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The natural history of these islands is eminently curious, and well

deserves attention.
—Charles Darwin, Journal of Researches1

Darwin’s Islands

Galápagos is a group of 13 islands and numerous smaller islets

and rocks that straddle the equator 1000 kilometers west of South
America.2 Like a cluster of forgotten crumbs on a well-swept floor
they are isolated, arid, and barely visible on the world map. But insig-
nificant they are not. Together they form a surprisingly heterogeneous
world of their own, occupied by an astonishing diversity of endemic
plants and animals. Their story, from their volcanic birth under the
sea, to their colonization by organisms from distant lands, and the
transformation of these colonists into diverse new species as the is-
lands themselves change and diverge, is the story of evolution itself.
Viewed in this light it is no wonder that Galápagos played a pivotal
role in the development of Charles Robert Darwin’s famous theory
of evolution by means of natural selection. For when Charles Dar-
win (referred to hereafter as Darwin) visited Galápagos for five short
weeks in 1835 his experiences and observations there transformed
his way of thinking about the natural world. They revolutionized our
understanding of life on Earth.
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2 Introduction

Galápagos inspired Darwin, and both the place and the man have
inspired us. It was Darwin and his theory of evolution that brought
each of us to Galápagos in the first place. I arrived in 1973 with my
parents, Drs. Peter and Rosemary Grant, at the start of their famous
long-term study of evolution in Darwin’s finches. Greg arrived nine
years later as leader of the 1982 Cambridge Darwin Centenary Galápa-
gos Expedition. Over the years we pursued university degrees in biol-
ogy, conducted independent ecological research on various Galápagos
species, and worked as naturalists in the islands. Our fascination with
Galápagos and Darwin swelled. Then one day in early 1996, atop the
most active and pristine volcano in the Galápagos archipelago, it took
on a new form. The two of us were on the summit of Isla Fernandina
(Narborough Island), making field observations on Darwin’s finches
and pledging a lifetime of adventure together. Wouldn’t it be extraor-
dinary, we agreed, to retrace Darwin’s footsteps through Galápagos,
to see the places he observed, to compare the wildlife he described
with what can be found at the same sites today, and to investigate how
else the archipelago has changed. We had little idea, at that point, of
how closely we would be able to follow his path. How much physical
detail had he recorded of the sites he had explored? Was it enough to
reveal the treks he had taken, define the paths he had walked? Would
we gain insights into the development of his ideas by following in his
footsteps? The possibility was beguiling, the challenge irresistible.
We took our brainchild to England and nourished it with litera-
ture. Today Darwin’s published works and transcriptions of some of
his unpublished notes and manuscripts are accessible on the World
Wide Web (www.darwin-online.org.uk and darwinlibrary.amnh.
org), but this service did not exist at the time of our expedition. At
the Cambridge University Library (CUL) we paged through volume
after volume of published and unpublished Darwiniana. We feasted
most heavily on Darwin’s original manuscripts in the CUL Darwin
Archive, for we guessed that Darwin’s first-hand notes, and espe-
cially those pertaining to the geology of the islands, would contain
the greatest number of clues to where he had landed and explored.
We were astonished to find that Darwin’s Galápagos geology notes
had gone virtually unnoticed. A full transcription of those 115 man-
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Introduction 3

uscript pages became our first-priority assignment. We also tran-

scribed the Galápagos portion of Captain Robert FitzRoy’s log of the
Beagle in the Public Records Office at Kew, and scoured his charts of
Galápagos in the United Kingdom Hydrographic Office (UKHO) at
Taunton. Back in Galápagos, at the Charles Darwin Research Station,
we prepped and primed and plotted our course through the archi-
pelago, based on all we had learned in England. Then, on October 19,
1996, one hundred and sixty-five years after HMS Beagle graced the
waters of Galápagos, we found ourselves flying over a choppy ocean
between our home island of Santa Cruz (Indefatigable) in the center
of the archipelago and Darwin’s starting point of Isla San Cristóbal
(Chatham Island) on the eastern perimeter. Our dream was literally
taking flight, with the two of us happily on board in search of what it
meant to be Darwin in Galápagos.

Why Darwin?

Darwin is one of the most celebrated naturalists and influential per-

sons in the history of mankind. Born a naturalist and an ambitious
one, he spent his life seeking to explain the great diversity of life that
exists in all its colors all around us, and to solve what contemporary
luminary Sir John Herschel called the “mystery of mysteries”—how
new species come into being.3 Through his keen powers of obser-
vation, his interest in all aspects of the natural world, his ability to
reason, and his rigorous approach to study, Darwin came up with
answers that “shook the world.”4 His theory of evolution by means of
natural selection not only restructured the entire science of biology, it
revolutionized the way people perceive themselves. It created a whole
new world of understanding.
Darwin will forever be credited with evolution, but no one pre-
tends he was the first to think up the idea. Author Loren Eiseley, in
his prize-winning book Darwin’s Century, likens the discovery of
evolution to “a new continent” that was glimpsed through lifting
fogs by “master mariners” well before voyager–naturalist Darwin fi-
nally established its reality.5 The first of these known “mariners” was
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4 Introduction

Anaximander, a 6th-century Greek philosopher who suggested that

man had sprung from some form of aquatic animal.6 The germ of an
evolutionary idea may well have entered the minds of acute observers
of nature even before this, in preliterate civilizations. What is known
is that, in the western world, evolutionary speculations took off dur-
ing the 18th century’s Age of Enlightenment, when learning from na-
ture began to replace unquestioning acceptance of dogma and myth.
Darwin’s own grandfather Erasmus contributed to this movement.
There was even a theory of evolution when Darwin arrived on the
scene, but the chief explanation for how it worked—Jean-Baptiste La-
marck’s idea of the inheritance of acquired characteristics—failed to
gain widespread acceptance at the time.
Darwin changed all that. By marshalling evidence in support of
evolution, and coming up with a new and ultimately convincing ex-
planation for how it works—natural selection—Darwin demonstrated
the fact of evolution. He transformed evolution from a radical, some-
what illusory idea into a coherent scientific theory of consequence.
First published in 1859, Darwin’s masterwork On the Origin of
Species by Means of Natural Selection, or the Preservation of Favoured
Races in the Struggle for Life (hereafter referred to as The Origin of
Species) is Darwin’s argument for evolution. In it he proclaims that
all species are modified and diversified descendants of a single com-
mon ancestor and reasons that species change because of the follow-
ing natural laws:7
1. Heritable variation exists among the individuals that comprise
populations of species.
2. There is a struggle to survive and reproduce, in which individu-
als compete for limited resources.
3. More individuals are born than survive this struggle for
existence.
4. Those individuals with variations that help them survive the con-
ditions of their local environment are the ones most likely to re-
produce and pass on their winning traits to the next generation.

As environments are neither uniform nor static it logically follows

that as populations move into new environments, whether through
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Introduction 5

time or space, they change. They form new species that diverge and
diversify in an ever-branching tree of life.
As asserted by evolutionist Julian Huxley, Darwin’s epochal book,
“altered the substance and the direction of human thought more
profoundly than any other publication of the age of print.”8 It trans-
formed the prevailing view of the world as a static, biblical place in-
habited by species created by God according to His will and design,
into a dynamic landscape run by nature and subject to the rules of
change. Ernst Mayr, another great evolutionist, goes as far as to say
publication of The Origin of Species “almost single-handedly effected
the secularization of science.”9
This is not to say that Darwin’s theory was accepted overnight. Dar-
win encountered major objections from scientists who doubted the
validity of natural selection as an explanation for evolution. Over the
15 years following the first publication of The Origin of Species, Darwin
revised his book five times to answer criticisms and clarify his argu-
ments.10 It took another 65 years for biologists to fully come to terms
with natural selection as a cause of evolution.11 Nonetheless, during
his lifetime Darwin succeeded in bringing about a scientific move-
ment that changed research programs and transformed the direction
of science. In the words of science historian Peter Bowler, Darwin’s
“great achievement was to force the majority of contemporaries to re-
consider their attitude towards the basic idea of evolution . . . despite
the fact that many found natural selection unconvincing.”12
The Origin of Species also made a huge, divisive impact outside the
scientific community. The working classes, eager to “wrest control . . .
from the old landed interests” hailed Darwinian evolution (or “Dar-
winism” as Darwin’s theory and corrupted interpretations of it be-
came known) as an endorsement for social progress and reform.13
Members of the ruling class, those whose social and political position
depended on maintaining the status quo, felt threatened. And because
Darwin’s theory offered secular answers for sacred questions—What
is life? Who are we? Where do we come from?—it kicked up a storm
of outrage among religious traditionalists. Even today, Darwin’s the-
ory of evolution remains contentious in society for its religious, so-
cial, and cultural implications.
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6 Introduction

Brilliant, ingenious, audacious, Darwin is loved and revered by

many, hated by others. Above all he fascinates. As Ernst Mayr once
said, “no biologist has been responsible for more—and for more
drastic—modifications of the average person’s worldview than
Charles Darwin. . . . Almost every component in modern man’s be-
lief system is somehow affected by Darwinian principles.”14 Indeed,
Darwin matters so much to human thought that he has been, and
continues to be, the subject of intense scrutiny by scientists, histo-
rians of science, philosophers, psychologists, and theologians alike.
A substantial body of work has been written on Darwin and his
life and seemingly no stone has been left unturned in an effort to
understand the man who redefined the science of biology and the
path he took to do so. In addition to a perennial supply of scientific
and historical papers produced by members of this “Darwin Indus-
try,” several outstanding books have been written by some of these
same scholars. They include Janet Browne’s comprehensive two-
volume biography15 that describes Darwin’s life, analyzes his achieve-
ments, and illuminates how his environment made him who he was;
Adrian Desmond and James Moore’s work of similar scope;16 and the
more concise, but insightful overviews of Richard Keynes,17 Randal
Keynes,18 Niles Eldredge,19 and David Quammen.20 These books ex-
amine parts or all of Darwin’s life—his childhood, his famous voy-
age round the world on HMS Beagle, his return to England, and the
subsequent steps that led him to become the most influential scientist
of the century—in the illuminating context of the Victorian era, the
social circles in which he moved, and the scientists with whom he
corresponded. Other authors adopt more oblique but equally reveal-
ing perspectives from which to explore the subject of Darwin and
his intellectual journey, angles that reflect their own particular in-
terests and expertise. For instance, Sandra Herbert investigates the
key role that geology played in Darwin’s thinking.21 Edward Larson22
and Peter Bowler23 choose the history of evolutionary thought as the
platform from which to examine Darwin and his theory.
By looking at Darwin’s life from historical, philosophical, and sci-
entific points of view, these authors paint a detailed picture of Dar-
win’s development as he changed from an ordinary creationist to an
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Introduction 7

extraordinary evolutionist. They vary on interpretation and the de-

gree to which various moments in Darwin’s life were important to
his theory. Yet all agree on one thing; Galápagos was key to Darwin’s
conception of evolution.

Why Galápagos?

Darwin spent only 5 weeks in Galápagos, a minute fraction of the

248-week voyage of HMS Beagle, yet his experiences in the archi-
pelago were of disproportionate importance to the development of
his scientific thinking. Quite simply Galápagos convinced Darwin
of evolution.24 It did not happen overnight. Rather, it took several
years for Darwin to fully recognize the significance of his Galápa-
gos observations. Nonetheless, his appreciation of three fundamen-
tal features of Galápagos—its isolation, the geographical distribution
of its organisms, and the affinities of these organisms to species on
the South American continent and between the islands themselves—
ultimately persuaded him that species could change. When he first
began putting his ideas on transmutation to paper in 1837, Darwin
declared the “S. American fossils —& species on Galapagos Archi-
pelago . . . [are the] origin (especially latter) of all my views.”25 He
maintained his emphasis on the importance of Galápagos, declaring
to fellow evolutionist Alfred Russel Wallace 22 years later, while pre-
paring for publication The Origin of Species, that the “Geographical
Distrib[ution] & Geographical relations of extinct to recent inhabit-
ants of S. America first led me to [the] subject [of evolution]. Espe-
cially case of Galapagos Islds.”26
Galápagos was not the only factor in Darwin’s conception of evo-
lution. As Darwin pointed out in the quotes above, his recognition of
the significance of Galápagos was influenced by his earlier observa-
tions on the mainland of South America and especially by his study
of geographical barriers as they relate to species distributions. Nor
was Galápagos central to the development of Darwin’s theory of evo-
lution. Three years after his visit to Galápagos Darwin hit upon the
idea of natural selection as an explanation for evolution, not from
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8 Introduction

contemplating Galápagos organisms but primarily from studying

domestic breeds and reading Thomas Malthus’s essay on human
population theory.27 He advanced his theory by spending decades
conducting original research and gathering facts on various groups
of organisms outside Galápagos, most notably on domestic pigeons,
barnacles, orchids, bees, and worms. Nevertheless, Galápagos was the
keystone of his conversion and the foundation for his understanding
of evolution. It was the Galápagos organisms he observed and col-
lected, and his recognition of their affinity to organisms found on
the South American continent and their representation as similar but
distinct species on the different islands of Galápagos, that persuaded
Darwin of the mutability of species and against their miraculous cre-
ation. As one author avows, “It cannot be maintained that without
Darwin the theory of evolution would not have come into being, but
it can be insisted that had Darwin not taken the voyage of the Beagle
to the Galápagos, it would have been seriously delayed.”28
That Galápagos was important to Darwin is a well-accepted fact.
The particulars—what about the islands stimulated Darwin, and how,
when, and why they influenced his ideas on evolution—are generally
not so well known. Indeed, they are the subject of ongoing study and
exciting debate. Like a pendulum the question of whether Darwin
was thinking in evolutionary terms while he was in Galápagos swings
back and forth, each oscillation provoking new investigations and
providing fresh insights into Darwin’s Galápagos experience. In 1966
Julian Huxley wrote, “It was on the Galápagos in the early autumn of
1835 that Darwin took the first step out of the fairyland of creation-
ism into the coherent and comprehensible world of modern biology,
for it was here that he became fully convinced that species are not
immutable—in other words, that evolution is a fact.”29
While correct in essence, Huxley’s words (and similar statements
from other authors30) oversimplify the process of Darwin’s conver-
sion to evolution, and misleadingly suggest a eureka-like moment
taking place in Galápagos. This has since been shown not to be the
case. By using Darwin’s spelling mistakes and changing orthography
as a means of dating his notes, historian Frank Sulloway has mapped
Darwin’s evolving attitudes. He has shown that Darwin first wrote
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Introduction 9

about the instability of species nine months after leaving Galápagos

and has argued that Darwin did not become convinced of the muta-
bility of species until he was back in England.31 Sulloway has made
great strides in elucidating the role Galápagos played in Darwin’s
thinking, and in debunking the myths of what Darwin did and did
not do in the islands.32
Yet the pendulum swings. While it is no longer contended that
Darwin reached an evolutionary standpoint while he was in Galá-
pagos, just how close he got, and how important Galápagos was, is
still under scrutiny. Darwin historian Sandra Herbert puts it this way:
“[A]t first Darwin’s experience in the Galápagos Islands was over-
emphasized as the turning point in his arrival at a transmutationist
position. Now that most readers have learned that Darwin did not
become an evolutionist until 1837, the Galápagos experience is pos-
sibly credited with too little.”33 After all, she reminds us, it was in
Galápagos that Darwin “recorded patterns of variation among spe-
cies on the islands . . . and that, ultimately, pushed him across the line
to a transmutationist position.”34
Darwin’s Galápagos experience was multifaceted and the effects
it had on his thinking multitiered. Thanks to the continued research
of scientists and Darwin historians, there is a widening apprecia-
tion of what Galápagos meant to Darwin, and when it did so. The
efforts of biologist Richard Keynes35 and botanist Duncan Porter36
have been particularly constructive. By examining Darwin’s zoo-
logical and botanical notes and specimens they have identified the
many Galápagos species that influenced Darwin’s developing ideas
and have shed light on how they did so. Paul Pearson37 and Sandra
Herbert38 have helped expose one of Darwin’s greatest geological dis-
coveries in Galápagos, and have suggested that Darwin’s developing
ideas on evolution were closely tied to his concurrent theorizing on
the origin and diversity of rocks. Darwin historian David Kohn and
colleagues have recently elucidated the early origins of Darwin’s in-
terest in variation (a key element of evolution by means of natural
selection) and have suggested that Darwin’s collecting activities in
Galápagos were influenced accordingly. They also suggest that al-
though Darwin was not thinking in evolutionary terms while he was
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10 Introduction

in Galápagos, he was operating within an intellectual framework to

allow him to recognize varieties as incipient species, and to appreci-
ate transmutation, soon after.39
In this book we take this rich cornerstone of Darwin’s career as
an evolutionist and put it under the magnifying glass. We examine
Darwin’s physical journey through Galápagos in unprecedented de-
tail. By taking a step-by-step tour of his visit we demonstrate just
how influential and inspirational it was. Using new facts drawn from
Darwin’s original unpublished notes, fresh insights from his sketches
and publications, the studies of modern-day scientists, the analyses
of Darwin historians, and our own intimate knowledge of the place,
we explore how Galápagos shaped Darwin and his theory and how it
defined the legacy he left behind. We show how Darwin’s Galápagos
experiences catalyzed his thoughts on evolution, how his Galápagos
collections provided him with persuasive evidence to support his
theory, and how his Galápagos observations fueled his speculations
and motivated some of his later experiments. In doing so we shed
light on the whole canvas of Darwin’s life and work.

Into the Wild

It has been said that the love of the chace [sic] is an inherent delight
in man,—a relic of an instinctive passion. —if so, I am sure the plea-
sure of living in the open air, with the sky for a roof, and the ground
for a table, is part of the same feeling. It is the savage returning to his
wild and native habits. I always look back to our boat cruizes [sic] &
my land journeys, when through unfrequented countries, with a kind
of extreme delight, which no scenes of civilization could create. —I
do not doubt every traveller [sic] must remember the glowing sense
of happiness, from the simple consciousness of breathing in a foreign
clime, where the civilized man has seldom or never trod.
—Charles Darwin, Beagle Diary40

Our Beagle on the first day of our own voyage of discovery was a
five-passenger Piper Aztec air ferry, but it metamorphosed over the
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Introduction 11

following weeks into whatever means of transport we could find and

afford: a fishing boat, the crews quarters of a cruise ship, a municipal
launch, a dinghy, our legs for walking, our arms for swimming. At one
point we really did travel on the Beagle. It was the current research
vessel of the Charles Darwin Research Station, and the sixth vessel of
that name to grace the waters of Galápagos since HMS Beagle. Dar-
win was our constant companion, speaking through his field notes,41
geology notes,42 zoology notes,43 ornithology notes,44 plant notes,45
diary,46 and specimen lists,47 copies of which we had obtained and, in
some cases (most notably the geology notes), transcribed in England.
Captain FitzRoy was also a commanding presence, showing us, from
the bearings and anchorages identified in his logs48 and charts,49
where to hit the shore and strike inland. Instead of spirit bottles and
collecting bags we carried cameras, a GPS (Global Positioning Sys-
tem) receiver 50 and compass, and a stack of photocopies.
It was already known, from published accounts of the voyage,
which four islands Darwin visited—San Cristóbal (Chatham), Flo-
reana (Charles), Isabela (Albemarle), and Santiago (James)—and
roughly where on each he explored. Frank Sulloway published a
rough outline of Darwin’s course through the islands but the details
of where Darwin landed and explored remained vague.51 Each is-
land is topographically and ecologically heterogeneous, its habitats
varied, and the distribution of its organisms uneven. Where Darwin
walked determined what he saw and what he saw influenced what
he thought. For the purposes of our study it was important to de-
termine as closely as possible where Darwin stepped ashore and to
define his movements inland and along the coast. Only in this way
could we compare the natural history Darwin observed with what
can be found in the same sites today. We could identify the sources
of his geological insights, the exact features that triggered his under-
standing of the physical processes governing evolution in Galápagos.
By gleaning clues from Darwin’s original notes, time and again we
were able to figure out where Darwin walked, the land formations he
examined, and the route he took to reach them. Indeed, on the very
first day of the expedition, from clues in Darwin’s forgotten geology
notes and the Beagle logs, we identified the cove at Cerro Tijeretas
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12 Introduction

(Frigatebird Hill) as Darwin’s first Galápagos landfall.52 For 165 years

this doormat of the most famous stopover on the voyage of the Beagle
and in the history of evolutionary thought had remained unidenti-
fied. And now we had elucidated its location. Never again would Dar-
win’s first landing spot be known only vaguely as somewhere on the
southern end of Isla San Cristóbal (Chatham Island). It was a confi-
dent start to our expedition, and one that gave us no modest feeling
of accomplishment. But not all the sites were so easy.
Despite traveling far faster than the Beagle, we took eight weeks
to cover the area Darwin did in five. To determine the limits of Dar-
win’s excursions we explored widely. As we pushed our way through
thickets, tripped over knife-like lava, trod sun-scorched beaches, and
clambered up crater after crater, we marveled constantly at Darwin’s
stamina. Not only were his hikes often long, some of the terrain was ex-
ceedingly rough. If he ever fell he never complained. We, on the other
hand, went home sporting a few new scars. Of course, Darwin was
only 26, and in fine form from his recent treks through South America!
One of the fascinating things about Darwin was his extraordinary
power of observation and reasoning. He noticed “things which easily
escape attention,” questioned them, and endeavored to understand
them.53 Fortunately for us, he wrote down such thoughts, albeit in
note form. Retracing his footsteps was like taking a guided walk
through the countryside “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.”54 Only instead of worms it was reptiles and instead
of through damp earth, it was over arid lava.
In the prologue of his book Fossils, Finches and Fuegians, author
Richard Keynes wrote, “When you have transcribed several hundred
thousand words of [Darwin’s] writings, concerned with places . . . not
too greatly changed 160 years later, you may once in a while almost
feel that you are talking to him.”55 Compared to Darwin’s great-
grandson we have transcribed but few words (and thank goodness,
for Keynes’s transcriptions made our footwork that much easier), but
we have certainly read and relived many. How many times did I not
look at Greg as he crouched to identify a plant, reach up to measure
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Introduction 13

the thickness of an ancient stream of lava visible in a cliff face, or hike

off toward a distant hill, and imagine, with allowances for costume,
I was seeing Darwin himself. Nine years later I was awarded a sense
of déjà vu when the British Broadcasting Corporation (BBC) filmed
Greg acting as Darwin’s double while we were working as on-site
script consultants for their three-part television series, Galápagos.
This time Greg was dressed for the part, with straw hat, waistcoat,
and hob-nailed boots, but because of his dark hair, brown eyes, and
beard—Darwin was fairer, blue-eyed, and possibly clean shaven in
1835—Greg was filmed from behind and at a distance, while another
man played Darwin close up.
During the expedition, we often caught ourselves asking Darwin’s
ghost out loud which way he had gone, and admonishing (to put it
mildly) the great man for not having made it clearer in his notes. For
not all the sites were named in Darwin’s day as they are today, and
some were not named at all. We had to feel our way along by match-
ing up landmarks with Darwin’s imagery. While our task was helped
enormously by the fact that Darwin was a geologist and generous in
his descriptions of land formations, his footsteps became faint wher-
ever outstanding geological features were lacking. Retracing Darwin’s
route, from hint to sometimes ambiguous or inconsistent hint, became
a veritable treasure hunt, frustrating at times, but infinitely reward-
ing in the process. Nor did we limit ourselves to walking in Darwin’s
own footsteps; we readily branched out to explore formations that
Darwin, having lacked the time and means to examine himself, had
nonetheless described from the decks of the Beagle. Islote Tortuga
(Brattle Islet) and Punta Cristóbal (Point Christopher) on Isla Isabela
(Albemarle Island) are two such places.
In 1996, the same year as our own expedition, an attempt was
made to follow Darwin’s route through mainland South America on
horseback. “All over the continent,” the rider wrote, “I found that ur-
ban growth made following Darwin’s precise routes dangerous and
sometimes impossible, it being no joke to ride a fairly wild horse
along main roads or through urban sprawls.”56 Other places had been
“swallowed” up by resorts. Fortunately Galápagos has been largely
spared this development. Several of the sites that Darwin visited now
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14 Introduction

have a National Park trail running through or near them, but the only
settlement from Darwin’s day (on Isla Floreana/Charles Island) actu-
ally has fewer residents today than it did then. Unfortunately, this
does not mean that Galápagos has been left untouched. Humans, and
the plants and animals brought with them, have wreaked havoc on
many of the islands, causing the local extinction of several endemic
species. In many places we came across invasive plants choking out
the native vegetation and saw the vandalistic signs of introduced
insects and mammals. Nowhere was the destruction more appar-
ent than in the highlands of Isla Santiago (James Island). Forests of
endemic Scalesia trees (Scalesia pedunculata) that once covered the
summits had been transformed into vast meadows by feral goats.
We saw herds of hundreds of them running about the hills. In stark
contrast, the native herbivores—the Galápagos tortoises that Darwin
had reported swarming through the damp undergrowth of the high-
lands—were few and far between. Since our expedition, and over the
first few years of the 21st century, the goats (and introduced pigs and
donkeys) have been eradicated from the island, and the native vegeta-
tion has started to reinvade. Although there are now new alien con-
tenders on Santiago—noticeably the invasive hill raspberry (Rubus
niveus) and the aggressive paper wasp (Polistes versicolor)—there is
much to rejoice about the natural state of Galápagos as a whole, for
most of the wildlife described by Darwin is still there, albeit in re-
duced population numbers. Despite the increasing threats of a grow-
ing human population, “Darwin’s Islands”57 are still, for now, worthy
of their epithet.

Darwin’s Legacies

The mind is its own place, and in it self [sic]

Can make a Heav’n of Hell, a Hell of Heav’n.
—John Milton, Paradise Lost58

Darwin lives on in Galápagos as nowhere else. The archipelago’s unique

plants, animals, and landscapes are responsible for its modern ranking
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Introduction 15

as a Natural World Heritage Site, a Biosphere Reserve, and a National

Park, but it is Darwin’s association with this same fauna, flora, and ge-
ology that gives Galápagos its iconic status. Darwin’s importance to the
islands is reflected in the numerous species and places in Galápagos
that bear his name (see appendix 2). It is also advertised in the titles of
a local research institution (Charles Darwin Research Station), a road
(Avenida Charles Darwin), a tour boat (M/V Darwin), and various
other businesses (Darwin Hotel and Charles Darwin Travel Agency,
for example) that operate in Galápagos. Indeed, the prominence of
Darwin’s name in Galápagos attests to the fact that the islands are “in-
exorably linked to the evolutionary views of Charles Darwin.”59
Darwin revolutionized not only science and the way we perceive
ourselves but also the way we view Galápagos. In the early days of its
discovery the archipelago was commonly regarded (or at least por-
trayed) as a “monstrous” heap of islands upon which “God had show-
ered stones”60 in some places, “brimstone”61 in others. They were
inhabited by “creatures . . . the ugliest in Nature,”62 “deformed fiends”
and “devils”63 and “imps of darkness,”64 all readily sacrificed by the
only men who dared frequent Galápagos waters—blood thirsty pi-
rates, whalers, sealers, convicts, and men of war. American novelist
and mariner Herman Melville summed up the gloomy impression of
these early visitors when he wrote:
Take five-and-twenty heaps of cinders dumped here and there in
an outside city lot, imagine some of them magnified into moun-
tains, and the vacant lot the sea, and you will have a fit idea of the
general aspect of the Encantadas, or Enchanted Isles . . . Man and
wolf alike disown them. Little but reptile life is here found: tor-
toises, lizards, immense spiders, snakes, and that strangest anom-
aly of outlandish nature, the iguana. No voice, no low, no howl is
heard; the chief sound of life here is a hiss.65

Darwin began exploring the islands with some of the same preju-
dice. But as he traveled through the archipelago, then reflected on all
he had seen, he realized that far from being dismal dumps of dust,
the islands had a fascinating tale to tell. Using the language of science
Darwin revealed the beauty of the archipelago’s youthful landscapes
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16 Introduction

and unique organisms. He lifted the veil of ignorance that had cursed
Galápagos for the past three hundred years, and exhibited them in
a new, secular light. In so doing he reclaimed the islands and their
inhabitants from human condemnation and bequeathed them long-
lasting fame and a life-saving future.
He did this in two ways. First, he revealed that the different islands
of Galápagos are tenanted by different organisms of common ancestry
adapted to different geographically isolated niches, and then issued a
challenge for future scientists to “determine to what extent the fact
holds good.”66 Numerous scientists picked up the gauntlet, expand-
ing on Darwin’s baseline collection and carrying out field studies to
further understand the biodiversity of the archipelago.67 Galápagos
has now become a magnet for evolutionary biologists who, thanks
to modern genetics, continue to reveal a greater amount of diversity
in the islands than Darwin could ever have imagined. It is thanks to
Darwin that Galápagos is now world famous and treasured as a “liv-
ing laboratory.”
Secondly, Darwin did the eminent service of inspiring the conser-
vation of the islands. For he anticipated “what havoc the introduc-
tion of any new beast of prey must cause in [Galápagos], before the
instincts of the aborigines become adapted to the stranger’s craft or
power.”68 His words were heeded by a handful of scientists who, on
the centenary of Darwin’s 1835 visit, initiated one of the first moves to
protect the wildlife of Galápagos.69 By 1959, one hundred years after
the first publication of The Origin of Species, two organizations com-
mitted to conserving the Galápagos ecosystems were up and running.
The Charles Darwin Foundation was founded to promote scientific
research and environmental education about conservation and natu-
ral resource management in the archipelago. The Galápagos National
Park Service dedicated itself to the protection by law of 97 percent of
the landmass of the archipelago.
The efforts of these two institutions, with the support of various
auxiliary conservation organizations,70 have, to date, successfully man-
aged to keep Galápagos one of the most pristine oceanic archipelagos
in the world. It has not been easy. Tightly regulated nature-oriented
tourism, introduced in 1967 with the hopes of it being a sustainable
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Introduction 17

economy compatible with Darwin’s legacy of science and conserva-

tion, now (2009) brings more than 150,000 visitors to the islands every
year. The sheer volume threatens the wildlife that the scheme was de-
signed to protect, for along with visitors and an increased awareness
of the importance of Galápagos, comes development, immigration,
invasive species and exploitation. Conservation measures struggle to
keep the threats in check by restricting access to and activities on the
uninhabited islands, prohibiting the harming and exportation of na-
tive wildlife, limiting the introduction of non-native plants and ani-
mals, conducting eradication programs to eliminate feral animals, and
controlling fishing activities. However, the magnitude of the problem
is such that on July 26, 2007, the United Nations Educational, Scien-
tific and Cultural Organization (UNESCO) added Galápagos to their
“World Heritage Site in danger” list. Now, with mounting pressures
from a spiraling human population, vigilance and understanding are
needed more than ever to ensure that Darwin’s islands are not de-
stroyed. It was through Darwin that people have come to appreciate
Galápagos, and it is through intimate recollection of his visit—what
he did, what he saw, what he interpreted—that its cherished worth
can hope to be maintained. It is to this end that this book is dedicated.

Two Tales in One

In the year 2000, after making repeated field trips to fine-tune Dar-
win’s paths in the islands, Greg and I published the results of our ex-
pedition as a scientific paper.71 We lectured on the subject, continued
with literature and field research, and designed an educational tour
of the islands based on the Beagle’s route through the archipelago. We
knew, however, that there was more to tell than where Darwin ex-
plored in Galápagos and a comparative analysis of what he saw there.
The significance of Darwin’s visit to Galápagos stretches beyond the
geographical boundaries of the archipelago and the temporal limits
of Darwin’s time there. The repercussions of his visit encompass the
world and exceed Darwin’s lifetime. The story of Darwin in Galápagos
was fabric for a book.
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18 Introduction

Darwin in Galápagos: Footsteps to a New World is primarily a tale

of two expeditions woven into one. Darwin’s visit to Galápagos on the
voyage of the Beagle provides the warp and our revisit provides the
weft. For without the second, and the research that made it possible,
there would be little to write about the first beyond what has already
been included in a chapter on Galápagos in a book about Darwin,
or a chapter on Darwin in a book about Galápagos. We have used
our unique and intimate knowledge of all the islands in Galápagos
to interlace a scenic backdrop to blend with the natural history facts
Darwin recorded. We have also applied the findings of modern sci-
entists and our own insights accumulated from over 50 combined
years of ecological research on various land birds, sea birds, reptiles,
mammals, invertebrates, and plants in Galápagos to give a modern
perspective to Darwin’s observations.
The story of Darwin in Galápagos would have little meaning with-
out some understanding of Darwin, the man. The maxim, “we are the
sum of all the moments of our lives,”72 cliché though it sounds, is key
to understanding Darwin’s visit to Galápagos. Everything Darwin
did, observed, collected, and thought in Galápagos was directed by
what he already knew, what he was in the process of discovering, and
what he yearned to understand better. A look at Darwin’s life before
Galápagos, and especially at his education both before and during
the voyage, is therefore needed to make sense of his Galápagos visit.
It is important to know that Darwin attended two of the most re-
spected universities in Great Britain, and was one of the most highly
trained men of his age.73 Before arriving in Galápagos he had been
introduced to a wide range of scientific ideas and was aware that the
scientific community, although heavily predisposed toward creation-
ism, was also debating the concept of transmutation. Indeed, it can
be said that many of the ideas Darwin developed were “lying fallow”
in some form or another in England before he sailed on the voyage of
the Beagle. It was the combination of Darwin’s university training, his
experiences during the voyage, and the “literary counsel” he received
along the way that enabled him to pull it all together.74
This book does not pretend to be a biography; it purposefully em-
phasizes one stage in his life. But in order to put Darwin’s Galápagos
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Introduction 19

experience in perspective with the rest of his life and the development
of his scientific thinking, we take a chronological pathway. Drawing
principally from Darwin’s autobiography, the Darwin biographies of
Janet Browne and Adrian Desmond and James Moore, and the criti-
cal analyses of James Secord, Peter Bowler, and other historians of
science, we provide glimpses of the experiences and ideas that helped
shape Darwin and prepare him for Galápagos. We start with Dar-
win growing up as a child naturalist in England during the Industrial
Revolution. We progress through his years at university, where he
learned the scientific theories of the day. We then examine how Dar-
win matured on the voyage of the Beagle, and how his experiences
in South America prepared him for what he would observe in Galá-
pagos. We next take a step-by-step exploration of Darwin’s journey
through Galápagos and beyond, and demonstrate how the islands
gradually changed his way of thinking. Finally, we look at Darwin af-
ter the voyage of the Beagle and show how Galápagos affected his life,
and work, back home. The greatest portion naturally takes place in
Galápagos for it is our principal aim to take the readers to the islands
and have them put on Darwin’s proverbial hiking boots and discover
Galápagos as he did . . . as we did.

To make Darwin’s visit to Galápagos a story rather than a fact sheet

we have used some poetic license. It is impossible to determine what
Darwin thought or how he felt at all times. He was a prolific writer75
but a factual one; his personal journal, at least in the second half of the
voyage, was often short on emotion, opinion, and the daily insignifi-
cances that one might expect to find in a diary. We have gleaned what
we can of his psychological state from his letters home, though they
were few and far between during the months surrounding his visit to
Galápagos. Having said this, the book is nonfiction, and wherever we
veer strongly away from known fact, we make clear our digression
and our reasoning for it.
In the book we refer to both Darwin’s diary and his journal. An
explanatory note is in order to distinguish between the two, because
Darwin confusingly used the word “journal” for both manuscripts.
Darwin’s diary is the “journal” of his activities and observations while
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20 Introduction

on the voyage of the Beagle. This Beagle diary is one of the most il-
luminating volumes about the voyage. Darwin’s granddaughter Nora
Barlow published the first edited version in 1933 and Darwin’s great-
grandson Richard Keynes published a second version in 1988. Even
though Darwin never published his diary himself, he used it and the
letters he had written during the voyage to compose a more polished
account of the Beagle voyage—his published Journal. This tome, orig-
inally entitled Journal and Remarks, was first published in 1839 as the
third and last volume of Captain FitzRoy’s Narrative of the Beagle.
Later that same year Darwin’s contribution was republished as a sepa-
rate book under a new title: Journal of Researches into the Geology
and Natural History of the countries visited during the voyage of HMS
Beagle round the world. In 1845, after most of the Beagle specimens
had been examined and described by taxonomists, Darwin rewrote
his Journal to include what he had learned from them. The title re-
mained the same, except the order of the words Geology and Natural
History was switched to reflect a change in emphasis. Later editions of
the same book were called The Voyage of the Beagle.76 To avoid confu-
sion, and for consistency’s sake, whenever we quote from Darwin’s
Journal we revert by default to the 1839 edition, unless the material
occurs only in later editions.
Except here in the Introduction we have chosen to use the original
English names of the Galápagos Islands, because they were the ones
used in Darwin’s day. We identify the modern Spanish name of each
island in the corresponding chapter headings, and in a table in ap-
pendix 1. Site names are also listed in this table. Species are identified
by their modern nomenclature (common name and scientific name),
unless otherwise stated.
For simplicity, throughout the book we use the terms “evolution”
and “transmutation” synonymously, defined as the process by which
populations of organisms change from one form into another.77 Dar-
win first used “transmutation” and then “descent with modification”
to express his developing ideas on biological change. While he used
the word “evolve” once at the end of the first edition of The Origin of
Species published in 1859, he did not adopt the term “evolution” until
the 1870s.78
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Introduction 21

Many people were instrumental in producing this book and we

have endeavored to give them full credit in a section devoted to their
acknowledgment. Here, however, we wish to make it clear that there
were really three authors to this book, just as there were three prin-
cipal leaders on our 1996 expedition. Darwin “wrote” much of this
book, and many of the quotations are his.