Biogeography of Vertebrates
RM Brown, University of Kansas, Lawrence, KS, USA
r 2016 Elsevier Inc. All rights reserved.
Glossary
Adaptive radiation The evolution of ecological and
phenotypic diversity within multiplying lineages, usually
associated with the rapid production of high species
diversity.
Ecological opportunity As organisms colonize new areas
and encounter environments (islands, habitats) in which
competitors or predators are absent, or in which new
food resources can be exploited, novel adaptive
processes can dive the evolution of new species as a
response.
Biogeography is the study of the evolution of geographical
distributions of organisms, a topic that has intrigued and be-
wildered theologians, philosophers, and biologists for thou-
sands of years. We can trace the origins of biogeographical
thought to early Greek philosophers like Plato (428–348 BC)
and Aristotle (384–322 BC) and, eventually, to scholars like
Linnaeus (1707–78) who first articulated theoretical concepts
of the nature of species (Essentialism and Typology) and then
logically asked why certain species occupied particular
places. A century prior to the publication of Darwin’s and
Wallace’s formulation of the idea of organismal evolution
by natural selection (Darwin, 1859; Wallace, 1860), Georges-
Louis Leclerc, Comte de Buffon (1707–88) clearly articulated
the fundamental principles of natural selection but delib-
erately deemphasized their significance in light of societal and
religious conventions of his day (sixteenth century Europe).
Buffon observed distributions of extant organisms but also
studied fossils, leading him to the inevitable conclusion that
life changes with time, and varies geographically, often in-
hibited by physical barriers. Based on his observations, Buffon
issued what has become known as Biogeography’s First Prin-
ciple (or ‘Buffon’s Law’), the idea that environmentally similar
(but isolated) regions of Earth have distinct assemblages
of animals. This simple observation, sometimes summarized
as The Uniqueness of Place (Lomolino et al., 2010), became so
influential that, in large part, it spawned an Age of Exploration
and more than a century of worldwide travel and discovery
by several generations of European cartographers, mariners,
botanists, and zoologists. Although the first generation of
these explorers was primarily composed of botanists, or
Phytogeographers, vertebrate zoologists soon joined their ranks
and the field of Zoogeography (biogeography of animals)
was born.
At the intersection of numerous physical sciences relating
to both life and the geographical template on which it has
evolved (Lomolino et al., 2010), the field of biogeography
addresses fundamental responses of life to the spatially vari-
able biotic and abiotic environment. As the focus of many
biogeographical hypotheses, physical barriers, environmental
Encyclopedia of Evolutionary Biology, Volume 1 doi:10.1016/B978-0-12-800049-6.00123-2
Gondwana The supercontinent, formerly composed of
today’s Africa, South America, India, Antarctica, Australia,
Arabian peninsula, New Zealand, Madagascar, New Guinea,
and other smaller landmasses.
Tetrapods The four-limbed vertebrates and descendants,
including the living and extinct mammals, birds,
amphibians, reptiles, and some extinct fishes.
Vicariance Passive divergence of evolutionary lineages,
resulting from the formation of a barrier or geological
processes; the splitting of formerly united species
distributions via continental fragmentation and drift.
gradients, atmospheric and temperature extremes, seasonal,
latitudinal and elevational variability, and interactions with
other organisms are often key to understanding both the ex-
tent and limits of species distributions. On land, vertebrate
animals respond in a wide variety of ways to environmental
limiting factors, geographic barriers, and the presence of other
species. The author’s purpose here is to review a small subset
of the many ways land vertebrates have responded – and
continue to respond – to the geographic template. The author
will discuss several bodies of work, which have influenced the
development of the field in important ways. However,
although biogeography might once have been viewed pri-
marily as an historical science, it is now characterized by the
melding of fundamental macroevolutionary and macro-
ecological principles, their associated bodies of theory, schools
of thought, and massive amounts of empirical data, now
rigorously evaluated in an increasingly integrative, hypothesis-
testing framework (Ladle et al., 2015). Most of these bio-
geographical elements became so influential as a result of the
careers, big ideas, and hard work of scores of individuals,
working in disparate disciplines. Thus, students of bio-
geography necessarily must constructively gather information
from a variety of sources to glimpse the ‘big picture’; it is only
relatively recently that textbooks with titles including the term
‘Biogeography’ have been published (e.g., Lomolino et al., 2010;
this volume). To understand the discipline’s foundational
concepts, the reader is referred to these texts as a first reference,
but then necessarily to the primary literature in geology,
geography, earth history, phylogenetic systematics, ecology,
and evolution.
The First Vertebrates
Vertebrates first appeared approximately 500 million years
ago, during the Cambrian explosion. The first forms were fish-
like jawless marine organisms with cartilaginous internal
skeletons, and bony, plated, external armor; these were the
Ostracaderms. True fish, with bony internal skeletons, and
211
212 Biogeography of Vertebrates
eventually functional jaws, first evolved more than 450
million years ago in the Ordovician, becoming quite common
in the Devonian. These early fish, the Gnathostomes, pos-
sessed moveable opposing jaws, teeth, paired appendages,
and a modern vertebrate immune system. The development
of an opposable and functional grasping jaw (derived from
anterior gill support arches), and many additional associated
feeding specializations, was a major advance that occurred
in the Gnathostome fishes approximately 420 million years
ago and allowed these organisms to exploit a broad array
of feeding environments, and evolving into possibly as
many as 65 000 to 75 000 species (Westneat et al., 2005;
Pough et al., 2009), including the Chondrichthyes (carti-
laginous fish) and Osteichthyes (bony fish) which are further
divided into the Sarcopterygii (lobe finned fish) and Acti-
nopterygii (ray finned fish) of today. The lobe finned fishes
include fossil and extant lungfish and coelacanths that we are
familiar with today, but it is a pair of fossil taxa, Eusthenopteron
and Panderichthys, that appears most closely related to the
series’ early tetrapods (e.g., Tiktaalik, Acanthostega, Ichthyostega,
Tulerpeton; 390–360 mya) that made the evolutionary transi-
tion from an entirely aquatic lifestyle to an at least partially
terrestrial, weight bearing, upright land vertebrate (Radinsky,
1987; Laurin, 2010).
Vertebrates invaded land at about 350 million years ago,
giving rise to many forms we know from extant lineages and
extinct fossils; possibly as many as 70 000 vertebrate species
have been formally characterized (with tens of thousands
more implied by trace fossils and other evidence). The next 10
million years was a period of major evolutionary radiations of
amphibians, giving rise to two large groups, the large and ro-
bust labyrinthodonts and their slender, delicate relatives, the
lepospondyls. Much of the initial Carboniferous and Permian
fossil amphibian diversity went extinct at the end of the Per-
mian (although a few lineages survived into the Triassic and
one into the Jurassic), and a variety of hypotheses exist to
explain how these dominant and diverse forms are related to
the amphibians of today (the Lissamphibia: frogs, sala-
manders, and caecilians; Duellman and Trueb, 1994; Marja-
nović and Laurin, 2007; Sigurdsen and Green, 2011).
Approximately 50 million years later, a lineage of labyr-
inthodonts evolved the amniotic egg and the ability to re-
produce away from water. Together with the evolution of
flexible scales composed of keratin, early reptiles were equip-
ped to invade the interior of landmasses, arid environments
(Laurin and Reisz, 1997).
Because of the timing of their initial diversification on land,
terrestrial vertebrate diversification was initially impacted by
the breakup of Pangaea and Gondwana (Figure 1), processes
that resulted in isolation and unique early radiations of land
vertebrates on individual landmasses. At 213 million years
ago, the Triassic–Jurassic extinction event resulted in the loss
of the labyrinthodont amphibians and most marine reptiles,
leaving the majority of terrestrial vertebrate lineages limited to
dinosaurs, who subsequently radiated evolutionarily into
many of the vacant niches left empty following these extinc-
tion events.
As landmasses moved farther and farther apart, their ver-
tebrate faunas became increasingly distinct taxonomically, and
biogeographic patterns can be related to habitat types and
Arabian
Peninsula
Madagascar
Africa
New
India Guinea
South America
Australia
Antarctica
New Zealand Tasmania
Figure 1 The breakup of the supercontinent Gondwana,
200–180 million years ago, with the outlines of today’s major
landmasses indicated. Adapted in part from Chatterjee, S.,
Goswami, A., Scotese, C.R., 2013. The longest voyage: Tectonic,
magmatic, and paleoclimatic evolution of the Indian plate during
its northward flight from Gondwana to Asia. Gondwana Research 23,
238–267.
paleoclimates to the extent that these can be reconstructed. The
assumption that early vicariance, or cladogenetic divergence as
a result of geological processes (splitting of formerly united
species distributions via continental fragmentation and drift)
primarily explained dinosaur distributions (Sampson et al.,
1998) was challenged by Sereno (1999), who suggested that
local extinction and dispersal may have been major ‘controls’
of dinosaur land distribution, bringing into question the issue
of whether a truly cosmopolitan dinosaur fauna ever existed.
However, Upchurch et al. (2002) demonstrated that major
vicariant divisions of dinosaur fauna were all a result of
(1) the separation of Asia from Pangaea in the Early–Mid
Jurassic; (2) the fragmentation of North America from
Gondwana; (3) the separation of Australia (formerly eastern
Gondwana) from Africa þ South America (western Gondwana);
and (4) the fragmentation of Africa from South America
during the Early–Mid Cretaceous. Meanwhile, the signature
of intercontinental dispersal and subsequent diversification
becomes apparent in the fossil record and numerous examples
of dinosaur dispersal have been documented (Rowe et al.,
2011; De Queiroz, 2014).
Following the Cretaceous–Tertiary (K–T) asteroid impact at
66 mya, the loss of two-thirds of all Earth’s diversity, and the
extinction of all non-avian dinosaur diversity, a small group of
terrestrial vertebrates (small mammals, birds, reptiles, am-
phibians) remained. In the absence of dinosaur and Mesozoic
reptile dominance, birds and mammals soon flourished and
occupied the majority of terrestrial niche space. The Paleogene
(66–23 mya) marked a period characterized by the onset of
many classic terrestrial mammal evolutionary radiations and
biogeographic range expansions, presumably in response to
the ecological opportunity presented by widespread extinc-
tions of the dinosaurs. These include the sudden and prolific
diversification of many clades of today: whales, bats, horses,
and primates (Alroy, 1999). Extinctions were selective in
some clades in particular areas. For example, marsupials
largely disappeared from North America, and Laurasian

multituberculate mammals (a large group that had survived
the previous 120 million years) went extinct, as did southern
hemisphere Gondwanatherian mammals. Chiropterans, mar-
supials, and Cetartiodactyla (whales and even-toed ungulates),
in contrast, may have diversified and expanded their
global distributions in response to the K–T extinction
(Bininda-Emonds et al., 2007).
Plate Tectonics, Continental Drift, and the
Paleotransport of Vertebrates
As discussed above, plate tectonics, rift formations, and the
breakup of Pangaea, Laurasia, and Gondwana (Figure 1) had
profound impacts on the distribution of a wide variety of early
land vertebrate lineages. These tectonic events initially struc-
tured vertebrate assemblages, rendering land vertebrate species
pools unique and identifiable on a continental scale. For
example, as the southern continents successively fragmented,
the taxonomically and biogeographically unique faunas
became isolated and developed their own faunistic identities
as a result of millions of years of isolation coupled with in situ
evolution and faunal exchanges facilitated by plate collisions
and landmass accretions. Mesosaurus, for example, was one
of the first aquatic reptiles inferred to be a coastal habitat
specialist, documented in fossil beds of southern Africa and
eastern South America. Part of the celebrated ‘Glossopteris’
floral/faunal assemblage, including Lystrosaurus (with an In-
dian, African, and Antarctica distribution) and Cynognathus
(distributed in South America and Africa), the group has been
cited as direct evidence in support of the hypothesis that the
southern continents were once joined and that Gondwanan
vicariance led to the primary patterns of geographical distri-
bution in terrestrial vertebrates (Laurin, 2010).
Phylogenetic analyses of extant taxa have also been used
to provide inference into the question of continental frag-
mentation and drift as a major initial causal factor for
understanding land vertebrate distributions. The lungfishes
(Lepidosireniformes) of South America, Africa, and Australia
might appear to be an ideal group with a classic Gondwanan
distribution; however, recent fossil evidence suggests that this
group was formerly widely distributed, with a modern day
distribution resulting from widespread extinction subsequent
to the breakup of the supercontinents. Despite the many re-
interpretations of previously assumed Gondwanan land ver-
tebrate distributions, paleotransport of vertebrates on drifting
continental fragments continues to be inferred in modern
phylogeny-aided biogeographical inference and involving
younger time scales (Bossuyt and Milinkovitch, 2001; Bossuyt
et al., 2006; Gamble et al., 2008; Blackburn et al., 2010; Siler
et al., 2012; Brown et al., 2016). In a recent review and ‘event-
based’ biogeographic analysis, Sanmartin and Ronquist
(2004) demonstrated the surprising result that animals more
closely followed order-of-Gondwanan landmass fragmen-
tation predictions than did plants. Other single-taxon analyses
have uncovered the apparent result of pure Gondwanan
vicariance as a result of the tectonic fragmentation of former
supercontinents. For example, Turner (2004) interpreted a
phylogenetic analysis and documented distributions of cro-
codyliform taxa (notably, including both fossil and extant
Biogeography of Vertebrates 213
taxa) as evidence for pure Gondwana fragmentation affecting
the primary diversification of a major class of vertebrates
during the Mid to Late Cretaceous (see also Gamble et al.,
2008, for a similar example in geckos).
Vicariance and Dispersal: Theory versus Data
The influential biogeographer Léon Croizat is noted for his
unwavering faith in the idea that most of the Earth’s disjunct
distributions could be explained by the breakup of former
supercontinents, leading to isolation, divergence, and the
formation of new lineages (vicariance). He famously sum-
marized this with his dictum: “earth and life evolve together”
(Croizat, 1962). Motivated and inspired by some of the
Planet’s most celebrated disjunct distributions, Croizat made
major advances in Panbiogeography as he strove to explain
shared patterns of endemism and provincialism. How could
closely related taxa (species, genera, families) come to occupy
separate and isolated geographical ranges on landmasses on
either side of the world’s major ocean basins? Operating in
the absence of rigorous phylogenetic methods or today’s
sophisticated and time-calibrated estimates of evolutionary
relationships, but with an appreciation of the emerging
understanding of plate tectonics, Croizat was led to the con-
clusion that disjunct distributions were the direct evidence of
vicariance. Focusing on patterns of endemism and disjunction
shared by many unrelated organisms with a simple procedure
he termed ‘track analysis,’ designed to summarize these in-
stances of shared disjunctions, Croizat was convinced that
virtually all groups of terrestrial plants and animals arrived at
their current distribution by paleotransport on the fragments
of former supercontinents. At the time, the standardization
of a formal, repeatable method for analyzing distributions
was novel and represented a substantial advance over the
descriptive and taxon-by-taxon, idiosyncratic biogeographical
studies that preceded. Croizat’s work also heavily influenced
the development of the school of Vicariance Biogeography
(Nelson, 1973; Nelson and Platnick, 1981), a dominant per-
spective in the field for more than 30 years. According to this
philosophical perspective, dispersal was virtually dismissed as
exceedingly rare or improbable, barely considered except in
passing, with the study of dispersal-related biology even ridi-
culed as being ‘unscientific’ and considered only by researchers
who were prone to fantasy. In much the same way that we
now look back at the period in the history of biogeography
when workers inferred hypothetical landmasses (such as land
bridges) on the basis of current day species distributions
as overly simplistic and naively misdirected, an emerging
mainstream of biogeographers similarly view Croizat’s Pan-
biogeography and the Vicariance Biogeography school as
an interesting but peculiar period in the history of the dis-
cipline. In retrospect, how could we possibly have dismissed
the possibility (and ignored the evidence) that organisms
do naturally disperse long distances, even across massive
geographical barriers such as mountain ranges, deserts, and
oceans (de Queiroz, 2005; Gillespie et al., 2012)? As Lomolino
et al. (2010) discuss, the period during which these ideas had
mass appeal (1960s through 1980s) marks a time of great
sociological and philosophical debate in systematics: a time
214 Biogeography of Vertebrates
in which the impact of many novel contributions was some-
what diminished by strong, domineering personalities, who
aggressively pursued winning in debate in order to promote
their beliefs. Vertebrates, especially those embodying relative
dispersal ability – amphibians, flightless birds, freshwater fish,
etc., – figured heavily into this heated discussion.
Patterns: Endemism, Provincialism, Cosmopolitanism,
and Disjunct Distributions
The search for explanations of biogeographic patterns –
emergent properties of species distributions – inspired the first
true biogeographers and contributed directly to the conception
of evolution by natural selection (Darwin, 1859; Wallace,
1860, 1876, 1881). As the nineteenth century drew to a close
and the newly acquired species distribution data from the age
of exploration became available, vertebrate biologists noted a
series of conspicuous shared patterns of species distributions.
The ‘Father of Biogeography,’ A. R. Wallace, and noted ver-
tebrate naturalist P. L. Sclater, are generally credited with
having defined the major terrestrial biogeographical regions or
‘realms’ of the planet, among many other distribution patterns
(Wallace, 1860, 1876; Sclater, 1858). Chief among these were
the recognition of vertebrate endemism, biogeographical
provincialism (or regionalism), and range disjunctions, all at
varieties of scale.
The discovery and recognition of mammal, bird, amphib-
ian, and reptile species with geographical distributions limited
to a particular landmass like a small island (and not occurring
anywhere else on Earth) is compelling evidence for the im-
portance of historical events (such as colonization of a far
away island by a vertebrate species) and the contribution of
isolation to the processes of speciation. For example, the ob-
servation that the primate species the Red Slow Loris (Loris
tardigradus) occurs only on Sri Lanka, that Melanesian Forest
Frogs (genus Cornufer) are largely endemic and confined to
islands of the Southwest Pacific, or that kangaroos and wal-
labies (marsupial mammals of the family Macropodidae) are
limited primarily to the Australian continent, are all examples
of the way in which varying taxonomic levels exhibit endem-
ism at different scales. These examples contrast strongly with
the nearly worldwide distribution of bats (order Chiroptera), a
true case of cosmopolitanism in a terrestrial vertebrate species.
When many species or even an entire biota are influenced
by such historical events, the resulting concordant patterns of
endemism are often referred to as biogeographical provincialism
or regionalism. Patterns of endemism are concentrated non-
randomly and usually are associated with particular areas. This
co-occurrence of many endemic forms is sometimes also as-
sociated with particular biotic characteristics. Provincialism is
a pattern that informs biogeographers about the significance of
geographic barriers and their impact on numerous related and
unrelated species. To return to the examples provided above,
biogeographic provinces have been defined by isolation of
biotas on islands (Bossuyt et al., 2006). As a result of isolation,
12 named primate species or subspecies are endemic to the
island of Sri Lanka, for example, and occur nowhere else
on Earth (Nekaris and de Silva Wijeyeratne, 2009). Similarly,
95 þ % of amphibians native to the Southwest Pacific’s
island archipelagos (the Solomon Islands, the Bismarcks,
Admiralties, and Fiji) are endemic to those landmasses
(AmphibiaWeb, 2015) and 90 þ % of Australian mammals
exhibit concordant distributions, primarily endemic to this
landmass (Nowak, 1999).
Empirical Insights into the Biology of Long-Distance
Vertebrate Dispersal
Well-known disjunct distributions in which vertebrate geo-
graphical ranges are discontinuous and usually divided by a
substantial geographical barrier have inspired some of the
most conceptually intriguing biogeographical studies and
the discipline’s most fervent debates. At the heart of these
heated debates have been two basic alternate biogeographical
hypotheses, namely the interpretation of biotic disjunctions as
necessarily the result of ancient vicariance versus the possi-
bility that these patterns lend support for the hypothesis of
recent dispersal (e.g., Yoder and Nowak, 2006; Sanmartin and
Ronquist, 2004). A third possibility in some systems, the in-
terpretation of a formerly more widespread distribution, fol-
lowed by extinction of populations or species in intermediate
areas, has been invoked in terrestrial vertebrates exhibiting
disjunct distributions on continents where climate change as-
sociated with glacial retreats caused shifts in major habitat
types (e.g., mesic versus xeric forests; Nielson et al., 2001).
The mechanisms of dispersal over marine barriers vary, and
some authors have emphasized organismal or ecological traits
that render certain vertebrates more prone to active or passive
transport (Carlquist, 1965, 1966). For vertebrates capable of
surviving exposure to salt water and sun, dispersal is achieved
by simply floating and traveling passively with currents
(Gerlach et al., 2006). For land vertebrates of low relative
dispersal ability and salt tolerance like nonvolant mammals,
amphibians, or freshwater fish, the common assumption is the
transport of animals on natural flotsam associated with strong
storms. These include floating islands (large portions of river
banks that have broken free, and floated out to sea), mats of
vegetation, and/or log jams and tangled masses of tree trunks,
which are often discharged from river mouths following
flooding on land (King, 1962; Krause et al., 1997). The
occurrence of floating debris fields, coupled to the discharge of
large amounts of freshwater, can create stratified freshwater
‘lenses’ (layers) of freshwater that form above denser salt
water, and create temporary freshwater currents – all of which
may favor the transport of salt-intolerance terrestrial ver-
tebrates. Such mechanisms have been either directly observed
(Censky et al., 1998) or inferred by testing predictions with
storm track data, localized flow patterns in the vicinity of river
deltas, information from prevailing ocean currents, and phy-
logeographic data (Measey et al., 2007; Bell et al., 2015). The
key to understanding the plausibility and importance of these
supposedly rare or ‘improbable’ events is the simple fact that
even seemingly unlikely phenomena may become easier for us
to consider when their ‘occasional’ successful occurrence is
multiplied by many millions of years (De Queiroz, 2014).
Several developments have contributed over the last three
decades to the unraveling of the dominance of the Vicariance
Biogeography paradigm; most of these directly involve

empirical studies of dispersal involving iconic land vertebrates
(e.g., Townsend et al., 2011). First, several of the most ardent
promoters of this perspective have retired, moved on to other
areas of research, or softened their perspectives. Additionally,
the development of modern model-based molecular phylo-
genetic analysis has changed our understanding of the
evolutionary relationships of most vertebrate clades, either
radically by turning evolutionary trees on their head (e.g.,
squamate reptiles; Townsend et al., 2004; Losos et al., 2012),
or by demonstrating widespread convergent evolution and
repeated evolution of vertebrate body form (such as repeated
evolution of elongate, limb-reduced/absent snake-like lizards),
which misled previous morphological trait-based studies
(Reeder et al., 2015). Third, biologists have developed a
better understanding of how vertebrates of seemingly low
relative dispersal ability actually do achieve overseas dispersal
(Carlquist, 1966; McDowall, 2002). Finally, as calibrated
molecular divergence date estimation has become more
reliable due to advances in methods and collection of large
robust multilocus datasets, many vertebrate groups of appar-
ent, ancient Gondwanan origin have simply turned out to be
too young. In these instances where disjunct groups on either
side of an ocean basin are many millions of years younger
than the dated fragmentation of the landmasses in question,
an explanation of vicariance becomes intractable.
One such clear case is illustrated by estimation of the split
between Old World and New World monkeys. For a vicariance
explanation to account for the distribution of these African
and American taxa, the divergence between the clades would
have to approximate the final separation of Africa and South
America (B110 mya). One recent time-calibrated phylo-
genetic analysis dated their divergence at somewhere between
50 and 26 mya (Springer, 2012), meaning that the ancestors
of New World monkeys (now about 55 species) crossed the
Atlantic at about 40 mya, much too late to permit the former
vicariance scenario.
Ratite birds (the Rheas and Tinamous of the Americas, the
Ostriches of Africa, the Emus and Cassowarys of Australasia,
and the Kiwis and extinct Moas of New Zealand) are a classic
and often invoked case of diversification via vicariance
(Cracraft, 1974). Their phylogenetic relationships (a mono-
phyletic clade), the preponderance of flightlessness in most
species, and their distributions on separate southern isolated
Gondwanan landmasses, has been invoked as a classic case of
diversification necessarily resulting from continental-scale
vicariance (Craw et al., 1999). Dispersal via powered flight in
contrast was comfortably discounted due to the clade’s ten-
dency toward large bulky bodies and the inability of extant
species to fly. At the center of these assumptions were the giant
extinct Moas of New Zealand, which vicariance biogeographers
confidently pointed out could not possibly have reached New
Zealand by any means other than overland colonization when
this island was still connected to Australia and Antarctica.
However, a recent phylogenetic analysis using a massive
genomic dataset recently has resulted in a major rearrange-
ment of the group’s family tree. Together with a analysis
of ancestral evolutionary state reconstruction Phillips et al.
(2010) estimated that flightlessness actually might not have
been the ancestral condition in this group, leading to a revised
interpretation of multiple recent losses of flight in individual
Biogeography of Vertebrates 215
lineages. This opens the door to the very real possibility that
the ancestors of these iconic large-bodied flightless birds may
have, in fact, dispersed to some of their current locations via
powered flight over open oceans – after which they sub-
sequently evolved large body sizes and lost the ability to fly.
The case emphasizes the importance of evaluating alternate
evolutionary hypotheses in conjunction with biogeographical
studies. And, just because many ratite birds are large and
flightless today does not necessarily mean that their ancestors
were giants, incapable of flight. And once these seemingly ‘self-
evident’ constraints are relaxed (i.e., giant flightless Moas
‘obviously’ could not have flown to New Zealand), biogeo-
graphers can consider the alternate explanations (in this case,
dispersal) for ratite Gondwanan distributions previously
assumed to have resulted from pure vicariance (Craw et al.,
1999; Phillips et al., 2010; De Queiroz, 2014).
Another frequently invoked case of Gondwanan vicariance
is the classic case of more than 1600 freshwater cichlid fishes
inhabiting southern portions on a number of continents. Be-
cause of their distribution and an early branching pattern
mirroring the order of fragmentation of Gondwanan land-
masses, cichlid fish have been widely accepted as a group,
whose distribution arose from Gondwanan vicariance, starting
at 135 mya. Friedman et al. (2013) presented a combined but
independent paleontological and molecular-clock estimate for
the time of the clade’s origin that soundly rejected the assumed
Gondwanan vicariance scenario in cichlid fishes. In this case,
both stratigraphic distribution of fossils and the age inferred
from a robust time-calibrated analysis of DNA sequence
divergences both agreed on a Paleocene (65–57 mya) origin of
the group, suggesting a primary role for dispersal in generating
the observed distribution of cichlid fish. Although the case of
cichlid fish left some questions unanswered (the Friedman
et al., 2013, analysis did not include fossil taxa), ichthyologists
generally agree that the Rainbowfishes, the Gobies, and the
Killifishes are most likely all pure Gondwanan in origin.
Yoder and Nowak (2006) reviewed the entire terrestrial
flora and fauna of Madagascar, a celebrated and spectacularly
isolated landmass, formerly tucked into the heart of Gon-
dwana (Figure 1), and often cited as a prime example of a
biota derived from pure vicariance, preventing exchange with
African fauna following the separation of the island from this
continent 135–125 million years ago. Reviewing all available
molecular divergence data the authors concluded that the vast
majority of the fauna – including freshwater fish and am-
phibians with their presumed low relative dispersal abilities –
was derived from descendants of more recent (Cenozoic)
transoceanic dispersal events from Africa.
Many similar and well-studied vertebrate groups, once
widely accepted as clear cases of Gondwanan or Pangaean
vicariance by virtue of their modern distributions and pre-
sumed poor dispersal abilities, have likewise failed to stand
the test of molecular phylogenetics and modern divergence
date estimation (Rowe et al., 2010; Townsend et al., 2011;
Friedman et al., 2013). Summarizing many of these studies,
De Queiroz (2014), with reference to Croizat’s dictum, stated:
“Earth and life evolve together—except when they don’t.”
With time and new advances, biogeographers have come to
routinely embrace necessarily more sophisticated analytical
approaches and much more pluralistic interpretations.
216 Biogeography of Vertebrates

However, the influence of Panbiogeography and Vicariance
Biogeography is still felt in curious ways. For example, only
recently have probabilistic modeling methods for bio-
geographic inference been developed with explicit parameters
that model long-distance dispersal to a novel area (such as a
distant island, not previously or currently occupied by the
clade of interest; Ree and Smith, 2008; Matzke, 2013a,b). The
assumption is that this methodological shortcoming may be
an historical and sociological artifact, dating back to a time
when mention of the word ‘dispersal’ would earn a young
biogeographer a swift and public rebuke from a siliverback
member of the Vicariance Biogeography school.
Biogeographic Barriers, Interchanges, Gradients, and
Filter Zones: Stepping Stones, Conduits, and Corridors
Biogeographic barriers, as discussed above, inspired the
scholarship that resulted in the birth of the field and the first
articulation of biogeography’s first principles (Wallace, 1860).
The inferred barriers themselves vary in assumed permeability,
from the absolute apparent turnover characterized by Wal-
lace’s Line (Huxley, 1868) to the various biogeographic lines
that divide the major realms of the planet (Wallace, 1860,
1876; Sclater, 1858). The universality of absolute bio-
geographic lines has been discussed at length (Mayr, 1944;
Simpson, 1977) and is tempered with a wide variety of
exceptions (Esselstyn et al., 2010; Lohman et al., 2011; Brown
et al., 2013).
The permeability of biogeographic barriers has been
studied at length, mostly in the context as the concept
was originally defined: impenetrable barriers across which
land vertebrate dispersal is limited by marine channels
(e.g., freshwater fish or amphibians, believed to be wholly
incapable of dispersal over salt water). For example, Esselstyn
et al. (2010) analyzed taxonomic and phylogenetic evidence
for all terrestrial vertebrates with ranges abutting either side
of the northern end of Wallace’s Line (i.e., Huxley’s Modifi-
cation of Wallace’s Line; Figure 2) and argued that the
immediate area (Palawan Island, and smaller landmasses
to the north and south of this elongate corridor) more accur-
ately meets the definition of a biogeographic filter zone and
not an absolute biogeographic line or barrier. Classic bio-
geographic filter or ‘transition’ zones have been defined by
linear island chains such as the Lesser Sunda islands of eastern
Indonesia, or the Alaskan Aleutian island chain, stretching
between distinct biogeographic provinces (Kricher, 2011;
Lomolino et al., 2010). In such systems, as the number
of faunal elements from one end of the chain declines
successively with distance form the source, the influence
and origin of inferred elements from the opposing region
increase (Figure 3; Carlquist, 1965; Lomolino et al., 2010).
Filter zones, whether island chains or matrices of suitable
habitat surrounded by inhospitable habitat (such as the ‘Sky

Mainland Huxley’s modification

Southeast of wallace’s line
Asia

The Philippines

Malay Wallace’s line

Peninsula
Weber’s line

Su Borneo Lydekker’s line

m
at
ra

Java
Bali Lombok

Australia

Figure 2 The Australasian faunal interchange zone is home to the most celebrated biogeographic lines (funal turnover zones or biogeographic
barriers) on the Planet. Wallace’s Line, as originally conceived (Wallace, 1860) illustrates the faunal turnover that occurs at the eastern edge of the
continental Asian landmass, Sunda Shelf (light blue shading, left). This feature corresponds to the land that was exposed during Pleistocene
periods of lowered sea levels. This biogeographic boundary is now recognized as the area of transition between Asian fauna and that of Wallacea,
the transitional faunal zone that separates Asian from Australian fauna. Lydekker’s Line is the equivalent feature on the Australian Sahul Shelf (light
blue shading, right), and Weber’s Line represents the geographic point of equivalency or balance between Asian and Australian faunas. Adapted
from Mayr, E., 1944. Wallace's Line in the light of recent zoogeographic studies. The Quarterly Review of Biology 19, 1–14; Simpson, G.G., 1977.
Too many lines; the limits of the Oriental and Australian zoogeographic regions. Proceedings of the American Philosophical Society 121, 107–120.

Indonesia Lesser sunda Island Chain
Figure 3 A terrestrial biogeographic filter zone is formed by areas of
suitable habitat (such as dry land), separated or surrounded by an
inhospitable matrix (sea water). The fauna of the Lesser Sunda islands
of Indonesia (inset) illustrate the concept well: a chain of small
landmasses separated by ocean channels, with the dispersal abilities
of species demonstrated by their successful colonization of more and
more distant islands. With dispersal from both ends of the chain of
islands contributing to each island’s fauna, the proportion of Asian
species (light blue) nearly balances that of Australian species (dark
blue) on midway through the chain. Adapted from Carlquist, S., 1965.
Island Life. Garden City, NY: Natural History Press.
Islands’ of the Madrean Archipelago; Manthey and Moyle,
2015) selectively illustrate the variation in relative dispersal
abilities of the various taxonomic groups and the degree to
which environmental filtering strongly structures community
assembly along edges of biogeographic transition zones
(Sommer et al., 2014).
Other biogeographic island chains have been viewed as
dispersal conduits or suitable habitat corridors, possibly
allowing ‘stepping stones’ dispersal or ‘island hopping’ from
one region to another (Carlquist, 1966). Dispersal conduits
between continents and island archipelagos have been iden-
tified in empirical studies. For example, Diamond and Gilpin
(1983) and Brown and Guttman (2002) (see also Brown and
Siler, 2013) identified multiple chains of islands, which have
facilitated faunal exchange of amphibians, reptiles, birds, and
mammals between the mainland Southeast Asian continent
and adjacent island archipelagos (see also Taylor, 1928; Inger,
1954). Comparative biogeographic analyses indicate that these
series of terrestrial stepping stones have served as the primary
source for mainland colonization of oceanic archipelagos
(Brown and Siler, 2013; Brown et al., 2013). The conservation
significance of stepping stones and corridors of suitable
habitat patches have been broadly discussed in insular terres-
trial systems in terms of their importance for long-distance
vertebrate dispersal – especially in light of species distri-
butional shifts anticipated under climate warming scenarios as
species track suitable habitats (Saura et al., 2014).
Insular Extremes: Spectacular Vertebrates of Isolated
Islands and Extraordinary Vertebrate Radiations in
Island Archipelagos
Studies of vertebrate island biogeography have involved some
of the planet’s most interesting and unique forms of life. Three
Biogeography of Vertebrates 217
(a) (b)
(c) (d)
Figure 4 An example of an adaptive radiation in frogs from an island
archipelago. Narrow-mouth frogs (genus Kaloula) have been
characterized as an adaptive radiation, which began as their ancestors
invaded an island archipelago (Blackburn et al., 2013). From an
inferred terrestrial mainland Asian habitat generalist, the Philippine
Kaloula evolved into small, ephemeral pool breeding montane
species ((a) Kaloula walteri), large-bodied open habitat burrowing
species ((b) Kaloula picta), delicate scansorial shrub-breeding species
((c) Kaloula conjuncta), and tree-hole cavity dwelling taxa ((d) Kaloula
kalingensis). Photographs by the author.
primary phenomena make this so. First, as previously dis-
cussed, the inferred origins of vertebrates on Earth’s most
isolated landmasses capture the imagination of evolutionary
biologists and beg for an historical explanation that is in-
herently interesting – whether an ancient vicariance scenario or
an inference of a spectacularly long-distance dispersal event.
Second, because of extreme isolation and the unique en-
vironments of islands, island vertebrates have evolved in many
cases into bizarre forms, with phenotypic novelty accentuated
and exaggerated by millions of years under unique selective
regimes. Celebrated examples include island gigantism in
tortoises, birds, and shrews, and at the other end of the spec-
trum, miniaturization in formerly large-bodied lineages like
mammoths. Other examples of the ‘Island Syndrome’ involve
inferences such as the secondary loss of flight in numerous
lineages of birds after having arrived on islands, or other novel
differences in life history, reproduction, demography, or be-
havior of island populations as compared to closely related
species on the mainland (Adler and Levins, 1994).
Third, the uniquely replicated environments and at times
unutilized resources (i.e., simplified food webs lacking top
predators) of insular systems appear to have had an extra-
ordinary impact on the process of diversification in vertebrates
of island archipelagos. A disproportionate portion of the Earth’s
well-studied and truly spectacular adaptive radiations are
found in islands or island-like archipelagos, and include highly
celebrated clades, such as Darwin’s finches in the Galapagos
(Grant and Grant, 2008), the Hawaiian honeycreepers (Lerner
et al., 2011), New World direct-developing frogs (Heinicke
et al., 2008), the mantelid frogs of Madagascar (Vieites et al.,
2009), the great cichlid fish radiations of African rift valley
lakes (Turner, 2007), lizards of the genus Anolis (Losos et al.,
1998; Mahler et al., 2010), and the markedly diverse rodents,
218 Biogeography of Vertebrates
narrow-mouth frogs, and forest frogs of the Philippines (Jansa
et al., 2006; Blackburn et al., 2013; Brown et al., 2015; Figure 4).
As such, it should be no surprise that islands not only
spawn the big radiations, but they also inspire the big ideas.
The geographic setting for much of our current speciation
theory was the island archipelagos of the Pacific (Darwin,
1859; Wallace, 1860; Mayr, 1942), and some of evolution and
ecology’s most conceptually compelling, classic ideas were
first conceived through the study of organisms on islands.
These include topics such as equilibrium biogeography theory
(Macarthur and Wilson, 1963), community assembly rules
(Macarthur and Levins, 1967), adaptive radiation (Glor,
2010), ecological opportunity (Mahler et al., 2010), evolution
of supertramps (Toussaint et al., 2013), taxon cycles (Wilson,
1961), and the Earth’s great speciators (Diamond et al., 1976;
Moyle et al., 2009). The importance of insular systems for
development of conceptual models of processes of evo-
lutionary biology cannot be over-emphasized (Vences et al.,
2009; R. Brown et al., 2013; J. Brown et al., 2014), and the
study of land vertebrate biogeography historically has pro-
vided the inspiration behind the study of the Earth’s most
iconic island systems.
Future Prospects
The study of land vertebrate biogeography is at a crossroads,
suggesting a synthetic paradigm shift has already begun with
the availability of genomic data, the conceptual merging of
traditions of historical versus ecological biogeography, and
dampening of oscillations of a philosophical pendulum
characterizing a history of debate between vicariance versus
dispersalist perspectives. Although among the major branches
of the Tree of Life the biodiversity of land vertebrates is
comparatively well known, their distributions are less well
documented, and an alarmingly large proportion of our
understanding of mammal, bird, amphibian, and reptile dis-
tributional limits are approximations based on centuries-old,
undocumented, or unverified observational data. This creates a
challenge of great urgency for biogeographers charged with
biotic inventories, surveys, and greatly needed follow-up sur-
veys, and represents a major conservation concern in the face
of species distributional shifts anticipated as a result of climate
and other environmental change. Nevertheless, modern trends
in the field of biogeography indicate that increasingly multi-
disciplinary approaches, massive integrative datasets, extensive
collaboration, and increasingly global coordination and co-
operation – where once competition prevailed – are becoming
standard practice in the research programs of most vertebrate
biogeographers. Because, relative to other groups, vertebrates
are so well studied, with many aspects of their phenotype,
genotype, and ecological characteristics understood by biolo-
gists, land vertebrates promise to serve as the model for
biogeographers interested in studying integrated organismal
biology through the application of multiple data streams.
See also: Land Animals, Origins of. Land Vertebrates, the Origin
and Evolution of
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