Island in the Cook Strait, in
1 894, the lighthouse keeper's cat brought in the first known specimen of
the Stephens Island wren. One of its descendants ate the last one.
The Galapagos were uninhabited when first seen by Europeans in
1 535. Half a million fossil bones of reptiles and birds have been found
in the lava tubes scattered across the landscape. In the eight thousand
years before the Spaniards arrived, each island lost, at most, three of its
native venebrates. In the four centuries since the onslaught began, the
rate of loss has gone up by a hundred times. Cats, pigs and ants have
done huge damage. Five hundred different foreign plants are on the is­
lands. The guava is a pest, as is a vine called the Curse of India. Black­
berries, too, are on the march. As they grow they shade out the botanical
equivalents of Darwin's finches. The cacti are unique to the archipelago,
but many have been strangled by vines. The giant tortoises that feed on
them are damaged in their turn. The Floreana flax, evolved on a single
island, now consists of a mere eight plants in the wild and three more in
cultivation. With a growth rate in the human population of 1 0 percent
a year the future is bleak.
The emergency goes further than the Galapagos. Half of all kinds of
bird in the world may disappear within the next three hundred years, a
rate of loss thousands of times that in most of the fossil record. About
one in twenty of all the people who have ever existed are alive today,
compared to a mere one in a thousand of the different kinds of animal
and plant. The fate of the defunct is as much testimony to the force of
natural selection as is the triumph of the survivors. Biologists often be­
moan the ecological crisis-the loss of the cod, the whale or the dodo­
as something outside the normal world of evolution. It is not: extinction
has happened millions of times before and is an exciting opportunity for
science. The years since Darwin have relived an evolutionary experiment
that shows how the more adapted will always drive out those less able to
cope.
96 D A RW I N ' S G H O S T

Divergence ofCharacter. Because all animals must compete with their rel­
atives, evolution favors things that differ from one another. The more it
can do so, the less each of its products is forced to depend on an asset in

short supply. The more diversified the descendants from any one species
become in structure, constitution and habits, by so much will they be
better enabled to seize on many and widely diversified places in the polity
of nature.
Canadian lakes are full of sticklebacks. They are recent arrivals, for the
whole country was covered by ice until a few thousand years ago. Most
sticklebacks live in the sea or in estuaries, and the fish have invaded fresh
water many times. They come in two forms, sometimes found in the
same lake. The first has a stocky body and lives in the shallows, where it
eats grubs, while the other grows longer and has a smaller mouth suited
to open water and a diet of surface swimmers. The two live in different
patches of an ecological quilt. Each is better off in its own habitat, and
each originated not long ago, each within its own lake system. Fish of dif­
ferent shape are already reluctant to interbreed. Each is, if not yet a sep­
arate species, well on the way to an identity of its own.
Small lakes contain but one kind of fish, of intermediate form. When
put into competition with the specialists, they fail. As soon as the chance
arises the sticklebacks split into two types with their own peculiar habits
and structure and exploit more of what the lake can offer. Canadian
salmon, too, come in distinct versions, one-the sockeye-migrating to
the sea before spawning, the other-the smaller kokanee-staying in the
lake for its whole life. Sockeyes moved to empty lakes quickly evolve a
new and reduced version that stays at home. This, too, is evidence of the
pressure to divide.
Natural selection has, built in, what may be called the principle of di­
vergence, causing differences, at first barely appreciable, steadily to in­
crease, and breeds to diverge in character both from each other and from
their common parent. The process is captured as a snapshot when, as
sometimes happens, variation that circulates through a population is par­
titioned into a series of distinct clones, each of which contains part of the
diversity of their sexual parent.
A certain New Zealand freshwater snail exists in both sexual and asex­
ual forms. Within a lake, the sexuals are accompanied by dozens of asex­
ual clones. The sexual form, in all its diversity, is found in most places.
 Natural Selection 97

Each clone, in contrast, lives at a certain depth and o n certain plants. It

is frozen into a separate place in the economy, unable to invade the
equally cramped niche next door. Its narrow life is testament to the fact
that competition will generally be most severe between those forms
which are most nearly related to each other. The snails show how finely
divided the evolutionary cake may be.
The snail was introduced into Europe in the 1 880s, on a load of ship's
ballast dumped into the Thames, close to where the dismal suburb of
Thamesmead now stands. It spread to fill most of the lakes and rivers of
Europe. It lacks any form of sex and has just the three clones first intro­
duced, rather than the hundreds found in its native land. They divide up
their waters quite amicably, and each occupies a wide range of habitats,
from ponds to estuaries. Without an infinity of competitors they can af­
ford to spread out and are not crowded into a smaller and smaller space
by their kin. Like any business, life must diversify its manufactures, or
fail. Evolution-like capitalism-must run to stay in the same place. If
the young overtake their parents, the parents have no choice but to find
another trade, or die.

That brutal fact launched the Industrial Revolution and drives the
economies of today. Commerce depends, like life itself, on a constant
input of energy. In modern London or Manchester, it flows not from the
coal-fired stations that poisoned the Victorians and their moths, but
from boilers fueled by uranium, gas or oil. Evolution is not mocked. Py­
lons carry the electricity across the land. Each is protected by a layer of
zinc, which drips onto the ground when it rains. Zinc, like copper or
lead, is poisonous to plants, but some have genes able to deal with it.
Natural selection has come up with the same response again and again:
under most pylons is a patch of grass with genes for zinc tolerance. Some
populations lack the right genes and cannot grow, but in most places
they cope with ease. In the archipelago of metal that fills all cities an evo­
lutionary experiment has been repeated thousands of times.
Zinc faces natural selection with nothing new. Man has long spread
his poisons. Four thousand years ago the production of lead reached a
peak not matched again until the Industrial Revolution. The records of
ice and peat show that in 1 979, the height of its use in gasoline, the air
98 D A RW I N ' S G H O S T

had fifteen hundred times the background level. .As a result, every road is
lined by a swath of lead-tolerant vegetation. Many soils (such as the ser­
pentine rocks of Jasper Ridge in California) are in any case awash with
metal. Dozens of plants have evolved to cope, some able to deal with
amounts sixty times more than those lethal for others. For selection, a
pylon is a minor provocation.
Cities, like pylons, do not last. The first was founded but ten thousand
years ago, and many have come and gone since then. Sometimes, the sole
evidence of their passing lies in evolution. Fourteenth-century Africa had
a culture based on copper, mined from the deposits-still the largest in
the world-of what is now the Congo and Zambia. Around today's
mines and smelters the soil is so full of metal that only plants with genes
for tolerance can survive. One, the copper flower, grows in dense violet
dumps on the most polluted soils of all.
Patches of that plant are found far from any habitation. They are the
tombstones of lost villages, the remnants of a forgotten Industrial Revo­
lution. Hundreds of copper crosses, used as money by the miners, are
buried beneath the violet blooms. The genes of the copper flower are
monuments to those who made the coins: all else has disappeared. They
are a reminder of how fleeting are the wishes and efforts of man, how
short his time, and in consequence how poor his products, compared
with those accumulated by nature during whole geological periods.
Natural selection is no more than a machine. What it makes depends
on what it has to work with and where it started. Evolution does its job
as well as it needs to, and no more. Sometimes, as in the balance of the
numbers of each sex, it does it well; but often it is satisfied with what
seems slapdash. Most of its products do not last. Who could ever have
designed a tree kangaroo? Clumsy as the animal may seem, it is infinitely
better adapted to the most complex conditions of life, and plainly bears
the stamp of far higher workmanship, than anything achieved by man.
Industry has begun to notice the superiority of nature. Nowadays, a
billion dollars' worth of copper a year is extracted not with furnaces, but
with a bacterium able to break down ore and to release the metal. The
bug obtains its energy by chemical means and may drive smelters to ex­
tinction. In Africa the metal accumulated by tolerant crops can be a har­
vest twice as valuable as wheat. It is even possible to sow plants on gold
mines to reap their treasure.
Man, in his factories for copper or soap, has begun to use the instru-
 Natural Selection 99

ment that shapes biology, but has rately matched the work of nature. In
spite of Otto Lilienthal's great work Bird Flight as the Basis for Aviation
and the long-and lethal-series of birdlike gliders and flappers that fol­
lowed, to day's airplanes do not have feathers. A few useful ideas have
been lifted: the inventor of the tunneling shield based it on the shipworm
that chews wood and passes the waste through its body, while the 1 874
patent for barbed wire stated that the invention was designed to look like
a thorn hedge. Like thorns themselves, the new product diversified into
(among many others) Griswold's Savage, Blake's Body Grip and Brink's
Stinger. The patentee almost lost his millions because rivals claimed that
he had invented nothing, but merely copied the living world.
Nature, though, starts in a different place and uses materials quite un­
like those available to man. It cannot smelt copper or make crosses, but,
with what it has, it works miracles. Man, his machines and Darwin's idea
may-given a few million yeats-clo almost as well.

Summary of Chapter. If during the long course of ages and under varying con­
ditions of life, organic beings vary at all in the several parts of their or­
ganisation, and I think this cannot be disputed; if there be, owing to the
high geometrical powers of increase of each species, at some age, sea­
son, or year, a severe struggle for life, and this certainly cannot be dis­
puted; then, considering the infinite complexity of the relations of all
organic beings to each other and to their conditions of existence, caus­
ing an infinite diversity in structure, constitution, and habits, to be ad­
vantageous to them, I think it would be a most extraordinary fact if no
variation ever had occurred useful to each being's own welfare, in the
same way as so many vatiations have occurred useful to man. But if vari­
ations useful to any organic being do occur, assuredly individuals thus
characterised will have the best chance of being preserved in the strug­
gle for life; and from the strong principle of inheritance they will tend to
produce offspring similarly characterised. This principle of preserva­
tion, I have called, for the sake of brevity, Natural Selection. Natural se­
lection, on the principle of qualities being inherited at corresponding
ages, can modify the egg, seed, or young, as easily as the adult. Amongst
many animals, sexual selection will give its aid to ordinary selection, by
assuring to the most vigorous and best adapted males the greatest num­
ber of offspring. Sexual selection will also give characters useful to the
males alone, in their struggles with other males .
Whether natural selection has really thus acted in nature, in modify-
1 00 D A RW I N ' S G H O S T

ing and adapting the various forms of life to their several conditions and
stations, must be judged of by the general tenour and balance of evi­
dence given in the following chapters. But we already see how it entails
extinction; and how largely extinction has acted in the world's history,
geology plainly declares. Natural selection, also, leads to divergence of
character; for more living beings can be supported on the same area the
more they diverge in structure, habits, and constitution, of which we see
proof by looking at the inhabitants of any small spot or at naturalised pro­
ductions. Therefore during the modification of the descendants of any
one species, and during the incessant struggle of all species to increase
in numbers, the more diversified these descendants become, the better
will be their chance of succeeding in the battle of life. Thus the small dif­
ferences distinguishing varieties of the same species, will steadily tend
to increase till they come to equal the greater differences between
species of the same genus, or even of distinct genera.
We have seen that it is the common, the widely-diffused, and widely­
ranging species, belonging to the larger genera, which vary most; and
these will tend to transmit to their modified offspring that superiority
which now makes them dominant in their own countries. Natural selec­
tion, as has just been remarked, leads to divergence of character and to
much extinction of the less improved and intermediate forms of life. On
these principles, I believe, the nature of the affinities of all organic be­
ings may be explained. It is a truly wonderful fact-the wonder of which
we are apt to overlook from familiarity-that all animals and all plants
throughout all time and space should be related to each other in group
subordinate to group, in the manner which we everywhere behold­
namely, varieties of the same species most closely related together,
species of the same genus less closely and unequally related together,
forming sections and sub-genera, species of distinct genera much less
closely related, and genera related in different degrees, forming sub­
families, families, orders, sub-classes, and classes. The several subordi­
nate groups in any class cannot be ranked in a single file, but seem rather
to be clustered round points, and these round other points, and so on in
almost endless cycles. On the view that each species has been indepen­
dently created, I can see no explanation of this great fact in the classifi­
cation of all organic beings; but, to the best of my judgment, it is
explained through inheritance and the complex action of natural selec­
tion, entailing extinction and divergence of character.
The affinities of all the beings of the same class have sometimes been
represented by a great tree. I believe this simile largely speaks the truth.
 Natural Selection 101

The green and budding twigs may represent existing species; and those
produced during each former year may represent the long succession of
extinct species . At each period of growth all the growing twigs have tried
to branch out on all sides, and to overtop and kill the surrounding twigs
and branches, in the same manner as species and groups of species have
tried to overmaster other species in the great battle for life. The limbs di­
vided into great branches, and these into lesser and lesser branches,
were themselves once, when the tree was small, budding twigs; and this
connexion of the former and present buds by ramifying branches may
well represent the classification of all extinct and living species in
groups subordinate to groups. Of the many twigs which flourished when
the tree was a mere bush, only two or three, now grown into great
branches, yet survive and bear all the other branches; so with the species
which lived during long-past geological periods, very few now have liv­
ing and modified descendants. From the first growth of the tree, many a
limb and branch has decayed and dropped off; and these lost branches
of various sizes may represent those whole orders, families, and genera
which have now no living representatives, and which are known to us
only from having been found in a fossil state. As we here and there see a
thin straggl ing branch springing from a fork low down in a tree, and
which by some chance has been favored and is still alive on its summit,
so we occasionally see an animal like the Ornithorhynchus or Lepi­
dosiren, which in some small degree connects by its affinities two large
branches of life, and which has apparently been saved from fatal com­
petition by having inhabited a protected station. As buds give rise by
growth to fresh buds, and these, if vigorous, branch out and overtop on
all sides many a feebler branch, so by generation I believe it has been
with the great Tree of Life, which fills with its dead and broken branches
the crust of the earth, and covers the surface with its ever branching and
beautiful ramifications.
 C H A P T E R V

LAWS O F VARIATION

Heredity, its myths and errors - Effects o f use and disuse; heredity as mem­
ory - The inheritance of privilege - Acclimatization, to heat and to poi­
sons - Atavism and the failure of the average - Genes as particles rather
than fluids - Mendel and the physical basis of inheritance - Mutation
and the rate of evolution - Correlation of growth; genes and develop­
ment - The simplicity of Mendel's laws and the complexity of the real
world - Genetics the foundation of the theory of evolution - Summary

George Spencer had the misfortune to live in Connecticut at the height

of the Puritan obsession with sex. In a childhood accident he had lost an
eye. When a one-eyed pig was born in the town, the culprit seemed ob­
vious and Spencer was accused of bestiality. Terrified (for this was a cap­
ital offense) he first admitted the crime, but then withdrew. Two
witnesses were, the law said, needed. So anxious were the magistrates to
hang him that his confession was accepted as the first and the pig as the
second.
Sex and confusion have long been bedfellows. For ten thousand years
of success for plant and animal breeders, its practice was quite divorced
from its theory. The laws of inheritance were quite unknown, and no­
tions now seen as absurd-the transfer of eyes from man to pig in­
cluded-were believed by everyone. Many of those ideas were
misguided, but some contain enough truth to explain (and perhaps to
excuse) the morass in which biology wallowed for so long.
It is easy to multiply (and to mock) curious beliefs about inheritance.
 Laws o/ Variation 1 03

His ignorance of the subject worried Darwin and led him, in his later
years, to complicate and confuse his ideas. Now genetics has become the
science that catches the collective imagination as does no other. To the
public it seems beautifully simple, but it is not.
Its laws are as elegant as is the idea of natural selection. Mendel started
a science that still rests upon his simple rules. Like evolution itself, it has
become more complicated since it began, and Darwin's perplexity makes
more sense today than it did fifty years ago. Many of his difficulties now
look like an honest attempt to find simple patterns in complicated situ­
ations. This chapter is less faithful to its original than are others, if only
because Darwin got it so wrong. Even so, with hindsight, The Origin
points at problems about heredity that are still scarcely understood.

Effects of Use and Disuse. The notion of the inheritance of acquired char­
acters does away with any need for a theory of evolution. In melanic
moths the camouflaged young were once said to result from a "powerful
impression on females during the all important period of life, viz., that
of propagation, coupled with an instinctive provision for the protection
of its future progeny." Humans followed the same rules, and the noble
Madeleine d'Auvermont assured her son's succession by her claim that
she had become pregnant when her husband was away, just by thinking
about him.
Before Mendel, all heredity was (as the French aristocracy noticed)
memory. The idea seemed to make perfect sense. The ostrich is exposed
to danger from which it cannot escape by flight, but by kicking it can de­
fend itself from enemies as well as any of the smaller quadrupeds. We
may imagine that the early progenitor of the ostrich had habits like those
of a bustard, and that as natural selection increased in successive genera­
tions the size and weight of its body, its legs were used more, and its
wings less, until they became incapable of flight. Such a notion is easy to
contemplate: but it is wrong.
Nature has plenty of instances of use and disuse. Blacksmiths have
thicker arms than bank clerks, but migratory birds put both of them in
the shade. Some birds double in size before their journeys (and, unlike
any human, increase the volume of their testicles by a hundred times in
spring) . Such characters are not themselves passed to the next genera­
tion. The young are heirs to an ability to grow large organs, rather than
1 04 D A RW I N ' S G H O S T

to the structures themselves. Fat parents have fat children, in the main,
not because stoutness is in the genes, but because they feed their off­
spring with a diet like their own. Fat people have fat cats, too, but no­
body blames that on DNA.
A moment's thought shows that the inheritance of acquired characters
must be common. Parents and their young nearly always share environ­
ments as much as they do genes. If identical plants are grown in pots of
soil from a few inches apart in a forest, the contrasts in habitat can cause
large differences in size and shape. As seeds fall close to their parent, the
fate of a young plant depends both on its heritage and on where it grew.
Although such effects last only a few generations, biologists are nowadays
wary about giving exclusive authority to genes. The conflict between
nurture and nature has lost much of its meaning. The attributes of most
interest to evolution-size, shape, or behavior-are influenced by both.
Those who first asked big questions about how species originate failed,
in part, because much of the answer lay in a smaller question about in­
heritance within each one.
The idea that a character acquired in an animal's lifetime can be
handed on was once anathema, dismissed with a story about Jews and
foreskins in the first lecture of every genetics course. It is now common­
place, but the notion is a detail on the edifice of genetics and not its foun­
dation.
All children get more than genes from their parents. The songs of
some birds are passed down the generations by education. Macaque so­
ciety is based on rank. Every female monkey knows her place (although,
now and again, she makes a furtive challenge in the hope of promotion) .
Her position comes from her mother, who helps her daughters to lord it
over those lower in the pecking order. Any female from a noble line, fee­
ble though she is, ranks above anyone from a lower stratum of society.
Status does not travel in the genes, but in the mind. The hierarchy set by
custom can last for years-and has a physical as well as a mental effect,
as low-grade animals have more heart disease than those higher in the so­
cial scale. All this might seem a slight exception to the great Mendelian
truth, but is a small part of the great range of characters whose inheri­
tance is in some way acquired.
Nutrients are passed to the next generation in egg or seed, diseases
strike before birth, and a mother's diet affects her young. Chemicals, as
 Laws o/ Variation 1 05

well as culture, are passed between generations (as doctors who deal with
babies born addicted to heroin know). To give a newborn female mouse
a hefty dose of thyroid hormone depresses her ability to make the right
amount of that crucial substance-and the effect is transmitted to her
offspring, who grow within her damaged body.
To breed from large or small mice can make one line twice the size of
another through artificial selection of the most conventional kind. How­
ever, a mouse's size also depends on its food. An animal with a good diet
will be larger than its twin on starvation rations. As a result, for young
animals, what matters most is how well fed their mother might be. Her
environment influences their fate. As might be expected, in inbred mice,
large and well-fed mothers have more young than their identical but
small and hungry sisters. However, the young of large mothers tend to
be small, because they grow up in a crowded womb and must struggle
for what milk is available. Because they are so tiny their own few off­
spring grow up uncrowded, well fed-and large. However good a
mouse's genes, the environment is enough to defeat them. The antago­
nism between nature and nurture controls their fate. Such complexity
baffled those who tried to work out the laws of genetics from the experi­
ence of animal breeders.
Plants are much the same. When flax plants are given fertilizer they
grow faster and have more branches with larger leaves. Simple enough;
but when the offspring of such plants are grown alongside others whose
parents were less fortunate, they too are more branched and leafy. The ef­
fect persists for several generations. The reason is straightforward. When
a plant is well fed, certain genes multiply and help it deal with the extra
food. Some of the copies are passed into the egg and reach the next gen­
eration, who benefit from their parents' happy lives.
Less fortunate parents can protect their offspring. Water fleas grow
thick helmets when exposed to the scent of their enemies, so that those
who escape when under fire the first time are ready for the next barrage.
A mother who has donned her armor equips her young (who are likely
to be born into her own perilous circumstances) to be hypersensitive to
risk, so that they protect themselves by growing a helmet at the merest
hint of danger. They grow up ready for an assault by the beast that ate
the previous generation.
Genes, too, have a memory of who transmitted them. It can make a
1 06 D A RW I N ' S G H O S T

difference whether they are passed on by a male or a female. Any father

is anxious to persuade his partner to invest as much as possible in his chil­
dren, while her own priority is to minimize what she provides in the
hope of more children later, perhaps with a different mate. One mouse
gene alters a mother's ability to care for her young. However, it exerts its
effects only through the copy inherited from the father: any new mother
turns off the version inherited from her own female parent. Males are, in
some way, controlling the effects of genes passed to the next generation;
but that control is lost when it passes through a female. All this and more
does away with the idea that the genes are sacred, safe from the insidious
effects of the outside world.

Acclimatization. Heat up a fruit fly and it may die; but if allowed to re­
cover and heated again it will cope better with the second shock. Plants,
too, can resist heat or cold if they are warmed or cooled before the main
challenge. They adapt to their new conditions with special proteins that
fight stress. These are switched on when danger threatens, to be ready
when it appears in its full ferocity.
Not all defenses need be kept at full alert at every moinent. Instead,
natural selection keeps much of its armor in reserve: with its troops stood
down until needed. A tan is no use at Christmas, but it does not take
much sunshine when spring comes to prepare the body for the next hot
day. In the same way, a sunbather who downs a stiff drink after a long
sober winter switches on a set of enzymes able to deal with it; which
means that, in time, more and more alcohol is needed to put him into
the right mood. The first drag at a cigarette does the same-and the to­
bacco hornworm, one of the few insects able to eat the plant, recoils in
horror at its first juvenile taste of the bitter leaves. Not until its anti­
nicotine enzymes have been activated can it settle down to its natural
diet. Tobacco itself, in retaliation, turns on a whole set of poisons after a
leaf has been damaged, to warn off later browsers.
None of this is much use when the stress is not there and the ability to
respond when called for is as honed by natural selection as is the response
itself Anyone interested in, say, the inheritance of dark skin, and un­
aware of the role of sunlight, would find it hard to sort out why some
people are brown and some not. Acclimatization even has an effect on
body structure. Flies given a sudden burst of high temperature as pupae
 Laws o/ Variation 1 07

have many deformities as adults. The errors are a side-effect. The pro­
teins that rush to the aid of the heat-stressed cell and ready it for another
bout have a second job; to insulate the body against the effects of genetic
damage. They act as a scaffold during development and contain minor
flaws to ensure that a perfect fly emerges. At times of danger, the need to
acclimatize takes their support away and the body reveals its inborn
weaknesses.

Distinct species present analogous variations; and a variety of one species

often assumes some ofthe characters ofan allied species, or reverts to some of
the characteristics ofan early progenitor. Most children are the average of
their parents, and a fat pig mated with a thin one tends to have young of
intermediate size. There are exceptions to this otherwise persuasive ob­
servation. It was a very surprising fact that characters should reappear
after having been lost for many, perhaps for hundreds of generations. But
when a breed has been crossed only once by some other one, the off­
spring occasionally show a tendency to revert in character to the foreign
breed. That single ugly fact, the reappearance of a character in a lineage
after it had seemed lost, was the foundation of the scientific study of in­
heritance.
Genetics is to biology what atomic theory is to physics. Its principle is
clear: that life is based on particles and not on fluids. Instead of the
essence of each parent mixing, with each child the blend of those who
made him, information is passed on as a series of units. The bodies of
successive generations transport them through time, so that a long-lost
character may emerge in a distant descendant. The genes themselves may
be older than the species that bear them.
Mendel was both lucky and a genius. His luck was favored by a pre­
pared mind: by his decision to study simple characters in a simple or­
ganism. His genius was to separate the products of inheritance from the
mechanism of heredity. He chose to work on peas, which have an odd
but useful sexual system. Like many garden plants, they exist as a series
of inbred families (or "pure lines"). These have been kept separate for so
long that, within a line, every plant is identical, but among lines the
plants are distinct. The lines diverge in many traits-flower color, plant
height, pea shape and the color of the pea included. What is more, peas
108 D A RW I N ' S G H O S T

are at once male and female, and (unlike many of that ilk) can fertilize
themselves.
His experiments now seem simple, but nobody, in ten thousand years
of agriculture, had tried them. Mendel took male sex cells-pollen­
from one pure line and used them to fertilize the eggs of another. He
looked at the various pairs of traits used to distinguish each inbred fam­
ily. For example, pea color in different lines was either yellow or green.
A plant from a yellow pure line crossed with another in which all peas
were green gave only offspring with yellow peas. That was itself remark­
able. It at once disproved the notion of blending, because all the seeds
looked like those of one parent and were not the average of the two. To
cross those progeny among themselves gave another useful result. Green
and yellow peas each appeared in the next generation, and whatever had
made the plants green was restored after it had lain hidden in a plant
whose own seeds were yellow. The agent of inheritance-the gene-had,
it seemed, an existence separate from that of its vehicle, the plant.
Mendel's ratios were always (given the accidents of sampling) the
same. In this second generation, there were three yellow peas to one
green. From this, Mendel deduced that his units came in two forms, or
"alleles. " Body cells contain a pair of alleles for each character, while each
pollen or egg cell receives just a single one. They combine in different
ways: two yellows, a yellow and a green, or two greens. One, the domi­
nant allele, can conceal the presence of its recessive partner. A yellow pea
can be made with two yellow alleles, or a yellow allele and a recessive
green. A recessive allele must be present in double copy to show itself,
and all green peas have two green alleles.
The yellow peas in Mendel's first generation descended from parents
of different color. Each has a single copy of the yellow allele, matched
with a single copy of the green. When those hybrid plants were inter­
crossed, simple arithmetic gives a ratio of a quarter with two yellow al­
leles, a quarter with two greens, and a half with one of each allele; to give
a proportion of three yellow to one green plant. Inheritance, he thought,
was explained.
Mendel's logic applied to every character, from flower color to plant
height. Even better, the pattern for each was independent of those of the
others. It made no difference to the three-to-one rule for yellow and
green peas if the two stocks also differed in flower color. Genetics seemed
 Laws o/ Variation 1 09

simple. It was based on independent particles, each coding for a single

attribute and each following its own path down the generations. The idea
explained how a character long lost could reappear: it had been hidden
by a matched copy of the dominant allele. A rare recessive allele almost
always suffers such a fate, as only in those unusual cases in which two
parents each carry a hidden copy is there a chance of its appearing in
their progeny.
Mendel saw the value of his work to the theory of evolution (as is
manifest in the marginal notes to his own copy of The Origin) . Although
his research was known to many biologists of his day it was, alas, ignored
as-at best-a discovery of interest only to those concerned with peas.
Later, Mendel went on to study hawkweeds, but was baffied by the
plant's (then unknown) ability to reproduce without sex. He could get
no results as elegant as those with peas. Discouraged, he gave up and be­
came an administrator.
Once, Darwin almost got it right. He noticed that the young from a
cross between two different stocks of pigeons were uniform, but that
when these mongrels were crossed for several generations then hardly
two were alike. Mendelism is, we now know, at work, as the stocks dif­
fer in several genes that later come together in many ways. For once, Dar­
win's insight failed him.
The importance of Mendel's work was at last realized in 1 900. Crosses
were soon made on a variety of plants and animals. In most cases they
followed his laws. Inheritance was, it appeared, quite simple. Of course,
it was not, and (as often happens in biology) the more that is known the
more there seems to be left to find out. Genetics still has its basis in
Mendelism, but on that firm foundation has grown a complex and often
enigmatic structure.

Sometimes, his famous ratios can shift. A cross between two Manx cats
gives not a three-to-one proportion in the next generation, but two ani­
mals without tails to each one tailed. The ratio emerges because a dou­
ble dose of the Manx allele (unlike a double dose of the allele for yellow
pea color) kills the one in four kittens unlucky enough to receive it.
Other odd patterns appear as different genes band together-as they
must-to build a living creature. Some abolish the effects of a whole
1 10 D A RW I N ' S G H O S T

string of others. A white cat, like an albino human or white whale, has a
biochemical quirk. It masks the presence of genes that would, given the
chance, make a patterned coat or a dark eye. In other cases, genes coop­
erate rather than compete. Persian cats have their elegant coats because
of a variant that reduces the intensity of the coat pigment laid down by
quite a different gene. When it comes to attributes such as size, shape or
behavior, many genes of large or small effect are involved.
Quite soon after the rediscovery of Mendel's work, there emerged an
important exception to his laws. Some characters are, it appeared, not in­
dependent. Instead, certain combinations tend to pass down the gener­
ations together. The link between the fellow travelers is sometimes
broken, but there remains an incomplete association between them. All
genes fall into one of several groups that are passed on in consort. Only
members of different groups follow Mendel's rule.
The discovery was the key to the physical apparatus of inheritance. In
fruit flies four such "linkage groups" emerged from crosses. They match
the four pairs of chromosomes, dense bodies in the cell nucleus. These
were the tangible signs of Mendel's magical particles. If two genes were
on the same chromosome they could be passed on together. If they were
not, they were inherited as autonomous units. The association was not
absolute because the chromosomes themselves break up and rejoin in
new combinations each time sperm or egg is made. It did not take long
to realize that the more a pair of genes deviated from independence, the
closer together they must be. The first genetic map was made by com­
paring the tendency of such pairs to stick together as the generations suc­
ceeded.
Chromosomes are complicated things, made up of hundreds of mol­
ecules. One element, DNA, seemed an unlikely candidate as the vehicle
of inheritance. It had a mere four different units (or "bases") and had
been dismissed as "the stupid molecule" as a result. A bold experiment in
which DNA was transferred between bacteria with colonies of different
shape showed it to be the crucial agent. The story of the famous double
helix and of the code for the structure of proteins is part of the cultural
inheritance of the twentieth century.
Now, all kinds of marvelous technologies are used to read the message
of the nucleic acids. The structure of any protein can be deduced from
the DNA sequence responsible, and this may hint at what each gene
 Laws of Variation 111

does. Already, many viruses and bacteria, together with yeast, have had
their genetic message laid bare. Worms and fruit flies, too, have had all
their units read off. The human map itself, three million DNA bases
long, is almost complete, less than fifty years after the discovery of the
double helix.
For most things more complex than bacteria, maps of the DNA do
not much resemble a chart based on crosses. Instead, the genes are full of
waste and redundancy. Some are interrupted by strings of material that
appear to code for nothing. All this is, perhaps, less remarkable to those
who do not come (as did many of the pioneers of molecular genetics)
from physics, but from biology. Anyone used to the muddle and waste
of evolution, its products cobbled together over long ages of expedience,
is not surprised to see in the genes themselves the same history of uneasy
compromise. The genome is as complicated, makeshift and imperfect as
the creatures it builds.

Genetics is the science of difference. Variety is the raw material of evolu­

tion, used up as natural selection takes its course. Once it has been con­
sumed, the Darwinian machine comes to a stop. Diversity is renewed by
chemical errors-mutations-made as DNA is copied. Geneticists were
once so impressed by mutation as to suggest that new forms of life arise
not through the accumulation of small changes but in great leaps. Evo­
lution was due to the instability of genes and genetics had, perhaps, de­
stroyed Darwin's idea.
It had not: mutation is the fuel rather than the engine of biological ad­
vance. The process involves mechanisms undreamed of in the science's
first days.
Some mutations are simple. They are no more than a change in the ge­
netic alphabet that alters the properties of a gene, or stops it altogether.
A single shift can persuade a growing protein to stop dead and, by so
doing, kill or much modify those who inherit it. Other mutations arise
from a sudden duplication or deletion of genetic material or of whole
chromosomes.
Mutation could once be studied only in bacteria or in fruit flies. Now,
those in humans are as accessible as those in any other species. The bones
of the Russian Royal Family, killed in 1 9 1 7, were identified through the
1 12 D A RW I N ' S G H O S T

fit of their mitochondrial genes to those of today's aristocrats. Embar­

rassingly enough, the tsar's own DNA did not quite match that of his
presumed relatives, because he carried two distinct types. One must have
arisen in his own lifetime. The United States Army records the genes of
its soldiers, and sometimes has cause to compare them with those of their
relatives. Again change is rapid, with a mitochondrial mutation in every
forty parent-child comparisons.
Much of the genetic damage lies at the feet of age and sex. Men are de­
fined by their ownership of a single gene. It has the modest task of per­
suading the early embryo to divide faster than it might. The early
stimulus sets it off down the path of masculinity. The gene sits on a cer­
tain chromosome, the Y, which has few other jobs to do. It is so small
that it does not mask genes carried on its partner, the X. Females have
two X chromosomes, which behave much like all others. In males,
though, every gene on the single X-whether dominant or recessive in
females-shows its effects (which is why there are more color-blind men
than women: the gene, a recessive, is on the crucial chromosome).
As a result, any new mutations in a man are likely to bear more heav­
ily not upon his sons (who receive only his Y) but upon his daughters,
who get one of their two X chromosomes from him. There lay the first
clue about sex and age. Among European royalty, the daughters of old fa­
thers die earlier than do those with younger sires. The difference is as
much as two years for a fifty-year-old compared to a thirty-year-old fa­
ther. For sons, parental age made no difference.
Males, it seems, are not just conduits for genes between females. In­
stead, they are responsible for most mutations. Males make sperm all the
time, while females make their eggs early on, releasing them when
needed. As a result, more cell divisions take place between the sperm in
one generation and that in the next than between egg and egg. In hu­
mans, a mother uses an egg separated by a couple of dozen cell genera­
tions from the egg that made her. A father, in contrast, makes sperm
separated by hundreds of generations of sperm from the cell responsible
for his own existence. The chance of error increases each time a cell di­
vides.
Older fathers (with many divisions behind each elderly sperm) have a
rate of error twenty times greater than do females. Males are, as a result,
the source of much of the raw material of evolution. To compare the
 Laws of Variation 1 13

males and females of related species of mammal reveals that the Y chro­
mosome-which spends its time in males alone-changes much faster
than does the X. In birds, in contrast (in which females rather than males
have an equivalent of the y) , the Y chromosome evolves at normal speed.
The accumulation of genetic change hence arises not from how sex is de­
termined, but from masculinity itsel£
Mutations are not just simple faults in a rigid set of commands, but
part of a flexible and inconstant system that works to its own rules. Most
creatures have a complex system of enzymes that repair DNA, which is
such an unstable chemical that it would decay without constant help.
They evolved, in effect, to reduce the mutation rate: indeed, if it could,
natural selection might act to eliminate it, halting evolution altogether.
In spite of the apparent chaos in the DNA, the rate of error for indi­
vidual genes is quite small, at about one in a million per generation. The
figure seems tiny, but in total is quite large. London has about two mil­
lion cats and each cat perhaps a hundred thousand genes. There may
hence be two hundred thousand genetic changes each year in that city
alone. Worldwide, any mutation is almost a certainty. If it is useful it will
at once be picked up by natural selection.
Does genetic accident limit the rate of evolution? In some senses, it
must. Pigs, after all, have not mutated to make wings. To increase the
number of errors can sometimes improve the ability to respond to a chal­
lenge. Plant breeders know this, and irradiate their seeds in the hope of
turning up new and useful forms. Even so, evolution has to wait for its
raw material before natural selection can get to work. If a mutation does
not happen, the process becomes impotent. Tsetse flies, for example, are
susceptible to most insecticides because they have not come up with the
genes to deal with them. Some agents of disease, too, have failed to
evolve their way around medical advance, although some day they may.
Thus, syphilis was for many years easy to treat with penicillin, but a re­
sistant strain has now been found in Africa. No doubt it will spread.
Resistance to insecticides, now ubiquitous, did not begin for a couple
of years after the first use of DDT. The brief respite before the pests could
fight back reflects the wait for a mutation. As soon as one appeared it
spread, showing that selection was indeed limited by the lack of a gene
to work with. One housefly evolved a sudden resistance to a certain
chemical. Each of its defiant billions, from Iran to Chile, carries the same
1 14 D A RW I N ' S G H O S T

genetic change, with the same length of DNA around it. Each copy must
descend from the same error within a single animal. Evolution seized its
chance as soon as it arrived, and the new gene took almost no time to fill
the world.
Mutation alone is not enough. On a farm sprayed with a dozen pesti­
cides, to have new genes able to protect against one or two--or eleven­
is not much help. Twelve changes in a row in the same family line is too
much to ask even of insects. The situation is saved by the most funda­
mental of all laws of variation: sex. Most of genetics is no more than the
scientific study of that eccentric pastime. Sex makes offspring unlike ei­
ther parent because they contain new combinations of genes. It allows a
favorable alteration in one family to get together with another in a sepa­
rate line. Without it, there would be a long wait in each lineage for the
second one to turn up. In an all-female mosquito, if such a thing existed,
the only safe individuals would be the direct descendants of the first to
strike lucky, rather than the multitude of otherwise unrelated animals
into which a new gene can spread through sex. Free exchange between
families has enabled some insects to resist twenty. poisons at once. Each
mutation appeared in a different line, and in distant parts of the world,
but soon got together. They outwit the best the chemical industry
can do.

Correlation of Growth. Inheritance is full of links among disconnected

characters. Darwin noted them, but was bafRed by what seemed a series
of incoherent facts. What can be more singular than the relation between
blue eyes and deafness, and the tortoiseshell color with the female sex in
cats; or, again, the relation between the hair and teeth in the naked Turk­
ish dog? It can hardly be accidental that if we pick out the two orders of
mammalia that are most abnormal in their dermal coverings-whales,
and armadillos, scaly anteaters and so on-that these are likewise the
most abnormal in their teeth.
Nobody can do everything. As a result, to breed from animals or crops
desirable for one reason often leads to failure in another part. An attempt
to breed mice as large as rats began in the 1 930s and lasted for thirty
years. It was abandoned with an animal less than twice the size of a typ­
ical mouse, because the much-selected beast was sterile. Much later,
genes for growth hormone were engineered into mice, and the rat-sized
 Laws o/ Variation 115

mouse became a reality. The failure of the mouse breeders involved an

unexpected correlation of growth among the genes responsible for size
and for sex and is a reminder of how little such traits are understood.
These patterns show, in their several ways, how one change can alter
many at once. In the 1 940s, a goat without forelegs was born in Russia.
It learned to stand upright and developed large hind legs, a curved spine,
an oval chest cavity and a thick neck. A single accident during develop­
ment had led to all these changes. Whether genes were involved in the
fate of the unfortunate goat nobody knows, but plenty of small changes
in DNA can have equally large and unexpected effects.
Often, the error takes place early. Blue-eyed cats are deaf because those
at first sight unrelated characters share an embryonic pathway. The gene
involved makes the dark pigment-melanin-that is responsible for skin
color. Melanin is found in many other parts of the body, brain, eyes and
ears included. Black cats are full of it, while pale animals have less. Those
with no pigment at all are white, with blue eyes. Melanin plays an unex­
pected part in the brain, for it guides the cells responsible for certain
nerve pathways to their correct places. As a result, a shortage of melanin
gives a cat a white coat and blue eyes-and makes it deaE
The bald and edentulous Turkish dogs tell another part of the genetic
story. A joint loss of hair and teeth is found in many animals (such as the
Chinese crested dog once popular, for symbolic reasons, with striptease
artists) . Darwin himself, fifteen years after The Origin, wrote of "a Hin­
doo family in Scinde, in which ten men, in the course of four genera­
tions, were furnished, in both jaws taken together, with only four small
and weak incisor teeth and with eight posterior molars. The men thus af­
fected have very little hair on the body, and become bald early in life.
They also suffer much during hot weather from excessive dryness of the
skin . . . Though the daughters in the above family are never affected,
they transmit the tendency to their sons: and no case has occurred of a
son transmitting it to his sons."
The problem comes because both teeth and hair are derived from the
same tissue: damage it, and each one suffers. Sweat glands are involved,
too, which is why those with the condition are uncomfortable in hot
weather. The gene makes a protein that exchanges signals between skin
cells as they develop. Tabby mice, as it happens, have the same error, and
it has been detected in a hairless dog. Darwin's insight was to note the
1 16 D A RW I N ' S G H O S T

pattern of inheritance of this strange mixture of attributes. They are

passed through females but seen most often in males. The pattern arises,
we now know, because the gene is carried on the X chromosome. The
hairless sons of Scinde are another hint about how close he came to the
mechanism of heredity.
His tie between tortoiseshell cats and the female sex is also explained.
The gene affects hair color, and codes either for orange or for black. It is
carried on the X chromosome. A male, with his single X, has either the
orange or the black allele and is an unremarkable animal. Many females
carry both the black and the orange version of the gene. They are not
pure orange or pure black, as might be expected. Instead, in some of their
skin cells the black allele is switched on, and in others its alternative. This
gives the tortoiseshell its patches of different colored hairs. The reason
lies, again, in development, as female cells use only one of their two X
chromosomes in any tissue, switched on at random.
In this age of faith in the power of genes, the public is as gullible about
the wonders of DNA as it once was toward the one-eyed pigs of Con­
necticut. Most ancient ideas are simply wrong. Nonetheless, many ap­
parently eccentric notions-use and disuse, acclimatization, the
correlation of different parts of the same animal, the reappearance of
characters long lost-turn out, in the light of modern biology, to have a
basis in fact. Filled with complexities and exceptions as it is, genetics re­
mains the rock upon which the edifice of evolution rests.

Summary Our ignorance of the laws of variation is profound. Not in one case
out of a hundred can we pretend to assign any reason why this or that part
differs, more or less, from the same part in the parents. But whenever we
have the means of instituting a comparison, the same laws appear to have
acted in producing the lesser differences between varieties of the same
species, and the greater differences between species of the same genus.
The external conditions of life, as climate and food, &c., seem to have in­
duced some slight modifications. Habit in producing constitutional dif­
ferences, and use in strengthening, and disuse in weakening and
diminishing organs, seem to have been more potent in their effects. Ho­
mologous parts tend to vary in the same way, and homologous parts tend
to cohere. Modifications in hard parts and in external parts sometimes af­
fect softer and internal parts. When one part is largely developed, per-
 Laws a/ Variation 1 17

haps it tends to draw nourishment from the adjoining parts; and every
part of the structure which can be saved without detriment to the indi­
vidual, will be saved. Changes of structure at an early age will generally
affe ct parts subsequently developed; and there are very many other cor­
relations of growth, the nature of which we are utterly unable to under­
stand. Multiple parts are variable in number and in structure, perhaps
arising from such parts not having been closely specialised to any par­
ticular function, so that their modifications have not been closely
checked by natural selection. It is probably from this same cause that or­
ganic beings low in the scale of nature are more variable than those
which have their whole organisation more specialised, and are higher in
the scale. Rudimentary organs, from being useless, will be disregarded
by natural selection, and hence probably are variable. Specific charac­
ters-that is, the characters which have come to differ since the several
species of the same genus branched off from a common parent-are
more variable than generic characters, or those which have long been in­
herited, and have not differed within this same period. In these remarks
we have referred to special parts or organs being still variable, because
they have recently varied and thus come to differ; but we have also seen
in the second Chapter that the same principle applies to the whole indi­
vidual; for in a district where many species of any genus are found-that
is, where there has been much former variation and differentiation, or
where the manufactory of new specific forms has been actively at work­
there, on an average, we now find most varieties or incipient species.
Secondary sexual characters are highly variable, and such characters
differ much in the species of the same group. Variability in the same
parts of the organisation has generally been taken advantage of in giving
secondary sexual differences to the sexes of the same species, and spe­
cific differences to the several species of the same genus. Any part or
organ developed to an extraordinary size or in an extraordinary manner,
in comparison with the same part or organ in the allied species, must
have gone through an extraordinary amount of modification since the
genus arose; and thus we can understand why it should often still be vari­
able in a much higher degree than other parts; for variation is a long­
continued and slow process, and natural selection will in such cases not
as yet have had time to overcome the tendency to further variability and
to reversion to a less modified state. But when a species with any extra­
ordinarily-developed organ has become the parent of many modified de­
scendants-which on my view must be a very slow process, requiring a
long lapse of time-in this case, natural selection may readily have suc-
1 18 D A RW I N ' S G H O S T

ceeded in giving a fixed character to the organ, in however extraordinary

a manner it may be developed. Species inheriting nearly the same con­
stitution from a common parent and exposed to similar influences will
naturally tend to present analogous variations, and these same species
may occasionally revert to some of the characters of their ancient pro­
genitors. Although new and important modifications may not arise from
reversion and analogous variation, such modifications will add to the
beautiful and harmonious diversity of nature.
Whatever the cause may be of each slight difference in the offspring
from their parents-and a cause for each must exist-it is the steady ac­
cumulation, through natural selection, of such differences, when benefi­
cial to the individual, that gives rise to all the more important
modifications of structure, by which the innumerable beings on the face
of this earth are enabled to struggle with each other, and the best adapted
to survive.
 C H A P T E R V I

D I F F ICULTIES ON THEORY

Difficulties on the theory of descent with modification - Absence or rarity of

transitional varieties - The fate of hybrids - Transitions in habits of life
and the origin of flight - A change of diet and a new existence - Or­
gans of extreme perfection - Organs of little apparent importance, from
caves to Everest - The neutral theory of molecular evolution -The star­
tling structure of the genome - Partial, profligate and promiscuous
DNA - The confederacy of life

Long before having arrived at this part of my work, a crowd of difficul­

ties will have occurred to the reader. Most have been noticed (and used)
by anti-evolutionists since the subject began. The lack of intermediates
between species, groups with forms distinct from their relatives, animals
and plants of strange and unique habits: all seem hard to explain on the
theory of slow and gradual change. Even worse, there have evolved some
structures that appear to play no part in the body's economy and-an­
other problem--complex organs that seem flawless. Can instincts be ac­
quired and modified through natural selection? What shall we say to so
marvelous an instinct as that which leads the bee to make cells, which has
practically anticipated the discoveries of profound mathematicians? And
how can we account for species, when crossed, being sterile and produc­
ing sterile offspring, whereas, when varieties are crossed, their fertility is
unimpaired? Behavior and sterility deserve chapters of their own: but all
these apparent difficulties, much used as fuel for the creationist diatribe,
are, in truth, each evidence for evolution.
1 20 D A RW I N ' S G H O S T

On the absence or rarity oftransitional varieties. Existence is divided into

its many kinds and seems to have been since it began. Why is nature not
all in confusion instead of the species being, as we see them, well defined?
The primeval soup has turned into minestrone. Why is life so lumpy?
Nobody, say the anti-evolutionists, has ever seen a species arise. That,
as it happens, is not true, but it is hard to deny that few creatures appear
to be in transition between one form and the next. Why should this be?
It has to do with natural selection and with the replacement of the old
by the new.
Evolution is, for most of the time, a race to stay in the same place. The
worst enemies of any animal are among its relatives and descendants,
who need the same things and may have evolved better ways to get them.
Unless a parent can keep up with its children, its fate is sealed. Most can­
not, and disappear. .As a result, at any time, just the tips of the twigs of
any evolutionary tree are on view. The branches have been replaced by
something better. Because the past has been wiped out by the present,
the ancients usually leave no clue to what their fate might have been.
Nowadays there are no animals halfway between whales and hip­
popotami, or houseflies and fruit flies, although Darwinism demands
that once there were.
The problem of the missing links arises in part because it is hard to see
who they may have been. The dinosaur whose descendants gave rise to
birds had arms a mere fraction longer than its doomed sibs and the same
may be true of some organ of some bird today that might be a step to­
ward a new and dramatic form of future life. Even so, neither the fossil
record nor the modern world is full of creatures caught in the act of a
change in lifestyle. Why should this be, as so many must have made a
shift from one existence to another?
Biologists are only too used to the criticism that part of an eye or an
ear is of no use and that evolution is as a result disproved. The argument
denies utility to a plate camera, a carbon microphone, or the ENIAC
computer of the 1 940s that used a room full of valves to provide little
more computing power than a digital watch. That to day's versions are
better than what went before says little about what they were worth in
the days ofDaguerre, Edison, or Alan Turing. For life, the problem of the
intermediates is more subtle: the eye, the voice or the brain were not de­
signed from scratch, but had to get from where they were to where they
are, step by step, while still doing their original job.
 Difficulties on Theory 121

It may be hard to get from one form to another without a middleman

who is worse at an old task although his descendants are better at a new
one. Evolution often faces the mountaineer's dilemma. Few peaks are a
straight slog upward to the summit. Instead, a climber has to lose some of
his hard-won gains by crossing a valley before he can reach the next high
point. Plenty of tasty butterflies gain protection from predators because
they mimic, almost exactly, the bright warning patterns of unrelated
species filled with poison. By so doing they flourish, but how could their
camouflaged ancestor have taken the first and imperfect step towards a
false advertisement? Any gene that made a savory insect easier to see must,
it seems, be disadvantageous, even if, in time, it leads to a new form of pro­
tection. Quite how the insects traversed the valley of death-in a sudden
leap, with a single gene pushing them most of the way, or by small changes
getting together almost by accident-is not clear. "What is certain is that
the intermediates were worse off than both their camouflaged ancestors
and their dishonest descendants and must certainly have disappeared.
From time to time, a natural experiment hints at how such forms meet
their demise. "When species that diverged not long ago form hybrids, an­
imals appear that are halfway between two well-adapted forms. They
hint at the dangers of obsolescence. Such creatures are doomed to death
or a narrow existence because all else is denied to them. Suppose that
three varieties of sheep are kept: one adapted to an extensive mountain­
ous region; a second to a narrow, hilly tract; and a third to wide plains at
the base. The great holders on the mountains or on the plains will im­
prove their breeds more quickly; and consequently the improved moun­
tain or plain breed will soon take the place of the less improved hill
breed. The intermediates, of sheep or anything else, are squeezed out.
Take the crow, denizen of mountain and plain (and devourer oflambs'
eyes) . Most crows are black; they are the carrion crow-"that loathsome
beast, which cries against the rain"-ofEngland, Wales and much ofEu­
rope. The crows of Scotland, Scandinavia and Eastern Europe (hooded
crows, as they are called) differ from their loathsome cousins as they have
pale gray sides. Apart from its flanks, the hooded looks almost the same
as its somber kin (although it prefers to live in rather different habitats).
"Where the two birds meet, they hybridize, to give a zone full of birds
halfway between the carrion and the hooded kind. The hybrids are re­
stricted to a narrow band between their improved descendants. In Scot­
land, the hoodeds take the high ground, the carrions the low, while the
1 22 D A RW I N ' S G H O S T

intermediates are confined, like the sheep of the foothills, to a narrow

strip in between. They have no chance of supplanting either common
form.
Victorian Cambridgeshire was full of hooded crows {as noted by
Charles Kingsley, the first person to turn Darwinism to theological
ends}. In The �ter Babies, the crows kill one of their number because
she will not steal eggs: "They are true republicans, these hoodies . . . so
that for any freedom of speech, thought, or action, which is allowed
among them, they might as well be American citizens of the new school."
By the 1 950s, those radical birds had retreated to the north of a line from
Glasgow to Aberdeen. Fifty years on, the hooded has been pushed far­
ther back, to Inverness. As the carrion crow expands its range, the
hooded retreats, because the two kinds are so similar that they cannot live
together. The hybrid zone, too, is on the move, but it stays as narrow as
it ever was and is, as before, confined to the foothills. The intermediates
relive the fate of an ancient and supplanted bird. They have no future be­
cause carrion and hooded crows are already so close in their needs that
no other animal can squeeze in.
What about the greater steps on the evolutionary road? Why are there
so few forms in transition not just between low and high ground but
from land to air? Again, selection has done its inexorable work. When we
see any structure highly perfected for any particular habit, as the wings
of a bird for flight, we should bear in mind that animals displaying early
transitional grades of the structure will seldom continue to exist to the
present day, for they will have been supplanted by the very process of nat­
ural selection. Each new form will tend to take the place of, and finally
to exterminate, its own less improved parent. If the improvement is great
it will not take long to complete the move from old to new. The chances
of survival for any ancestor are small indeed.
Most animals able to fly do the job quite well. Those who hesitated on
the boundary of the new medium have gone, but it is possible to guess
at what they were like. For bats or birds, the task is simple. Their wings
are modified arms. Plenty of animals behave as the forefathers of the
eagle or the vampire bat might have done. They glide from high to low,
to save energy and to avoid the dangers of the ground. Lizards, frogs, ro­
dents and even a remarkable flying lemur {at first misclassified as a bat
rather than a primate} all go in for it. A flap of skin between forelegs and
 Difficulties on Theory 1 23

hind legs, enlarged feet or a flattened body all help. Part of an airfoil,
even in the form of enormous toes, is a great deal of use. To change a fore­
limb into an organ of flight is no great task. Each bone in a bird's or a
bat's wing has its match in the rabbit's foreleg or the whale's flipper.
However, nothing today looks like a creature halfway to a bird or bat.
Bats are, the molecules show, related to rabbits, an eminently terrestrial
group, but there are no living hints at what the ancient rabbit-bat might
have looked like. In the family of squirrels, though, we have the finest
gradation from animals with their tails only slightly flattened, and from
others with the skin on their flanks rather full, to the so-called flying
squirrels that have their limbs and even the base of the tail united by a
broad expanse of skin, which serves as a parachute and allows them to
glide through the air to an astonishing distance from tree to tree. The bat
wing membrane still reveals traces of an apparatus originally constructed
for gliding. If such a sequence from land-bound life to expert flight can
be found among different animals today, why should there not have been
such a gradual change among the ancestors of today's bats or birds? Any
improvement in the ability to glide or flap would soon put paid to a less
effective foregoer.
Most flying animals are not birds, but insects. Like angels, they grow
wings without losing their arms. The structure of an insect wing gives no
clue about what its ancestor may have been, and insects have no obvious
equivalent of the squirrels to suggest what their predecessors lived
through.
The first airborne insects to be preserved as fossils, more than three
hundred million years ago, were already blessed with magnificent wings
that carry most of the struts and airfoils used by modern dragonflies to
fly with such skill. The distant and unknown parents of those aeronauts
might have had simple skin folds that helped their bearers to glide, or
clumps of hair that kept them afloat. Perhaps, instead, the first step was
via a central-heating radiator. To aim a flat plate at the sun soaks up en­
ergy, to turn away loses it. In time the solar panel grew to become useful
in another sphere. The earliest wings could even have emerged from flaps
used by aquatic insects to absorb oxygen and hijacked by evolution as
aids to flight; but it is hard to see how an animal that lives in water took
to the air.
All those routes to a new world have analogs-the curse of evolu-
1 24 D A RW I N ' S G H O S T

tion-today. Crickets leap upward and glide to ground. For tiny insects,
the air is viscous enough to allow them to drift through the summer sky
supported by tufts of bristles. Butterflies, in contrast, open their wings to
the sun to warm up and take off. Any organ able to do such things may
have allowed a feeble flight. Which one actually did the job once rested
on speculation.
DNA proves what the first fliers were. Shrimps are distant relatives of
insects. A search through their genes reveals a set almost identical to
those that help make the wing of a fruit fly. The shrimp versions are ac­
tive not in the body wall (as might be true if the glider, balloonist or cen­
tral-heating theories were right); but in a set of specialized limbs used as
gills. The wing, they show, must have evolved from a jointed leg, used
first not for waving, but to stop its owner from drowning. A few animals
still follow the ancient ways. Those primitive beasts, the stoneflies, de­
velop in streams and emerge as adults to the surface. They use raised ap­
pendages based on gills to sail or to skate until they reach land. That, no
doubt, is what the first wings looked like, and there, among the stone­
flies, they remain, as implausible steps on the road to flight.

On the origin and transitions of organic beings with peculiar habits and
structure. Natural selection can make new organs and new forms to re­
place what went before. The theory of evolution has to do much more
than that. It must explain how some animals adopt habits quite different
from those of their relatives. Such dramatic moves could not, it seems, be
achieved by gradual change, as there must be a shift in the dozens of
characters that separate the old version from the new.
We sometimes see individuals of a species following habits widely dif­
ferent from those of others and might expect that such individuals would
occasionally have given rise to new species, having anomalous habits, and
with their structure either slightly or considerably modified from that of
their proper type. Plenty of animals take up unlikely opportunities as
they arise. To imagine from the behavior of an eccentric bear that a whole
race could, in time, be rendered more and more aquatic, till at last a beast
was produced as monstrous as a whale, is a little much to ask even of nat­
ural selection. Even so, a small shift in behavior can have large effects on
the future of those who make it. Of cases of changed habitats it will suf­
fice merely to allude to the many British insects that now feed on exotic
 Difficulties on Theory 1 25

plants. Each has had to adapt to new food and to a new place to mate
and to spend its days.
The Europeans who settled the United States saw the chances on offer.
Apples were planted wherever they would grow and soon developed into
huge orchards. In the 1 860s, in the valley of the Hudson, the crop was
attacked by a new pest, the apple maggot fly. It was at first assumed to be
a European immigrant but was, in fact, a native that had changed its
habits. A local insect was able to evolve, within a century, a new calling.
It now does millions of dollars of damage each year.
The apple pest is an altered form of the hawthorn fly. The first of those
animals to visit the new host found a mountain of food. A simple shift
led to a long chain of consequences and to a step toward a new species
with an identity of its own. It was the latest of many in its family. One
fly made the crucial move, and billions of its descendants profited.
The flies mate where they feed, which itself cuts off the apple visitors
from their ancestors. They were faced with new challenges to which they
had to adapt. Apples appear earlier in the year than hawthorn, the fruit
takes a whole hot summer to ripen and has its own defensive chemicals.
To make up for that, a maggot deep inside an apple is safe from the par­
asitic wasps that attack those on the small fruits of the original host and,
as an added bonus, it does not need to compete with the many local in­
sects who enjoy a meal of hawthorn. As a result, there has been much
evolution among the descendants of the first hawthorn fly to take to a
new diet, and the apple pest has now altered so much that it almost never
meets its forefathers. Given the choice, it will fly to an apple rather than
a hawthorn and-although it has no great genetic differences from its
ancestor-the apple flies emerge two weeks before their original form.
This reduces the chances of sex (although, in the lab, the two pests still
fall on each other with enthusiasm).
Other, more distant, relatives hint at a similar history. One, the dog­
wood fly, mates with the apple maggot in the laboratory, but not in na­
ture. The blueberry maggot fly never breeds with any other and artificial
hybrids are unfit. Yet another lives on wild laurels, is unable to cross with
any of its kin, and contains genes found in none of them. For each mem­
ber of the group, a small change of habit was the first step to an identity
of its own. Now, the hawthorn fly is testing its ability to live on cultivated
cherries. If it succeeds {and the U.S. Department of Agriculture is doing
1 26 D A RW I N ' S G H O S T

its best to stop it), that native American will assume yet another person­
ality.
Many insects have evolved alongside the plants upon which they feed.
A plant diet drove them to diversify. Three hundred thousand kinds of
beetles are known, far more than any other group. They feed, in the
main, on flowering plants, themselves evolved from ancestors without
flowers. A pedigree based on genes shows that the beetles followed their
food. Those near its root still eat conifers or cycads, ancient and conser­
vative plants, much older than anything with flowers. Their DNA shows
that, for the first hundred million years of beetle history, not much hap­
pened. Not until the plants themselves blossomed could the beetles di­
versify their habits and set off, with the flowers, down a tangle of
evolutionary roads.
Sometimes, the footprint of the crucial individual with a novel
lifestyle remains in its descendants. Like the wheel or the thermionic
valve, it shows how a single idea can lead to a whole range of new prod­
ucts.
The world is full of poisons. Some, like deadly nightshade, are lethal,
but others (such as the nicotine in tobacco) we have learned to love. The
body, too, generates wastes that must be made safe. A specialized group
of genes does the job. They began with the first poison of all. Oxygen ap­
peared in the atmosphere two billion years ago. It was fatal to the life of
those days (as it still is to the bacteria that have not evolved to deal with
it). Anything able to remove the deadly gas was picked up by natural se­
lection. There soon evolved a protein that attached the oxygen molecule
to other chemicals and reduced its malign effects.
Then, long ago, there was a neat inversion of evolutionary logic. Oxy­
gen became friend rather than foe and most organisms began to use it to
fuel their lives. The defensive protein was utilized not to make the gas
safe, but to add it to noxious substances and to render them harmless.
One change of habit generated a family of genes that branched into
thousands of different forms. Every mammal has at least two hundred
versions of the protective molecule, insects many more. Without their
defenses they would at once fall victim to the chemical world in which
they live.
Some of the new proteins result from an arms race between animals
and plants. Every new plant poison was met by an animal molecule
adapted to deal with it and, as one party changed its habits, its opponent
 Diffic ulties on Theory 1 27

followed. Some plants are ahead in the race (which is why we do not eat
rhubarb leaves), but some animals can attack a plant denied to others be­
cause they alone can handle what it makes. The tobacco budworm can
break down nicotine and flourishes on tobacco plants, a single leaf of
which will kill a man.
Swallowtail butterflies are spectacular, their bright colors improving
the forest's gloom and the glare of the savanna. Each of the two hundred
different kinds has a host plant of its own, kept more or less to itself be­
cause other insects avoid them. All their hosts make a toxin called
coumarin. This keeps most grazers off, but is defeated by the evolution­
ary shift that came to their progenitor. The evidence of an ancient tran­
sition is in their DNA. Each species has a gene that can destroy
coumarin. Although parts of its structure have changed, the small section
that attacks the poison has not altered over millions of years. A single
mutation in an ancient insect opened the gates of an evolutionary
citadel. It allowed hundreds of new kinds of butterfly to evolve and to ex­
ploit a new and diverse set of circumstances.
Whole groups of animals may undergo such unexpected shifts in rou­
tine and evolve in directions quite different from their ancestors. Can a
more striking instance of adaptation be given than that of a woodpecker
for climbing trees and for seizing insects in the chinks of the bark? Yet in
North America there are woodpeckers that feed largely on fruit, and oth­
ers with elongated wings that chase insects on the wing; and on the plains
of La Plata, where not a tree grows, there is a woodpecker, which in every
essential part of its organization, even in its coloring, in the harsh tone
of its voice, and undulatory flight, tells its close blood-relationship to our
common species; yet it is a woodpecker which never climbs a tree.
Swallowtails, woodpeckers and many other creatures all show how na­
ture can change in an arbitrary way. For each, the world now has a set of
products that it did not know it needed. Evolution, given the chance of
a better hole, always goes to it. If nobody else can get in, so much the bet­
ter. A peculiar habit, or a novel structure, opens a world of opportunity
that can be exploited in a myriad ways.

Organs ofextreme perfection and complication. The perfection of life has

often been used to prove the existence of a Creator. Griffith Hughes, rec­
tor of St. Lucy's, Barbados, put the case in his Natural History ofBarba­
dos of 1 750. In the island's animals he "traced the Workmanship of a
1 28 D A RW I N ' S G H O S T

Divine Architecture . . . without Defect, without Superfluity, exactly fit­

ted and enabled to answer the various Purposes of their Creator, to min­
ister to the Delight and Service of Man, and to contribute to the Beauty
and Harmony of the universal System . . . For instance, the Potato­
Louse, which is so small that it is scarce discernible! . . . Yet every Part
that is necessary to animal Life is as truly found in one of them as in Be­
hometh or Leviathan . . . What less than infinite Wisdom and Power,
could dispose a little Portion of Matter, almost too small to be viewed by
the naked Eye, into that infinite Variety?"
The notion that existence is so flawless as to need a designer was much
taken up by the Victorians and is still trotted out today. The logic is
empty. Perfection is relative and, for potato lice or anything else, depends
on subjective judgment. The eye, the ear-even the toenails-all rebut
the "argument from design," as the claim of Griffith Hughes and his
many successors is known.
Although the belief that an organ so perfect as the eye could have been
formed by natural selection is more than enough to stagger anyone, yet
in the case of any organ, if we know of a long series of graduations in
complexity, each good for its possessor, then, under changing conditions
of life, there is no logical impossibility in the acquirement of any con­
ceivable degree of perfection through natural selection.
The American politician William Jennings Bryan-the victor in the
trial ofJohn Scopes, convicted in 1 925 for teaching evolution-liked to
mock the subject: ''A piece of pigment or, some say, a freckle appeared
upon the skin of an animal that had no eyes. That piece of pigment or
freckle converged the rays of the sun upon that spot and when the little
animal felt the heat on that spot it turned the spot to the sun to get more
heat. The increased heat irritated the skin-so the evolutionists guess,
and a nerve appeared there and out of the nerve came the eye . . . Can
you beat it? And it happened not once, but twice!"
In fact, the eye happened not twice but fifty times, and the problem
ofhow to extract information from light has been solved in a dozen ways.
The eye is as intricate as it needs to be, and no more. Its apparent per­
fection does not destroy but upholds the theory of evolution. There are
many sequences of eyes in different creatures. Each hints at the stages
that even an organ as complex as our own must have passed through be­
fore it gained the moderate abilities it can claim today.
 Difficulties on Theory 1 29

Many animals have a single lens used to focus light on to a plate able
to convert it into nerve impulses. Humans, worms, jellyfish, snails and
spiders all do the job in much the same way. As paparazzi know, the big­
ger the lens the better it sees, and mice have eyes larger in relation to their
body size than are our own. Even a tiny device does quite well. The sim­
ple eye of a spider, like the complex organ of the mouse, tells enemy from
friend at thirty times its bearer's length.
All eyes reflect what history has demanded and are restricted by what
it provides. The human eye is complicated enough, with a hundred mil­
lion rods, used in dim light, and three million cones, responsible for
color vision. Each rod contains thousands of proteins that transform
light into signals via a molecule that crosses the membrane in a sevenfold
zigzag. Three color-sensitive pigments pick up the red, green and blue el­
ements of a scene. Our eye is imperfect, but we are used to what it can­
not do. The world has plenty of white flowers, but only to us. Bees can
see in the ultraviolet, and to them the plants are full of detail. All cam­
eras correct for the colored fringes that surround an image passed
through a lens; our own sneaks around the problem with a shortage of
blue-light receptors in the center of its field {a fact noted by the Impres­
sionists, who blurred their blue flowers}.
In spite of such adjustments, evolution will not produce absolute per­
fection, nor do we always meet, as far as we can judge, with this high
standard under Nature. Everything has been modified, but not per­
fected, for its present purpose. Any structure that evolves has to cope
with its past. Because the chief part of the organization of every being is
simply due to inheritance it is, for most of the time, impossible to get to
one place from another. Every animal is limited in what it can do by what
it starts with.
The eye is a servant of that inflexible rule. That of mammals has a
weakness that has dogged it since its earliest days. It began as a patch of
cells and was later formed into a cup. As a result, the light must pass
through the wires that take information to the brain before it reaches the
retina. No camera that put the sensitive part of the film on the wrong
side and then had to compromise to cope would sell. The feeblest de­
signer could improve it {which is why we have eyeglasses and micro­
scopes}.
Any insect would be astonished by our ability to see. Their eyes are
1 30 D A RW I N ' S G H O S T

built with not one but hundreds of lenses, each of which concentrates
light upon a sensor. That set of tiny and cheap cameras is a forceful state­
ment of what evolution cannot do. Because of where they began, insect
eyes are limited in what they are. They specialize in the big picture and
are no good at details. Insects are Nature's victims. k any movement
could mean death, they have a bird's-eye view of the world, every object
in their sights, any activity at once detected. However, the most suspi­
cious insect is ten times less able to identify the fine points of an adver­
sary than is a spider with a crude but more effective organ of vision. Their
worldview was described by the first scientist to take photographs
through insect eyes as "a picture about as good as if executed in rather
coarse wool-work and viewed at the distance of a foot."
Whatever the limitations of its raw material, natural selection has im­
proved their eyesight as far as it can. Nocturnal insects have large lenses
that increase sensitivity by a hundred times, while dragonflies have more
cameras, with a patch of tight-packed small units able to pick up prey
against the sky. Bees go on long journeys and have an upright strip of
sensors adapted to the vertical world of trees and branches. The world of
water-skaters, by contrast, is flat and their eyes have a horizontal band,
suited for the watery plain upon which they swim. Sex comes in, too.
Male flies have acute vision to stalk a potential mate to whom her suitor
is a distant blur. Even the molecules change to fit. Light causes ions to
rush across a membrane and sparks off a train of impulses passed to the
brain. Fast fliers, faced with a stream of new information, have an ion
channel able to respond at once, while that of their relatives with a more
tranquil existence is slower.
k insects battle to improve a feeble design, evolution does its best, but
that best is not very impressive. The eye of the dragonfly or the water­
skater has triumphed, but only because all its competitors are worse. For
sight, excellence is in the eye of the beholder.
In the context of evolution, perfection is not necessary. If the eye were
only a hymn to the supreme powers of a deity called natural selection it
would be no more persuasive as evidence than was William Paley's cele­
brated watch as proof of the existence of God. His book multiplied ex­
amples of flawless design, and, with no other idea of whence it came,
turned to a Great Designer. Unfortunately for him, the song of the eye
has many discordant notes. They show it to be not the work of some
 Diffic ulties on Theory 131

great composer but of an insensible drudge: an instrument, like all oth­

ers, built by a tinkerer rather than by a trained engineer.

Organs oflittle apparent importance. Natural selection can give rise to or­
gans of extreme complexity (eyes included) and has often done so. Any
increase in the ability to see is useful and is at once seized upon. But what
of organs of little apparent use? How can evolution explain the origin of
simple parts, of which the importance does not seem sufficient to cause
the preservation of successively varying individuals?
Part of the answer is: important to whom? We might not be able to see
the point of a structure, but it may be crucial to its owner. The giraffe has
a tiny tail, which looks like an artificial fly-flapper. It seems at first in­
credible that this could have been adapted for its present purpose by suc­
cessive slight modifications, each better and better, for so trifling an
object as driving away flies.
Flies suck blood and carry parasites. The tsetse fly makes it impossible
to raise cattle in much of southern and central Mrica. Its attacks on live­
stock are so fierce that it once forced humans, all over that continent, to
carry their own loads as no pack animals could survive. The flies still
cause a billion dollars' worth of damage a year. Whether a cow is black,
cream colored or patchy might be thought a most trivial character-but
some breeds are more liable to attack because tsetse are attracted to large
blocks of continuous color and avoid animals with patterns. Even the
limited protection given by a flapper is of some help. The tail, the coat,
or even the eyelashes of the cow or the giraffe are, trivial as they appear
to us, crucial to its defenses.
Scientists often dismiss organs as unimportant because they have not
bothered to find out what they do. Once, many of the endocrine glands
of the body-the pineal, or the thymus-were shrugged off as mere use­
less relics, rather than as the masters of its internal economy. Even the eye
has unexpected tasks. It is used to see with, but it does much more. Many
cave animals are blind. Few things seem less useful than an organ of sight
in a place with no light. In the dark, the eye at once loses its impor­
tance-or so it might seem. In fact, darkness reveals uses that are other­
wise invisible. The blind mole rat of Israel has the smallest eyes of any
mammal, sealed beneath the skin. Even the parts of its brain involved in
vision are much reduced. However, its organ, diminished as it is, is still
1 32 D A RW I N ' S G H O S T

an eye. It has kept the remnants of sight in a place where it can see noth­
mg.
Some of the eye's nerves go, not to the visual centers of the brain, but
to the hypothalamus, a place much involved in the control of tempera­
ture, of feeding and of sex. A single three-second burst of illumination
will set the brain clock, and in mole rats enough light crosses the eyelids
in the brief moments when they kick earth out of their burrows to tell
them how long the day is and at what time of year to breed. Even the
sightless need the remnants ofwhat once allowed them to see. To destroy
a blind eye disrupts the rhythm of existence. The organ has, it seems,
powers revealed only after generations of darkness.
Even so, plenty of cave animals have the remnants of eyes that seem to
be of no use at all. In many fish, even the nerve connection with the brain
has gone. In some of the crabs the stalk for the eye remains, though the
eye is gone; the stand for the telescope is there, though the telescope with
its glasses has been lost. These organs may be real relics, of no impor­
tance, but how is it possible to prove a lack of use?
Nowhere is it easier to dismiss the value of any structure than in DNA
itsel£ Great tracts seem to have no function and the molecule has mil­
lions of sites that differ at random from one individual to the next-or
so it appears. It is easy (and may be fair) to see most of its changes as be­
side the evolutionary point. Can all the millions of differences between
two mice or two fruit flies-or the many more that separate insects from
mammals-have evolved to cope with a struggle for existence? Perhaps,
some say, most of the DNA is an organ of small importance, whose pres­
ence is not noticed by Darwin's machine. If so, much of life is neutral
ground upon which natural selection enacts its rare battles.
That view is supported by a surprising fact: that the parts of the body
that vary most are those that appear to be least important. Blood clots are
made when small proteins link together in response to damage. For
much of the time each one floats in the plasma, its ability to bind
checked by a short section that blocks the crucial site. After a cut, the
plug is snipped out by a special enzyme, the molecules link up and the
clot forms. Most of the protein does not vary at all, but the stopper, with
its simple job, is filled with diversity. Natural selection surely cannot act
to retain differences in the part of the molecule with the least exacting
task. Most of the changes in the blocking section probably have no effect
 Diffic ulties on Theory 1 33

on how it works and merely accumulate with time. In DNA as a whole,

the parts that make no protein vary more than those that do, and the
more embedded in the machinery of the cell any protein may be, the less
variable is its gene. Diversity, it seems, builds up where it does no harm,
but is excluded from places where it might cause trouble.
True champions of Darwin find it painful to admit that most varia­
tion under nature is a spectator at the evolutionary play. Even so, the idea
is now accepted, almost by default. The molecular clock and the trees
used to link distant beings assume that change in DNA measures no
more than the passage of time. As most clocks and trees make at least
some sense, perhaps the belief is fair.
However, the clock itself hints that to dismiss most diversity as ran­
dom noise may be a mistake. In sharks, a sluggish group, the clock ticks
slowly; in bony fish with the same length of life at greater speed; and in
the primates (ourselves included) for some reason is slower than in other
mammals. Something is changing the clocks, but what it is, nobody
knows.

All selection-on behavior, on color, or on proteins-is, in the end, on

DNA. Almost never do we know where in that giant molecule it is at
work. Most of the genetic material does a job that is quite obscure and
in our ignorance it becomes as easy to dismiss the value of any change as
it was to belittle the tail of the giraffe. Sometimes, though, a molecule's
structure and function come together, to show how dangerous it can be
to deny use to any characteristic until it has been studied in detail.
The red pigment of the blood, hemoglobin, has many tasks. Its most
important job is to pick up oxygen in the lungs and move it around the
body. The molecule consists of two pairs of protein chains, folded
around each other. The oxygen fits into a cleft between them and escapes
as the chains move apart when the blood reaches the sites where it is
needed. The bar-headed goose migrates across the summit of Everest at
five miles above sea level, a place where humans die within a few hours.
Andean birds spend much of the year in conditions almost as tough.
Each bird's hemoglobin stays in its oxygen-binding shape long after that
of others has opened up. The bar-headed goose alters a single DNA let­
ter, which removes a crucial contact between the chains and makes the
1 34 D A RW I N ' S G H O S T

protein more eager to bind oxygen. The Andean goose alters the same
junction to the same effect, but with an alteration in the opposite chain.
Andean vultures become avid for the life-giving molecule with quite a
different set of mutations. The three subtle pieces of genetic engineering
are each as direct a response to natural selection as are the wings of the
melanic moth. Without an intimate knowledge of hemoglobin it would
be easy to dismiss them as random noise.
Even so, one bar-headed goose differs from the next, or from its An­
dean cousins, in millions of sites throughout the DNA. Can selection
have crafted them all? It seems improbable. For most of the time natural
selection must act as a policeman rather than as an architect. It does not
adapt every molecule to each shift in the environment, but spends its ef­
forts on a purge of mutations that interfere with the smooth operation
of the body. After all, a random change rarely improves a device that al­
ready works well. To hit a heart-lung machine with a hammer does not
often increase the oxygen supply (although, sometimes, it might). In
evolution, most changes are for the worse, and most selection acts to pre­
vent modification and not to promote it.
To keep a police force on the alert-and to punish those it does not
approve of-is expensive. If every one of the millions of inherited differ­
ences influenced their carriers' ability to stay alive or to reproduce, al­
most everyone would be unlucky in what they drew; one of their genes
would fail the test and the rest (advantageous as they might be) would
pay the price when selection struck down their bearers. Such a stringent
application of the Darwinian rules might make it impossible for a pop­
ulation to sustain itself Many biologists hence assume that variations
neither useful nor injurious would not be affected by natural selection,
and would be left a fluctuating element. Most diversity, on this neutral
view of the world, is mere froth on the surface of the great Darwinian
sea: random noise in a system whose important parts are fine-tuned by
selection.
But, others counter, if selection works on what seem at first sight triv­
ial characters like the blind eye of a mole rat, it could influence all diver­
sity, slight as its importance might appear. So far (apart from the odd
exception, such as the blood proteins of mountain birds) there is little ev­
idence one way or another. Once, the action of selection was denied in
things that seemed as trifling as the color of moth wings or the structure
 Difficulties on Theory 1 35

of hemoglobin. Now, those are seen as the great proofs o f its power. Per­
haps the same will happen to all changes at the DNA level-but perhaps
not.

Hemoglobin has many messages for evolution. It bears on all the sup­
posed difficulties of descent with modification and shows most of them
to be false. However, the molecule and its fellows also raise problems that
are hard to accommodate within a Darwinian framework. Perhaps the
greater number are only apparent, and those that are real are not fatal:
but the natural history of DNA may need explanations beyond those of
The Origin ofSpecies. In spite of the fuss about its supposed defects, that
great work has done well at explaining the diversity of plants and ani­
mals. It does rather worse when it comes to the structure of genes.
First, there is not much fit between the size or complexity of the body
and that of the genes that build it. A whale has three thousand million
DNA bases, but a certain salamander less than an inch long has twenty
times as many. Other animals have shed such genetic paraphernalia alto­
gether. The fugu, or puffer fish, has all its genes packed into a length of
DNA an eighth of our own, while small green algae-whole nucleated
cells, with all their machinery-contain a mere three hundred and eighty
thousand DNA bases. Their genes are jammed together and even over­
lap, with the end of one marking the start of the next. That, it seems, is
the minimum needed to make a cell. Why does a mammal need ten
thousand times as much and the salamander twenty times more? Sala­
manders are more complicated than seaweeds, but surely not that much.
There also seems to be a pressure for change within the DNA itselE
The Malthusian world of "severe labour . . . excesses of all kinds . . .
wars, plague and famines" includes plants and animals as well as the un­
deserving poor. For all of them natural selection involves a clash between
individuals for survival or for sex. Now, biology's attention is being
drawn to another conflict: to a war beneath the skin, to the struggles
among genes. DNA has, it appears, its own agenda that may conflict
with the interests of its carriers. Some of the molecular battles can be ex­
plained in familiar terms, but some follow rules that seem at first sight
quite alien to Darwinism.
In the 1 960s, geneticists noticed an odd result in their crosses. When
1 36 D A RW I N ' S G H O S T

1 38 D A RW I N ' S G H O S T

between them, arise from simple change over time? Why has there been
no divergence among copies within the midwife or its cousin, given that
the molecule has altered so much since they split? All this is not a mere
aberration among toads. The two common European mice each contain
twenty thousand copies of a particular section of DNA. It is the same in
all its details within each one, but different between them, although they
severed relations a mere few thousand years ago. There has, it seems, been
a sort of ethnic cleansing of the genes. Just one version is allowed to re­
main within a species, whatever happens when different species are com­
pared.
The dissidents can be purged in many ways, forcible conversion in­
cluded. DNA is surrounded by a priesthood of enzymes anxious to cor­
rect its smallest errors. Without them, it would soon fail. If the repair
enzymes are biased in their belief about what the correct message should
be, then that version of the genetic creed is bound to take over. Other
methods of genetic purification can homogenize a DNA sequence. In
places with many copies of a particular string of letters, the segments
tend to mispair, rather like the teeth of a zipper done up too quickly. One
version may as a result have a built-in tendency to increase at the expense
of the other and to drive it out.
Such behavior hints that genes have an evolutionary agenda of their
own. Perhaps, to parts of the DNA, species are no more than a place to
live, great continents of animals linked by sex. Different species (such as
southern and northern midwife toads) may look much the same, but
from the molecule's point of view each is an island isolated from its
neighbors by a sexual barrier. As a result, each evolves to its own internal
rules.
Startling as this might be for Darwinians, other patterns are even more
unexpected. Certain genes make great leaps across the living world. He­
moglobin itself crops up in unexpected places. A few insects (such as the
midge larvae found in stagnant water) have the molecule and are able, as
a result, to take up habits widely different from their allies. How did a
protein from mammals get into an insect?
Vertebrate DNA also has a curious distribution. A certain piece is
found in sheep, goats and cows-and in several snakes and a couple of
lizards. No other animal has anything like it. An ancestor of the boa con­
strictor, the viper and the rattlesnake was the source from which this no-
 Difficulties on Theory 1 37

After all, an organism whose retina is inside-out has n o right to complain

about the oddities of its genes. Eyes and genes each evolve through a se­
ries of compromises. Tapeworms hijack the machinery of the gut, which
has to cope with its unwelcome visitor-why should not transposable el­
ements do the same with the cell?
Darwinism is not mocked by movable or repetitive DNA, but it still
has a lot of explaining to do. Physics, after relativity, could no longer ac­
count for the behavior of atom, apple and universe in the rational terms
of Newton. Within genetics, too, is hidden a world of turmoil that may
need new kinds of answers. Evolution lacks its Einstein-and, some say,
does not need one-but its theories, successful as they have been, are
sorely tested by the natural history of the genes.

Descent with modification predicts that two groups apart for many years
will, on the average, differ more than two that split not long ago. For
bones, or behavior, or blood proteins, the rule works well. In parts of the
DNA it fails so badly as to give pause to Darwin's most dedicated sup­
porters.
Consider the case of the midwife toad, notorious for its failure to dis­
prove Mendelism when the inheritance of an acquired character (a black­
ened foot) was found to turn on the furtive injection of ink by a corrupt
biologist. Now the toad has become an icon for those who hope to tran­
scend Darwin.
Toads have a set of genes that makes parts of the machinery used to
translate the message of the DNA. Hundreds of copies of the genes are
present in every animal; and each one has exactly the same structure
(which is odd, given the tendency of any multiplied gene to diverge from
its fellows) . The midwife toad has a blood relation, the northern mid­
wife, much alike in appearance. That too has a gene family with an iden­
tical job (as do most of its relatives). Each is derived, no doubt, from
some ancient shared ancestor.
The structure of the DNA is quite unexpected. Every duplicate of the
much-repeated gene in the midwife toad has precisely the same sequence
as every other; while the northern midwife has just as many copies, all
identical-but all different from those of its relative. How can this pat­
tern of absolute identity of repeats within a species, but fixed differences
1 38 D A RW I N ' S G H O S T

between them, arise from simple change over time? Why has there been
no divergence among copies within the midwife or its cousin, given that
the molecule has altered so much since they split? All this is not a mere
aberration among toads. The two common European mice each contain
twenty thousand copies of a particular section of DNA. It is the same in
all its details within each one, but different between them, although they
severed relations a mere few thousand years ago. There has, it seems, been
a sort of ethnic cleansing of the genes. Just one version is allowed to re­
main within a species, whatever happens when different species are com­
pared.
The dissidents can be purged in many ways, forcible conversion in­
cluded. DNA is surrounded by a priesthood of enzymes anxious to cor­
rect its smallest errors. Without them, it would soon fail. If the repair
enzymes are biased in their belief about what the correct message should
be, then that version of the genetic creed is bound to take over. Other
methods of genetic purification can homogenize a DNA sequence. In
places with many copies of a particular string of letters, the segments
tend to mispair, rather like the teeth of a zipper done up too quickly. One
version may as a result have a built-in tendency to increase at the expense
of the other and to drive it out.
Such behavior hints that genes have an evolutionary agenda of their
own. Perhaps, to parts of the DNA, species are no more than a place to
live, great continents of animals linked by sex. Different species (such as
southern and northern midwife toads) may look much the same, but
from the molecule's point of view each is an island isolated from its
neighbors by a sexual barrier. As a result, each evolves to its own internal
rules.
Startling as this might be for Darwinians, other patterns are even more
unexpected. Certain genes make great leaps across the living world. He­
moglobin itself crops up in unexpected places. A few insects (such as the
midge larvae found in stagnant water) have the molecule and are able, as
a result, to take up habits widely different from their allies. How did a
protein from mammals get into an insect?
Vertebrate DNA also has a curious distribution. A certain piece is
found in sheep, goats and cows-and in several snakes and a couple of
lizards. No other animal has anything like it. An ancestor of the boa con­
strictor, the viper and the rattlesnake was the source from which this no-
 Diffic ulties on Theory 1 39

madic gene got into the progenitor of today's farm animals forty million
years ago. Within both snakes and mammals, its evolution fits the stan­
dard patterns of relatedness, so that the transfer must have happened just
once. Rattlesnakes and sheep pale when compared to plants. A piece of
mobile DNA hidden within a gene for part of the metabolic machinery
has mounted an assault on a whole range of vegetation. It started off in
a fungus, but has invaded a thousand or more hosts, from coffee to fox­
gloves to bananas, picked off at random from the three hundred thou­
sand kinds of plants with flowers.
Although the idea that genes can move between such different places
is unexpected enough, such long-distance commerce is everywhere.
Plant and animal cells are complicated things. As well as the nucleus with
its DNA and RNA, they contain mitochondria, the power generators;
various whips and lashes used to row themselves about, and complicated
cisterns that contain their raw material. They may, it seems, be ancient
confederacies of creatures once separate but now living together. For ex­
ample, mitochondria have their own DNA, quite different from that of
the nucleus; not a string of millions of bases but a small closed circle.
When put onto the grand evolutionary tree, it groups not with animals,
but with bacteria (whose own genes are arranged in a loop) . An ele­
phant's mitochondria are, as a result, more similar to the inhabitants of
its guts than they are to most of its own DNA.
Elephant cells, like those of all other advanced creatures, must descend
from an ancient society made up of several members. The genetic struc­
ture of the mitochondrion is close to that of the agent of typhus, the dis­
ease that killed twenty million people at the time of the First World War.
Typhus is transmitted by lice and can live only inside cells. Its agent (like
mitochondria themselves) cannot make its own amino acids, but feeds
off its host. Long ago, it seems, two creatures-one now a mitochon­
drion, the other its enclosing cell-patched up their differences and lived
in harmony. In time, the treaty ripened into an indissoluble bond, and
each party found it impossible to live without the other.
That has now widened into a federation. Different parts of the cells of
trees or elephants trace their ancestry to a whole range of ancient beings.
The whips and lashes came from another bacterial group, the ancestors
of the bacterium that causes syphilis. In the same way, the green elements
of plant cells descend from bacteria able to use energy from sunlight.
1 40 D A RW I N ' S G H O S T

Other parts of the cellular machinery, internal pipework included, were

also independent, although their own genes are now lost to the nucleus.
Perhaps the nucleus itself was once a separate organism, engulfed by an­
other. Mer all , it contains both DNA and RNA, the latter used as the
genetic material by a few creatures today.
A crucial hint of how wide the confederacy of life might spread comes
from bacteria themselves. E. coli, common in our own guts, has had its
entire complement of bases laid out. Great segments of its DNA speak a
language different from the rest, give evidence of a deep split in the bac­
terial family tree. A fifth of that creature's genes come from elsewhere
among the bugs.
The emerging insights into molecular anatomy oflife show that, three
hundred million years ago, gene exchange was universal. In some senses,
species themselves were late arrivals on the evolutionary scene for, in
those distant days, free trade ruled, with genes leaping from one form of
life to another. All genomes of all higher creatures are a patchwork of
parts that started in different places and retain traces of a bastard ances­
try from the earliest times. The structure of a thousand genes is known
from a wide enough range of beings, from bacteria to yeast and worms,
to trace the remote past. They group not-as simple descent with mod­
ification would predict-by those who bear them, but by what they do.
One set-whether it finds itself in plants, animals or bacteria-orga­
nizes, operates and edits the information kept in DNA. The other does
household tasks such as repair, food preparation, waste disposal and
moving around. The information branch resembles the genes of simple
bacteria that pump out methane, while the rest of the genetic material
has been assembled from many places. The housekeeping genes have, it
seems, been hopping about almost since life began, while the data
processors are less mobile (perhaps because they have to communicate
with others) .
Life is much more fluid than it once seemed. Trees, by most defini­
tions, have a trunk; but there are plenty of exceptions. Banyans, for in­
stance, have stems that run into one another to make a tangled mass.
Trees of genes look much the same. They show that not only is the cell a
coalition, but the genes themselves descend from separate founders and
have shuffled around in a way unimagined before the advent of molecu­
lar biology.
 Difficulties on Theory 141

The structure of DNA raises problems so grave for the theory of evo­
lution that it is hard to reflect on them without being staggered. Genet­
ics shows how Darwinism can explain what seems at first inexplicable. It
is also a useful reminder that a science without difficulties is not a science
at all.

Summary of Chapter. We have in this chapter discussed some of the difficul­

ties and objections which may be urged against my theory. Many of them
are very grave; but I think that in the discussion light has been thrown
on several facts, which on the theory of independent acts of creation are
utterly obscure. We have seen that species at any one period are not in­
definitely variable, and are not linked together by a multitude of inter­
mediate gradations, partly because the process of natural selection will
always be very slow, and will act, at any one time, only on a very few
forms; and partly because the very process of natural selection almost
implies the continual supplanting and extinction of preceding and inter­
mediate gradations. Closely allied species, now living on a continuous
area, must often have been formed when the area was not continuous,
and when the conditions of life did not insensibly graduate away from
one part to another. When two varieties are formed in two districts of a
continuous area, an intermediate variety will often be formed, fitted for
an intermediate zone; but from reasons assigned, the intermediate vari­
ety will usually exist in lesser numbers than the two forms which it con­
nects; consequently the two latter, during the course of further
modification, from existing in greater numbers, will have a great advan­
tage over the less numerous intermediate variety, and will thus generally
succeed in supplanting and exterminating it.
We have seen in this chapter how cautious we should be in conclud­
ing that the most different habits of life could not graduate into each
other; that a bat, for instance, could not have been formed by natural se­
lection from an animal which at first could only glide through the air.
We have seen that a species may under new conditions of life change
its habits, or have diversified habits, with some habits very unlike those
of its nearest congeners. Hence we can understand, bearing in mind that
each organic being is trying to live wherever it can live, how it has arisen
that there are upland geese with webbed feet, ground woodpeckers, div­
ing thrushes, and petrels with the habits of auks.
Although the belief that an organ so perfect as the eye could have been
formed by natural selection, is more than enough to stagger any one; yet
1 42 D A RW I N ' S G H O S T

in the case of any organ, if we know of a long series of gradations in com­

plexity, each good for its possessor, then, under changing conditions of
life, there is no logical impossibility in the acquirement of any conceiv­
able degree of perfection through natural selection. In the cases in which
we know of no intermediate or transitional states, we should be very cau­
tious in concluding that none could have existed, for the homologies of
many organs and their intermediate states show that wonderful meta­
morphoses in function are at least possible . For instance, a swim-bladder
has apparently been converted into an air-breathing lung. The same
organ having performed simultaneously very different functions, and
then having been specialised for one function; and two very distinct or­
gans having performed at the same time the same function, the one hav­
ing been perfected whilst aided by the other, must often have largely
facilitated transitions.
We are far too ignorant, in almost every case, to be enabled to assert
that any part or organ is so unimportant for the welfare of a species, that
modifications in its structure could not have been slowly accumulated by
means of natural selection. But we may confidently believe that many
modifications, wholly due to the laws of growth, and at first in no way ad­
vantageous to a species, have been subsequently taken advantage of by
the still further modified descendants of this species. We may, also, be­
lieve that a part formerly of high importance has often been retained (as
the tail of an aquatic animal by its terrestrial descendants), though it has
become of such small importance that it could not, in its present state,
have been acquired by natural selection-a power which acts solely by
the preservation of profitable variations in the struggle for life.
Natural selection will produce nothing in one species for the exclusive
good or injury of another; though it may well produce parts, organs, and
excretions highly useful or even indispensable, or highly injurious to an­
other species, but in all cases at the same time useful to the owner. Nat­
ural selection in each well-stocked country, must act chiefly through the
competition of the inhabitants one with another, and consequently will
produce perfection, or strength in the battle for life, only according to the
standard of that country. Hence the inhabitants of one country, generally
the smaller one, will often yield, as we see they do yield, to the inhabi­
tants of another and generally larger country. For in the larger country
there will have existed more individuals, and more diversified forms, and
the competition will have been severer, and thus the standard of perfec­
tion will have been rendered higher. Natural selection will not necessar­
ily produce absolute perfection; nor, as far as we can judge by our limited
faculties, can absolute perfection be everywhere found.
 Diffic ulties on Theory 1 43

On the theory of natural selection we can clearly understand the full

meaning of that old canon in natural history, "Natura non facit saltum."
This canon, if we look only to the present inhabitants of the world, is not
strictly correct, but if we include all those of past times, it must by my
theory be strictly true.
It is generally acknowledged that all organic beings have been formed
on two great laws-Unity of Type, and the Conditions of Existence. By
unity of type is meant that fundamental agreement in structure, which we
see in organic beings of the same class, and which is quite independent
of their habits of life. On my theory, unity of type is explained by unity of
descent. The expression of conditions of existence, so often insisted on
by the illustrious Cuvier, is fully embraced by the principle of natural se­
lection. For natural selection acts by either now adapting the varying
parts of each being to its organic and inorganic conditions of life; or by
having adapted them during long-past periods of time: the adaptations
being aided in some cases by use and disuse, being slightly affected by
the direct action of the external conditions of life, and being in all cases
subjected to the several laws of growth. Hence, in fact, the law of the
Conditions of Existence is the higher law; as it includes, through the in­
heritance of former adaptations, that of Unity of Type.
 C H A P T E R V I I

INSTINCT

Instincts and habits, inborn and learned - Conflict, cooperation and compro­
mise - Slave-making instincts - Natural instincts of the cuckoo and os­
trich - Simple rules make complex habits: cell-making and social habits
of the hive bee - Gradual shifts to new societies - Cannibalism -
Neuter or sterile birds, rats and insects - The evolution of behavior
through natural selection and kinship - Summary

The American Kennel Club standard for the Chinese Shar-Pei has as its
ideal an animal "regal, alert, intelligent, dignified, lordly, scowling, sober
and snobbish." The Rottweiler, in contrast, should be "calm, confident
and courageous . . . with a self-assured aloofness that does not lend itself
to immediate and indiscriminate friendships."
Breeds differ in how highly strung they are, how much they snap at
children and in their fondness for barking. When it comes to how easy
each type is to housebreak. or how much they enjoy simple destruction,
all are about the same. Everyone knows that terriers are excitable, that pit
bulls bite and that all pups urinate on the carpet. Temperament, like size
or shape, is in the genes. It is known that a cross with a bulldog has af­
fected for many generations the courage and obstinacy of greyhounds;
and a cross with a greyhound has given to a whole family of shepherd
dogs a tendency to hunt hares.
An action, which we ourselves should require experience to enable us
to perform, when performed by an animal-more especially by a very
young one, without any experience-and when performed by many in-
 Instinct 1 45

dividuals in the same way, without their knowing for what purpose it is
performed, is usually said to be instinctive.
Instincts are of the highest importance to each animal. They show re­
markable adaptations, from the migrations of birds to the slave-making
castes of ants. If behavior varies and is inherited, it has no choice but to
evolve. Apparent miracles of complexity in animals act hence need no ex­
planation beyond those involved in how they are built. Behavior reveals
the force of natural selection in a manner that mere appearance may not.
Can habits or instincts, intricate and flexible as they may be, follow
the same evolutionary rules as color, shape or size? That might seem un­
likely, for behavior is often learned or comes from a simple reflex. In the
end, though, every action of every animal is a product of genes, however
much the environment determines what those genes might do. There is
no mystery about such things. Behavior comes from brains; and brains,
like hearts or kidneys, are made by genes. Thomas Henry Huxley, in a
stark appraisal of the human condition, claimed that the brain secretes
thought, as the liver bile. Our own brain is made of millions of cells, each
obedient to the message of its DNA. Within it, half the genes of the body
are at work. They are as open to change as are others with less noble tasks.
Genes make brains; and brains make behavior. Thus is instinct trans­
mitted across the generations.
Many animals show inherited differences in how they comport them­
selves. Even bacteria "behave," in some general sense, as they interact
with one another, with their food or with their host. One soil microbe
has, like a lion, a social life. It congregates into swarms and, when things
get tough, makes a special structure that splits up to form a new colony.
Behavior may be as variable as is shape or size. Some wild mice have a
gene for "neophobia" -fear of the new, be it food or nest hole-not
found in their fellows, while melanic moths prefer to sit in more cam­
ouflaged places than do their light-colored sibs. �uch traits can be em­
braced by natural selection as much as can an increase in a molecule's
capacity to bind oxygen.
Not all behavior is in the genes, as habit or custom rules much of what
animals can do. Any dog can learn to stand on its hind legs. It is not done
well only because bipedalism is laid over an ancestral wish to stay on all
fours. Birds can do more: a certain American nutcracker stores as many
as thirty thousand seeds a year, each one in a hole in a tree. Every bird
1 46 D A RW I N ' S G H O S T

learns the map of its homeland and can return to a seed six months after
it srowed it away. In the same way, parrots can be trained to talk and
monkeys to manipulate signs in a strained and distant imitation of
human grammar.
A little dose of judgment or reason comes into play even in animals
very low in the scale of Nature. Take, as an instance, the sea anemone, a
creature that seems safe from most forms of intellect because it lacks a
brain. Anemones are among the simplest of all animals, mere sacks of
flesh with a fringe of tentacles. They release larvae into the water. These
settle, divide and grow into colonies of identical partners. One Pacific
kind lives in clonal groups separated from each other by strips of bare
rock. Within a colony all is peaceful, but among them rages constant
conflict.
All anemones have sting cells (some of which secrete a poison that can
kill a man). The wars of the clones are unending. Each has its own per­
sonality. Some are aggressive, while others are calmer but respond at once
to attack. Some clones do not fight back, but instead throw more soldiers
into the front line as their members are killed. Colonies are able to settle
only next to those against whom they have some chance in a fight. In
time, a resentful truce emerges and battle starts only when a newcomer
arnves.
Anemone society-brainless as it is-shows how all animals, even the
lowliest, have an existence beyond that written into DNA. For some
creatures, habit rules most of what they do. Each of the thirteen species
of Galapagos finch is born with a fixed taste for its own food, be it fruits,
or seeds, or insects, picked from the ground or branches, or gouged out
with a cactus spine. One is a vampire, drawing blood as it pecks at the
backs of seabirds. All members of each species go for more or less the
same behavior, whatever it might be. The Cocos Island finch, in its home
three hundred miles off South America, has a different attitude toward
life. Individual birds vary in diet almost as much as do separate species of
their famous relatives. Some eat insects, others nectar, or fruit or seeds,
or even snails and small lizards. Each sticks to what it likes.
Cocos Island, unlike the Galapagos, contains but a single kind of
finch; but it is an animal with a talent for discovery. Each bird picks up
its tastes in its first days. It copies its elders and, by holding to its own
task, reduces the amount it has to remember. It inherits not a fixed set of
 Instinct 1 47

actions, but the ability to adapt to many. With a change of habit, a sin­
gle species fills as many gaps in the economy of nature as does a whole
group on the Galapagos themselves.
Learning, knowledge, habit, tradition-or whatever else it might be
called-can itself be inherited. For much of the year, humpback whales
live as two separate populations around the Poles, but all move to the
Equator to breed. The northern and southern groups mate at random as
they frolic in the tropical waters, but the calves follow their mothers to
one Pole or the other and learn to feed where she takes them. A history
held in the mind rather than in the genes splits them to the opposite ends
of the earth.
Sometimes, a new habit can be seen as it begins. Somewhere, a blue
tit was the first to open the top of a milk bottle, an idea that spread
through much of Europe. There are other local cultures. Thus, the
chimps of the Cote d'Ivoire use rocks to crack nuts, a talent absent from
their kin a few hundred miles away.
The culture of birds and apes comes from their history and their abil­
ity to learn. How they behave is built upon what evolution has provided
(which, in the brain of the chimpanzee, is quite a lot) . In humans, biol­
ogy grants even more, but the same laws apply. If Mozart, instead of play­
ing the pianoforte at three years old with wonderfully little practice, had
played a tune with no practice at all, he might truly be said to have done
so instinctively. Of course, he did not. His ability to learn the piano came
from his ancestors. He played as he did because he was Mozart. Genes set
the limits even to genius. Although that is not a term to be applied to an­
imals, what they do is also constrained by what they inherit. Even a brain
tiny in comparison to that of Mozart can achieve remarkable things.

Slave-making instinct. One of the most powerful testaments to natural se­

lection is to observe that the instinct of each species is good for itself, but
has never, as far as we can judge, been produced for the exclusive good of
others. Some animals may seem to be good Samaritans; but in truth they
never are. Even the many cases of apparent cooperation-plants and
their pollinators or the mitochondria that live inside cells-evolved
through a mutual manipulation that can soon go wrong. The balance of
cost and benefit is always calculated, and although many creatures suffer
at the hands (or teeth) of their enemies, none selflessly help their friends.
148 D A RW I N ' S G H O S T

Some parasites admit their lowly status and creep into the body of
their host. Others give the impression that they do not cheat but rule;
while yet others set up societies in which two or more creatures seem to
live in harmony. Different though each appears, all are somewhere on the
path that links conflict to cooperation. Where they end up depends on a
struggle among instincts as vicious as that against the elements.
Malthusian theory, inexorable as it was in condemning the poor to
starve, saw the common good as superior to self-interest. Darwin's view
was simpler and more ruthless. To him, evolution had no common­
wealth; self-interest is what matters. He was right. There is no charity in
Nature.
The ants run the gamut from conflict to conquest to apparent coop­
eration. A few insinuate themselves into the colonies of others and per­
suade their hosts into a free meal. With no work to do, the hangers-on
become feeble, with thin skins and mouths reduced to tubes. Some
evolve to become, like fleas, tiny travelers on the skin of a larger ant. Oth­
ers are more confident. Certain ants raid the nests of others and seize
them as slaves. The ant is absolutely dependent on its slaves; without
their aid, the species would certainly become extinct in a single year. The
males and fertile females do no work. The workers or sterile females,
though most energetic and courageous in capturing slaves, do no other
work. They are incapable of making their own nests, or of feeding their
larvae. When the old nest is found inconvenient, and they have to mi­
grate, it is the slaves which determine the migration and actually carry
their masters in their jaws.
Energetic and courageous though the slave-makers might appear, they
are parasites-for they could not live without their lackeys. Slave­
makers, with their fearsome jaws, are as much dependent on their vic­
tims as are their feeble relatives. Some ants enslave their own kind. Thus,
honeypot ants often attack neighbors and carry off their workers. More
often, they steal from other species; the Amazon ants, for instance (wide­
spread in Europe) thieving from the red ants around them. The colony
is attacked, its guards killed and the cocoons taken back home. There,
when they hatch, they labor for their masters, unaware that they are help­
ing to pass on foreign genes. Some slave-makers are more ingenious.
Rather than risk an attack, they spray the slave colony with an alarm sig­
nal and, as its guards flee in panic, steal the cocoons. One American
 Instinct 1 49

driver of slaves takes advantage of two of its neighbors: a small kind to

act as a worker and a larger insect to serve as its soldiers.
Slaves and their owners have obvious conflicts of interest, as one gains
at the expense of the other. Other relationships seem at first sight to be
benign. Ants and aphids have evolved a mutual dependence; the ants
gain from the sweet secretions of the aphids (and even "milk" them to
obtain it) and the aphids of those species, lacking soldiers of their own,
trust in the jaws of the ants to keep their enemies at bay. Neither party is
a philanthropist. The aphids provide not just sugars, but vital nutrients
that the ants would otherwise lack. Their excreta-the "manna" of the
Bible-is nine tenths sugar (and, as bees take advantage of what is left by
ants, much of our own honey is made from their droppings) . So dose is
the relationship between ant and aphid that some queen ants carry
aphids with them when they found a new colony, to be sure of their
source of food. The aphids, in turn, have bacteria in their guts that help
them digest food. Kill them off with antibiotics, and the aphids-and
their ants-will die. Some plants help with modified leaves in which the
ants nest (and, by stinging, keep out other insects anxious to eat the
leaves). A network of what seems like kindness is in fact a web of mutual
exploitation.
Ants have progressed from slavery into domestication. The New
World has hundreds of species that depend on fungus gardens, carefully
tended by special castes. The first farmers appeared fifty million years
ago. They fertilize the small mushrooms in their nests and pick out weeds
or spray them with herbicides. When a queen sets off to found a new
colony she carries a small packet of fungi to set up a garden of her own.
The farmers depend on their harvest to such a degree that they have lost
their own stomach enzymes and depend on the fungi to digest food.
Every ant colony has its own done of fungi, distinct from that on the
neighboring farm. Ancient though it is, their association seems to be an
open one, with both parties benefiting from it. Some of the farmed
mushrooms are identical to others that live free and have been picked up
only recently.
Friendship is, of its nature, unobtrusive. When two parties live in har­
mony, neither makes much fuss and mutual dependence may pass un­
observed. Lichens, consortia of fungi and algae, were once thought to be
single creatures. Only when the relationship ends and each struggles to
1 50 D A RW I N ' S G H O S T

gain the upper hand is attention drawn to the conflict. When the cost of
cooperation outweighs its benefits, society soon breaks down. The soil
bacteria that behave so harmoniously in their native home can grow in
liquid culture in the laboratory. Within a thousand generations, the an­
imals no longer meet to feed or share a reproductive structure. Instead
they dwell as sturdy loners, each happy to live and breed in indepen­
dence. The genes for cooperation have been replaced by others that do
the job more cheaply.
Why should a slave submit to its master, or a fungus allow itself to be­
come a commodity? The balance is finely adjusted. Clones of identical
fungi are passed down from parent colony to offspring, reducing the
chance of conflict in a well-adjusted system. Both parties have reduced
their options through their mutual dependence, which helps it to persist.
Some fungi, however, creep in as weeds and are suppressed with a special
chemical that kills them alone.
Slavery goes well beyond the insects. Plenty of animals persuade oth­
ers to bring up their young. About one kind of bird in a hundred lays its
eggs in the nest of another (and some fish even dump their eggs in the
mouths of others who tend them). A battle-a microcosm of many other
struggles-rages between exploiter and exploited, with sometimes one
party ahead and sometimes the other. Because it is unresolved it hints at
the tactics involved before a compromise is reached. In birds, the habit
has evolved at least half a dozen times (cuckoo ducks, for instance, take
advantage of gulls) . It can cost the host a lot-a whole brood pushed out
or stabbed to death by a sharp beak. Some parasites, such as the cowbirds
of Mrica and North America, dump their eggs on any of dozens of kinds
of hosts and almost every nest may be filled with unwelcome visitors.
Others, the cuckoos included, are more refined birds and concentrate on
just a few victims.
A tiny reed warbler, faced with a cuckoo, feeds a chick much larger
than herself She defers to this foreign adolescent although it does not re­
semble her own young. Her true offspring, once satiated, stop begging;
but the cuckoo is more persistent and forces her to keep up the food sup­
ply. The alien, when it demands food, makes a noise not at all like that
of a single warbler chick but one that is close to the cacophony of a whole
nest of young warblers.
The evolutionary arms races of the avian world go on at full speed.
 Instinct 151

North America has just one common parasitic bird, Mrica thirty or
more. As a result, almost all the perching birds ofMrica can identify (and
eject) foreign eggs, while almost none of their American equivalents are
able to do so. The process can work in reverse. The village weaverbird in
its native Africa rejects most cuckoo eggs, but since it was introduced
into the Caribbean (which has no cuckoos) in the eighteenth century it
has lost its ability to judge. Education also plays a part, as first-time par­
ents are happier to accept a cuckoo's egg than are their more cynical el­
ders who have brought up a brood without an intruder.
The parasites can fight back, with a range of eggs that mimic those of
their chosen host. The huge range of the common cuckoo, from Mrica
to Asia, contains dozens of races, each specialized to lay a particular egg.
Some hosts are able to identify the foreigner and eject it or abandon the
nest, but others are less fastidious. In most places, the cuckoo mimics its
hosts: blue eggs for the redstart, say, or spotted gray, like those of the reed
warbler.
Sometimes, the parasites turn their attention to the most obvious vic­
tim. The occasional habit of birds laying their eggs in the nests of other
birds of the same species is not very uncommon, and this perhaps ex­
plains the origin of a singular instinct in the ostriches. For several hen os­
triches unite and lay a few eggs first in one nest and then in another. This
instinct, however, of the American ostrich-the rhea-has not as yet
been perfected, for a surprising number of eggs lie strewn over the plains.
Half a dozen females share a nest, a simple hollow within the square mile
or so of a male's territory. One female is the boss. She mates with that
male and lays and guards her own eggs. Then, the extra females-some
of them mated by the fortunate male-wander in and add to the pile.
They act like huge and flightless cuckoos, as they stroll off and take no
part in parental care. So well do the parasites perform that a nest may
hold forty eggs-far too many for a lone bird to mind.
The nest-holder is no fool. She can tell her own eggs from those of the
spongers, and keeps hers in the middle of the group, with the aliens
pushed to the edge. The foreigners pay the price: just half of the parasites'
eggs hatch, while almost all those of the owner do. At three pounds, an
ostrich egg is a good meal, and plenty are eaten, but those at the edge go
first because not even a hungry jackal can manage more than one. The
rootless females dump their eggs because they have failed to find a terri-
1 52 D A RW I N ' S G H O S T

tory, or have lost their own nests. As a result, it pays them to lay even if
most of the eggs are destroyed. The holder suffers their attentions not be­
cause they share her genes, but because of the help it gives to her own
eggs. No benevolence is required.

Cell-making Instinct of the Hive Bee. The beehive is a metaphor for

human society. Shakespeare was impressed: bees are "Creatures that by a
rule in nature teach / The act of order to a peopled kingdom. / They have
a king, and officers of sorts: / Where some, like magistrates, correct at
home; / Others, like merchants, venture trade abroad; / Others, like sol­
diers, armed in their stings, / Make boot upon the summer's velvet buds;
/ Which pillage they with merry march bring home / To the tent-royal
of their emperor: / Who, busied in his majesty, surveys / The singing ma­
sons building roofs of gold; / The civil citizens kneading up the honey; /
The poor mechanic porters crowding in / Their heavy burdens at his nar­
row gate; / The sad-eyed justice, with his surely hum, / Delivering o'er to
executors pale / The lazy yawning drone."
It does not take much to cause bees or any other animals to act in a
complicated, or even an elegant, way. Three simple and familiar rules are
called for: variation, inheritance and natural selection. The variation may
be in the genes or in past experience and the vehicle of inheritance can
be DNA or memory, but whatever the machinery, a complex pattern of
behavior soon evolves. Bee society also shows how the principles of eco­
nomics can be modified by kinship and lead to actions that at first sight
seem to violate the very laws of evolution.
Shakespeare tells better than most the story of the beehive. He is mis­
taken only in the sex of the ruler. The monarch of the hive is not a male
but a single female, a fact not realized until just before Shakespeare's
death: "We must not call the Queen 'Rex'; the Bee-state is an Amazon­
ian or feminine Kingdom" (a notion rejected by the bee master to King
Charles II on the grounds that it denied the divine right of kings).
The queen, who lays nearly all the eggs, is helped by several thousand
female workers, accompanied at times by a lesser number of males called
drones. She depends on the workers for food and can lay two thousand
eggs a day. The workers construct the nest and feed the young, either
with the secretions of a special gland, or with honey or pollen. Some are
builders, others undertakers who throw out the dead. When the queen
dies, another is elected by feeding her with a royal jelly that transforms a
 Instinct 1 53

lucky larva's status. Once a year, the queen allows the males a chance for
sex. When their job is done they are judged useless, and killed or thrown
out.
For bees and other insects, social life pays. Just one insect in fifty lives
in such a way, but those that take up a shared existence may flourish. The
Brazilian rainforest contains as much biomass in the form of ants (a
highly social group) as in birds, mammals, snakes and lizards combined.
The habit is fairly new. Insects appeared about half a billion years ago,
but stayed solitary until termites turned up three hundred million years
later. Although bees and ants evolved a hundred million years ago the
balance of each society is still ready to tip whenever circumstances allow.
As in Adam Smith's hypothetical factory for the manufacture of pins
(in which each workman took up his own task and much improved the
factory's productivity), an ant's nest, a termite mound or hive has divi­
sion of labor. It is hard to examine the exquisite structure of a comb, so
beautifully adapted to its end, without enthusiastic admiration. How
could such strange behavior evolve? Elephants never forget-but bees?
Their brains are tiny: why are they so smart?
Karl Marx got it more or less right: "What distinguishes the worst ar­
chitect from the best of bees is this, that the architect raises his structure
in imagination before he erects it in reality." His statement brings out the
pragmatic nature of evolution, in a hive or anywhere else. Bee society has
no plans. Instead, its complexity comes from some elementary rules. All
that is needed is for each bee to know what goes on in its neighborhood,
to pass its knowledge to others, and to have a threshold at which it
changes from one behavior to another. The hive is an information soci­
ety. Any action generates feedback and there soon emerges a system that
far transcends the actions of those within it.
Such laws determine the colony's architecture. It has been remarked
that a skillful workman, with fitting tools and measures, would find it
very difficult to make cells of wax of the true form, though this is per­
fectly effected by a crowd of bees working in a dark hive. Grant whatever
instincts you please, and it seems at first quite inconceivable how they
can make all the necessary angles and planes, or even perceive when they

are correctly made. But the difficulty is not nearly so great as it at first ap­
pears: all this beautiful work can be shown to follow from a few very sim­
ple instincts.
Each hive, with its hundred thousand cells, has three concentric re-
1 54 D A RW I N ' S G H O S T

gions. A central brood area full oflarvae is surrounded by a rim of pollen

cells, and then by a wall of cells filled with honey. The design might, per­
haps, result from some hidden blueprint of what a hive should look like,
deep within each bee's brain. The truth asks less of insect intellect.
When an empty comb is put into a hive, the cells are at first filled at
random. Then, in time, order begins to emerge. The queen wanders
about as she lays her thousands of eggs. She follows a simple rule-lay an
egg close to a cell that is already full-and does it with such consistency
that, within a few hours, 70 percent of her products are placed next to an
occupied cell. This soon leads to a mass of eggs near the center, where she
will more often recross her path.
The workers have laws of their own. They store pollen and honey at
random, but remove it more quickly from cells near a larva. Pollen is
used at ten times the rate of honey and in places with more brood cells
it runs out faster. This leads to more turnover in a pollen cell, while those
that contain honey, the long-term reserve, are soon blocked. The single
place left for a pollen delivery is near the brood, where demand soon
empties each cavity for reuse. To make the three-level pattern of the
comb all that is needed is for each bee to test the contents of the cell next
door. Soon there appears a structure that seems-but is not-well de­
signed. Its simple rules mean that small shifts in behavior cause large
changes in the system as a whole.
The hunt for food shows how information and stupidity work to­
gether to make what seems like intelligence. A hive has thousands of for­
agers who comb the landscape for miles around. They come in two
forms. Scouts buzz around alone, while the more abundant recruits,
guided by the scouts' dance, sally out when needed. Scores of bees ap­
pear, as if by magic, at a rich source, while a smaller patch is visited by
fewer diners. The regulations are simple. If a scout finds a small meal as
she Hies about, she eats it and goes on her way. If the item is large enough,
she goes back home and, with her famous waggle dance, tells her fellows
where the food is. They respond and, quite automatically, more join in
as each traveler returns with the good news. When the food has gone, its
appeal diminishes, the dance stops, the crowd disperses and a new hunt
begins.
The better the source, the longer the waggling goes on. Because each
bee reacts at once to a dance, long bouts attract many foragers while
 Instinct 155

short ones persuade only a few bees to leave home. When hungry times
set in, the scouts become less fastidious and give lengthy dances even for
poor food. All this makes the colony at once efficient and flexible.
The cybernetic hive can get quite complicated. The workers divide
their labor by age and by experience. The youngest bees clean out the
cells and nurse the brood. Some then become undertakers and eject the
dead, while others venture out as hunters. Every bee has a set task that
changes with age. As the foragers grow older they move from a juvenile
taste for sweet nectar to a more refined preference for pollen. Modest
changes in the conduct of single bees have major effects on society. Un­
dertakers take up the pastime in middle age, but their juniors will join in
to help clean up after a disaster. A shortage of pollen means that the
young turn to it and more is gathered. If not enough nectar can be
found, then older bees stick to their juvenile habits and continue to
search for the sweet substance.
Genes also play a part. In some colonies, queens mate with several
males. Quite why is not clear (although it might help protect against dis­
eases that would kill all members of a group of identical animals), but to
do so means that several lines of descent are present. DNA tests show
that, within a hive, scouts and recruits are dissimilar, so that instinct as
well as habit rules their behavior. In the same way, some bees are born
with a taste for pollen, others for nectar.
What seems so complex-bee society-obeys simple rules. They
allow each colony to be efficient but adaptable. A shift in preference ad­
justs the economy to cope with whatever hits it. Genetic diversity means
that evolution can seize an opportunity as soon as it arises. The most
wonderful of all known instincts, that of the hive bee, can thus be ex­
plained by natural selection having taken advantage of numerous, suc­
cessive, slight modifications of simpler instincts.
For bees, cooperation pays-but is it enough to explain the most re­
markable property of all, the surrender of sex to a single female? To do so
is the mark of the highest form of society. Queen bees and ants gain a
great deal from their elevated role. They are the longest lived and most
fecund of insects, with some ant queens lasting thirty years. Those of
African driver ants may each give birth to three hundred million young,
almost all of them sterile workers who pillage the forest and bring back
food for the family. Any queen also produces some males, and a few vir-
1 56 D A RW I N ' S G H O S T

gin queens of her own who fly forth in search of a possible new home­
although almost none succeed.
All this is fine for the ruling female, but what about the workers? How
can natural selection favor sterility-apparently the deadest of evolu­
tionary ends? That is a special difficulty, which might appear insuperable
and actually fatal to the whole theory. By good fortune, nature has pre­
served some hints as to how barrenness can be preferred. The answer lies
in economics, modified by family interest.
A series of stages from selfishness to cooperation remains as a reminder
of the steps taken on the road from solitary insect to the Byzantine world
of the hive. Ninety-five percent of the twenty thousand different kinds of
bee dwell not in colonies but in proud independence, in burrows, empty
snail shells, or plant stems. Each carpenter bee, for instance, makes a
small nest inside a twig or shoot, lays an egg therein, and provides it with
a ball of food. A bigger ball means a better chance for her hungry larva.
Force two females to share a nest, and things become more compli­
cated. Independence at once gives way to collaboration. The household
chores are portioned out as one bee stays close to the nest and guards it,
while the other-with no task left to do at home-spends her time in the
search for rations. Are bees quite so quick-witted? Indeed they are not,
but a division of labor does not need much logic. All that is required is
that once a task is over another should loom larger in a bee's mind. A
head start for one of the partners in a certain department-as a nest
guard, say-means that the other gets on with what is left. The mental
rule of doing what is left undone leads the bees, with no conscious effort,
to work together for the good of the larva. Who hunts and who guards
may turn on tiny differences in size or habit, but once the decision is
made, the rest must follow. With a series of such choices a society soon
emerges.
In a carpenter bee from Israel, most females set up home in pairs. The
partners are almost always mother and daughter. The mother is the re­
productive boss. She eats the eggs laid by her daughter, whose sole job is
to raise her own young sisters. The system turns on property rights and
social convention, because the bee who gets there first (always, needless
to say, the mother) lords it over the next to arrive. But how can her
daughter's actions be sustained? A small genetic change might let her
have grubs of her own. Indeed, in some years daughters set up in inde­
pendence. Why, most of the time, are they so generous?
 Instinct 1 57

It all has to do with kinship, and a shortage of places to live. Except in

the best years, good nest sites are hard to find, and any shelter is at risk
of a takeover by a rival. A home with a guard is safer and has, on average,
more young. The daughter gains because, although by guarding she loses
any hope of progeny of her own, her help gives her mother a chance to
lay another egg. As a result, copies of her own genes, contained within
her young siblings, profit.
Evolution favors teamwork, not through goodwill but because of in­
creased efficiency in multiplying DNA. Selection always acts through
genes. As mothers and daughters have such a high proportion in com­
mon, the balance of profit, loss and risk means that, for most of the time,
it pays the daughter to help her parent rather than to strike out as a
single-parent family. When the economic equilibrium changes (as it does
in a particularly favorable year) she at once abandons her mother and
makes a home of her own.
Hive-bee society, with its single dominant female, began in much the
same way, with added property rights as represented by the hive. Once
capital is inherited, a complex society makes even more sense. The first
females to take possession can control their sibs and force them to sup­
port their young.

Bees are a reminder that individuals are just part of the evolutionary
equation. As all in the end die, the fate of the genes is detached from
those who transmit them. Most members of most species passed away
long ago. Their job as conduits through which genes travel through time
is over. Natural selection acts through the medium of DNA, rather than
on the flesh of those who bear it. In other words, it is interested in kin­
ship.
Animal breeders know that well. They prefer to breed from the best­
the fattest pig, perhaps. However, a pig can be fat for two reasons: good
food, or good genes. To breed from an animal obese just because it ate
well is a waste of time. It is, experience shows, better to pick out not the
grossest of all pigs but those of more modest form from fatter families.
Their household may not contain the most corpulent beasts of all but, as
a group, they are better endowed in their genes than are most. The broth­
ers and sisters of moderately chubby pigs may not themselves be unduly
large, but to breed from them is a better way to make progress than just
1 58 D A RW I N ' S G H O S T

to select the few prize specimens out of many. Selection on the genes, and
not the individuals, does the job: a thin pig of good family may well have
fat genes struggling to get out.
The key to success-in bees as much as in pigs-lies in the blood. The
closer the affinity, the better the value of one individual predicts that of
another. The second cousin of a fat pig is a less credible candidate for ge­
netic greatness than is his brother. Kin selection, as it is called, is as im­
portant in the real world as on the farm. It explains behavior as
strange-and more-as that of the beehive.
Kinship can pay an individual to reduce its own chances if it improves
the prospects of other members of its family. It can lead to the evolution
of animals that cannot reproduce, and to others that kill their own kind.
Such actions may, to the casual or the creationist glance, appear to fly in
the face of reason, to be evidence against evolution itsel£ Strange though
instincts may appear, not one has been produced for the sole benefit of
other animals. If it had, it could not be explained by Darwinism. Gen­
erosity or selfishness emerge from natural selection. Once attention is di­
rected to the genes as well as to their bearers, such eccentric behavior
makes perfect sense.
There is more to life than kinship. Some animals care little for the fate
of those who share their heritage. The nine-banded armadillo always has
identical quadruplets. They behave to one another as they do to a foreign
armadillo, in spite of their genetic similarity. In the same way, some bees
treat their kin as they do anyone else. Perhaps these animals live in places
where no help is needed; but perhaps the opportunity for sharing has
never turned up. Benevolence needs a gene as much as black wings do
and, if it does not arise, then such behavior will not evolve.
The laws of the animal world are ruthless. Plenty of parents kill their
children, and plenty of children murder their sibs. For them, the eco­
nomic part of the argument looms large and selfishness pays: if only one
can survive, shared genes do not matter. Sometimes, though, three or
four offspring can pull through and the erstwhile murderers behave as
gently as anyone else-at least, to those of common blood.
Many creatures grow up with their sibs: in a nest, in the corpse of the
mouse in which a burying beetle lays her eggs, or as seeds in a common
fruit. Quite often, a mother has more young than have a realistic chance
of clinging to life. Perhaps, in an exceptional year, several might, or a few
may replace brothers or sisters who die by accident. Most of the time,
 Instinct 1 59

though, the fate of the extras is as a meal for their luckier kin, their nurs­
ery more an ambush than a sanctuary. The surplus is no more than a bi­
ological insurance policy, and when times look good their parents
begrudge the cost of cover. Often, the firstborn are favored from the mo­
ment of birth--or before. Birds pass testosterone to their eggs and those
with more of the hormone grow more quickly as chicks. Piglets are born
with sharp teeth. Nipples nearer the mother's head give more milk, and
the larger young seize these and fight off their sibs. Again, the rules of
cost and benefit determine who lives and who dies. Both are important,
but inheritance is often the key. Among bears and lions many young are
cared for and survive-unless another male takes over the group, when
most are killed by the intruder.
Animals are forced to live with their relatives for many reasons. In
most cases, ecology is to blame. With nowhere to go, the only choice is
to stay at home. For a termite, home is a fallen log. It may be crowded,
but to set off into the unknown in the hope of setting up another branch
of the family business is risky. Other animals live in burrows that take
years to build, or in a colony filled with thousands of workmen, so that
a pioneer has almost no chance of a residence of his own.
Once confined with one's relatives, kinship and wealth loom large.
Like families gathered at Christmas, social animals are poised between
cooperation and conflict. The more distant the tie, and the cheaper the
gift, the more the chance of a quarrel. A distant aunt is welcomed if she
is rich enough, but a cousin will be spurned if he drinks more than he
brings.
Shared blood can be a great help in passing on genes-and in per­
suading some of their carriers to accept the supreme sacrifice. Take, for
example, the aphid. Most aphids, most of the time, are celibate. Sex hap­
pens just once a year, on a tree. In some, the fertilized female lays her eggs
in a gall, a growth in the wood used as a retreat in which her asexual
progeny-identical copies of herself-develop. As the weather warms,
these multiply, without benefit of males, into millions.
Although aphids seem innocuous enough, some have soldiers to de­
fend their homes. Such animals have jaws, or horns, or claws, or sharp
mouth parts. They may be small, but are fierce enough to drive off mice
and even to bite through human skin. All are sterile. They give up their
own reproductive future to protect that of their gall-mates.
The explanation lies in inheritance. All descendants of a female are
1 60 D A RW I N ' S G H O S T

identical twins. As a soldier shares all its genes with those it guards, it
makes perfect sense for it to be sacrificed if, by so doing, it increases the
chances of its sibs. Soldiers, like the teeth of tigers, are a defense organ, a
specialized part of a mass of genetically identical cells. In a few aphids, a
single gall contains the product of several females. Such aphids never
have soldiers. They would make no sense: why sacrifice yourself for
someone who does not carry your DNA?
Self-sacrifice does not always demand absolute identity. Plenty of an­
imals give up their future to help brothers, sisters, parents or more dis­
tant relatives. In a closed community, common ancestry becomes the
rule. Any animal living in isolation-in an Alpine valley, or a fallen log­
has no choice but to mate with its kin, because there is nobody else
around. If (as in inbred lines of mice) this goes on for long enough the
residents become almost as similar to one another as are members of an
aphid clone. Selection measures the value of its subjects in the currency
of the genes they carry and estimates their worth in relation to that of
others.
The naked mole rat has a society as unattractive as its name. It lives in
underground colonies of up to three hundred in arid parts of southern
Mrica. The soil is so hard and the roots upon which the rats feed so scat­
tered that no solitary rat could survive. Within a colony just one female
reproduces. She is the largest inhabitant of all and has sex with a few of
the biggest males. Her fellows, both male and female, raise the young,
dig tunnels and, with a special class of fighters, attack (often at the cost
of their own lives) marauding snakes and foreign mole rats. In a good
year, when the soil softens after rain, the queen rat has twenty young at
a time and the workers can dig for half a mile a month.
In spite of the dangers of childbirth, the queen (and her favored males)
live for many years. So do the workers, if they are kept alone, in a zoo. In
the colony, most die young. Their lives are full of stress-not just because
of the snakes and the digging, but because they are bullied by the queen
and her partners. So intense is the social pressure that the hormones
needed for sexual development are shut down. An animal removed from
the community at once becomes mature, with a puberty that lasts a
week, and a dead empress is soon replaced by the largest of her inferiors.
However, for most of the time the queen keeps her fellows in a perpetual
and bitter adolescence. DNA is the key. It shows that every inhabitant of
 Instinct 161

a mole-rat colony is a close relative. The society has become a great ex­
tended family. By helping their relatives the workers and fighters pro­
mote their own biological heritage.
Charity can go a long way. Most of the time, animals have the same
number of sons and daughters, as each passes on the genes with equal ef­
ficiency. Under some circumstances, though, the balance can change.
One way to ensure the right mix is quite straightforward. Mothers do
not hesitate to kill the sex less able to pass on genes. The marsupial
mouse, a small and fierce Australian carnivore, has an interesting sex life.
Males live for only a year and die exhausted after copulation. Some fe­
males, too, last for just twelve months. Those fortunate enough to sur­
vive another season are old and tired and make worse mothers than they
were when young. In their first year, mothers kill all their daughters but
one, to give their well-brought-up and aggressive sons a chance. In their
second try at reproduction, though, mothers destroy not daughters but
sons, who, because they are so feeble compared to the competitors, do a
worse job at handing on her genes than do her newborn and relatively
healthy female offspring.
Other creatures are more subtle in planning their families. Parrots in
cages have runs of twenty sons or twenty daughters in a row. How-or
why-they do it, nobody knows. Rank may play a part, for mothers high
in society can choose what sex best transmits their DNA.
The Seychelles are an island paradise that, like Paradise itself, has been
abused by those who live there. Among the survivors is a small bird
called-with some lack of imagination-the Seychelles warbler. By 1 988
fewer than four hundred were left, all on the barren island of Cousin.
Then, a small group was moved to a more luxuriant islet, Aride.
On Cousin, males outnumber females. The new colonies, in contrast,
have many more female birds. On the new and fertile island parents have
more daughters than sons, while those who still starve in their native
home prefer male progeny. Again, it is a matter of passing on the inher­
ited message. Young females do not look for mates of their own but stay
close to their parents and help them to raise their young (who are, need­
less to say, their own brothers and sisters) . When food is abundant, this
is helpful. The parents lay more eggs and the genes of the generous
daughters gain as a result. When food is short, the balance changes and
the spinsters become a nuisance. They eat more than they can give-and
1 62 D A RW I N ' S G H O S T

their parents retaliate. Plenty of food makes it better to bear daughters

who can lend a hand. A shortage favors investment in sons who fly off
and find mates. In favorable habitats nine out of ten chicks are female,
in starvation territory just one in five. Birds moved from poor habitats
on Cousin to the luxury of Aride at once switch their sexual preference
toward daughters.
A wasp's nest, a beehive or an anthill has many more females than
males. Most, like the warblers, seem to commit evolutionary suicide.
They have no offspring of their own, but help others to reproduce. If
evolution depends on inherited differences in the ability to copy DNA,
how can sterile forms arise?
Again, the balance of cost and kinship is all, and a mother invests in
the sex best able to transmit her genes. The fate of the childless matters
less than what they can do for those who have children. Natural selec­
tion, sometimes helped by odd patterns of relatedness, solves the para­
dox of the infertile. It can lead to sterility, to evolution in animals that
bear no young and to individuals that put their sexual efforts into help­
ing others.
Bees, ants and wasps have an unusual way of deciding who is born
male and who female. It gives mothers control over their progeny and
leads to unexpected patterns of relatedness. They can make it worthwhile
to abandon sex, to slave on behalf of others of one's own kind, Of-now
and again-to murder them.
Unlike mole rats (or humans), male bees have but a single copy of all
their chromosomes whereas females have two. As a result, a female has
twice as many versions of every gene as does a male. A simple rule deter­
mines bee sex. Any individual with two different copies of a special sex
gene is female while any with but a single form is male. Any animal with
a solitary set of chromosomes is hence doomed to masculinity.
The queen can choose the sex of her progeny. She stores sperm and
can decide whether or not to use it. Unfertilized eggs lead to sons (with
just her own inheritance), but those that are fertilized, with genes from
her mate, make daughters. In this way the mother controls her related­
ness to her descendants. A female shares less of a heritage with her daugh­
ters than she does with her sisters.
The logic is simple. Human sisters have half their DNA in common.
A pair of bee sisters, in contrast, shares every gene passed on by their fa-
 Instinct 1 63

ther but have a 50 percent chance that any gene comes from their
mother. Sisters share not half their genes, but three quarters (all their fa­
ther's genes plus half their mother's) . Any female bee is hence more re­
lated to a sister than to any daughter she might herself have. Given the
iron rules of kinship, it then pays her to help raise those sisters, rather
than to have daughters of her own. The apparent reproductive suicide of
a female worker is, as a result, a matter of biological self-interest.
Such strange patterns of relatedness may alter the economic balance,
but do not themselves cause sterility. Termites make that point. They
look like ants (although they are not much related to them) and live in
huge colonies, most of whose members are sterile workers. Some are
ruled by single females, in a society even more rigid than that of a bee­
hive.
However, termites determine the sex of their offspring in a way not
much different from our own, with males and females differing only in
a single chromosome. In consequence, a termite mound has both male
and female workers. Only when blood ties are distorted, as in ants or
bees, does the burden of sterility fall on females alone. Unlike bees, in
which such behavior has evolved many times, all termites are social, so
that the habit started long ago, and stuck. Perhaps the strange pattern of
relatedness between mother and offspring in bees does make it easier for
a solitary animal to take up socialism when necessary.
Within a beehive all is not, as Aesop had it, sweetness and light. Not
only is there sterility but murder and cannibalism. Deplorable though
that seems, such acts help to understand the evolution of behavior. As
any stockbroker knows, kindness is hard to analyze. How does a donor
lose and a recipient gain? Greed is simpler. Often, losers forfeit every­
thing and winner takes all-which makes it easier to work out the bal­
ance sheet.
Cannibalism puts paid to the comfortable idea that Nature is not re­
ally red in tooth and claw. Thousands of animals go in for it and death
by fellows can be the main cause of mortality. Some fish eat 90 percent
of their peers. Walleye (a sport fish of North American lakes) eat each
other tail first, with whole chains of fish busy at a shared meal. The pas­
time has many excuses, and kinship can be one. For most cannibals,
blood is thicker than water. Given the choice, they keep themselves, their
relatives and their biological heritage alive at the expense of others. The
1 64 D A RW I N ' S G H O S T

losers, although they may not realize it, show the logic of martyrdom, of
death so that others might live.
Many animals assess the genetic cost of a slaughter of the innocents.
Often, males kill and eat the young of any unprotected female. A preg­
nant female mouse faced with a new male makes the best of a bad job­
the imminent death of her pups-with a preemptive strike. She
reabsorbs her fetuses, in a prenatal feast that gives the meal to her, rather
than to the hopeful male. Slugs show how costs and benefits depend on
kinship. Hermaphrodites though they are, sex often proceeds in a miti­
gated way, as each animal fertilizes the other. A box of outbred (and
hence unrelated) slugs turns into a loathsome soup as the survivors feed
on the corpses of the rest. Inbred animals, in contrast, the offspring of
self-fertilization, respect their own DNA held within their comrades and
refrain from that macabre feast.
Such rules can lead to murder in the beehive. Both queens and work­
ers invest in the next generation, as they lay eggs or help to raise larvae.
Because of the uneven relationship between the sexes, attitudes to the
young by the ruling and the working class are different. Two sisters share
three quarters of their genes; a brother only a quarter of his sister's her­
itage, through their maternal connection. For a worker (a sterile female),
the best strategy is hence to give three times as much attention to sisters
as to brothers. Then, her genes are transmitted with equal efficiency
through either sex. Queen bees, in contrast, are related to the same de­
gree to sons and to daughters. For them it makes sense to put the same
investment into each, as all the young pass on a mother's heritage with
equal efficiency.
In other words, within a colony, sons are worth more than brothers. It
is in the queen's interests for the hive to contain equal numbers of males
and females, but from the workers' point ofview the proportion of males
should never rise above one in four. Most of the time, the workers have
their way, because there are lots of them and they hold most of the
weapons. They kill and eat what they see as the excess among their broth­
ers. This increases their own biological chances and gives them a free
meal into the bargain. Life in a hive can become quite gothic. Unhappy
with attacks on their investment some queens fight back against worker
control and-with a certain ingenuity-give birth to sons who are saved
from murder because they masquerade as females.
 Instinct 1 65

The problem of sterility is solved when seen through the eyes of kin­
ship. But another great difficulty for the idea of gradual change through
natural selection lies in the workers' differing widely from both the males
and the fertile females in structure, yet they are absolutely sterile so that
they could never have transmitted successively acquired modifications of
structure or instinct to their progeny. The neuters of several ants differ,
not only from the fertile females and males, but from one another, some­
times to an almost incredible degree, and are thus divided into two or
even three castes. The castes, moreover, do not generally graduate into
one another, but are perfectly well defined. The honeypot ant of Ameri­
can deserts has workers whose simple job is to hang like great barrels of
sugared water from the ceiling, to be tapped when needed, while others
have castes with huge heads to block the nest entrance against intruders.
It may seem odd that sterile animals can evolve a variety of forms, but
this too comes from indirect natural selection.
The problem is no harder (or easier) to solve than the origin of soldiers
in aphids, or teeth in tigers. With all the genes of an organism the same
(albeit confined within a gall rather than a cloak of flesh) the focus shifts
from evolution to development: cells with the same DNA can make or­
gans as different as teeth and toenails, or plant suckers and soldiers. It is
all a matter of division of labor.
In bees or ants, the sterile castes do not share all their genes with their
mother. However, they have enough in common to enable the colony to
control-with as little conscious effort as the embryo of a tiger-the
growth of its members, using cues of place, time and expedience. Certain
ant castes have risky lives. In some, a twentieth die each hour as others
attack them; but the same is true of tiger skin cells as they are worn away.
Some castes vary in size, just like the teeth of tigers. Among the leaf­
cutter ants of South America are individuals that differ by three hundred
times in weight, from tiny fungus gardeners to massive soldiers. The sol­
diers, expensive as they are, do not appear until the colony is at least a
hundred thousand strong. Like those of a young cub, the first days of a
colony, with its shared inheritance, must be devoted to food rather than
defense. That is risky, but unavoidable. Not until her extended family is
well established will the daughters of the queen be enabled to grow into
her teeth and claws.
The beehive or anthill, a society that appears to cooperate, is in fact
1 66 D A R W I N ' S G H O S T

full of compromise and deceit; a place patrolled at all times by the ge­
netic police, who punish those who break its laws. They are microcosms
of evolution. Natural selection scrutinizes the value of every animal using
an exchange rate based on kinship, analyzes the market, and comes up
with whatever best transmits the DNA. Some individuals give up their
own future because it is contained within someone else, while others kill
to ensure that their heritage persists. Selfishness and charity are closer
than they seem. They follow biological rules which are, in the end, no
different from those that determined the evolution of bees or mole rats
themselves.

The ethical dilemmas posed by the ant or the cuckoo-so useful to sa­
vants who read human behavior into Nature-began long ago. The
Shakespearian fashion for society as a beehive has been succeeded by the
sociobiological fad of seeing it as a converted monkey house. George
Romanes, a friend of Darwin, made a fifty-point scale of intellect.
Worms were at step eighteen as they could "feel surprise or fear," and
dogs and apes at twenty-eight because of their "indefinite morality, along
with the capacity to experience shame, remorse, deceit and the ludi­
crous." Homo sapiens, in all its moral variety, occupied steps twenty-nine
to fifty.
The moral lessons of Nature, if such they are, began long before
mankind. Dinosaurs had societies as eccentric as those of birds today.
Some were cannibals: a two-hundred-million-year-old fossil has the
skeleton of a member of its own kind within its ribcage. Others were as
generous as the Seychelles warbler. Great herds of eighty or more giant
brontosaurs left tracks that reveal a life based, like that of elephants, on
a few adults and many young. Parental care started long ago. The pat­
terns in which fossil dinosaur eggs were laid show that some young were
fed by their parents, in great rookeries that lasted for millennia, while
others fled at once, to live a solitary existence. A few species laid eggs be­
neath mounds of rotten vegetation that warmed as it decayed. The mod­
ern birds-the megapodes-with the same habits have not changed
much from their dinosaur ancestors. Some make hillocks thirty feet
across, while others are happy with mounds of grass clippings. A few live
in huge colonies on volcanoes, their eggs hatched by the earth's heat. For
 Instinct 1 67

some megapodes, two brothers cooperate to help a female build her

mound. Dinosaurs might have been just as generous.
Although those great animals have gone, they followed the same rules
of instinct and habit as did their descendants. No doubt they could feel
surprise or fear and had, perhaps, some capacity for shame, remorse or
even deceit. We will never know: nobody will ever peer into the mind of
a dinosaur (or, for that matter, of a megapode) . Because the laws of de­
scent with modification act on behavior as much as on bones, the in­
stincts, if not the souls, of dinosaurs and all animals make as much
biological sense as does everything else in their lives: a tale told by varia­
tion, by struggle and by inherited differences in the ability of genes to
copy themselves.

Summary I have endeavoured briefly in this chapter to show that the mental
qualities of our domestic animals vary, and that the variations are inher­
ited. Still more briefly I have attempted to show that instincts vary
slightly in a state of nature. No one will dispute that instincts are of the
highest importance to each animal. Therefore I can see no difficulty,
under changing conditions of life, in natural selection accumulating
slight modifications of instinct to any extent, in any useful direction. In
some cases habit or use and disuse have probably come into play. I do
not pretend that the facts given in this chapter strengthen in any great
degree my theory; but none of the cases of difficulty, to the best of my
judgment, annihilate it. On the other hand, the fact that instincts are not
always absolutely perfect and are liable to mistakes;-that no instinct
has been produced for the exclusive good of other animals, but that each
animal takes advantage of the instincts of others;-that the canon in nat­
ural history, of "natura non facit saltum" is applicable to instincts as well
as to corporeal structure, and is plainly explicable on the foregoing
views, but is otherwise inexplicable-all tend to corroborate the theory
of natural selection.
This theory is, also, strengthened by some few other facts in regard to
instincts; as by that common case of closely allied, but certainly distinct,
species, when inhabiting distant parts of the world and living under con­
siderably different conditions of life, yet often retaining nearly the same
instincts. For instance, we can understand on the principle of inheri­
tance, how it is that the thrush of South America lines its nest with mud,
in the same peculiar manner as does our British thrush: how it is that the
1 68 D A R W I N ' S G H O S T

male wrens (Troglodytes) of North America, build "cocknests," to roost

in, like the males of our distinct Kitty-wrens, a habit wholly unlike that
of any other known bird. Finally, it may not be a logical deduction, but
to my imagination it is far more satisfactory to look at such instincts as
the young cuckoo ej ecting its foster-brothers, ants making slaves,-the
larvae of ichneumonidae feeding within the live bodies of caterpillars,
not as specially endowed or created instincts, but as small consequences
of one general law, leading to the advancement of all organic beings,
namely, multiply, vary, let the strongest live and the weakest die.
 C H A P T E R V I I I

HYBRIDISM

Sex and the marketplace - Laws of sterility - What keeps species apart? -
Natural selection against hybridism - The sterility of crosses, within and
between species - Cues of identity - Rules of estrangement, from brain
to gene - Sex and species: male ardor and female discretion - Fertility
of varieties when crossed - Laws of sterility and the genetics of species
barriers - Promiscuity and new kinds of plants - Breaking the fron­
tiers - The future of species

Adam and Eve fell into temptation when they ate the tomato of the tree
of knowledge. They did, that is, if one takes the European view of celes­
tial botany. For most of its nations, the tomato was such an exotic fruit
that it deserved a noble title. The French call it the pomme d'amour, the
Italians the pomo daro-the golden apple-and the Croats the paradis (a
name that explains itself) . Ridiculous, perhaps, but whatever Eden's fruit
may have been it was not an apple (which is rare in the Levant) and at
one time was painted as a tomato, an orange or a fig, fruits succeeded in
sacred art for a time by the banana.
Tomatoes, figs, oranges and bananas have different names because
they have different genes. They often grow near one another and have at
least the chance to mate, to exchange their DNA, and to merge into
some kind of amalgamated plant. They do not-but why not?
Sex, the trap into which the inhabitants of Eden fell, is a biological
marketplace. Every species is, more or less, a sexual republic. Within
each, open exchange is the rule and every inhabitant has a chance to
170 D A RW I N ' S G H O S T

barter its DNA with any other. The world as a whole is broken into fief­
doms-species-separated by barriers to genetic free trade. To under­
stand the origin of species we need to know what these frontiers are and
how they arise.
The differences between apples and tomatoes (or anything else) can be
measured from their DNA. Most of the changes so revealed have ap­
peared by simple descent with modification since an ancient break in
sexual relations or are relics of a shared and distant ancestor. They have
nothing to do with the barriers themselves. The real impediments to the
marriage of tomato and apple are genetic, but are impossible to test be­
cause the two never mate. That is Darwin's dilemma. It denies to biolo­
gists the main tool of their trade. Rather like nations who find it
impossible even to discuss their differences, a lack of intercourse between
species makes it hard to establish what keeps them apart.
The failure of tomatoes, apples and figs to cross is not some magical
property but comes from descent with modification. Somehow, a con­
tinuous process-natural selection-has given rise to discontinuous en­
tities called species. How it does so was once a mystery; but biology is
beginning to reveal the truth about what species are and how they arise.
Sterility itself cannot be selected for, as success would spell the auto­
matic end of the line involved. It can, nonetheless, emerge from other
evolved changes (as it does in bees, in which reproductive failure pays be­
cause it helps others to pass on DNA) . The sterility of species when
crossed also comes from indirect selection, in this case as a result of the
workings of evolution as it builds creatures able to cope with what the
environment throws at them.
Many genes are involved in that vital task. Within a species, in differ­
ent places selection may call for (or be offered) different mixtures of mu­
tations to do its job and may come up with subtly different products.
Quite often, one local blend does not combine well with others. Some­
times-as in the two inherited errors that jointly cause the smoky gray
fur of the Persian cat-the nature of the interaction is known, but more
often it is not. If the failure of adapted mixtures to work together be­
comes complete, the populations find it impossible to exchange genes
when they meet, and each becomes, in effect, a new species.
DNA's inability to copy itself without mistakes-mutation-means
that evolution is inevitable. Natural selection does no more than capital­
ize on that fact. Species, too, are by-products of the Mendelian machin-
 Hybridism 171

ery. They emerge from the apparatus of inheritance, from the ways that
genes join forces to do their job. If genes for size, shape or behavior work
together only in the right combinations, then, as different mixtures build
up in different places, the origin of species becomes inevitable. Once es­
tablished they may evolve further, but, of their nature, species happen by
accident.

Laws governing the sterility offirst Crosses and ofHybrids. The first line of
the United States Army Mule Training Manual is, it is said, "First, catch
the animal's attention by striking it smartly between the ears with a stout
stick." Mules, the offspring of a cross between a donkey and a horse, are
famous for their obstinacy, but are sterile and are, as a result, no threat to
the integrity of their parental species. In a few creatures, though, distinct
species can mate and may have fertile offspring when enticed to do so in
captivity. Ducks do it, geese are happy to exchange genes and pheasant
species have been hybridized to make a great variety of colors and forms.
The fact that the barriers among life's divisions can so easily be reversed
by man shows that they are not irrevocable. We must hence look at steril­
ity not as an indelible characteristic, but as one capable of being removed
by domestication.
Biological frontiers, plastic though they can be, have an identity that
extends beyond simple difference. Strange alloys can be made with in­
gredients from different creatures. Shoots of certain plants can be grafted
onto the roots of another and a pig's heart valve can be transferred, with
medical wizardry, to a man. Such mixtures may survive but are made
without benefit of sex. Their success does not much depend on the re­
latedness of those involved; a pig valve does better than one from a rab­
bit, although rabbits are more akin to ourselves.
The ease with which genes can be exchanged by mating, in contrast,
fits-like evolution itself--a hierarchy of biological change. Consensual
sex yields to grudging acceptance of a less than desirable mate, to rare li­
aisons with unlikely partners and at last to complete reproductive failure.
Only a small proportion of all species-perhaps one in a hundred-will
accept genes from another. There are, though, plenty of groups in which
sterility when crossed is so different in degree and graduates away so in­
sensibly, and the fertility of pure species is so easily affected by various
circumstances, that for all practical purposes it is most difficult to say
where perfect fertility ends and sterility begins.
1 72 D A R W I N ' S G H O S T

The dusky salamander lives in the southern Appalachian Mountains,

and likes to stay at home. It has been in its fastnesses for many years, each
population evolving to its own rules. Animals from within ten miles or
so mate promiscuously, but those offered the choice of a local partner or
one from far away almost never choose the stranger. Plants, too, may pre­
fer a neighbor to a distant individual who is adapted to other things (al­
though for them the choice involves the growth of pollen toward egg
rather than the rejection of an ardent swain).
Sometimes, simple choice fails, and hybrids appear. Some are sterile,
some so feeble that they die almost at once, while others do well for most
of the time. If they do so for long enough, one species may be absorbed
into another. Human interference is often to blame. John Constable,
when he painted the meadows running down to the River Stour, saw
plenty of poppies and thistles in his hayfields. The cows and sheep that
ate the fodder in winter had to cope with that less than nutritious part of
their diet. Nowadays, many of his fields are filled not with scarlet or blue
flowers, but with yellow: the blooms of poisonous plants. They descend,
in part, from plants introduced to the Oxford Botanic Garden from
Sicily in 1 794. Britain already had a native relative, the groundsel; and
the Oxford ragwort as it spread along the cinders of the new railways­
a place not unlike its native volcanoes-found a local partner with which
to mate. Two centuries later, the mongrels have found an identity of their
own and have become a pest. Many of Constable's great English oaks,
too, have been succeeded by hybrids between native plants and those
brought in from the Continent. ,Bastards as they might be, they seem
perfectly able to cope with their new home.
However well such hybrids may survive, and however stable they
might appear, they can fail Nature's test when life gets too hard. The is­
land of Daphne Major in the Galapagos has four kinds of ground
finches, the small, the medium and the large, together with the cactus
finch. It contains a hundred or so of each of the first two, a mere dozen
or so large ground finches and even fewer cactus finches. Their lives have
been followed for twenty years. These birds, the icon of all evolution
textbooks, show why species adapted to different ways of life stay sepa­
rate. It is because their hybrids do not survive the scrutiny of natural se­
lection.
The frontiers between finches can leak. About one union in fifty is be-
 Hybridism 1 73

tween birds of different kinds (and, on some islands, they mate with
birds not counted as Darwin's finches at all) . Such illegitimate behavior
is born of desperation. With a mere half dozen cactus finches on Daphne
Major, some are forced to turn to another species for any chance of sex.
Finches recognize one another by the shape of their beaks and by song.
Rather like cows and sheep, they learn who is who at their parents' knee.
This Freudian relationship can go wrong, as some males fail to pick up
their father's song (perhaps because their natal nest is close to that of a
noisy foreigner) and as a result court females of the wrong sort. Their
startled mates accept the alien and produce intermediate young.
At first sight, their progeny are no worse off than those of legitimate
unions and, in most years, survive and reproduce as well as anyone else.
If the four kinds of finch on Daphne were to carry on in this way, before
long they would fuse into an average bird. Fossils, however, show the
finches to have been distinct for thousands of years. Natural selection
against the crossbreeds has put a stop to the urge for union. It destroys
animals that do not fit into their evolved place in the economy and en­
sures that life stays in separate compartments.
Every few years, the Pacific winds change direction. No longer do they
blow warm water west toward Australia. Instead, tropical seawater moves
east toward South America and as far north as California. The current­
El Nino, the Christ Child-arrives around Christmas. It brings rain and
storms to the Americas and drought to Australia. The warm waters rise
by six inches along American coasts, and because they contain far less
oxygen and food than does the usual cold current, the numbers of fish
and of the birds that feed upon them collapse.
All this leads to inconvenience and more. The El Nino of 1 998 was
the strongest of the twentieth century. It caused twenty billion dollars'
worth of damage. Rain in East Africa swept away villages and farms.
Even Microsoft had to cancel its annual party because of storms along
the California coast.
The Galapagos in an El Nino has a huge increase in rainfall-by up to
seven times that of a normal year. The sea warms and loses much of its
nutrition. On the land, nature explodes. The plants are thick and lush,
the finches breed several times in succession, and hybridism flourishes.
As a result, many birds halfway between species appear.
As El Nino ebbs away, drought follows the torrential rain. Most of the
1 74 D A R W I N ' S G H O S T

finches die and, at once, selection sets to work. It reserves its greatest
punishment for the hybrids. For large ground finches the main item of
diet after a drought is a seed called the caltrop, after the spiked iron ball
used to trip horses in medieval battles. In such difficult times, finches
that have shifted down in size because they accepted genes from a lesser
relative find no food soft enough to eat, as it has already been taken by
an even smaller form. The offspring of matings with a larger species do
just as badly, as their beaks are not tough enough to manage what is left
by their more adapted relatives.
Excavations in Peru show that fishermen were washed out of their
campsites by an EI Nifio twelve thousand years ago, with major disasters
every few centuries since then. One struck in the 1 780s, leading to failed
harvests in Europe and to Marie Antoinette's dismissive comment: "Let
them eat cake." The hybrid finches' lesson about the baleful force of evo­
lution has-like that of the aristocracy-been repeated many times.
Each species has evolved to deal with life in separate compartments. The
sexual frontiers may leak when times are good, but, eventually, the wan­
dering genes are stopped by the onslaughts of natural selection on those
who bear them.

Causes ofthe sterility offirst Crosses and ofHybrids. Sex and species have a
lot in common. Asexual plants and animals are a classifier's nightmare,
because they exist as a range of forms, each grading into the next. Dan­
delions are divided into thousands of distinct kinds, fitted to where they
live and blended into an almost continuous series. Botanists often give
up and refer to such groups as "aggregates" or "complexes."
Sexual species are rather easier to define. They can be marked out in
genetic terms, as reproductive units, isolated from each other. Courtship
is a series of challenges and responses. Within a species, a male has at least
a chance of persuading his mate to accept his donation. However, a gene
that tries to pass between two species faces a sexual obstacle race in which
one barrier-on the first lap or at the finish-is impassable. The ob­
structions are those facing any uxorious male; but when a partner is of
the wrong kind they cannot be penetrated. They range from distance, to
learned preferences, to barriers embedded within the DNA. Each one in­
volves, in one way or another, a breakdown in the biological teamwork
needed to build a relationship, fertilize an egg, or produce a healthy off­
sprmg.
 Hybridism 1 75

Much of natural history is the scientific study of mating barriers and,

like bird-watching, soon degenerates into somewhat of a list. Species are
divided from each other in many ways-by space, by time, by mating
preference, by the inability to fertilize an egg or produce healthy young,
or by the sterility of offspring. The hurdles at which the sexual athletes
fall are as various as life itsel£ Those involved may never meet, or may
mate at their own special time or season. Males and females of different
kinds may choose not to pair, or may-with more or less enthusiasm­
mate but fail to make a fertile egg. The geographical checks can be as nar­
row as the few inches between different orchids upon which certain bees
feed or as wide as the ocean that separates American and European
species of gull. When it come to time, some Hies mate in the morning
and some in the evening and some crickets in the spring and others in
the autumn; but two kinds of cicadas in North America emerge and mate
every thirteen or every seventeen years. The difference ensures that they
almost never get together (in spite of a certain confusion every couple of
centuries).
Color, song, scent and more all play a part in settling who is, and who
is not, acceptable. The differences may be obvious to our own eyes or ears
(which is helpful to bird-watchers) but quite often they are not. Some
crickets make chirps attractive to their own kind and to ourselves, but
others retain their identity with sounds too low to be heard by humans.
More barriers await any hopeful sperm that has penetrated too far.
Some females mount an attack on sperm recognized as foreign, while
others, if mated both by a male of their own kind and an alien, much pre­
fer the local product. Plants do the same, with a refusal by a female to
allow foreign pollen tubes to grow through the wall of tissue that must
be breached before the egg is reached.
The cues may be on the sex cells themselves. The eggs of many marine
animals have a protective membrane into which a hole is punched by the
first successful sperm. Every kind has its own receptor. Each female
change is matched by a male equivalent. In abalones, the lock is a re­
peated sequence of DNA, which changes at great speed. It binds many
copies of the sperm's identity molecule. Although the seven abalones of
Californian waters look much the same, their egg receptors are quite dif­
ferent. The females evolve, the males are forced to follow them, and the
chances of inappropriate sex are much reduced.
The signals that separate species--color, scent or song-are often
1 76 D A R W I N ' S G H O S T

those used as statements of excellence by males in their struggle for fe­

males. For many plants, the process involves a third party, a pollinator
who must be persuaded to act as their brain and mobile sex organ. Bees,
flies, birds, lemurs and tree kangaroos are all pressed into service. The
plant strikes a bargain with its emissary. Pollinators are rewarded with
food, a place to sit (as in the lotus, whose metabolic heat keeps its flower
warm while beetles copulate within), or with a sexual signal (as in the
bees that pick up their own sex cues from orchids) . Much of this comes
from the need to avoid sexual ambiguity. Plants want their pollinators to
be busy, hungry and faithful, while the other party prefers to get as much
as it can from its host with the least effort and to spread its favors as
widely as possible.
Any failure to hybridize involves a loss of communication somewhere
in the biological conversation that is sex. As in any dialog, two individu­
als are involved (with, in the case of plants, a go-between); and if they
speak different languages things will not get far.
Sometimes, it is a matter of simple recognition. Many birds follow the
first object they see, be it parent or biologist, and take it as a role model.
Such creatures-ducks, doves and finches-tend to be divided into
many different kinds. For gull chicks, the colour of the spot on the
parental bill is the cue. When herring gull eggs are brought up by lesser
black-backed gulls-birds with red rather than yellow bill spots-the
young see themselves as members of the latter group and, when adult,
prefer to mate with black-backeds. Such boundaries turn more on the
ability to see who is who than on any deeper incompatibility. That may
be why birds differ so little in their genes, with the average genetic di­
vergence among relatives, from eagles to gulls, half that of mammals.
Fragile though they seem, boundaries held in the mind can be quite
secure. Some Mrican lakes contain hundreds of kinds of perch-like fish
called cichlids. The males are bright in color, the females dull. Each
species has its own signal. The transparent waters of some of the great
Mrican lakes allow the messages to flash from fish to fish and to keep
them separate. Although the frontiers are guarded by preference alone,
they allow dozens of different kinds to live together. Clear lakes have
more species than do those with muddy water. Each trademark diverges
most when in places with a danger of confusion, and a cichlid with red
males tends to share a section of shore with others whose males are blue.
Reds and blues put together in a tank stay apart. Females are faithful
 Hybridism 1 77

to their own-as long as they can tell who they are. In red light, which
makes it hard to separate the colors, a female will accept whatever male
courts her and confusion reigns. When the two parts of the incompati­
bility system-male signal and female choice-break down, what were
once species become, at best, varieties of the same one. That has been the
fate of many of the cichlids of Lake Victoria. As forests were felled and
farmers moved in, soil was washed into the lake. Victoria has lost her
clarity and her fish have become confused. Species that have been sepa­
rate for thousands of generations have, as a result, amalgamated into a
federation.
If a female cichlid (or anything else) chooses an inappropriate mate­
a feeble member of her own kind, or the wrong species-she pays a price
as her progeny do not have the correct genes. Males are less worried
about sex with an alien, as each act is so cheap that it is worth wasting
seed upon barren ground on the chance that, now and again, one suc­
ceeds.
The number of species within any group is, as a result, related to re­
productive habits. Where sexual selection rules, with lots of competition
among males and choice by females, speciation is rampant. There are
seven species of flowering plant to every one without flowers, because
each flower attracts a limited range of visitors and allows the plant to
keep its genes to itsel£ Ducks, with males and females very different be­
cause of sexual selection, have two dozen kinds in Britain alone, while
swans, in which pairs may bond for life, have far fewer. As was true for
President Coolidge's celebrated and insatiable rooster, male guppies will
mate repeatedly if presented with a succession of new females, but lose
interest when stuck with the same one.
When it comes to the origin of such species, females (with their
evolved ability to choose a mate) are, as a result, often in the driving seat.
When males take the wheel, the urge to split is reduced. A male let loose
on those who have not gained the ability to fight him off can often
breach the boundary. Take, as an example, the cuckoo. The common
cuckoo-the bird with the repetitive note, the "word of fear, Unpleasing
to a married ear"-is found from Japan to Ireland. The bird, widespread
as it is, has resisted the temptation to diverge into numerous kinds found
among many nest parasites. That is odd, given that cuckoo females are
divided into distinct races, each able to mimic the eggs of its own host.
The answer lies in cuckoo cuckoldry; in its male's insistence on sex
178 D A R W I N ' S G H O S T

with any female, whether or not she belongs to the race that brought him
up. Female cuckoos bear more allegiance to a particular host, be it red­
start or warbler, than do their males. As a result, a cuckoo who was him­
self brought up by warblers may father an egg found in a redstart nest.
Egg color itself is inherited down the female line, so that females stay
with the bird by whom they were fostered. As males are so much less
faithful, their promiscuity explains why the cuckoo has not split into
many species, each true to its dupe.
Africa, in contrast, has dozens of species of egg-dumping finches, each
tied to a single host, and quite unalike in appearance. Each species of par­
asite concentrates on a single victim, and the chicks mimic the bright
mouth markings, cheeps and movements of those of its unwitting step­
parent. The barriers are built in a finch's early days. The male finch learns
its foster-mother's song and this cue of identity is used by its own fe­
males, who will mate only with birds who sing the right one. Such forced
fidelity means that the finches have broken up into many species rather
than remaining as an entity bound together by masculine ardor.
When males are unleashed onto unsuspecting females, there may be
evolutionary mayhem. More than a million mallard ducks are released in
the United States each year to allow every hunter the kill that is an Amer­
ican's right. The male, with its green head and dark purple breast, is a for­
ward bird. Some female ducks, such as its close relative the pintail (whose
own male is quite distinct, with a long tail and chestnut head), have long
lived with it and can cope with its ardor. Although they can be prevailed
upon to interbreed in captivity, they never do so in the wild. Wherever
it goes (and it has traveled all over the world) , the male mallard makes its
intentions clear. The Australian black duck, the New Zealand gray, the
Florida mottled and the Hawaiian duck have all suffered from its ad­
vances. Each-once a separate species more distinct in its genes from the
mallard than is the pintail-has begun to hybridize with the invader.
Soon, they will be absorbed into it and disappear from the bird books.
This has led to calls for mass slaughter. Although activists have de­
nounced what they call "avian eugenics," the death of hybrids is backed
by the Royal Society for the Protection of Birds. Bird-watchers have an
austere view of existence: that which cannot be pigeonholed should be
shot. They ignore the evolutionary message hidden in the duck soup:
that the origin of species can result from the struggle between the sexes
as much as against the elements.
 Hybridism 1 79

The joint action of genes as they respond to the demands of selection

means that those who accept an incompatible partner pay a penalty. As
discord grows, it begins to pay each party to stick to its own kind as, if
negotiations are bound to fail, it is cheaper to break them off at once
rather than to go through the whole painful business of courtship. Hy­
bridism itself becomes a liability and the barriers begin to rise with an
impetus of their own.
Mines have grasses that-like the copper flower in Africa-grow in
places where their neighbors would shrivel. The grass responsible for the
scent of new-mown hay is among them. Its internal economy changes in
many ways as a result of metal tolerance and the mine populations can­
not compete except on the mines themselves. There they prevail as their
neighbors are poisoned. Grasses exchange genes with the help of the
wind. In a wild Welsh valley, an ancient lead mine is surrounded by un­
polluted pasture. As the wind howls over the workings it blows pollen
from the nontolerant population onto the mine, and a great cone of male
sex cells from the mine plants into the downwind meadow. Neither set
of genes does well except on its own patch. Selection has, as a result,
acted to reduce the amount of exchange. The mine plants flower a week
earlier than their neighbors, which limits the chance of either sort of
pollen finding a partner whose offspring would be unfit. The plants have
taken a first step toward a persona of their own. Should such discrimi­
nation continue, in time they might become quite unable to accept genes
from outside and would be, in one sense at least, a new species.
Monkey flowers, with their showy yellow blossoms, have gone further.
They have split into tolerant and nontolerant kinds. A cross between the
two gives young that sicken and die even on ordinary soil. Even popula­
tions from different mines may lose much of their tolerance when inter­
crossed. Although what is selected for is an ability to grow on poisoned
ground, sterility emerges as an incidental. When tested together the
genes give each plant a newly evolved identity.

Fertility ofVarieties when crossed, and oftheir Mongrel offiprings. In Berlin,

Friedrichstrasse meets Zimmerstrasse at a very ordinary road junction
across which traffic flows freely. A decade ago this was Checkpoint Char­
lie, one of the few gaps in an otherwise impenetrable barrier a hundred
miles long. Even a boundary as firmly defined as the Berlin Wall can,
with suitable manipulation, leak or collapse; and the same can happen to
1 80 D A R W I N ' S G H O S T

the frontiers between species. As a result, biologists are now beginning to

understand not just the displays that keep them separate, but the genes
responsible for their very existence. Again and again, it seems, new forms
have arisen because genetic systems that have, like the German Federal
and Democratic republics, faced different challenges and evolved differ­
ent ways of dealing with them are unhappy when brought together.
For species, as for nations, descent with modification builds obstacles,
almost as an afterthought, as populations diverge. As selection goes on
there emerge biological principalities that are reluctant to trade with
their former partners. Each contains its own set of genes that have never
been tested except in their own surroundings. The longer two societies
have been apart, the more they become fitted to local conditions and the
greater the chance of a mismatch. The greater the divisions forced by
time and circumstance, the less the chance of a new gene feeling at home
in the company of others. The process feeds on itselfand, soon, local en­
tities evolve toward the full statehood, which, when achieved, stops all
exchange between their citizens.
Fruit flies are evidence of how past distaste helps to guard a species'
virtue. For them, familiarity has bred contempt. How long ago two
species of fly last shared an ancestor can be worked out from how differ­
ent they have become in their DNA. Some pairs split not long ago, while
others have lived their own lives for millions of generations. When tested
in the laboratory, those with a recently established identity are more re­
luctant to mate with their relative if the two species live in the same place
than if they have never experienced each other's company except in a bot­
tle. For flies that live as neighbors natural selection against those who
made sexual mistakes meant that mild reproductive failure soon ripened
into real aversion. Only the blocks to mating itselF-and not to devel­
opment or to the fertility of hybrids-are involved. Those first barriers
in the obstacle course were, it seems, reinforced because they were so
often tested by hopeful males forcing their unwanted attentions on a re­
lated species.
The best place to track down what in the DNA is responsible for the
origin of species is among fruit flies. They will mate in vials and breed in
a couple of weeks. Their chromosomes can be seen through a micro­
scope, and thousands of genes have been tracked down. Even better, the
group has many species, some of which will cross in the laboratory. Their
 Hybridism 181

willingness to compromise their reproductive purity is a great help in un­

derstanding what keeps species apart.
The rules of inheritance hint at the truth. Male flies have a single
X chromosome, matched by a small Y. All genes on the X manifest their
presence in males, whether or not they are recessive in females. The same
is true in mammals, in which the male has an X and a separate Y chro­
mosome and the female two Xs. In butterflies and birds, in contrast, the
pattern is the other way round, as females are (in effect) XY and males
xx. Although DNA structure shows that the actual chromosomes in­
volved in the various groups have different origins, sex determination
works in much the same way, but in reverse.
There is a consistent pattern when two species are crossed. If, among
the offspring, one sex is absent or sterile, it is nearly always that with two
different kinds of sex chromosomes-males in mice or fruit flies, females
in birds or butterflies. Because different sexes are affected in each case,
the pattern does not exist because males are more sensitive about sex, but
must reflect some deeper biological truth.
All this supports the idea that a gene useful at home can be harmful
when mixed with others to whom it is a stranger. The wrong blend leads
to disaster, most of all when genes of one of the partners are exposed in
all their nakedness, as happens, in one sex, for those on the X. The ex­
posure of hidden genes to an unfamiliar set of companions is what
counts. Fruit flies can be manipulated to put foreign X chromosomes­
one from each of two species-in a single female. She is fertile, although
the lack of harmony of her alien X with her other chromosomes is no
greater than that of a hybrid (and sterile) male. However, in the experi­
mental female, the mismatched DNA is masked by her second X chro­
mosome, while in the males it is exposed to do its worst. Simple
dominance hides most of the genes that might be dangerous out of con­
text; but in male mammals (or female birds) their failure to get on with
the neighbors is revealed.
Several related species of fly have been hybridized in the laboratory in
the hope of chasing down what might be keeping them apart. The gen­
eral picture is clear: many genes are involved, each with a small effect on
its own, but each liable to a violent interaction when put into combina­
tion with others from its relative-just as expected if species are well­
suited communities of genes, easily disrupted by an intruder not used to
1 82 D A R W I N ' S G H O S T

their ways. Occasionally, hybrid sterility seems to need rather few

changes in the DNA, as the mongrel flies can be saved with a single gene
introduced into one of the parents. Quite how this snatches the progeny
from the jaws of death or childlessness is not known, but the ability of
one segment of DNA to do the job suggests that the block may be rather
simple. In certain plants, too, the avoidance of hybrids by pollinators can
be tracked down to a mere half-dozen genes or so, each responsible for
flower color, shape or scent.
Although many speciation genes have been found among animals, few
give any clue about how they work. Evolution, though, makes unlikely
bedfellows and can give birth to unexpected offspring. Part of the truth
about origin of species comes from research into cancer.
Cancer is a genetic disease of body cells. It arises from a failure of the
machinery that controls cell division. In the absence of brakes, the sys­
tem runs wild. It is not a single illness but many, each the result of a
change somewhere in a cascade of instructions that rules the life and
death of cells. Cell division, like the U.S. Constitution, has many checks
and balances. Some genes promote the process, others slow it down.
They exchange signals about when and where to act. An equilibrium be­
tween suppressors and promoters allows the body to grow and to sustain
itself, but if, for some reason, an enthusiast for division refuses to re­
spond to a command to stop, the result is uncontrolled growth.
Two tropical fish, the swordtail and the platyfish, are each decorated
with a series of attractive black spots. When put together in a tank, they
can hybridize (although they never do so in the wild). In the progeny of
a hybrid and its parent, the elegant spots turn into large and fatal black
lumps called melanomas. Their young inherit from one parent a gene for
a molecule promoting growth, and from the other a partner that should
keep it in check. In such fish, the second fails to suppress the first, too
much of the growth molecule is made, and cancer sets in. The two ele­
ments work in harmony when they have long lived together, but are
doomed to conflict when inherited in the wrong combination. The bat­
tle kills hybrids and keeps the species apart.
The genes for cell division did not evolve to separate different kinds of
fish. Their breakdown in hybrids is an incidental to evolution within
each one. The crucial genes have equivalents in humans and are of much
interest to medicine (which is why so much is known about them). They
also tell a tale about frontiers. Since swordtail and platyfish parted, each
 Hybridism 1 83

has diverged in the machinery of cell control. Two genes-each perfectly

at home in its own milieu, but lethal in combination-are enough to
keep them pure.
Plants work to rules rather different from those of animals. They are
more ready to accept a foreign mate, and, quite often, the offspring of
such a liaison find themselves with combinations of genes that fit so well
together that the new mixtures flourish, even at the expense of their par­
ents. Hybridization becomes a fast track to a new existence, rather than
(as in animals) a crack through which DNA leaks to dilute the prospects
of a hopeful species. A glance at chromosomes shows that many higher
plants-wheat and rice included-arose in this way, with complete sets
of genes from two ancestors that, long ago, got together, found that they
were able to work in harmony and flourished. Other plants arise by the
crossing of two kinds, which then reshuffle their heritage to make a new
one. A wild sunflower of the American West is, its molecules show, the
bastard progeny of two others. It looks somewhat different and lives in
drier places, but is in every other way a mixture of its parents. The an­
cestral genes stay in large blocks in the daughter form, as it arose too fast
for them to be broken up by sex before the new plant gained an identity.
Crosses between the parents in a greenhouse can make plants resembling
the new one in a mere five generations. Plant species can, it seems, orig­
inate at some speed, with no need for the long probation involved in new
forms of animal life. Their genes are more ready to cooperate than are
those of animals.

Hybrids and Mongrels compared, independently of their fertility. Mon­

grels-crosses between mere varieties-are uncertain beasts, mixtures of
the attributes of their parents. They attract little attention. Hybrids, the
results of matings between distinct species, have always been special;
enigmatic creatures ready to be explored by evolution. Aristotle believed
that a camel could mate with a panther to produce a giraffe, while ap­
pian, a more radical thinker, argued that to cross that animal with a spar­
row made an ostrich. That is fantasy, but the progress from mongrel to
hybrid and beyond now transcends the imagination of even botanists
and Greeks. It throws new light on the nature of species themselves.
Take the tomato. The fruit was once shunned because it was thought
to be poisonous, and the leaves are indeed filled with noxious stuff called
solanine. The Latin name Lycopersicon means "wolf-apple," and the
1 84 D A R W I N ' S G H O S T

plants were once grown for ornament alone (although Thomas Jefferson
is known to have eaten one). The crop has now increased to eight mil­
lion tons a year. The tomato seems a simple, not to say banal, item of
diet. It is more than that, as artificial selection has made a splendid vari­
ety of sizes, shapes, colors and tastes. Farm tomatoes differ among one
another more than do their wild relatives and, like farm animals, tell a
tale of how species arise.
The tomatoes (and there are many kinds) are natives of South Amer­
ica. They belong to a diverse and talented group of plants. Their relatives
include the deadly nightshade and the mandrake (known for its aphro­
disiac properties, its forked root of human form, and its lethal shriek
when uprooted) . Tobacco and the potato are not far away. Wild toma­
toes are sexual, but will accept genes only from a separate individual of
their own kind. The farm version is self-fertile, which places it behind a
barrier. If its own pollen is there, why accept an outsider?
The readiness of the tomato and its relatives to hybridize has long
been known. The great Kolreuter, whose accuracy has been confirmed by
every subsequent observer, proved the remarkable fact that one variety of
the common tobacco is more fertile, when crossed with a widely distinct
species, than are the other varieties. Crosses such as his have tracked
down just what prevents sex in the tomato family. They show that the
crucial shift to self-fertilization in the cultivated plant involves but a sin­
gle gene. It also controls the number, shape and size of the flowers. As
pollen success and flower form keep the plants apart, the barriers be­
tween tomato species may result from a simple interaction between this
gene and others.
The dozens of kinds of wild tomato can be divided, on the basis of
flower and fruit, into three large groups. Many flourish in places where
their domestic relative would die. Botanists have traveled from Chile to
the Galapagos in the search for new kinds able to resist their problems of
wilt, blight, mildew, rot, spot, speck, aphids, nematodes, cold, drought
and salt. On the farm, their genes would be useful indeed.
The U.S. Department of Agriculture has mounted an assault on the
tomato and (as an incidental) on the laws of hybridism. In greenhouse or
field, pollen and egg from wild tomatoes were tested for the ability to
cross with cultivated plants. Almost none of the thousands of attempts
succeeded. Most of the time, no plants emerged, and in the few that did,
 Hybridism 1 85

the progeny were sterile. In a very few, there appeared a fertile hybrid.
These rare individuals had, by chance, inherited an unusual mix from
each parent that, instead of the conflict that killed most of their kin, had
combinations of genes that were able to cooperate. A few of the genes,
detached &om their jealous fellows, are willing to work in an alien back­
ground. They are useful indeed.
Each fertile hybrid is a Checkpoint Charlie between sexual republics.
Through it, by means of a cross with one of its parents, DNA can pass.
Often the combinations fail, but dozens of variants that affect disease re­
sistance, fruit color and sugar content have now been moved from the
wild onto the farm. Many of the new genes mix and reshufRe quite hap­
pily with those of their domestic host, to produce even better mixtures
in later generations. Such limited free trade has already been a spectacu­
lar success. Selective breeding improves the yield of farm tomatoes by a
few percent a year, but wild genes can push it up by halE Even when a
wild plant seems feeble in comparison with its improved relative, it may
contain DNA that works wonders in its new home. Wild tomatoes are
green rather than red, but some have a powerful enzyme early in the pig­
ment pathway that, once placed in conjunction with domestic machin­
ery further down the production line, turns the fruits a brilliant scarlet.
Genes from different species, put together, transcend what either one can
do alone; given the chance they do not conflict to produce sterility, but
work together to escape from their uncooperative neighbors.
Botanical diplomacy has moved genes a long way. One group of wild
tomatoes is so distinct from the domestic kind that it refuses to cross
with it, however many attempts are made. It will, though, allow its genes
to enter another wild tomato. The intermediary can be induced to mate
with the farm variety. Its DNA moves down a chain of allies to a desti­
nation far from its native home. To an evolutionist, that is no great sur­
prise. All tomatoes share an ancestor and, by retracing their pedigree,
breeders do no more than reverse the history of their species.
Now, though, a real Common Market of biology has arrived. Frontiers
can be penetrated not with sex, but with technology. The nature of mon­
grels, hybrids and even species has been called into question. To fuse cells
of potatoes and tomatoes together in a bottle moves whole chromosomes
at a single step. The bastard cells give a new plant: a tomato-potato, never
seen before. Genetic engineers can do much more. With technical
1 86 D A R W I N ' S G H O S T

tricks-DNA attached to viruses or blasted into an alien home on tiny

bullets made of gold-they can separate genes from their native back­
ground and move them from anywhere to anywhere else. Sometimes the
deported DNA makes a protein, but more often-as evidence that genes
can work with others from their own, but not from alien, populations­
it does not. Even so, hybrids of the most unexpected kind are becoming
commonplace. They may be sterile but, in this new and open world, who
cares?
Tomatoes were pioneers in this libertarianism. Supermarket tomatoes
look good, but have no taste. Fruits ripened on the vine are tasty but soft
and difficult to transport. The Calgene company altered the gene re­
sponsible for maturation to allow them to remain on the bush and to
soak up sugar for a few crucial extra days. Now these Flavr Savr fruits can
be picked when they are red (rather than green) and taste as tomatoes are
supposed to taste. In 1 994, the u.S. Food and Drug Administration de­
fined them to be as safe as tomatoes bred by conventional means, in spite
of their acceptance of an alien piece of DNA. They did not even need a
special label, as the Flavr Savr "maintains the essential characteristics of
traditionally developed tomatoes." But is that true? Is this new fruit, with
its new heritage, even the same species as before?

One of the persistent myths of the Middle Ages was that of the Lamb of
Tartary, the Borametz. It was both an animal and a plant. The fruit of a
tree rather like a melon burst open to reveal a little lamb within, with a
fleece of surpassing whiteness. It was attached by its navel to a root in the
earth, around which it ranged until it had eaten all the grass, whereupon
it died. Erasmus Darwin himself hailed the creature that "eyes with mute
tenderness her distant dam, / And seems to bleat-a vegetable lamb."
Not even in medieval Tartary did sheep mate with melons (and the
Borametz was probably the "wool-bearing tree" of Plato, now known as
cotton) . Consider, though, today's tomato and its many relatives. Ge­
netic engineers have, they claim, inserted into them not just genes for
ripening, but one for an antifreeze, obtained from an Antarctic fish. The
plant should delight the most ardent free marketeer, as it marks the re­
moval of what seemed an impenetrable barrier to trade. Oscar Wilde was
fond of green carnations, but he had to depend on dye to achieve the
 Hybridism 1 87

right effect. Now the flower comes in a blue version, made by putting a
petunia gene into that normally pink, red or yellow plant. Other plants
contain genes from bacteria that give them resistance to insects, and scor­
pion toxins have been put into viruses to kill caterpillars. There may not
yet be vegetable lambs, but scientists have bred whole flocks of sheep
bearing human genes. Technology has broken the sex barrier; the Bo­
rametz is no longer the fantasy it was, and the distinction between mon­
grels and hybrids has gone forever. The engineers have globalized the
genes and have moved DNA to places it could once never have reached.
However, biotechnology, with its twenty-first-century powers, has an
eighteenth-century view of what species are. So confident are the techni­
cians of the safety of their products that each one is seen as no more than
an arbitrary mix of independent lengths of DNA. A protein that has
been proved to be harmless to man moved to a foreign plant used as food
does not even have to be tested in its new home: the new plant is, say
the authorities, "substantially equivalent" to the old. Their view takes
no account of the notion of species as interacting groups of genes, the
properties of one-as shown so starkly in the cancers of hybrid fish­
depending upon the others with which it is placed. With even greater
disregard for Darwinism, those who market unnaturally tasty tomatoes
or herbicide-resistant crops claim their new commodity to be distinct,
stable and free of exchange with others, so that DNA moved in can never
leak out. Public concerns are, they say, unreasonable: their modified
plants are safe and-in spite of the evidence of the readiness of plants to
hybridize-each species represents a Platonic ideal unable to exchange
genes with others.
The engineers deny the central facts of evolution: that the action of a
gene can depend on the species in which it finds itself and that all species
were once varieties (which means that the boundaries between many can
still be breached) . Already, genes for herbicide resistance put into oilseed
rape have seeped into wild mustards and radishes. Many other candi­
dates for manipulation-beets and rice included-are certain to mate
with their un domestic relatives as soon as they meet them and to pass an
alien heritage to unexpected places.
The experience of the tomato suggests that industry's optimism about
the nature and stability of species is unfounded. Those who cast down
the barriers to hybridism will soon be reminded of what evolution can
1 88 D A R W I N ' S G H O S T

do; and the very fact of their permeability is testimony of their origin in
descent through modification.

Summary of Chapter. First crosses between forms sufficiently distinct to be

ranked as species, and their hybrids, are very generally, but not univer­
sally, sterile. The sterility is of all degrees, and is often so slight that the
two most careful experimentalists who have ever lived, have come to di­
ametrically opposite conclusions in ranking forms by this test. The steril­
ity is innately variable in individuals of the same species, and is
eminently susceptible of favourable and unfavourable conditions. The
degree of sterility does not strictly follow systematic affinity, but is gov­
erned by several curious and complex laws. It is generally different, and
sometimes widely different, in reciprocal crosses between the same two
species. It is not always equal in degree in a first cross and in the hybrid
produced from this cross.
In the same manner as in grafting trees, the capacity of one species or
variety to take on another, is incidental on generally unknown differ­
ences in their vegetative systems, so in crossing, the greater or less fa­
cility of one species to unite with another, is incidental on unknown
differences in their reproductive systems. There is no more reason to
think that species have been specially endowed with various degrees of
sterility to prevent them crossing and blending in nature, than to think
that trees have been specially endowed with various and somewhat anal­
ogous degrees of difficulty in being grafted together in order to prevent
them becoming inarched in our forests.
The sterility of first crosses between pure species, which have their re­
productive systems perfect, seems to depend on several circumstances;
in some cases largely on the early death of the embryo. The sterility of
hybrids, which have their reproductive systems imperfect, and which
have had this system and their whole organization disturbed by being
compounded of two distinct species, seems closely allied to that sterility
which so frequently affects pure species, when their natural conditions
of life have been disturbed. This view is supported by a parallelism of
another kind;-namely, that the crossing of forms only slightly different
is favourable to the vigour and fertility of their offspring; and that slight
changes in the conditions of life are apparently favourable to the vigour
and fertility of all organic beings. It is not surprising that the degree of
difficulty in uniting two species, and the degree of sterility of their
hybrid-offspring should generally correspond, though due to distinct
 Hybridism 1 89

causes; for both depend on the amount of difference of some kind be­
tween the species which are crossed. Nor is it surprising that the facility
of effecting a first cross, the fertility of the hybrids produced, and the ca­
pacity of being grafted together-though this latter capacity evidently
depends on widely different circumstances-should all run, to a certain
extent, parallel with the systematic affinity of the forms which are sub­
jected to experiment; for systematic affinity attempts to express all kinds
of resemblance between all species.
First crosses between forms known to be varieties, or sufficiently alike
to be considered as varieties, and their mongrel offspring, are very gen­
erally, but not quite universally, fertile. Nor is this nearly general and
perfect fertility surprising, when we remember how liable we are to argue
in a circle with respect to varieties in a state of nature; and when we re­
member that the greater number of varieties have been produced under
domestication by the selection of mere external differences, and not of
differences in the reproductive system. In all other respects, excluding
fertility, there is a close general resemblance between hybrids and mon­
grels. Finally, then, the facts briefly given in this chapter do not seem to
me opposed to, but even rather to support the view, that there is no fun­
damental distinction between species and varieties.
 C H A P T E R I X

O N THE IMPERFECTION O F THE

GEOLO G I CAL REC O RD

On the absence of intermediates, today and as fossils - On decay - On the

vast lapse of time, as inferred from depos �tion and denudation, and in
other ways - On the power of water - On means of preservation - On
the sudden appearance of whole groups of species - On gaps in the
record as revealed by genes - On the Cambrian explosion and before -
On the lost history of ancient times - On the fate of the dead.

A murder victim becomes an ecosystem in his own right. Within hours

after a body is dumped in an English woodland, blowflies lay eggs
around its eyes and mouth. Soon, they are joined by flesh flies, which
drop larvae onto the skin. As maggots burrow in, gut bacteria work their
way out. The intestine bursts and its contents attack other organs. The
cadaver liquefies. It begins to ferment, with a strong smell of cheese.
This, in turn, attracts carnivorous beetles, and a fly called the cheese skip­
per (named from the larva's odd habit of jumping into the air when dis­
turbed) joins the fray. Three months after death, five hundred kinds of
insect may feast upon the corpse.
The sordid details depend on the weather and where the body is hid­
den. They are of interest to the police, as they make it possible to work
out the date of death. If blowfly larvae are still around, then the murder
was within the past week. Cheese skippers are evidence of an older crime.
Forensic entomology has solved many murders. A thirteenth-century
Chinese work, The Washing Away of Wrongs, describes how flies settled
on a certain sickle and proved it to be the weapon. Now, science has
 On the Imperfection of the Geological Record 191

joined the study of decay. At the Anthropological Research Facility in

Tennessee, cadavers are put in wire coffins above a concrete slab, or
buried at different depths, to relive the history of death. Corruption has
five stages-fresh, bloat, active rot, post-decay and dry remains. An in­
fant is reduced to a skeleton in a mere six summer days, while an adult
takes three weeks. The bones themselves can last much longer but, a few
years later, the chances are that not many will be left. The gravedigger in
Hamlet has it about right: "How long will a man lie i'th'earth ere he rot?
. . . He will last you some eight or nine year." For murder victims the re­
turn to dust is swift. The same is true for all corpses, human or not. As a
result, most of what evolution has made has moldered away.
Death, decay and dissolution all help to solve its greatest puzzle. The
fossil record-in defiance of Darwin's whole idea of gradual change­
often makes great leaps from one form to the next. Far from the display
of intermediates to be expected from slow advance through natural se­
lection, many species appear without warning, persist in fixed form, and
disappear, leaving no descendants. Geology assuredly does not reveal any
finely graduated organic chain, and this is the most obvious and gravest
objection which can be urged against the theory of evolution.
The unexpected chronicle of ancient life is not proof that the idea of
slow change is of its nature flawed. Much of the reason lies in the record
itself Few bones rest in peace and most of those who die decay almost at
once. Even the few that are preserved are soon washed away. As a result,
great periods leave no token and dramatic events stay unrecorded. The
archive of the rocks is a series of snapshots, taken at long intervals with a
badly focused camera. As in a Victorian photograph album, the past ap­
pears as a series of apparently unrelated images. Worse, these are them­
selves faded, torn, stained, lost and muddled by the passage of time. Even
a figure as prominent-and long-lived-as Charles Darwin himself has
left, a century or so after his passing, a mere dozen or so likenesses. The
lives of most of his contemporaries are quite unrecorded.
This dismal picture has a few exceptions. Geologists have unearthed
whole pages of the past. Everyone knows about the dinosaurs of the
American West and the fossil bird Archaeopteryx, in which even the de­
tails of feathers are retained. Two kinds of elephant exist today, but a
hundred and sixty-five species of their extinct relatives are preserved,
many as complete skeletons, to make a seamless set oflinks between past
1 92 D A R W I N ' S G H O S T

and present. The remains of our own ancestors-unknown at the time

of The Origin--are firm proof that we descend from apes. Any of those
bones is conclusive evidence of evolution and is in itself enough to de­
molish the creationist case. The smugness shown by students of living
creatures when they decry the gaps in our knowledge of those long gone
skates over our ignorance of the modern world, with, some say, only one
living species in a hundred yet described.
However, so persuasive is the fait accompli revealed by the relics of
prehistoric beings that it is easy to forget how little the record can say
about the machinery-rather than the pattern-of evolutionary change.
That leads to noisy (and much-publicized) disputes between students of
the ancient world and that of today. Most arise because each party asks
different questions of the past and is satisfied with different kinds of an­
swers. Not much is left of the great tree of life that fills with its dead and
broken branches the crust of the Earth. Today's creatures are its twigs,
and so few remnants of its limbs have been preserved that to work out
what controlled their growth when they were themselves still young may
be impossible.
Fossils have long been accepted as messages from history. The ancients
saw shark teeth as the petrified tongues of snakes fallen from the sky, and
amber as congealed wildcat urine. The curled fossils called ammonites
(named after Ammon, an Egyptian god with ram's horns) were thought
to be snakes decapitated by a saint and turned to stone. They were sold
as curios with carved heads attached.
Ammonites (relatives of today's squids and nautiluses) are not petri­
fied snakes, but those who found them could make no other tie to the
animals of their own day. Their sellers recognized that to link the past
with the present the details must be filled in. That is still true. The num­
ber of intermediate and transitional links, between all living and extinct
species, must have been inconceivably great. Almost none are found (al­
though, as in the elephants, or in the ancient snakes with legs, those
silent few are eloquent indeed) . Where have they all gone?
Their fate is a matter of time and chance. In the Old Testament, the
Song of Deborah tells that "the mountains melted from before the Lord."
In biblical days her vivid image was a statement that God is indestruc­
tible and that to Him even the landscape does not last. On evolution's
span, the ground is a turbulent sea and the dead are subjects of its uneasy
 On the Imperfection of the Geological Record 1 93

movements. Their remains disappear at once, or are borne many miles

from where they fell. They may sink below the surface to rise again, and
be churned up with the relics of animals who expired far away and long
ago. Most of the evidence has gone forever. Those who hope to use the
fragments of the past as the key to the workings of the present will, as a
result, almost always fail.

On the lapse of Time. The passage of time, so great as to be quite inap­

preciable by the human intellect, disturbs all graves. Time once seemed
a domestic thing. Deborah lived in the twelfth century before Christ,
quite soon after what the writers of the Old Testament saw as the start of
the universe. The passing of the ages could be measured out in just one
way, in lifetimes. To count the scriptural generations (as listed in the
margin of many Bibles, Darwin's included) would date the Earth. Arch­
bishop Ussher's estimate for its origin was, famously, 4004 B . C . (which
means that, for real believers, the sixth millennium passed in 1 997).
When John Burgon described Petra as a city "half as old as time," he
meant it.
Even those who dismissed such notions could not imagine how far
they would have to go in search of yesterday. The great French naturalist
Georges Cuvier noted that the five-thousand-year-old ibis mummies
brought back by Napoleon from Egypt were identical to their fellows
alive today. That, he felt, showed that animals could not change, as five
millennia seemed to him to encompass most of the Earth's history.
Copernicus and Galileo put an end to the idea that space could be
measured in human terms. The geologist James Hutton did the same for
time. In 1 788, on a visit to Siccar Point near Edinburgh, he realized that
the rocks were far more ancient than anything contemplated by theol­
ogy. A layer of sandstone rested on the ground-down and tilted edge of
a still older bed of slate and grit-which must have been deposited and
worn away long before. "The mind," he wrote, "began to grow giddy by
looking so far into the abyss of time." The Earth, it became evident, was
old: it had "no vestige of a beginning-no prospect of an end." Since his
day, geology has made it clear how incomprehensibly vast has been the
span of the years.
The notion of ancient landscapes shaped by unhurried change was at
first denied. Henry de la Beche, founder of the Geological Survey of
1 94 D A R W I N ' S G H O S T

Great Britain, circulated a sketch called "Cause and Effect" in which a

baby urinates at the head of a gorge and the products of his labor flow
down the middle. His nurse comments: "Bless the baby! What a walley
he have a-made!" The belief that species were immutable productions
was almost unavoidable as long as the history of the world was thought
to be of short duration. Only if there has been enough time for slow
change can Nature mold its products into novel forms by gradual selec­
tion, without the need to make leaps.
How old are the Earth's dead inhabitants? Because their rest has been
so much disturbed, it is difficult even to sort out the sequence in which
they perished, let alone the date of their demise.
The first job is to put the layers in order. The task seems easy: those on
top are young, and those buried deep are the oldest. However, our planet
is stoked by fires and from the start has suffered from indigestion. What
were its guts are on its surface and what was once its skin has been driven
into its depth. Because the Earth was built from within, its rocks were
lifted or buried, its beds turned over and their order reversed. The Lizard
Peninsula, in the south of Cornwall, is a piece of its molten interior
squirted to the surface. It has no match anywhere in Europe. Even worse,
modern material is often worn away to show the ancient strata below.
The first geological map of Britain, made in 1 8 1 5, managed to trace only
one thin seam of limestone called the Cornbrash across southern Eng­
land. Now, after years of exploration, the logic of the whole world has
been laid bare. Layers of material can be linked with others far away, to
rebuild the globe.
A glance at any geological map shows how much has been lost. Cer­
tain strata can be followed for thousands of miles. In a few places, the
beds are almost as thick as when they first settled, but in others they have
been worn thin or have gone altogether. If the maximum depth of each
separate layer across the globe is added together, our planet should be
cloaked in sedimentary rock a hundred miles thick. Most of its coat is a
thousand times thinner because a history written in sand has been torn
to pieces, its fossils with it.
Time can be measured only by change, in the flesh of a decaying
corpse, in the tension in a watch spring, or in the landscape itself The
first attempt to age our planet (rather than just to order the pages of its
biography) compared the saltiness of the sea with that of the rivers feed-
 On the Imperfection of the Geological Record 195

ing it. If the sea was once fresh, and its salt came from the land, it should
be possible to work out how long the ocean had taken to become saline.
The estimate-of a hundred million years-was a deathblow to those
who measured time in human terms. It was the first hint of the unimag­
ined antiquity of life.
It was far too low, because salt is laid down in great deposits on the sea
bed and is returned to the land as it rises. We live not a hundred million
but four thousand five hundred million years since the Earth was spun
from dust and rock around the sun. The evidence comes not from our
own planet, but from its satellite. The Moon Hew off its parent after a
giant impact. Because it stayed small, cold and undisturbed it gives a bet­
ter picture of the past than does its parent. A quick trip by the Apollo XI
mission was enough to date it. The Earth's turmoil makes it harder to
trace its own origin. Its oldest rocks, found in Greenland and Western
Australia, are just under four billion years old.
For living things, the answer to time lies not with deposition, but with
decay, and not of murder victims, but of atoms. Radioactive materials
throw off parts of their structure and, as they do, change into other forms
of matter. How long it takes is measured by the "half-life"-the period
needed to convert half the original into its derivative. This varies from
the four and a half billion years needed to change uranium-238 (the ra­
dioactive version of the element) to lead, to the twelve years or so for the
shift of tritium (a form of hydrogen) to helium.
Some chemicals are useful in dating the past. Potassium has a half-life
of rather more than a billion years as it breaks down to argon. A com­
parison of daughter with parent says when the decay began. The inactive
gas produced as the element breaks down shows when the rocks were
made (or at least when they were last melted and the gas driven off).
Carbon- l 4-a radioactive form of the atom upon which life is based­
has, in contrast, a half-life of a mere five thousand seven hundred and
thirty years. It is made when nitrogen is bombarded by cosmic rays, high
in the atmosphere. Plants and animals all pick up traces of carbon- 1 4,
which begin at once to decay. When the animals themselves die, the rare
form of the element is no longer replaced and the proportion left is a key
to the time of their demise.
To explore the past it helps to have benchmarks. Volcanoes help be­
cause their ash blankets the ground to give layers that can be read as a se-
1 96 D A R W I N ' S G H O S T

ries of pages upon which history is written. The explosion of Mount St.
Helens in 1 980 generated so much debris that the Columbia River was
blocked and ships were trapped upstream. The cloud of particles circled
the globe. Fossils are mixed into the ash itself, or are trapped in the lay­
ers of rock between each band. Some of the first evidence of our own an­
cestors is in a line of two-million-year-old footprints left as two upright
primates strolled across an African plain coated with soft volcanic dust,
stopped, and looked to the east. Lake beds, too, are blank sheets upon
which the past is recorded. Each spring, they accumulate a layer of silt.
The Green River beds of Wyoming contain twenty million of these an­
nual bands. They allow its fossils to be ordered with great accuracy.
Such evidence has aged the Earth. Its many layers have been named
(some-the Cambrian, the Ordovician, and the Silurian-after Wales
and its ancient tribes) and dated. Eighty percent of history is in the Pre­
cambrian, an era once thought to be barren. The sixth or so of the past
into which most fossils are crowded itself represents a lapse of time im­
possible to imagine in human terms. Even much briefer periods are hard
to comprehend. Fifteen million years ago, life on Earth was much as it is
today; but fifteen million years hence, because of the imperceptible creep
of coastal California along the San Andreas fault, the Santa Monica Free­
way in Los Angeles will run directly into the San Francisco Bay Bridge as
the City of Angels migrates northwards.
A scheme is under way to build a clock-the Clock of the Long
Now-in a remote desert, driven by a twisting pendulum powered by
the expansion and contraction of a metal bar as night follows day. The
hands will move once every year, it will chime once a century, and every
thousand years a cuckoo will pop out. To geology, that clock is a mere
stopwatch, and the Long Now but an instant in the narrative of our
planet.

On the poorness of our Paleontological collections. The Natural History

Museum in London owns six million fossils of marine snails. It seems an
impressive collection, bur is not. A square yard of sea floor can accumu­
late a hundred of those unpretentious animals a year, and within any ge­
ological period-some tens of millions of years-could make enough
shells to swamp all the world's museums. The crust of the Earth is itself
a gigantic depository, but its collections have been made only at intervals
 On the Imperfection of the Geological Record 1 97

of time immensely remote. They are a reminder o f how many of the

products of evolution have been lost. Any single place retains mere frag­
ments of the past, either a large part of a small time or, more often, a few
glimpses of a much longer period.
The greatest destructive force is water. American builders shift eight
billion tons of earth a year. The rain does much more. It consumes the
countryside and planes great mountains smooth. Water destroys, but it
also builds. As it does, it edits the record of the past.
At the time of The Origin the California Gold Rush was in full swing.
In its first days, miners were content to use technology not very different
from that of the Greeks and the Golden Fleece to trap small particles of
gold from rivers; not with animal skins suspended in the stream to filter
out the metal but by running a trickle of water over the gravel in a pan.
A good miner could work through a cubic yard of pebbles a day and be­
come modestly rich from his efforts.
In 1 853, Edward E. Matteson had the idea of using high-pressure
water from the mountains to wash out the gold. It was a spectacular suc­
cess. Two men could break up fifteen hundred tons of ore a day. The
Nevada City Transcript declared that the miners had "broken into the in­
nermost caves of the gnomes, snatched their imprisoned treasures, and
poured them, in golden showers, into the lap of civilized humanity."
Matteson's nozzles, sixteen feet long, were called the Dictator, the Mon­
itor or the Giant. In five years the miners built five thousand miles of
ditches and flumes to feed them. By the end of the boom (when Matte­
son died bankrupt), twenty-five million ounces of gold had been hosed
from the gravels of California.
As the dirt ran downstream, it covered thousands of acres of farmland
and blocked San Francisco Bay. Within five years, a century's worth of
sediment was washed from the mountains and laid down in the plains.
Even today, mile-wide gullies in the Sierra Nevada are monuments to a
single decade in which man used a tiny part of Nature's energies. Now,
the miners have moved to Alaska, and geologists comb through the de­
tritus blasted from the rocks by today's Monitors and Giants in a search
for the bones of extinct mammals.
The land is also demolished as the sea pounds it. The Japanese are des­
perate to save the remains of their distant and decayed island of Okino­
torishima. At high water it consists of two lumps of coral, each fifteen
1 98 D A R W I N ' S G H O S T

feet across. Those tiny fragments allow Japan to claim a coastal economic
zone of a hundred and fifty thousand square miles, with all its fish and
minerals. They have spent millions of dollars to build a steel-reinforced
concrete wall around the rocks (not in touch with them, as it would then
be classed as artificial and could not qualify as a territorial claim) . The
force of the waves means that the task is hopeless. Soon, Japan will lose
another great swath ofwhat was once its empire; and another page of fos­
sil history will be lost from view.
Water shows its real force when it turns to ice. The California miners
soon learned to pour water into cracks high in the mountains in daytime
and to let it freeze at night. The stress on the rock as it expanded was that
of a sledgehammer swung with the force of ten men. Great blocks of ore
could be shattered with a bucket dipped into a stream.
The rain can melt rock as well as smash it. The granite columns of
Egyptian temples lie in the desert, their inscriptions almost as sharp as
when they were cut. Roll a column over and it is blank. Damp in the soil
has reacted with the stone, turned it to clay, and wiped off its message.
In 1879, two obelisks that had stood for three thousand years in the ruins
of Thebes were shipped to New York and to London to allow the public
to admire their hieroglyphs. A century later, the New York stone is un­
adorned (although its London twin, Cleopatra's Needle, has survived
rather better).
The Pyramids were time machines, designed to project the ego of a
Pharaoh into the future. They are a better monument to how fragile even
stone can be. Their place in geological history is secure, as Herodotus
identified the ancient shells found in the rock from which they are made
as the remains of lentils discarded by their builders (Voltaire, in turn,
thought that fossil fish in the Alps were the remnants of pilgrims' packed
lunches). Their structure is less safe. The earliest tombs, those at Mem­
phis, were built of brick and have long gone. They were succeeded by the
Pyramids themselves, the first built about four and a half thousand years
ago. Until the time of William the Conqueror, many were covered with
polished limestone that shed the rain as it fell. Their covers were removed
to build mosques in Cairo, and water began to soak into their structure.
Many are now mere stumps, covered by the debris of their own destruc­
tion. The Great Pyramid has lost sixty thousand cubic yards of mater­
ial-and its innumerable fossils-since its shield was removed.
 On the Imperfection of the Geological Record 1 99

The Pyramids are young compared to the hills from which they were
quarried. Those seem almost timeless; but that, too, is an illusion. Most
mountains arise as the plates of the Earth's crust strike one another. The
Himalayas began a mere fifty million years ago-after the death of the
dinosaurs-as India crashed, at a foot a year, into Asia. Mountains de­
termine their own fate because they make their own climate. Because it
faces the wind and the rain, the Indian slope of the Himalayas is gashed
by huge valleys like the Khumbu Ice Fall, up which the first climbers
struggled, while Tibet-half the size of the United States and most of it
higher than America's highest point-is dry, Hat and cold. With no rain,
it has lost less of its substance.
Mountains float, like icebergs, upon the mantle of the Earth, with
most of their mass miles below the surface. They are buoyed up in a sea
oflighter material, their summits renewed from below. This makes them
older but less sturdy than they seem. As glaciers grind millions of tons
from their path, the peaks thrust even higher. The Appalachians are a
foot taller than when first seen by Europeans because so much soil has
been lost from the valleys; the mountains more or less unchanged in
shape, but their material washed away and replaced from within.
In the end, the rain always wins. Great mountains are popular, but
rare. The mean height of the land across the world is two and a half thou­
sand feet and most of it is Hat. The turmoil of the peaks means that those
who die there will soon be gone. Australia is the most exhausted country
of all, its low hills in the last stage of decay. Its fossils include those of
alpine plants, remnants of a more elevated past. The rest of antipodean
history has been swept into the sea. Sooner or later, the Himalayas, too,
will be gone. Their inhabitants will follow.
The remnants move downstream to the ocean. A slow river carries
more earth than it does water. Every stream is an engineer. Its silt settles
into sediment and forms the rocks that cover three quarters of the globe.
In their turn, these are disturbed by new downpours. As it falls, the mud
builds new soil. It was explained to Darwin in South America that the
bones of giant sloths deep below the surface proved how, in an earlier age,
they lived in holes. The idea is seductive, but the truth is simpler. Given
long enough, mountains-and plains-How, to use the biblical
metaphor, like rivers. They take their contents, vandalize them, and bury
them in the refuse. That is stirred up and reburied as gain follows loss.
200 D A R W I N ' S G H O S T

As the remains of a shattered landscape pour downhill, the globe's sur­

face is reshuffied.
The Aswan Dam was completed in 1 970, and Lake Nasser began to
fill behind it. The Nile slowed down and stopped. As it did, it dumped
its load of sand, silt and corpses onto Nasser's Hoor. Below the dam, the
pure water ran faster than before and picked up more silt. It scoured the
stream bed, with the loss of five hundred bridges and of many miles of
the Delta. To build the dam disturbed the balance of deposit and with­
drawal in the mud bank.
Cairo sits on a huge rift-bigger than the Grand Canyon-which was
cut and then filled with sediment by an ancient Nile. The Grand Canyon
itself began at about the same time, as the Colorado River sawed through
the Earth's crust. Around it the land was uplifted to give the mile-high
cliffs of today as the river cut through their soft rocks. Like those of the
Nile, the Colorado's waters are valuable. Thirty years ago, the Glen
Canyon Dam domesticated the river to feed Los Angeles. Its lake, like all
others, has begun to fill with mud. Below it, where once were sandbanks
and rapids, are thickets of mesquite and a channel clogged with boulders.
Concerned by the changes, ecologists opened the dam for a week to feign
a Hood. Millions of gallons of water poured out and the river rose by
twenty feet. The effects were dramatic. Huge boulders shifted for hun­
dreds of yards as fans of sand were destroyed and re-formed as fresh
beaches. The Canyon was born again after decades of decline. Now the
Sierra Club is calling for its two-hundred-mile-long lake to be drained to
allow Nature a permanent return.
Among the debris were thousands of bones, any of which might in
time have become a fossil. They were churned by the torrent into a soup,
to settle as a new and arbitrary mix. A week of Hood muddled the evi­
dence of thirty years. In the Grand Canyon's long history, such deluges
must have happened uncounted times.
Science can be magnificent in its simplicity. How better to test the
movements of the uneasy Earth than to simulate the fate of a giant sloth,
dead fifty thousand years? Take the bones of slaughtered cows, mark, and
scatter around the landscape. Leave for a decade or so, and return to see
how many remain, where they might be and what company they keep.
The experiment was carried out around the famous fossil sites of the East
Fork River in Wyoming. Its results were clear. Most of the bones have
 On the Imperfection of the Geological Record 201

gone forever. Many o f the survivors have moved and are buried deep in
sediment. The remains of cows that expired in the 1 970s mingle with
those of animals last seen ten thousand years before. A geologist yet un­
born would have a strange image of today's America, with herds of cattle
pastured among the mammoths.
Sometimes, life adds to the confusion of the landscape. In a few places
the shells of sea snails are mixed with the bones of land mammals and
birds. The snails were picked up by ancient seabirds who dived for food
and left the fragments around their nests. Even the famous human fos­
sil, the Taung Child, is so mixed with the bones of other animals that its
remains may have been taken by a bird of prey to feed its young.
The turmoil of the rocks means that fossils are not laid down in neat
sequence in an ordered world. Instead, they fall into a universe of change.
The passenger pigeon flourished in North America as late as the Civil
War. At the time of the Mayflower, nine billion were alive-more than all
the birds of America today. A single flock was said to have been a mile
wide and two hundred long. So stupid was the bird and so ruthless its
pursuers that the last, a female called Martha, died in Cincinnati Zoo in
1 9 14. Nobody has ever seen the fossilized bones of a passenger pigeon.
Without a written record we would never know of its existence. Untold
numbers of other beings have accompanied it into oblivion. Like the
products of past labor, too much has been made for it all to be preserved.
The hope of survival for any corpse depends on what its owner was,
and how and where it died. Some places give the dead a chance, but oth­
ers are less kind. Almost all fossils are mere replicas of animals and not
their flesh. Their bones or shells are transformed after death as they be­
come saturated by minerals and turn to stone. The image, not the real
thing, is saved, and must be petrified at some speed if it is to have a
chance. In Victorian times stone bowler hats were a popular souvenir of
a visit to a cave. All that is needed is to drip limestone-rich water onto a
hat for a few months and its shape is preserved. Already, half a century
after it went out of fashion, almost as many survivors of that strange
headwear are preserved in stone as in felt. A thousand years hence there
will be many more.
An animal stifled in mud will last better than its cousin destroyed by
fire, and a cadaver has more hope in a chalky sea than an acid marsh. One
of the oldest of all fossil beds, the Burgess Shale, was buried half a billion
202 D A R W I N ' S G H O S T

years ago. The animals were infiltrated by soft clay, which made perfect
images of parts of their bodies. Whole communities were embalmed in
a fall of mud onto the sea bottom after a flood. Archaeopteryx, in contrast,
died on land. It was swept into an anoxic sea, where it was conserved in
exquisite detail, along with many of the plants and animals living along­
side it.
Sometimes a triumph of the frail allows the animal itself to survive.
Human remains in peat bogs retain the hair, clothes and pained expres­
sion of their original owners. As the flesh was tanned by acids in the soil,
the bones were dissolved to leave mere bags of skin. Animals can topple
into lakes of pitch such as the La Brea Tar Pits of Los Angeles or stick to
tree gum that turns into amber. All the household means against decay­
deep frozen mammoths, Spanish woolly rhinos marinated in a salty
swamp, ground sloths in South American caves reduced to beef jerky by
slow evaporation-help to conserve these Mona Lisas of the fossil record.
Some animals have more hope of immortality than do others. A crab
has a better chance of a monument than does the sea anemone that grows
upon it. Worms and jellyfish rot at once, while snails, corals and the like
have better prospects. Two thirds of marine animals have soft bodies. Al­
most none are known as fossils. A solitary ancient specimen of krill (the
food of great whales) has been found, although some shoals contain a
hundred million tons of the tiny creatures, the weight of the world's an­
nual fish catch. The survivor had been eaten by a fish and fossilized
within it.
In Klagenfurt, in Austria, is a stone fountain carved in 1 590. It depicts
a beast mythical in that it has wings and breathes fire. The head looks
strangely like that of the ancient European rhinoceros. The sculptor, it is
said, based it on fossils found around the town. The Klagenfurt fountain
is one of many reconstructions of the past in which artistic imagination
takes precedence over scientific fact. The Burgess Shale remains were so
decayed and scattered that single animals were at first identified as sev­
eral. One made a simultaneous entry into science as a jellyfish, a sea
squirt and a beast rather like a crab. Not until a complete specimen was
found was it recognized as a being quite unlike any alive today and
named as Anomalocaris.
The La Brea Tar Pits or the German quarry where the remains of Ar­
chaeopteryx were found are great museums of ancestors. Both places, like
 On the Imperfection of the Geological Record 203

the Louvre or the Hermitage, hold but a few moments of evolutionary

effort. Just seven skeletons of Archaeopteryx (plus an isolated feather)
have ever been found. As in a museum, the accidents of preservation give
a biased view of the past, with too many gold ornaments and too few clay
pots. The accuracy of the collection can be checked against those alive
today. How many of each modern kind are found as fossils?
Sometimes, the assembly of the dead is not much different from that
of those who survived. In California, where the sea floor has risen to
make terraces along the coast, two thirds of the marine snails on the
modern shoreline are in the fossil beds which hence gives a reasonable
image of how things used to be. In most places, though, the dead are not
an accurate sample of the living. Three quarters of the large mammals in
the savanna of East Mrica leave their bones for at least a year, but smaller
beasts disappear almost at once. Although living shrews are far com­
moner than their giant neighbors, the fossils do not reflect reality; the
bones of elephants scattered around the Mrican landscape are overrepre­
sented by five hundred times compared to those of shrews. Museums are
short of fossils not because their curators are lazy but because they have
set themselves such a difficult task. What they have is a glimpse of the
unexpected worlds of the past, and too often one so fragmented that it
says little about what those ancient universes must really have been like.

On the sudden appearance of whole groups ofAllied Species. The abrupt

manner in which whole groups suddenly appear in certain formations
has been urged as a fatal objection to the belief in the transmutation of
species. If numerous species, belonging to the same genera or families,
did start into life all at once, the fact would be a severe blow to the the­
ory of descent with slow modification.
Students of ancient barnacles, bony fish, mammals and birds have
each, at one time or another, proclaimed their favorite animal's abrupt
arrival in the rocks as evidence that gradual change is not enough to ac­
count for its first appearance. All have been proved wrong by finds of
forms much older than the supposed pioneers. Now, genes-descend­
ing, as they do, from a past more ancient than any fossil-can help fill
gaps in the record. Quite often, they show how a sudden emergence may
hide years of obscurity in the wings. We continually overrate the perfec­
tion of the geological record and falsely infer, because certain genera or
204 D A R W I N ' S G H O S T

families have not been found beneath a certain stage, that they did not
exist before that stage.
Archaeopteryx was discovered just two years after the publication of
The Origin. As the first "missing link" it caused a sensation and was at
once seized upon as proof that birds must have arisen in a single step. In­
deed, its descendants, today's birds, do not appear as a well-established
group until after the demise of the dinosaurs, millions of years later.
However, Archaeopteryx is not as unique as it seems. New finds show
from whence it came and where it went. Its direct ancestor was a small
meat-eater that, like Velociraptor (famous for its role in the film Jurassic
Park), ran upright on two legs. Until the late 1 990s, the record between
those land-bound animals and Archaeopteryx itself consisted of an enor­
mous gap. In 1 997 an intermediate was found in Patagonia. It cemented
the tie between birds and dinosaurs. Unenlagia ("half-bird," in a local
language) lived ninety million years ago, long after its famous relative. As
its shoulder joint pointed not downward but to the side, Unenlagia, al­
beit earthbound for most of the time and without feathers, could move
its forelimb with a full upstroke.
Soon after the discovery of Unenlagia, a turkey-sized, carnivorous (and
emphatically flightless) dinosaur with short arms was found. It had un­
mistakable feathers, barbs and all, fanning from its tail. In the absence of
wings, they could not be used for flight, and may have been courtship
displays or even insulation for the animal's rear end. The tail was short
and fused to the pelvis, a feature that would later be useful in the air. In
almost every other respect the animal is a dinosaur. Birds can no longer
claim sole ownership even of feathers, let alone of an identity that sprang
fully formed into the fossil record. The shift to bird from dinosaur called
for fewer changes than had earlier taken place among dinosaurs them­
selves and were, much later, to happen on the long road to the birds of
today.
There still remains a great gap in the record of modern birds. The
divers have long been suspected from their behavior and their anatomy
to be primitive; in addition, they turn up as fossils earlier than do their
fellows. Most birds of modern form appear somewhat later than the
divers, at almost the same time, some sixty-five million years ago-a
hundred million years after Archaeopteryx. Cormorants, owls and geese
turn up first, to be followed by the first penguins, the earliest parrots, and
then by the preserved remains of a whole host of feathered beings. In
 On the Imperfection of the Geological Record 205

spite of their noble ancestry, modern birds seem to have been sudden ar­
rivals on the fossil scene. Only a few hints of the long-lost avian past re­
main. The sandy shores of ancient seas a hundred million years ago
echoed to the cries of the waders who left their tracks as a memorial.
Those birds must have looked much like modern oystercatchers or red­
shanks, forty million years before the first fossil of any wading bird.
The lost birds that walked the sand left biological footprints in the
DNA of their descendants. They can be used to estimate when they
began. The new avian pedigree, based on a set of slowly evolving genes
associated with cell division, shows how much has been lost. Penguins
and albatrosses are close and fall into a group with the shearwaters and
the ancient divers. This puts the joint ancestor of the four at a time ear­
lier than the first fossil divers, and long before the demise of the di­
nosaurs. The rhea, the ostrich and the moa (together with the domestic
chicken) are so genetically distinct from other birds that their branch of
the avian family must have flourished even earlier. Fossil hunters, skep­
tical about such a gap, were persuaded by the discovery of a dead parrot
in Wyoming from a time when dinosaurs flourished. Birds of familiar
form may, it seems, go back a hundred million years before the first bird
fossil of modern appearance.
They are animals of the Southern Hemisphere, for most of their an­
cestors are found on the remnants of the great broken continent of
Gondwana (the progenitor of Australia, Antarctica and South America) .
The oldest fossils of ostriches, parrots, pigeons, songbirds, divers and
penguins are all on that land mass. In the still scarcely explored southern
part of the world there were many plumed and aerial descendants of di­
nosaurs, with diverse habits, flitting around their giant and earthbound
cousins. Our knowledge of their fossils, and of those of most other crea­
tures, is, it seems, so sketchy that it is as rash to dogmatize on the suc­
cession of organic beings throughout the world as it would be for a
naturalist to land for five minutes on some one barren point in Australia
and then to discuss the number and range of its productions.

On the sudden appearance ofgroups ofAllied Species in the lowest known

fossiliferous strata. For geology, life came in with a bang. The oldest fos­
sils known in 1 859-and until the 1 940s-were from the Cambrian sys­
tem of rocks, which began some five hundred and fifty million years ago.
Most of the major divisions of today's animal kingdom-the phyla-ap-
206 D A R W I N ' S G H O S T

pear just after the start of the Cambrian. Before then, nothing is to be
found. If evolution is gradual, how could so many forms spring all at
once into existence?
The nineteenth century, with its dislike of drama, appealed to a lost
and ancient world in which these groups had evolved, but left no rem­
nant. There was little evidence from before that time and what life there
was seemed quite unlike that of today. Stromatolites are limestone
mounds. Some are huge-a hundred yards across. Two billion years ago,
they made great reefs. Then they disappeared, with no obvious descen­
dants. Now, a few of their much-reduced builders have been found alive
on the shores of Western Australia; each stromatolite not an individual,
but an ecosystem a few inches tall, built by microbes. Not much else
seems to remain from before the Cambrian and most modern groups can
trace their " history no further than that time.
The Earth might then have been in the midst of a great evolutionary
adventure, the "Cambrian Explosion." Within a mere five million years,
there appear in the record snails, starfish and animals with jointed limbs
(whose descendants include the insects, spiders and crabs). Perhaps this
reflects some crucial change in DNA. After all, some mutations can at a
single bound change a fruit fly antenna into a leg. Might a great burst of
genetic creativity have driven a Cambrian Genesis and given birth to the
modern world?
The idea gained weight with the discovery in the 1 940s in the Edi­
acara Hills of South Australia of some shadowy and mysterious fossils.
They came from a time before that famous era and were the relics of soft­
bodied beasts that look not at all like life today-further evidence, per­
haps, of an eruption of novelty in the Cambrian itsel£ Such animals have
now been found all over the world. The youngest are just below the base
of the Cambrian, and the oldest go back as much as six hundred million
years.
That era was, without doubt, a busy time for evolution. In its operatic
landscape there appeared strange beings whose names-Anomalocaris or
Hallucigeni�testify to their odd appearance. An inhabitant of the
Burgess Shale beds, Opabinia, had five eyes and a long nozzle, a body
plan not seen since. But was it a period of unique innovation, with so
many new groups leaping into existence so quickly that selection could
never have made them? The answer is, with little doubt, no. First, the
Burgess Shale animals are less peculiar than is sometimes claimed. Hal-
 On the Imperfection of the Geological Record 207

lucigenia, with its strange spikes on its back, had, it turned out, been
drawn upside down. It has relatives among those drab animals the velvet
worms, still common in rotten logs in the Southern Hemisphere and no
more different from other animals with jointed limbs than are today's
barnacles.
The Cambrian Explosion, so called, is a failure of the geological record
rather than of the Darwinian machine. Its radical new groups reflect not
a set of exceptional events, but something more banal: the first appear­
ance of animals with parts capable of preservation. Before then, there
were soft creatures that decayed as soon as they died. Why shells ap­
peared all of a sudden is not certain. Perhaps the first predators evolved
and drove their prey to don expensive armor, or perhaps a surge of oxy­
gen enabled animals to grow large enough to need a skeleton. Whatever
the reason, the Cambrian marks the origin of a fossil record, rather than
of modern life.
Take those evolutionary celebrities, the trilobites, the first animals to
lay claim to jointed limbs. They are close to the roots of a tree that later
grew branches as flamboyant as the insects and a living fossil called the
horseshoe crab. If-as the record suggests-trilobites burst into exis­
tence within five million years at the base of the Cambrian, one brief
event changed the whole direction of evolution. In fact, a closer look
shows that among the earliest to be preserved were many distinct kinds.
Such diversity shows that trilobites had a past dating to long before that
famous era. What made them seem new was no more than their skele­
tons. Their predecessors had died and decomposed, but their more solid
descendants were preserved in millions. The Cambrian was a busy time
for trilobites, but it marked their middle age and not their infancy.
Now, at last, the youth of the world has been revealed. Newly discov­
ered fossils hint at a history older than anything imagined by those whose
inspiration begins with the Cambrian. Bit by bit, they have pushed life
further into the past; and each discovery is a new hint at the untold bil­
lions that have disappeared.
Phosphorites are rocks laid down in shallow seas. They are much
mined for fertilizer, and, in some places, preserve soft-bodied fossils in
fine detail. Great beds are found around the Yangtze gorges in China,
where they hide remains of lives from before even the Ediacarans.
Their ancient world was full of diversity. The Chinese beds, five hun­
dred and seventy million years old, hold the remnants of sponges, long
208 D A R W I N ' S G H O S T

thought of as primitive. To the amazement of those who found them

they also preserve the embryos of other animals, each cell seen in all its
detail. These oldest embryos look much like those of modern insects or
worms, animals with a left side and a right. This feature was, the first fos­
sils show, established in a distant past, quite unrecorded until a crucial
find was made.
Most unexpected of all are the seaweeds. Soft though they are, many
of those plants are in the phosphorite beds. They are almost as varied in
form as are their equivalents today. Those ancient plants have a hidden
history that began long before the Yangtze beds. One is almost identical
to a modern seaweed, so that this unlovely creature lays claim to the
longest of all pedigrees. The record of past seaweeds has always seemed
scanty, but this single collection shows it to be far more incomplete than
anyone had feared. Unlike the Ediacara, who have no modern equiva­
lents, such fossils show how much is missing from the early history of the
life of today.
Some billion-year-old rocks in central India have sinuous grooves on
their surface that look rather like those made today by worms creeping
on the bottoms of muddy ponds. Perhaps they are the traces of such
beasts, their bodies lost, but their spoor remaining. To burrow through
mud needs muscles and an internal cavity filled with liquid, and some
geologists question whether an animal so advanced could exist so far into
the past. If it did, it marks a gap in the record of worms as long as the
whole time from the Cambrian to the present.
A few discoveries from an unknown land prove that biology's sup­
posed preface, the Cambrian, is in fact well into its plo t . So deficient was
the evidence of earlier times that nobody guessed it was there. Single­
celled creatures stretch back even further. The balance of isotopes in car­
bon trapped in Greenland rocks three thousand eight hundred million
years old resembles that of the modern bacteria that live on methane.
They evolved when Earth was in its infancy and show that it took several
times longer to develop a cell with a nucleus than it did for life to appear.
Chemistry hints three billion years' worth of fossils older than the Chi­
nese seaweed are waiting to be discovered.
More familiar places are a token of how much has been lost. On Lon­
don's southern fringe are low hills of chalk, now improved by suburbs,
but with Darwin's own house still hidden at their foot. Those North
Downs give way to a wooded and intricate landscape, the Kentish Weald,
 On the Imperfection of the Geological Record 209

until, twenty miles farther on, a second range of hills, the South Downs,
takes over. At Beachy Head the Downs reach the English Channel. On a
clear day, France can be seen. That, too, is white. It continues the system
that built southern England. The Channel Tunnel, indeed, owes its exis­
tence to a band of solid chalk beneath the waves.
Once, long ago, the North and South Downs-and the Bas Boulon­
nais in France-were one. They have been worn down in what seems to
us an age, but was in terms of evolution an instant. Their chronicle is that
of the world, of the piling up and throwing down of rock, of times below
the sea and above it, and of great sections of the earth, with their fossils,
gone forever.
It is an admirable lesson to stand on the North Downs and to look at
the distant South Downs; for, remembering that no great distance to the
west the northern and southern escarpments meet and close, one can
safely picture to oneself the great dome of rocks which must have cov­
ered up the Weald within so limited a period as since the latter part of
the chalk formation. The history of southern England is written into that
VIew.

Its chronicle is one of shallow seas that dried and left their remains be­
hind. Nowadays Kent is a bowl with chalk hills on either side. Not long
ago the bowl was inverted, a peak with slopes running down to the sea.
It was evidence of a fold in the Kentish earth, made when the Alps began
to build. The strain buckled a whole series of beds. They begin with coal
from more than two hundred million years ago. Soon after the coal
forests, much ofWestern Europe sank. A great sea covered Kent. In it set­
tled sand and clay. The waters ebbed and rose until, a hundred million
years ago, the land sagged again. Over the whole sea bottom a thick sheet
of calcareous mud was laid. It became chalk.
Chalk does not just contain fossils-it is fossils, the shells of tiny ani­
mals called foraminifera, mixed with billions of tiny plates of lime se­
creted by marine algae. Scattered among them are the teeth of extinct
sharks, the shells of nautiluses, the bones of the odd pterodactyl, and
many of the sea urchins that once lived on the ocean floor. The white ex­
panse also has a few foot-long boulders, stones carried in the stomachs of
ancient reptiles as an aid to digestion. Apart from those remnants, the
rock is pure, as little was washed into it from the arid deserts that sur­
rounded the seas over southern England.
Then, at the end of the dinosaurs (the first of which was found in the
210 D A R W I N ' S G H O S T

Weald), the sea fell back and the land rose. Kent became an island of
chalk. Its rocks, and the sand and clay below, buckled upward. Five mil­
lion years ago, the land dried for the last time and rose still higher.
Nowadays, the chalk survives only as the North and South Downs.
The central part of Kent is made of older rocks: the coals, sandstones and
clays that once lay beneath the white shroud. The rest of the winding
sheet has gone. Most disappeared within a mere two million years. In the
ice ages, the soft rock was shattered as water froze within it. Rain could
not seep away, because the soil was so hard, but made great rivers that
carved valleys, now dry, through the hills. The top of the dome was
skimmed off. The denudation goes on. Sunken roads-holloways-run
through steep-sided trenches twenty and more feet deep. They were
formed by the horses, cows and carts that wore paths through the soft
rocks laid bare when the topmost layer was lost. The cliffs still fall into
the sea in great landslides, one of which, in 1 9 1 5, moved a train and its
passengers fifty yards out onto the beach at Folkestone.
The Channel itself was made as the sea destroyed the earth. The wa­
ters of the Dover Straits beat at the base of cliffs and cut them back like
a giant saw until the rocks collapsed under their own weight. A few
months before the millennium, hundreds of thousands of tons fell off
Beachy Head, and what had been a boat trip to the lighthouse became a
rocky scramble. The soft chalk moves at a great rate. The Belle Tout light­
house was built on the cliff top in 1 834. The edge moved back by fifty
feet a century (in spite of its owners' efforts to block rabbit burrows to
save it) and the precipice moved to ten feet away from the front door. Al­
though it has now, at great expense, been moved fifty feet inland, the
tower's fate is, within a geological instant, certain.
Deep grooves on the bed of the Channel show that the first breach in
the land bridge was made half a million years ago. The Thames was
pushed south by glaciers until its waters filled a huge lake behind the
ridge connecting England to France. The lake burst through and tore
away great blocks. Soon, the sea fell back as the climate cooled and the
bridge was rebuilt. Seven thousand years ago, the waters rose again and
wore away the rocks to make the present Channel. It gave Britain the
Straits of Dover, as a statement of national identity and a shield against
InvasIOn.
The Kentish landscape and its cousins across the Channel show how
 On the Imperfection of the Geological Record 21 1

fragile is the record of death over even a moment of evolutionary time.

They are the key to evolution's great anomaly, the difference in pattern
between past and present, between fossils and the plants and animals of
today.

From an aircraft window on an evening flight over the battlefields of Bel­

gium and northern France, great lakes of shadow fill the valleys, while the
hills, low as they are, are picked out in the sunset. Eighty years ago that
difference of a few feet meant life or death. It determined the results of
the great battles of the First World War, Ypres, Passchendaele and the
Somme. Passchendaele is notorious for the first use of poison gas, the
Second Battle of the Somme as the last gasp of the war. Each was, in ad­
dition, an experiment in paleontology.
To visit the trenches is to test the fate of the dead. The battlefields
show how incomplete is the record of the rocks, even when less than a
century old.
Before the Third Battle of Ypres in 1 9 1 7 the Germans held the high
ground and could look over the Allied lines. At the Somme, in March
1 9 1 8, the order was reversed and the British line was dug through a line
of low hills. The Passchendaele plain is a flat clay, its hills made of sand.
Further south-where the Somme was fought-chalk takes over. The
landscape is much the same as that of the Weald and the Downs.
The soldiers soon learned their geology. Clay became sodden and
drowned them in thousands, while sand collapsed as they dug their
trenches. Chalk was more reliable and huge dugouts could be built to es­
cape the shells. At Passchendaele a layer of clay passed from the British
lines to deep beneath those of the enemy. Miners were set to work to bur­
row through it. On 7 June 1 9 1 7 twenty huge mines went off at once.
The blast-the biggest non-nuclear explosion in history-blew a hole
two hundred feet across and forty feet deep. Ten thousand Germans were
killed and most of the remainder fled. Then, it rained. Tanks were
bogged down and the battle became another frightful stalemate. At the
Somme, the British were on the defensive and their line was broken. The
troops retreated but held the enemy a few miles farther back.
At Passchendaele, the slate of history has been wiped almost clean.
The visitor notices little, as the soft soil has swallowed up the trenches
212 D A R W I N ' S G H O S T

and most of the craters. The site of the giant mine is now an unremark­
able pond called the "Pool of Peace." Few tourists come because there is
not much to see.
They prefer the Somme, where the chalk has kept a better record.
Even so, just a few of the miles of trenches are still visible. Most are now
ditches and mounds rather than the crisp military structures of 1 9 1 8.
Their fate-and that of those who died in them-is that of the chalk that
once covered Kent. The erosion at Passchendaele reflects the loss of the
Weald's softer and more ancient rocks. Quite soon, no remnant will be
left of either field of war to remind visitors of the conflict. In less than a
century the surface of the earth has been so blurred that the record of the
past is almost lost.
Most of the bodies of the Somme were recovered and crowded into
cemeteries. Passchendaele still spits out the dead from its dreadful mires.
Its cemetery-the largest war grave in the world, Tyne Cot, named for
the thousands of Northumbrians who died in the battle-contains
twelve thousand graves. Three times as many are recorded only as ''A sol­
dier of the Great War-Known unto God" or as a name on a shared
monument; their bodies sunk so deep into the mud as to be unrecogniz­
able, or to be lost altogether. The Somme cemeteries are almost as full,
but more of their graves have names. Even so, to dig them up, as is some­
times done in the interests of good order, reveals, quite often, nothing
that can be identified as human.
Sixty billion people have lived since man appeared in modern form.
To excavate every graveyard-and every fossil site-in the world would
turn up no more than a minute fraction of their remains. The lost armies
of the dead have a moral for evolution. They are a reminder that the ge­
ological record is a history of the world imperfectly kept and written in
a changing dialect; of this history we possess the last volume alone. Of
this volume, only here and there a short chapter has been preserved; and
of each page, only here and there a few lines. However grand the monu­
ments, and however firm the hope of eternal life, in the depths of time a
Pharaoh in a pyramid has as little chance of immortality as does a soldier
stamped into a bloody swamp. The history of ancient Egypt, of the pres­
ent century-and of the existence of our own species-will soon be gone
forever: but what fragments might remain may allow some future histo­
rian to guess at the forgotten struggles that built his own world.
 C H A P T E R X

O N THE GEOLO G I CAL SUCCESS I O N

O F O RGANIC BEINGS

O n the power o f gradual change - Species once lost do not reappear - On

disaster caused by earthquakes, deluges and heavenly bodies - Gradual
and sudden change in the fossil record - On the true rate of evolution
and the sense of time - On fossil genes and the affinities of the past with
the present - On the state of development of ancient forms - On the
succession of the same types within the same areas - Summary of pre­
ceding and present chapters.

It takes three hours to travel the hundred and fifty miles of freeway from
Thebes to Memphis (Memphis, Tennessee, that is) . The journey can also
be done by river steamer, a trip that lasts a couple of days. The boat of
course is slower, but has to travel three times farther as the mile-wide
Mississippi meanders across a flat landscape. The river Meander itself, a
Turkish stream a twentieth the size of its American cousin, does the
same. Even rain does not take the shortest path downhill as it trickles
down a road.
Heraclitus (who lived near the Meander) had evolutionary views: no
man, he said, can step in the same river twice. Streams-always the same
but constantly in flux-were a metaphor for life. In biology, too, slow
change can have great consequences.
Streams evolve through a balance of forces. The bed shifts as it erodes
one bank and dumps its remains on the other. It returns when its loops
are cut off as the water finds a more direct route downhill. Complexity­
meandering-is opposed by simplicity, the shortest path to the sea.
214 D A R W I N ' S G H O S T

Raindrop, Meander and Mississippi follow the same rules. Measure­

ments of dozens of real rivers, and computer simulations of many more,
show that the relationship between their shortest possible path across a
plain and their actual length is always the same. It is pi, the ratio between
the circumference of a circle and its diameter. Each river, whatever its
size, goes a little more than three times farther than it needs on its way
to the sea.
As it does, it shifts uneasily. When Paul visited Miletus on his journey
to the Ephesians, the city was at the mouth of the Meander. Now, the
river passes it by. The efforts of the u.s . Army Corps of Engineers are all
that keep the Mississippi itself in place. Without them its mouth would
be at Morgan City, Louisiana, rather than at New Orleans. Rivers are as
inconstant as is their water. Stand on the bank for long enough, and you
will drown, because the stream will come and get you. How soon (sec­
onds for a trickle of rain and years for the Mississippi) is determined by
a simple and general law.
Darwin's theory makes the same appeal to unity in space and time.
Nature has no need to make sudden leaps. Instead, varieties, species and
the larger divisions of nature emerge-like raindrops and rivers-from a
process that acts without regard to scale. The very title of his greatest
work-On the Origin of Species by means of Natural Selection, or the
Preservation ofFavored Races in the Struggle for Life-joins the course of
change within a species to the grander process that causes new forms to
appear. DNA is a river from the past. If the trickle seen today is driven
by selection, why should its majestic passage through time not obey the
same rules?
It is possible to argue that all the great leading facts in paleontology
seem simply to follow on the theory of descent with modification. Old
forms are supplanted by new and improved forms, produced by the laws
of variation that still act around us, and preserved by natural selection.
Like the bed of a great river, the course of evolution is in its substance
simple.
As streams move on, they abandon their earlier path. Why should not
natural selection, too, have over the years led the inhabitants of each suc­
cessive period of the world's history to beat their predecessors in the race
for life? If some have improved, others must pay the price. On this view,
new forms are destroyed by the process that forges them. A species once
 On the Geological Succession of Organic Beings 215

lost can never reappear, even if the very same conditions of life, organic
and inorganic, should recur.
The rocks are filled with evidence that forms which do not become in
some degree modified and improved will be liable to be exterminated.
Until a few years ago we knew little of our own roots, as they, like most
others, have suffered that very fate. Once, the vertebrates-the group to
which mammals, birds and fish belong-seemed to have had a timorous
childhood. Today's versions of their predecessors-the lancelet and the
hagfish-feed on soft tissues. The lancelet and the young hagfish filter
particles from the water, while the hagfish adult is more interested in the
flesh of dead or crippled fish. They are not, perhaps, a noble set of an­
cestors for the lords of creation.
Now, the image of our past has changed and the lost world from which
we descend has at last been revealed. An abundant but enigmatic fossil
was the key. Many chunks of limestone contain thousands of tiny
pointed objects. Conodonts, as they are called, were discovered in Rus­
sia in the nineteenth century and have since turned up all over the world.
Because they are so widespread, and because their shape changes over ge­
ological time, they are much used to identify from which layer a partic­
ular rock might come. The animal who made them was quite unknown.
Like the tooth of the Buddha at Kandy, what remained was not enough
to reconstruct what might have been. The many guesses as to its appear­
ance included a version that looked rather like a jelly roll studded with
razor blades. All were, at best, implausible.
In 1 982, in some rocks from the shore near Edinburgh, were found
the preserved remains of the animal itself It had a soft eel-like body a few
inches long, with large paired eyes, a stiff rod down the back, and tail
fins. The conodonts themselves were not separate beings, but its teeth.
Before these dozen or so specimens-the first examples of the animals
that made the tens of millions of conodonts seen by geologists over a cen­
tury and more-the vertebrate skeleton was thought to have started as a
set of defensive plates on the body of a primitive fish. The first vertebrates
were, it seemed, victims, prey rather than predators.
The complete conodonts changed all that. The first sign of the skele­
ton was, it seems, in the mouth. Sets of conodont teeth, when pieced to­
gether, look as if they were used to shear flesh. The conodonts flourished
and diverged before they were driven out. More-and larger--conodont
216 D A R W I N ' S G H O S T

animals have now been found, from Wisconsin to South Mrica. One, the
size of a small fish, even preserves a pair of eyes (themselves as first clas­
sified as the remains of a plant) . The conodonts prove that our predeces­
sors were not grazers, sifters or suckers, but carnivores.
Now, the conodonts have gone and can never return; but their de­
scendants do all that they did, and much more, with grazing geese, sift­
ing whales, sucking lampreys and dogs that eat meat. Such animals are
more diverse and have more complex relations to their organic and inor­
ganic conditions of life, in land, sea and air, than did their predecessors.
Conodonts had a long and varied history, but were at last driven out. Ex­
tinction, as environmentalists so often remind us, is forever, and we will
not see their like again even if the vertebrates themselves should someday
disappear.
The idea of a dignified passage of origin, divergence and departure
has, for conodonts and many other vanished creatures, much in its favor.
In the real world, however, rivers or raindrops-or species-are often de­
flected from mathematical virtue. The unexpected intervenes, and the
beautiful simplicity that links their actual story to that set by theory dis­
appears.
In December 1 8 1 1 a gigantic earthquake struck between Memphis
and Thebes. It was equal in power to the combined energy of all those in
North America since. The shock was enough to ring bells in Boston.
Columns of coal dust rose from the ground, and few houses within two
hundred miles were left upright. The crew of the first steamboat on the
Mississippi had moored to an island. When they awoke, it was gone:
"The numerous large ponds . . . were elevated above their former banks
. . . A lake was formed on the opposite side of the river, in the Indian
country . . . It is conjectured that it will not be many years before the
principal part, if not the whole of the Mississippi, will pass that way."
Although a ten-mile lake-today's Reelfoot State Park-appeared that
day, the river disdained it as a new course. Even so, it shifted its path in
many places as islands sank and bluffs collapsed. One cataclysm did
more to determine the Mississippi's fate than had centuries of gradual
change. Now, the river faces another catastrophe: control by man. Al­
though the state line between Arkansas and Mississippi still meanders
along the bed of an abandoned stream, the waters themselves flow down
a straight and narrow channel.
 On the Geological Succession of Organic Beings 217

Life, too, has had its disasters. Why should they not, at least in part,
determine patterns in the fossil record? No doubt, gradual selection was
important in the rise and fall of many beings. But could not ancient
tragedies of geology or genetics sometimes have done the job? The record
has many sudden deaths and just as many unexpected entrances. Some,
no doubt, are there only because the fossil record is incomplete, but some
might be the biological equivalent of the Mississippi earthquake. Are dis­
asters needed and, if so, how often do they happen? Are they a merely de­
structive force, or might they be a help to evolution? Such questions are
at the heart of the disagreements between paleontologists and those who
study the plants and animals of today.

On Extinction. Many creatures disappear because they are replaced by

something better. That process, busy all over the world, is itself evidence
of natural selection. Before Darwin, the nineteenth century had pre­
ferred to see the past swept away by acts of God. Disasters were an excuse
for not thinking. Charles Lyell was scornful: "We are told of general cat­
astrophes and a succession of deluges, of the alternation of periods of re­
pose and disorder, of the refrigeration of the globe, of the sudden
annihilation of whole races of animals and plants, and other hypotheses,
in which we see the ancient spirit of speculation revived, and a desire
manifested to cut, rather than patiently to untie, the Gordian knot."
As any visitor to the Galapagos can see, plenty of extinction takes place
in the Darwinian way, as an old order gives way to a new. However, lots
of life has gone out with a bang. General catastrophes there have cer­
tainly been; disorders greater than the Mississippi Earthquake that have
wiped out the well-crafted products of evolution over much of the Earth.
Any animal caught up in such an event, however well adapted it may
have been, has every chance of annihilation. In geological terms, even the
great glaciations (slow as they appear to us) were sudden, and the oblit­
eration of much of the Northern Hemisphere over a few thousand years
is, on the scale of the rocks, abrupt enough to count as a rude interrup­
tion of a gradual climatic trend.
When it comes to deluges, the Black Sea tells part of the tale. It was
once a small freshwater lake, the Bosphorus no more than a dry ridge be­
tween two basins. Then, to the north, a river carved a valley. To the
south, a slip in the earth's surface made a deep fjord. Five and a half thou-
218 D A R W I N ' S G H O S T

sand years ago, as the sea rose, the Mediterranean broke through and the
Black Sea filled. It did so over a cascade the size of two hundred Niagara
Falls. The roar could be heard three hundred miles away and in some
places the shore moved back by a mile a day.
The salt waters were an evolutionary tragedy for the animals of the
Black Sea plains, however well they were adapted to their humble lives.
Five hundred years later, the Mediterranean itself died, leaving a thin
layer of rot on its floor as evidence of a holocaust. Soon after the
pharaohs arrived there was a time of heavy rain. It ran off the denuded
land to cover-and smother-the ocean with fresh water. No oxygen
could get through as the layers did not mix. Like today's Black Sea (in
which a layer of oxygen-free salt water sits below a fresher cap), the
Mediterranean was dead in its depths. The whole process took sixty years
and swept away thousands of species. Given today's pollution, the sea
could die again just as quickly.
Flood legends are common in the Middle East. The earliest, that of
the Epic of Gilgamesh, dates from about the time of the inundation of
the Black Sea basin. Noah's Flood, much derided by evolutionists, may
be one of many sudden reverses that changed the course of their science.
On Friday 9 October 1 992 a meteorite damaged the right rear fender
of a 1 980 Chevrolet Malibu, the property of Michelle Knapp of Peeks­
kill, New York. The rock was sold for fifty-nine thousand dollars and the
car, damage and all, for ten thousand.
The chance of real harm from a celestial body seems tiny. In spite of
the threats of Apocalypse ("a star from heaven, burning as it were a
lamp"), the sole recorded case of a direct hit was in Alabama in the 1 9 50s
when a lady asleep on a couch was bruised by a meteorite that bounced
off her hip.
In the end, though, disaster is inevitable. A trillion comets orbit the
sun. The gravel that surrounds them appears, should it hit our atmo­
sphere, as a shower of meteors. The Earth gains a ton in weight every
hour from their dust. Two thousand asteroids big enough to destroy civ­
ilization orbit nearby. Given the likelihood of a hit in the course of a life­
time, the safety of aircraft, and the few who fly, the chance of an average
person meeting death from a comet is higher than that of death in an air
crash. Apocalypse is, sooner or later, guaranteed. The United States gov­
ernment even established a committee (chaired by the then vice presi-
 On the Geological Succession of Organic Beings 219

dent J . Danforth Quayle) to find out how to deflect or destroy any ce­
lestial body audacious enough to threaten to land on the United States.
There are plenty of reminders of the danger. In 1 996 an asteroid missed
Earth by a mere quarter of a million miles. A direct hit would have been
equivalent to an explosion of all the globe's atomic weapons at once. As
Voltaire said of such an event: "What a Disaster would it be for our Earth
. . . The Idea of two Bombs, which burst on clashing together in the Air,
is infinitely below what we ought to have of such an encounter as this."
Meteor Crater in Arizona was not recognized as such until 1 929, giv­
ing rise to a wave of cosmic fears to go with the Wall Street crash. The
greatest crater of all is further south, near the port of Progreso in the Yu­
catan Peninsula of Mexico. The surface shows little sign of the object six
miles across that struck with the force of five billion Hiroshimas sixty­
five million years ago. Its testimony is buried under a mile of limestone.
An ancient seabed in New Jersey reveals, at just that time, a three-inch
layer of glassy spheres, the remains of the liquefied rock ejected from the
crater. All over the world, the rocks of those days have high levels of the
element iridium (brought, perhaps, by the meteor), together with much
soot.
The seventh-century Chinese philosopher Li Ch'un Feng had it that
"comets are vile stars. Each time they appear . . . something happens to
wipe out the old and establish the new." The Yucatan disaster wiped out
an old regime forever and helped to establish the modern world. Below
the layer marking its arrival are the remains of tiny dinoflagellates and
foraminifera-hard-shelled animals present in billions in the ocean­
while above it, all are gone. No new marine animals appeared for thou­
sands of years and, after the bang, the seas were almost dead. The
dinosaurs, too, disappeared at about that time.
The mass extinction is itself smaller than the cataclysm marking the
end of the Paleozoic period two hundred and forty million years ago,
when nine-tenths of all marine species disappeared; an event that has set
the sea's ecological agenda ever since. The oceans lost much of their oxy­
gen, perhaps because of a huge volcanic eruption in Siberia and an out­
pouring of sulfurous gas that poisoned much of the world.
There have been more recent cataclysms. In central Siberia is the fifth­
largest impact crater on earth. The rock that splashed out is thirty-six
million years old. At almost the same time another comet struck on what
220 D A R W I N ' S G H O S T

became the eastern coast of the United States to make the Chesapeake
Bay. The two big bangs marked the appearance of ice sheets in Antarc­
tica and the greatest extinction since the loss of the dinosaurs. It marks
the boundary between two geological periods, when the first simple
horses, deer and whales gave place to mammals close to those of today.
Three extinctions at the time of a geological disaster must be more
than a coincidence. But how sudden were the bangs--or were they roars,
or even distant rumbles? In one sense, they were abrupt indeed. At the
famous collision of Yucatan, everything within a mile of the impact was
extinct within a second. Within a minute the Americas were shaken by
earthquakes and nine minutes later most of the continent's forest burst
into Hames. Later the same day, huge areas across the world were de­
stroyed by tidal waves, and at sunset a nine-month night of smoke and
dust began. Voltaire underrated the disagreeable effects of even a small
meteor.
Even so, for most of the Earth's inhabitants the explosion meant dis­
comfort rather than death. It took a hundred thousand years before all
the dinosaurs were gone, and the last ammonite lingered on for as long
again. Many bivalves {mussels and the like} disappeared at about the
same time-but their holocaust happened thousands of years before the
impact. Indeed, the dinosaurs themselves had begun to decline before
their fate was {perhaps} sealed by an agent from outer space. Land plants
and fish seem unperturbed by the blow and their history continues more
or less as normal through those troublesome times. What is more, some
marine animals rose, like Lazarus, from their graves, long after the disas­
ter. They were dead to all appearances, but resurfaced as a statement of
the record's deficiencies rather than of divine assistance.
It is, nevertheless, certain that an event out of the ordinary hit the
world sixty-five million years before the present. It took two million years
to get back to normal, and the new normality was noticeably different
from the old, with many novel kinds in the place of those who had gone.
Evolution's progress {if such it is} is not uninterrupted. Catastrophe on a
scale unknown to history has played a part. Whether it had a construc­
tive, rather than a merely lethal, effect is another issue. Some claim that
mass destruction led to biological explosions as the survivors evolved to
fill the gaps. If they are right, cataclysms drive change as much as does
slow modification. Certainly, some patterns in fossils suggest that grad-
 On the Geological Succession of Organic Beings 22 1

ualism is not enough and that the river of life has suffered many earth­
quakes, of several kinds.

On the Forms ofLife changing almost simultaneously throughout the World.

A world designed by a physicist (or a god) might start simple and grow
more complicated and stop when it reached perfection. Life has never
been like that. A year after The Origin, the geologist John Phillips recog­
nized three peaks of diversity among fossils, each followed by a sudden
extinction and a gradual climb back. The record has long periods in
which not much happens, followed by episodes of rapid transformation.
In spite of arguments about just how flat were the plains and how tall
the peaks, most geologists accept that there were indeed three high points
of variety. They came at the height of the Cambrian around half a billion
years ago; in the Paleozoic from around four hundred to two hundred
million years ago; and in the modern period, which marks the richest
mix of species ever seen. What caused such deviations from divine (or at
least arithmetic) perfection?
The idea that birds and mammals had to wait for the demise of the di­
nosaurs before they could come into their own is wrong. The genes show
that, for both, there were tens of millions of years of profitable change
before the heavenly body struck. All the modern groups were well on
their way before that vexatious event. Even so, most lineages appear in
the record quite suddenly. One section of marine fossils in upstate New
York is typical: it stretches over a hundred million years and all its many
kinds stay the same for millions of years before most alter over just a few
hundred thousand. For marine fossils with a long enough history­
corals, snails and small bottom-living creatures-nine tenths of all
records are like that: a sudden appearance followed by millions of years
of tedium until the line disappears or emerges unchanged today. A few
alter slowly, in true Darwinian style, as proof that the patterns found in
the others are not due to some error of sampling, but the general picture
is that life is calm for most of the time. One tadpole shrimp is so static
that it has kept the Latin name used for its ancestors of a million years
ago, but it copes so well with the modern world that it is a pest of irri­
gated rice fields.
Proponents of gradual evolution, of stability and of revolution can
each cali on convincing evidence to support their views. No universal
222 D A R W I N ' S G H O S T

pattern unfolds within the rocks. Now that the fossil record is so much
better understood than once it was, evolutionists can no longer point to
imperfection as the sole explanation for its structure. Some lineages alter
in what seems a non-Darwinian way, not gradually but with episodes of
calm punctuated by change. Whether such conservatism lays bare some
inherent reluctance to adapt, or whether it shows merely that not much
evolution is needed because Nature's challenges tend to stay the same, we
do not know. However Darwinians may protest {and they do}, millions
of generations of inertia scarcely fit his image of life as poised for an in­
stant response to any challenge. The argument between supporters of
evolution as unhurried Victorian progress and those who hold the mod­
ern view of history as boredom mitigated by panic is unresolved.
In part that is because the question asks too much of a fossil record so
battered by the accidents of time; but it also turns on a disagreement
about the meaning of words. What is gradual, and what instantaneous?
Zhou En-Lai, when asked his opinion of the French Revolution, said
that "It is too early to tell." His was a long view of the past, which to
Western eyes seems quite out of proportion. The Nationalist regime in
China had by then occupied just a couple of decades in a political con­
tinuum from the Shang dynasty, four millennia before. The events since
the Bastille, dramatic as they appeared to those involved, were an instant
in history, and perhaps an unremarkable one. It is all a matter of scale.
Much of the argument about whether the fossil record shows slow or
rapid change depends on what those terms signify.
The Turkana Basin in East Africa holds the remains of many ancient
primates, together with those of our own tool-using predecessors. All are
preserved in strata separated by well-dated layers of volcanic ash. Spec­
tacular as these bones might be, they are rare and say little about gradual
or interrupted evolution.
In the same Koobi Fora beds are preserved millions of shells of twenty
or so different kinds of freshwater snail. Their fossils have remained al­
most undisturbed since they were laid down. At first sight, what they re­
veal is not at all the pattern expected of slow and successive modification.
Instead-as is true for so many marine fossils-long periods of stability
are interrupted by sudden bursts of change. A new variety appears, per­
sists unaltered, and disappears as quickly as it came. This is not just a
matter of the inadequacy of the record, as huge numbers of the inhabi-
 On the Geological Succession of Organic Beings 223

tants of these lakes are preserved. The intermediates are around for just
a tiny part of each lineage's history. How can this be, if nature does not
make leaps?
The problem comes, as Zhou En-Lai saw, from the sense of time.
Those who peer into its depths find it hard to see things in proportion.
When one referee in nature's race is used to a stopwatch and the other to
Big Ben, disputes are to be expected. An instant to a paleontologist may
appear an infinity to those who study life today. In the Turkana fossils the
"intermediates" last for just a tiny part of the duration of their ancestor
or descendant forms. As a result, they seem a classic case of an evolu­
tionary leap in the snail pedigree; a pattern quite unlike anything ex­
pected from slow and successive modification.
However, that moment represents, for different lineages, between five
thousand and fifty thousand years. The snails have one or two genera­
tions each year. A blink of a paleontologist's eye hence covers about
twenty thousand mollusk generations. For dogs, that is equivalent to
forty thousand years, a period longer than it took to evolve chihuahua
from wolf by conventional selection. Why appeal to anything else in the
case of the snails?
Natural selection can certainly do the job of transforming one snail
into another. But why, if it can, are its energies confined to such a short
part of the history of each kind? And why are the snails immovable for
so long when they have the capacity to evolve? Is something more needed
to explain such unexpected patterns? That is the argument of "punctu­
ated equilibrium"-the notion of evolution as stasis interrupted by sud­
den change, a pattern that might, perhaps, result from some intrinsic
property of the organism, rather than of the environment in which it
lives. It points at a great disparity between what life can do and what,
over evolutionary history, it does.
A universal measure of the rate of evolution that can be used on fruit
flies over a few weeks in the laboratory or on dinosaurs over the millen­
nia points up the contrast. A unit of change per million years is called a
"Darwin." It is based on how much the average size of any feature (cor­
rected for the absolute size of the structure itself) alters with the years.
Laboratory experiments on flies can generate values of over a hundred
thousand Darwins, and selection on the farm often gives rates of several
tens of thousands. Transplant experiments {such as those of the guppies
224 D A R W I N ' S G H O S T

moved from a dangerous stream to a calmer place) give rates of change

of up ro fifty thousand Darwins. English sparrows were introduced into
the United States a century ago and have filled the Americas. As they
spread, they evolved a pattern similar to that in the Old World, with
heavy birds with short limbs in the cold north and lighter and more
graceful animals in hot places. The legs of the birds from the warm south
have lengthened at around a hundred Darwins (which is about 5 percent
in a century) . Although it is not known to what extent the shift in spar­
row shape is due to genes rather than to a direct response to the envi­
ronment, evolution at that speed would bring forth a sparrow with the
legs of an ostrich in just ten thousand years.
Fossil rates of evolution are far smaller. To paleontologists, horses are
a classic of rapid modification, as their teeth altered to keep up with a
shift from soft leaves to hard grasses. Over several million years, they
changed at a tenth of a Darwin-which means that it took millions of
generations for the average height of the tooth to shift from that of the
smallest to the largest present in the population at any time. In the
deeper record-the hundred and fifty million years of the dinosaurs in­
cluded-evolution of shape or size is thousands of times slower still. Life,
it seems, has done less evolving than it could. Why, we are by no means
sure.

On the affinities of extinct species to each other, and to living forms. The
genes of today link every plant and animal together in an unbroken chain
of ancestors that descends from the ancient past. Now, they can do more,
for some of the molecules of life are themselves preserved as fossils.
A coal mine is a hecatomb of past existence, full of the remnants of an­
cient trunks and leaves. Oil is a degraded form of chlorophyll, the green
material of plants. About a fiftieth of all rocks-limestone, chalk and so
on-that fell as sediment from oceans is made up of biological remains;
and such rocks contain ten thousand times more organic material than
does the whole of life today. Some of the ancient material preserves its
structure. Chitin, the solid material of the insect skeleton, has been
found in twenty-five-million-year-old remains from a lake bed, and the
amino acids in scallop shells may last for six times as long.
Snail shells, too, retain a genetic record of the past. They keep their
marks for thousands of years. Those buried, unnoticed, by the ancient
 On the Geological Succession of Organic Beings 225

farmers who erected the great monuments of Silbury Hill and Avebury
Trusloe in southern England were dug up, just a few feet below their de­
scendants, when the sites were excavated. The genes for shell pattern had
changed. Ancient populations from colder times are darker in color, be­
cause dark shells are favored at times when it pays to soak up the sun's
heat.
The idea that ancient DNA could be preserved gained fame from the
film Jurassic Park, in which dinosaurs were re-created with the help of
fossil DNA. The molecule certainly can persist after death. It was first
found in a nineteenth-century museum specimen of a quagga, an extinct
zebra. More turned up in a two-thousand-year-old mummy dried in
Egyptian sands (exciting the Copts, who hoped to test their claim to
pharaonic descent). There were reports of its presence in a hundred-and­
twenty-million-year-old weevil preserved in amber and in dinosaur
bones themselves.
That was every evolutionist's dream. What better than to have the
genes of ages past to compare with those of today? Reality made an un­
welcome appearance when it emerged that almost all so-called ancient
DNA results from contamination with modern material (a real problem,
given the sensitivity of the methods used to search for it.) The molecule
cannot last for long. One measure of its frailty lies in proteins. After
death, they break down into individual building blocks called amino
acids. These come in two mirror-image forms, which in life are biased to
the left. The leftward inclination begins to disappear after death, with a
slow chemical switch back to a mixture of the two. The more equal the
proportion of right- and left-handed amino acids, the older a fossil must
be. DNA decays at about the same rate. Any fossil whose building blocks
have lost their natural left-handed bias must hence, for simple reasons of
chemistry, have lost all its DNA.
Almost all supposedly ancient DNA fails the test. As Hamlet's
gravedigger points our, "Water is a sore decayer of your whore-son dead
body." Whatever its parenthood, any DNA in an animal whose body lies
in a wet place is at once destroyed. Ice, in contrast, can be a positive help
(which is why pathologists have searched for the genes of the 1 9 1 8 flu
virus in the frozen remains of Alaskan fishermen) . Genuine (albeit frag­
mented) DNA is in the bones of the giant sloths that led Darwin to call
the pampas "one wide sepulchre of these extinct gigantic quadrupeds." A
226 D A RW I N ' S G H O S T

North American relative from twenty thousand years ago left feces filled
with DNA from its last meal of grapes, flavored with mint. Frozen Sibe­
rian mammoths from forty thousand years earlier seem, from their genes,
to have been more related to African than to Asian elephants.
Rare though its remains may be, DNA has left plenty ofless direct ev­
idence of its history, for although the molecule itself may disappear, its
footprints remain in the bones themselves. Lungfish are living fossils: an­
imals with an agile and creative past that nowadays have sunk into con­
servatism. Long ago, they slowed down, and they have stayed unchanged
for hundreds of millions of years, while their relatives moved on.
Bone contains many cells, all with a nucleus. The hard material
squeezes each one so that its size is a measure of how much DNA it once
contained. Early in lungfish history, the size of the cell nuclei-and the
amount of genetic material-began to creep up. Soon, the animals had
hundreds of times as much as did their relatives. As it did, evolution
slowed. Now, the lungfish are stuffed with DNA (most of it with no ap­
parent function) and their evolution has stalled altogether. The fit be­
tween DNA content, a lethargic lifestyle and evolutionary sloth is
widespread. To copy that chemical takes energy. Bacteria are speedy and
have no excess genetic material, while salamanders, torpid as they are, are
filled with DNA. Plants, too, have a close fit between habit and nucleic
acid content. All weeds have small genomes, while more established
plants are packed with DNA and can take a month to make a single egg
cell. Whether an indolent life allows the amount of genetic material to
build up, or whether the extra dose itself slows down evolution, nobody
knows.
Fossils show that the same happened to the lungfish. Whatever slowed
it down, the imprint of lost genes left in its bones shows the affinity of
an ancient species to its descendants. Although life, as usual, does not live
up to what Hollywood can do, the tragedy of the lungfish links today's
genetic material to that of long ago.

On the State ofDevelopment ofAncient Forms. The idea of a present in de­

cline from the past has been around since history began. Homer and Vir­
gil deplored the decrease in stature of the human race, and Greek actors
in the role of mythical heroes played them on stilts. The notion was sup­
ported by fossils. The bones of the monster Polyphemus, found in Sicily
 On the Geological Succession of Organic Beings 227

in the fourteenth century, joined many others hung in churches as proof

of the Genesis claim of "giants in the earth in those days. " Polyphemus's
bones were in fact those of fossil elephants. His single giant eye (as leader
of the Cyclopes he was blinded by Odysseus when cast ashore on Sicily)
came from the large nostril in the center of an elephant skull, which re­
sembles a huge eye socket. Other ancient elephant bones were paraded
around Europe reassembled into saints. They led a French anatomist to
calculate that, given the rate of decline since the Creation, Adam must
have been a hundred and thirty feet high.
When it comes to evolution, size still counts. Buffon had claimed that
in America dogs lost their bark and "all animals are smaller." Even men
lacked virility: the American Indian was "feeble . . . He has small organs
of generation . . . and no ardour whatever for his female . . . Nature, by
refusing him the power of love, has treated him worse and lowered him
deeper than any animal." Jefferson sent Buffon measurements of bears
and beavers and, as final proof, a stuffed moose, but not until the first all­
American mastodon was exhibited in Paris was the reputation of the
New World saved.
The notion of decline was succeeded by the idea of advance, by the as­
sumption that today's plants and animals are more developed than were
their ancestors. Nature, it seemed, moves inexorably from microbe to
man, in Pope's "Vast chain of being! Which from God began, / Natures
aethereal, human, angel, man, / Beast, bird, fish, insect, what no eye can
see, / No glass can reach; from Infinite to thee." But has life improved,
and, if it has, are its latest models better than what went before?
The theory of evolution by natural selection differs from other ideas
about the past because it has no inbuilt need for progress. Richard Owen,
inventor of the word dinosaur, founder of the Natural History Museum
in London and polymath to the British government, was unabashed
about what history was for. When it came to fish, for example, "those
species, such as the nutritious cod, the savoury herring, the rich­
flavoured salmon, and the succulent turbot, have greatly predominated
at the period immediately preceding and accompanying the advent of
man; and they have superseded species which were much less fitted to af­
ford mankind a sapient and wholesome food." His view that life changes
for man's convenience contrasts with Darwin's stark-and accurate­
warning that no animal exists for the good of another. The evolution ma-
228 D A R W I N ' S G H O S T

chine can do no more than adapt its products to whatever they face at
any instant. It has no inbuilt direction, be it forward, backward or side­
ways.
In Italy, such was the force of habit that it was until 1 923 customary
to drive on the left-hand side of the road in town, and on the right in the
country. The Darwinian machine, too, once started, has plenty of iner­
tia. Life might be short on progress but it has plenty of inertia. The Eu­
ropean ancestor of domestic cattle, the aurochs, was much hunted before
it had the sense to form an alliance with its enemy. The last one died in
1 627. It was black, six feet high, and had fearsome horns. There have
been many attempts to re-create it with crosses between fighting bulls
and other breeds. Although these animals resemble their noble an­
tecedent, they are a cheap copy: like it in shape, but quite different in
their genes. To reverse through the maze of descent is impossible.
The vehicles of evolution often become trapped in a one-way system.
In 1 9 1 0 a German miner found a beautiful fossil of a chicken-sized aer­
ial reptile. Although one expert thought this improbable and removed
the wings as mere remnants of a dead fish it was indeed the earliest ver­
tebrate to fly, twenty million years before the next animal took to the air.
Bony rods stiffened a pair of curved wings made from skin flaps. These
opened like a Japanese fan and allowed the animal to glide a hundred
times its own length, with an approach quite unlike that of bats or birds.
However, the Japanese fan went nowhere-because, unlike a modified
arm, there was nowhere much to go.
Sex, too, is a habit that tends to stick. Some are able to return to the
innocence of Eden when the chance arises, but many others have taken
an irrevocable step along the sexual road. Pines stay with it because the
chloroplasts (the green factories that use sunlight as a source of energy)
are passed through pollen, while fish are forced to hold on to their males
because the sperm provides the machinery of cell division. Virgin birth
is unknown among mammals because males put a stamp upon genes that
pass through sperm. Certain fish use sperm of another species to activate
an egg, but they, too, are imprisoned by sex; as although they do not use
his genes, they need the male to make the crucial sperm. A state of de­
velopment descending from the most ancient forms is, it seems, hard to
escape.
Even so, in one particular sense the more recent forms must be higher;
 On the Geological Succession of Organ ic Beings 229

for each new species is formed by having had some advantage in the
struggle for life over other and preceding forms. From the extraordinary
manner in which European productions have recently spread over New
Zealand we may believe that in the course of time a multitude of British
forms would exterminate many of the natives. Under this point of view,
the productions of Great Britain may be said to be higher than those of
New Zealand. Yet the most skillful naturalist from an examination of the
species of the two countries could not have foreseen this result.
Some trends do suggest progress, but on a limited scale. The tiny
worms who left evidence of their ancient hunts as tunnels in mud that
later became stone started off searching at random, but-four hundred
million years later-were tracking back and forth across the bottom in a
more purposeful way. In mammals, most ancient species are smaller than
those alive today. At the end of the dinosaurs, the largest mammal was
the size of a cat, but now we have the elephant and the whale. The di­
nosaurs themselves grew from cats to eight-ton giants within a few tens
of millions of years. However, the leviathans did not last. Their only sur­
vivors are now known as birds and have suffered a drastic decrease in size.
Not just the dinosaurs lost out. The giant sloths of the Americas have dis­
appeared; modern horsetails are tiny compared to those of the coal
forests and in spite of signs of progress in fossil behavior, plenty of worms
today blunder at hazard across lake bottoms. Evolution has no escalator
of increase, be it in smartness or in size.
The idea of evolution as a ladder is (or ought to be) dead, but life has
certainly got more complicated since it began. Today's bacteria are not
much different from those of two billion years ago; but now we have an­
imals and plants as well. Some organs have become more intricate than
before. Insect limbs have evolved from simple jointed tubes into pincers,
paddles and other things, while the mammalian brain is more folded
than it was. Other structures (such as the skull, with fewer bones in hu­
mans than in ancient fish) have stalled. Plenty of animals simplify their
lifestyles, with males as parasites inside their females. In parasites them­
selves, all is lost but the organs of digestion and sex.
Complexity sends mixed messages about progress along the evolu­
tionary road. The earliest sponges had a mere half-dozen cell types, but
humans have hundreds. There has, even so, not been much increase in
that measure of complication over millions of years, since fish have about
230 D A R W I N ' S G H O S T

the same number as ourselves. Sponges have become more colonial since
they began, but the corals are nowadays more solitary than they were. In
economics, in the long term, as Keynes said, we are all dead. Biology, too,
never thinks of the future and what matters happens now, in an ecolog­
ical instant, with no grand plan.
Too often, the notion ofprogress is used as a code word for perfection,
the chain of being in a different guise. The term should be employed
with caution. Some see an arrow of time in biology, as in physics, but in
the opposite direction-a relentless tendency to improve, j ust as the uni­
verse has a built-in trend toward chaos and disorder. That is too opti­
mistic. Some lineages get more complicated, some simpler, and much of
life has to struggle to stay in the same place. If everyone is evolving, no­
body can afford to stop, and there may be constant change with no over­
all advance at all. Although living things have become more complicated
in the past four billion years, the issue ofwhich form is higher and which
lower usually depends on who asks the question. Evolution does not
need progress. After all, transfixed by time's arrow, all its products will
soon be gone.

On the succession of the same TJpes within the same area, during the later
tertiary periods. Science fiction sees the ancient world as much like today,
with added dinosaurs. For biology, though, the past is another country,
a story of different worlds as much as of different beings.
Evolution is, rather like history itself. a drama in which different ac­
tors succeed to the same role. The world has fifty or so large carnivores­
lions, wolves, jackals, bears and more. Wherever they are found, they fall
into four groups with different habits. Some, big cats included, eat only
meat. Hyenas eat meat and crush bone. Jackals have wider tastes, and
take half their food as roots and vegetables, while bears have shifted fur­
ther toward vegetarianism. Each faction has its own skull structure, and
any skull can at once be ascribed to the group to which it belongs.
There is more than one way to skin a cat. Five million years ago, in
what is now Yellowstone National Park, all the groups were present, but
different animals played each part. Hyenas had not yet appeared. In­
stead, a large dog went in for bone crushing. Bear-like animals roamed
the land. They were not real bears, but huge raccoons with a taste for
berries and roots as well as flesh. In spite of much extinction and the ap-
 On the Geological Succession of Organic Beings 23 1

pearance of new forms, the four elements, made up at various times of

quite dissimilar beasts-bear-dogs, bear-raccoons and a lost set of cat­
like meat-eaters called nimravids-Iasted for millions of years. Birds
show the same consistency. Today's vultures, from the Americas or from
Mrica, fall into three guilds, each with distinct heads and beaks. They are
rippers, gulpers and scrapers; and scrapers, gulpers and rippers from
quite a different set of birds are preserved in the La Brea Tar Pits.
However, the evolutionary play has had such a long run that some
changes in its scenery are inevitable. Five hundred million years ago the
air had twenty times as much carbon dioxide as it contains now. This led
to a natural "greenhouse effect" which was reversed two hundred million
years later when the level of the gas dropped. Oxygen, too, has swung be­
tween extremes. Twice as much of the gas as today allowed the growth of
enormous plants, of spiders the size of a book, and of scorpions a foot
long. A later burst led to the development of aerial reptiles such as Quet­
zalcoatlus, with wings forty feet across. Oxygen's abundance allowed
many animals to burn energy at a rate great enough to persuade them
into the air. In today's attenuated atmosphere, nothing so large could
carry the burden of gravity. Even the days of the Earth have changed. As
the moon saps its neighbor's rotational energy, the globe slows its spin.
Corals have daily and annual surges of activity, and growth rings from
four hundred million years ago show that there were then four hundred
days a year. Whatever its state of development, the pace of life in those
short and energetic days was faster than it is today.
The geological record is the court oflast appeal for all theories of evo­
lution. Although biologists still argue about how the process works, fos­
sils make it impossible for anyone, biologist or not, to deny that it
happened. Cuvier, faced with a pile of bones of extinct animals from the
Paris Basin, said, "We will take what we have learned of the comparative
anatomy of the living and will use it as a ladder to descend into the past."
Now, the fossils themselves provide, if not a ladder, a set of hesitant steps
through time and a proof of a past more dramatic than anything he
imagined.
The days when fossil hunters were, like the College of Heralds, en­
gaged in a futile search for missing links have gone. The record of ancient
beings is the most forceful statement of what evolution can do. A cen­
tury ago, geology was at the center of the conflict between science and
232 D A R W I N ' S G H O S T

belief For a time, its patterns seemed so different from those of today as
to lead to scientific arguments that were almost theological in their in­
tensity. The disagreements are still there; but now that those who study
ancient and modern forms of life both speak the language of descent
with modification they may, some day, be solved.

Summary of the preceding and present Chapters. I have attempted to show that
the geological record is extremely imperfect; that only a small portion of
the globe has been geologically explored with care; that only certain
classes of organic beings have been largely preserved in a fossil state;
that the number both of specimens and of species, preserved in our mu­
seums, is ahsolutely as nothing compared with the incalculable number
of generations which must have passed away even during a single for­
mation; that, owing to subsidence being necessary for the accumulation
of fossiliferous deposits thick enough to resist future degradation, enor­
mous intervals of time have elapsed between the successive formations;
that there has probably been more extinction during the periods of sub­
sidence, and more variation during the periods of elevation, and during
the latter the record will have been least perfectly kept; that each single
formation has not been continuously deposited; that the duration of each
formation is, perhaps, short compared with the average duration of spe­
cific forms; that migration has played an important part in the first ap­
pearance of new forms in any one area and formation; that widely ranging
species are those which have varied most, and have oftenest given rise to
new species; and that varieties have at first often been local. All these
causes taken conjointly, must have tended to make the geological record
extremely imperfect, and will to a large extent explain why we do not find
interminable varieties, connecting together all the extinct and existing
forms of life by the finest graduated steps .
He who rejects these views on the nature of the geological record, will
rightly reject my whole theory. For he may ask in vain where are the num­
berless transitional links which must formerly have connected the
closely allied or representative species, found in the several stages of the
same great formation. He may disbelieve in the enormous intervals of
time which have elapsed between our consecutive formations; he may
overlook how important a part migration must have played, when the for­
mations of any one great region alone, as that of Europe, are considered;
he may urge the apparent, but often falsely apparent, sudden coming in
of whole groups of species . He may ask where are the remains of those
 On the Geological Succession of Organic Beings 233

infinitely numerous organisms which must have existed long before the
first bed of the Silurian system was deposited: I can answer this latter
question only hypothetically, by saying that as far as we can see, where
our oceans now extend they have for an enormous period extended, and
where our oscillating continents now stand they have stood ever since the
Silurian epoch; but that long before that period, the world may have pre­
sented a wholly different aspect; and that the older continents, formed of
formations older than any known to us, may now all be in a metamor­
phosed condition, or may lie buried under the ocean.
Passing from these difficulties, all the other great leading facts in
palaeontology seem to me simply to follow on the theory of descent with
modification through natural selection. We can thus understand how it is
that new species come in slowly and successively; how species of differ­
ent classes do not necessarily change together, or at the same rate, or in
the same degree; yet in the long run that all undergo modification to some
extent. The extinction of old forms is the almost inevitable consequence
of the production of new forms. We can understand why when a species
has once disappeared it never reappears. Groups of species increase in
numbers slowly, and endure for unequal periods of time; for the process
of modification is necessarily slow, and depends on many complex con­
tingencies. The dominant species of the larger dominant groups tend to
leave many modified descendants, and thus new sub-groups and groups
are formed. As these are formed, the species of the less vigorous groups,
from their inferiority inherited from a common progenitor, tend to be­
come extinct together, and to leave no modified offspring on the face of
the earth. But the utter extinction of a whole group of species may often
be a very slow process, from the survival of a few descendants, lingering
in protected and isolated situations. When a group has once wholly dis­
appeared, it does not reappear; for the link of generation has been bro­
ken.
We can understand how the spreading of the dominant forms of life,
which are those that oftenest vary, will in the long run tend to people the
world with allied, but modified, descendants; and these will generally
succeed in taking the places of those groups of species which are their
inferiors in the struggle for existence. Hence, after long intervals of time,
the productions of the world will appear to have changed simultaneously.
We can understand how it is that all the forms of life, ancient and re­
cent, make together one grand system; for all are connected by genera­
tion. We can understand, from the continued tendency to divergence of
character, why the more ancient a form is, the more it generally differs
234 D A R W I N ' S G H O S T

from those now living. Why ancient and extinct forms often tend to fill up
gaps between existing forms, sometimes blending two groups previously
classed as distinct into one; but more commonly only bringing them a lit­
tle closer together. The more ancient a form is, the more often, appar­
ently, it displays characters in some degree intermediate between groups
now distinct; for the more ancient a form is, the more nearly it will be re­
lated to, and consequently resemble, the common progenitor of groups,
since become widely divergent. Extinct forms are seldom directly inter­
mediate between existing forms; but are intermediate only by a long and
circuitous course through many extinct and very different forms . We can
clearly see why the organic remains of closely consecutive formations
are more closely allied to each other, than are those of remote formations;
for the forms are more closely linked together by generation: we can
clearly see why the remains of an intermediate formation are intermedi­
ate in character.
The inhabitants of each successive period in the world's history have
beaten their predecessors in the race for life, and are, in so far, higher in
the scale of nature; and this may account for that vague yet ill-defined
sentiment, felt by many palaeontologists, that organisation on the whole
has progressed. If it should hereafter be proved that ancient animals re­
semble to a certain extent the embryos of more recent animals of the
same class, the fact will be intelligible. The succession of the same types
of structure within the same areas during the later geological periods
ceases to be mysterious, and is simply explained by inheritance.
If then the geological record be as imperfect as I believe it to be, and
it may at least be asserted that the record cannot be proved to be much
more perfect, the main obj ections to the theory of natural selection are
greatly diminished or disappear. On the other hand, all the chief laws of
palaeontology plainly proclaim, as it seems to me, that species have been
produced by ordinary generation: old forms having been supplanted by
new and improved forms of life, produced by the laws of variation still
acting round us, and preserved by Natural Selection.
 C H A P T E R X I

G E O G RAPH I CAL DISTRI BUTI O N

O n geography and change - Present distribution cannot b e accounted for by

differences in physical conditions - The five mediterraneans: their in­
vaders and their residents - Importance of barriers - Means of dispersal
by land, sea and air, and by changes of climate - Centers of evolution -
Dispersal during the glacial period - The uneasy Earth and the geogra­
phy of life.

In the Arizona desert in the early 1 990s an island was built. Biosphere
Two, as it was called (the Earth itself was Biosphere One), tried to isolate
itself from the pollution and vice around it. The plan was to create a
world that never was-an unadulterated place in which man and nature
could live in harmony. Eight Biospherians set up a community sufficient
unto itself, an ecosystem in the balance that had once, they claimed,
ruled our planet. The immense greenhouse was sealed off from the air as
a two-hundred-million-dollar microcosm of diversity, from desert to
rainforest to million-gallon ocean.
Within a year, its inhabitants faced reality. Microbes in the soil caused
the amount of carbon dioxide to shoot up and the level of oxygen to fall
to that on the summit of Mont Blanc. Vines strangled whole sections of
the Biosphere as other plants died out. The animals had even less success.
Nineteen of twenty-five kinds of vertebrate perished, as did all the insect
pollinators (which meant that most of the plants were doomed). The
"desert" grew grass and the water could be kept clean only by cutting
great mats of algae. In 1 994 the Biosphere was abandoned.
236 D A R W I N ' S G H O S T

The United States itself was established for the same reason as that
great greenhouse. It was a new world, isolated from the evils of the old;
a chance to start again in harmony with nature. Much of what its pio­
neers saw was familiar. Laurel, walnut and ivy; robins, blackbirds and
larks, all were there.
Although the new England seemed much like that left behind, it was
not. The American laurel is a poisonous heath plant, unrelated to Euro­
pean plants of the same name, and the robin is a thrush. The settlers'
names were based on nostalgia rather than biology. Each colony-under
glass, and under God-faced the unpalatable truth that when life is iso­
lated, it changes.
The Origin begins with travel: "When aboard HMS Beagle, as natu­
ralist, I was much struck with certain facts in the distribution of the in­
habitants of South America . . . " That sentence is full of meaning. It was
the door to what became a science of its own. Darwin realized that to
prove the fact of evolution all that is needed is to go somewhere else. As
he saw, some of its best evidence comes from maps, with the geography
of life "a grand game of chess with the world for a board."
There is a striking parallelism in the laws of life throughout time and
space: the laws governing the succession of past times being nearly the
same as those governing at the present time the differences in different
areas. Why should Australia be the only place with kangaroos? Do we
need St. Patrick to explain the absence of snakes from Ireland? And why
is coal, the remnant of tropical forests, found in the Antarctic? All this
makes sense if existence altered as it moved. Geography is an escape from
the sad truth that (except through fossils) we cannot visit the past.
In considering the distribution of organic beings over the face of the
globe, the first great fact that strikes us is that neither the similarity nor
the dissimilarity of the inhabitants of various regions can be accounted
for by their climatic or other physical conditions. There is hardly a cli­
mate or condition in the Old World that cannot be paralleled in the
New. Notwithstanding this parallelism in the conditions of the Old and
New Worlds, how widely different are their living productions!
The wine counter of any supermarket (except in France) shows how
different places can, given the chance, support identical forms of life, al­
though, most of the time, they do not. It may sell twenty different
Chardonnays. They taste much the same, although some are smoky and
 Geographical Distribution 237

others redolent of butter, peach or passionfruit-subtle contrasts, but

important to those old and rich enough to disguise their favorite drug.
Some bottles are expensive-Chablis can cost eighty dollars-others as
cheap as South Mrican or Chilean labor can make them. Those who buy
them are out of fashion, and Britain has an Anything but Chardonnay
club, founded to stamp out the ubiquitous invader.
Distant parts of the world have millions of vines that grow the grape.
Only an expert can separate the best French from, say, the best Australian
product of that name. Even the botdes tend to look the same. Whatever
the price, and from wherever they come, their labels often show pantiles
and vineyards set among scrub-covered hills. From France (the home of
Chardonnay) to Western Australia (where it was first grown only twenty
years ago) , via Chile, California and the Western Cape, the weather and
the landscape are similar. The wine trade makes, although the oenophile
might deny it, an identical product in each location.
The Chardonnay grape flourishes in these scattered places because
each has the same weather and soil. They are the five great Mediterranean
ecosystems of the globe. All are about the same distance from the equa­
tor, and all have wet winters and hot dry summers. Vines, first domesti­
cated around the Mediterranean itself, now grow in every region lucky
enough to share its climate.
Although much has been displaced by grapes, each of the five has its
own native vegetation-maquis or garrigue in the Mediterranean, fyn­
bos in South Mrica, chaparral in California, mattoral in Chile, kwongan
in Australia. Whatever the local flora might be called, it is made up of
hard, spiny and fire-resistant shrubs. To wander through maquis, kwon­
gan or fynbos is to sniff the heady scents of nature to the sound of birds
while being torn apart by thorns. Such ecosystems, wherever they might
be, represent some of the most diverse communities on earth. The
Mediterranean itself has twenty thousand kinds of higher plant (a tenth
of all those in the world) . The five communities taken together contain
a greater variety of plants than the whole of tropical Africa and Asia com­
bined. The birds, too, are marvelous. Almost as many kinds breed
around the Mediterranean as in the whole of the rest of Europe. A wet
winter and a hot dry summer are, for some reason, great promoters ofbi­
ological variety.
Similar as they seem, the native plants and animals of each place (un-
238 D A R W I N ' S G H O S T

like those introduced by man) are in reality quite unalike. Each faces
much the same conditions-that, after all, is why vines do well-but
their resemblance, in thorns or in song, is superficial. If we compare large
tracts of land in Australia, South Africa and western South America, we
shall find parts extremely similar in all their conditions, yet it would not
be possible to point out three faunas and floras more utterly dissimilar.
Those natural communities, and their twins in the Mediterranean and
California, have almost no biological affinity. Instead, all have their own
residents, descended from indigenous ancestors and adapted to a regime
of fire and hard grazing. In southern Europe, dwarf oaks do the job, in
South Africa heathers dominate, and in Australia most of the trees are
wattles. Chilean spines are as likely to be those of cacti as of shrubs. The
plants descend from separate sets of rainforest ancestors that flourished
before the last glacial epoch.
Nature's work is in great contrast to our own. The natives look simi­
lar but are in fact distinct, while the wine tastes the same because it is the
same. Faced with the same conditions, ready for exploitation by the nar­
row and commercial mind of man, biology has molded different sets of
inhabitants to do the same job. Their resemblance is on the surface, and
a glance at the actors in each ecological play, similar though their plots
might be, shows that all have their own history. The likeness of fynbos,
kwongan, mattoral, chaparral and maquis reflects five independent re­
sponses to the same force of natural selection. The locals-unlike the
vineyards-make the case for evolution.
Except in gardens, there are no cactuses around the Mediterranean, or
wattles in Chile. Such plants are confined to their native land because
they are confined behind barriers-the Sahara or the Andes-that pre­
vent them from spreading farther. All plants or animals face obstacles
that exclude them from places where they might otherwise do well. No
two marine faunas are more distinct, with hardly a fish, shell or crab in
common, than those of the eastern and western shores of South and
Central America; yet these great faunas are separated only by the narrow,
but impassable, Isthmus of Panama. Any plant or animal will migrate to
wherever it can, given the chance. As soon as an impediment is removed,
life pours through and destroys any less adapted forms in the way. To
breach a barrier is to experiment with evolution.
In the 1 980s, Israelis were forced to give up bathing. Their beaches
 Geographical Distribution 239

were clogged with immigrants: poisonous jellyfish, twenty i n every cubic

yard of water. They choked power-station inlets and forced fishermen to
stay in port because their nets were filled with decaying flesh. The culprit
was a native of the Red Sea that had broken through a five-million-year­
old barrier.
The Mediterranean has an unexpected history. It has been not one,
but several, seas (and, now and again, deserts). Five million years ago, it
was dry, because the last of many great evaporations had left a layer of
salt a mile thick across its floor. Some of its inundations were from its
eastern end and brought warm-water plants and animals that were well
adapted to the tepid waters of what is, of its nature, a subtropical ocean.
The Mediterranean's modern waters, though, came from the Atlantic,
over the great Falls of Gibraltar. Because it last filled from the west, the
Mediterranean is now a warm sea filled with the descendants of plants
and animals from a cold ocean. At the end of the last ice age, just eigh­
teen thousand years ago, there were polar bears in the South of France,
and even today the Mediterranean has its own whales and seals.
The Suez Canal was opened in 1 869. It was the successor to several
earlier links with the Red Sea, the first made three thousand years ago by
Rameses II of Egypt (the builder of the Abu Simbel temples, themselves
inundated by today's engineers) . The present Canal is half a mile across
in parts of its hundred-mile length. It acts like a giant new Mississippi as
its waters flow downhill from the Red Sea to its younger cousin. The con­
nection between the seas allowed their animals, confined for millennia,
to move.
The traffic was one-way. Three hundred Red Sea natives made it to the
Mediterranean, almost none the other way. The migration continues at
a rate of ten new forms a year, with the newcomers flourishing at the ex­
pense of the locals. A third of the Israeli catch now consists of Red Sea
fish and the wave of aliens has reached Sicily. Many fish are in decline.
The survivors manage only because they leave space for the invaders. The
local mullets, for example, live in deeper waters than do their newly ar­
rived relatives.
The immigrants succeeded because they are more adapted to today's
Mediterranean than are its natives. The locals have had no time-nor, in
the absence of competition, much need-to respond to the challenges
presented by their home's warm and salty waters since the sea last filled.
240 D A R W I N ' S G H O S T

As a result, they were soon driven out by the subtropical outsiders. The
breach in the barrier allowed animals to swarm into the new space. In
time they will fill it-as far as they are able, and until they are stopped
by others more suited to the conditions. Wherever it arises, and however
it travels, a species will move on until something restrains it. As it does,
it must evolve or die.
All this means that the continents generate their own mixtures of in­
habitants. On each one, successive groups of beings, specifically distinct
yet clearly related, replace each other. On the plains of La Plata we see the
agouti and bizcacha, animals with nearly the same habits as our hares and
rabbits, but they plainly display an American type of structure. We look
to the waters and we do not find the beaver or muskrat, but the coypu
and capybara, rodents of the American type. We see in these facts some
deep organic bond, prevailing throughout space and time. The natural­
ist must feel little curiosity who is not led to inquire what this bond is.
The bond is inheritance, modified by natural selection. Such commu­
nity of descent casts its net wide. The world can be divided into great do­
mains-the New World Arctic and tropics, their equivalents in the Old
World, Mrica, the Far East and Australia, with more beneath the sea.
Each has its own identity, shared not just by mammals, but by insects,
snails, worms and trees. The differences are not absolute, and many crea­
tures range over more than one province, but the existence of such huge
areas of affinity is itself evidence of a shared past.

Means of Dispersal Once evolved, plants and animals face a constant

struggle against the pressure of their own numbers. They move as far as
they can, by their own efforts or with the help of others. Some can travel
for huge distances by land, sea or air, while others are confined to the
place where they were born.
Often, a hopeful migrant faces an impassable barrier. The capacity of
migrating across the sea is more distinctly limited in terrestrial mammals
than perhaps in any other organic beings; and, accordingly, we find no
inexplicable cases of the same mammal inhabiting distant points of the
world. For other animals the sea is a highway. Pytheas of Massalia was the
first sailor to venture beyond the Pillars of Hercules {and the first Greek
to visit the British Isles}. As he explored the Atlantic shore of Spain he
noticed that the ocean flowed south, like an immense river, an okeanos.
 Geographical Distribution 24 1

His river runs on as the Canary Current, part of a great girdle of water
that flows around the North Atlantic. The Gulf Stream has the force of
three hundred Amazons; a river forty miles across that takes a month to
cross the Atlantic and is but one of many great conveyor belts bearing
flotsam across the world.
All oceans have their currents. About a tenth of their waters are always
on the move. Most flow on the surface (although slow streams in the
deeps take a thousand years to carry icy water from the Antarctic to the
Galapagos). The top ten feet of the ocean store as much heat as does the
entire atmosphere. The movements of water are started by the rotation
of the Earth. Because water at the Equator is warm, it expands. As a re­
sult, the sea in the Caribbean is three inches higher than at Newfound­
land. The warm water runs downhill toward the poles and is twisted in
a clockwise direction in the Northern Hemisphere (and its reverse on the
other side of the equator) as the world turns. The wind obeys the same
rules, and its storms help generate the sea's drift as they spin round the
globe.
Some animals float to a new home. The surface of the water between
the Galapagos Islands carries dozens of insects. Most will drown, but a
few will make the journey. Others hitch lifts on the many vehicles that
pass by. Every day, ten million pieces of garbage-bottles, bags and plas­
tic sheets-are dumped from ships. From the land, the sea receives much
more. The island of Pitcairn is three thousand miles from the nearest
mainland. Its best-known detritus was the Bounty mutineers, who landed
in 1789. The island was so remote that their refuge was not discovered
until all but one had died. Nowadays, its beaches are as filthy as any in
Europe, with a piece of rubbish every yard. Pitcairn has the European
mix of buoys, bottles and bags, but a relative shortage of disposable dia­
pers. The empty liquor bottles suggest that many of the migrants come
from South America. Not all the flotsam is useless, as the local land crabs
are fond of shoes as shelters, but it is a dismal reminder of how the most
remote places have been forced to join the modern world.
Some marine debris is still mysterious: why are twice as many left
shoes as right washed ashore in Holland, while the opposite is true for
Scottish beaches? All has a message for evolution. Nowhere is isolated.
Given the chance, plants and animals will float, fly or drift through the
air to reach the most remote parts of the earth.
242 D A R W I N ' S G H O S T

Plenty of animals travel on rafts (shoes and bottles included). Off the
coast of Cuba float substantial islands of vegetation that may bear ma­
ture trees. On Christmas Island, three thousand miles from the Ameri­
cas, American redwood and walnut are used as firewood because so many
come ashore. Canoe builders on coral atolls in the deep Pacific once de­
pended on logs that had floated halfvvay across the world. A complex eti­
quette determined who got the biggest. The natives procured stones for
their tools, solely from the roots of drifted trees, these stones being a
valuable royal tax. Many of the unsinkable vagrants bore not just stones,
but animals and plants. Glaciers, too, deliver their contents as they float
from the ice caps. Antarctic boulders on the sea floor off Cape Town were
brought within the past ten thousand years. Among them are the re­
mains of penguins. However, the sea can be a formidable barrier. Few
freshwater fish can manage to cross a strait more than a couple of miles
across, elephants stop swimming at thirty miles, and tortoises and snakes
cannot manage more than five hundred.
To find a raft is not enough, because the travelers must survive their
journey. Most land plants find it hard to deal with seawater. Their seeds
do rather better. Some simple experiments prove how tough they are.
Often, dried seeds do best. To dry stems and branches of many different
plants and to place them on seawater shows that some when dried
floated; for instance, ripe hazelnuts when dried floated for ninety days
and then germinated. As many currents run at sixty miles a day, plants
might be floated across miles of sea; and, when stranded, if blown to a
favorable spot by an inland gale, they would germinate.
Seeds are the genetic memory of the plants that bore them. Many sur­
vive not for weeks, but for years, with germination sparked off by a
change in the environment.
An unplanned test of the power of the seed began when the wheat­
fields of northern France were abandoned after the economic collapse
that followed the Franco-Prussian War of 1 870. The crop was in those
days full of weeds. Poppies were everywhere. The plant can generate
thirty thousand seeds in a square yard of soil. After the French collapse
the farms of Flanders stayed grazed and flowerless until 1 9 14. Then, the
land was cultivated again-not with plows, but with swords, shells and
blood. Once disturbed, the poppies bloomed at once. A quarter of a cen­
tury later, in the next round of human folly, the Natural History Mu-
 Geographical Distribution 243

seum in London was bombed. The fire hoses caused many seeds to ger­
minate, among them a mimosa, collected in China in 1 7 13, and revived
in a sudden Hood two centuries on and five thousand miles away. The
poppy is much used as a symbol of war's destruction; but it bears a more
hopeful message about how well life can survive in the face of adversity.
With a third of the world's plants in danger, there is a new interest in
conservation. The toughness of seeds is a great help-and is a reminder
of their vital role as containers for genes. A Millennium Seed Bank at
Kew aims to store dried and frozen seeds of a tenth of the world's kinds
of plants (together with the whole of the British Hora). Most should last
for centuries. Simply to dry the seeds of beet, rice or elm allows them to
survive for a decade and more. Some weed genes allow the plants to lie
low even in good times and to sprout over months rather than all at
once-which is useful when it comes to long and risky journeys.
Plenty of travelers Hy, rather than Hoat, across the globe. Every few
years, after an Atlantic storm, dazed North American birds reach Europe.
Most die, but the survivors carry a cargo. The crops of birds do not se­
crete gastric juice and so do not in the least injure the germination of
seeds. All the grains do not pass into the gizzard for twelve or even eigh­
teen hours. A bird in this interval might easily be blown five hundred
miles, and hawks are known to look out for tired birds. Some bolt their
prey whole, and after an interval disgorge pellets, which, as seen in ex­
periments made in the Zoological Gardens, include seeds capable of ger­
mination.
Locusts soar upward as the sun sets and are caught in the winds of the
upper air, to move fifty miles a night. They concentrate on the edge of
weather systems to form plagues as they descend to earth. Less conspic­
uous things also move-as anyone who suffers from hay fever knows.
Every summer afternoon, on the Costa del Sol, a great cloud of mari­
juana pollen descends from the illicit fields of Morocco. A constant rain
of pollen and spores from South America, mixed with the odd seed and
insect, falls on Signy Island, on the edge of the Antarctic. Few places on
the Antarctic continent itself are warm enough for mosses and liverworts
to grow, but where they can-around hot springs and the like-they do,
evidence that nowhere, remote though it may seem, is safe from mi­
grants. The presence of the same form in distant places is not evidence
that it was created twice, but that it can move.
244 D A R W I N ' S G H O S T

Dispersal during the Glacial Period. The identity of many plants and ani­
mals on mountain summits, separated from each other by hundreds of
miles of lowlands, where the alpine species could not possibly exist, is
one of the most striking cases known of the same species living at distant
points, without the apparent possibility of their having migrated from
one to the other. The pattern arises not because each peak is a separate
factory for the same product, but because a brutal landlord has broken
up great estates of the living world.
Their inhabitants are a relic of ancient ice sheets, now retreated. Gla­
ciers were the first hint that science and the Bible do not coincide. How
could even a believer in Noah's Flood account for the Scottish boulders,
too big for any conceivable deluge, that turned up in Wales, where geol­
ogists found great chunks of the Scottish island ofAilsa Craig? At a con­
ference in Glasgow in 1 840 the young Swiss geologist Louis Agassiz
persuaded the assembled skeptics that these "erratic blocks" (later the
subject of a celebrated spoonerism) were moved not by water, but by ice.
Now, glaciers are known to explain many enigmatic patterns of distrib­
ution.
Most of the familiar landscapes of today are a statement of the power
of frozen water. The evidence of past ice ages is everywhere around us.
The ruins of a house destroyed by fire do not tell their tale more plainly
than do the mountains of Scotland and Wales, with their scored Hanks,
polished surfaces and perched boulders, of the icy streams with which
their valleys were lately filled.
Ice is the commonest rock in the solar system, but Earth is the sole
planet upon which it exists in consort with water itsel£ The balance of
the two has waxed and waned, with rapid swings between ice ages over
the past three million years. Great waves of cold are not common. Bouts
of glaciation hit seven hundred million years ago, and five hundred mil­
lion years later, but for most of the time the world has been warmer than
today. It has cooled for a hundred million years. When the last long win­
ter began, northern Alaska had ferns and lush forests of ginkgo, and the
banks of the Thames were covered by a subtropical forest, with crocodiles
and turtles in its waters.
The Pleistocene, the period just before our own, had sixteen cycles of
cold and warmth. At times, glaciers covered more than a quarter of the
land. Each glaciation lasted about a hundred thousand years, with brief
 Geographical Distribution 245

incursions of warmth, in one of which we live. In the most recent cold

period, ice reached to Missouri and central England and the tropics
cooled and dried. As the sea fell, it exposed thousands of square miles of
land. Water is heavy stuff. The lake behind the Hoover Dam depressed
the deserts around it by six feet. The cloak of ice, too, forced the land to
sink and, after its burden was shed, it rose again. Arctic Canada has ele­
vated itself by a thousand feet-and it still bounds upward at an inch a
year. Scotland's rise is less precipitate, but its release from a frigid past
means that the British Isles are tipping over, with Scotland on an upward
path and the south of England falling into the sea.
When the last glaciation ended, a mere dozen millennia ago, the sum­
mer temperature rose by eight degrees Celsius-the difference between
London and Lagos. Great changes in temperature over shorter times are
recorded in tree rings, in ice cores, and in the sediment of Arctic lakes.
The brief period since the beginning of agriculture has, they show, been
a time of unusually stable climate. Before then, the weather changed
drastically every thousand years or so. Sometimes, within a single year,
the snowfall doubled over the whole of the northern hemisphere as the
temperature fell by ten degrees.
Lesser cycles of cold and warmth continue, and animals and plants
still travel in their wake. The "little ice age," which began in the sixteenth
century and lasted to Victoria's day, explains both the Dickensian
Christmas and the failure of explorers from Cabot to Franklin to find a
sea route around North America. It has switched into today's global
warming. A peak of heat in 1 945 has been succeeded by another. Three
of the five warmest years in England (and the records go back to 1 659)
have been since 1 988, with 1 998 the warmest year ever recorded for the
entire planet.
Satellite photographs show that spring in the temperate parts of the
Northern Hemisphere now starts a week earlier than it did in the 1 960s.
A network of schoolchildren tracks the key events-the first snowdrops,
the earliest oaks to Hower and the first cuckoo. They follow a noble tra­
dition. The descendants of Robert Marsham, of Norfolk, pinned up a
chart of the arrival of spring in 1 736 and kept it for two hundred years.
It recorded how often-and how late in the year-the chamber pot had
a skim of ice. Frozen urine was more of a feature of an eighteenth­
century Easter than of that festival today.
246 D A R W I N ' S G H O S T

Why is the climate so unstable? In part it turns on the Earth's orbit, an

ellipse whose shape changes with a rhythm of about a hundred thousand
years. At its most extreme it causes winters to be colder and summers
hotter as the planet moves farther from and closer to its source of heat.
A series of cold winters is enough to tip the globe into frost. The se­
quence is modulated by a shorter cycle in the Earth's tilt and wobble. As
forests bloom-and later rot-in warm periods, carbon dioxide and
methane escape into the air. These allow energy from the sun to pour in,
but trap the ground's own long-wave radiation to give a "greenhouse ef­
fect" that pushes the temperature up still further. Because ice can pass
through a glacier at five miles a year, it does not take much of a drop in
temperature for it to race across the landscape, or much of an increase for
an ice cap to shrink.
For much of the time the climate is poised on edge. The end of the last
European ice age was marked by two sudden spurts of cold. In each, for
a few centuries, glaciers returned to the Pyrenees and the Alps. The short
interval of warmth between the ice's last breaths lasted a mere moment,
but was enough to allow Mediterranean beetles to flourish in Britain be­
fore they were again wiped out. Today's concern about heat will be suc­
ceeded by anxiety about a new ice age if the warm and salty Gulf Stream
is displaced by cold fresh water from what used to be the ice cap-which,
in parts of Greenland, has lost three feet of its depth in only a decade.
The last great interglacial period, ten thousand years long, ended in just
four centuries as the Atlantic conveyor belt was stopped by melted ice.
As ice retreated at the end of its most recent advance, it forced the an­
imals and plants of cold climates to withdraw with it. Most made it back
to the far north or south, but some were marooned in mountain ranges.
They moved up the valleys as the weather warmed. Now, each range has
its glacial relicts, identical or almost so to those of distant mountains. For
them, the evidence of shared descent remains, but as fragments. They
serve as a record, full of interest to us, of the former inhabitants of the
surrounding lowlands. Such refugees show that the difficulties in believ­
ing that all the individuals of the same species, wherever located, have de­
scended from the same parents are not insuperable.
The evidence of climatic change is in the inhabitants of the mountains
as much as on their scarred flanks. The varying hare (so called because it
goes white in winter) is found in Northern Canada and in Scandinavia-
 Geographical Distribution 247

and in the Alps, a thousand miles away. Insects, too, follow the ice. Two
thousand different kinds are known from Britain's glacial times. Almost
all have gone from these islands, but fewer than twenty are altogether ex­
tinct. Instead, they survive-not in today's subtropical England but in
Siberia, Arctic Canada or the high mountains of Europe. What was once
the commonest dung beetle in England now lives mainly in Tibet. Genes
show that many other forms now widespread (and with no obvious sign
of past disaster) were fractured. In Europe, oaks and grasshoppers have
deep divisions, with a great split between the DNA of west and east
which reflects their isolation and advance from refuges in Spain and in
Turkey. The plants and animals of the north are less variable than are
their southern relatives. As they moved step by step to colonize Europe,
they went through one bottleneck after another. As a result, the genes of
a few formed the fate of their many successors.
Forests are always poised to move. In a churchyard near Edinburgh
live a group of trees known as "the walking yews of Ormiston." Legend
has it that, with some supernatural help, they have shifted from place to
place in response to crisis. And, of course, they have. Yews put out new
shoots from old branches where they touch the ground. When the old
trunk dies, its branch replaces it. In time it grows into a new plant, yards
away from its previous incarnation. No miracle is required.
As glaciers fall back, trees press on behind. In North America, spruce
was the first to migrate, followed by pine. Each traveled at a great rate­
a mile every three years. Behind came the chestnuts. Their progress was
more dignified, at a hundred yards a year, but for a large tree with heavy
seeds that is almost a sprint. Birnam Wood, with its pines, could have
marched on Dunsinane (ten miles away) well within Macbeth's lifetime.
The forest journeys continue. In the Alps, the remains of a conflict
more recent than the Somme were revealed when shelters dug deep into
the ice in 1 944 were exposed to view. The retreating glacier at Glacier
Bay in Alaska has withdrawn by sixty miles since it was first seen by Eu­
ropeans in 1 750. It took just a century for mature forest to cover its
empty bed. The birds and insects of the north arrived almost at once.
The climate may have changed too fast for them to evolve out of trou­
ble, but instead they moved.
Life's geography is like that of nations. It seems natural that a state's
frontiers should be determined by the wishes of its inhabitants and those
248 D A R W I N ' S G H O S T

of its neighbors, but that ignores the facts of politics. Most Britons have
the good fortune to live in a territory that defines itself, on an island. Few
countries-or animals-are as lucky. History is a record of how difficult
it can be to divide up the landscape.
Species and countries each vary in what they occupy. The Devil's Hole
pupfish fills a single freshwater spring in Death Valley. It has its being in
a space the size of a large room (and was once reduced to a global habi­
tat of a bucket when its home dried up and its occupants were rescued
by an alert conservationist). The blue whale, in contrast, roams all the
world's oceans. In much the same way, Russia, the largest political unit
in the world, is a million times bigger than Nauru, the smallest.
The range of most birds, flies or plants in the tropics is smaller than
that of those in the north. The same is true of states, with tropical na­
tions a quarter the size of those overlapping the Arctic. The geography of
animals and plants turns on hard times long ago. As the ice retreated,
only those able to survive in the chill landscapes left behind could follow.
The homelands of those inconspicuous birds, the warblers, vary by a
hundred times. Those with the largest ranges are in the north, because
they could move with the birch or pine forests that marched behind the
ice. Birds whose haunts (such as rhododendron thickets) are restricted to
warmer places are perforce confined to smaller spaces. Most rare plants
and animals are found in the tropics, because they have not been able to
expand into the space available to their hardier kin.
The Garden of Eden is always painted as a lush and sultry place. From
biology's point of view, too, the tropics are a great and ancient city, with
more inhabitants, more energy, more water, more production and, be­
cause the land has not been wiped clean by glaciers, more time for spe­
cialists to evolve than in the icy north and south. A fifth of all the world's
kinds of plant are found on a two hundredth of its land. Such centers of
origin are tropical, in Madagascar, Malaysia or Central America. Arctic
Canada possesses ten kinds of ant, compared to two thousand in the
same area of tropical South America; and Hong Kong, at four hundred
square miles, has more kinds of birds, mammals, insects and plants than
the whole of the British Isles. Such gradients from Pole to Equator are as
old as the fossil record itself They prove the claim of Adam Smith that
division of labor is proportional to demand. Although a workman who
made only nails would never survive in a Scottish village, in a large town
such a narrow specialist would prosper.
 Geographical Distribution 249

The great tropical factories of life and the rigors of the last ice age
mean that the north is a new nation for all its inhabitants . .As plants and
animals followed the glaciers as they retreated they filled vast tracts of
country. Many had no real need to change, because their new homes
were no colder than those they had just left. Evolutionists often see the
environment as rigid and life as flexible. The story of the ice shows that,
quite often, the opposite is true, and that conditions change more rapidly
than those who suffer them. Life is always on the lookout for somewhere
more comfortable. It is much easier to migrate than to evolve. Northern
roots lie in southern parts, and the inhabitants of half the world are tes­
timony to a catastrophe of just ten thousand years ago. Much as Scottish
Nationalists might disagree, to cut themselves off from their neighbors
may destroy their chances of survival when the cold returns.

Some journeys do seem improbable. Why are there ostriches in Mrica,

emus in Australia and rheas in South America? All are large, aggressive
and-above all-flightless. Their genes show them to be relatives who
descend from a wingless ancestor. How did they make the trip? Trees,
too, show unexpected patterns. The southern beech of South America
has close kin in Australia, in South Mrica and even in the Antarctic, pre­
served beneath the ice. Ten thousand miles separate these places but their
forests are almost the same. Pheasants, electric fishes and tree frogs each
have ties between their members in Australia, South America and New
Guinea, with more distant links even as far as Europe, but none are much
good at ocean crossings. Three thousand miles of sea are a barrier for an
eagle, let alone a heavy beech seed or a flightless bird.
The geography of the sea is even harder to understand. Some of its in­
habitants are found at the northern and the southern ends of the Earth
but nowhere in between. Their distribution has lasted for two hundred
million years so that the simple notion that they floated across the equa­
tor on some recent iceberg does not work. Mussels, scallops and whelks
all have close relatives in Arctic Canada, Siberia, Chile and the Antarc­
tic. How can this be?
Such patterns might seem to be evidence against descent with modi­
fication. It is incredible that individuals identically the same should ever
have been produced through natural selection from parents specifically
distinct. To find the same plants and animals in isolation at opposite ends
250 D A R W I N ' S G H O S T

of the globe leads some to conclude that the same species must have been
independently created. The truth is simpler, but almost as startling.
Scott of the Antarctic froze to death eleven miles from safety. His
sledge was weighed down with thirty pounds of rock dragged by ex­
hausted men for hundreds of miles. The rocks helped kill Scott, but held
a crucial clue about history. Some contained fossils of a tropical tree, oth­
erwise found in India and Mrica.
Much later, the skull of a thirty-foot dinosaur from eighty million
years ago was found in Madagascar. It is quite different from those of
Africa, a hundred miles away. With its rough surface and prominent
horn it looks like others found in South America and India. In today's
atlas, all this makes rather little sense. Perhaps, though, today's atlas is not
the one to use.
Some odd patterns of shared geology suggest that an earlier edition is
called for. The folds of the Appalachian Mountains in the eastern United
States can be traced to a parallel set of rocks in Ireland and Brittany. In
the same way, parts of India, South Mrica and tropical South America
bear scars made at about the same time by ancient glaciers.
Once there was talk of land bridges between the continents, used as
highways for trees, dinosaurs and more on their global journeys. The
connections rose and fell almost on demand. So many links between dis­
tant places were needed to explain the distribution of animals and plants
that the seas were filled with theoretical Atlantises. Those who drew the
maps bridged every ocean and united every island to some mainland. In
an attempt to explain the wide distribution of plants and animals, the re­
motest islands were seen as the wrecks of sunken continents.
The truth is more remarkable, for the land itself, rather than the seas,
is on the move. Continents now far apart were once part of the same
mass. They have split and taken their inhabitants with them, to become
great arks of rock that wander the globe and, now and again, collide. The
ancient movements of a fluid Earth explain much of the geography of
plants and animals today.
The fragments of what were once c�ntinents travel at about the rate
that fingernails grow but, in time, that is enough to shift them for great
distances. The expansion of the Atlantic has been measured from satel­
lites (leading to a rare joke by the Duke of Edinburgh: that at last Britain
knows where it stands in relation to the United States) . If the Pilgrims

-
 Geographical Distribution 25 1

were to repeat their transatlantic trip, they would find themselves out of
place, not because Puritanism and stovepipe hats would seem eccentric
in America's new conformism, but because America itself has moved on.
Their landing place, Plymouth Rock, is fifty feet farther west than it was
in 1 620. William Bradford, ifhe stepped off the Mayjlo wer today, would
get his feet wet.
Leonardo da Vinci had noticed that the continents would fit together
like jigsaws. A Frenchman, Antonio Snider-Pellegrini, revived the idea in
the year before the publication of The Origin, in his forgotten work Cre­
ation and its Mysteries Revealed. Unfortunately for him, he used the ob­
servation as a proof of Noah's Flood rather than the key to the Earth's
structure. In 1 9 1 2, the first great world continent was given the title of
Pangaea, the universal land, long before evidence of its existence was
found. The idea was ridiculed by one geologist as "utter, damned rot!"
because there seemed no force able to drive a land mass across the globe.
Although some claimed that the continents had been pushed by the
Earth's spin as they plowed through the ocean floor, that made no sense.
Our planet is less solid than it seems. Deep inside, the core is liquid,
and on the surface-rather like the lines of movement in a pot of boil­
ing porridge-are upwellings of molten rock. Most reach the surface in
long chains of submarine mountains. These midocean ridges were dis­
covered in the nineteenth century by the Atlantic Telegraph Company,
whose engineers laid a cable between Europe and North America. They
assumed that the sea floor was flat, but their connection broke within
weeks because it was suspended between peaks higher than the Alps.
The forty thousand miles of ridge mark the lines where the fluid con­
tents of the globe spew into the seas. As they congeal, the fresh rock
moves outward to make a new ocean floor. The continents float on the
Earth's liquid core, their keels embedded deep within it. They are pushed
apart as the sea floor spreads. The flow from the center means that the
edges of the oceans are cooler and thicker than the submerged ridges, and
sink into the mantle below, with the younger rock dragged behind. The
constant gain and loss of rock means that the ocean floor is young com­
pared to the land. Near the ridges it dates from the past million years or
so, at the edges of the continents from about two hundred million years
ago. Most of the sea bottom is younger than the dinosaurs.
Iceland has the misfortune to sit astride the lava factory, the Mid-
252 D A R W I N ' S G H O S T

Atlantic Ridge itself The country gets bigger by the day. Most of the is­
land is less than twenty thousand years old. In 1 783, huge rifts were
formed as a hundred square miles of new rock belched from below. A
cleric of the time wrote that: "Those terrors that fell over and upon us I
can hardly describe . . . In the middle of the Hood of fire great cliffs and
slabs of rock were swept along, tumbling about like large whales swim­
ming, red-hot and glowing." One Icelander in five died of suffocation by
noxious gas, or of famine. The infernal stench reached Europe, causing
widespread fears of imminent damnation.
When two plates meet out in the ocean, the older, cooler and heavier
slips beneath its neighbor. As its mass slips back into the liquid core it
pulls the ocean Hoor behind it. Along the line of sinking rock a trench is
formed, and volcanoes burst through to give great circles of islands. They
include the Aleutians, part of the "Ring of Fire" around the western Pa­
cific (an ocean increasing in width by two yards a decade). When two
continents crash, a mountain range is thrust toward the skies. That is
why the summit of Everest is formed of rocks made in a shallow sea.
The atlas of the past holds many surprises. To reconstruct it, all that is
needed is to subtract from the map of today the ocean Hoor made since
the date in question. The key is in the Earth's magnetic field. Every few
hundred thousand years, what was the North Pole becomes the South
until the poles reverse once more in their endless dance. The switch is
recorded in the rocks. Great stripes of magnetic reversal across the bot­
tom of the ocean mark their movements. As band after band of older ma­
terial is taken away, the continents reveal their ancient positions. Before
about hundred and eighty million years ago, no ocean Hoor is left and
the information must come from matched geological sequences in dif­
ferent places.
The land itself bears evidence of its journeys. At St. Martin in western
Canada, Manicougan in the east, and Rochechouart in France are three
comet craters dating from two hundred and fourteen million years ago.
At first, their shared age seems a coincidence. Rearrange the continents
to their position at the time of the collision, and the craters lie on a
straight line, proof of the breakup of a comet just before it struck and of
the slower decay of the once solid earth upon which it fell.
Leonardo was right. The Americas and the Old World were at one
time joined. So were many other places. Five hundred million years ago,
 Geographical Distribution 253

there was but Pangaea, a single mass of land. This was later separated by
the Tethys Sea into two great continents-Gondwanaland (made up of
much of the present India, South America, Africa, Australia and Antarc­
tica) and Laurasia (now North America, much of Southern and Western
Europe, and Asia) and a smaller one, Baltica (Northern Europe and
Scandinavia) . A hundred million years ago, Gondwanaland itself broke
up. Several sections declared their independence and drifted northward
to make parts of Europe, Tibet and two pieces of China. Madagascar,
with its unexpected dinosaurs, is a piece broken off Gondwana long after
Africa gained its identity; part of a lost world stranded two hundred
miles off a foreign land when its birthplace was shattered by movements
of the crust.
The face of the Earth was not much like that of today until the ex­
tinction of the dinosaurs. Not until fifty million years ago did Australia
separate from Antarctica and could Europe, adrift from Greenland, at
last cement its relationship with Asia. By then, the world could be navi­
gated, more or less, with a modern chart.
As the continents drift like slow ships, they take their passengers­
alive and long dead-with them. An animal that started its journey in
what is now Australia may have links with others in Africa or South
America. Penguins and flightless birds like the ostrich and the rhea are
scattered across South America, Australia, South Africa and Antarctica
because new oceans have divided their ancient home. DNA shows the
rhea of South America and the Australian emu to be more similar than
either is to the ostrich. Genetics and geology tell the same story, for their
tale is that of the continents. South America split from Africa, but stayed
in contact with Australia via a junction across the Antarctic.
Evolution itself reconstructs the movements of the land. The first fos­
sils of birds and mammals, each evolved from separate sets of reptiles, are
separated by a giant gap from their modern forms. A clock based on the
genes of today times their great radiations at a hundred million years ago,
when Pangaea had broken up and the world was more fractured than it
has ever been. Europe, the Americas and Africa were each divided into
several islands. Dinosaurs, frogs and toads also split into a variety of
forms at that time. The histories of life and of the continents are, it
seems, close companions.
The restless lands acted not just as arks and cradles, but as funeral
254 D A R W I N ' S G H O S T

ships. The fossils of plants and animals that met their end on the same
land mass have moved to form postmortem alliances with others that ex­
pired much later. They are a reminder that the Earth-and its inhabi­
tants-have been in turmoil since they began and a proof of dramatic
changes in the atlas at a time long before the ostrich split from its flight­
less cousins.
In spite of today's resurgence of Celtic racial thinking, Scotland and
England seem quite similar places. Their rocks show that this is not at all
the case. The ancient limestone around Durness in the far north of Scot­
land and the matching beds in northern England each contain many fos­
sils of trilobites. The Scottish versions are covered with lumps and warts.
English rocks of the same age hold nothing like them, although they
have many trilobites of their own. The Scottish fossils are in fact more
similar to the trilobites found in parts of North America than they are to
their relatives a few miles to the south. The rocks of Scotland tell the tale
of an upheaval that formed these islands five hundred million years ago.
The Stone of Scone is an emblem of Scottish identity. The nation's
kings were crowned upon it until it was removed to Westminster Abbey
in 1 296 to do the job for England. Seven hundred years later, faced with
a slump in popularity north of the border, the Conservative government
sent it home in a political association of object and nation more typical
of Serbia than Britain. It can now be viewed in Edinburgh Castle, at
£5.50 a time. Dr. Johnson's comment on another piece of Celtic geology,
the Giant's Causeway in Ulster, comes to mind: "Worth seeing, yes; but
not worth going to see."
According to legend, the Stone was already well traveled. It was the
pillow upon which Jacob slept and dreamed his celebrated dream of a
ladder of angels on their way to heaven. It passed with divine help to
Egypt, Sicily, Spain and Ireland, and enabled the Scots, in the Declara­
tion of Arbroath in 1 320, to define themselves as a Lost Tribe of Israel.
What of the real movements of the Stone over that vast passage of
time? Jacob's Scottish pillow is made of pale yellow sandstone. Three
hundred and fifty million years ago, it began its existence in shallow trop­
ical waters as silt and sand were washed from a range of mountains. Over
many years the grains consolidated into the rock from which the stone
was made. The journey from its brief exile in London, up the slope of
glacial rubble known as the Royal Mile, to the granite island in the ice
 Geographical Distribution 255

upon which sits Edinburgh Castle, seems fair restitution, the righting of
a geological wrong.
But geology is above politics. Two hundred million years before the
stone was formed, Scotland itself was on the move. Much of the country
was on holiday abroad. A vanished ocean called Iapetus stretched be­
tween England and its northern neighbor and split Ulster from Eire. The
rocks that were to become what Nationalists insist on calling the Stone
of Destiny were then in North America. Soon, Iapetus-an ocean as
broad as but three times older than the Atlantic-began to close.
Until it did, England was part of a separate continent, Avalonia
(named for the island to which Morgan Ie Fay carried King Arthur) .
Quite where its northern shore ran all those years ago is hard to say, be­
cause most has been buried by the sediments scraped from its floor to
make the Southern Uplands of Scotland as the ocean closed.
A series of cracks in the landscape-faults-can be traced across
northern England and Ireland. These, together with chemical differences
in the rocks of the ancient continents, hint at where the frontier used to
be. In Ireland the boundary was south of today's political line. It started
north of Dublin and reached to the mouth of the Shannon. On the
mainland, the northernmost remnants of Avalon are near the town of
Moffat in the Scottish Borders.
Wherever their encounter, the desire for Scotland to reunite with En­
gland was such that the two halves of this island charged toward each
other at the extraordinary speed of a foot a year. Scotland skated so
quickly across the globe to fall into the arms of its southern neighbor be­
cause its roots are deeper within the liquid earth than are those of most
continents. When their marriage was consummated, life crossed the gap,
with invasions and extinctions as great as those of the Mediterranean
from the Red Sea, millions of years later.
The clash of Caledonia with Avalon threw up a range of mountains as
high as the Himalayas on the north side of the narrowing strait between
them. Their stumps are the Scottish Highlands, and their eroded remains
the source of the sand that settled, in a shallow sea full of giant sea scor­
pions, to become the Stone of Scone.
Scottish Nationalists may hope to reopen Iapetus and reverse the Pa­
leowic Act of Union, but geology shows that the real home of the Stone
of Destiny is in Newfoundland. There, Scotland's trilobites evolved and
256 D A R W I N ' S G H O S T

died; their fossils a reminder of how that nation-and every other-has

moved around the globe far more than its inhabitants did when they
were alive. Such postmortem journeys also explain the bands of coal and
fossil coral that link Central America, southern Europe and much of
Asia. They are remnants of a time when all were still part of a great trop­
ical continent.
And what of the future? If the continents continue to move at their
present rate, in fifty million years America will be close to Asia as the At­
lantic broadens and the Pacific gets narrower. Australia will rush north­
ward to collide with Japan, and the eastern part of Africa will declare
independence as the Rift Valley becomes a sea. The Straits of Gibraltar
will soon close, and the Mediterranean may again dry into a salty plain
before it disappears for ever.
Many lands have moved around the charts. Atlantis has come and
gone, but the Island of Buss (supposed to have been discovered in mid­
Atlantic by the Frobisher Expedition in the sixteenth century and big
enough to take three days to sail around) shrank and shifted as successive
travelers failed to find it. It was demoted first to a reef and then to the
fable it had always been. Maida Island, spotted near Newfoundland at
about the same time, ended up in the Caribbean on a map published as
late as 1 906.
Land bridges, Atlantises, and all the other myths dreamed up to ex­
plain the distribution of plants and animals are less remarkable than the
truth: that the world has evolved as much as have its inhabitants.
 C H A P T E R X I I

G E O G RAPH I CAL D ISTRIBUTION-CONTINUED

Babies as islands - New lands and their colonists - On the nature and in­
habitants of oceanic islands and their relation to the mainland - Evolu­
tion by accident - The plants and animals of Hawaii - Freshwater
lakes - The fragility of islands and the grand simplification of life -
Summary of the last and present chapters.

A newborn baby enters the world and is at once corrupted. At birth, its
body is sterile, free of the molds, bacteria and mites that infest the rest of
us. From a bacterium's point of view, the child is a large, fertile and un­
derexploited island. Within moments, it is colonized. Some of the in­
vaders come from its mother's guts, others from the midwife or the
proud father. The pioneers are a small sample of the millions of bacteria
that float through human entrails. For a time the infant bowel is volatile
and unsettled. As any parent knows, that has dramatic effects on the
ecology of the territory upon which the invaders make their home.
Within a few days, the child contains ten times as many bacteria as it
does cells of its own. From its neighbors, clean as the household might
be, comes a rain of immigrants. People, some say, grow to look like their
dogs. In fact, what most unites any family (pets included) is the contents
of their guts. The younger a baby, and the more isolated from a source of
new bacteria, the lower-and more distinctive-its internal diversity.
Because of migration, any archipelago, of entrails or anything else, be­
gins to resemble the nearest mainland. In a large town many people have
the same bowel contents, while isolated villages diverge. Cities far
258 D A R W I N ' S G H O S T

apart-great continents of bacteria-have a flora of their own (as soon

becomes obvious to those who travel to them). The gut of a Suffolk vil­
lager at once links him to the intestines of London, while those of a
farmer in Bangladesh show his affinity to the entrails of the great city of
Dacca.
The eruptions of travelers and babies, messy as they are, reflect an in­
ternal struggle for existence, the colonization of a new territory from a
larger expanse nearby. What arrives is a matter of chance, dictated by
how far from the source the new island might be and how good each mi­
grant is at making the journey. Some bacteria triumph by changing to
deal with new conditions, while others fail in the battle and disappear. In
time the digestive system settles down as its ecology changes. The process
has a lesson for evolution, because the history of guts is not much differ­
ent from that of islands, of mountains or of great lakes.
The farther an island is from the mainland, the harder it becomes for
new animals and plants to arrive. As a result, each contains just a sample
of what is present on the nearest continent. The most isolated patches of
land are the most diminished of all .
To an agent of disease, each host is an island. If the archipelago of vic­
tims is small, the infection may disappear. Iceland had too few people to
support a permanent population of measles viruses until after the Second
World War. If a place is too remote, a disease may never arrive-which
is why Britain has no rabies. Even in Europe, the virus must pass between
islands-hosts-to stay alive and to reach a new victim before the last
sinks dead beneath the metaphorical waves. Where foxes are rare, each is
so isolated that the disease cannot gain a hold. Rabies can be controlled
with increased isolation. To shoot-or to immunize-foxes moves the
survivors farther from a source of infection. For a fox, a city is a conti­
nent, and the animals are so common that neither vaccines nor bullets
can save them from invasion.

On the morning of 27 August 1 883 a gigantic explosion shook the

world. The bang was heard from Sri Lanka to central Australia. Its blast
traveled four times around the globe, and the tidal wave reached Dover.
Krakatau had exploded, with the force of ten thousand Hiroshima
bombs.
 Geographical Distribution-continued 259

The volcano sits in the Sunda Strait, between Sumatra and Java, o n a
line of strain in the Earth's surface. As a fulcrum of a gigantic geological
lever, the strait is always in turmoil. The Javanese Book ofKings records a
huge eruption in A. D . 4 1 6. There have no doubt been many more. Be­
fore the bang, Krakatau had been covered with forest. The Endeavour, on
Captain Cook's first voyage, visited in 1 77 1 : "We saw that there were
many houses and much Cultivation upon Cracatoa, so that probably a
ship might meet with refreshments who chose to touch here." After
Cook's death in Hawaii, his comrades revisited the island and the expe­
dition artist made a sketch of its rich landscape.
The eruption put paid to all that. The pressure gauge of the Jakarta
gasometer is the record of its story. On the day of the explosion it rose
and sank as the blast passed over. Then, the sea fell back and the sky dark­
ened as ash rained down. Its tsunami drowned thirty thousand and the
stream of red-hot fragments that boiled across the sea incinerated many
more. Dust was carried around the world and led to spectacular sunsets.
Fire engines were called out in New York State to fight what seemed a
giant blaze in the distance. Tennyson himself asked, "Had the fierce ashes
of some fiery peak / Been hurled so high they ranged about the globe? /
For day by day, thro' many a blood-red eve. / . . . / The wrathful sunset
glared."
A week after the eruption two thirds of the island was gone. The rest
was covered by red-hot pumice. The first visitor wrote that "in spite of
all my searching I could find no sign of plant or animal life upon the
land, except a very solitary small spider; this strange pioneer of the revival
was in the process of spinning its web." The island was almost sterile.
Life did not take long to return. By the end of the century grasses and
cane to the height of a man covered Krakatau. A survey a few years later
in what had become a dense fig forest revealed more than eight hundred
different kinds of animal. Nowadays, to a casual glance, existence on
Krakatau seems the same as that on the mainland. It is not. Although its
slopes are covered with trees filled with bats, eagles and woodpeckers,
pythons and monitor lizards, they lack the monkeys, cats and frogs abun­
dant on Java itself, thirty miles away. Otters are its only land mammals.
Evidence of Krakatau's tenants before the cataclysm comes from a
small collection of snails and plants from 1 867 and from the sketch made
by the Cook expedition. Although the island now has nineteen kinds of
260 D A R W I N ' S G H O S T

snail, they include none of the five collected before the eruption. Of the
half-dozen plants, two never made it back. The Cook picture shows four
recognizable plants (a grass, a fern and two trees), just three of which
have returned.
The fliers and swimmers got there first. The earliest plants were fol­
lowed by worms that floated across in rotten tree trunks, and by moni­
tor lizards (themselves often seen out at sea). Grass seeds arrived by air,
together with a surfeit of orchids and ferns. Then came birds, their guts
loaded with the seeds of figs and other trees, and bats with other seeds
stuck to their fur. In time, a new community emerged.
The history of Krakatau is an experiment in evolution. It differs from
other isolated islands only in that its birth was seen by man. Its deformed
mix of plants and animals makes sense because its history is known. For
most islands, the evidence of the past lies only in the present, in the fact
that their communities have a different shape from those elsewhere. Ge­
ography, not St. Patrick, is to blame for the lack of snakes in Ireland.
Frogs are absent from all oceanic islands-Hawaii, Madeira, the Galapa­
gos-.and the commonest mammals (apart from those introduced by
man) are bats. In the same way, such places have grasses in abundance,
but not many trees.
If frogs or snakes appeared on Earth through the agency of a benefi­
cent architect, why should they not be brought into being on Hawaii or
Ireland as much as on the mainland? They are absent not because the
Creator has a devious mind, but because they cannot cross the sea. How­
ever, islands are also great manufactories of life. If New Guinea is taken
as the world's largest, islands make up a thirtieth of the land surface.
They contain, however, about one in six of all known species. Such places
are proof ofwhat Darwinism can do and those most isolated give the best
evidence of all.

On the Inhabitants ofOceanic Islands. Some islands are too remote for any
but the most determined migrants. The farther from a source, the slower
the rate of colonization, until, at last, a piece of land very distant from a
continent contains far fewer forms than does an equivalent area of main­
land. Although in oceanic islands the number of kinds of inhabitants is
scanty, the proportion of endemics (those found nowhere else in the
world) is often extremely large. If we compare the number of the en­
demic land shells in Madeira, or of the endemic birds in the Galapagos
 Geographical Distribution-continued 26 1

archipelago, with the number found on any continent, and then com­
pare the area of the islands with that of the continent, we shall see that
this is true. The inhabitants of the most distant islands have been isolated
for so long that descent with modification has done its work undis­
turbed . .As a result, they lay claim to forms of existence-related to but
distinct from those of the nearest mainland-found nowhere else.
For less mobile creatures, evolution can proceed apace. Madeira is in­
habited by a wonderful number of peculiar land shells, whereas not one
species of seashell is confined to its shores. Though we do not know how
seashells are dispersed, yet we can see that their eggs or larvae, perhaps at­
tached to seaweed or floating timber, or to the feet of wading birds,
might be transported far more easily than land shells, across three or four
hundred miles of open sea.
Krakatau is young and close to its source of immigrants. Although its
inhabitants are a biased sample of those on Java, none is unique. More
distant islands, in contrast, are full of change. Evolution did not spring
from Darwin's brow as soon as he saw the Galapagos finches (the best he
could find to say about them was that: "their general resemblance in
character and the circumstance of their indiscriminately associating in
large flocks, rendered it almost impossible to study the habits of partic­
ular species . . . They appeared to subsist on seeds") . Even so, the birds
helped form his later ideas, and islands are still among the best evidence
of how evolution works.
They come in many shapes and sizes. Most are the remains of
drowned-or raised-continents. At the height of the last ice age so
much water was trapped in the glaciers that there was a fifth more dry
land than today. The Hebrides, Martha's Vineyard and many more
formed as the sea reclaimed its own. What once were hills became is­
lands. Cyprus, Crete and Corsica have the same history. .As sea levels
changed, the barrier at the Straits of Gibraltar allowed the Mediterranean
to dry (and Mrican apes to reach Gibraltar). Then, as the Atlantic rose,
the straits turned into a marine waterfall that filled it within a century.
Yet other islands rose into existence as the ice was shed. The sea bottom
around Finland-a country that is a dilute solution of land in water­
still rises by half an inch a year and forms archipelagoes as it goes. Places
such as these are mere fragments of a broken continent and do not dif­
fer much from their parent nearby.
When The Origin was published, Alfred Russel Wallace was in In-
262 D A R W I N ' S G H O S T

donesia (from whence he had sent the famous letter that caused Darwin
to rush into print). He noticed a strange pattern of distribution among
its inhabitants. A line-later called Wallace's Line-drawn across the
archipelago, with the Philippines, Borneo, Java and Bali to its north and
west, and Lombok, Sulawesi and New Guinea to the south and east, sep­
arated two biological worlds. One side had ties to Asia, with its fruit
thrushes, weaverbirds and tigers, and the other to Australia, with cocka­
toos, honey eaters and kangaroos. In some places a mere fifteen miles of
sea separated the two provinces.
Wallace almost got it right: he wrote of the archipelago, "I believe the
western part to be a separated portion of continental Asia, the eastern the
fragmentary prolongation of a former Pacific Continent." He appealed
to the drowned land of Lemuria, now sunk in the Pacific, to explain the
links with Madagascar. In fact, the history of Gondwana is to blame. The
country east of his Line was part of that continent in its last days, while
that to the west was part of Asia. The ancient division remains today. No
lost continent is required, just one that moves around. As Wallace no­
ticed, the narrow straits that made his Line were deep and passages of
shallow water were no barrier even to animals that could not swim. That
is a relic of the end of the last glaciation. The shallows are mere valleys
filled with water, while the deeps mark the edges of ancient continents.
Some islands have no ties of any kind. They are isolated by vast oceans
and emerge, as sterile as a newborn baby, from the sea as lava boils from
below. Hawaii is above a hot spot, a spring of liquid rock that wells up
from the Earth's core and waves like a plume of smoke as it makes its way
toward the surface. As the sea floor grinds over it, volcanoes burst
through. Each new expanse is borne to the west as the plate moves on
and, as it moves, is worn away. Hawaii is the highest mountain on Earth,
almost a mile higher than Everest when measured from its base on the
ocean floor to the summit of Mauna Loa, fourteen thousand feet above
the sea. It is one of fifty great volcanoes. The most distant, the Emperor
Sea Mounts, stretch almost to Siberia and are eroded stumps deep below
the surface. Hawaii itself is less than a million years old, the farthest sea
mount more than seventy million. To the east of Hawaii, as yet unre­
vealed, another member of the group-to be named Loihi-is under
construction. It will break the surface in thirty thousand years. As it
moves on, the chain is a conveyor belt of evolution.
 Geographical Distribution-continued 263

Distant islands, wherever they might be, have much in common. Each
has a balance of species distinct from that of the mainland, and tends to
resemble places further from the Equator than does the nearest conti­
nental point of the same latitude. Thus, the Canaries have a rather
Mediterranean feel (although they are off the coast of Africa) while the
Galapagos are deserts compared to the equatorial forests found at the
same latitude in South America. Island plants become more interested in
sex; with many more species having separate males and females than on
the mainland. Their animals, oddly enough, tend to lean the other way,
with many sexual species abandoning their males. Not much of this is
understood, but remote islands all share one property that gives them a
special place in the case for evolution.
The most striking and important fact for us in regard to the inhabi­
tants of islands is their affinity to those of the nearest mainland, without
being actually the same species. On the Galapagos archipelago, situated
under the equator, between five hundred and six hundred miles from the
shores of South America, almost every product of the land and water
bears the unmistakable stamp of the American continent. The close
affinity of most of the birds to American species in every character, in
their habits, gestures and tones of voice, is manifest. The naturalist, look­
ing at the inhabitants of these volcanic islands in the Pacific, distant sev­
eral hundred miles from the continent, easily feels that he is standing on
American land. It is obvious that the Galapagos Islands would be likely
to receive colonists from America and that such colonists would be liable
to modifications; the principle of inheritance still betraying their origi­
nal birthplace.
All islanders resemble the inhabitants of the nearest mainland and not
those of comparable islands far away. The Cape Verdes have animals and
plants resembling those of Africa and not the Galapagos. They show that
isolation, not insularity, is what causes change. Distant islands-unlike,
say, the zinc-coated archipelago of electricity pylons, whose surrounding
plants each evolves an identical response to the same poison-are differ­
ent because each received their own mix of immigrants from their main­
land. Necessity-evolution-has molded what chance provided.
All new islands-marine volcanoes, mountaintops left empty by the
ice, or lakes that open as the Earth's crust moves-are colonized by a
sample of the mainland population. If the sample is small (as it often is)
264 D A R W I N ' S G H O S T

then a roll of the genetic dice itself may cause the new arrivals to differ
from those left behind. That means immediate change through the acci­
dents of travel. What is more, the first generations in a new home will be
few in number. As a result, and at random, more genes will be lost, be­
cause their bearers fail to reproduce.
Any population that goes through a bottleneck has no choice but to
evolve. Cats tell the story. Cities are filled with feline diversity-tabby,
black and white, orange, long-haired and all the rest. Islands are differ­
ent. Every cat on the remote French Dependency of Kerguelen, in the
southern Indian Ocean, is black (some mitigated by splashes of white) ,
no doubt because the few arrivals carried only those genes. On the
equally French territory of St. Pierre et Miquelon, fifteen miles off New­
foundland, the cats are different again. Their genes are not like those of
the nearby mainland, but a reduced sample of those of Bordeaux, two
thousand miles away.
To stay alive in a distant land is not easy. As those who tried to intro­
duce birds to New Zealand found, a hundred or more of each form were
needed to survive. The much-touted idea that a whole community could
descend from a single seed or pregnant female is in most cases not real­
istic. Any traveler able to reach a safe haven will, given time, arrive in
some numbers. There are a few exceptions, like the solitary (and some­
what dazed) macaque found afloat on a log after the Krakatau explosion,
but most colonists need, like the Pilgrim Fathers, a reasonable group to
have a chance of success.
Genetic variation in animals that live both on a continent and on an
island-mammals, birds or snails-shows most insular populations to
be less variable than their progenitor. Even so, the reduction is modest.
Darwin's finches are themselves so diverse that they must descend from
a solid nucleus of birds that made the journey from South America. The
idea of whole species arising in such places almost by accident-a sort of
supply-side evolution-has been much explored by the mathematicians
who cluster around the margins of Darwinism. Most of the evidence is
against it. Necessity-natural selection-forms their inhabitants, as it
does those who stay at home. An empty and remote volcano is full of op­
portunities. Animals and plants soon make use of them, to give new
kinds that might find it hard to live in the competitive world of their an­
cestors.
 Geographical Distribution-continued 265

Accident does play a part in who ends doing what. On different

Caribbean islands, the lizards of the treetops look much the same but are
not in fact related. DNA shows that one island's climber may belong to
quite a different family of lizards from the climbing lizard on another.
Each has (like the partridges and hares that go white in winter) evolved
the same independent solution to an ecological challenge, starting from
a different place. No doubt the first to arrive in a new place filled a gap
and denied it to later arrivals.
Most mainland trees (apart from those with buoyant seeds, such as the
coconut) have no hope of an oceanic journey. An herbaceous plant,
though it would have no chance of successfully competing in stature
with a fully developed tree, when established on an island and having to
compete with herbaceous plants alone, might readily gain an advantage
by growing taller and taller and overtopping the other plants. If so, nat­
ural selection would often tend to add to its stature and thus convert it
first into bushes and ultimately into trees. As they reach to the sky, the
most improbable plants become large and woody, a habit denied to them
back home. Daisies are fond of the pastime, as are many more. Tree let­
tuces grow on Madeira and the Cape Verde islands; tree cabbages, tree
asters and tree sneezeweeds on St. Helena; tree sunflowers and tree flea­
banes on the Galapagos; and tree celeries are scattered over the Azores,
Canaries and Cape Verde islands. The mountaintops of East Mrica­
Mount Kenya and Mount Elgon-have their unique tree lobelias. All
rise above their competitors and shade out the plants below.
Because many island insects are flighdess, and others-such as butter­
flies or bees-do not often cross the sea, the plants have to adapt to a new
sexual landscape. Hawaii has a mere half-dozen kinds of moth, two but­
terflies and no bumblebees. Most island flowers are not the showy dis­
plays of the mainland, but tend to be small and inconspicuous. The
commonest colors are white, green and yellow, in lands as far apart as
New Zealand and St. Helena. As windblown seeds-like the pollinators
themselves-might blow away, many plants reduce the wings or para­
chutes that spread the seeds of their land-bound ancestors.
Diversity explodes in distant places, on the Hawaiian archipelago
most of all. Almost a thousand flowering plants are found only on the
chain. Eight hundred distinct kinds of fruit flies, a third of the world
total, live there, and the beedes have burst into even mor'� variety.
266 D A R W I N ' S G H O S T

The oldest survivor, Kauai, is a mere five million years old. The string
of drowned and more ancient lands to the west shows that much of the
chain's evolution happened in lands now beneath the sea. Even so, the
Big Island of Hawaii, the youngest member of the group, has eighty
plants of its own, each of which must have evolved in the million years
since it rose above the waves. One group, the lobelioids, has a great range
of forms. More than a hundred kinds are found on the archipelago, a
ninth of the native flora of the Hawaiian chain. None of them is present
anywhere else on Earth, and most are restricted to single islands in the
group. Their DNA shows them all to be close kin, but in shape and size
they vary from low rosettes, to vines and to trees twenty feet high. A
small fleshy cliff dweller whose seeds blow in the wind has as its nearest
relative a large tree with seeds dispersed by the birds that eat its fruit.
Their diversity is a monument to evolution and to other inhabitants,
now lost. Many of Hawaii's plants (unlike those on most islands) have
red, purple or blue flowers. Those garish blooms evolved to attract polli­
nators-the archipelago's own birds, the honeycreepers. Twenty-three
different kinds of honeycreeper have been seen by Europeans, fifteen of
which survive. Most have long, thin curved beaks. They live on nectar
and specialize in their own sweet flower. Other unique birds of the chain
had a less amicable relationship with the plants. Some lobelioid leaves
and stems are covered to chest height with thick spines to protect them
against their enemies. Now, the grazers have gone, but no t long ago
Hawaii had its own giant herbivorous ducks and geese. The thorns were
needed on the lower leaves alone, as the great birds were flightless. They
were killed off, with many of the honeycreepers, by the first human set­
tlers a thousand years ago.

The world has many other islands; of people, of land and of water. A
third of all species of fish live in fresh water, although lakes and streams
cover only one part in a hundred of the Earth. Even so, most lakes and
ponds, isolated as they might seem, are in effect members of the same
shared continent; only the true islands of water (as isolated as those of
land) have been hotbeds of evolution, and for the same reason.
Most freshwater fish live risky lives, because a sudden drought may
put paid to their home. Streams and ponds are temporary places, liable
 Geographical Distribution-continued 267

to dry up and to re-form as valleys are worn away and as rainfall comes
and goes. Their inhabitants are, in most places, quite similar, living on
watery islets separated by dry land though they might, because such
places are less separate than they seem. Floods can move fish from stream
to stream, and changes in the level of the land cause rivers to flow into
one another. Most freshwater plants and animals are good at travel; as
spores, attached to the legs of birds, or as live adults (the fish and frogs
dropped by whirlwinds included). As a heron moves from one pond to
the next, any egg stuck to its leg is guaranteed to arrive in a favorable
place. Nature, like a careful gardener, has taken her seeds from a bed of
a particular nature and has dropped them in another equally fitted for
them.
The zebra mussel is a modest beast-a freshwater snail the size of a
thumbnail. Its home is in the basins of the Black, Caspian and Aral seas.
There, mussels are everywhere. Their larvae float, or stick to flotsam or
the legs of birds. The Volga Boatmen dragged plenty of the animals with
them as they heaved their barges upstream. As trade increased, the mus­
sels moved west. They reached the Danube by 1 800 and the Thames
soon after. In 1 988, a colony was seen in Lake Saint Clair, near Detroit.
It had arrived as a stowaway, in water carried by a ship as ballast (millions
of gallons of which are moved from port to port each year). The zebra
mussel was the most successful of the four hundred kinds of animal
found in the tanks. In the 1 970s hundreds of Russian ships came to the
Great Lakes to collect the grain that the Soviet Union could not grow for
itsel£ Some carried mussels. Within ten years the animal had spread over
the eastern United States. It took almost no time for the animals to fill
the Mississippi. On the way they have driven out their native equiva­
lents, the purple wartyback, the shiny pigtoe and the monkeyface in­
cluded.
However, a few distant lakes have long been safe from such immi­
grants. They have evolved, like Hawaii, an identity of their own. They
are ancient and remote, filled with unique animals and plants, veritable
volcanoes of water separated by great oceans of land. Their inhabitants
have been isolated for millions of years. Like those of other islands, they
have exploded into a diversity of form, and, like them, they are now
threatened by the greatest traveler of all.
Lake Tanganyika was first seen by Europeans in 1 858, when the
268 D A R W I N ' S G H O S T

British explorers Burton and Speke reached its eastern shore in their
quest for the source of the Nile. It lies, with its fellows Lake Victoria and
Lake Malawi, in the Great Rift Valley. Its arid shores are home to some
of the most spectacular animals in the world. They live out their strug­
gle for existence for a global audience of television viewers.
The cameras would find much more diversity beneath the water. Each
of the great lakes has thousands of species found nowhere else. They are
the finest evolutionary microcosm on earth. As the longest and, at almost
five thousand feet, the second deepest lake in the world (after Baikal, it­
self the home to a strange fauna of its own), Lake Tanganyika contains a
sixth of the globe's fresh water.
It was formed as rivers flowed into the basin left by two continental
plates as they shifted. Tanganyika is among the oldest oflakes, first filled
twelve million years ago. Below about three hundred feet, the water is
dead. Its sides are steep and most of the inhabitants are confined to a nar­
row layer around the shore. They include fifteen hundred unique kinds
of animals and plants-fish, sponges, crabs, snails and even a freshwater
jellyfish. Many look rather like sea creatures, and the idea that the lake
was once joined to the ocean led to the first scientific expedition. Its evo­
lution, however, happened within its own confines.
The lake has almost as many kinds of freshwater fish as does the whole
of northern Europe. Catfish, eels and perch have evolved an assortment
of shapes, sizes and behaviors. Although just a small part of the shore has
been explored, two hundred members of one group, the cichlids, are
known. All except five are found in Lake Tanganyika alone. Lake
Malawi, which shares much the same history, has a thousand different
kinds.
Within the lakes are the fishy equivalents of the elephants, lions and
hyenas that roam the nearby plains. Cichlids vary in size from an inch to
two feet or so. Some scrape algae off the rocks, others browse on snails or
on the animals of the bottom. Some, almost like whales, filter the water
for tiny particles of food. Members of another group, with no land-based
equivalent, sneak up on fish from behind and tear scales from their bod­
ies. They have two forms, one a specialist on the left flank, the other on
the right. All cichlids care-like lions or elephants-for their young. In
some, females hold them in their mouths (and are open to the attack of
fish cuckoos, which dump their own eggs in another's maw). Others use
 Geographical Distribution-continued 269

abandoned snail shells as homes for their brood. Their brains as well as
their bodies have changed, as the fish eaters are better endowed with gray
matter than grazers on plants or snails. All this is evidence for what nat­
ural selection can achieve when left to do its work.
Most of the fish inhabit rocky outcrops or patches of gravel close to
the shore. They are sedentary beasts, and almost never venture across the
sand separating each patch. They live on islets within the large lake isle
itsel£ Even the small Lake Nagugago, separated from the main Lake Vic­
toria by a spit of sand within the past fifteen hundred years, has several
cichlids of its own. As the waters rise and fall over a few years, many tiny
lakes have appeared and disappeared around the shores of their great par­
ents. They were, like the drowned stumps of the Hawaiian chain, part of
the engine that drives diversity.
Lake Tanganyika itself almost dried up twelve thousand years ago and
split into three smaller bodies of water. This division led to an outbreak
of evolution. The ancient breach in its waters is reflected in today's fish,
who are separated into distinct lineages that arose within the reduced
lake and have diverged further in their own tiny homelands. Each lineage
has its specialists-a grazer, a grinder or a filterer-adapted to the op­
portunities on offer. Lake Victoria dried up altogether at the same time,
and its three hundred cichlids have evolved since that disaster.

Archipelagoes are the workshops of evolution, their products a micro­

cosm of its every stage. What they make is fragile. Islanders do well in
their own limited marketplace, but once exposed to the outside world are
soon driven to extinction. Because they have adapted free from outsiders
most are, like the flightless geese of Hawaii, doomed. Their fate comes
from a simple law of economics: the need to compete. The products of
islands, like those of other small factories, find it hard to survive in the
global economy. More than half the mammals lost during historical
times lived in such places, as did nine tenths of the extinct birds.
Integration destroys diversity, as solitude promotes it. Islands are dan­
gerous places, because they are small and are open to accident. Any loss
that might elsewhere be replaced is final. On a patch of land, things are
liable to go wrong more often than on a larger tract, and there is nowhere
to hide from disaster. Its inhabitants are at constant risk from rude out-
270 D A R W I N ' S G H O S T

siders. The rudest of all is man. He filled the farthest points of the hab­
itable Earth a mere thousand years ago, when boats reached New
Zealand. As soon as he arrived, he had no compunction in exploiting
what he found.
William Strachey wrote an account of his shipwreck on Bermuda in
1 609. His book influenced Shakespeare, whose The Tempest, written two
years later, mentions the archipelago as the "still-vexed Bermoothes."
Strachey noticed a bird that "for their cry and whooting, wee called the
Sea Owle . . . Our men found a prettie way to take them, which was by
standing on the Rockes or Sands by the Sea side, and hollowing, laugh­
ing, and making the strangest out-cry that possibly they could: with the
noyse whereof the Birds would come flocking to that place, and settle
upon the very armes and head of him that cryed." The Sea Owle is no
more.
The first travelers noted the tameness of the birds of the Galapagos; it
is not complete, for the finches are wary of familiar enemies, and will
perch on a person if a hawk flies over. Their evolved confidence in the
face of the unknown was a mistake, and they, with many others, have
paid the price. The number of seabirds on Ua Huku in the Marquesas
dropped from twenty-two to four after man arrived two thousand years
ago. The first New Zealanders destroyed eight kinds of moa, together
with parrots, owls and many others. In Polynesia, two thousand unique
terrestrial rails were lost, and the last Wake Island rail, first described in
1 903 by Lord Rothschild, was devoured by Japanese soldiers in 1 945: a
career from discovery to extinction of four short decades.
Ancient lakes, too, have paid for their sheltered lives. Ten million peo­
ple live around Lake Tanganyika, and thousands of tons of fish are taken
each year. The Nile perch, introduced as a food fish into several of the rift
lakes, has driven many of the native cichlids to extinction. The perch is
now exported to Europe and will no doubt be moved to wherever it can
grow, so that many cichlids yet unknown will soon be gone.
At the time of the Beagle most islands were as isolated as they had been
since they began. Now, everything has changed. The world's land
masses-like its economies, with five trillion dollars' worth of goods
traveling by sea each year-have become connected into a single body.
So have their animals and plants. Isolation is now a rare commodity, and
the most distant scrap of land has joined the continents. In the United
States, one plant species in ten has come from elsewhere, in Britain al-
 Geographical Distribution-continued 271

most half-and o n Ascension Island in the Atlantic more than 8 0 per­

cent of plants are immigrants. The destruction of the locals and the ho­
mogenization of the world's ecology-a biological Coca-Colonization to
go with its cultural equivalent-is not yet complete, but soon will be.
The larger any piece of land, the more forms of life it contains, but the
number does not rise in exact relationship to its area. In that fact is a dis­
mal message for the future. If all the world's islands were to be joined to­
gether into a supercontinent-a new Pangaea-its magnificent expanse
would, given the relationship between the size of islands and the num­
ber of species, contain half the kinds of animal that now exist.
As travel brings the earth together, many of the products of evolution
will disappear. Darwin, were he to circle the globe today, would find it a
less remarkable place than it was when the Beagle made its voyage. That
grand simplification is itself evidence of the power of his machine.

Summary of last and present Chapters. In these chapters I have endeavoured

to show, that if we make due allowance for our ignorance of the full ef­
fects of all the changes of climate and of the level of the land, which have
certainly occurred within the recent period, and of other similar changes
which may have occurred within the same period; if we remember how
profoundly ignorant we are with respect to the many and curious means
of occasional transport,-a subject which has hardly ever been properly
experimented on; if we bear in mind how often a species may have
ranged continuously over a wide area, and then have become extinct in
the intermediate tracts, I think the difficulties in believing that all the in­
dividuals of the same species, wherever located, have descended from
the same parents, are not insuperable. And we are led to this conclusion,
which has been arrived at by many naturalists under the designation of
single centres of creation, by some general considerations, more espe­
cially from the importance of barriers and from the analogical distribu­
tion of subgenera, genera, and families.
With respect to the distinct species of the same genus, which on my
theory must have spread from one parent-source; if we make the same al­
lowances as before for our ignorance, and remember that some forms of
life change most slowly, enormous periods of time being thus granted for
their migration, I do not think that the difficulties are insuperable;
though they often are in this case, and in that of the individuals of the
same species, extremely grave.
As exemplifying the effects of climatal changes on distribution, I have
272 D A R W I N ' S G H O S T

attempted to show how important has been the influence of the modern
Glacial period, which I am fully convinced simultaneously affected the
whole world, or at least great meridional belts . As showing how diversi­
fied are the means of occasional transport, I have discussed at some lit­
tle length the means of dispersal of fresh-water productions .
IT the difficulties be not insuperable in admitting that in the long
course of time the individuals of the same species, and likewise of allied
species, have proceeded from some one source; then I think all the grand
leading facts of geographical distribution are explicable on the theory of
migration (generally of the more dominant forms of life), together with
subsequent modification and the multiplication of new forms . We can
thus understand the high importance of barriers, whether of land or
water, which separate our several zoological and botanical provinces. We
can thus understand the localisation of subgenera, genera, and families;
and how it is that under different latitudes, for instance in South Amer­
ica, the inhabitants of the plains and mountains, of the forests, marshes,
and deserts, are in so mysterious a manner linked together by affinity,
and are likewise linked to the extinct beings which formerly inhabited
the same continent. Bearing in mind that the mutual relations of organ­
ism to organism are of the highest importance, we can see why two areas
having nearly the same physical conditions should often be inhabited by
very different forms of life; for according to the length of time which has
elapsed since new inhabitants entered one region; according to the na­
ture of the communication which allowed certain forms and not others to
enter, either in greater or lesser numbers; according or not, as those
which entered happened to come in more or less direct competition with
each other and with the aborigines; and according as the immigrants
were capable of varying more or less rapidly, there would ensue in dif­
ferent regions, independently of their physical conditions, infinitely di­
versified conditions of life,-there would be an almost endless amount of
organic action and reaction,-and we should find, as we do find, some
groups of beings greatly, and some only slightly modified,-some devel­
oped in great force, some existing in scanty numbers-in the different
great geographical provinces of the world.
On these same principles, we can understand, as I have endeavoured
to show, why oceanic islands should have few inhabitants, but of these a
great number should be endemic or peculiar; and why, in relation to the
means of migration, one group of beings, even within the same class,
should have all its species endemic, and another group should have all
its species common to other quarters of the world. We can see why whole
 Geographical Distribution-continued 273

groups o f organisms, a s batrachians and terrestrial mammals, should be

absent from oceanic islands, whilst the most isolated islands possess
their own peculiar species of aerial mammals or bats. We can see why
there should be some relation between the presence of mammals, in a
more or less modified condition, and the depth of the sea between an is­
land and the mainland. We can clearly see why all the inhabitants of an
archipelago, though specifically distinct on the several islets, should be
closely related to each other, and likewise be related, but less closely, to
those of the nearest continent or other source whence immigrants were
probably derived. We can see why in two areas, however distant from
each other, there should be a correlation, in the presence of identical
species, of varieties, of doubtful species, and of distinct but representa­
tive species.
As the late Edward Forbes often insisted, there is a striking paral­
lelism in the laws of life throughout time and space: the laws governing
the succession of forms in past times being nearly the same with those
governing at the present time the differences in different areas. We see
this in many facts. The endurance of each species and group of species
is continuous in time; for the exceptions to the rule are so few, that they
may fairly be attributed to our not having as yet discovered in an inter­
mediate deposit the forms which are therein absent, but which occur
above and below: so in space, it certainly is the general rule that the area
inhabited by a single species, or by a group of species, is continuous; and
the exceptions, which are not rare, may, as I have attempted to show, be
accounted for by migration at some former period under different condi­
tions or by occasional means of transport, and by the species having be­
come extinct in the intermediate tracts. Both in time and space, species
and groups of species have their points of maximum development.
Groups of species, belonging either to a certain period of time, or to a
certain area, are often characterised by trifling characters in common, as
of sculpture or colour. In looking to the long succession of ages, as in now
looking to distant provinces throughout the world, we find that some or­
ganisms differ little, whilst others belonging to a different class, or to a
different order, or even only to a different family of the same order, dif­
fer greatly. In both time and space the lower members of each class gen­
erally change less than the higher; but there are in both cases marked
exceptions to the rule. On my theory these several relations throughout
time and space are intelligible; for whether we look to the forms of life
which have changed during successive ages within the same quarter of
the world, or to those which have changed after having migrated into dis-
274 D A R W I N ' S G H O S T

tant quarters, in both cases the fonns within each class have been con­
nected by the same bond of ordinary generation; and the more nearly any
two fonns are related in blood, the nearer they will generally stand to
each other in time and space; in both cases the laws of variation have
been the same, and modifications have been accumulated by the same
power of natural selection.
 C H A P T E R X I I I

M UTUAL AF FINITIES O F O RGANI C BEIN G S ;

M O RPHOLOGY; EM B RYOLOGY;

RUDIMENTARY O RGANS

CLASSIFICATION, and the hidden order of life - Arbitrary and Natural sys­
tems - Common descent the key to classification - Cladistics and the
rules of arrangement - The comparative anatomy of the genes and the
new tree of life - An identical present may conceal a separate past. MOR­
PHOLOGY, theme and variations in related creatures, and in their several
parts - The switches of growth - Monstrous animals and plants. EM­
BRYOLOGY, laws of, how the embryo reveals a past lost in the adult. RUDI­
MENTARY ORGANS; the cost of unwanted structures, and their origin
explained - Summary

A celebrated (and perhaps apocryphal) Chinese encyclopedia of the

tenth century classifies plants and animals as follows: (a) those that be­
long to the Emperor, (b) embalmed ones, (c) those that are trained, (d)
suckling pigs, (e) mermaids, (f) fabulous ones, (g) stray dogs, (h) those
that are included in this classification, (i) those that tremble as if they
were mad, (j) innumerable ones, (k) those drawn with a very fine camel's­
hair brush, (1) others, (m) those that have just broken a Hower vase, (n)
those that resemble Hies from a distance.
To the author of The Celestial Emporium of Benevolent Knowledge-­
and no doubt to his readers-that catalog made sense. It is hard to com­
prehend today. There may be a certain affi nity among beasts that break
Hower vases or tremble as if mad, but to the modern eye they form a less
natural group than do, say, suckling pigs, stray dogs or even those that
from a distance resemble Hies. The philosopher Michel Foucault re-
276 D A R W I N • S G H 0 S T

marked of the list that: "In the wonderment of this taxonomy, the thing
that is demonstrated in the exotic charm of another system of thought is
the limitations of our own." Biology has begun to show how limited have
been our own ideas about the great emporium of existence.
Man is a classifying animal. His world is so full of objects that he must
reduce their number by arranging them in groups. They can be sorted in
many ways-red or blue, large or small, safe or dangerous, near or far.
The brain can be seen at work as it puts objects into pigeonholes. As peo­
ple think of tools, or names, or vegetables, they activate specific areas of
gray matter. Drill, mallet and sandpaper all involve the same few cells;
Tom, Dick and Harry another set. Those who have difficulty with the
names of tools have the same problem with those of animals because the
system for sorting the two is in the same place.
The notion that a spirit level and a file fall into the same category must
be learned, because a group called "tools" makes sense only in context.
Apart from utility, such objects have little in common. Why should the
brain not separate things belonging to the Emperor from those with
other owners? In China, after all, to do so might be a matter of life and
death. Even so, to sort by ownership does not say much else about any
object, be it palace, Pekingese or porcelain teapot. They are united by a
single property that does not overlap with others.
A thousand years after the oriental list, life can still be organized in
many ways. All but one are useful but biased, because they are artificial.
The Catholic church once classified, for culinary reasons, the capybara as
a fish. That makes sense for hungry travelers on a Friday, but is not much
use to students of South American mammals. To define animals as edi­
ble, or endangered, or cute is a great help in the context of kitchens, zoos
and pet shops, but members of such groups have little else in common.
One arrangement, and only one, reveals the order hidden in nature.
This classification is evidently not arbitrary like the grouping of stars in
constellations. Something more is included than mere resemblance. It is
based on a natural structure, on common descent from a distant past. As
a result, it reveals the truth of evolution in a way denied to all others.
The contrast is laid bare in the natural history museums of Paris. The
Zoology Museum and the Museum of Comparative Anatomy trace their
descent from the Jardin des Plantes, founded by Louis XIII. They were,
for much of their history, repositories of Nature, present and past; or-
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 277

dered, rational and deductive, their contents arranged by the rules of bi­
ological classification. Each exuded the same grave nineteenth-century
air as does The Origin (although the image of Darwin was hard to find
in either).
The Museum of Zoology sank into decline and was closed in 1 965 as
its specimens crumbled in their cases. Then, in 1 994, its Grand Gallery
of Evolution was reborn as part of the French reinvention of their capi­
tal. The museum's public face has changed. The stuffed animals that once
decayed in organized grandeur are now arranged as a series of spectacles:
the Mrican savanna, the ecological crisis, or the plants and animals use­
ful to man. Spotlights abound, with animated exhibits popular with chil­
dren.
The new museum's logic, splendid as its interior may be, is that of a
Chinese encyclopedia. The members of each display are united by a sin­
gle property; each lives on the Mrican plains, has been domesticated, or
is threatened by extinction. To know that does not say much more about
what they are. In one exhibit, an elephant and a locust are next to each
other, in another a camel and a goose.
The nearby Gallery of Comparative Anatomy and Paleontology is un­
renewed. Around its walls are hundreds of pickled guts and through its
dusty halls march a myriad of bones, in progress from fish to man (not a
Frenchman, but an Italian). Marbles and bronzes-An Orang-Utan
Strangling a Native of Borneo, Man Triumphant over Two Bear�press
home the message of progress (although the unfortunate victim of the
orang has for some reason a snail crawling up his leg). The edifice, with
its cast-iron beams and Beaux-Arts style, has much charm but is for most
of the time almost empty. The distractions of its modernized neighbor
are what attract the crowds.
Forlorn as it is, the Anatomy Museum has stayed true to the spirit of
evolution. It sets out its animals by affinity. The Linnaean system has a
hierarchy of order-the American wolf, for instance, belongs in the
kingdom Animalia, the subkingdom Metazoa, the phylum Chordata,
the subphylum Vertebrata, the superclass Tetrapoda, the class Mam­
malia, the subclass Theria, the infraclass Eutheria, the cohon Ferungu­
lata, the superorder Ferae, the order Carnivora, the suborder Fissipeda,
the superfamily Canoidea, the family Canidae, the subfamily Caninae,
the genus Canis, the species lupus and the subspecies occidentalis (perhaps
278 D A R W I N ' S G H O S T

fortunately, the Caninae-dogs, dingos, foxes and wolves-are not di­

vided, as are some groups, into separate tribes, and the genus Canir­
dogs, dingos and wolves-has not been split into subgenera).
Linnaeus invented his system as a bookkeeper might, as a scheme for
filing similar objects. God created, but Linnaeus arranged, and how he
did his job was criticized from the earliest days. A reviewer of 1 759 com­
plained, in anticipation of today's museologists, that dogs should not go
with foxes and wolves but with horses, as both are found in the farmyard.
His scheme has lasted well (although it has been hijacked by wits who
have sneaked in names such as Ba humbugi for a snail, and Agra vation
and Agra phobia for a pair of beetles). It is, like the Zoology Museum, to
some degree arbitrary, most of all in its higher ranks. The eight thousand
or so kinds of birds are divided into a hundred and sixty families, while
sixty thousand species of parasitic wasps have to fit into a single family in
their part of the catalog. Nevertheless, his system contains a great truth:
that the world has a pattern. Linnaeus did better in drawing life's big pic­
ture than did the tribal experts who are so good at recognizing different
species of bird. Once above taxonomy's basic unit, the species, their cat­
egories become bizarre. For one set of people, all birds bar one go to­
gether, but the cassowary fits, in the rural mind, with rats and mice
because it is brown and lacks wings. The Linnaean system, in contrast, is
arranged as a universal plan that hints that, from the first dawn oflife, all
organic beings resemble each other in descending degrees, so that they
can be classed in groups under groups.

As a result, a taxonomy based on one property predicts much of what

else its members will share. All animals that give milk also have hair. In
the same way, all those with hard external skeletons and air tubes (insects
included) have jointed limbs, and no animal both gives milk and has its
skeleton on the outside. Plants show the same harmony, as those with
flowers all have seeds enclosed in a fruit rather than directly exposed. To
put animals and plants in order reveals a long chain of affinity-an un­
seen connection-between different creatures, distant as they might ap­
pear.
Propinquity of descent-the only known cause of similarity of organic
beings-is the bond, hidden as it is by various degrees of modification,
which is partially revealed to us by our classifications. All true classifica­
tion is genealogical. Just as in a human family, with its brothers, sisters,
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 279

cousins and second cousins, it hints at descent from shared ancestors fur­
ther and further in the past. The patterns seen by biologists as they
arrange their world are, unlike those of tenth-century China, evidence
for a system outside the mind of the classifier.
The logic of life can be seen without bothering with evolution at all .
Linnaeus and his Papuan cousin each used descent with modification as
the key to order, without realizing it. To classifiers, of whatever persua­
sion, relatedness is always more important than mere appearance. Every­
one includes within the same species its males, females, adults and
young, however different they appear. The growth stages of certain plants
are so distinct as to be recognizable as members of the same kind only by
experts. The Killarney fern was written off as extinct in the nineteenth
century. It is in fact still widespread-but most of the survivors remain
in another part of the lifecycle as a green mat on moist rocks. As soon as
that was realized, its status as an endangered species was removed.
What unites Killarney ferns is not shape or size, or habit, for the dif­
ferent stages are quite unalike, but common parenthood. Once continu­
ity of descent-the bond that draws together all members of a group--is
established, mere lists of similar objects can be abandoned. If descent is
used to unite individuals of the same species, then their arrangement into
higher classes, however distinct they are, must also involve an element of
shared ancestry. Thus the grand fact in natural history of the subordina­
tion of group under group, which, from its familiarity, does not always
sufficiently strike us, is fully explained.
The cicadas have been around for three hundred million years. Their
image is as a background to the Mediterranean summer, but their capi­
tal has always been in Australia, where the various kinds-the greengro­
cer, the double drummer, the floury baker and many more-are much
collected by children. Cicadas show how group becomes divided into
group as evolution proceeds. That family of a thousand or so insects is
spread over the tropical and subtropical world, in Old World and New,
and has so been for almost three hundred million years. Individual gen­
era (the second-lowest level in the Linnaean system) are more confined:
thus, three quarters of the Australian genera are restricted to their native
continent. Species within a genus are even more localized, with two hun­
dred and forty-six of the two hundred and fifty Australian forms found
only there. The arrangement of cicadas-in a museum case or a child's
280 D A R W I N ' S G H O S T

collection-hence retraces, with no need for conscious effort, their

shared ancestry. Life's logic, its grandest fact, reflects its history.

All classifications need rules that do not depend on the arbitrary interests
of a curator. Biology now has a statistical machine to order the world and
to search for the structure of relatedness over time.
The new science of cladistics means that ancient forms are not seen
merely as missing links on the way to today, to be inserted at the right
point in some great chain of being. Instead, the extinct and the extant are
put into the same system. Cladistics maps out affinities as impartially as
it can. It depends on a single idea: that groups sharing traits not present
in others must descend from a common ancestor. The more characters
that are added to the mix, the further the root is pushed into the past.
Men and whales are related because they have warm blood, four limbs
(reduced to a relic in whales) and hair. Whales, men and fish are joined
because all have backbones (although fish are not hairy and are excluded
from man's more immediate family) . Those three fall into the same
group as the oak as all have a cell nucleus enclosed by a membrane. Bac­
teria are outside even that capacious assembly because, unlike the others,
their DNA floats free inside the cell.
Cladistics, a German invention, has strict rules and a complex vocab­
ulary. It can, if not carefully used, give erratic results and is still filled with
argument about just what should be plugged into its analyses. It has, nev­
ertheless, transformed our view of the world.
The central rule of the new science is that only shared characters, each
derived from the same ancestor, can be used to decide relatedness. As a
result, ancestral characters are of no use in working out who is kin. Many
animals-humans, chimps and iguanas-have five fingers, but others
manage with fewer; thus, horses have a single toe on each foot. A whole
set of shared patterns-hair, milk and so on-show that five fingers came
first and that horses lost them later. In spite of their hands, humans are
not closer to iguanas than they are to horses and the five-fingered hand
is of no use in deciding where they should sit. In contrast, all horses, and
no other mammals, have a single toe. For them, the hoof is a shared de­
rived character. It places them in a group evolved from a common an­
cestor. The less any part of the organization is concerned with special
 Mutual Affin ities,· Morphology; Embryology; Rudimentary Organs 28 1

habits, the more important it becomes for classification . .As a result, at­
tributes only found in a single species-the single tusk of the narwhal, or
the erectile penis of the female hyena-are of little use as they, too, con­
tain no information about descent.
An impartial judge is often called for when trying to sort out a family
problem. To trace the affinities of a set of species it helps to appeal to a
creature so detached from the animals whose relationship is disputed
that it can act as an outgroup, a reference point against which to com­
pare their similarities and differences. Captain Cook, on his first sight of
a kangaroo, saw that it had "a long tail which it carried like a grey hound,
in short I should have taken it for a wild dog, but for its walking or run­
ning in which it jumped like a Hare or dear." The platypus was even
more baffiing. To the contemporary naturalist Thomas Bewick it ap­
peared "to possess a three fold nature, that of a fish, a bird and a
quadruped, and is related to nothing that we have hitherto seen." Al­
though the story was for a time confused by the alleged discovery of fish
with pouches, the antipodeans have now been put in their place. The
kangaroo and the mouse bear live young; the platypus and the birds lay
eggs. By appealing to what is clearly a distant relative-a fish, say, or a
frog-it becomes clear that to lay eggs is the ancestral habit, and that
live-bearing comes later. Kangaroos and mice are hence closer than either
is to a bird or fish, and the platypus, a further set of characters shows,
is-in spite of its birdlike beak (a derived nose)-in a group whose an­
cient members took the first step on the road to kangaroos.
The cladist's art is enlivened by the comparative anatomy of the genes.
DNA provides millions oflinks to the past. A fluorescent probe based on
the genes of one animal can be used to search for its match in another.
Wherever a few thousand bases correspond, the bait is taken up and
makes a lurid blob on the chromosomes. Man and pig, or man and cow,
share more than fifty long sequences. All are evidence of common de­
scent as persuasive as are live young, milk or hair.
The genes of many creatures have now been read from end to end.
They reveal groups within groups beyond the imagination of earlier nat­
uralists. A hundred thousand genes have been located in humans, and a
fifth as many in mice. PIGMAP, the cartography of swine, is up to five
hundred or so, as are the maps of cats and cows. A tiny worm, the first
animal to have had all its DNA letters read, has nineteen thousand and
282 D A R W I N ' S G H O S T

ninety-nine genes altogether, all of them tracked down. Several bacteria

and single-celled parasites have also been deciphered as has the fruit fly.
Most genetical landscapes have a lot in common. Whole sections are
the same in mice and men, and two thousand human genes have exact
homologues in mice. A trudge along the DNA shows more than half of
a certain mouse chromosome to be more or less identical in the arrange­
ment of genes to one of our own; and cows are even more like us. Half
of all plant genes have a mouse equivalent. The worm shares a fifth of its
own heritage with yeast (from which it split a billion years ago) . Unex­
pected parallels emerge from distant places. A gene that in humans
causes an inherited disorder of the nervous system has an exact match in
yeast (which has no nerves at all) .
The scattered information on bones, leaves and DNA can be put into
a common frame. Because cladistics considers so many characters and so
many forms of life at once, the science is a statistical nightmare. Thirty­
five million pedigrees are possible when just eleven different species are
compared. Even so, it has changed our view of life's tree, from its famil­
iar branches to its deepest roots.
There was once disagreement about how the mammals fit together.
Some were an obvious ragbag. The insectivores included hedgehogs,
moles and shrews, all put in the same pigeonhole on the basis of their
joint fondness for insects, the shape of the skull and their absence from
South America. Other categories, such as the carnivores (dogs, cats and
the like) made more sense, while yet other alliances (such as the marriage
of elephants with manatees) seem at first sight odd but hold up when
enough characters are looked at. The deeper roots of the mammalian
family were quite unknown.
Bones and molecules, objectively arranged, reveal the truth. The in­
sectivores as an entity disappear altogether and their members shuffie off
to other places. The hedgehog is on a first and separate branch in the
mammalian family, and the elephant shrew, the golden mole and the
aardvark (all once included as insectivores) join manatees, elephants and
hyraxes in a conjunction of mammals that evolved in Mrica. Other
mammals, too, change their alliances. Not only do whales group with
hippopotami, but dogs and cats join on as more distant members of their
coalition. Humans and apes are, it transpires, quite close to rabbits and
bats.
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 283

Before cladistics, four-legged vertebrates as a whole-lizards, kanga­

roos and mammals-were thought to descend from the ancient lobe­
finned fish, most of which disappeared four hundred million years ago.
Their fins do have a structure at their axis that might have turned into
legs. The discovery of a member of this group, the coelacanth, off the
coast of Mrica in the 1 930s was hailed as a "missing link" between fish
and ourselves. Cladistics showed this to be untrue. The coelacanth is not
on the same branch as vertebrates with four legs. The honor belongs in­
stead to another great group, the lungfish, who flourished at the same
time. Although most modern forms of these fish lack fins altogether (and
even those of their fossils are not at all leg-like), an objective look at skele­
tons puts them closer to ourselves. Now, the molecules have made the
case: the coelacanth is indeed further from today's four-legged animals
than is any lungfish.
Cladistics sometimes simplifies the past. It shows that all land plants,
diverse as they are, are members of a single stock and (unlike the animals,
who did the job dozens of times) moved from sea to land but once, a fact
upon which the fossils are silent. Sometimes it makes things more com­
plicated. The reptiles are, it transpires, a group no more natural than are
those of the Chinese encyclopedia. Reptiles include lizards and croco­
diles, but a whole host of characters show crocodiles to be closer to birds
(not, of course, classified as reptiles) than to their supposed sibs. Croco­
diles are related to dinosaurs; and cladistics shows that birds are di­
nosaurs who grew wings and flew away. Indeed, to ask "Is it a bird or a
dinosaur?" is like asking "Is it an apple or a fruit?"
The birds still have a tie with their newly revealed kin. Crocodiles
(and, no doubt, their dinosaur forebears) lay eggs in nests, can chirp and
determine sex in a peculiar way. If an egg is incubated at low tempera­
tures it develops into a male, at higher temperatures into a female.
Armed with the new insight into their past, an audacious biologist raised
chicken eggs at high temperatures and increased the proportion of fe­
males, which, for an animal in which most males are wasted, is of much
interest to farmers.

The basic rule of evolution remains, however strange its products. On

the principle of the multiplication and gradual divergence in character of
284 D A R W I N ' S G H O S T

the species descended from a common parent, together with their reten­
tion by inheritance of some characters in common, we can understand
the excessively complex and radiating affinities by which all the members
of the same family or higher group are connected. They hint at a shared
past older than any fossil.
To track down the most distant links and draw a new pedigree of life
needs characters that are almost universal; genes at work so deep within
the cell as to resist almost all change and to retain clues about their ear­
liest origin. A hunt for similarity through the dozen and more DNA se­
quences now known in all their detail, and the hundreds for which long
segments have been worked out, reveals the very framework of existence.
About a thousand genes are shared by every organism, however simple or
complicated. Although their common ancestor must have lived more
than a billion years ago, their shared structure can still be glimpsed. It
shows how the grand plan of life has been modified through the course
of evolution.
Such genes show that animals with backbones are close to starfish; and
that worms and snails live on a different branch from insects and round­
worms (both of whom shed their coats as they grow). Their shared root
lies within an ancestor who lived hundreds of millions of years before any
life appeared in the rocks.
Biology's greater divisions are at first sight self-evident. Men and
chimps are close kin, each is less related to worms, and bananas and bac­
teria are quite separate. However, the new taxonomy has transformed the
tree oflife into an exotic plant. Men and chimps are indeed more related
than are men and bananas, but humans, insects and plants are, the DNA
shows, all mere twigs on the same branch. Its trunk has suffered some
radical changes of shape.
Living beings were once divided into five kingdoms of more or less
equal size (animals, plants, fungi, protozoa-such as the familiar
amoeba-and bacteria). Bacteria were out on somewhat of a limb, as
their genes are not contained in a cell nucleus. They seemed otherwise
not much more distinct from other beings than plants were from ani­
mals. Now, a radical new logic has emerged. The DNA reveals that plants
and animals lie close together. Mushrooms deserve a branch of their own,
closer to animals than to plants. Most of the tree belongs not to the lords
of creation, or even to mushrooms, but to the bacteria and their previ-
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 285

ously unrecognized relatives. They put humankind i n its place, near ba­
nanas.
The difference between men and mice, or mice and plants, is tiny
compared to the gulf separating all three from other organisms such as
bacteria. The familiar branches of existence (on one of which men, mice
and plants reside) are in reality unimportant twigs. There is no need to
travel to distant places to see extraordinary beings. Any zoology text
claims that there are more kinds of insect than of anything else; but
squash a fly and thousands of microbes unknown to science will be
squeezed from its gut. Gardens are familiar, biology under control, but a
handful of suburban soil contains a myriad undescribed bacteria, quite
distinct from the plants and animals around them.
Such minute beings are a diverse and remarkable group. Species
thought to be quite similar (such as the agents of botulism and of an­
thrax) are, the genes show, as distinct as men and maize. Escherichia coli,
a bacterium present in billions in Out guts (and the cause of outbreaks of
food poisoning), has about four and a half million DNA units in its ge­
netic instructions. They code for four thousand or so proteins. Although
some are similar to those of other beasts, almost half are quite different
and give no sign of what they do.
One set of genes is found everywhere. It translates the information
coded in the DNA and allows it to make proteins. The job is so essential
that such structures changed little over millions of years. To put them
through the cladistic machine shows how biology has failed to notice
some fundamental splits among its subjects.
Nature's new pedigree has three great domains. One encompasses or­
ganisms whose DNA is contained within a nuclear membrane, together
with, a little further away, a variety of single-celled gut parasites. The bac­
teria and-quite new-the archaea, tiny entities once seen as a mere sub­
division of bacteria but in fact distinct, are each in a class of their own.
Their dominion is divided not into a mere five kingdoms, but into
dozens.
Some are found in unexpected places. The water in Octopus Spring,
near the Old Faithful geyser in Yellowstone National Park, is hot, clear
and alkaline. It contains "pink filaments"-a whole new group. Their ex­
istence needs no oxygen. The nearby Obsidian Pool, whose waters are
filled with iron, hydrogen sulfide and carbon dioxide, has scores of novel
286 D A R W I N ' S G H O S T

groups of archaea. Undiscovered empires live within this single pond.

They are so various that so far they have just been given numbers. The
great naturalist Lionel Walter Rothschild is commemorated in the scien­
tific names of fifty-three birds, eighteen mammals, three fish, two rep­
tiles, a hundred and fifty-three insects, three spiders, a millipede and a
small worm. These new beings will need a host of Rothschilds and whole
departments of classics to give them the titles they deserve.
The world beneath the surface has always seemed an alien place, with
a mythology of its own. Who sang the national anthem to Queen Vic­
toria from the sewers beneath Buckingham Palace? What became of the
plan to pipe their contents to the suburbs for sale in corner shops for
garden use, or of Thelma Ursula Beatrice Eleanor, the first baby born
on a tube-train? Like all cities, London is a vast and curious three­
dimensional world unknown to those who live on its surface. Its buried
inhabitants are beyond the imagination of any tube passenger.
Underground London is a Victorian city, built when life was lived in
the atmosphere-a layer no thicker than the varnish on a school globe­
and when even the oceans seemed dead. Objects sank by weight, with a
layer of cannons suspended beneath drowned ships, themselves below
the bodies of their sailors. The biology of the abyss did not begin until
the Challenger Expedition of 1 872, which revealed a hidden world be­
neath the waves. A century later, our understanding of the deep oceans­
and of the interior of the Earth itself--has undergone a sea change. A
strange and rich universe stretches far beneath our feet.
Around the ocean vents in the midocean ridges is a new world. In
1 977, the submarine Alvin dived two miles below the Pacific. There, lava
pours from the sea bed. "Black smokers," mineral-rich streams of super­
heated water, gush into the sea. Other vents, "white smokers," are sur­
rounded by mats of bacteria. So rapid is the flow that the entire volume
of the oceans circulates through the Earth's crust every five million years.
As it does, it carries heat and chemicals from deep within.
Such places are home to many bizarre beings. They include bright red
tubeworms twelve feet long. The tentacles of these Vestimentifera
(worms in dinner jackets, to translate their Latin name) absorb hydrogen
sulfide from the hot water, and forward it to dense masses of bacteria
within their own bodies. A smoker may need a dinner jacket, but a
mouth and gut is optional.
 Mutual Affin ities; Morphology; Embryology; Rudimentary Organs 287

DNA shows that the worms, like oaks, are mere versions of ourselves.
The real discoveries are among their microscopic helpers. They, with
their fellow microbes of the vents, are quite distinct from worms, oaks
and the whole world of the surface.
One of the vent archaeans lives ten thousand feet down, at a pressure
of two hundred atmospheres. Its genes come in three closed circles and
code for seventeen hundred proteins, most of which have no known
matches elsewhere. It needs only inorganic compounds to stay alive, and
makes methane as waste. Some of its genes are more like our own than
those of bacteria, but this novel beast has many unconventional proper­
ties of its own.
The enzymes of such strange beings are forced to work in hot places.
They might be useful to bleach paper, or to make jeans with a "stone­
washed" look. Other archaeans turn up just as deep, but in the icy cold,
as mats on the bones of drowned whales. They were first discovered by a
u.S. Navy submarine as it searched for a lost missile. The skeleton of a
whale supports more kinds of life than does the richest hot vent and can
take ten years to decay. Its inhabitants make enzymes able to work at low
temperatures, which is useful for companies interested in cold-water de­
tergents. They now fish for new forms in the ocean trenches, with dead
whales attached to buoys as bait. Their catch might include the bugs that
ate the Titanic, a wreck already covered with "rusticles," and, within a
century or so, doomed to return to a mound of inorganic sludge. A vast
lake of fresh water has been found beneath the Antarctic ice and scien­
tists are debating whether to drill into it in the hope of finding yet an­
other universe of life.
More oddities are hidden beneath our feet. They made their presence
known with the collapse of the Cairo sewers soon after they were built.
A new bug had eaten the concrete. It was the first of many. Novel king­
doms have now been found two miles into the earth. As some survive at
a temperature of a hundred and thirty degrees Celsius, such forms may
exist another mile further down. Dead bacteria gush from ocean vents,
evidence that the land under the deepest seas is alive. Some unique crea­
tures were discovered in a search for oil, ten thousand feet under Vir­
ginia. They live in what was a stream, buried for two hundred million
years below layers of sediment. They have been isolated from the world
since the days of the dinosaurs.
288 D A R W I N ' S G H O S T

The Underworld-like much of the sea, but hot where that is cold­
is an austere place. Its denizens use not oxygen but minerals such as iron
ore to burn the carbon used for food. They are a million times less abun­
dant than the inhabitants of the topmost layer of soil, and divide not
every twenty minutes but once a century. Even so, the newfound land in­
side the Earth holds one part in a thousand of the entire mass of life. Re­
mote as it seems, the underground universe impinges on our own.
Natural gas is the product of archaea, who cause change and decay in the
deepest mines (which should worry those who hope to use them to store
nuclear waste) . Life beneath the land, as yet little studied, already looks
so distinct from even its submarine cousins that the gulf between
mankind and the animals and plants around us will, in comparison,
shrink still further.
Genes mean that the whole of existence can now be arranged into a
single natural system. Evolutionists are obsessed with pedigrees. Their
desire to draw family trees based on flimsy information with man on the
topmost twig has led them into many blind alleys. Now, at last, it is pos­
sible to draw a true map of relatedness; a plan seen from above, with no
declaration of where life's journey begins or how high or low each branch
might be. Like the London Underground, much of its substance is hid­
den away, but the map cares not at all for that.
The map of the Underground twists the metropolis out of shape to
suit the perception of travelers. The distance between stations is exag­
gerated in well-known parts at the expense of distant suburbs. As a re­
sult, a place like Wimbledon-larger than the West End or the City of
London-dwindles into apparent insignificance. To a tourist, it seems
scarcely to exist. The plan of the Paris Metro, in contrast, is less biased as
it shows the real intervals between stops, how far each district is from the
center, and the size of every suburb.
Before DNA, our image of the map of life was as distorted as that of
a tourist on the Underground, with too much attention paid to its fa­
miliar sectors-animals, plants and mushrooms-and very little to its re­
mote extremities. The genes do for life what the Plan du Metro does for
Paris: they reveal the real and unexpected contours of the great city of ex­
istence.
The accompanying diagram will aid us in understanding this perplex­
ing subject. It shows the plan of life as revealed by one set of highly con­
servative genes, those involved in the machinery that helps assemble
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 2B9

proteins. Crowded and unfamiliar neighborhoods cover-as do the sub­

urbs of any large town-most oflife's territory. Two districts, the animals
and the plants, are known to all. Like Westminster and Chelsea they
loom large in the eyes of visitors, but represent only a small part of the
whole. Even the fungi are on the same branch line as those celebrated
places. The rest of town is much farther away than it seems to the unin­
formed traveler, and is little known to anyone apart from a few cartogra­
phers (and those who live there) .
On the true chart of the genes, the bacteria occupy a great Wimble­
don of life: a large neighborhood of their own, with the archaea in the
suburb next door. Three groups known as the diplomonads, trichomo­
nads and microsporidians are diffused through a nearby district. They
sound obscure, but among them are the gut parasite that causes severe
cases of travelers' diarrhea and a single-celled creature responsible for in­
flammation of the vagina. Although they have a cell nucleus, such crea­
tures lack mitochondria.
The rest of town is filled with creatures blessed with both those useful
structures. Kinetoplastids-single-celled animals with a tail that lashes
them through the water-include agents of disease such as the try­
panosome that might, perhaps, have infected Darwin in South America
and led to his many years of invalidism. Next door is a thriving group of
amoeba-like creatures; and two separate groups of slime molds (much
used in the study of development). Alveolates include the ciliates (single­
celled animals covered with fine mobile hairs), the agent of malaria, and
the dinoflagellates, creatures enclosed within the solid shells that make
up much of the chalk that covers southern England.
Some areas traditionally seen as one are, like the East End of London,
in fact several, of different character. The algae-seaweeds, waterweeds
and the green film found on treetrunks-comprise three separate groups,
each with an identity as distinct as is that of the animals from the plants.
Red algae, on a branch of their own, include the seaweeds much eaten in
Japan, together with others that make reefs. The brown algae contain fa­
miliar seaweeds such as kelp, but they belong with diatoms, tiny shelled
creatures that abound in the ocean, in a group called the stramenopiles.
Green algae, found in freshwater ponds, are different again, and live close
to the familiar plants. Plants, animals and fungi are near neighbors in a
well-explored but minor part of the metropolis of life.
An Underground map does not show where any journey begins. How-
 tv
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DIAGRAM OF DIVERGENCE OF TAX A

Genealogy redrawn .from original by M L. Sogin

 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 29 1

ever, the chart of life suggests that the first genes were those of the sim­
ple creatures of the underworld, the archaea, lacking cell nuclei and mi­
tochondria as they do. The idea makes sense: after all, three and a half
billion years ago, the surface was bombarded with meteorites big enough
to evaporate the oceans and flooded with ultraviolet strong enough to
sterilize an elephant. If so, man-with most of the rest of creation-as­
cended from Hades rather than stepping down from the Garden of
Eden. Genetics has drawn a new map of hell, with humankind firmly
placed in the suburbs.

Evolution can sometimes hide a true pedigree. It would be foolish to clas­

sify flies and bats as the same because both have wings. Their similarity
comes not from shared descent but from separate answers to the same
problem. It is shallow indeed, based on external appearance rather than
essential structure, and as a result is not much use in taxonomy. Charac­
ters are of real importance for classification only insofar as they reveal de­
scent. Analogical or adaptive characters, although of utmost importance
to the welfare of the being, are almost valueless to the systematist.
That can sometimes be very clear. The females of a certain moth and
of the Mrican elephant each make the same complex sex pheromone.
This attracts males of either kind (which must be riskier for the moth
than for the other partner in the relationship). For some reason, the mol­
ecule hints at sex, and insect and elephant have each taken it up through
different routes.
In evolution, parallel lines often converge. As they do, they conceal
real patterns of affinity. Polar bears, Arctic insects and the leaves of north­
ern plants are all covered in fur, but nobody calls them relatives because
they share a taste in coats. The fish of those icy waters show how the dis­
tinction between true alliance and separate solutions to the same prob­
lem can be concealed by the amazing efficiency of natural selection.
The Antarctic perch has an antifreeze, a remarkable protein that al­
lows it to live at two degrees below zero, a temperature at which most fish
would turn to ice. The gene responsible resembles another that makes a
digestive enzyme in the fish's own stomach. That job came first. Its role
was to prevent the stomach from freezing solid, but when the substance
was manufactured in the liver as well, the Antarctic perch came up with
an antifreeze for its whole body.
292 D A R W I N ' S G H O S T

The protein (or another identical in structure) is also found in the

Arctic cod, a fish from the other end of the earth-which is a surprise,
because the two are but distant relatives. All other evidence-from
anatomy, from geology and from geography-agrees that they split forty
million years ago, long before the Antarctic froze. They differ in their
body structures and in many of their genes.
The antifreeze is, in fact, a separately evolved adaptation in each fish.
The Arctic and Antarctic proteins, specialized as each is and with almost
the same structure, are not evidence of common descent but-like the
furry coats of polar bears and northern plants-of a shared solution to
an evolutionary challenge. Natural selection has done its job so well that
it has fashioned two identical molecules at opposite ends of the earth.
Their convergence shows how animals belonging to two most distinct
lines of descent may readily become adapted to similar conditions and
thus assume a close external resemblance; but that such resemblances will
not reveal-will rather tend to conceal-their blood relationship to their
proper lines of descent. For the two protective proteins, an identical pres­
ent conceals a separate past.

Morphology. The members of the same class, independently of their

habits of life, resemble each other in the general plan of their organiza­
tion. This resemblance is often expressed by the term "unity of type," or
by saying that the several parts and organs in the different species of the
class are homologous. The whole subject is included under the general
name of morphology. This is the most interesting department of natural
history and may be said to be its very soul. What can be more curious
than that the hand of a man, formed for grasping, that of a mole for dig­
ging, the leg of the horse, the paddle of the porpoise, and the wing of the
bat, should all be constructed on the same pattern and should include
the same bones, in the same relative positions?
Today's students despise comparative anatomy (as the study of such
structures is now called) . Many refuse to cut up animals on what they see
as moral grounds and the remainder find the whole business tedious and
irrelevant. They move with relief to the double helix. Although they may
not realize it, they are practicing anatomy without a license. The nine­
teenth century used bones, but now we have molecules. Much of today's
biology is no more than the study of the structure of genes in related or­
ganisms. Instead of cutting up guts or skulls, biologists now sequence the
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 293

DNA that makes them. To do so reveals most oflife to be a set of themes

and variations.
All complex beings are built from a series of repeated and modified
segments. Sometimes that is plain. The earthworm is made up of dozens
of sections, many of which look much the same, but others are modified
to include eyes, sex organs and the like. It can regenerate its whole body
from a fragment of its original. Such an ability raises profound issues.
The eighteenth-century Swiss naturalist Charles Bonnet was much con­
cerned: "Must we admit that there are as many souls in these Worms as
there are portions of these same Worms?" The biological questions are
easier to answer than are the theological. For the earthworm, each seg­
ment retains within itself the essence of a complete animal; proof that,
different as they appear, each part of its body is constructed on the same
plan. Natural selection, during a long-continued course of modification,
has seized on a certain number of the primordially similar elements,
many times repeated, and has adapted them to the most diverse pur­
poses.
Just the same thing goes on among the molecules. An inbuilt tendency
to alter in number can make Darwinian sense. Aphids resistant to pesti­
cides do the job with a vast increase in the dose of the genes able to break
the chemicals down. Duplication is also favored because one copy can
concentrate on part of the task in hand while another sets off on its own.
The hemoglobins live as families, changed versions of an original, each
now a specialist at a different job. Long ago, there was but a single trans­
port molecule, which doubled up several times to give two great groups
of genes. One member of each family makes the hemoglobin found in
adults while others produce versions useful in the fetus, which must ca­
jole oxygen from its mother's blood. Each protein has its own task but
retains, like the segments of a worm, the stamp of its shared ancestry.
The strange resilience of the mouse hints as to how much repetition
there must be even within the mammalian frame. A technical trick
makes it possible to knock out particular genes. Sometimes, these in­
complete mice are in deep trouble and die young. More often, they live
blithely on, although what seem essential parts of their machinery-a
gene for collagen, the structural material of much of the body, or for an­
other that passes signals around the cell-have been removed. Duplica­
tion is a useful insurance policy against the wiles of geneticists; but how
and why these extra copies evolved, nobody knows.
294 D A R W I N ' S G H O S T

In biology, repetition is everywhere. Worms, insects and leeches are

built of a series of modules, and even in higher animals elements of such
a plan can plainly be seen. Parts many times repeated are eminently li­
able to vary in number and structure. A glance at the teeth of any verte­
brate shows how a simple and much-duplicated element has been
modified for its job of biting, tearing or chewing. The mammal body
does, nevertheless, retain a certain individuality in its various parts: a cat's
skull has not, it seems, much in common with the tip of its tail.
Vertebrates-the group to which humans, snakes, birds and frogs be­
long-have some modest relatives. They include the lamprey and the
lancelet. The lancelet looks like an anemic anchovy fillet, but has no eyes,
no ears, no jaw, no discernible skull even. The Chinese thought it to be
a maggot grown from the corpse of a divine crocodile bearing the god of
literature, but many Western scholars denied even that it had a brain. In
the days when students could stand the sight of blood, both lamprey and
lancelet were much dissected. To anatomize their genome rather than its
products discloses their role in history and hints at how mammals gained
the organ of literature itselE
The mammalian skull has scores of bones. The eye and ear are tri­
umphs of complexity and the brain so elaborate as-so far-to be un­
able to understand itselE The lancelet is less pretentious. It has changed
little from the ancient animals that gave rise to humans, snakes and birds.
The great nineteenth-century German biologist Ernest Haeckel was con­
vinced of its closeness to the earliest vertebrates. So certain was he of its
importance that lancelets on caviar were served on his sixtieth birthday
(although the menu also makes the enigmatic claim that this was fol­
lowed by a main course of Archaeopteryx with sauerkraut).
Where did our skull come from and why should the brain be enclosed
in a box composed of so numerous and such extraordinarily shaped
pieces of bone? Is it a modified piece of an ancient body, or an extra struc­
ture bolted onto an insensible frame as a container for a new and mighty
organ of thought? Because lancelets lack that noble structure it was long
thought that vertebrates evolved the skull as a new element, a sort of
hood ornament attesting to the power of the machine within. The pat­
tern of repeats shows that, instead, vertebrates have an old head on young
shoulders.
The genes in charge of the machinery for making skulls are arranged
 Mutual Affin ities; Morphology; Embryology; Rudimentary Organs 295

i n groups. One set is found in all vertebrates, from lancelet to man. A

young fish or a human embryo has a series of regular blocks of tissue that
develop into the complicated bones and muscles of the body. In the adult
the simplicity of their recurring structure is lost, but the genes show that
there are parts in the same individual which become unlike and serve for
diverse purposes.
A test of a young lancelet with a genetic probe derived from mammals
shows that, although the animal lacks a head, it has the DNA later used
to make it. Brainless though it seems, the genes that in other creatures
make a discrete forebrain are also at work. Each is active at the same stage
as in mice or chickens. The advance guard of the brain and skull have, it
seems, been around since the start. Some of the details have changed.
The lancelet has a single light-sensitive organ at the tip of its head-but
a gene almost identical to that for the twin eyes of mammals builds its
lone eye. Cyclopes though they may have been, we do not stand head
and shoulders above our ancestors.
The road to mammals called for other changes. It takes more to make
a mouse than a lancelet; and mice or humans have extra copies of all
lancelet genes, because, somewhere on the road from their common an­
cestor, their number doubled, twice over. One group, the ray-finned
fishes, had yet another doubling after they split from the lineage that led
to mammals.
All genes need to know not just what to make, but where, when and
how often to make it. Changes in time or place can have great effects on
body form as a structure loses its way on the inborn map. Some muta­
tions alter not the shape or function of an organ, but where and when it
appears. By so doing, they allow the primordial elements to make their
presence felt. In fruit flies, single mutations can cause the middle section
of the body to duplicate itself, to give a fly with four rather than two
wings. To make a thorax takes thousands of genes, but a simple com­
mand can, it seems, set the whole army into motion. The additional or­
gans appear because an order is given in the wrong place and the local
cell machinery obeys it.
The switches in control of development are called the homeobox
genes. They are arranged in groups of ten or so. Invertebrates have a sin­
gle copy of each group, stretching over a hundred thousand or so DNA
bases. Mammals have four, with many tasks in the embryo (and others
296 D A R W I N ' S G H O S T

in the adult; with mutations in homeoboxes leading to baldness and even

to leukemia). Ray-finned fish have seven or eight copies of the crucial
segments. They lay claim to twenty-five thousand species, from sturgeon
to salmon-as many as all other vertebrates combined. Such fish put
mammals to shame, with seahorses, flatfish, anglerfish, cichlids, eels and
thousands more. Perhaps their extra homeoboxes, the masters of devel­
opment, allow them to experiment with new and eccentric sets of body
form.
The switches are arranged in series, each in charge of a separate part of
the body's battalions. Their sequence, from front to back, is the same as
that of the organs for which they are responsible. k a result, homeoboxes
lead from the front. Those nearer the head can control structures behind
them, while those farther back have less influence on parts of the body
further forward. Changes in such genes persuade different segments to
develop into head, heart or tail. The pattern of central command is com­
mon to mice, fruit flies, snails and every other animal. Ownership of a
homeobox may, indeed, be as good a way as any other of defining what
an "animal" might be. They decide what their bearer will look like.
Baleen whales have a slight quirk near one of the structures, with four
DNA bases missing from part of the mechanism. When transferred to a
mouse embryo, the tiny change moves the activity of certain neck seg­
ments farther back, explaining, perhaps, the whale's great head and enig­
matic smile.
Plants, too, are built on a multiplied plan. It is familiar to almost
everyone that in a flower the relative position of the sepals, petals, sta­
mens and pistils, as well as their intimate structure, are intelligible in the
view that they consist of metamorphosed leaves, arranged in a spire.
Monstrous plants often give direct evidence of the possibility of one
organ being transformed into another. Any gardener understands the
power of monstrosity: roses, in their native state a plain and unremark­
able flower, have been persuaded to double up their petals from the five
or so in wild roses to ten times as many in varieties such as 'Elizabeth Tay­
lor' and 'Blushing Bride.' k in the multiwinged fruit flies, such freaks
arise through changes in genes that control the number of copies of a
compound structure.
Flowering plants grow in a simple way, from a group of cells at the tip
of each shoot. k these divide, they make small clusters of active cells,
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 297

each destined to be a leaf or a petal. Different though the Hower of a

snapdragon or a rose might appear, the first as a mirror image, with a left
side and a right, the second arranged like the spokes of a wheel, those or­
gans are at an early stage of growth exacdy alike. The DNA also shows
that, complicated although the parts of a Hower might be, they are built
to simple instructions. Various mutations change the number and posi­
tion of parts such as petals and some can persuade a two-sided Hower to
take up a disc shape. To damage just two crucial genes can persuade a
Hower to develop as a simple leaE A mutation in another member of its
triumvirate of control can double its entire structure. A garden rose's
complexity depends, like that of a worm, on a set of modified and re­
peated units, honed by natural selection from simple parts. Life has, it
seems, developed like opera, from short pastorals with a single motif and
a limited range of players to the great music dramas of the garden, the
farm and the forest.
Biology has also begun to reveal a complex set of themes and varia­
tions within the genes themselves. Many proteins are built from a series
of modules shared with others that do quite different jobs. Thus, the nu­
merous proteins involved in blood clotting are made up of four distinct
sections, shuffied together in various ways to make a useful device. Not
all the members of the clotting chain have all the units, but all share
pieces in common. Some elements of the blood-clot apparatus also ap­
pear in genes that control the growth of cells or digest meat. They
demonstrate how hidden motifs may appear, in true Wagnerian style,
deep within the cellular plot, and how the complex structures of today
retain-like the rose, the earthworm, or the song of the Rhine Maidens
as Valhalla burns-distant echoes of a simpler past.

Embryology. "We saw the Emperors standing all together huddled under
the Barrier cliff some hundreds of yards away. The litde light was going
fast: we were much more excited about the approach of complete dark­
ness and the look of wind in the south than we were about our triumph.
After indescribable effort and hardship we were witnessing a marvel of
the natural world, and we were the first and only men who had ever done
so; we had within our grasp material which might prove of the utmost
importance to science; we were turning theories into facts with every ob­
servation we made,-and we had but a moment to give."
298 D A R W I N ' S G H O S T

Thus Apsley Cherry-Garrard in his extraordinary book The WOrst

Journey in the WOrld. It is the tale of a trip through the Antarctic winter
of 1 9 1 1 to collect the eggs of the Emperor penguin; part of the journey
(ill-prepared as only a British expedition could be) that ended with the
failure to reach the Pole and the death of Captain Scott, weighed down
by his precious rock specimens, a few miles from safety.
But why risk death to fight in total darkness across the ice? What was
this "material which might prove of the utmost importance to science"?
The answer lies in The Origin, in its claim that each animal relives its an­
cient history as an embryo. To biologists of Cherry-Garrard's generation
the story of an individual as it developed was that of the species to which
it belonged. To read the narrative that begins when sperm meets egg and
ends with death was to retrace the past. The eggs of the Emperor pen­
guin, he believed, held the due to the origin of birds: "These three em­
bryos from Cape Crozier . . . were striven for in order that the world may
have a little more knowledge, that it may build on what it knows rather
than on what it thinks."
Biology, unlike physics, has few laws and those made are soon broken.
In 1 866 Haeckel came up with his "biogenetic law" : new forms are, he
said, the result of extra stages added on to the development of those that
went before. He published sketches of the embryos of birds, reptiles and
mammals (somewhat doctored to support the idea) that seemed to prove
his case. The Emperor penguin was the most primitive avian form,
pushed to the edge of the world by competition from more recent-and
more advanced-relatives to the north. Its embryo would reveal the ori­
gin of birds.
After many troubles, and a studied rudeness by the Natural History
Museum when Cherry-Garrard tried to deposit his specimens, the eggs
turned out to be well on in their growth and of no use as a test of
Haeckd's theory. We now know that penguins split off from a winged
ancestor forty million years ago and are no more primitive than is the al­
batross. The science upon which the Antarctic travelers depended was
wrong, but life before birth reveals more about evolution than they ever
dreamed. The question that seemed so important to naturalists of their
day-the development of a complex organism from a simple egg-is on
its way to a solution.
The Origin makes no mention of "evolution," a word whose sense has
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 299

gone full circle since it began. In its Latin root it referred to the unfold­
ing of a scroll. As Cicero so memorably put it: "Quid poetarum evolutio
voluptatis affert'-' "What pleasure does the reading of the poets pro­
vide!" The word was first used in biology to describe the changes in shape
of an embryo as it developed. Not until much later did "evolution" begin
to suggest the gradual transformation of one form into another. Now, its
definitions have come together and the study of development is un­
rolling the scroll of biological history.
The power of the unborn was recognized within five years of The Ori­
gin. Charles Kingsley's The water Babies is a tale of a young chimney
sweep drowned, transformed into an infant and washed clean of his sins.
If evolution culminated in that miracle, the Englishman, why could it
not bring forth the greater miracle, the soul-and where better to find it
than in an English baby? The drowned Tom "when he woke . . . found
himself swimming about in the stream . . . having round the parotid re­
gion of his fauces a set of external gills, just like those of a sucking eft,
which he mistook for a lace frill, till he pulled at them, found he hurt
himself and made up his mind that they were part of himsel£" He relives
his innocent past as a young newt or eft, then a sinful child, and-at
last-enters into the post-evolutionary state with which the scroll of life
should end. Not for the last time, science and theology were in perfect
harmony.
The new embryology is based, like that of Charles Kingsley, on a
search for resemblances among young animals that may be lost later in
development. Now at last the embryo is revealed as a picture, more or
less obscured, of the common parent form of each great class of animals.
It uncovers hidden patterns of relatedness because an embryo, sheltered
as it is from the need to adapt to the world outside, may retain more of
its past than does an adult. An embryonic elephant has kidneys rather
like those of a manatee, adapted to life in water. Perhaps long ago an am­
phibious mammal split into two, one taking to the water while the other
returned to land (but still fond of a bath and of squirting itself with a
trunk that was once a snorkel) .
The case is different when an animal during any part of its embryonic
career is active, and has to provide for itself Then, natural selection acts
to modify even the youngest, and their history is lost. The period of ac­
tivity may come on earlier or later; but whenever it comes on, the adap-
300 D A R W I N ' S G H O S T

tation of the larva to its conditions of life is just as perfect and as beauti­
ful as in the adult animal. Take, for instance, clams, oysters and the like.
All look much the same as adults, but their larvae are quite distinct.
Some float for months as plankton, while others develop within their
mothers' bodies; some are hunters, while others burrow into the flesh of
fish. Selection can, when called upon, work as hard on the embryo as on
any other stage of life.
A few animals have the choice of whether to advance into adulthood
or to stay as juveniles. Tom went backward into the water with his "set of
external gills" that allowed him to give up breathing air. The axolotl, a
North American eft, goes the other way. In some places, it spends its time
in lakes, with gills as impressive as any water baby's, and reproduces in
this stage. In others, it grows through its adolescence and clambers onto
land, where it has habits close to those of any other salamander. Crosses
in the laboratory reveal that a single gene controls the difference between
them.
Some axolotls can be prompted to abandon their childish habits with
a dose of thyroid hormone, which makes them grow legs. Others resist
such unwanted maturity and stay in the water-baby stage even after a
dose of the chemical. Somewhere in the chain of development is a switch
that turns on the biological factory needed to make a land axolotl. It re­
veals the hidden hand that rules the embryo's fate and shows how a few
pieces of DNA can direct a series of underlings that, when set into ac­
tion, make legs, lungs and everything else a terrestrial animal might need.
Such hidden potential is everywhere. The new embryology shows that
most tissues in most creatures have the capacity to develop into quite un­
expected structures. Most of the time they do not (although plenty of
creatures have a change in lifestyle with age at least as great as does the
axolotl), but, now and again, they reveal their powers and, by so doing,
their evolutionary past. The evidence lies in a hierarchy of ancient com­
mand.
Often, that past stretches amazingly wide, with similar genes in con­
trol of very different structures. One ancient gene has spawned a long­
lasting dynasty of power. The descendants of the first Duke of Habsburg
found themselves, over seven centuries, in control of Austria, Spain and
Hungary and of parts of Germany, Italy and even of Mexico. Although
their nations varied from semi-democracies to the revolutionary chaos of
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 30 1

Central America each was ruled by a member of the same family and fol­
lowed, with more or less enthusiasm, the instructions of their hereditary
monarch. Certain genes, it seems, have much the same history.
The eyes of mice and of fruit flies are quite unalike. One is a camera
with a single lens, the other a whole host of small devices arranged into
a complex machine. Fruit flies and mice each have mutations that lead
to the loss or great reduction of the eye. Although their organs are so dif­
ferent, the gene that reduces or removes the eye in flies is almost identi­
cal in structure to that which reduces its size in mice.
A certain fly mutation affects a length of DNA that-like the master
switch of the axolotl-turns on a whole series of eye genes. When it is
persuaded to act !fi unsuitable parts of an embryo, eyes grow in unlikely
places, such as the wing or the leg. Quite remarkably, the normal version
of the mouse-eye gene, moved to a fly embryo, induces the growth of
eyes, with the typical fly structure, wherever it lands-legs, wings or any­
where else. It steps into its ancient and dictatorial role and the cells of its
new host respond. The emperor of the eye is much the same in mammals
and insects, although these groups last shared an ancestor a billion years
ago. Either an ancient light-sensitive structure-a patch of cells-took
two different routes down the lines that led to flies and to mice, or two
separate visual organs elected to obey the same ruler. Whatever the truth,
to grow fly eyes under the instructions of a mouse gene shows how uni­
versal is the history revealed before birth.
Other parallels in the development of flies and backboned animals
prove how far the genetic gridlines stretch. The ear, like the eye, has an
unexpected ancestry. Fruit flies cannot hear much, but are easily irritated
because they are covered by bristles that sense vibration. The same genes
are at work in the inner ears of fish, chickens and mice. In the same way,
the gene specifying where the rear half of each fly segment should be does
the same job in the young lancelet. Genes switched on at the tips of the
young limbs of a variety of creatures reveal another deep and ancient
connection; a footrace that ended up with wings, a starfish's tube feet,
horses' hooves and the hollow legs of crabs, all descended from some sim­
ple and primitive limb.
One of the more outlandish suggestions of the old anatomy was that
humans and their relatives are invertebrates turned upside down. In
1 822, the French biologist Geoffroy St. Hilaire (several times referred to
302 D A R W I N ' S G H O S T

in The Origin, but more famous in France as the guardian of the first gi­
raffe to enter the country) saw a dissected lobster lying on its back. He
was at once struck by its apparent similarity to a man lying on his stom­
ach. In both, the spinal cord is above the intestine, and the esophagus
above the heart. Perhaps, he thought, life had Hipped over on the way to
the vertebrates. The idea seemed absurd: French biology again on its own
eccentric path, with the rest of the world out of step. A century and a half
later, the anglophones had reason to be embarrassed when a gene active
in the cells of a frog embryo that make part of its back turned out to be
at work on the lower surface of the body in developing fruit Hies. There
has indeed been a great somersault on the way to ourselves.
For eyes, ears, limbs, and deciding which way is up, the embryo is left
as a sort of picture, preserved by nature, of the ancient and less modified
condition of each animal. Its image is proof of the course of evolution
from a time older than the earliest fossil.

Rudimentary, atrophied, or aborted organs. The mass is a Christian eter­

nal. As an exchange with God it has its roots in the earliest Church.
Wherever he might be, a believer finds himself at home and-except for
the language differences that emerged with the abandonment of Latin­
part of a universal ceremony.
Not, however, in Ethiopia. In a minor triumph of comic prose, Eve­
lyn Waugh (who visited the country in 1 930) described the responses of
an expert to his first experience of a local Mass: " 'That was the offertory
. . . No, I was wrong; it was the consecration . . . I think it is the secret
Gospel . . . the Epistle . . . I have noticed some very curious variations in
the Canon of the Mass . . . particularly with regard to the kiss of peace.'
Then the Mass began."
His liturgical confusion has a message for biology. The Ethiopian
Church was cut off for a thousand years, safe from the reforms that seized
the rest of Christendom. As its rite passed down the generations, the er­
rors grew until whole sections made no sense even to those who cele­
brated it. They are of interest to theologians as they reveal a history
elsewhere swept away by the march of progress. All structures degenerate
as soon as their job is done. Once selection ceases, the chaos of nature
sets in and evolution loses its way. A relic may hence be better evidence
of the past than is the most exquisite adaptation.
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 303

A glance around the world shows general decay: parasites reduced to

sacks of guts and genitals, useless organs such as the human appendix,
and great stretches of DNA that make no protein. The sea squirt, after
an active life, settles on the sea floor and, like a professor given tenure,
absorbs its brain. Whole communities may turn to degeneracy. More
than a hundred thousand different kinds of animals-spiders, insects,
fish and salamanders-live in caves. Most are pallid imitations of their
open-air relatives: slow, blind, calm and solitary when compared to their
cousins who live in the glare of day.
Such characters, bearing as they do the stamp of inutility, have long
been seen as links with the past. A traveler to Tibet spoke of a "species of
human being with short straight tails, which, according to report, were
extremely inconvenient to them, as they had to dig holes in the ground
before they could attempt to sit down." That organ (and a few children
are born with tails) was much used in pre-Darwinian times to make a
link between humans and apes. A structure that has lost its purpose
proved descent from an ancestor to whom it was still useful.
A complicated structure can work in only one way but can go wrong
in many. A cross between blind cave fish from different places gives off­
spring with larger eyes than either parent. The fish from each cave have
lost different parts of the machinery and to put two of them together
cobbles together something better than either can manage on its own.
Other rudimentary organs, too, sometimes retain their potentiality and
are merely not developed. Why is the milk of human kindness made by
only half the population? After all, men have nipples and the capacity to
use them. Males given certain chemicals lactate with no difficulty. Even
a heavy dose of alcohol can do the same as the liver loses its ability to sup­
press each man's guilty secret, his female hormones. Teenage boys, in a
natural desire to see what might be done with their bodies, now and then
stimulate their own nipples and (no doubt to their amazement) may
eject milk. The Dyak fruit bat is the sole mammal to reach the logical
conclusion, as its males suckle their young, but as the males of many oth­
ers invest lots of care in children it seems strange that they do not go the
whole hog and make food themselves.
A man's pert but useless nipples are his real stamp of inutility. They
have no job because of evolution. Many males leave at fertilization and
do not hang around for birth. Their energies are invested in more sperm
304 D A R W I N ' S G H O S T

rather than in the results of an earlier fling. There is no market for their
milk. Because the female is already committed through an expensive
pregnancy to her child, it does not pay her to abandon it in the hope of
success with another. Selection has turned her into a milk machine. The
male's job, if any, is to protect his child as he keeps a weather eye open
for what else might turn up. His nipples are just a reminder that he is of
common stock with his partner.
An organ serving two purposes may become rudimentary or utterly
aborted for one, even the more important purpose, and remain perfectly
efficient for the other. Tiny bones inside the ear of mammals-the ham­
mer, the anvil and the stirrup, as they arev�alled-act as a series of levers
to transmit sound waves from the eardrum to the sense cells. They trace
their evolution to organs in ancient fish, useful in quite different ways,
but now decayed. The most primitive fishes had not jaws but supports
called gill arches. The first jaws were made of hijacked copies of these
structures, although the rest stayed on as struts for gills. As fish evolved
toward mammals, the arches were lost, but some of the tiny bones of the
middle ear remain as their descendants. The rest of the ear has the same
utilitarian history. Fish have a lateral line, a network of receptors that
stretches along the body and detects changes in water pressure. The sen­
sory part of our own organ of hearing descends from the ancient pressure
sensor of fish (and, the genes show, in the end from a more distant an­
cestor shared with the irritable hairs of flies), but all other signs of its
presence have gone.
Plant parts, too, can have a change of career. Climbers may use re­
duced leaves (as in the grape), leaflets (the trumpet vine) or parts of the
flower (blackberries) as they clamber up their supports. Each tendril or
hook has lost one job to take up another, as evidence of the expedience
of evolution when offered a structure that might be induced to help in a
new way. Natural selection does not hesitate to pick up and use whatever
becomes available. But, for most of the time, decay is just decay and life
has, with a sigh of relief, given up what it does not need.
Rudimentary organs, such as the upper jaws of whales, can often be
detected in the embryo, but afterward wholly disappear. Kiwis, while still
in the egg, have wings as big as those of embryonic albatrosses; but they
never grow. The instructions may be retained even when the structures
themselves are lost. A bird can be persuaded to grow teeth in its devel-
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 305

oping beak if a few appropriate mouse cells are moved nearby to switch
on those long-dormant genes. Once again, economics is at work. As the
main expense of making wings or teeth comes later in life, natural selec­
tion is not much interested in making cuts in places where no real sav­
ings are made.
There is a message for evolution hidden in its discarded baggage: it is
heavy, and costs a lot to carry. An unencumbered traveler can use the en­
ergy in other ways. The biggest burdens are shed first. The speed with
which they are lost hints at the cost of keeping them. H. G. Wells, in
The mtr of the Worlds, has his Martians reduced to mere gutless heads
that suck the blood from men and women. They come from an old, dry
planet, short of oxygen. On Earth, they struggle against gravity-a "cope
of lead. » They have abandoned all but the capacity to think.
Take, in contrast, that most artificial of beasts, the chicken. It has
taken the opposite route to the Martians. It joined the household in Asia
and was used to divine the future (entrails were most reliable) long be­
fore the Romans turned it into food. Because their wild relatives, jungle
fowl, are still with us, today's chickens show how evolution has cast off
its burdens. They have become animated machines to turn food into
flesh, with lighter bones, weaker muscles and smaller brains than jungle
fowl. Why spend energy on thought when man does it for you? Brains
have been sacrificed to make guts. Chickens have intestines three times
as long as those of their ancestors, but much reduced intellects.
Jungle fowl, unlike their descendants, can still fly. Wings are expensive
and have been abandoned again and again. Females are more liable to
lose them than are males. A simple but brutal experiment shows why.
Cut off the wing of a young cricket, and she lays more eggs. Birds, too,
have shed the organ gained with such effort by their dinosaur ancestors.
Some are unique to coral islands a mere hundred thousand years old and
have cast off much of the structure of their wings in that time. The dodo,
a turkey-sized flightless pigeon, went further along its evolutionary path
than anything achieved by the breeders. Madagascar was once full of ele­
phant birds (flightless, as the name suggests), most Pacific islands had
their own ground rails, and the Galapagos retains its flightless cor­
morant. In New Zealand, more than thirty different birds are confined
to the ground-moas, kiwis, ducks, geese, rails, crows, wrens and a par­
rot. With no enemies to flee from, and nowhere to go, why fly?
306 D A R W I N ' S G H O S T

Many flightless forms are sluggish in comparison to their winged kin.

Small islands are hungry places and a slowdown may be the only way to
cope. Island birds lose more than wings. Most are smaller than their
mainland relatives-sometimes half the size-and can consume a fifth as
much energy. The bones of reduced elephants found in Cyprus gave
Jonathan Swift the idea for Lilliput, a land of insular dwarfs. Lethargy
needs less food, and the torpid survive where those full of energy would
starve. Six-ton mammoths were reduced to two-ton midgets on Wrangel
Island in the Arctic Ocean, and red deer marooned on Jersey after the ice
age withered in a mere six thousand years to the size of a St. Bernard dog.
They were forced to shrink or starve. Sex, too, is a costly pastime. Pota­
toes kept as clones can still flower, but their sexual advertisements are fee­
ble compared to those of their wild relatives. For potatoes advertising is
expensive. Its hoardings soon rust when not burnished by natural selec­
tion.
Given all this emphasis on efficiency, it is a surprise to find great tracts
of repetition and decay within the DNA. There are not just thousands of
repeats of the same message, but hundreds of dilapidated ruins of what
were once working genes. Such pseudogenes, as they are known, are
everywhere, in mammals at least. The hemoglobin gene family has half a
dozen, each corrupted almost beyond recognition. Long ago, a mutation
destroyed the switch that turns the gene on, or inserted an instruction
that it should stop doing its job, or damaged its ability to edit its mes­
sage. Some pseudogenes are the remnants of viral attack and have been
read back into the DNA from an edited version of the genetic message,
to be scattered where they fall. Evolution at once lost interest: as soon as
the gene stopped work it was, in effect, invisible. Such structures sit for
millions of years and crumble until their shape can barely be discerned.
Why do rotting genes persist and why is life happy to accept them
when it so soon gets rid of unwanted wings or eyes? Perhaps DNA is
cheaper than the flesh and bones it makes. Pseudogenes, like other rudi­
mentary organs, retain more of their family relatedness than do those still
scrutinized by natural selection. A pedigree based on decay reveals de­
scent with modification better than one in which most changes have
been removed. As a result, true relatedness is often best revealed in char­
acters that may be considered rudimentary or atrophied; both in DNA
and in the leg bones that draw the resemblance between whales and deer.
 Mutual Affinities; Morphology; Embryology; Rudimentary Organs 307

In their functional parts, the antifreeze proteins of the Arctic cod and
Antarctic perch are the same. They say nothing about the affinity of the
two fish. However, the genes for antifreeze contain sections of DNA that
make no product. Such DNA-a rudimentary or useless characteristic
indeed-reveals the real patterns of kinship. The Arctic cod's inclusions
are quite different from those of the Antarctic animal. Unlike the protein
itself, they tie it to its true relatives, who include the fish pursued with­
out mercy by man.
Such withered lengths of DNA may be compared to the letters in a
word, still retained in the spelling, but become useless in the pronuncia­
tion. They serve as a clue in seeking for its derivation. A message that has
become gibberish, and is transmitted-like the Ethiopian mass-with­
out regard for meaning, retains more of the past than one visible to nat­
ural selection, the great editor who removes all errors and eliminates
history as he does so.
When seen through the lens of evolution, facts as disparate as the true
affinities of birds, the life of the deep sea vents, the multiplied structure
of the earthworm and the nipples of male mammals all come into com­
mon focus as evidence of descent with modification. Together they bear
persuasive witness to change: enough to establish the truth of evolution
even without the mass of supporting evidence given elsewhere in these
pages.

Summary. In this chapter I have attempted to show, that the subordination of

group to group in all organisms throughout all time; that the nature of the
relationship, by which all living and extinct beings are united by com­
plex, radiating, and circuitous lines of affinities into one grand system;
the rules followed and the difficulties encountered by naturalists in their
classifications; the value set upon characters, if constant and prevalent,
whether of high vital importance, or of the most trifling importance, or, as
in rudimentary organs, of no importance; the wide opposition in value
between analogical or adaptive characters, and characters of true affin­
ity; and other such rules;-all naturally follow on the view of the com­
mon parentage of those forms which are considered by naturalists as
allied, together with their modification through natural selection, with its
contingencies of extinction and divergence of character. In considering
this view of classification, it should be borne in mind that the element of
308 D A R W I N ' S G H O S T

descent has been universally used in ranking together the sexes, ages,
and acknowledged varieties of the same species, however different they
may be in structure. H we extend the use of this element of descent,­
the only certainly known cause of similarity in organic beings,-we shall
understand what is meant by the natural system: it is genealogical in its
attempted arrangement, with the grades of acquired difference marked
by the terms varieties, species, genera, families, orders, and classes.
On this same view of descent with modification, all the great facts in
Morphology become intelligible,-whether we look to the same pattern
displayed in the homologous organs, to whatever purpose applied, of the
different species of a class; or to the homologous parts constructed on the
same pattern in each individual animal and plant.
On the principle of successive slight variations, not necessarily or
generally supervening at a very early period of life, and being inherited
at a corresponding period, we can understand the great leading facts in
Embryology; namely, the resemblance in an individual embryo of the ho­
mologous parts, which when matured will become widely different from
each other in structure and function; and the resemblance in different
species of a class of the homologous parts or organs, though fitted in the
adult members for purposes as different as possible. Larvae are active
embryos, which have become specially modified in relation to their
habits of life, through the principle of modifications being inherited at
corresponding ages. On this same principle-and bearing in mind, that
when organs are reduced in size, either from disuse or selection, it will
generally be at that period of life when the being has to provide for its
own wants, and bearing in mind how strong is the principle of inheri­
tance-the occurrence of rudimentary organs and their final abortion,
present to us no inexplicable difficulties; on the contrary, their presence
might have been even anticipated. The importance of embryological
characters and of rudimentary organs in classification is intelligible, on
the view that an arrangement is only so far natural as it is genealogical.
Finally, the several classes of facts which have been considered in this
chapter, seem to me to proclaim so plainly, that the innumerable species,
genera, and families of organic beings, with which this world is peopled,
have all descended, each within its own class or group, from common
parents, and have all been modified in the course of descent, that I
should without hesitation adopt this view, even if it were unsupported by
other facts or arguments.
 I N T E R L U D E

ALMOST LIKE A WHALE?

Difficulties o n the theory o f humans as apes - Evidence that Man is a product

of evolution - Recapitulation of circumstances in which selection works
upon ourselves - Causes of the general belief in the immutability of
man - How far the theory of natural selection may be extended - Ef­
fects of its adoption on the study of Man - Concluding remarks

The lines of doggerel that begin this book are a parody. They mock a
work called The Progress of Civil Society; A Didactic Poem in Six Books,
written in 1 796 by Richard Payne Knight (whose botanical brother An­
drew is, by coincidence, mentioned in The Origin). His instructive poem
is a broad-a universal-survey of humankind set in terms of the animal
world and annotated with the apparatus of Enlightenment scholarship.
Hookham Frere's burlesque ascends from aerial mackerel and aquatic
bears into a kind of gothic dementia (''Ah! Who has seen the mailed lob­
ster rise, I Clap her broad wings, and soaring claim the skies? I When did
the owl, descending from her bower, I Crop, midst the fleecy flocks, the
tender flow'r, I Or the young heifer plunge, with pliant limb, I In the salt
wave, and fish-like strive to swim?") . On the way it makes a serious point
about the limits to biology in human affairs.
The parody has marginal comments and notes to match the dozens
that decorate Payne Knight's futile attempt to use science to explain the
obvious. The annotation to the first line (in which a feathered race skims,
in leaden verse, the air) reads, simply, « l Birds fly."
Humans are a footnote to biology. Evolution is often seen as a progress
310 D A R W I N ' S G H O S T

toward civil society, an overture to Homo sapiens. That view ignores the
substance of life in favor of its details. The last chapter of The Origin of
Species is an abstract of the book's "long argument" in simple, even ur­
gent, terms. It returns with almost poetic intensity to the woodpeckers,
wasps and oaks that make Darwin's case, and contains its most famous
sentence: "Light will be thrown on the origin of man and his history."
His final statement appears as an appendix to the present work.
My own last chapter, unlike its original, is not a digest of the whole,
but a detail of history, a test of how our own past is illuminated by evo­
lution, of whether man follows the rules that govern the whale and the
AIDS virus.
When Wallace asked Darwin whether he planned to include evidence
from humans in The Origin, he answered, "I think I shall avoid the
whole subject, as so surrounded with prejudices; though I fully admit
that it is the highest and most interesting problem for the naturalist."
He was right. In 1 859 not a single human fossil was known, but now
thousands have been found. Genes reveal the true relatedness of people,
peoples and primates. Darwin's entire argument can be made in human
terms and The Origin restated with evidence taken from our own evolu­
tion. Variation, a struggle for existence, natural selection, fossils, em­
bryos, geography, comparative anatomy and instinct; all the elements
are there. The evidence that mankind evolved is impossible to resist. We
know so much about our own biology that the progress (if such it is) to­
ward the animal that civilized society has a lesson for evolution as a
whole. Its logic is that of The Origin itself, and can be presented in the
same format to do what Darwin could not: to make his case with
human examples.

Man has tamed many creatures, but has himself become the most do­
mestic animal of all . A fifth of the Earth is farmed, as proof of how the
world has been forced into the household. The passage was not easy. Al­
though cows gained from a move to the field, for one species farms re­
duced the quality of life. Before agriculture, the people of the Middle
East ate a hundred and fifty kinds of plants. Afterward, that fell to half a
dozen or so. Those who tilled the soil were smaller and less healthy than
their ancestors, with worse teeth and signs of anemia written in their
 Almost Like a Whale? 311

bones. Women's elbows were ruined by constant grinding and men paid
the price in social stress, with far more beaten to death or stabbed than
before.
The malign effects of farming are almost over. Since the Second World
War, Europeans have grown by half an inch a decade. One man in four
in Britain is six feet tall. In fifty years that is likely to be the average and
we will have regained the physique, if not the psyche, of the hunter­
gatherers. People who stayed as hunters or fishermen (Pacific islanders
included) find it hard to deal with the excesses of the Western diet and
suffer from diabetes as a result. Domestication put pressure on man's
genes, as it did on those of his plants and animals.
Cities did further damage to those who built them and provided nat­
ural selection with a new forum. In the Middle East, hunters lived in set­
tlements hundreds strong. Nine thousand years ago-before the great
Black Sea flood-came the first metropolis. Catal Hiiyiik in Turkey had
a population of ten thousand, crammed into tiny houses decorated with
images of volcanoes and of headless men attacked by vultures, with the
remains of ancestors buried beneath their floors. For the first time, in­
fection found a pool of people in which to sustain itself, and selection
sprang into action. For much of its history, even London could not
maintain its numbers except by movement from the countryside, so per­
vasive was disease. Illnesses such as cystic fibrosis today might be the rem­
nants of genes that, long ago, protected against the cholera which swept
through the new nurseries of pestilence.
However domestic man may have become, he is (unlike many of his
servants) still filled with diversity. Our variation under nature is such that
we have a great ability to tell ourselves apart. Francis Galton, Darwin's
cousin, set out to spot a typical English face-the John Bull of those
days-in Kensington Gardens. He failed because each person he saw was
so unalike. A distinctive face is remembered better than others, with
Prince Charles noticed sooner than Tony Blair. Much of the variation is
inherited, and to study faces in families shows that rather few genes are
involved.
Diversity is in faces, on cells and within the DNA. The sections that
code for useful proteins-the eye lens, or the framework of the cell­
vary not at all, while others, such as the short segments that make the
"DNA fingerprint," differ to such a degree that no two people who have
312 D A R W I N ' S G H O S T

ever lived (or ever will live) are alike. Molecular biology once had a Lin­
naean view of man: that to establish his nature it is enough to study a sin­
gle spe�imen (although the chart of human genes was drawn from a sort
of chimera in which lengths of DNA were lifted at random from differ­
ent people). Rather late in the day, evolution has been granted a say, with
a first step to a world map of variation. Human DNA has about three
million variable sites. A grid of a hundred thousand points has been set
up as a tool in the search for the parts of the genome that affect suscep­
tibility to disease.
Damaged genes are as universal in humans as in their dogs and may
be revealed in much the same manner. When people mate with relatives,
they bring forth unhealthy children. The children of cousins have a sur­
vival rate some 3 percent lower than average. They have been unlucky in
the biological lottery and inherit double copies of DNA that wins no
prizes. A simple sum based on the health of inbred children shows that
everyone must carry single copies of one or two genes that, if present in
double dose, would kill them. As is the case for domestic animals and
plants, inbreeding reveals a hidden universe of human variation.
Most of the diversity separates one person from the next. Some,
though, reflects differences among populations. The center of variety is
in Africa, with trends of decrease through Europe and Asia to the small
islands of the Pacific and the New World. Within Mrica live peoples as
distinct as the Zairean pygmies and the tall Nilotics. Europe, in contrast,
can do no better than Greeks and Swedes. Between people and between
places, Mrica is different. Two Mricans are, on the average, more dis­
similar than two Europeans, and two Mrican villages a few miles apart
can be as distinct as whole nations elsewhere. By comparison with that
continent, the rest of the world is monotonous indeed.
By comparison with other mammals, however, man himself is a te­
dious beast. No alien primate watcher would gain much from the tax­
onomy of humans. Chimps have two species-the common chimpanzee
and the bonobo-and gorillas are divided into mountain and lowland
kinds, but man and aardvark are unique among mammals as solitary
species, each within a single genus. Our relatives have evolved more than
we have because they have been around for much longer. Man is a new­
comer and has not had time to build up the diversity found in other pri­
mates. The most remarkable thing about humankind is how uniform it
 Almost Like a Whale? 313

is. Eight tenths of all our variation is among individuals within a popu­
lation, a tenth or so from the undoubted differences that exist among
populations on the same continent-the Greeks and the Irish, or the
Kenyans and Nigerians-and about the same small proportion from the
divergence between Europe, Asia, the Americas, Australia and Mrica.
The world is divided by politics, but it is united by genes; and our vari­
ation under nature is more confined than that of any comparable crea­
ture.

Man is also amazingly common. He has released himself from the strug­
gle for existence. The abundance of any creature is related to its size, with
fewer whales than mice. The fit is consistent, and almost all mammals
from mouse to whale sit on the same line (cars follow the same rule, with
more Toyotas than Cadillacs) . Civilization has removed the constraints
that limit all other animals and man is ten thousand times more plenti­
ful than expected from his size. In spite of many claims that resources are
near exhaustion (in 1 865 that Britain's coal was almost finished, in 1 972
that the world's oil would be gone within a decade) , most minerals are
still abundant and prices are falling. There have been apocalyptic fore­
casts of famine, which did not arrive; and food production continues to
go up as, most of the time, the crops get cheaper. Even so, with man al­
ready using almost half the total production of all plants, a limit must be
reached. On ecological grounds Americans are the most evolved of all ,
for each needs as much energy as a sperm whale. Their nation is already
so advanced that it would take two complete Planet Earths to feed the
world to the level of the United States.
If humans were on the curve that ties mice to elephants, the global
population would be around half a million (rather than six billion) and
Britain would contain the same number of people as it does seals. The
explosion began with agriculture and speeded up with industry. Al­
though the rate of increase has begun to slow, our numbers are almost
certain to double before they begin to fall. Today's respite from the
Malthusian struggle will not last.
There never has been a rest from natural selection. Genes are checked
at all times for their ability to cope; and many people fail the examina­
tion. In the Western world only one baby in a hundred dies from an in-
314 D A R W I N ' S G H O S T

herited disease and, given a survival rate of98 percent to the age of thirty,
it might seem that natural selection has little raw material left to play
with. However, genes were involved in far more deaths a century ago. For
most of history, half of all children died from cold, from starvation or
from infection. Although squalor may have been the direct cause, DNA
was busy behind the scenes because some babies failed and some suc­
ceeded by virtue of what they inherited.
Cold, hunger and disease are, of course, still hard at work. Nineteen
out of every twenty deaths of children are in the Third World, compared
to half the mortality at the age of seventy. Although smallpox vanished
twenty years ago, and polio should be extinct within a decade, not all the
news is good: there was no cholera in South America for a century, but
now it has a million cases a year. Many societies live on the edge. In West
Mrica, children born at the time of the rains, the "hungry season," are
ten times more liable to die in their teens than are others, as they never
recover from the experience. Because genes influence how well they sur­
vive, and because death in childhood counts for much more than one de­
layed, their demise is fuel for selection.
Many of the trends in human size, shape and appearance that cross the
world may reflect past natural selection; but-even for striking differ­
ences such as skin color-we are embarrassingly ignorant about what it
might have been. For other patterns (such as the absence of the B blood
group in the New World) we know even less. Sometimes there is a hint
of the truth. Some people are unable to deal with nature's poisons and
get cancer as a result. The genes involved are related to those that allow
insects to live on toxic plants. Many people are lacking in one of these
crucial segments of DNA. As a result, they find it hard to break down the
drugs used to treat heart conditions and a dose helpful to most is lethal
to them. Selection is, once again, at work, but for humans-as for other
creatures-whether it influences the mass of diversity, from fingerprints
to DNA, is as yet unknown.
The same is true of sexual selection. Darwin, in a letter twenty years
before The Origin, weighed the case for and against marriage: "Chil­
dren-(if it Please God)-Constant companion, (& friend in old age)
who will feel interested in one,-object to be played with. better than a
dog anyhow . . . but terrible loss of time.-cannot read in the
evenings-fatness & idleness-less money for books &c-if many chil­
dren forced to gain one's bread . . . banishment & degradation into in-
 Almost Like a Whale? 315

dolent, idle fool." In the end uxoriousness triumphed over literature and
he married his cousin (who had ten children to pass on the Darwinian
legacy) . In humans, too, selection works on sexual success as much as on
survival, with a balance of cost and benefit for each party to the repro­
ductive debate.
In elephant seals and flowers, males have more variation in the num­
ber of sexual encounters than do females, with plenty of chance for male
competition and female choice. Men prefer young, attractive and faith­
ful women, while women look for the same in their own partners, but a
decent bank balance helps. That revelation by the new science of socio­
biology has been claimed to result from sexual selection. Without evi­
dence of genetic variation in looks or wealth it is hard to know whether
the rules that govern elephant seals also apply to humans. Human be­
havior is certainly susceptible to a change in the balance of the sexes. In
China, because of selective abortion, a hundred and fourteen boys are
born to every hundred girls. This has led to a generation of frustrated
bachelors-and to an increase in the value of females, who are either
forced into marriage or attracted with a large bride price. In time, no
doubt, the attraction of sons will wane as they are forced to stay single.

The days of confusion about human inheritance have been succeeded by

what seems like clarity. It takes about a hundred thousand genes to make
a man (about the same number of parts needed to build a modest exec­
utive jet) and the Human Genome Project-the scheme to read the
order of the DNA bases-is almost complete, less than fifty years after
the discovery of the double helix. A single laboratory can now read off
millions of DNA bases a year and (in some at least) the information is
fed on to the Internet, to protect it from those who hope to patent genes.
Two people out of three in the Western world die for reasons connected
to the genes they carry (most of them because their inheritance is less
able to cope with the environment-tobacco, alcohol, salt, sugar or
stress-which they face). So important has genetics become in medicine,
and so far has it advanced, that to study humans is often the first (rather
than the last) resort in understanding the science. As a result, our own
inborn weaknesses illustrate the apparent difficulties of evolutionary the­
ory better than anything else.
Malaria kills two million people a year. Its agent is a single-celled par-
316 D A R W I N ' S G H O S T

asite passed on by mosquitoes. All the parties-man, mosquito and par­

asite-have responded to natural selection. Among the mosquitoes, in­
secticide resistance is everywhere. In the parasite, resistance to drugs
means that death rates from the illness have risen by ten times and that
in some places medicine is a single remedy from defeat. In spite of great
hopes for vaccines or new treatments in a continent in which some states
spend half their wealth on arms, most of Mrica cannot even afford to
control the insects.
Man, too, has done a lot of evolving in the face of malaria. How he
did it rebuts many of the supposed problems of Darwinism.
What once appeared a single illness is in fact four, caused by parasites
not much related to each other. Each does its damage as it invades the
red cell and spreads to liver and brain, with fevers, anemia and worse.
Some groups are highly susceptible. When the disease moved to the New
World, one tribe was reduced in numbers from fifteen thousand to sev­
enty within a century. Within Mrica, malaria's native continent, too,
some individuals-and some peoples-deal with it far better than do
others.
Evolution is to blame. Malaria is a great test of Darwinism. It shows
how that theory's apparent weaknesses are in fact its best support. An
ever-changing enemy is fought with whatever comes to hand, and dif­
ferent people achieve the same end through different means. Often, the
same gene arises in distant places and has tasks unrelated to disease.
Sometimes, resistance imposes a terrible burden on those involved; but
that matters not at all if the cost of the defenses is outweighed by the ben­
efits they bring.
The parasite spends much of its time inside red blood cells, where it
feeds on hemoglobin. A change in the structure of the molecule can put
paid to it. The genes involved protect those who inherit a single copy, but
they also cause the commonest inherited diseases in the world, because
in double dose they damage their carriers. One person in fifteen, world­
wide, has a copy of an altered hemoglobin, and tens of millions suffer as
they pay the price for the advantage that such genes give to their kin.
The symptoms of sickle-cell anemia were described in 1 670. They
come from a single change in the DNA, a shift in a certain base that al­
ters the stability of the oxygen-carrying molecule. In those with two
copies of the gene, the red cells collapse into a crescent shape when oxy-
 Almost Like a Whale? 317

gen is short. As the cells buckle, the parasite may die; but the sickled cells
clog tiny blood vessels and lead to pain, kidney failure, heart problems
and more. The gene maintains itself because children who inherit just
one copy resist malaria and are otherwise well. Their unhealthy rela­
tives-those condemned by the laws of Mendel to inherit two damaged
versions of the hemoglobin gene-pay the price for the survival of their
more plentiful relatives. The spread of the sickle-cell gene depends on a
balance of kinship, cost and benefit as relentless as that of the beehive,
and involves as little conscious effort.
Millions of Mricans trace their descent from the single individual
within whom the mutation happened long ago. Sickle-cell is also found
in India, but there the DNA around the crucial site reveals an origin not
in Mrica but in an ancient Asian. For the sickle-cell gene (as for the an­
tifreeze of the Arctic and Antarctic fish), an identical present conceals an
independent past.
Hemoglobin has hundreds of other changes that, in their several ways,
help defeat the parasite. Some involve single shifts in its building blocks,
while others cut out great lengths of the protein chain. Changes in quite
separate genes alter the red cell's economy to make things hard for the in­
vader. Some render cells more sensitive to wastes so that infected cells,
and their unwelcome visitor, die. The identity molecules on white blood
cells also protect against malaria's worst effects as some versions allow the
immune system to destroy the parasite before it enters the brain. Certain
peoples resist infection for reasons quite unknown. In the Sudan, the Fu­
lani live mixed among the Rimaibe (who used to be their slaves). Every­
one is bitten at least once a night-but most Fulani stay clear of the
disease, while almost all their neighbors suffer from it.
The Duffy blood group has two forms. The agent of malaria uses one
variant as a docking site. The other, common in West Mrica, gives pro­
tection against the disease (a fact discovered when blacks were infected
with malaria in an attempt to treat syphilis but failed to contract it) . The
Mrican version removes the whole structure and leaves the parasite baf­
fled. The Duffy receptor remains on all other tissues, but on the red cells
it is switched off. The protein also serves another purpose: in the brain,
where it helps to direct cells to where they should be. In West Mricans it
has become rudimentary or utterly aborted for its docking job, but re­
mains perfectly efficient for the other.
318 D A R W I N ' S G H O S T

Such diversified adaptations in the same species are just part of the war
against infection. Some antimalaria genes were first recognized as dis­
eases in their own right; as organs of small importance, indeed. People of
Mrican origin are susceptible to high blood pressure because they are bad
at clearing salt from the body. Salty blood is, it transpires, a defense
against the parasite. Many suffer from a high (and often fatal) level of
iron in the liver. Iron, too, kills the agent of malaria, and the liver disease
is a side-effect. All this is evidence of the many unknown laws of corre­
lation of growth, which, when one part of the organization is modified
and the modifications are accumulated by natural selection, will cause
other changes, often of the most unexpected nature.
In a few places-East Anglia, the United States, around the Mediter­
ranean-the parasite has been driven out. The protective genes are still
there, and will long remain (as they will, perhaps, after the disease has
gone altogether from the world) . They will be seen as relict characters,
evidence of a battle against a defeated enemy. Some, like sickle-cell, will
show themselves as apparent flaws that have slipped through the net of
natural selection. Because the history of the illness is known, such genes
can, instead, be recognized as evidence of selection's efficiency and its un­
canny ability to offset a benefit against what it costs.

Society is much the same. Human behavior is, without doubt, crafted by
evolution, and we have inborn instincts as strong as those of any animal.
Our species has long been seen as an ape remade by thought: to one ex­
pert, "The ancestors of the Gorilla and the Chimpanzee gave up the
struggle for mental supremacy because they were satisfied with their cir­
cumstances." Humans, it appears, were not. The brain makes us human.
It has doubled in size in the past two million years. Its progress was not
smooth, as some of our forerunners had brains larger than our own (al­
beit on a heftier frame). Bodies became smaller, but the brain did not fol­
low. Our intellect might result not, as we flatter ourselves, from the
benefits of a great mind, but a small body.
Most of what makes primates what they are resides within the skull.
Our relatives are pretty smart: but why? Perhaps it is because the fruit
trees favored by many are patchily distributed and a mental map is
needed to remember where they are. Social life, too, needs gray matter to
 Almost Like a Whale? 319

tell who is who and how to treat the neighbors. Comparative anatomy
hints at the past. The bigger the group, the more complex the society.
The size of the brain fits that of the community, with a relationship
much better than anything to do with what a particular species eats. So­
ciety, not shopping, swelled our heads.
Their contents are expensive. A newborn baby uses more than half its
energies on what is in its skull. The human brain is two pounds heavier
than that of a similar-sized mammal-a hefty pig, say; bur we do not use
much more energy than pigs. Men have smaller guts than other primates
and have, unlike chickens, invested in brains over bellies. The brain's
costs come early on, as the organ reaches almost adult size by the time a
child goes to school. As Winston Churchill said, "There is no finer in­
vestment for any community than putting milk into babies. " A giant in­
tellect explains why so much is needed.
Far more genes are active there than in any other tissue, and they can
mutate and evolve as much as others. Many mental disorders have a sim­
ple genetic basis. Genes also affect personality, with claims of inborn
variation in how sociable or misanthropic people might be and in their
ability to deal with words or with tools. When it comes to intelligence,
schools mold what nature provides (which is why Eton stays in business);
but without some foundation in reason, the most expensive education is
wasted. Variation in a gene controlling the growth of cells (brain cells
included) plays a small part in differences in intelligence. Other such
variants will, no doubt, be found. Whatever might emerge, man's be­
havior-like that of the Cocos finch-will always be defined more by
habit than by instinct; by what he learns rather than what he is.

Behavior apart, man is-of course-not much more than just another
primate. No doubt, the laws of hybridism that keep him separate can be
blamed to some degree on intellect. There are rumors of test-tube crosses
between men and chimps. One such creature made it to the front pages,
although on close examination he turned out to be just an unusually ra­
tional ape. A cross between the two primates might at least be possible.
Humans and chimps share 98.8 percent of their DNA, humans and go­
rillas rather less. Their close kinship is proved-as in whales and hip­
pos-by their fellow-travelers. All three carry the same set of viral
320 D A R W I N ' S G H O S T

hitchhikers in the same place in their genes, while those of orangs, gib­
bons and baboons are more and more different. In chromosomes, too,
we are similar. The main difference between the great apes and ourselves
is that two chimpanzee chromosomes are fused and three more are
reshuffled (which means that any hybrid would be sterile).
Our affinity to chimps should be seen in context. It leaves thousands
of genes unaccounted for; and, in any case, the diversity of mammals as
a whole hides genetic conservatism, with a mere thirty or so large sec­
tions of DNA reordered when humans are compared with cats. One cell
surface molecule has changed in man alone. It acts as a docking site for
diseases such as malaria (to which humans are uniquely susceptible) and
as a means of communication between cells-those in the brain in­
cluded. Perhaps it played a part in man's struggle for mental supremacy
and helped keep him distinct from his kin.
Man may differ little in his anatomy from other primates, but his past
has a certain interest if only on grounds of familiarity. Darwin was cau­
tious in The Origin, but in The Descent ofMan, published twelve years
later, could afford to be direct: "Man is descended from a hairy
quadruped, furnished with a tail and pointed ears, probably arboreal in
its habits, and an inhabitant of the Old World." To console those in­
sulted by such a notion, he noted that if the human pedigree was no
longer "of noble quality," at least it was "of prodigious length."
Now, we have fossils as proof of his claim. The human record is, like
all others, incomplete, although it has become much less so in the past
half-century. Dozens of Neanderthals have been found. Another almost
complete set of bones comes from the famous "Lucy," who lived more
than three million years ago. These are exceptions. Primate remains are
as liable to upheaval as are those of less noble beings. Hundreds of fossil
teeth, and not much more, are found along the Lower Omo River in
Ethiopia, because teeth are tough and last when other relics have gone.
Most human fossils are small fragments of bone, upon each of which
great mountains of theory have been built.
Lucy, for example, was smaller than her fellows. Perhaps, some sug­
gest, males were then much larger than females and life was based on sex­
ual battles. However, it now seems that there were two different-sized
species around at the time, so that her supposedly unequal society disap­
peared as soon as more fossils were found. Other finds have also caused
 Almost Like a Whale? 32 1

much speculation. On one set of remains was built a theory of man as a

killer ape (in the language of science, an "osteodontokeratic culture") be­
cause the skulls were damaged and bones scored as if by knives. In fact,
the cracks were the accidents of time, and the cuts are grooves made as
bones tumbled down a riverbed.
Science needs theories, but-with its shortage of facts-human evo­
lution has rather more than it can cope with. Take the art of walking.
Lucy and her predecessors could stand erect (although the organ of bal­
ance in her inner ear is more like that of apes than humans, so that per­
haps they did not do so very well) . It is still a delicately poised talent, as
anyone who overdoes the Chardonnay finds out. But why did we change
from horizontal animals who, now and again, took to the vertical, to an
upright beast who must struggle against gravity with a body that evolved
at ninety degrees to where it usually finds itself?
In the days when man was seen as striving to reach a manifest destiny,
the answer was obvious: our ancestors stood up to free their hands for
tasks more important than walking. Of course, the idea is foolish: it is
like saying that the brain evolved in order to watch television. Evolution
did not plan ahead to the days of The Jerry Springer Show. Man stood up,
as he did everything else, for pragmatic and short-term reasons. But what
could they have been?
Perhaps a vertical animal found it easier to hunt, or to follow herds of
grazers in the hope of plunder. Perhaps our ancestor was himself hunted
and, by getting on his hind legs, could see danger in good time. There
may have been a change of diet toward fruit high on trees or, as the cli­
mate dried, a change in habits as food became more scattered and had to
be carried back to a distant camp. Being erect is a sexual display, and a
male who could keep upright for longer might do better with the fe­
males. Those who lie down on a hot day expose more to the sun (which
is why sunbathers rarely do so standing up) and, in the tropics at least,
pay the price. All these ideas are reasonable but, given that the event hap­
pened five million years ago, are hard to test. With so little from the past,
anthropology is one of the few sciences in which it is possible to be fa­
mous for having an opinion, and until more facts emerge such specula­
tion is bound to remain.
Incomplete as it is, and overinterpreted as it may be, the record of the
past is forceful evidence of the reality of human evolution. We descend,
322 D A R W I N ' S G H O S T

with all other mammals, from a rat-sized creature of a hundred and sixty
million years ago whose descendants lived modest lives around the feet
of the dinosaurs until those giants were wiped out. The first fossil pri­
mates are found soon after that event in the warm and wet Mrica of sixty­
five million years ago. Some six thousand kinds have lived since then
(and two hundred or so remain today, from the quarter-pound mouse
lemur to the gorilla, a thousand times heavier). Once, the world had
many more species of apes than of monkeys, but now just five great apes
are left (one of which is us) while monkeys flourish; proof that there was
no inevitable progress toward mankind.
About fifteen million years after the emergence of the first primate, the
predecessors of apes appear. A molecular clock based on inserted viruses
suggests that the lines to chimp, to gorilla and to humans split some five
million years ago. A dozen or so hominines-as the branch upon which
we belong is called-have lived since then (although how many are real
entities is hard to assess) . Over that period, there appeared Ardipithecus
(four and a half million years old and of intermediate form); several
kinds ofAustralopithecus (an Mrican primate from around a million years
later which was, roughly speaking, a human below the neck but an ape
above); and Homo habilis (an animal defined as having crossed the cere­
bral Rubicon of brain size needed to qualify for our own family) . It may
have been the first tool-user nearly two and a half million years ago. It
was followed by Homo erectus (a large-brained ape that looked rather like
a man). Each seems to have emerged first in Mrica and most spread, at
one time or another, to Asia and to Europe.
The first members of our own species, Homo sapiens, arose about a
hundred and fifty thousand years ago as large, thick-skulled, but recog­
nizably human apes. By the time the miseries of the ice ages were over we
were, more or less, ourselves, with smaller brains in a thin skull on a slim
and elegant body. Why we shrank is not certain. Perhaps, as in the dwarf
mammoths on islands, shortage of food did the job, or perhaps a shift to
a kinder society cut down the need for a sturdy frame.
It once seemed natural to arrange the fossils in a sequence that clam­
bered up an evolutionary tree in single file to man. Every find had a name
and a place in the hierarchy of the almost-human. Two species, almost
by definition, could not live together, as the cultural superior would at
once drive out its brutish ancestor.
 Almost Like a Whale? 323

As more bones turn up, the story becomes less clear. For much of the
time, two or more kinds lived together in an uneasy coexistence rather
like that of men and chimps today. In East Mrica, for instance, two
species of large-brained Homo lived alongside smaller-brained Australo­
pithecines of several types, with perhaps half a dozen forms present at
once. Some enigmatic fossils, such as the "Black Skull" found at Lake
Turkana, combine primitive and advanced features and suggest that pat­
terns of change were complex indeed.
In spite of a century's claims of the discovery of "missing links," it is
quite possible that no bone yet found is on the direct genetic line to our­
selves. With so many kinds to choose from, so few remains of each, and
such havoc among their relics, none of the fossils may have direct de­
scendants today. The proportion of the people alive even a hundred
thousand years ago who contributed to modern pedigrees is small, and
the chance that any surviving relic, one among lost billions, belonged to
that elite is quite minute.
Neanderthals are the most familiar of fossils. Once dismissed as the re­
mains of diseased Napoleonic soldiers, and then hailed as our immediate
predecessors, their true history is one of a dead end on the road of human
evolution. With their stocky bodies, they were adapted to cold; and they
evolved in Europe or the Middle East, their home until they disappeared
thirty thousand years ago. They used stone tools, but in an uninspired
way, and stayed apart from their intellectual neighbors.
Although Neanderthals, at first sight so similar to modern humans,
were once placed on the last rung before mankind, fossil DNA hints that
they may not even be on the same ladder. Their mitochondrial genes are
quite distinct from our own. They were not the ancestors of human
genes but followed a separate path. For mitochondria, at least, Nean­
derthals and ourselves split half a million years ago.
Today's molecules also hint that our immediate ancestors have gone
forever. The longer the history of any population, the more variation it
contains, because there has been more time for mutations to build up.
The more abundant it is, the less the chance of an accidental loss of the
new variants as they arise. Any large and ancient group of animals hence
contains lots of different inherited forms, many of which-given time­
become common. A new or sparse population, in contrast, has little di­
versity and the altered forms are rare.
324 D A R W I N ' S G H O S T

The story of genes is rather like that of surnames. The probable date
of the shared ancestor of everyone with a certain surname depends on the
size and age of the population. Half a dozen Sidebothams in a tiny village
are almost certain to stem from the same recent ancestor. Six carriers of
that noble name chosen at random from among the many Sidebothams
of London will have to go much further back to find the ur-Sidebotham
from whom they descend. A city in which most people have the same
name is likely to have expanded from a small village, while one with
thousands of surnames traces its origin to a huge and variable ancient
populace. Genes, like names, can be used to make guesses about the past.
They show that, compared to other primates, humans are not very di­
verse and most variants are rare. Chimpanzees are three times more dis­
tinct one from the other than are men, with fifty times as much
divergence among separate populations. The logic of the genes shows
that chimps, now only a couple of hundred thousand strong, were once
common and that the human race, abundant as it is today, was scarce. Its
average size over most of its history may have been a mere ten to twenty
thousand people. Our immediate ancestors were a small band who oc­
cupied a few hundred square miles and, more than likely, left no fossils
at all. To draw the human family tree reveals an explosion of change, a
starburst like that of AIDS viruses, with its center a hundred thousand
years ago. It may mark the expansion and spread of modern Homo sapi­
ens, but the chances of finding where we came from are small indeed.
Wherever the Garden of Eden may have been, man became a traveler
as soon as he escaped it. Although a shortage of bones makes it hard to
be certain, there were several journeys out of (and perhaps even back to)
Mrica.
The great arena of evolution was in Asia, Europe and Mrica. The hab­
itable world was not filled until a thousand years ago, with the settlement
of New Zealand. Many barriers stood in the way. Some, narrow though
they are, proved hard to cross. The New World, most agree, was not
reached until about fourteen thousand years ago, across the Bering Land
Bridge between what is now Siberia and Alaska. Within a thousand years
people had reached southern Chile (where hints have been found of an
occupation twice as old) . By contrast, humans have been in Australia for
fifty thousand years, and crossed the Sunda Strait-recognized by Wal­
lace as a break between the continents-to do so.
The simplest of all barriers is distance. Although that has been de-
 Almost Like a Whale? 32 5

feated by the wheel, it is easy to forget how isolated all of us once were.
Numbers shot up after farming began, but even then, most people stayed
at home. Although the conventional view of history is of rape and pil­
lage-men, from Attila the Hun onward, forcing their genes on to
women-DNA reveals another past. The genes that pass through fe­
males, on the mitochondria, are less localized than those of the male
chromosome, the Y. Women, it seems, have traveled more than men
(perhaps because they move to find a husband in the next village) .
Like Red Sea fish in the Mediterranean, men destroy as they move on.
Alfred Russel Wallace noted that "we live in a zoologically impoverished
world, from which all the hugest, and fiercest, and strangest forms have
recently disappeared." The culprit is plain. Humans reached Australia
fifty thousand years ago. They came across tortoises as big as a Volkswa­
gen Beetle, carnivorous kangaroos, and flightless birds twice the size of
an emu. Within ten thousand years, all were gone. The first Americans
were even more efficient. Throwing sticks, atlatls, increased the leverage
on a spear and gave it the power of a Magnum rifle. Five hundred years
after the hunters reached the Great Plains, mammoths, camels and
horses were extinct or almost so and the survivors-wild sheep and
bison-were much reduced in size. Their demise is a reminder that, for
most species, extinction at the hands of a successor is inevitable.
Travel as he might, man is a lowland animal. Eleven of the world's fif­
teen largest cities are on the coast. Mountain-and northern-peoples
have been pushed around by ice ages as much as have dung beetles. In
China, the tribes of the hills are distinct while the masses of the coastal
plains are more uniform. Even in Italy, hill villages are, because of their
isolation, more different from one another in their genes than are the
cities of the plain.
DNA reconstructs the history of human migration. The trends across
the world reflect bonds of shared descent, as modified by natural selec­
tion. With few exceptions, people from northern places are taller and
broader than those from the tropics. What counts is the relation between
the mass of the body and its surface area. To stay warm it pays to be
spherical. Eskimos (and Neanderthals) have thick bodies with short
limbs, while most Mricans are slimmer with longer arms and legs. Pat­
terns of body shape are consistent to north and south, and those in skin
color (little though we understand them) have parallels in the New
World and the Old.
326 D A R W I N ' S G H O S T

On islands, too, people have changed, more by the accidents of mi­

gration than by natural selection, as most islands have not been occupied
for long. All over the world, isolated by water, by mountains or by big­
otry, they have evolved. By chance, certain genes that are rare at home
took a trip and at once became relatively common. The inhabitants of
several Pacific islands suffer ftom inherited blindness, and religious iso­
lates such as the Amish of North America also have much genetic disease.

Human evolution can, of course, be studied in the same objective way as

that of any animal. Genetics, geology, and geography illuminate our
past. Morphology--comparative anatomy-is the most powerful tool of
all. Queen Victoria noticed as much on her first visit to London Zoo, in
1 842, and was not amused: "The Orang Outang is too wonderful . . .
He is frightful and painfully and disagreeably human." She was right.
Anti-AIDS medicines are tested on chimps because their bodies are so
like ours. The cladistic rules put humans into the same family as chimps
and gorillas, as all derive from a common ancestor not shared by any­
thing else. The laws that put birds with crocodiles cannot be broken sim­
ply to satisfy our wish to be in a class of our own.
Man is a primate, and in some ways not a very special one. He can do
more than any other creature, but has not changed much to do so. The
strangest thing about human evolution is how little there has been.
Nothing else is so widespread and nobody fills so many gaps in the econ­
omy of nature. Many animals carry out tasks almost as wonderful as
those achieved by ourselves, but through biology rather than intellect.
For them, success at one task means failure at all others. In the past hun­
dred thousand-in the past hundred-years, human lives have been
transformed, but bodies have not. We did not evolve, because our ma­
chines did it for us .& Darwin put it in The Descent ofMan: "The high­
.

est possible stage in moral culture is when we recognize that we ought to

control our thoughts." Human progress has made a simple but crucial
move, from body to mind. That mind is built from genes but what it can
do has long transcended DNA. Human are-of course--constrained by
biological history, as pigs are limited in the ability to fly by their ances­
tors' lack of wings. We are all branches on a common tree and share de­
scent with primates and, for that matter, with pigs. Biology tells us that
 Almost Like a Whale? 327

we evolved, but when it comes to what makes us human is largely beside

the point. There might be inborn drives for rape or for greed, but Homo
sapiens, unique among animals, need not defer to them. To say that the
genes are in control of our behavior has no more intellectual content
than to claim that people go to work because their legs drag them to the
office. This has not stopped those unable to explain society by other
means from pressing evolution into service.
Many sociologists (and a few biologists) hope for a comparative
anatomy of the mind; but that can never succeed. When it comes to what
makes us different from other creatures, science can answer all the ques­
tions except the interesting ones. The human intellect stands alone. As
there is nothing else like it, the rules of classification come into play. If
an object is one of a kind, it is impossible to know where to put it. The
problem with the mind, or any uniquely human attribute, is simple: it
is, like the narwhal's tusk, unique.
To understand how far a body part has come, we must know where it
started from. That can be done for cell membranes, for gills, breasts, kid­
neys or opposable thumbs, because some animals have them and some
do not. Their pattern-present in groups who descend from a shared an­
cestor and not in others-is a map through the past. It is the problem of
language: if there were but one world tongue (as, with the spread of En­
glish, there may one day be) it would be impossible without further evi­
dence to tell from whence it came. Shared derived characters-words
common to Welsh, English and French, or even to Chinese-are needed
to untangle the hierarchy of evolution. The conscious mind is different:
we all have one, but, as far as we can tell, nothing else does. As a result,
to speculate about its evolution is largely futile.

This book began with mankind's latest challenge, an attack by a virus

that came from an ape. We have evolved in response to AIDS through
the most familiar kind of natural selection, but the disease also shows
how humans are different; how we transcend the biological rules that
apply everywhere else.
Some HIV-positive people survive for decades, for good evolutionary
reasons. Their luck results from inherited diversity; not in viruses, but in
men. The survivors tend to carry certain forms of the receptor lock into
328 D A R W I N ' S G H O S T

which the HN key fits so that the virus finds it hard to break in to their
cells. Access is denied in many ways, with a whole series of bolts and bars,
each of which comes in several versions. One person in six inherits an al­
tered form of part of the defense mechanism. For them, the onset of
symptoms is delayed for years. About one in twenty is lucky enough to
have two copies of this altered gene. This shuts out the virus altogether.
A variant of a second receptor is present in about a quarter of all Asians,
and rather fewer Mricans and Europeans. Although its bearers may pick
up the virus, they last much better than average. If-as it will-the dis­
ease continues to kill, only those with a heritage that allows them to sur­
vive for long enough to have children will succeed. Their genes will
spread and perhaps, in time, man (like his primate relatives) might evolve
resistance to AIDS.
There has, however, been another, uniquely human, response to the
illness that far surpasses what evolution can do. Minds have changed
much more than genes. For the first few years after the outbreak, the gay
community was in denial. AIDS tests were criticized as a step toward
concentration camps and the New York Health Commissioner banned
them. However, as news of the disaster spread homosexuals changed
their habits in the quickest response to a health threat ever seen. Bath­
houses closed, the numbers of partners dropped, and unprotected anal
sex went out of fashion. The rate of transmission among gay men plum­
meted by nine tenths and, in the developed world, continues to do so. In
Europe the number of new cases went down by half between 1 993 and
1 998. In parts of Mrica, too, there has been a tenfold rise in condom use
within a decade and a matching fall in the number of infections. A
change in behavior means that, in time, evolution will be beside the
point.
That is true for most of what makes us human. The many attempts to
explain the oddities of human life-sex life included-in terms of that
of bees or chimpanzees show how easy it is to fall unthinking into the ca­
pacious arms of Darwin. Many universals link the human race. Most,
like our need for vitamin C or for company, descend in modified form
from our ancestors. There are also differences among people and among
societies that seem to cry out for an evolutionary explanation. The task
is impossible, because-like the mind itself--there is no outgroup, noth­
ing apart from ourselves with which to compare them.
 Almost Like a Whale? 329

That elementary fact is often forgotten. For some, to explain any pat­
tern of society all that is needed is to stir in a Darwinian nostrum. If the
anthropological souffie fails to rise-reach for another bottle . .As in the
kitchen, the ingredients can be varied to taste. Mix them with enough
enthusiasm and, with a single bound, life is explained. Its infinite vari­
eties are justified with adaptive stories to fit.
In fact, most are an incidental to the healing power of lust, greed or
political expedience. Sexual fidelity or promiscuity seems, according to
taste, the natural thing to do; families are often a joy, and patriotism
arouses some, but we do not need natural selection to tell us why. So
much light has now been thrown on the origin of man and his history
that the limits of what biology can say about his present condition are
starkly exposed.
Such vulgar Darwinism is of interest only in its diversity. Evolution is
a political sofa that molds itself to the buttocks of the last to sit upon it.
Alfred Russel Wallace, in old age, turned to socialism. He saw biology's
real message: ''All shall contribute their share either of physical or men­
tal labour, and . . . shall obtain the full and equal reward for their work.
The future progress of the race will be rendered certain . . . by a special
form of selection which will then come into play." Marxists in their thou­
sands agreed. In China they named their children "Natural Selection" or
"Struggle for Existence" in homage to the Law of Nature that was to
transform society.
Marx's monument in Highgate is opposite the grave of Herbert
Spencer, who coined that unfortunate phrase "the survival of the fittest"
and the notion of Social Darwinism, an idea much used to explain the
excesses of capitalism. One devotee saw millionaires as "naturally selected
in the crucible of competition." The steel magnate Andrew Carnegie
agreed. "Before Spencer," he said, "all for me had been darkness, after
him, all had become light-and right." Sidney Webb, an important fig­
ure in the early history of the British Labour party, spoke in overtly Dar­
winian terms of "the inevitability of gradualism."
Spencer and Wallace-and even Webb-at least knew their biology.
The acidulous English literary critic F. R. Leavis once derided what he
called the culture of the Sunday papers: a civilization based on color sup­
plements. The new culture of the science pages uses a nodding acquain­
tance with evolution to promote an ethical agenda. For some social
330 D A R W I N ' S G H O S T

theorists, science is no more than fuel for parable, its evidence selected in
neat accord with political bias. As for Carnegie, biology makes every­
thing light-and right. It is the universal excuse.
Evolution deserves better than that; and this book has, I hope, shown
as much. It has not been a simple homage to Darwin, much as he de­
serves it. He had a wonderful phrase to describe those unaware of his the­
ory: they "look at an organic being as a savage looks at a ship, as at
something wholly beyond his comprehension." Nowadays, nobody can
use the defense of ignorance when faced with the need to understand
life's diversity: evolution forces all of us to look at the world in a new way.
Some of Darwin's ideas survive while others have been proved wrong.
The Origin ofSpecies endures as a work of art as much as of science. Its
message remains. Man, the highest of animals, and the most exalted ob­
ject which we are capable of conceiving, emerged from the war of nature,
from famine and death, as much as did all others. Humans, alone, have
gone further. As a result, much of what makes us what we are does not
need a Darwinian explanation. The birth of Adam, whether real or
metaphorical, marked the insertion into an animal body of a post­
biological soul that leaves no fossils and needs no genes. To use the past
to excuse the present is to embrace Payne Knight's pathetic fallacy, that
society can be explained in terms of the animal world. However, the new
insight that biology gives into our history releases us from the narcissism
of a creature that is one of a kind. It shows that humans are part of cre­
ation, because we evolved.
 C H A P T E R X I V

RECAPITULATION AN D CONCLUS I O N

Recapitulation of the difficulties on the theory of Natural Selection - Reca­

pitulation of the general and special circumstances in its favour - Causes
of the general belief in the immutability of species - How far the theory
of natural selection may be extended - Effects of its adoption on the
study of natural history - Concluding remarks.

As this whole volume is one long argument, it may be convenient to the reader
to have the leading facts and inferences briefly recapitulated.
That many and grave objections may be advanced against the theory of de­
scent with modification through natural selection, I do not deny. I have en­
deavoured to give to them their full force. Nothing at first can appear more
difficult to believe than that the more complex organs and instincts should
have been perfected not by means superior to, though analogous with, human
reason, but by the accumulation of innumerable slight variations, each good
for the individual possessor. Nevertheless, this difficulty, though appearing to
our imagination insuperably great, cannot be considered real if we admit the
following propositions, namely,-that gradations in the perfection of any organ
or instinct, which we may consider, either do now exist or could have existed,
each good of its kind,-that all organs and instincts are, in ever so slight a de­
gree, variable,-and, lastly, that there is a struggle for existence leading to the
preservation of each profitable deviation of structure or instinct. The truth of
these propositions cannot, I think, be disputed.
It is, no doubt, extremely difficult even to conjecture by what gradations
many structures have been perfected, more especially amongst broken and
failing groups of organic beings; but we see so many strange gradations in na-
3 32 D A RW I N ' S G H O S T

ture, as is proclaimed by the canon, "Natura non facit saltum," that we ought
to be extremely cautious in saying that any organ or instinct, or any whole
being, could not have arrived at its present state by many graduated steps.
There are, it must be admitted, cases of special difficulty on the theory of nat­
ural selection; and one of the most curious of these is the existence of two or
three defined castes of workers or sterile females in the same community of
ants but I have attempted to show how this difficulty can be mastered. With re­
spect to the almost universal sterility of species when first crossed, which
forms so remarkable a contrast with the almost universal fertility of varieties
when crossed, I must refer the reader to the recapitulation of the facts given at
the end of the eighth chapter, which seem to me conclusively to show that this
sterility is no more a special endowment than is the incapacity of two trees to
be grafted together; but that it is incidental on constitutional differences in the
reproductive systems of the intercrossed species. We see the truth of this con­
clusion in the vast difference in the result, when the same two species are
crossed reciprocally; that is, when one species is first used as the father and
then as the mother.
The fertility of varieties when intercrossed and of their mongrel offspring
cannot be considered as universal; nor is their very general fertility surprising
when we remember that it is not likely that either their constitutions or their
reproductive systems should have been profoundly modified. Moreover, most
of the varieties which have been experimentised on have been produced under
domestication; and as domestication apparently tends to eliminate sterility, we
ought not to expect it also to produce sterility.
The sterility of hybrids is a very different case from that of first crosses, for
their reproductive organs are more or less functionally impotent; whereas in
first crosses the organs on both sides are in a perfect condition. As we contin­
ually see that organisms of all kinds are rendered in some degree sterile from
their constitutions having been disturbed by slightly different and new condi­
tions of life, we need not feel surprise at hybrids being in some degree sterile,
for their constitutions can hardly fail to have been disturbed from being com­
pounded of two distinct organisations. This parallelism is supported by an­
other parallel, but directly opposite, class of facts; namely, that the vigour and
fertility of all organic beings are increased by slight changes in their condi­
tions of life, and that the offspring of slightly modified forms or varieties ac­
quire from being crossed increased vigour and fertility. So that, on the one
hand, considerable changes in the conditions of life and crosses between
greatly modified forms, lessen fertility; and on the other hand, lesser changes
in the conditions of life and crosses between less modified forms, increase fer­
tility.
 Recapitulation and Conclusion 333

Turning to geographical distribution, the difficulties encountered o n the

theory of descent with modification are grave enough. All the individuals of the
same species, and all the species of the same genus, or even higher group,
must have descended from common parents; and therefore, in however distant
and isolated parts of the world they are now found, they must in the course of
successive generations have passed from some one part to the others. We are
often wholly unable even to conjecture how this could have been effected. Yet,
as we have reason to believe that some species have retained the same specific
form for very long periods, enormously long as measured by years, too much
stress ought not to be laid on the occasional wide diffusion of the same species;
for during very long periods of time there will always be a good chance for wide
migration by many means. A broken or interrupted range may often be ac­
counted for by the extinction of the species in the intermediate regions. It can­
not be denied that we are as yet very ignorant of the full extent of the various
climatal and geographical changes which have affected the earth during mod­
em periods; and such changes will obviously have greatly facilitated migra­
tion. As an example, I have attempted to show how potent has been the
influence of the Glacial period on the distribution both of the same and of rep­
resentative species throughout the world. We are as yet profoundly ignorant of
the many occasional means of transport. With respect to distinct species of the
same genus inhabiting very distant and isolated regions, as the process of
modification has necessarily been slow, all the means of migration will have
been possible during a very long period; and consequently the difficulty of the
wide diffusion of species of the same genus is in some degree lessened.
As on the theory of natural selection an interminable number of intermedi­
ate forms must have existed, linking together all the species in each group by
gradations as fine as our present varieties, it may be asked, Why do we not see
these linking forms all around us? Why are not all organic beings blended to­
gether in an inextricable chaos? With respect to existing forms, we should re­
member that we have no right to expect (excepting in rare cases) to discover
directly connecting links between them, but only between each and some ex­
tinct and supplanted form. Even on a wide area, which has during a long pe­
riod remained continuous, and of which the climate and other conditions of life
change insensibly in going from a district occupied by one species into another
district occupied by a closely allied species, we have no just right to expect
often to find intermediate varieties in the intermediate zone. For we have rea­
son to believe that only a few species are undergoing change at any one period;
and all changes are slowly effected. I have also shown that the intermediate
varieties which will at first probably exist in the intermediate zones, will be li­
able to be supplanted by the allied forms on either hand; and the latter, from
334 D A R W I N ' S G H O S T

existing in greater numbers, will generally be modified and improved at a

quicker rate than the intermediate varieties, which exist in lesser numbers; so
that the intermediate varieties will, in the long run, be supplanted and exter­
minated.
On this doctrine of the extermination of an infinitude of connecting links,
between the living and extinct inhabitants of the world, and at each successive
period between the extinct and still older species, why is not every geological
formation charged with such links? Why does not every collection of fossil re­
mains afford plain evidence of the gradation and mutation of the forms of life?
We meet with no such evidence, and this is the most obvious and forcible of
the many objections which may be urged against my theory. Why, again, do
whole groups of allied species appear, though certainly they often falsely ap­
pear, to have come in suddenly on the several geological stages? Why do we
not find great piles of strata beneath the Silurian system, stored with the re­
mains of the progenitors of the Silurian groups of fossils? For certainly on my
theory such strata must somewhere have been deposited at these ancient and
utterly unknown epochs in the world's history.
I can answer these questions and grave objections only on the supposition
that the geological record is far more imperfect than most geologists believe.
It cannot be obj ected that there has not been time sufficient for any amount of
organic change; for the lapse of time has been so great as to be utterly inap­
preciable by the human intellect. The number of specimens in all our muse­
ums is absolutely as nothing compared with the countless generations of
countless species which certainly have existed. We should not be able to
recognise a species as the parent of any one or more species if we were to ex­
amine them ever so closely, unless we likewise possessed many of the inter­
mediate links between their past or parent and present states; and these many
links we could hardly ever expect to discover, owing to the imperfection of the
geological record. Numerous existing doubtful forms could be named which
are probably varieties; but who will pretend that in future ages so many fossil
links will be discovered, that naturalists will be able to decide, on the common
view, whether or not these doubtful forms are varieties? As long as most of the
links between any two species are unknown, if any one link or intermediate va­
riety be discovered, it will simply be classed as another and distinct species.
Only a small portion of the world has been geologically explored. Only organic
beings of certain classes can be preserved in a fossil condition, at least in any
great number. Widely ranging species vary most, and varieties are often at first
local,-both causes rendering the discovery of intermediate links less likely.
Local varieties will not spread into other and distant regions until they are con­
siderably modified and improved; and when they do spread, if discovered in a
 Recapitulation and Conclusion 335

geological fonnation, they will appear as if suddenly created there, and will be
simply classed as new species. Most fonnations have been intennittent in their
accumulation; and their duration, I am inclined to believe, has been shorter
than the average duration of specific forms. Successive fonnations are sepa­
rated from each other by enonnous blank intervals of time; for fossiliferous for­
mations, thick enough to resist future degradation, can be accumulated only
where much sediment is deposited on the subsiding bed of the sea. During the
alternate periods of elevation and of stationary level the record will be blank.
During these latter periods there will proably be more variability in the fonns
of life; during periods of subsidence, more extinction.
With respect to the absence of fossiliferous fonnations beneath the lowest
Silurian strata, I can only recur to the hypothesis given in the ninth chapter.
That the geological record is imperfect all will admit; but that it is imperfect
to the degree which I require, few will be inclined to admit. If we look to long
enough intervals of time, geology plainly declares that all species have
changed; and they have changed in the manner which my theory requires, for
they have changed slowly and in a graduated manner. We clearly see this in
the fossil remains from consecutive formations invariably being much more
closely related to each other, than are the fossils from fonnations distant from
each other in time.
Such is the sum of the several chief objections and difficulties which may
justly be urged against my theory; and I have now briefly recapitulated the an­
swers and explanations which can be given to them. I have felt these difficul­
ties far too heavily during many years to doubt their weight. But it deserves
especial notice that the more important objections relate to questions on which
we are confessedly ignorant; nor do we know how ignorant we are. We do not
know all the possible transitional gradations between the simplest and the
most perfect organs; it cannot be pretended that we know all the varied means
of Distribution during the long lapse of years, or that we know how imperfect
the Geological Record is. Grave as these several difficulties are, in my judg­
ment they do not overthrow the theory of descent with modification.
Now let us turn to the other side of the argument. Under domestication we
see much variability. This seems to be mainly due to the reproductive system
being eminently susceptible to changes in the conditions of life so that this
system, when not rendered impotent, fails to reproduce offspring exactly like
the parent-fonn. Variability is governed by many complex laws,-by correla­
tion of growth, by use and disuse, and by the direct action of the physical con­
ditions of life. There is much difficulty in ascertaining how much modification
our domestic productions have undergone; but we may safely infer that the
amount has been large, and that modifications can be inherited for long peri-
336 D A R W I N ' S G H O S T

ods. As long as the conditions of life remain the same, we have reason to be­
lieve that a modification, which has already been inherited for many genera­
tions, may continue to be inherited for an almost infinite number of
generations. On the other hand we have evidence that variability, when it has
once come into play, does not wholly cease; for new varieties are still occa­
sionally produced by our most anciently domesticated productions.
Man does not actually produce variability; he only unintentionally exposes
organic beings to new conditions of life, and then nature acts on the organisa­
tion, and causes variability. But man can and does select the variations given
to him by nature, and thus accumulate them in any desired manner. He thus
adapts animals and plants for his own benefit or pleasure. He may do this me­
thodically, or he may do it unconsciously by preserving the individuals most
useful to him at the time, without any thought of altering the breed. It is cer­
tain that he can largely influence the character of a breed by selecting, in each
successive generation, individual differences so slight as to be quite inappre­
ciable by an uneducated eye. This process of selection has been the great
agency in the production of the most distinct and useful domestic breeds. That
many of the breeds produced by man have to a large extent the character of
natural species, is shown by the inextricable doubts whether very many of
them are varieties or aboriginal species.
There is no obvious reason why the principles which have acted so effi­
ciently under domestication should not have acted under nature. In the preser­
vation of favoured individuals and races, during the constantly-recurrent
Struggle for Existence, we see the most powerful and ever-acting means of se­
lection. The struggle for existence inevitably follows from the high geometri­
cal ratio of increase which is common to all organic beings. This high rate of
increase is proved by calculation, by the effects of a succession of peculiar
seasons, and by the results of naturalisation, as explained in the third chapter.
More individuals are born than can possibly survive. A grain in the balance
will determine which individual shall live and which shall die,-which vari­
ety or species shall increase in number, and which shall decrease, or finally
become extinct. As the individuals of the same species come in all respects
into the closest competition with each other, the struggle will generally be most
severe between them; it will be almost equally severe between the varieties of
the same species, and next in severity between the species of the same genus.
But the struggle will often be very severe between beings most remote in the
scale of nature. The slightest advantage in one being, at any age or during any
season, over those with which it comes into competition, or better adaptation
in however slight a degree to the surrounding physical conditions, will turn the
balance.
 Recapitulation and Conclusion 3 37

With animals having separated sexes there will in most cases be a struggle
between the males for possession of the females. The most vigorous individu­
als, or those which have most successfully struggled with their conditions of
life, will generally leave most progeny. But success will often depend on hav­
ing special weapons or means of defence, or on the charms of the males; and
the slightest advantage will lead to victory.
As geology plainly proclaims that each land has undergone great physical
changes, we might have expected that organic beings would have varied under
nature, in the same way as they generally have varied under the changed con­
ditions of domestication. And if there be any variability under nature, it would
be an unaccountable fact if natural selection had not come into play. It has
often been asserted, but the assertion is quite incapable of proof, that the
amount of variation under nature is a strictly limited quantity. Man, though
acting on external characters alone and often capriciously, can produce within
a short period a great result by adding up mere individual differences in his
domestic productions; and every one admits that there are at least individual
differences in species under nature. But, besides such differences, all natu­
ralists have admitted the existence of varieties, which they think sufficiently
distinct to be worthy of record in systematic works. No one can draw any clear
distinction between individual differences and slight varieties; or between
more plainly marked varieties and subspecies, and species. Let it be observed
how naturalists differ in the rank which they assign to the many representative
forms in Europe and North America.
If then we have under nature variability and a powerful agent always ready
to act and select, why should we doubt that variations in any way useful to be­
ings, under their excessively complex relations of life, would be preserved, ac­
cumulated, and inherited? Why, if man can by patience select variations most
useful to himself, should nature fail in selecting variations useful, under
changing conditions of life, to her living products? What limit can be put to
this power, acting during long ages and rigidly scrutinising the whole consti­
tution, structure, and habits of each creature,-favouring the good and rej ect­
ing the bad? I can see no limit to this power, in slowly and beautifully adapting
each form to the most complex relations of life. The theory of natural selection,
even if we looked no further than this, seems to me to be in itself probable. I
have already recapitulated, as fairly as I could, the opposed difficulties and
objections: now let us turn to the special facts and arguments in favour of the
theory.
On the view that species are only strongly marked and permanent varieties,
and that each species first existed as a variety, we can see why it is that no line
of demarcation can be drawn between species, commonly supposed to have
338 D A R W I N ' S G H O S T

been produced by special acts of creation, and varieties which are acknowl­
edged to have been produced by secondary laws. On this same view we can un­
derstand how it is that in each region where many species of a genus have been
produced, and where they now flourish, these same species should present
many varieties; for where the manufactory of species has been active, we might
expect, as a general rule, to find it still in action; and this is the case if vari­
eties be incipient species. Moreover, the species of the large genera, which af­
ford the greater number of varieties or incipient species, retain to a certain
degree the character of varieties; for they differ from each other by a less
amount of difference than do the species of smaller genera. The closely allied
species also of the larger genera apparently have restricted ranges, and they
are clustered in little groups round other species-in which respects they re­
semble varieties. These are strange relations on the view of each species hav­
ing been independently created, but are intelligible if all species first existed
as varieties.
As each species tends by its geometrical ratio of reproduction to increase
inordinately in number; and as the modified descendants of each species will
be enabled to increase by so much the more as they become more diversified
in habits and structure, so as to be enabled to seize on many and widely dif­
ferent places in the economy of nature, there will be a constant tendency in
natural selection to preserve the most divergent offspring of any one species.
Hence during a long-continued course of modification, the slight differences,
characteristic of varieties of the same species, tend to be augmented into the
greater differences characteristic of species of the same genus. New and im­
proved varieties will inevitably supplant and exterminate the older, less im­
proved and intermediate varieties; and thus species are rendered to a large
extent defined and distinct objects. Dominant species belonging to the larger
groups tend to give birth to new and dominant forms; so that each large group
tends to become still larger, and at the same time more divergent in character.
But as all groups cannot thus succeed in increasing in size, for the world would
not hold them, the more dominant groups beat the less dominant. This ten­
dency in the large groups to go on increasing in size and diverging in charac­
ter, together with the almost inevitable contingency of much extinction,
explains the arrangement of all the forms of life, in groups subordinate to
groups, all within a few great classes, which we now see everywhere around us,
and which has prevailed throughout all time. This grand fact of the grouping
of all organic beings seems to me utterly inexplicable on the theory of creation.
As natural selection acts solely by accumulating slight, successive,
favourable variations, it can produce no great or sudden modification; it can
act only by very short and slow steps. Hence the canon of "Natura non facit
 Recapitulation and Conclusion 339

saltum," which every fresh addition to our knowledge tends t o make more
strictly correct, is on this theory simply intelligible. We can plainly see why
nature is prodigal in variety, though niggard in innovation. But why this should
be a law of nature if each species has been independently created, no man can
explain.
Many other facts are, as it seems to me, explicable on this theory. How
'
strange it is that a bird, under the form of woodpecker, should have been cre­
ated to prey on insects on the ground; that upland geese, which never or rarely
swim, should have been created with webbed feet; that a thrush should have
been created to dive and feed on subaquatic insects; and that a petrel should
have been created with habits and structure fitting it for the life of an auk or
grebe! and so on in endless other cases. But on the view of each species con­
stantly trying to increase in number, with natural selection always ready to
adapt the slowly varying descendants of each to any unoccupied or ill­
occupied place in nature, these facts cease to be strange, or perhaps might
even have been anticipated.
As natural selection acts by competition, it adapts the inhabitants of each
country only in relation to the degree of perfection of their associates; so that
we need feel no surprise at the inhabitants of any one country, although on the
ordinary view supposed to have been specially created and adapted for that
country, being beaten and supplanted by the naturalised productions from an­
other land. Nor ought we to marvel if all the contrivances in nature be not, as
far as we can judge, absolutely perfect; and if some of them be abhorrent to our
ideas of fitness. We need not marvel at the sting of the bee causing the bee's
own death; at drones being produced in such vast numbers for one single act,
and being then slaughtered by their sterile sisters; at the astonishing waste of
pollen by our fir-trees; at the instinctive hatred of the queen bee for her own
fertile daughters; at ichneumonidae feeding within the live bodies of caterpil­
lars; and at other such cases. The wonder indeed is, on the theory of natural
selection, that more cases of the want of absolute perfection have not been ob­
served.
The complex and little known laws governing variation are the same, as far
as we can see, with the laws which have governed the production of so-called
specific forms. In both cases physical conditions seem to have produced but
little direct effect; yet when varieties enter any zone, they occasionally assume
some of the characters of the species proper to that zone. In both varieties and
species, use and disuse seem to have produced some effect; for it is difficult to
resist this conclusion when we look, for instance, at the logger-headed duck,
which has wings incapable of flight, in nearly the same condition as in the do­
mestic duck; or when we look at the burrowing tucutucu, which is occasion-
340 D A R W I N ' S G H O S T

ally blind, and then at certain moles, which are habitually blind and have their
eyes covered with skin; or when we look at the blind animals inhabiting the
dark caves of America and Europe. In both varieties and species correction of
growth seems to have played a most important part, so that when one part has
been modified other parts are necessarily modified. In both varieties and
species reversions to long-lost characters occur. How inexplicable on the the­
ory of creation is the occasional appearance of stripes on the shoulder and legs
of the several species of the horse-genus and in their hybrids ! How simply is
this fact explained if we believe that these species have descended from a
striped progenitor, in the same manner as the several domestic breeds of pi­
geon have descended from the blue and barred rock-pigeon!
On the ordinary view of each species having been independently created,
why should the specific characters, or those by which the species of the same
genus differ from each other, be more variable than the generic characters in
which they all agree? Why, for instance, should the colour of a flower be more
likely to vary in any one species of a genus, if the other species, supposed to
have been created independently, have differently coloured flowers, than if all
the species of the genus have the same coloured flowers? If species are only
well-marked varieties, of which the characters have become in a high degree
permanent, we can understand this fact; for they have already varied since
they branched off from a common progenitor in certain characters, by which
they have come to be specifically distinct from each other; and therefore these
same characters would be more likely still to be variable than the generic
characters which have been inherited without change for an enormous period.
It is inexplicable on the theory of creation why a part developed in a very un­
usual manner in any one species of a genus, and therefore, as we may natu­
rally infer, of great importance to the species, should be eminently liable to
variation; but, on my view, this part has undergone, since the several species
branched off from a common progenitor, an unusual amount of variability and
modification, and therefore we might expect this part generally to be still vari­
able. But a part may be developed in the most unusual manner, like the wing
of a bat, and yet not be more variable than any other structure, if the part be
common to many subordinate forms, that is, if it has been inherited for a very
long period; for in this case it will have been rendered constant by long-con­
tinued natural selection.
Glancing at instincts, marvellous as some are, they offer no greater difficulty
than does corporeal structure on the theory of the natural selection of succes­
sive, slight, but profitable modifications. We can thus understand why nature
moves by graduated steps in endowing different animals of the same class with
their several instincts. I have attempted to show how much light the principle
 Recapitulation and Conclusion 341

of gradation throws on the admirable architectural powers of the hive-bee.

Habit no doubt sometimes comes into play in modifying instincts; but it cer­
tainly is not indispensable, as we see, in the case of neuter insects, which leave
no progeny to inherit the effects of long-continued habit. On the view of all the
species of the same genus having descended from a common parent, and hav­
ing inherited much in common, we can understand how it is that allied
species, when placed under considerably different conditions of life, yet
should follow nearly the same instincts; why the thrush of South America, for
instance, lines her nest with mud like our British species. On the view of in­
stincts having been slowly acquired through natural selection we need not
marvel at some instincts being apparently not perfect and liable to mistakes,
and at many instincts causing other animals to suffer.
If species be only well-marked and permanent varieties, we can at once see
why their crossed offspring should follow the same complex laws in their de­
grees and kinds of resemblance to their parents,-in being absorbed into each
other by successive crosses, and in other such points,-as do the crossed off­
spring of acknowledged varieties. On the other hand, these would be strange
facts if species have been independently created, and varieties have been pro­
duced by secondary laws.
If we admit that the geological record is imperfect in an extreme degree,
then such facts as the record gives, support the theory of descent with modifi­
cation. New species have come on the stage slowly and at successive intervals;
and the amount of change, after equal intervals of time, is widely different in
different groups. The extinction of species and of whole groups of species,
which has played so conspicuous a part in the history of the organic world, al­
most inevitably follows on the principle of natural selection; for old forms will
be supplanted by new and improved forms. Neither single species nor groups
of species reappear when the chain of ordinary generation has once been bro­
ken. The gradual diffusion of dominant forms, with the slow modification of
their descendants, causes the forms of life, after long intervals of time, to ap­
pear as if they had changed simultaneously throughout the world. The fact of
the fossil remains of each formation being in some degree intermediate in
character between the fossils in the formations above and below, is simply ex­
plained by their intermediate position in the chain of descent. The grand fact
that all extinct organic beings belong to the same system with recent beings,
falling either into the same or into intermediate groups, follows from the living
and the extinct being the offspring of common parents. As the groups which
have descended from an ancient progenitor have generally diverged in char­
acter, the progenitor with its early descendants will often be intermediate in
character in comparison with its later descendants; and thus we can see why
342 D A R W I N ' S G H O S T

the more ancient a fossil is, the oftener it stands in some degree intermediate
between existing and allied groups. Recent forms are generally looked at as
being, in some vague sense, higher than ancient and extinct forms; and they
are in so far higher as the later and more improved forms have conquered the
older and less improved organic beings in the struggle for life. Lastly, the law
of the long endurance of allied forms on the same continent; of marsupials in
Australia, of edentata in America, and other such cases,-is intelligible, for
within a confined country, the recent and the extinct will naturally be allied by
descent.
Looking to geographical distribution, if we admit that there has been during
the long course of ages much migration from one part of the world to another,
owing to former climatal and geographical changes and to the many occasional
and unknown means of dispersal, then we can understand, on the theory of de­
scent with modification, most of the great leading facts in Distribution. We can
see why there should be so striking a parallelism in the distribution of organic
beings throughout space, and in their geological succession throughout time;
for in both cases the beings have been connected by the bond of ordinary gen­
eration, and the means of modification have been the same. We see the full
meaning of the wonderful fact, which must have struck every traveller, namely,
that on the same continent, under the most diverse conditions, under heat and
cold, on mountain and lowland, on deserts and marshes, most of the inhabi­
tants within each great class are plainly related; for they will generally be de­
scendants of the same progenitors and early colonists. On this same principle
of former migration, combined in most cases with modification, we can under­
stand, by the aid of the Glacial period, the identity of some few plants, and the
close alliance of many others, on the most distant mountains, under the most
different climates; and likewise the close alliance of some of the inhabitants of
the sea in the northern and southern temperate zones, though separated by the
whole intertropical ocean. Although two areas may present the same physical
conditions of life, we need feel no surprise at their inhabitants being widely
different, if they have been for a long period completely separated from each
other; for as the relation of organism to organism is the most important of all
relations, and as the two areas will have received colonists from some third
source or from each other, at various periods and in different proportions, the
course of modification in the two areas will inevitably be different.
On this view of migration, with subsequent modification, we can see why
oceanic islands should be inhabited by few species, but of these, that many
should be peculiar. We can clearly see why those animals which cannot cross
wide spaces of ocean, as frogs and terrestrial mammals, should not inhabit
oceanic islands; and why, on the other hand, new and peculiar species of bats,
 Recapitulation and Conclusion 343

which can traverse the ocean, should s o often b e found o n islands far distant
from any continent. Such facts as the presence of peculiar species of bats, and
the absence of all other mammals, on oceanic islands, are utterly inexplicable
on the theory of independent acts of creation.
The existence of closely allied or representative species in any two areas,
implies, on the theory of descent with modification, that the same parents for­
merly inhabited both areas; and we almost invariably find that wherever many
closely allied species inhabit two areas, some identical species common to
both still exist. Wherever many closely allied yet distinct species occur, many
doubtful forms and varieties of the same species likewise occur. It is a rule of
high generality that the inhabitants of each area are related to the inhabitants
of the nearest source whence immigrants might have been derived. We see this
in nearly all the plants and animals of the Galapagos archipelago, of Juan Fer­
nandez, and of the other American islands being related in the most striking
manner to the plants and animals of the neighbouring American mainland; and
those of the Cape de Verde archipelago and other African islands to the
Mrican mainland. It must be admitted that these facts receive no explanation
on the theory of creation.
The fact, as we have seen, that all past and present organic beings consti­
tute one grand natural system, with group subordinate to group, and with ex­
tinct groups often falling in between recent groups, is intelligible on the theory
of natural selection with its contingencies of extinction and divergence of
character. On these same principles we see how it is, that the mutual affinities
of the species and genera within each class are so complex and circuitous. We
see why certain characters are far more serviceable than others for classifica­
tion;-why adaptive characters, though of paramount importance to the being,
are of hardly any importance in classification; why characters derived from
rudimentary parts, though of no service to the being, are often of high classifi­
catory value; and why embryological characters are the most valuable of all.
The real affinities of all organic beings are due to inheritance or community of
descent. The natural system is a genealogical arrangement, in which we have
to discover the lines of descent by the most permanent characters, however
slight their vital importance may be.
The framework of bones being the same in the hand of a man, wing of a bat,
fin of the porpoise, and leg of the horse,-the same number of vertebrae form­
ing the neck of the giraffe and of the elephant,-and innumerable other such
facts, at once explain themselves on the theory of descent with slow and slight
successive modifications. The similarity of pattern in the wing and leg of a bat,
though used for such different purposes,-in the j aws and legs of a crab,-in
the petals, stamens, and pistils of a flower, is likewise intelligible on the view
344 D A R W I N ' S G H O S T

of the gradual modification of parts or organs, which were alike in the early
progenitor of each class. On the principle of successive variations not always
supervening at an early age, and being inherited at a corresponding not early
period of life, we can clearly see why the embryos of mammals, birds, reptiles,
and fishes should be so closely alike, and should be so unlike the adult forms.
We may cease marvelling at the embryo of an air-breathing mammal or bird
having branchial slits and arteries running in loops, like those in a fish which
has to breathe the air dissolved in water, by the aid of well-developed
branchiae.
Disuse, aided sometimes by natural selection, will often tend to reduce an
organ, when it has become useless by changed habits or under changed con­
ditions of life; and we can clearly understand on this view the meaning of rudi­
mentary organs. But disuse and selection will generally act on each creature,
when it has come to maturity and has to play its full part in the struggle for ex­
istence, and will thus have little power of acting on an organ during early life;
hence the organ will not be much reduced or rendered rudimentary at this
early age. The calf, for instance, has inherited teeth, which never cut through
the gums of the upper jaw, from an early progenitor having well-developed
teeth; and we may believe, that the teeth in the mature animal were reduced,
during successive generations, by disuse or by the tongue and palate having
been fitted by natural selection to browse without their aid; whereas in the calf,
the teeth have been left untouched by selection or disuse, and on the princi­
ple of inheritance at corresponding ages have been inherited from a remote pe­
riod to the present day. On the view of each organic being and each separate
organ having been specially created, how utterly inexplicable it is that parts,
like the teeth in the embryonic calf or like the shrivelled wings under the sol­
dered wing-covers of some beetles, should thus so frequently bear the plain
stamp of inutility! Nature may be said to have taken pains to reveal, by rudi­
mentary organs and by homologous structures, her scheme of modification,
which it seems that we wilfully will not understand.

I have now recapitulated the chief facts and considerations which have thor­
oughly convinced me that species have changed, and are still slowly changing
by the preservation and accumulation of successive slight favourable varia­
tions. Why, it may be asked, have all the most eminent living naturalists and
geologists rejected this view of the mutability of species? It cannot be asserted
that organic beings in a state of nature are subject to no variation; it cannot be
proved that the amount of variation in the course of long ages is a limited quan­
tity; no clear distinction has been, or can be, drawn between species and well-
 Recapitulation and Conclusion 345

marked varieties. I t cannot be maintained that species when intercrossed are

invariably sterile, and varieties invariably fertile; or that sterility is a special
endowment and sign of creation. The belief that species were immutable pro­
ductions was almost unavoidable as long as the history of the world was
thought to be of short duration; and now that we have acquired some idea of
the lapse of time, we are too apt to assume, without proof, that the geological
record is so perfect that it would have afforded us plain evidence of the muta­
tion of species, if they had undergone mutation.
But the chief cause of our natural unwillingness to admit that one species
has given birth to other and distinct species, is that we are always slow in ad­
mitting any great change of which we do not see the intermediate steps. The
difficulty is the same as that felt by so many geologists, when Lyell first in­
sisted that long lines of inland cliffs had been formed, and great valleys exca­
vated, by the slow action of the coast-waves. The mind cannot possibly grasp
the full meaning of the term of a hundred million years; it cannot add up and
perceive the full effects of many slight variations, accumulated during an al­
most infinite number of generations .
Although I am fully convinced of the truth of the views given in this volume
under the form of an abstract, I by no means expect to convince experienced
naturalists whose minds are stocked with a multitude of facts all viewed, dur­
ing a long course of years, from a point of view directly opposite to mine. It is
so easy to hide our ignorance under such expressions as the "plan of creation,"
"unity of design," &c., and to think that we give an explanation when we only
restate a fact. Any one whose disposition leads him to attach more weight to
unexplained difficulties than to the explanation of a certain number of facts
will certainly reject my theory. A few naturalists, endowed with much flexibil­
ity of mind, and who have already begun to doubt on the immutability of
species, may be influenced by this volume; but I look with confidence to the
future, to young and rising naturalists, who will be able to view both sides of
the question with impartiality. Whoever is led to believe that species are mu­
table will do good service by conscientiously expressing his conviction; for
only thus can the load of prejudice by which this subject is overwhelmed be
removed.
Several eminent naturalists have of late published their belief that a multi­
tude of reputed species in each genus are not real species; but that other
species are real, that is, have been independently created. This seems to me a
strange conclusion to arrive at. They admit that a multitude of forms, which till
lately they themselves thought were special creations, and which are still thus
looked at by the majority of naturalists, and which consequently have every
external characteristic feature of true species,-they admit that these have
346 D A R W I N ' S G H O S T

been produced by variation, but they refuse to extend the same view to other
and very slightly different forms. Nevertheless they do not pretend that they
can define, or even conjecture, which are the created forms of life, and which
are those produced by secondary laws. They admit variation as a vera causa in
one case, they arbitrarily reject it in another, without assigning any distinction
in the two cases. The day will come when this will be given as a curious illus­
tration of the blindness of preconceived opinion. These authors seem no more
startled at a miraculous act of creation than at an ordinary birth. But do they
really believe that at innumerable periods in the earth's history certain ele­
mental atoms have been commanded suddenly to flash into living tissues? Do
they believe that at each supposed act of creation one individual or many were
produced? Were all the infinitely numerous kinds of animals and plants cre­
ated as eggs or seed, or as full grown? and in the case of mammals, were they
created bearing the false marks of nourishment from the mother's womb? Al­
though naturalists very properly demand a full explanation of every difficulty
from those who believe in the mutability of species, on their own side they ig­
nore the whole subject of the first appearance of species in what they consider
reverent silence.
It may be asked how far I extend the doctrine of the modification of species.
The question is difficult to answer, because the more distinct the forms are
which we may consider, by so much the arguments fall away in force. But some
arguments of the greatest weight extend very far. All the members of whole
classes can be connected together by chains of affinities, and all can be clas­
sified on the same principle, in groups subordinate to groups. Fossil remains
sometimes tend to fill up very wide intervals between existing orders. Organs
in a rudimentary condition plainly show that an early progenitor had the organ
in a fully developed state; and this in some instances necessarily implies an
enormous amount of modification in the descendants. Throughout whole
classes various structures are formed on the same pattern, and at an embry­
onic age the species closely resemble each other. Therefore I cannot doubt that
the theory of descent with modification embraces all the members of the same
class. I believe that animals have descended from at most only four or five
progenitors, and plants from an equal or lesser number.
Analogy would lead me one step further, namely, to the belief that all ani­
mals and plants have descended from some one prototype. But analogy may be
a deceitful guide. Nevertheless all living things have much in common, in their
chemical composition, their germinal vesicles, their cellular structure, and
their laws of growth and reproduction. We see this even in so trifling a cir­
cumstance as that the same poison often similarly affe cts plants and animals;
or that the poison secreted by the gall-fly produces monstrous growths on the
 Recapitulation and Conclusion 347

wild rose or oak-tree. Therefore I should infer from analogy that probably all
the organic beings which have ever lived on this earth have descended from
some one primordial form, into which life was first breathed.

When the views entertained in this volume on the origin of species, or when
analogous views are generally admitted, we can dimly foresee that there will
be a considerable revolution in natural history. Systematists will be able to
pursue their labours as at present; but they will not be incessantly haunted by
the shadowy doubt whether this or that form be in essence a species. This I feel
sure, and I speak after experience, will be no slight relief. The endless dis­
putes whether or not some fifty species of British brambles are true species
will cease. Systematists will have only to decide (not that this will be easy)
whether any form be sufficiently constant and distinct from other forms, to be
capable of definition; and if definable, whether the differences be sufficiently
important to deserve a specific name . This latter point will become a far more
essential consideration than it is at present; for differences, however slight, be­
tween any two forms, if not blended by intermediate gradations, are looked at
by most naturalists as sufficient to raise both forms to the rank of species.
Hereafter we shall be compelled to acknowledge that the only distinction be­
tween species and well-marked varieties is, that the latter are known, or be­
lieved, to be connected at the present day by intermediate gradations, whereas
species were formerly thus connected. Hence, without quite rejecting the con­
sideration of the present existence of intermediate gradations between any two
forms, we shall be led to weigh more carefully and to value higher the actual
amount of difference between them. It is quite possible that forms now gener­
ally acknowledged to be merely varieties may hereafter be thought worthy of
specific names, as with the primrose and cowslip; and in this case scientific
and common language will come into accordance. In short, we shall have to
treat species in the same manner as those naturalists treat genera, who admit
that genera are merely artificial combinations made for convenience. This may
not be a cheering prospect; but we shall at least be freed from the vain search
for the undiscovered and undiscoverable essence of the term species.
The other and more gen�ral departments of natural history will rise greatly
in interest. The terms used by naturalists of affinity, relationship, community
of type, paternity, morphology, adaptive characters, rudimentary and aborted
organs, &c. , will cease to be metaphorical, and will have a plain signification.
When we no longer look at an organic being as a savage looks at a ship, as at
something wholly beyond his comprehension; when we regard every produc­
tion of nature as one which has had a history; when we contemplate every com-
348 D A R W I N ' S G H O S T

plex structure and instinct as the summing up of many contrivances, each use­
ful to the possessor, nearly in the same way as when we look at any great me­
chanical invention as the summing up of the labour, the experience, the
reason, and even the blunders of numerous workmen; when we thus view each
organic being, how far more interesting, I speak from experience, will the
study of natural history become!
A grand and almost untrodden field of inquiry will be opened, on the causes
and laws of variation, on correlation of growth, on the effects of use and dis­
use, on the direct action of external conditions, and so forth. The study of do­
mestic productions will rise immensely in value. A new variety raised by man
will be a far more important and interesting subject for study than one more
species added to the infinitude of already recorded species. Our classifications
will come to be, as far as they can be so made, genealogies; and will then truly
give what may be called the plan of creation. The rules for classifying will no
doubt become simpler when we have a definite object in view. We possess no
pedigrees or armorial bearings; and we have to discover and trace the many di­
verging lines of descent in our natural genealogies, by characters of any kind
which have long been inherited. Rudimentary organs will speak infallibly with
respect to the nature of long-lost structures. Species and groups of species,
which are called aberrant, and which may fancifully be called living fossils,
will aid us in forming a picture of the ancient forms of life. Embryology will re­
veal to us the structure, in some degree obscured, of the prototypes of each
great class.
When we can feel assured that all the individuals of the same species, and
all the closely allied species of most genera, have within a not very remote pe­
riod descended from one parent, and have migrated from some one birthplace;
and when we better know the many means of migration, then, by the light
which geology now throws, and will continue to throw, on former changes of
climate and of the level of the land, we shall surely be enabled to trace in an
admirable manner the former migrations of the inhabitants of the whole world.
Even at present, by comparing the differences of the inhabitants of the sea on
the opposite sides of a continent, and the nature of the various inhabitants of
that continent in relation to their apparent means of immigration, some light
can be thrown on ancient geography.
The noble science of Geology loses glory from the extreme imperfection of
the record. The crust of the earth with its embedded remains must not be
looked at as a well-filled museum, but as a poor collection made at hazard and
at rare intervals, The accumulation of each great fossiliferous formation will be
recognised as having depended on an unusual concurrence of circumstances,
and the blank intervals between the successive stages as having been of vast
 Recapitulation and Conclusion 349

duration. But w e shall b e able to gauge with some security the duration o f these
intervals by a comparison of the preceding and succeeding organic forms. We
must be cautious in attempting to correlate as strictly contemporaneous two
formations, which include few identical species, by the general succession of
their forms of life. As species are produced and exterminated by slowly acting
and still existing causes, and not by miraculous acts of creation and by cata­
strophes; and as the most important of all causes of organic change is one
which is almost independent of altered and perhaps suddenly altered physical
conditions, namely, the mutual relation of organism to organism,-the im­
provement of one being entailing the improvement or the extermination of oth­
ers; it follows, that the amount of organic change in the fossils of consecutive
formations probably serves as a fair measure of the lapse of actual time . A
number of species, however, keeping in a body might remain for a long period
unchanged, whilst within this same period, several of these species, by mi­
grating into new countries and coming into competition with foreign associ­
ates, might become modified; so that we must not overrate the accuracy of
organic change as a measure of time. During early periods of the earth's his­
tory, when the forms of life were probably fewer and simpler, the rate of change
was probably slower; and at the first dawn of life, when very few forms of the
simplest structure existed, the rate of change may have been slow in an ex­
treme degree. The whole history of the world, as at present known, although of
a length quite incomprehensible by us, will hereafter be recognised as a mere
fragment of time, compared with the ages which have elapsed since the first
creature, the progenitor of innumerable extinct and living descendants, was
created.
In the distant future I see open fields for far more important researches. Psy­
chology will be based on a new foundation, that of the necessary acquirement
of each mental power and capacity by gradation. Light will be thrown on the
origin of man and his history.
Authors of the highest eminence seem to be fully satisfied with the view that
each species has been independently created. To my mind it accords better
with what we know of the laws impressed on matter by the Creator, that the pro­
duction and extinction of the past and present inhabitants of the world should
have been due to secondary causes, like those determining the birth and death
of the individual. When I view all beings not as special creations, but as the
lineal descendants of some few beings which lived long before the first bed of
the Silurian system was deposited, they seem to me to become ennobled. Judg­
ing from the past, we may safely infer that not one living species will transmit
its unaltered likeness to a distant futurity. And of the species now living very
few will transmit progeny of any kind to a far distant futurity; for the manner
350 D A R W I N ' S G H O S T

in which all organic beings are grouped, shows that the greater number of
species of each genus, and all the species of many genera, have left no de­
scendants, but have become utterly extinct. We can so far take a prophetic
glance into futurity as to foretell that it will be the common and widely-spread
species, belonging to the larger and dominant groups, which will ultimately
prevail and procreate new and dominant species. As all the living forms of life
are the lineal descendants of those which lived long before the Silurian epoch,
we may feel certain that the ordinary succession by generation has never once
been broken, and that no cataclysm has desolated the whole world. Hence we
may look with some confidence to a secure future of equally inappreciable
length. And as natural selection works solely by and for the good of each being,
all corporeal and mental endowments will tend to progress towards perfection.
It is interesting to contemplate an entangled bank, clothed with many plants
of many kinds, with birds singing on the bushes, with various insects flitting
about, and with worms crawling through the damp earth, and to reflect that
these elaborately constructed forms, so different from each other, and depen­
dent on each other in so complex a manner, have all been produced by laws
acting around us. These laws, taken in the largest sense, being Growth with
Reproduction; inheritance which is almost implied by reproduction; Variabil­
ity from the indirect and direct action of the external conditions of life, and
from use and disuse; a Ratio of Increase so high as to lead to a Struggle for Life,
and as a consequence to Natural Selection, entailing Divergence of Character
and the Extinction of less-improved forms. Thus, from the war of nature, from
famine and death, the most exalted object which we are capable of conceiving,
namely, the production of the higher animals, directly follows. There is
grandeur in this view of life, with its several powers, having been originally
breathed into a few forms or into one; and that, whilst this planet has gone cy­
cling on according to the fixed law of gravity, from so simple a beginning end­
less forms most beautiful and most wonderful have been, and are being,
evolved.
 F U RT H E R R E A D I N G

There is no obvious reason why the theory of evolution should attract the finest sci­
entific writers, but it is so. From Darwin to the great popularizers of today, natural
selection-and what it implies-has inspired the best of all scientific literature.
Darwin stands alone. Not many people with today's literary tastes would open
The Origin for pleasure (although it contains much to enjoy) . The Voyage ofthe Bea­
gle, though, is a classic of travel writing that reads as well now as it did in 1 839.
When it comes to Patagonia, Darwin was the Bruce Chatwin of his day. His Auto­
biographical Sketch, brief though it is, is also a simple and often moving account of
a long life well spent. There is a multitude of biographies. Some emphasize the so­
cial context of Darwin's life and (inevitably) the coincidence between the theory of
evolution by natural selection and the affiuent Victorian world in which he was
brought up, while others are more concerned with his own story and his scientific
views, detached ftom political speculation. Janet Browne's extensive biography,
Charles Darwin, with its first volume, Voyaging {Princeton University Press, 1 996),
i s comprehensive, clear and dispassionate. Adrian Desmond and James Moore's
Darwin (Norton, 1 994) puts his ideas firmly into the context of his own society.
Darwin has been well served by his successors. Some of science's most gifted au­
thors-Richard Dawkins, Jared Diamond and Stephen Jay Gould-have produced
a series of books that illuminate his work, bring it up to date and put it into his­
torical, literary and social context. My own favorites, among many, are Dawkins's
Climbing Mount Improbable (Norton, 1996), Diamond's The Third Chimpanzee
1 993) and Gould's Dinosaur in a Haystack (Harmony, 1 996).
(HarperPerennial,
Evolution suffers from a surfeit of both anecdotes and mathematics. The former
have had a fairer run of it (and many are displayed in Richard Milner's Encyclopae­
dia ofEvolution, published by Facts on File, New York and Oxford, 1 990). For a
real insight into the subject it is hard to escape from at least a nod at mathematical
352 Further Reading

reality. The standard texts on the modern theory of evolution are Douglas Fu­
tuyma's Evolutionary Biology (Sinauer Associates, Sunderland, Mass. , 3d ed. , 1 998) ,
John Maynard Smith's Evolutionary Genetics (Oxford University Press, 2d ed. ,
1 998) and Mark Ridley's Evolution (Blackwell Science, Oxford, 2 d ed. , 1 996) . All
give a solid introduction to the subject (and to the many problems that have been
so lightly skipped over in my own book) . In most modern texts (unlike The Origin
itself) plants do not get a fair say. That balance is corrected by K. J . Niklas's The
Evolutionary Biology ofPlants (University of Chicago Press, 1 997) .
As the theory of evolution encompasses the whole of the biological sciences,
there are dozens of popular works that illuminate part of the Darwinian story.
Richard Fortey in his Life: An Unauthorized Biography (Knopf, 1 998) gives a vivid
account of evolution's raw material, the advance of existence through time. Matt
Ridley's The Red Queen (Penguin USA, 1 995) discusses an aspect of the subject not
enough considered in my own book: natural selection as a race in which not all shall
have prizes. His Genome: The Autobiography ofa Species in 23 Chapters (Fourth Es­
tate, 2000) says more about the story of human evolution as revealed by the genes.
Any science writer has to mine the scientific literature, technical and tedious as
it often is, in the hope of extracting a few gems. This book has been no exception:
and at times through the thousand and more books and papers consulted here it ap­
peared that the seam was almost exhausted. I list just a few of the most recent (and
more accessible) publications as an entree to the literature; all appeared in the 1 990s
and through them it is possible to refer back to earlier work. Each week, the jour­
nals Science and Nature keep up-to-date with the almost limitless world of evolu­
tion. Trends in Research in Ecology and Evolution produces reviews of advances in the
field. The main technical journal is Evolution, complemented by The American
Naturalist, The Proceedings ofthe National Academy ofSciences and The Philosophi­
cal Transactions ofthe Royal Society ofLondon. More specialist journals include The
Journal ofMolecular Evolution. The Journal ofAnimal Ecology, Behavioural Ecology
and Sociobiology, Systematic Biology, Paleobiology, and many more. The Cottage Gar­
dener and Country Gentleman's Companion (much consulted by Darwin) is, alas, de­
funct.
In science, the Internet is overtaking the printed word, and there are several ex­
cellent sites that keep bang up-to-date with modern biology. Among them are the
WWW Virtual Library: Evolution (http://golgi.harvard.edu/biopages/evolution.
html) ; MendelWeb (http://www.netspace.org/MendelWeb) ; the Society for the
Study of Evolution (http://lsvl.la.asu.edu/evolution) ; the Tree of Life (http://
phylogeny.arizona.edultree/phylogeny.html) ; and the Natural History Museum in
London (http://nhm.ac.uk) . There is also the regularly updated and highly infor­
mative HMS Beagle at http://biomednet.com/hmsbeagle. No doubt others will
come and go.
 Further Reading 353

Introduction
Gatesy, ] . , "More DNA support for a CetacealHippopotamidae clade: the blood­
clotting protein gene gamma-fibrinogen, " Molecular Biology and Evolution 14:
537-43, 1 997.
Goudsmit, ]., Viral Sex: The Nature ofAIDS, Oxford University Press, 1 997.
Hoelzel, R. , "Genetics and ecology of whales and dolphins," Annual Review ofEcol­
ogy and Systematics 2 5 : 377-99, 1 994.
Hooper, E., "Sailors and star-bursts, and the arrival of HN," British MedicalJour­
na1 3 1 5 : 1 689-9 1 , 1 997.
Korber, B., Theiler, ]., and Wolinsky, S., "Limitations of a molecular clock applied
to considerations of the origins of HN- l ," Science 280: 1 868-7 1 , 1 99 8 .
Molla, A., e t al. , "Ordered accumulation o f mutations i n HIV protease confers re­
sistance to ritonavir, " Nature Medicine 2: 760-66, 1 996.
Montgelard, c., Catzeflis, F. M., and Douzery, E., "Phylogenetic relationship of
artiodactyls and cetaceans as deduced from the comparison of cytochrome B and
1 2S rRNA mitochondrial sequence," Molecular Biology and Evolution 14:
5 50-59, 1 997.
Nikaido, M., Rooney, A. P., and Okada N., "Phylogenetic relationships among
certartiodactyls based on insertions of short and long interspersed elements: Hip­
popotamuses are the closest extant relatives of whales, " Proceedings of the Na­
tionalAcademy ofSciences 96: 1 026 1-66, 1 999.
Thewissen, ] . G . M . , and Fish, F. E., "Locomotor evolution in the earliest
cetaceans: Functional model, modern analogues, and paleontological evidence,"
Paleobiology 23: 482-90, 1 997.

Chapter I: Variation Under Domestication

Bradley, D. G., et al. , "Mitochondrial diversity and the origins of Mrican and Eu­
ropean cattle," Proceedings of the National Academy of Sciences 93: 5 1 3 1 -3 5 ,
1 996.
Croke, v. , The Modern Ark: The Story of Zoos, Past, Present and Future, Charles
Scribner's Sons, New York, 1 997.
Frankham, R. , "Conservation genetics," Annual Review of Genetics 29: 305-27,
1 993.
Kendrick, K. M., et al., "Mothers determine sexual preference," Nature 395:
229-30, 1 998.
Ostrander, E. A. , and Giniger, E., "Insights from model systems: What man's best
friend can teach us about human biology and disease," American Journal of
Human Genetics 6 1 : 475-80, 1 997.
The Domestic Dog: Its Evolution, Behaviour and Interactions with People,
Serpell, ].,
Cambridge University Press, 1 99 5 .
Tudge, c., Last Animals at the Zoo, Oxford University Press, 1 992.
Vila, c., et al. , "Multiple and ancient origins of the domestic dog," Science 276:
1 687-89, 1 997.
354 Further Reading

Chapter II: Variation Under Nature

Barrate, E. M . , et aI. , "DNA answers the call of pipistrelle bat species, " Nature 387:
1 38-39, 1 997.
Berry, R ]., and Bronson, F. H., "Life history and bioeconomy of the house
mouse," Biological Reviews 67: 5 1 9-50, 1 992.
Din, w., et al. , "Origin and radiation of the house mouse-clues from nuclear
genes, " Journal ofEvolutionary Biology 9: 5 1 9-39, 1 996.
Garner, M., " Identification of yellow-legged gulls in Britain," British Birds 90:
25-62, 1 997.
Mallet, ]., ''A species definition for the modern synthesis," Trends in Research in
Ecology and Evolution 1 0: 294-99, 1 99 5 .
Meffe, G., and Carroll, C. R , Principles ofConservation Biology, 2d ed. , Sinauer As­
sociates, Sunderland, Mass., 1 997.
Rhymer, ] . M., and Simberloff, D., "Extinction by hybridisation and introgres­
sion, " Annual Review ofEcology and Systematics 27: 83- 1 09, 1 996.
Shuker, K., The Lost Ark: New and Rediscovered Animals of the Twentieth Century,
HarperCollins, London, 1 993.

Chapter III: Struggle for Existence

Begon, M., Harper, ] . L., and Townsend, C. R, Ecology: Individuals, Populations,
Communities, 3d ed. , Blackwell Science, Oxford, 1 996.
Behrenfeld, M . ]., and Kolber, Z. S., "Widespread iron limitation of phytoplank­
ton in the South Pacific," Science 283: 840-43, 1 999.
Boonstra, R. , et al. , "The impact of predator-induced stress on the snowshoe hare
cycle," Ecological Monographs 79 : 47 1-94, 1 998.
Budiansky, S . , Nature's Keepers: The New Science ofNature Management, Phoenix,
London, 1 996.
Green, R. E., "The influence of numbers released on the outcome of attempts to
introduce exotic bird species to New Zealand, " Journal ofAnimal Ecology 66:
25-3 5 , 1 997.
Greenwood, ] . ] . D., et al. , "Relations between abundance, body size and species
number in British birds and mammals," Philosophical Transactions ofthe Royal
Society B 35 1 : 265-78, 1 996.
Kurlansky, M., Cod: A Biography ofthe Fish That Changed the World, Penguin, 1 998.
Newton, I . , "The contribution of some recent research on birds to ecological un­
derstanding," Journal ofAnimal Ecology 64: 675-96, 1 99 5 .
Rebertus, A . ] . , e t al. , "Blowdown history and landscape patterns i n the Andes of
Tierra del Fuego, Argentina," Ecology 78: 675-96, 1 99 5 .
Rhymer, ] . M . , and Simberloff, D., "Extinction b y hybridisation and introgres­
sion," Annual Review ofEcology and Systematics 27: 83- 1 09, 1 996.
 Further Reading 355

Chapter IV: Natural Selection

(various authors) , '�timicrobial resistance," British MedicalJournal3 17: 609-90,
1 998.
Bell, G . , The Basics o/Selection, Chapman and Hall, New York, 1 997.
Frank, 1. G., "Evolution of genital masculinization: Why do female hyaenas have
such a large 'penis'?," Trends in Research in Ecology and Evolution, 1 2: 5 8-62,
1 997.
Henson, S. A. , and Warner, R. R. , "Male and female alternative reproductive be­
haviors in fishes," Annual Review o/Ecology and Systematics 28: 5 7 1 -92, 1 997.
Jennions, M. D., and Petrie, M., "Variation in mate choice and mating preferences:
A review of causes and consequences," Biological Reviews 72: 283-327, 1 997.
Kearns, K. A., Inouye, D. w., and Waser, N. M., "Endangered mutualisms: The
conservation of plant-pollinator interactions," Annual Review o/Ecology and Sys­
tematics 29: 83- 1 1 2, 1 998.
Lawton, J. H., and May, R. M., Extinction Rates, Oxford University Press, 1 99 5 .
Linhart, Y. , and Grant, M. c., "Evolutionary significance of local genetic differen­
tiation in plants," Annual Review o/Ecology and Systematics 27: 237-77, 1 996.
Majerus, M. E. N., Melanism: Evolution in Action, Oxford University Press, 1 998.
Reznick, D. N . , et al. , "Evaluation of the rate of evolution in natural populations
of guppies," Science 275: 1 934-37, 1 998.
Schluter, D., "Ecological speciation in post-glacial fishes," in Grant, P. R. , ed. , Evo­
lution on Islands, Oxford University Press, 1 998.
Welch, A. M . , Semlitsch, R. D., and Gerhardt, H. c., "Call duration as an indica­
tor of genetic quality in tree frogs," Science 280: 1 928-30, 1 998.
Williams, G. c., Natural Selection: Domains, Levels and Challenges, Oxford Univer­
sity Press, 1 992.

Chapter V: Laws of Variation

Ellegren, H., and Fridolffson, A.-K. , "Male-driven evolution of DNA sequences in
birds,"Nature Genetics 1 7 : 1 82-84, 1 997.
The Platypus and the Mermaid and otherfigments o/the Classifying Imag­
Ritvo, H . ,
ination, Harvard University Press, Cambridge, Mass. , 1 997.
Rossiter, M., "Incidence and consequences of inherited environmental effects," An­
nual Review o/Ecology and Systematics 27: 45 1-76, 1 996.

Chapter VI: Difficulties on Theory

Averof, M . , and Cohen, S . M., "Evolutionary origin of insect wings from ancestral
gills," Nature 3 8 5 : 627-30, 1 997.
Berenbaum, M. R. , Favret, c., and Schuler, M. A. , "On defining 'key innovations'
in an adaptive radiation: cytochrome P450s and Papilionidae, " American Natu­
ralist 148: S 1 39-5 5 , 1 996.
Farrell, B. D., " 'Inordinate fondness' explained: Why are there so many beetles?"
Science 28 1 : 5 5 5-59, 1 998.
356 Further Reading

Feder, ]. L., et al. , "The effects of winter length on the genetics of apple and
hawthorn races of Rhagoletis pomonella, " Evolution 5 1 : 1 862-76, 1 997.
Golding, G. B., and Dean, A. M., "The structural basis of molecular adaptation,"
Molecular Biology and Evolution 1 5: 355-69, 1 998.
Lawrence, ]. G . , and Ochman, H., "Molecular archeology of the Escherichia coli
genome, " Proceedings ofthe NationalAcademy ofSciences 95: 941 3-1 7, 1 998.
Herre, E. A. , et al. , "The evolution of mutualisms: Exploring the paths between
conflict and cooperation," Trends in Research in Ecology and Evolution 1 4: 49-53,
1 999.
Kazazian, H. H., and Moran, ] . v. , "The impact of 11 retrotransposons on the
human genome," Nature Genetics 19: 1 9-24, 1 998.
Land, M. E, "Visual acuity in insects," Annual Review ofEntomology 42: 147-77,
1 997.
Lewin, B., Genes V7. Oxford University Press, 1 997.
Rivera, M. c., et al. , "Genomic evidence for two functionally distinct gene classes,"
Proceedings ofthe National Academy ofSciences 95: 6239-44, 1 998.

Chapter VII : Instinct

DreHer, c., "Division of labor between scouts and recruits: Genetic influence and
mechanisms," Behavioral Ecology and Sociobiology 43: 1 9 1-96, 1 998.
HoHdobler, B., and Wilson, E. 0., Journey to the Ants: A V0yage ofScientific explo­
ration, Belknap Press of Harvard University Press, Cambridge, Mass. , 1 994.
Klein, N. K. , and Payne, R. B., "Evolutionary associations of brood parasitic finches
(Vidua) and their host species: Analyses of mitochondrial DNA restriction sites,"
Evolution 52: 566-82, 1 998.
Mock, D. w. , and Parker, G. A. , "Siblicide, family conflict and the evolutionary
limits of selfishness," Animal Behaviour 56: 1-10, 1 998.
Rothstein, S. I., ''A model system for coevolution: Avian brood parasitism," Annual
Review ofEcology and Systematics 2 1 : 48 1-508, 1 990.
Seeley, T. D., "Honey bee colonies are group-level adaptive units," American Nat­
uralist 1 50, supp.: S22-41 , 1997.
Stern, D. L., and Foster, W. A., "The evolution of soldiers in aphids," Biological Re­
views 7 1 : 27-79, 1 996.

Chapter VIII: Hybridism

Grant, P. R. , and Grant, E. R. , "Genetics and the origin of bird species,"Proceed­
ings ofthe NationalAcademy ofSciences 94: 7768-75, 1 997.
Howard, D. ]., and Berlocher, S . H., Endless Forms: Species and Speciation, Oxford
University Press, 1 998.
Marchetti, K. , Nakamura, H., and Gibbs, H. L., "Host-race formation in the com­
mon cuckoo," Science 282: 471-72, 1 998.
Orr, H. A. , "Haldane's Rule," Annual Review of Ecology and Systematics 28:
195-2 1 8, 1 997.
 Further Reading 357

Parker, G . A., and Partridge, L., "Sexual conflict and speciation," Philosophical
Transactions ofthe Royal Society B 353: 261-74, 1 998.
Seehausen, 0., Van Alphen, J. J. M., and Witte, E, "Eutrophication that curbs sex­
ual selection," Science 277: 1 808-1 1 , 1997.
Swanson, W. J., and Vacquier, V. D., "Concerted evolution in an egg receptor for a
rapidly evolving abalone sperm protein," Science 28 1 : 710-1 2, 1 998.
Turelli, M., "The causes of Haldane's Rule," Science 282: 889-9 1 , 1 998.
Ungerer, M. c., et al. , "Rapid hybrid speciation in wild sunflowers," Proceedings of
the NationalAcademy ofSciences 95: 1 1757-68, 1 998.

Chapter IX: On the Imperfection of the Geological Record

Asian, A., and Behernsmeyer, A. K. , "Taphonomy and time resolution of bone as­
semblages in a contemporary fluvial system-the East Fork River, Wyoming,"
Palaios 1 1 : 41 1-2 1 , 1 996.
Cooper, A. , and Penny, D., "Mass survival of birds across the Cretaceous-Tertiary
boundary: Molecular evidence," Science 275: 1 1 09-13, 1 997.
Fortey, R. A., Briggs, D. E. G., and Wills, M. A. , "The Cambrian evolutionary 'ex­
plosion' recalibrated," BioEssays 19: 429-34, 1 997.
Gallois, R. w., British Regional Geology: The Wealden District, 4th ed., HMSO,
London, 1 992.
Kidwell, S. M., and Flessa, K. w., "The quality of the fossil record: Populations,
species and communities," Annual Review ofEcology and Systematics 26: 269-99,
1 995.
Kumar, S., and Hedges, S. B., ''A molecular timescale for vertebrate evolution," Na­
ture 392: 9 1 7-1 8, 1 998.
Padian, K. , and Chiappe, L. M., "The origin and early evolution of birds," Biolog­
ical Reviews 73: 1-42, 1 998.
Xiao, S . , Zhang, Y., and Knoll, A. H., "Three-dimensional preservation of algae
and animal embryos in a Neoproterozoic phosphorite," Nature 39 1 : 553-55,
1 998.

Chapter X: On the Geological Succession of Organic Beings

Aldridge, R. J., and Purnell, M. A., "The conodont controversies," Trends in Re­
search in Ecology and Evolution 1 1 : 463-67, 1 996.
Briggs, D. E. G., and Crowther, P. R. , eds., Paleobiology: A Synthesis, Blackwell Sci­
ence, Oxford, 1 990.
Conway Morris, S., "Ecology in deep time," Trends in Research in Ecology and Evo­
lution 10: 290-94, 1 995.
Gokasan, E., et al. , "On the origin of the Bosphorus," Marine Geology 140:
1 83-99, 1 997.
Jackson, J. B. c., and Cheetham, A. H., "Tempo and mode of speciation in the
sea," Trends in Research in Ecology and Evolution 14: 72-77, 1 999.
McShea, D. w. , "Metazoan complexity and evolution: Is there a trend?," Evolution
50: 477-92, 1 996.
358 Further Reading

Maynard Smith, ]., and Szathmary, E., The Major Transitions in Evolution, W H.
Freeman, Oxford and New York, 1 995.
Rosenzweig, M. L., "Tempo and mode ofspeciation," Science 277: 1 622-32, 1 997.

Chapter XI : Geographical Distribution

Alley, R. B., Lynch-Stieglitz, J., and Severinghaus, J. P., "Global climate change,"
Proceedings ofthe NationalAcademy ofSciences 96: 9987-88, 1 999.
Brown, J. H., Stevens, G. C, and Kaufman, D. M., "The geographic range: Size,
shape, boundaries and internal structure," Annual Review ofEcology and System­
atics 27: 597-623, 1 996.
Cocks, L. R. M., McKerrow, W S., and van Staal, C R, "The margins of Ava­
lonia," GeologicalMagazine 134: 627-36, 1 997.
Cox, C B., and Moore, P. D., Biogeography: An Ecological and Evolutionary Ap­
proach, 5th ed., Blackwell Science, Oxford, 1 993.
Davis, G. W , and Richardson, D. M., eds., Medite"anean-1jpe Ecosystems: The
Function ofBiodiversity, Springer Verlag, Berlin, 1995.
Hamblin, W K., and Christiansen, E. H., Earth's Dynamic Systems, 7th ed., Pren­
tice Hall, Upper Saddle River, N.J., 1 995.
Spanier, E., and Galil, B. S., "Lessepsian migration: A continuous biological
process," Endeavour 1 5: 102-6, 1 99 1 .
Van Oosterzee, P. , Where Worlds Collide: The Wallace Line, Cornell University Press,
Ithaca, N.Y., 1 997.

Chapter XII : Geographical Distribution-continued

Frankam, R., "Do island populations have less genetic variation than mainland
populations?" Heredity 78: 3 1 1-27, 1 997.
Garrison, T., Oceanography: An Invitation to Marine Science, Wadsworth Publish­
ing, Washington, 1 996.
Grant, P. R., Evolution on Islands, Oxford University Press, 1 998.
Johnson, L. E., and Padilla, D. K., "Ecological lessons and opportunities from the
invasion of the zebra mussel Dreissena polymorpha," Biological Conservation 78:
23-33, 1 996.
Rossiter, A., "The cichlid fish assemblages of Lake Tanganyika: Ecology, behaviour
and evolution of its species flocks," Advances in Ecological Research 26: 1 87-252,
1 995.
Steadman, D . , "Prehistoric extinctions of Pacific island birds: Biodiversity meets
zooarcheology," Science 267: 1 123-3 1 , 1 995.
Thornton, I., Krakatau: The Destruction and Re-Assembly of an Island Ecosystem,
Harvard University Press, Cambridge, Mass., 1 996.
Whittaker, R. J., Island Biogeography: Ecology, Evolution and Conservation, Oxford
University Press, 1 998.
 Further Reading 359

Chapter XIII: Mutual Affinities of Organic Beings ; Morphology;

Embryology; Rudimentary Organs
Fong, D. w., Kane, T. c., and Culver, D., "Vestigialization and loss of nonfunc­
Annual Review ofEcology and Systematics 26: 249-68, 1 995.
tional characters,"
1 0 years," Sci­
Gale, M . D . , and Devos, K . M., "Plant comparative genomics after
ence 282: 656-59, 1 998.
Jackson, S . , and Diamond, J . , "Metabolic and digestive responses to artificial selec­
tion in chickens," Evolution 50: 1 638-50, 1 996.
Katz, L. A. , "Changing perspectives on the origin of eukaryotes." Trends in Research
in Ecology and Evolution 1 3: 493-97, 1998.
Logsdon, J . M . , and Doolittle, W. F. , "Origin of antifreeze proteins: A cool tale in
molecular evolution," Proceedings of the National Academy of Sciences us 94:
3485-1 87, 1 997.
McNab, B . K. , "Energy conservation and the evolution of flightlessness in birds,"
American Naturalist 144: 628-42, 1 994.
Pace, N. R. , ''A molecular view of microbial diversity and the biosphere," Science
276: 734-40, 1 997.
Shubin, N . , Tabin, c., and Carroll, S., " Fossils, genes and the evolution of animal
limbs, " Nature 388: 639-48, 1 997.
Tatusov, T. , Koonin, E. v., and Lipman, D. J., ''A genomic perspective on protein
families," Science 278: 63 1-37, 1 997.
Voss, S. R. , and Shaffer, H. B . , ''Adaptive evolution via a major gene effect: Paedo­
morphosis in the Mexican axolotl," Proceedings of the National Academy ofSci­
ences 94: 14185-9, 1 997.
Williams, R. w. , "Plant homeobox genes," BioEssays 20: 280-82, 1 998.

Interlude: Almost Like a Whale?

Boyd, R. , and Silk, J. B . , How Humans Evolved, W. W. Norton and Company, New
York, 1 997.
Harpending, H. c., et al. , "Genetic traces of ancient demography, " Proceedings of
the NationalAcademy ofSciences 95: 1961-67, 1 998.
Lewin, R. , Principles ofHuman Evolution: A Core Textbook, Blackwell Science, Ox­
ford, 1 998.
Modiano, D . , et al. , "Different responses to Plasmodium falciparum malaria in West
Mrican sympatric ethnic groups," Proceedings ofthe NationalAcademy ofSciences
93: 13206-1 1 , 1 996.
Ruff, C. B . , "Morphological adaptation to climate in modern and fossil hominids,"
Yearbook ofPhysical Anthropology 37: 65-107, 1 994.
Science 282: 1446-50, 1 998.
Pringle, H., "The slow birth of agriculture,"
Smith, M. w., et al. , "Contrasting genetic influence of CCR2 and CCR5 variants
on HIV- l infection and disease progression," Science 277: 959-65, 1 997.
Stoneking, M . , "Women on the move," Nature Genetics 20: 2 1 9-20, 1 998.
 INDEX

aardvark, 282, 3 1 2 algae, 1 3 5 , 1 47, 1 49, 289

Abu Simbel, 239 alleles, 1 08
acclimatization, 1 06-7 alligators, 78
acquired characteristics, 1 03-6 alveolates, 289
Adam, 1 69, 227 Alvin (submarine), 286
Africa: AIDS, 6-7, 1 2; copper, 98; dogs, Ambulocetus, 1 6- 1 7
26; genetic diversity, 3 1 2; metal America: East and West, 238-9;
extraction, 98; parasitic birds, immigrants, 5 5-6; New World,
1 50-1 , 1 77-8; use of antibiotics, 236, 324
90 American Kennel Club, 30, 3 1 , 1 44
Agassiz, Louis, 244 amino acids, 70, 224, 225
agouti, 240 Amish people, 326
agriculture, 23-4, 33-4, 3 1 0-1 1 ammonites, 1 92
AIDS, 2-1 6; appearance, 4; drugs, 7-9; ampicillin, 90
evolution, 2-4, 1 2-14, 34; HN Anatolian Shepherd dog, 29
mutation, 7, 8, 9; origins, 8-1 3 , Andean birds, 1 34
1 4- 1 5 ; progress o f illness, 5-6, angler fish, 82
8-9, 327-8; responses to, 4-5, anoles, 73
328; spread, 6-7 Anomalocaris, 202, 206
Ailsa Craig, 244 Antarctic perch, 29 1 , 307
Akbar, Emperor, 3 5 Antarctica: coal, 236; ice sheet, 220;
Alabama, legislation, 1 plant life, 243; separation from
Alaska: Bering Land Bridge, 324; Australia, 253
Glacier Bay, 247; plant life, 244 antelope, 46
albatrosses, 205 antibiotics, 89, 90, 98-9
alcohol, 106 antifreeze protein, 29 1 -2, 307
Aleuts, 252 ants: African driver, 1 5 5 ; Brazilian
362 Index

ants (conta )
rainforest, 1 53; Caribbean, 58; guts, 90, 1 39, 1 49, 257-8, 285;
Galapagos, 95; honeypot, 1 49, resistance to antibiotics, 89-92,
1 65 ; parasites, 1 48-9; relationship 1 39
with aphids, 1 49, 1 68; relationship Bagehot, Walter, xxviii
with fungi, 1 49-50; sterility, Bahamas, lizards, 73
1 65-6 Baikal, Lake, 268
aphids, 1 49, 1 59-60, 1 65 baleen whales, 296
Appalachian Mountains, 1 99, 250 Baltica, 253
apple maggo t fly, 125 bananas, 284
apples, 1 24-5, 1 69-70 Barbados: ants, 58; natural history,
Judhae� 286, 287, 288, 29 1 1 27-8
Archaeopteryx. 1 9 1 , 202. 203, 204 barnacles, 60, 82, 203
Arctic: cod, 292, 307; wolf, 25 Basilosaurus. 1 7
Ardipithecus. 322 basset hounds, 32
Argentina, catde, 58 bats, 47, 260, 282, 303
Aristode, 35, 80, 1 83 bears: black, xviii ; diet. 230; young, 1 59
armadillo, nine-banded, 1 58 beaver, 240
asses, 33 Beccari, Odoardo, 1 03
Aswan Dam, 200 beeches, 62, 249
atavism, 1 07 bees: cell making, 1 53-4; dance, 1 54-5;
Adantic: expansion, 250- 1 ; ridge, DNA, 1 55 ; hive society, 1 52-8,
25 1-2 339; island species, 265;
Adantis, 250, 256 pollination, 79, 1 76; sex and
atrophied organs, 302-7, 344 sterility, 1 62-3, 1 64-5; sight, 1 29,
aurochs, 228 1 30; sting, 339
Australia: age of rocks, 1 95, 1 99; beedes: distribution, 247; evolution,
ecosystem, 237, 238; extinct life­ 1 26; island species, 265; kinds of,
forms, 325; flighdess birds, 325; 48, 94; life with relatives, 1 59
human presence, 324, 325; Bell, Thomas, xxviii-xxix
imports, 56-7; jarrah, 65; Bering Land Bridge, 324
kangaroos, 236, 325; kwongan, Berkdey, George, Bishop, xxvii
237, 238; pre-Cambrian fossils, Berlin Wall, 1 79
206; separation from Antarcti� Bermuda, birds, 270
253 Bewick, Thomas, 28 1
Australopithecus, 322 Bible, 1 , 1 92, 1 93
Avaloni� 255 bighorn sheep, 94
axolod, 300 Biosphere Two, 235
birds: behavior, 145-6; chromosomes,
1 1 3, 1 8 1 ; classification, 283;
baboons, 1 5, 80 evolution, 1 20, 1 66-7, 204-5; first
bacteria: behavior, 145; classification, appearance, 204-5; Mediter­
289, 307-8; metal extraction, 98; ranean, 237-8; parasitic, 1 50-1 ,
DNA, 1 40, 226, 285; evolution, 1 78
89-92, 229; in aphids, 1 49; in Birnam Wood, 247
 Index 3 63

bison, European, 34 Cairo, sewers, 287

bizcacha, 240 Calgene company, 1 86
Black Sea, 2 1 7- 1 8 California: chaparral, 237, 238; condor,
Black Skull, 323 56; Death Valley, 248; ecosystem,
black smokers, 286 237-8; fossils, 1 97; Gold Rush,
blackberries, 304 1 97; plants, 94
blindness, 340; inherited, 326 Cambrian era, 205-6, 22 1
blood: clots, 1 32; groups, 3 1 4; pressure, Cambrian Explosion, 206-8
317 camouflage, 73, 84, 1 2 1
bloodhounds, 32 Camus, Albert, 4
blowflies, 1 90 Canada: fishing, 60; fish, 96
blue whale, 1 5, 239, 248 Canary Current, 24 1
blueberry maggot fly, 1 25 Canary Isles, 263
blue tits, 8 1 , 83, 1 47 cancer, 1 82, 1 87, 3 1 4
bone cells, 226 Canis familians, 2 5
Bonnet, Charles, 293 Canis lupus, 39
bonobo, 3 1 2 cannibalism, 1 63-4
Borametz, 1 86-7 Cape Verde Islands, 263
Bosphorus, 2 1 7 capybara, 240, 276
Bounty. HMS, 24 1 carbon, 1 3 , 1 9 5
bowerbirds, 84 carbon dioxide, 23 1 , 2 3 5 , 246
Boxgrove Man, 26 Caribbean: birds, 1 5 1 ; lizards, 73,
Bradford, William, 25 1 , 265
brains, 1 4 5 , 229, 294, 295, 3 1 8, 3 1 9 Carnegie, Andrew, 329
B�il: ants, 1 53; coffee, 63 carnivores, 230, 282
Bryan, William Jennings, 1 28 carpenter bee, 1 56, 1 57
Buccleuch, Duke of, 77 Catahoula leopard cowhog dogs, 29
Buffon, Comte de, 227 Catal Hiiyiik, 3 1 1
bulldogs, 30 cats: city, 264; classification, 282;
Bulo Burti boubou shrike, 43 deafness, 1 1 4, 1 1 5 ; edible, 2 1 ;
Burchell, William, 2 1 , 22 Galapagos, 9 5 ; genes, 28 1 ; island
Burgess Shale, 20 1 , 202, 206 populations, 264; London, 1 1 3 ;
Burgon, John, 1 93 Manx, 1 09; Persian, 1 70;
Burton, Richard, 268 tortoiseshell, 1 1 4, 1 1 6; viral
burying beede, 1 5 8 diseases, 1 5 ; wild, 48
Buss, Island o f, 256 came: defenses against flies, 1 3 1 ; DNA,
butterflies: camouflage, 1 2 1 ; 33, 1 38; domestication, 24, 33;
chromosomes, 1 8 1 ; classification, face recognition, 33; genetic map,
46, 47; flight, 1 24; Hawaii, 265; 28 1 ; in Americas, 58; selection, 33;
swallowtail, 1 27 teeth, 344; types of, 33
cave animals, 1 32, 303, 339-40
cells, 1 39-40; division, 1 82-3
C57Bl mice, 50, 5 1 chalk, 209- 1 2
Cabot, John, 67, 245 Challenger Expedition ( 1 872) , 286
cacti, 95, 238 change: gradual, 2 1 3-14; rate of,
3 64 Index

change (cont'd )
223-4; simultaneous, 22 1-3; cladistics, 280-3, 326
sudden, 22 1-3 clams, 299
Channel Tunnel, 209 Clark, William, 92
characteristics: acquired, 1 03-6; classification: Chinese, 275-6; cladistics,
reverting to, 1 07 280-3; comparative anatomy of
Chardonnay grape, 236-7 genes, 283-9; Darwin on, 346-7;
Charles II, King, 3 1 , 1 52 difficulties, 42-9; Linnaean system,
Checkpoint Charlie, 1 79 276-9; map of genes, 288-9 1
Chediak-Higashi Syndrome, 1 7- 1 8 Cleopatra's Needle, 1 98
cheese skippers, 1 90 climate, 63, 245-7, 271-2
cheetahs, 35 climbing plants, 304
Cherry-Garrard, Apsley, 298 clones, 96, 146
Chesapeake Bay, 220 coal, 236, 256, 3 1 3
chestnut trees, 247 coal mines, 7 1 , 224
chevrotains, 48 coconuts, 265
Chiang Kai-shek, 222-3 Cocos Island finches, 1 46
chickens, 74, 283, 305 cod, 67, 74, 9 5 ; Arctic, 292, 307
child mortality, 3 1 3- 1 4 coelacanths, 94, 283
Chile: beeches, 62; ecosystem, 237, 238; coffee, 63-4
human presence, 324; mattoral, collections, paleontological, 1 96-7
237, 238 collies, 29
chimpanzees: AIDS virus, 1 3-14; Colombia, coffee, 63
ancestors, 3 1 8; diversity, 324; Colorado River, 200
DNA, 3 1 9; human kinship, Columbus, Christopher, 58
1 3- 1 4, 284, 326; penis size, 86; comets, 2 1 8-20, 252
species, 3 1 2- 1 3 competition, 339
China: balance o f sexes, 3 1 5; condor, California, 56
classification, 275, 276; forensic conodonts, 2 1 5- 1 6
entomology, 1 90; fossil record, Constable, John, 1 72
207-8; hill tribes, 325; mice, 49; continents, 250-6
pandas, 43 Cook, Captain James, 259, 28 1
Chinese crested dog, 1 1 5 Copernicus, 1 93
chitin, 224 copper, 97, 98, 1 8 1
cholera, 3 1 1 , 3 1 4 copper Bower, 98, 1 79
chows, 32 Copts, 225
Christmas, 1 59, 1 73 corals, 22 1 , 230, 256
Christmas Island, 242 corgis, 29
chromosomes, 44, 1 1 0- 1 3 , 1 8 1-2 cormorants, 204, 305
Churchill, Winston, 3 1 9 Cornbrash, 1 94
cicadas, 1 75 , 279 corpses, decay of, 1 90-1
Cicero, 299 correlation of growth, 1 1 4- 1 5, 347-8
cichlids, 1 76-7, 268-70 Costa del Sol, 243
ciliates, 289 cougars, Texas, 53
cities, 97-8, 3 1 1 courtship, 83-4
 Index 365

cowbirds, 1 50 David, Pere Armand, 43

coyotes, 26 DDT, 1 1 3
coypu, 240 de la Beche, Henry, 1 93-4
crabs, 1 32, 206, 24 1 , 30 1 de Tocqueville, Alexis, 7 1
creationist movement, 1-3 deer, 46
crickets, 1 75 Defoe, Daniel, 7 1
crocodiles, 244, 283 Devil's Hole pupfish, 248
crows, 1 2 1-2 diatoms, 289
Cruft's, 24, 30 dingo, 26
cuckoo ducks, 1 50 dinoflagellates, 289
cuckoos, 1 50- 1 , 1 66, 1 68, 1 77, 1 78 dinosaurs: bird ancestry, 1 20, l 66,
Curse of India, 95 204-5; classification, 283;
Cuvier, Georges, 1 43, 1 93, 23 1 digestion, 209; distribution, 250,
Cyprus, elephants, 306 262; DNA, 225 , 226; end of, 209,
cystic fibrosis, 3 1 1 2 1 9-20; fossil record, 1 9 1 ;
lifestyles, 1 66-7; size, 229;
varieties, 253
Daguerre, Louis, 1 20 disease, 3 1 1 , 3 1 4; genetic, 326
daisies, 265 dispersal: during Glacial Period,
dandelions, 1 74 244-56; means of, 240-4
Darwin, Bernard, xxii distribution, geographical, 27 1-4, 333,
Darwin, Charles: correspondence, xxiii; 34 1-3
The Descent ofMan, 320, 326; divergence of character, 96-1 0 1 , 337-9,
family, xxi-xxii; on classification, 350
346-7; on common parentage, diversity, 1 33, 1 34, 3 1 1 - 1 2
307-8, 348-9; on difficulties of his DNA: amount o f, 226; ancient, 225-6;
theory, 1 4 1 -3; on embryology, bases, 1 35-6; changes, 1 32-3;
343-4, 348; on first crosses, decay, 306; discovery, 1 1 0-1 1 ; dog
1 88-9, 332-3; on geographical feces, 3 1 ; fingerprint, 3 1 1 ; fossil,
distribution, 27 1 -4, 333, 342-3; 224-6, 323; HIY, 5, 7; identi­
on geological record, 232-3, fication by, 43-5; maps, 1 1 1 ;
333-5, 34 1 -2, 348-9; on heredity, mitochondrial, 26, 39, 1 39, 325;
1 02-3; on hybrids, 1 88-9, 332-3; movable, 9 1 -2, 1 3 5-9; natural
on instincts, 1 67-8, 340- 1 ; on history of, 135; preservation, 225;
intermediate varieties, 333; on repair, 1 1 3, 138-9; repetition, 306;
marriage, 3 1 4- 1 5 ; on morphology, sequencing machines, 44;
347; on natural selection, 98-1 0 1 , venebrate, 1 38; whales, 1 7-1 9 ,
1 4 1-3, 33 1-2, 337-4 1 , 350; on 135
rudimentary organs, 343-4, 347, Dobermann pinschers, 32
348, 349; The Origin ofSpecies, dodo, 95, 329
xvii-xx, 2 1 4; prose style, xix-xx; dogs: bipedalism, 145; Chinese crested,
reading of Malthus, 57-8 1 1 5; classification, 27-8, 38-9,
Darwin, Roben, xxii 282; DNA, 26, 39; copulation, 78;
Darwin, Erasmus, xxii, 1 86 domestication, 23, 25-9, 32, 33;
dating, 1 94-6 evolution, 223; feral, 27, 38;
366 Index

dogs (cont'd )
genetic disorders, 32; herding, 29; emperor penguins, 297, 298
naked Turkish, 1 1 4, 1 1 5; acquired emus, 249, 253
characteristics, 1 03; sacred, 22; Engels, Friedrich, 57
selection, 32; wild nature, 38 Escherischia coli, 285
dogwood fly, 125 Eskdale, sparrowhawks, 77
dolphins, 1 5, 46 Eskimos (Inuit), 29, 325
domestication, 2 1-33, 335-6 evolution: use of term, 298-9
Douglas fir, 75 extinction, 92-5, 2 1 7-2 1
doves, 1 76 eyes, 1 28-3 1 , 1 42, 300- 1
dragonflies, 1 23, 1 30
du Chaillu, Paul, 1 4
ducks: flightless, 339; Hawaiian, 266; Fang people, 1 4
imprinting, 1 76; sexual selection, farming, see agriculture
1 77; species crosses, 1 7 1 , 1 78 fathers, age of, 1 1 2-1 3
Duffy blood group, 3 1 7 ferns, 244, 250, 260, 279
Dumas, Alexandre, 1 0, 67 fertilizer, 64
dung beetle, 247 figs, 260
dunnock, 1 5 1 finches: Cocos Island, 1 46; egg-dump­
ing, 1 78; Galapagos, 95, 1 46, 1 72,
1 73, 260- 1 , 270; zebra, 85
eagles, 65 Finland, 26 1
ears: development, 30 1 , 304; inner, 49, fire, 62
32 1 first crosses, 1 7 1-9, 1 8 8-9, 332-3
Earth: dating, 1 94-5; ice, 244; magnetic First World War, 2 1 1-12, 242
field, 252; rotation, 23 1 fish: ear development, 304; freshwater,
earthquakes, 2 1 6- 1 7 266-9; genes, 295, 296; hybrids,
earthworms, 293 1 87; ray-finned, 295, 296; sex, 82;
East Fork River, 200 sperm, 228
Eden, Garden of, 1 69, 248, 29 1 , 324 flatworms, 3 0 1
Ediacara fossils, 206, 207, 208 flax, 95, 1 05
Edison, Thomas, 1 20 flies, 1 06-7, 1 30, 1 3 1 , 1 75
Ein Mallah, burial, 26 flood legends, 2 1 8
electric fishes, 249 flowers: colors, 340; island, 265, 266;
elephant birds, 305 structure, 296-7
elephants: bones, 203, 227; flycatchers, 1 79
classification, 282; dwarf, 306; flying, 1 22-4
embryos, 299; mitochondria, 1 39; flying fish, xvii
reproduction, 65; sex pheromone, food chains, 64-5
29 1 ; swimming, 242; transitional Forbes, Edward, 273
links, 1 92; types of, 47; zoo, 37 fossil record: confusion, 20 1 ; evidence
Ellis Island, 5 5 , 68 of extinction, 93; incompleteness,
embryology, 297-302, 307-8, 343-4, 1 9 1-3, 334-5, 348-9; slow or
348 rapid change, 1 9 1 , 22 1 -2, 333-5,
embryos, fossil, 207-8 34 1-2
 Index 3 67

Foucault, Michel, 275-6 fossil,

224-6; homeobox, 295-6;
foxes: rabies,
258; selective breeding, 28 human, 323-4; map, 288-9 1 ;
Franco-Prussian War, 242 speciation, 1 8 1
Franklin, Benjamin, 57 genetic engineering, 1 86-8
Franklin, John, 245 genetically modified plants, 1 86-7
Freke, Dr., xix genetics: laws and principles, 1 03,
Frere, John Hookham, xvii, 309 1 06-7, 108-9; medical, 3 1 5- 1 6
friendship, 1 49-50 Geological Survey o f Great Britain,
Frobisher, Martin, 256 1 93-4
frogs: classification, 47; oceanic islands, geology, 1 93-6
260; sexual selection, 83, 84; George I, King, xxiv
varieties, 253 Germany, 42
fruit flies: acclimatization, 1 06; Ghanaian Times, 1 2
chromosomes, 1 1 0, 1 8 1 ; diversity, Giant's Causeway, 254
44; eyes, 300- 1 ; hearing, 30 1 ; Gibraltar, 239, 26 1
laboratory breeding, 1 80, 223-4; Gilgamesh, Epic of, 2 1 8
mobile DNA, 1 35-6; mutation gingko, 244
rate, 1 1 3, 224; natural selection, giraffes, 78, 1 3 1 , 1 83, 30 1
1 80; sex and death, 75; sexual Glacial Period, dispersal during,
selection, 83 244-56, 333
fruitbat, Dyak, 303 Glacier Bay, Alaska, 247
fugu, 1 35, 1 36 glaciers, 242-5
Fulani people, 3 1 7 Glen Canyon Dam, 200
fu ndamentalists, 92 goats: bipedalism, 1 1 5; DNA, 33, 138;
fungi, 46, 1 39, 1 49-50 domestication, 24; face
recognition, 33
Gondwanaland, 205, 253, 262
Galapagos: American species, 263-4; gonorrhea, 92
birds, 260- 1 , 270, 305; Darwin's gorillas: affinity to humans, 1 3-14, 326;
description, xxi; extinctions, 95, ancestors, 3 1 8; as food, 1 4; DNA,
2 1 7; finches, 95, 1 46, 1 72, 1 73, 3 1 9; lowland, 3 1 2; mountain, 46,
260- 1 , 270; frogs, 260, 263; ocean 3 1 2; penis size, 86; zoo diet, 35
currents, 1 73, 24 1 ; plankton, 64 Grand Canyon, 200
gales, 62 grape: Chardonnay, 236-7; climbing,
Galileo, xviii, 20, 1 93 304
Galton, Francis, 3 1 1 grasses, 1 79, 260
geese: Andean, 1 34; bar-headed, 1 34; grasshoppers, 247
first appearance, 204; Hawaiian, greenhouse effect, 23 1 , 246
266, 269; hybrid, 1 7 1 Greenland: age of rocks, 1 95 ; colonists,
genes: as particles, 1 07, 1 09; behavior, 56; fossil record, 208; temperature,
1 45-6; brains, 145, 320; 246
classification, 28 1-2, 283-6, groundsel, 1 72
288-9 1 ; crosses and hybrids, grouse: black, 73; red, 73, 78; snow, 60;
1 8 1-4; development and, 1 1 5-16; testosterone, 78, 80
disease, 3 1 3-14; diversity, 3 1 2; guava, 95
368 Index

Gulf Stream, 24 1 , 246 homeobox genes, 295-6

gulls, 40, 4 1-2, 54, 1 50, 1 76 Homer, 226
guppies, 74-5, 223-4 hominines, 322
Homo ereetus, 322
Homo habilis, 322
Habsburgs, 300 Homo sapiens, 322, 324
Haeckel, Ernest, 294, 298 honeycreepers, 266
hemoglobin, 1 33, 1 3 5 , 1 37, 1 38, 306, honeypot ants, 1 49, 1 68
3 1 6- 1 7 Hoover Dam, 245
hagfish, 2 1 5 hormones, 1 6 1 ; see also testosterone
Haldane's Rule, 1 8 1 horses: domestication, 33; edible, 2 1 ;
halibut, 67 evolution, 224; hooves, 30 1 ; in
Hallucigenia, 206-7 America, 58-9; striped, 340
hares: European, 239; snowshoe, 60; horsetails, 229
varying, 246 Hudson Rock, 4
Harvey, William, xxvii Hudson's Bay Company, 60
Hastings Rarities, 4 1 Hughes, Griffith, 1 27-8
Hawaii: bird extinction, 266, 269, 270; Human Genome Project, 3 1 5
frogs, 260; humpback whales, humans: brain, 3 1 8- 1 9 ; diversity,
1 7-1 8, 34; insects, 265; origins, 3 1 1-12; effects of farming,
262; plants, 265-6 3 1 0-1 1 ; evolution, 309- 1 0 ,
hawkweed, 1 09 321-3, 324-5 ; male nipples,
hawthorn fly, 125-6 303-4; migration, 324-6; size,
Hearne, Samuel, xvii-xviii 3 1 0, 3 1 1 ; tails, 303; walking,
heathers, 238 321-2
Hebrides, Outer, 65 humpback whales, 1 8, 34, 60, 1 47
hedgehogs, 282 Hutton, James, 1 93
Henry Patrick, 57 Huxley, Thomas Henry, 73, 145
Henry VI, King, 35 hybrid(s} : ape-human, 3 1 9; fertility,
Heraclitus, 2 1 3 1 79-84, 1 8 5-6; fish cancers, 1 87;
herbicide resistance, 1 88 legal status, 53; mongrel
hermaphrodites, 80, 88 comparison, 1 82-8; position of,
Herodotus, 1 98 1 2 1 -2; sterility, 1 7 1-2, 1 74-9,
herons, 267 332, 345; survival, 1 72-4; zones,
Himalayaeetus, 1 6, 1 9 51
Himalayas, 1 99 hyenas: diet, 230; spotted, 79-80
hippopotami, 1 9, 282 hypothalamus, 1 32
Hispaniola, ants, 58 hyrax, 282
HN (Human Immunodeficiency
Virus) : defenses against, 327-8;
HIV- 1 , 1 0- 1 6; HN-2, 1 0, 1 3-14; Iapetus, ocean of, 255
genes, 5, 7; origin of, 1 1-12; ice, 244-7
subtypes, 1 1 , 1 3 ; types, 1 0-1 1 , 14; Iceland: lava, 25 1 -2; origins, 252-3
see also AIDS ichneumonidae, 1 68, 339
hive-bee society, 1 52-8 inbreeding, 1 60, 3 1 2
 Index 36 9

incest, 83-4 distribution, 236; giant, 65; tree,

India, fossil record, 208 98, 1 76
Indonesian archipelago, 261-2 Kaposi's sarcoma, 5
Industrial Revolution, 7 1-3, 97-8 kelp, 289
insecticides, 79; resistance to, 1 1 3-14, Kennel Club of England, 3 1
315 Kent, chalk, 2 1 0
insects: classification, 284; island, 265; Kenya, antibiotics, 90
sexual selection, 80- 1 ; sight, 130 Kerguelen, cats, 264
instinct, 1 1 9, 145, 340- 1 ; cell-making, Ketengban people, 86
1 52-4; slave-making, 1 47-5 1 , Kew, Millennium Seed Bank, 243
1 67-8 Keynes, John Maynard, 230
insurance, 75 Killarney fern, 279
intelligence, 3 1 9, 327-8 kinetoplastids, 289
intercrossing, 87-8 King Charles spaniels, 3 1
intermediate forms, 1 4 1 , 1 9 1 , 223, 333, Kingsley, Charles, 1 22, 299
34 1 -2 kinship, 1 57-66, 307
International Commission on kiwis, 304
Zoological Nomenclature, 38, 39 Klagenfurt fountain, 202
Inuit (Eskimos), 29, 325 Knapp, Michelle, 2 1 8
investment, 74 Knight, Richard Payne, 309, 330
Ireland: ancient forest, 62-3; Knight, Thomas Andrew, 309
continental boundary, 255; snakes, Kolreuter, Josef Gottlieb, 1 84
236, 260 Koobi Fora beds, 222
iridium, 2 1 9 Krakatau: eruption, 258-60, 264;
Irish wolfhounds, 32 inhabitants, 260, 26 1
iron, liver levels, 3 1 8 krill, 202
islands, oceanic, 260-7 1 , 273, 342-3
Israel: j ellyfish, 238-9; mole rat, 1 3 1
Italy: feral dogs, 27; hill villages, 325 La Brea Tar Pits, 25, 202, 203, 23 1
La Plata, animals, 240
laboratory mice, 49-50
jackals, 230 Labrador dog, 28
Jackson Laboratory, 49-50 Lamb of Tartary, 1 86
Japan: moths, 72 lamprey, 294
jarrah, 65 lancelet, 2 1 5 , 295
Jefferson, Thomas, 92, 1 99, 227 language, 9-1 0
jellyfish, 238-9; freshwater, 268 Lathrop, Abbie, 49-50
Jews, 1 04 Laurasia, 253
Johnson, Samuel, 254 lead, 97, 1 79
jungle fowl, 305 leaf-cutter ants, 1 65-6
Jurassic Park (film) , 204, 225 learning, 1 47
Leavis, F. R, 329
leeches, 294
kangaroos: carnivorous, 325; Leigh, Isaac, 43-4
classification, 28 1 , 325; Lemuria, 262
370 Index

lemurs, 1 76, 262 Maida Island, 256

Leonardo da Vinci, 25 1 , 252 maize, 24, 33, 47
leopard frog, 47 malaria, 3 1 5- 1 6, 3 1 7- 1 8 , 320
Lewis, Meriwether, 92 Malawi, Lake, 268
Li Ch'un Feng, 2 1 9 mallard ducks, 1 78
lice, 3-4, 1 39 Malthus, Thomas, 57-8, 60, 66, 68, 76,
lichens, 1 49 1 48
Lilienthal, Otto, 99 mammals: classification, 28 1 -3; early,
Linnaeus, Carolus, 25, 39, 50, 278-9 220-2; genes, 295; size, 229
Linnean Society, xxviii mammoths, 202, 226, 306
lions, 35, 37, 1 59, 230 manatees, 282
Live Stock Journal and Fancier's Gazette, Manchester: growth, 7 1 ; industry, 7 1 ,
21 72, 97; moths, 72, 90
liverworts, 243 mangabey, sooty, 1 4
Lizard Peninsula, 1 94 Manx cats, 1 09
lizards: Caribbean, 73, 265; Marie Antoinette, 1 74
classification, 283; clones, 1 58; marijuana pollen, 243
DNA, 1 38; kinship, 1 58; leg market, 63
length, 73; monitor, 260 marriage: age at, 76; Darwin on, 3 1 4
lobster, 301-2 Marsham, Roben, 245
locusts, 243 marsupial mice, 1 6 1
logger-headed duck, 339 Marx, Karl, 57, 1 53, 329
London: black wildlife, 72; cats, 1 1 3; Mass, Catholic, 302, 307
East End, 289; Great Plague, 6; mastodons, 92, 227
growth, 7 1 -2; industry, 97; moths, Matteson, Edward E., 1 97
72; population, 3 1 1 ; Tower, 35; Mauna Loa, 262
Tube, 286, 3 1 1 ; Zoo, 34 Mayflower, 2 5 1
London, Jack, 38 Meander, River, 2 1 3- 1 4
Longfellow, Henry Wadsworth, 62 Mediterranean: ecosystems, 237-40;
lotus, 1 76 history, 239
Louis XIII, King, 276 megapodes, 1 66
Lucy (primate), 320-1 melanin, 1 1 5
lungfish, 93-4, 226, 283 melanomas, 1 82
lungs, 1 42 Melville, Herman, xxi, xxvii
Lyell, Charles, 92, 2 1 7, 345 memory, 1 03, 1 0 5-6
lynx, 60 Mendel, Gregor, xxiv, 49, 1 03, 1 07- 1 0
meningitis, 8 9
mental disorders, 3 1 9
macaques: society, 1 04 mesonychids, 1 6
Madagascar: birds, 305 ; plants and Meteor Crater, Arizona, 2 1 9
animals, 262; resistance to meteorites, 2 1 8- 1 9, 29 1
antibiotics, 92 Mexico: cattle, 58; Yucatan crater, 2 1 9,
Madeira: frogs, 260; land shells, 260, 220
26 1 ; seashells, 260, 26 1 mice: breeding, 48-9, 105, 1 1 4-1 5,
magnetic field, 252 1 82-3; chromosomes, 1 8 1 , 282;
 Index 371

diet, 1 05 ; diversity, 48-52; DNA, mussels: distribution, 249; freshwater,

1 38, 282; eyes, 300-1 ; fetus 43-4, 267i larvae, 299; zebra, 267
reabsorption, 1 64; genes, 1 06, 293; mutation, 1 1 1-1 5, 1 70-1
identification, 44; inbred, 1 60; myoglobin, whale, 70
laboratory, 49-50; marsupial, 1 6 1 ;
neophobia, 1 4 5 ; sexual strategy,
78; size, 1 1 4- 1 5 ; tabby, 1 1 5; Nagubago, Lake, 269
terrible, 46; varieties, 48-52; Napoleon Bonaparte, 1 93
white-footed, 36; wild races, 50 national identity, 42
midge larvae, 1 3 8 Natural History Museum, London, 1 96,
midwife toad, 1 37 227, 242-3
migration, human, 324-6 natural selection: action of, 70- 1 ; action
Milton, John, xxiii on the descendants from a
mimosa, 243 common parent, 96; circumstances
mines, 1 79 favorable to, 89-92; compared to
missing links, 1 20, 250, 323 man's selection, 98, 337-8;
Mississippi River, 2 1 3- 1 4, 2 1 6- 1 7 divergence of character, 96-7,
mitochondria, 26, 3 9 , 1 39, 288, 323, 338-9; extinction caused by, 92-5;
325 illustrations of its action, 7 1 -6;
moa, 205, 305 organs of extreme perfection,
Moby Dick, 1 7, 1 8, 20 1 27-3 1 ; organs of linle apparent
moles, 340; golden, 282 importance, 1 3 1 -3; origin and
mole rats, 1 32, 1 60, 1 62 transition of organic beings with
mongrels, 1 82-8, 332 peculiar habits and structure,
monkey flowers, 1 79 1 24-7; sexual selection, 76-87,
monkeys: number of species, 321-2; 3 1 4- 1 6; 336-7; speed of action,
training, 1 46; viruses, 1 3- 1 5 74
Moon: dating, 1 95; effect o n rotation of Neanderthals, 320, 323, 325
Earth, 23 1 nematodes, 46, 28 1 -2, 284
morphology, 292-7, 308, 326, 347 New Bedford, whale pon, 68
mosquitoes, 1 1 4, 3 1 6 New Guinea, dogs, 26
mosses, 243 New York, tuberculosis, 9 1
moths: island species, 265; melanic, New York Coffee, Sugar, and Cocoa
145; peppered, 72, 84, 9 1 ; sex Exchange, 63
pheromone, 29 1 New Zealand: arrival of man, 270, 324;
Moustache Gang, 84 flowers, 265; flighdess birds, 305;
Mozan, Wolfgang Amadeus, 1 47 imponed birds, 56-7, 264; snails,
mules, 1 7 1 96
mummy DNA, 225 Newfoundland: fish stocks, 67, 68;
Murray Collection, 89, 9 1 Maida Island, 256; Stone of Scone,
Mus domesticus, 50, 5 1 255
Mus musculus, 50, 5 1 Newton, Isaac, xxvii, 1 37
mushrooms, 284 Nicholas II, Tsar, 1 1 1- 1 2
music, 1 47 nicotine, 1 06, 1 27
muskrat, 240 Nile River, 200
372 Index

Nile perch, 270 parasites, 1 48-9, 229, 3 1 5- 1 6

nimravids, 23 1 Paris: Anatomy Museum, 277; Basin,
Nifio, EI, 1 73-4 23 1 ; Metro, 288; Zoology
nipples, male, 303 Museum, 276, 277, 278
Noah, 48, 2 1 8, 244, 25 1 parrots, 1 46, 1 6 1 , 204, 205
North Downs, 208-9 Passchendaele, batde ( 1 9 1 7) , 2 1 1 - 1 2
North Pole, 252 passenger pigeon, 20 1
Nova Scotia duck-tolling retriever, 3 1 Paul, aposde, 2 1 4
nucleus, 1 39-40, 289 peacocks, 80, 84, 86, 8 8
nutcracker, 145 peat bogs, 202
Pekingese, 29
penguins: distribution, 253; earliest
oaks, 238, 247 appearance, 204; emperor, 297,
Obsidian Pool, 285-6 298; fossil record, 205, 242
ocean: currents, 240- 1 ; floor, 2 5 1 -2; penicillin, 89, 92, 1 1 3
vents, 286 penis size, 86-7
Octopus Spring, 285 peppered moths, 72, 84, 9 1
oil, 224, 3 1 3 perch: Antarctic, 29 1 , 307; Nile, 270
oilseed rape, 1 88 perfection, 1 27-3 1 , 350
Okinotorishima, 1 97 pesticides, 1 1 3- 1 4, 3 1 5
Opabinia, 206 pets, 1 4- 1 5, 28
Oppian, 1 83 phallocarp, 86
orangutans, 1 3 , 326 pharaoh hound, 29
orchids, 65, 79, 1 75, 260 pheasants, 1 7 1 , 249
oryx, Arabian, 34, 37 Philadelphia Zoo, 35, 36
ostriches: distribution of flighdess birds, Philip, Prince (Duke of Edinburgh), 87,
249, 253; evolution, 1 03; fossil 250
record, 205; hybrid theory, 1 83; Phillips, John, 22 1
nesting habits, 1 5 1 phosphorites, 207
Owen, Richard, 227 pigeons, 22-3, 72, 86, 1 09, 205, 340
owls, 204 pigs: breeding, 1 57-8; domestication,
Oxford Botanic Garden, 1 72 24; Galapagos, 9 5 ; gene map, 28 1 ;
oxygen, 1 26, 1 33, 23 1 , 235 sexual misconduct, 1 02; trials of,
oystercatchers, 85, 205 2 1 -2
oysters, 299 Pilgrim Fathers, 67, 250- 1 , 264
Pillars of Hercules, 240
pineal gland, 1 3 1
Pakicetus, 1 6 pine trees, 228, 247
Paleozoic era, 22 1 pintail ducks, 1 78
Paley, William, 87, 1 30 pipistrelle bat, 47
palms, 47, 63 Pitcairn Island, 24 1
Panama, Isthmus of, 238 plague bacillus, 92
panda, giant, 43 plankton, 64
Pangaea, 25 1 , 253 plants, structure, 296-7
panthers, Florida, 53 plasmids, 9 1-2
 Index 373

Plato, 1 86 Rameses II, 239

platyfish, 1 82 rats, 95
platypus, duck-billed, 28 1 ray-finned fishes, 295, 296
Pleistocene, era, 244 red deer, 306
Pliny, 49, 50 Red Sea, 239
pneumonia, 4, 92 redstatt, 178
poisons, 97-8, 1 26-7 reed warblers, 1 50
polio, 3 1 4 reptiles, aerial, 1 23; classification, 283
pollen, dispersal, 243 retrovirus, 5, 1 9
pollination: Biosphere Two, 235; Rewa, Maharaja o f, 36
cooperation and parasitism, 1 47-8, rhea, 205, 249, 253
1 75-6; islands, 265, 266; male rhinoceros, 22; woolly, 202
genes, 79-80 rice, 23, 24, 1 83
Polynesia, rails, 270 Ritonavir, 8
Polyphemus, 226-7 �A, 5, 8-9, 1 3 , 140
Pope, Alexander, 227 Roanoke, colony, 56
poppies, 1 72, 242 Romanes, George, 1 66
population size, 56-6 1 , 3 1 3 Roosevelt, Kermit, 43
potato lice, 128 Roosevelt, President Theodore, 35
potatoes, 24, 1 85-6, 306 Rothschild, Lionel Walter, Lord, 270,
Priesdey, Joseph, xxii 286
primates: brains, 3 1 8; DNA, 3 1 9-20; Royal Society for the Protection of
evolution, 1 33; guts, 3 1 9; penis Birds. 1 78
size, 86-7 rudimentary organs, 302-7, 344, 347,
Przewalski's horses, 36 348
pseudogenes, 306 Russian Royal Family, 1 1 1 - 1 2
pterodactyls, 92 Rutherford, Lord, 53-4
puffer fish, 1 3 5 rye, 23
punctuated equilibrium, 223
Pyramids, 1 9 8-9, 2 1 2
Pyrenean Mountain dog, 29 St. Helena, 265
Pytheas of Massalia, 240 St. Hilaire, Geoffroy, 3 0 1 -2
St. Pierre et Miquelon, cats, 264
salamanders, 1 3 5 , 1 72, 226
quagga, 225 salmon, 78, 96, 296
Quayle, J. Danforth, 2 1 9 Salvinia, 59, 6 1
Quetzalcoatlus. 23 1 San Francisco, AIDS, 4 , 8, 1 4
saola, 46
scallops, 243, 249
rabbits, 56, 240, 282 Scopes, John, 128
rabies, 258 scorpions, 23 1
radioactive materials, half-life, 1 95 Scodand: continental movement,
ragwon, l 72 254-6; rocks, 2�5
rails, terrestrial, 270, 305 Scott, Captain Roben Falcon, 250, 298
Raleigh, Sir Walter, 56 sea anemones, 146
374 Index

sea dephants, 3 1 5 smallpox, 3 1 4

Sea Owle, 270 Smith, Adam, 1 53, 248
sea salt, 1 94-5 Smithsonian Institution, 39
sea scorpions, 255 snails: classification, 284; evolution,
seals, 239 222-3; fossil record, 1 96, 206,
seashells, 26 1 24 1 ; freshwater, 96-7, 222;
seaslugs, 80 Krakatau, 259; marine, 1 96, 22 1 ;
seaweeds, 208, 242, 289 sexual and asexual forms, 96-7;
seed: banks, 34, 243; dispersal, 24 1-3, shells, 224-5; struggle for
265 existence, 59-60
Sdborne, swifts, 59, 77 snakes: coloring, 84; DNA, 1 39;
septicemia, 89 Irdand, 236, 260; swimming, 242
sex cells, 8 1-2, 1 75 Snider-Pellegrini, Antonio, 25 1
sexual sdection, 76-87, 3 1 4-1 5, 337 snow grouse, 60
Seychelles warblers, 1 6 1 -2 Social Darwinism, 37, 38, 329
Shakespeare, William, 1 52, 270 Society for the Propagation of Horse
sharks, 1 33 Flesh as an Article of Food, 2 1
Shaw, George Bernard, 70-1 Somerville, Lord, 23
sheep: bighorn, 94; DNA, 1 39; Somme, Second Battle ( 1 9 1 8) , 2 1 1 , 2 1 2
domestication, 23, 24, 33; South Africa: ecosystem, 237, 238;
evolution, 1 2 1 ; face recognition, fynbos, 237, 238
33; sdection, 33; sheepdogs, 29; South Downs, 209
types of, 33 South Pole, 252
sheepdogs, 29 Soviet Union: AIDS, 6; whaling, 68
shipworm, 99 soybeans, 33
shrew, dephant, 282 sparrow: dusky seaside, 52; evolution,
shrimps, 1 24, 22 1 224; sexual dominance, 84
Shulgia, King, 35 sparrowhawks, 77, 83
Siberia: Bering Land Bridge, 324; species: breed and, 38; classification,
craters, 2 1 9 42-3; crosses, 1 1 9; definition, 53;
sickle-cell anemia, 3 1 6- 1 7 dominant, 338; extinct, 2 1 4- 1 5 ,
Sierra Club, 200 224-6; frontiers, 1 70, 1 80-1 ;
Signy Island, 243 geographical distribution, 242-3;
Silbury Hill, 225 modification of, 346-7; numbers
Silurian era, 233, 334, 335, 349-50 of, 46; reproduction, 338;
size: evolution, 226-7; island creatures, reversion to earlier characteristics,
305-6; sexual sdection, 82 1 07; sexual, 1 74; subspecies, 47-8;
skin color, 1 06 sudden appearance of groups of
skulls, 229, 230, 294 allied, 203- 1 2; variations, 1 07;
slave-making instinct, 1 47-5 1 , 1 68 varieties or, 48-9, 5 1-3, 334,
sled dogs, 29 337-8, 345-6; see alro hybrid
slime molds, 289 Speke, John Hanning, 268
sloths: giant, 1 99, 225 , 229; ground, Spencer, George, 1 02
202 Spencer, Herbert, 329
slugs, 80, 88, 1 64 sperm, 78-9, 8 1-2, 1 75, 228
 Index 375

sperm whale, 1 6 Tertullian, 57-8

spiders, 206, 23 1 testosterone, 78, 80, 1 59
spong�, 207, 229-30 Tethys Sea, 1 6, 253
spor�, dispersal, 243, 267 tetracycline, 90, 9 1 , 92
squash, cultivated, 23 Tham�, River, 2 1 0, 244
squirrels, 72, 83 thrush�, 1 68, 341
stag beetl�, 78 thymus, 1 3 1
starfish, 206, 284, 3 0 1 Tibet: dung beetle, 247
Stephens Island wren, 95 Tierra del Fuego, diet, 26
sterility: hybrids, 1 1 9, 1 7 1 -2, 1 74-9, tigers, white, 36
332, 344; insect societi�, 1 62, Time magazine, 32
1 64-6; selection, 1 70 toads: midwife, 1 37; varieti�, 253
sticklebacks, 96 tobacco, 1 06, 1 27, 1 84
Stone of Scone, 254-6 tobacco budworm, 1 27
Strachey, William, 270 tobacco hornworm, 1 06
stramenopil�, 290 Toft, Mary, xxiv
stromatolit�, 206 tomato�, 1 69-70, 1 83-8
struggle for existence, 59-68, 33 1 , 336, tortois�: giant, 9 5 , 325; swimming,
350 242
sturgeon, 296 transitional varieties, 1 20-4, 1 4 1 , 1 9 1 ,
subspeci�, 47 222-3, 333-4, 34 1 -2
Suez Canal, 239 transplants, 1 7 1
Sunda Strait, 324 tree frogs, 84, 249
sunfish, bluegill, 85 tree kangaroos, 98, 1 76
sunflower, 1 83 trees, 1 40, 265
sunlight, 1 06 trilobit�, 207, 254, 25 5-6
swans, 1 77 Trinidad, guppi�, 74
Swift, Jonathan, 306 trumpet vine, 304
swifts, 59, 77 tsetse fli�, 1 1 3, 1 3 1
swim bladder, 1 42 tuberculosis, 9 1
swordtail, 1 82-3 tubeworms, 286
syphilis, 1 1 3, 1 39 Turing, Alan, 1 20
Turkana, 222-3, 323
turtl�: Kemp's Ridley, 56; Tham�, 244
tails, human, 303 Tutankhamen, 4, 29
talapoin monkeys, 1 4 Twain, Mark, 66
Tanganyika, Lake, 267-70 Type, Unity of, 1 43
tapeworms, 1 37 typ�, suc�sion of same, 230-1
Taung Child, 20 1 , 320 typhus, 139
Taxol, 34
taxonomy, 42-8
Teilhard de Chardin, Pierre, xxviii Ua Huku, 270
temperature changes, 244-5 UK400 Club, 54
Tennyson, Alfred, Lord, 66, 259 underground life, 288
termit�, 1 53, 1 59, 1 63 Unenlagia, 204
376 Index

ungulates. 1 9 water skaters, 1 30

United States: AIDS. 4 . 6 . 7 . 8 ; apples. waterweeds , 289
1 25; Army. 1 1 2; Centers for watdes. 86, 238
Disease Control. 4; Corps of Army Waugh, Evdyn, 302
Engineers. 2 1 4; creationism. 1-2; weaverbird. 1 5 1
Department of Agriculture. 1 25. Webb. Sidney, 329
1 84; dog breeding, 24-5. 30; Wedgwood, Josiah, xxii
Endangered Species Act. 52; Fish weevils: black long-snouted, 6 1 ; in
and Wildlife Service. 52. 53; Food amber, 225
and Drug Administration. 1 86; Wells, H. G., 305
fruit flies. 1 36; New World. Wen Wang. Emperor, 35
235-6 whales: baleen, 296; beluga, 1 7; blue,
Unity of Type. 1 43 1 5-16, 239. 248; classification.
use and disuse. effects of, 1 03-6 282; discoveries, 46; DNA, 1 7- 1 9,
Ussher. Archbishop James. 1 93 1 35; drowned, 287; evolution,
1 5-18; extinction, 95; genes. 296;
humpback, 1 8, 34, 60, 1 47;
varieties: fertility, 1 1 9; mice, 48-9; or kinship, 1 9-20; Mediterranean.
species. 48-9. 5 1-3, 334-5, 239; origin. 2-3; retroviruses, 1 9;
337-8. 345-6; ttanSitional, 1 20-4 sperm. 1 6, 1 7; teeth , 304; whaling,
VelociraptIJr, 204 67-8; white, 1 7
vdvet worms. 207 wheat, 24, 65, 1 83
vents, ocean. 286 whelks, distribution, 249
vertebrates, 294 White. Gilbert, 59
Vian, Boris, xxii white smokers, 286
Victoria, Lake, 1 77, 268, 269 Whittington. Dick, 71
Victoria. Queen. 24, 286, 326 Wdde. Oscar, 1 86
Virgil, 226 wings, 1 2 1-4, 305-6
virgin birth, 228 wolves: Arctic, 25; carnivore role, 230;
Virginia: colony, 56, 57; fishery, 67 dog kinship, 23, 26-7, 37-9;
volcanoes, 1 9 5-6 rdationship with humans, 25-6;
voles, red-backed, 6 1 survival, 23
Voltaire, 1 98. 2 1 9, 220 woodpeckers, 1 27
vulrures: Andean, 1 34; types. 23 1 worms: brain absorption, 303;
classification, 284; fossil record,
208. 229; nematode, 46; strucrure,
Wake Island rail. 270 294; vdvet, 207
walleye fish, 1 63 Wrangel Island. mammoths, 306
Wallace, Alfred Russel , xxiv. xxix, wrens, 95, 1 68, 305
26 1-2, 3 1 0, 324, 325, 329
Wallace's Line, 262
warblers, 1 50. 1 6 1-2, 1 78 Yangtze fossil beds, 207-8
wasps: coloring, 84; family gender. 1 62; yeast, 282
sterility, 1 62 yellowhammers, 56
water buffalo. 33 Yellowstone National Park, 230, 285
 Index 377

yew trees, 247 zebra mussd, 267

Ypres, 2 1 1 zebras, 2 1
Yucatan crater, 2 1 9, 220 zebu, 33
zinc, 97, 98
Zoological Society of London, 34
zebra finches, 85 zoos, 34-7
 A B O UT T H E AUT H O R

STEVE JONES is professor of genetics at University College London. He regularly ap­

pears on British TV and radio and wrote and presented a hugely successful BBC TV
series called In the Blood. His previous books include The Language of Genes, win­
ner of the prestigious Rhone-Poulenc Science Book Prize. He lives in London.
 A B O UT THE TYP E

This book was set in Garamond, a typeface originally designed by the Parisian type
cutter Claude Garamond (1480-1 561). This version of Garamond was modeled
on a 1 592 specimen sheet from the Egenolff-Berner foundry, which was produced
from types assumed to have been brought to Frankfurt by the punch cutter Jacques
Sabon (d. 1 580) .
Claude Garamond's distinguished romans and italics first appeared in Opera Ci­
ceronis in 1 543-44. The Garamond types are clear, open, and elegant.