I.

3
The Evidence for Evolution
Gregory C. Mayer

OUTLINE Homology. The correspondence, determined by their rel-

ative positions and connections, of organs in different
1. The fossil record
organisms, which is indicative of affinity; the cause of
2. Comparative biology
this correspondence is inheritance from a common
3. Biogeography
ancestor.
4. Evolution in action
Oceanic Island. An island that has never been connected
5. Evolution as fact and theory
to a mainland and thus has received its fauna and
flora over water by occasional means of transport.
The evidence for evolution was first comprehensively as- Phylogenetic Tree. A representation of the history of life,
sembled by Charles Darwin, who succeeded in convincing with branching indicating the splitting of lineages,
essentially all his scientific contemporaries of the fact of and the connection of the branches indicating the pas-
descent with modification. A signal factor in Darwin’s sage of genetic information and materials from one
achievement was that he was able to weave together nu- generation to the next.
merous strands of natural history—paleontology, sys- Progression. The pattern in the fossil record in which
tematics, embryology, morphology, biogeography—into earlier forms of life differ from later forms, with major
a coherent framework. Since Darwin, genetics has joined groups first appearing in the record in a generalized
this synthesis, and, in a development that might have form and later as more diversified members of the
surprised Darwin, evolution in natural populations has same group. Some of the earlier forms may become
proven to occur sufficiently rapidly that it can be observed extinct, and later forms may more closely resemble
on human timescales. The most direct evidence of evo- modern forms. Not to be confused with progress, a
lution comes from the fossil record, in which the dynamic different concept, according to which evolution pro-
changes of life over time are recorded, including many ceeds toward some externally defined goal.
transitions between major taxa. A host of phenomena in Speciation. The splitting of a lineage into two or more
comparative biology (e.g., systematics, morphology, em- daughter lineages reproductively isolated and evo-
bryology, genomics) and biogeography that otherwise lutionarily independent from other lineages.
appear inexplicable or anomalous are readily explained Tetrapods. The group of four-limbed vertebrates com-
under the hypothesis of descent with modification. In prising amphibians, reptiles, mammals, and birds;
addition, direct observation of natural and artificial pop- includes species that have secondarily lost their limbs,
ulations shows the process of evolutionary change in ac- such as snakes.
tion. Together, these sources of evidence lead to a “con- Unity of Type. Similarities among organisms leading differ-
silience of inductions” that makes the fact of evolution ent ways of life under diverse conditions of existence,
one of the most securely established generalizations in going beyond any functional need for similarity.
science.
The evidence for evolution, it has been remarked, is not
the result of some crucial experiment but something
GLOSSARY
more like the contents of the American Museum of
Adaptation. A feature of an organism that fits it to its con- Natural History. And so it is: the evidence for evolution
ditions of existence, giving rise to similarity among comes from a plethora of biological and geological
organisms leading the same or similar ways of life. subdisciplines—systematics, paleontology, stratigraphy,
 Evidence for Evolution 29

geochronology, biogeography, morphology, botany, zo- Despite these adaptive similarities, sharks and whales
ology, embryology, genetics—many of which find their differ in many features, such as their respiratory, circu-
objects of study in the vast and varied collections of nat- latory, and reproductive systems, that mark them as
ural history museums. belonging to different major groups of organisms—fish
It is the diversity of these sources of evidence, all and mammals, respectively.
leading to the conclusion that life on earth has under- Darwin provided in his theory of descent with mod-
gone a long history of descent with modification, that is ification a unified explanation of the geological (progres-
the great strength, and the most striking aspect, of the sion), structural (unity of type), and functional (adapta-
evidence for evolution. The varied sources of evidence tion) phenomena. Unity of type reflects inheritance of
are all brought into the unified explanatory scheme of features from common ancestors, and the changes from
evolution, forming what the philosopher William the common ancestor are due to modification. The most
Whewell (1794–1866) called a “consilience of induc- important means of modification, natural selection,
tions,” each piece of evidence reinforcing the whole. leads to adaptation. And when played out over geolog-
Charles Darwin (1809–1882) in On the Origin of ical time, descent with modification leads to progres-
Species used precisely such a form of argumentation, sion. Once admitted, descent with modification and
marshaling the disparate facts of geology, systemat- common ancestry would also account for the hier-
ics, morphology, embryology, and biogeography to archical relationships revealed by systematics and for
support his theory of descent with modification. the distributions of organisms across the surface of the
globe. It was Darwin’s triumph to combine geological,
morphological, embryological, systematic, and biogeo-
Progression, Unity of Type, and Adaptation
graphic evidence into a single explanatory theory.
By the time Darwin began his scientific career, it was This chapter considers the classes of evidence ad-
already well established that the earth was old and that duced by Darwin to support his theory of descent with
the fossil record was progressive, that is, that earlier modification, including examples from post-Darwinian
forms of life differed from later forms, that some of the disciplines such as genetics, and adds as an additional
earlier forms had become extinct, and that later forms class of evidence observations of evolution in action, a
more closely resembled modern forms. It was also be- class that was largely unavailable to Darwin.
coming clear that in this progression not only were older
forms replaced by newer ones but later forms were in
1. THE FOSSIL RECORD
some way related to earlier ones. Thus, a major group
would appear in the fossil record in a generalized form
Progression
and would be succeeded by more diversified members of
the same group. The earliest fossils are of simple (prokaryotic), single-
In addition to progression, two other great, but un- celled, photosynthetic bacteria that lived in mats called
explained, classes of phenomena occupied biologists at stromatolites 3.5 billion years ago (see chapter II.11).
this time: unity of type and adaptation. Unity of type One and a half to 2 billion years ago, more complex,
refers to the similarities among organisms living differ- organelle-bearing (eukaryotic) but still single-celled
ent ways of life, similarities that extend far beyond any forms appear, and then multicellular forms. The origins
functional needs. The same basic skeletal plan of the of eukaryotes and multicellularity are not well docu-
forelimb—a humerus, then radius and ulna, then car- mented in the record, and molecular data suggest that
pals, then metacarpals, then phalanges—occurs in all they may have occurred considerably earlier than the
tetrapods, even though the limbs might appear quite record shows (see chapter II.12). In the last part of the
different externally and be used for very different func- Precambrian, about 600 million years ago, diverse forms
tions (figure 1). Such similarities extend to embryolog- of marine invertebrates appear, and then in the Cambrian
ical features as well: all tetrapod embryos, for example, many more invertebrate groups arise, as well as the first
have four limb buds, even if the adults (e.g., whales, vertebrates (see chapter II.15).
snakes) lack one or two sets of limbs. The earliest vertebrates, such as the recently dis-
Adaptation refers to those features of organisms that covered Myllokunmingia and Haikouichthys, which
suit them to their conditions of existence (see chapter look remarkably like previously hypothesized general-
III.1; for a nuanced discussion of terminology, see ized vertebrates, were soft-bodied jawless forms that
chapter II.6), which may be shared by organisms with lived about 525 million years ago. Diverse jawless fishes
similar ways of life. Thus, sharks and whales share flat- with mineralized hard tissues followed. The first jawed
tened tail flukes and dorsal fins, both features being of fishes appear in the Late Ordovician, about 445 million
obvious functional importance for aquatic organisms. years ago. The bony fishes, or Osteichthyes, the most
30 Introduction

Human arm

Humerus Radius Wrist bones

Ulna and digits

Theropod dinosaur

Seal

Bird

Lizard

Pterosaur

Penguin

Bat

Humpback whale
Figure 1. Unity of type and adaptation, illustrated by the forelimbs though the size, shape, and relative proportions of the elements
of tetrapods. The pattern of one bone (humerus), two bones (radius have been modified for varied ways of life, including walking, flying,
and ulna), many bones (wrist and digits) is present in all, even grasping, and swimming. (Copyright Kalliopi Monoyios.)

diverse living group of jawed fishes, are first found in of organisms through geological time. Moreover, as one
the Late Silurian, about 420 million years ago. The first moves toward the present, many taxa go extinct and
four-legged vertebrates (tetrapods) are the approxi- others dwindle in diversity, and the array of major
mately 365 million-year-old amphibians Acanthostega groups comes to progressively resemble that of today.
and Ichthyostega from the Devonian (see chapter II.17).
Reptiles, the first tetrapods to be independent of water
Transitions
(amniotes), make their appearance in the Pennsylvanian,
about 315 million years ago. In the Mesozoic era, the last Darwin did not have closely spaced transitional forms
two major tetrapod groups arise: the first mammals are that could demonstrate evolutionary continuity between
known from the Triassic/Jurassic boundary (about 200 major groups; he attributed their absence to the im-
million years ago), and the first bird is from the Late perfections of the geological record. The record is indeed
Jurassic, about 150 million years ago. imperfect: only hard parts of organisms are readily fos-
The fossil record of vertebrates, then, exemplifies the silized, only certain sedimentary environments are con-
sequential origin and diversification of the major groups ducive to fossilization, fossils must survive erosion and
 Evidence for Evolution 31

metamorphism and, finally, must be exposed on the

surface and discovered. As a consequence, fewer than
q
1 percent of all species that ever lived are represented in
the fossil record, and the record is spotty temporally,
geographically, and taxonomically (see chapter II.9). For
example, only about 30 localities have yielded important d an ar
fossil faunas of tetrapods from the 60 million years of the
Carboniferous, when the first major radiation of am- Early synapsid
phibians occurred.
Despite the imperfections, the transitional forms that
Darwin hoped might be found soon started turning up.
In 1861, Archaeopteryx was discovered and became the
first of the proverbial “missing links” to be found. q
Clothed in the feathers of a bird with broad wings, it
nonetheless had the long bony tail, toothed jaws, and d ar
an
free, clawed fingers of a reptile. T. H. Huxley (1825–
Advanced synapsid an
1895) saw it clearly as intermediate between reptiles and
birds and suggested a relationship to dinosaurs, a sug-
gestion now well documented. Extraordinarily well-
preserved fossils show not only the close skeletal resem-
blance of Archaeopteryx and early birds to theropod q
dinosaurs but also that feathers are not peculiar to birds; ar
they were present on quite a few nonflying dinosaurs as d
well. an
Morganucodon
A few years later, Richard Owen (1804–1892) and
E. D. Cope (1840–1897) both recognized that certain
ancient reptiles (synapsids) bore an affinity to mammals. ar

The origin of mammals from the synapsids is now known q

in exquisite detail (figure 2). Reptiles have several bones
in the lower jaw, and one of the more posterior ones, the an
articular, forms the jaw joint with the quadrate bone of
the skull. Mammals have a single bone in the lower jaw,
d
the dentary (also present in reptiles), which articulates
with the squamosal bone of the skull. A long series of
Advanced mammal
fossils, beginning with fully reptilian forms in the Penn-
Figure 2. Homology of jaw bones of reptiles and ear bones of mam‐
sylvanian, lead gradually to the mammals at the Triassic/ mals. In early synapsids the lower jaw comprises the tooth-bearing
Jurassic boundary. The dentary enlarges, becoming the dentary (d) and several postdentary bones, including the angular (an)
largest bone of the lower jaw. The quadrate and articular and articular (ar). The latter articulates with the quadrate (q), form‐
become smaller. Eventually, the dentary contacts the ing the reptilian jaw joint. In advanced synapsids the latter bones
squamosal, leading to several forms, such as Probainog- are reduced, while the dentary enlarges. In the earliest mammals
(Morganucodon) these bones are reduced further, and the dentary
nathus, that have a dual jaw joint—both the old reptilian makes contact with the upper jaw, forming the mammalian jaw joint.
one and the new mammalian one. The transition is so In advanced mammals the angular, articular, and quadrate detach
gradual that it becomes a matter of convention to decide from the jaw entirely, becoming the tympanic, malleus, and incus
which form is the first “mammal”: Morganucodon is of the mammalian ear. (After D. Davis 1991, K. Kardong 2012, and
R. Carroll 1988.)
often so regarded, but it, too, has a dual jaw joint. In later
forms the quadrate and articular detach from the jaw and
become two of the three mammalian ear bones. Many
other features that change during this transition—for (figure 3). Osteolepiforms were “typical” fish, with dor-
example, the dentition becomes cusped, a secondary sal and anal fins, rounded heads with short snouts, and
palate forms, the ilium becomes rod shaped—are like- their shoulder girdles connected to their heads. But their
wise documented in the fossil record. pectoral and pelvic fins extended from the body in fleshy
Another well-documented transition is that between lobes, and within the lobe was a skeleton that, starting at
the lobe-finned osteolepiform fishes and tetrapods the base (the end near the body), had a pattern of “one
32 Introduction
planktonic protists, such as foraminifera and radiolar-
ians, that can be recovered by extracting cores from the
seabed. The shells of dead individuals rain down onto the
ocean floor, forming an essentially continuous sedimen-
tary record. In these organisms, such as the foraminif-
Acanthostega gunnari
erans in the genera Globorotalia and Contusotruncana,
and the radiolarian genus Eucyrtidium, such fine-scale
changes can be observed, and in the latter even the
split into two species from an original one has been
recorded.
There are many other examples of transitional forms
Tiktaalik roseae in the fossil record—such as ancestral whales and snakes
with hindlegs—and more are being discovered every
year.

2. COMPARATIVE BIOLOGY
Eusthenopteron foordi

Figure 3. Transition from lobe-finned fish to tetrapods. The lobe- Unity of Type
finned osteolepiform fish Eusthenopteron has the tetrapod-like “one
bone, two bones, many bones” pattern in its fin. Tiktaalik, the “fish- While the fossil record provides direct evidence of change
apod,” has lost the dorsal and anal fins, the head is free of the shoulder of life over time, comparisons among organisms, either
girdle, the snout is elongated with the eyes directed upward, and there living or extinct, give evidence that the link between the
is a wrist joint in the forelimb. Acanthostega, one of the first tetrapods,
has digits but retains the caudal fin of its fish ancestors. The bones of
different forms at different times is genealogical. Primary
the left forelimb shown in gray are, from darkest to lightest, the hu- among these comparisons are the observed similarities in
merus, the radius, and the ulna, respectively; more distal elements fundamental structure among organisms referred to as
are unshaded. (Copyright Kalliopi Monoyios.) “unity of type.” These similarities, which extend from
morphology to development to the genome, are homol-
ogies, that is, similarities due to inheritance from a
bone, two bones, many bones.” This is the same pattern common ancestor (see chapter II.6).
as in tetrapods: the plan that exemplifies the unity of type Famous homologies are the “one bone, two bones,
of the tetrapod forelimb extends to osteolepiform fishes. many bones” pattern of the tetrapod limb (figure 1), and
In Panderichthys, from about 380 million years ago, the the jaw and ear bones of reptiles and mammals, both
head and body are flattened, the snout is elongated, and mentioned previously (figure 2). In the case of the limbs,
the eyes are on top of the head; the anal and dorsal fins the strong similarity of the same bone among the various
have been lost. In the remarkable Tiktaalik from 375 tetrapods allows the homologous bones to be easily rec-
million years ago, the shoulder girdle (equivalent to our ognized: for example, humerus, radius, ulna. The sim-
collarbone and shoulder blades) has been freed from the ilarity in plan is not accounted for by the functional
skull—Tiktaalik had a neck; and there is a joint within requirements of the limbs but by inheritance from a
the “many bones” of the forelimb—it also had a wrist. common ancestor. In the jaw and ear, the homologies are
Ten million years later we have the first actual tetrapods, traced with the aid of fossil and embryological (see the
Acanthostega and Ichthyostega, which have legs with section Markers of History) evidence. The angular and
toes but retain some of the gill-cover bones and the articular bones of the reptilian lower jaw are homologues
caudal fin of their fish ancestors. of the tympanic and malleus bones of the mammalian ear,
The origins of birds, mammals, and tetrapods rep- while the quadrate of the reptilian upper jaw is the incus
resent fairly large changes in morphology and way of life, of the mammalian middle ear.
and the intermediate forms bridge the differences be- These patterns can be seen not only in the skeleton
tween these major taxa. The much smaller transitions but in the genome itself at the cellular level. The human
that occur as one species evolves into another are also genome contains 23 pairs of chromosomes, whereas the
documented in the fossil record, but continuous sedi- genome of great apes has 24 pairs. The difference of one
mentary deposition is necessary to record such fine-scale pair can readily be accounted for: human chromosome 2
temporal events. Such conditions are not common but is homologous to two ape chromosomes, which have
occur most often in the fossil record of shelled marine become fused in the human lineage. The homology has
 Evidence for Evolution 33

Tetrapods Amniotes Squamates Anoles

Toepads

Hemipenis

Amniotic membrane

Four legs

Figure 4. Phylogenetic tree of anole lizards showing nested evolved hemipenes, in turn are nested within amniotes. And finally,
homologies. The common ancestor of all tetrapods possessed four anoles are characterized by the possession of expanded toepads.
legs, a trait that, with modifications, was passed on to all of its Anoles thus possess toepads, hemipenes, amniotic membranes,
descendants. One of these descendants evolved the amniotic and four legs, each trait inherited from a successively more distant
membrane, thus nesting the amniotes (reptiles, birds, and mam- ancestor, and each marking a more inclusive group. The nested
mals) within the tetrapods. Squamates (snakes and lizards), which boxes in the figure indicate the named clades.

been confirmed by the discovery of remnants of the central the branching history of life, represented by the phyloge-
(centromere) and end (telomere) portions of the ape netic tree (see chapter II.1). When a lineage divides, giving
chromosomes within the human chromosome, showing rise to a new branch in the tree of life, the characteristics of
that the latter arose from what were originally two the splitting lineage are passed on to its descendants.
chromosomes. Modifications may occur in a descendant lineage, which
At perhaps the most basic level, the near universality will in turn be passed on to its, and only its, descendants.
of the genetic code is another homology that argues for Each homology is the origin of a new feature, shared with
the common ancestry of all life. The genetic code as- descendants but not with collateral relatives. These nested
sociates each amino acid with a codon of three nucleo- sets of homologies, which are exactly what one would
tides, which carries the information for the making of expect from a process of descent with modification, are
proteins. But the identity of the three nucleotides is ar- powerful evidence for evolution.
bitrary, so the code’s universality cannot be due to For example, possession of a toepad, composed of
functional constraint but arises as a legacy of the code laterally expanded scales at the end of the digit, char-
established in distant progenitors. acterizes lizards of the genus Anolis (figure 4). Anoles
also have a hemipenis (oddly named, as this means that
they have two penises, rather than half a penis, as the
Common Ancestry
word might imply) as the male intromittent organ, but
Although the common ancestry indicated by the genetic this organ characterizes a larger group, the squamates
code embraces all (or nearly all) living beings, homol- (lizards plus snakes), within which anoles are nested. The
ogies do not generally have such a wide distribution. squamates, in turn, are nested within a yet-larger group
Rather, homologies characterize smaller groups, and these characterized by the presence of an amniotic membrane
groups are nested within larger homology-characterized around their embryos. The amniotes (reptiles, birds, and
groups, which in turn are nested within yet larger such mammals) also have four legs, a trait shared with am-
groups, and so on. This nesting of homologies results from phibians as well, which together make up the tetrapods.
34 Introduction
The tetrapods, also in turn, share features of the limb along the jaw corresponding to those occupied by the
with certain fishes, forming again a more inclusive group. homologous jaw bones of reptiles. The “thumb” of pan-
Continuing in this manner, the bony fishes, vertebrates, das (figure 6) is not homologous to other mammals’ inner
chordates, and deuterostomes constitute successively digits, nor is it even a digit: it is a modified radial sesamoid
larger groups within which anoles are nested, and this bone, pressed into service as a makeshift digit, in an ex-
nesting is indicative of the history of common ancestry. quisite example of tinkering—that is, the modification of
Eventually all, or almost all, life can be subsumed available structures, rather than an engineering ideal. In
within the nested tree. For organisms as divergent as the Australian lungfish (Neoceratodus), there is a single
insects and vertebrates it is hard to recognize morpho- dorsal lung, unlike in other lungfish (and tetrapods), in
logical homologies, but genetic data show that homol- which the lungs are paired (figure 7). In all lungfish (and
ogies of gene and genome structure can be recognized tetrapods), the lung attaches to the ventral part of the
in, for example, the presence of homologous sets of de- esophagus. In Australian lungfish, the pneumatic duct
velopmental regulatory genes (Hox genes) in both ar- travels up alongside the right side of the esophagus to the
thropods and vertebrates. While nestedness holds true dorsally positioned lung, seeming to trace the course of its
for eukaryotes, there is some question whether at the morphological modification. Confirming this movement,
base of the tree of life transfer of genetic material be- the left pulmonary artery, instead of going directly to the
tween prokaryotes may be so common as to obscure or dorsal lung, travels down the left side of the esophagus,
efface the nested pattern (see chapter II.11). curls under the esophagus, and follows the pneumatic
duct up to the lung.
Such examples are not limited to morphology: pseu-
dogenes, nonfunctional versions of genes that are func-
Markers of History
tional in related species or even in other copies in the
Among the most striking evidences of evolution are the same organism, occur commonly in organism’s genomes
features of organisms that appear to reflect a constraint: (see chapters V.3 and V.5). For example, in most
organisms inherit from their ancestors a developmental mammals, vitamin C is synthesized by a battery of en-
system, and evolutionary modifications must take that zymes. In primates, which get vitamin C in their diet, a
inherited system as a starting point. Indeed, organ- gene for one of the necessary enzymes is present as a
ismal features give every indication of having arisen by nonfunctional pseudogene, so that the vitamin is not
“tinkering” with this inherited preexisting developmen- synthesized. The broken gene is a relict from earlier
tal system, and make sense only as a result of descent mammals that do use it in their synthesis of the vitamin.
with modification. In primates, the gene has been disabled by a mutation,
This constraint is evident in the similarities among but the now-inactive gene remains as a marker of pri-
the embryos of jawed vertebrates. The embryos go mates’ forebears.
through a stage in which they resemble one another in
the possession of pharyngeal arches, limb buds, tails, and
3. BIOGEOGRAPHY
other traits. After this stage, embryos diverge, develop-
ing into the varied forms they will become as adults. Historically, the biogeographic evidence for evolution
What perhaps is most striking is that structures present was crucial, because it was the evidence that convinced
in the embryo are not always present in the eventual Darwin himself. Many features of the distribution of
adult. Thus, humans and apes have an embryonic tail organisms that are anomalous or merely curious under
that largely disappears in the adult; only a bony vestige a hypothesis of special creation became explicable and
remains internally. Humans develop a coat of fine fur, expected under the hypothesis of descent with mod-
the lanugo, which is lost just before or shortly after birth. ification. Under the latter hypothesis, related species
All four limb buds develop in tetrapods, such as whales should occur in geographically connected areas or in
and snakes, that in the adult lack one or both pairs of areas that could have been reached by a common an-
limbs. In some snakes, a vestigial leg is still visible ex- cestor of the related species. The connectedness and
ternally (figure 5). In each case, the eventual adult form “reachability” of areas are determined chiefly by the
develops from a shared state and subsequently passes distance and geographic barriers between them, and
through stages shared with smaller nested groups until the ability of organisms to cross such barriers. Geo-
finally arriving at its own specific state. graphic conditions that either present barriers to dis-
Many structures show the traces of ancestry. In fetal persal (e.g., the ocean around islands) or facilitate it
mammals, the bones that eventually become the bones (e.g., the continuous land of continents), and organ-
of the middle ear begin their development in positions isms’ abilities to overcome or utilize these geographic
 Evidence for Evolution 35

Figure 5. External hindlimb of a snake, the ball python (Python is underlain internally by rudiments of the femur and, deeper in the
regius). Located just lateral to the anal scale, the keratinous claw is body, the pelvis. (Photo by G. C. Mayer.)
here shown slightly pushed away from the body by a probe. The claw

barriers and bridges, are thus key determinants of the have been many successful introductions of verte-
distribution of life on earth. brates, including land birds, reptiles, amphibians, and
land mammals (figure 8). In its features, the fauna of
Bermuda is typical of oceanic islands.
Islands
All these characteristics are readily explained by the
Bermuda is a small group of coral islands in the North hypothesis of descent with modification. As a geologically
Atlantic, 1000 km to the east of North America. Sitting oceanic island, Bermuda lacks organisms that cannot
atop a long-extinct volcanic platform and surrounded cross 1000 km of ocean (land mammals and amphibians)
by waters of abyssal depth, islands such as Bermuda and is inhabited by descendants of a limited number of
are called oceanic—they have never been connected by successful colonists, whose characteristics, such as the
land to a mainland. The native non-marine vertebrate ability to fly, have permitted cross-oceanic dispersal. The
inhabitants of Bermuda are few—several land birds, nearest relatives of these colonists reside in the adjacent
migratory bats, a lizard, a terrapin—and show closest landmass of North America, because the islands are most
affinity to forms from the North American mainland. accessible from there by “occasional means of trans-
Some are identical or nearly so to the North American port” (as Darwin called them). Some of the colonists
forms, whereas others are distinct endemic species have been isolated sufficiently long to have diverged into
found nowhere else. Several major groups common on endemic forms (e.g., the lizard), whereas others more
the mainland are lacking entirely—there are, for ex- recently arrived have not so diverged (e.g., the terrapin).
ample, no nonflying terrestrial mammals and no am- The success of invasive introduced forms shows that the
phibians. Subsequent to human colonization, there island fauna was, as Darwin put it, insufficiently stocked
36 Introduction

Right
pulmonary
artery

Lung

Left pu
lmon
Gut

ar y
ar
te
ry
Figure 7. The lung and its arterial blood supply in the Australian
lungfish (Neoceratodus). Note that the pneumatic duct swings around
the gut to make the lung dorsal, and the left pulmonary artery, after
branching from the dorsal aorta, follows it down and around (instead of
going straight to the lung). (After E. S. Goodrich 1909.)

found elsewhere only in the Americas, where they are

much less diverse. In Australia, the marsupials occupy all
Figure 6. The right manus in the brown bear (Ursus arctos; left) and
the major habitats and have a rich fossil record going back
the giant panda (Ailuropoda melanoleuca; right). The panda’s thumb is tens of millions of years. Having been isolated in Aus-
not a true digit but a modified radial sesamoid bone. (After D. D. Davis tralia, the marsupials have prevailed and diversified into
1964.) most ecological roles: herbivores (kangaroos), carnivores
(thylacine), burrowers (wombats), climbers (koala), gli-
by the “creative force”: it was difficulties of dispersal, ders (phalangers), and many more. The only native pla-
not habitat suitability, that caused the fauna to be re- cental mammals are bats and murid rodents. Bats can fly,
stricted in species richness and taxonomic diversity. and rodents are able to colonize across water barriers
(although not ones as wide as those that can be crossed by
bats).
Continents
Geographic distributions that might seem anomalous
On continents, the accessibility of adjacent areas leads, in under the hypothesis of descent with modification are
general, to related forms being found throughout. Dar- explicable when the movement of the continents is taken
win noted this for the mammals of South America. The into account. For example, Cynognathus, a terrestrial
current mammals of the varied South American habitats synapsid of the Triassic, about 240 million years ago,
—tropical, temperate, alpine—are all related to one an- has been found in South America, Africa, and Antarctica,
other and not to mammals of similar habitats in distant all now widely separated by seas unlikely to be crossed
places. Although organisms can sometimes disperse over by a stout, meter-long animal like Cynoganthus. But in
large distances and across barriers (as in the case of many the Triassic all these continents were connected in a sin-
oceanic islands), dispersal occurs more readily between gle landmass, Gondwana. Far from being a problematic
nearby areas, so that the biota of nearby areas, such as case, Cynognathus shows again the importance of his-
the different habitats of South America, have similar torical connectedness, both genealogical and geological,
compositions. Similarly, the fossil South American mam- in the distribution of organisms.
mals Darwin found were related to the ones now present.
Thus, related organisms live in places they can reach by
4. EVOLUTION IN ACTION
dispersal, and descend from the previous inhabitants.
This pattern is also well illustrated by the mammal
Changes within Populations
fauna of Australia, a continent that became isolated from
the other continents tens of millions of years ago. Most of Although fossils record descent with modification over
the mammals of Australia are marsupials, a group now billions of years, evolution often occurs quickly enough
 Evidence for Evolution 37

Figure 8. Endemic and invasive oceanic island species. Left: The native to Middle and South America, and introduced to Bermuda in
endemic Bermuda skink (Eumeces [Plestiodon] longirostris) is the 1885. It is one of several exotic amphibian and reptile species that
only extant native terrestrial reptile of Bermuda. Its nearest rel‐ have thrived and even become pests there, after the barrier to dis-
atives are from the nearest mainland, North America, 1000 km to persal was overcome by human agency. (Skink photo by Richard
the west. Right: A Bermudian specimen of Bufo (Rhinella) marinus, Ground; toad photo by G. C. Mayer.)

to be observed within one or a few human lifetimes, The use of pesticides and antibiotics has frequently led
and such cases allow the full panoply of evolutionary to the undesired evolution of resistance in the targeted
mechanisms—mutation, gene flow, genetic drift, and organisms, including rodents, insects, bacteria, and virus-
natural selection—to be seen in action. es. Multiple-antibiotic-resistant strains of bacteria have
Darwin’s chief examples of observed evolution in- become a major health problem, and one of the greatest
volved the work of animal and plant breeders in pro- difficulties in treating acquired immune deficiency syn-
ducing and elaborating the features of domesticated drome (AIDS) has been the rapid evolution of resistance
organisms (see chapter VIII.5). Studies of such species by the human immunodeficiency virus (HIV). In the evo-
show the importance of selection in establishing and lution of HIV, high mutation rates, short generation
fixing desired characteristics. The many varieties of do- time, large population sizes, and strong selection have all
mestic dog, differing so much in size, shape, and be- combined to make the virus adapt extremely rapidly to
havior, have all been produced from the wolf in the last drugs. Drug mixtures, which attack the metabolism of
few thousands of years. Corn, one of the most highly the virus in different ways simultaneously, have proven
modified and economically important organisms ever more effective, as the multiple mutations required for
created by humans, was developed by selective breeding resistance to all the drugs are less likely to occur.
from a wild species of grass. Species introduced into new geographic areas, whether
Rapid evolution has also unintentionally been caused by humans or by natural means, are likely to find them-
by humans, who changed the environment in ways that selves in new environments and thus are more likely to
prompted evolutionary responses from affected organ- undergo evolutionary changes. Such introductions have
isms (see chapters VIII.2 and VIII.3). Industrial melanism, indeed produced many examples of divergence in fea-
the evolution of darker coloration in animals living in tures such as size, shape, and coloration. One example
environments darkened by pollution, is widespread in a is the house sparrow (Passer domesticus), introduced
variety of insects in industrialized areas of both Europe into North America from Europe about 1851. By the
and North America. When pollution controls have been middle of the twentieth century, the sparrow had
enacted, the evolutionary change has been reversed, and geographically differentiated in coloration, size, shape,
the lighter-colored forms have again increased in fre- and physiology in a manner parallel to that of native
quency. Both the spread of melanism and its reversal have species.
been observed in Britain in the moth Biston betularia over Natural populations have also been observed evolv-
a period of about 150 years. ing in response to natural environmental changes (see
38 Introduction
chapter III.7). Perhaps the best-studied case is that of
5. EVOLUTION AS FACT AND THEORY
Darwin’s finches (Geospiza) in the Galápagos, where
populations have been carefully tracked over decades. It is sometimes noted pejoratively that evolution is a
During this time span, repeated episodes of morpholog- “theory.” This pejorative usage confuses a vernacular
ical evolution have occurred in response to climatically notion of “theory” as something uncertain or conjec-
induced variations in the food supply. Further, the ge- tural, with the word’s scientific usage. In science, a theory
netic basis of these changes has been demonstrated by is not a mere conjecture but a connected series of prop-
observations in nature. These studies show that natural ositions supported by, and explanatory of, many and
populations are not evolutionarily static but can adap- varied lines of evidence. We refer to the “germ theory of
tively track changes in the environment. disease” not because we are unsure that microbes cause
disease but because the theory is a set of high-level and
powerful generalizations that account for and are, in
Speciation
turn, supported by a huge amount of data.
Speciation, the origin of new lineages reproductively iso- Regarding evolution, it was Darwin who first con-
lated and evolutionarily independent from other line- vincingly assembled the many and varied lines of evi-
ages, is a key part of evolution (see Section VI: Specia- dence that could be accounted for by, and provide evi-
tion and Macroevolution). Without it, evolution might dence of, descent with modification. His many successors
occur within a lineage, but there would be no increases in have carried on his work, and Darwin would have been
biodiversity. Divergence of lineages is enhanced by their both pleased and astonished by the further lines of evi-
geographic separation (since it reduces or eliminates the dence that have been brought to bear. In his own lifetime,
homogenizing force of gene flow). Isolated populations transitional fossils began to be found, and we now have
may differentiate independently—including in character- them in abundance. Darwin’s own attempts at for-
istics affecting reproductive compatibility—to the point mulating principles of inheritance failed, so he would
that they can no longer exchange genes if brought again have been gratified by the explosive growth of genetics
into contact. The insensible gradation over space of pop- and now genomics filling in what he did not know.
ulations varying from mere geographic isolates to Crucially, the facts of genetics could have turned out to be
highly differentiated populations approaching genus- incompatible with Darwin’s evolutionary views but, in-
level distinction provides evidence for speciation by geo- stead, his ideas have passed this important test. He would
graphic isolation, and there are many examples, such as in perhaps have been most astonished by the evidence for
kingfishers of the genera Tanysiptera and Halcyon on the the rapidity with which evolution can occur, including
islands of the southwest Pacific. In most cases though, the formation of new species within a human lifetime.
divergence takes too long to be observed within a human As briefly reviewed here, all these lines of evidence, all
lifetime. Laboratory studies have shown that incipient leading to the same conclusion, serve to make descent
reproductive isolation can arise in separated populations with modification one of the most securely established
undergoing differential adaptation, and that reproductive high-level generalizations in science and allow us to
isolation can be enhanced by selection against individuals speak confidently of the fact of evolution.
who hybridize with the “wrong” population.
In some cases, though, speciation occurs sufficiently FURTHER READING
quickly to be observed in the wild within a human life-
time. Speciation by polyploidy (i.e., the duplication of Carroll, R. 2009. The Rise of Amphibians: 365 Million Years
of Evolution. Baltimore: Johns Hopkins University Press.
chromosome sets) is an important mode of splitting in
Includes an account of the origins of vertebrates and
plants (see chapter VI.9). A well-studied case is the recent reptiles, as well as of the origin of amphibians.
origin of the British salt marsh grass Spartina anglica, a Coyne, J. A. 2009. Why Evolution Is True. New York: Viking
polyploid derived from the natural crossing of the in- Penguin. An account of the evidence for evolution for a
troduced American species S. alterniflora (diploid chro- general audience.
mosome number 2n = 60) with the native S. maritima (2n Dawkins, R. 2009. The Greatest Show on Earth. New York:
= 62). Spartina alterniflora was accidentally introduced Free Press. Another account of the evidence for evolution
in 1829. In 1870, sterile hybrids with S. maritima were for a general audience.
Futuyma, D. J. 2013. Evolution. 3rd ed. Sunderland, MA:
first recorded. Sterility of the hybrids, due to mismatch-
Sinauer. The leading textbook of evolutionary biology.
ing of the parental chromosomes, was overcome by Grant, P. R., and Grant, B. R. 2008. How and Why Species
chromosome doubling, and the fertile S. anglica (2n = Multiply: The Radiation of Darwin’s Finches. Princeton,
122) was first recorded in 1892. The new species has NJ: Princeton University Press. An account of the detailed
since spread along coastlines throughout Britain. evolutionary studies conducted by the authors and their
 Evidence for Evolution 39

associates over four decades, including several episodes of amphibian transitional form Tiktaalik, and of the traces
closely observed evolutionary changes. of common ancestry within vertebrates (including man),
Mayr, E. 2001. What Evolution Is. New York: Basic Books. A and of the common ancestry of vertebrates with other
summary of the evidence for evolution and its mechanisms animals.
by the “Darwin of the twentieth century.” Young, D. 2007. The Discovery of Evolution. 2nd ed. Cam-
Prothero, D. R. 2007. Evolution: What the Fossils Say and bridge: Cambridge University Press. Intended for a general
Why It Matters. New York: Columbia University Press. A audience, this superb and richly illustrated history serves
richly illustrated account of the fossil record emphasizing as a text for evolutionary biology itself, because it in-
transitions between major groups. troduces and explicates not just the ideas and historical
Shubin, N. 2008. Your Inner Fish. New York: Pantheon figures but the evidence on which the major discoveries of
Books. A popular account of the discovery of the fish- evolutionary biology are based.