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THE RISE OF AMPHIBIANS: 365 MILLION YEARS OF EVOLUTION by Robert L.

Carroll

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A Review of “The Rise of Amphibians: 365 Million Years

of Evolution”
a
J. Sébastien Steyer
a
CNRS–Muséum national d’Histoire naturelle, Département Histoire de la Terre, CP38, 8 rue,
Buffon, 75005, Paris, France

Available online: 08 Nov 2011

To cite this article: J. Sébastien Steyer (2011): A Review of “The Rise of Amphibians: 365 Million Years of Evolution”, Journal
of Vertebrate Paleontology, 31:6, 1406-1407

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 Journal of Vertebrate Paleontology 31(6):1406–1407, November 2011
© 2011 by the Society of Vertebrate Paleontology
THE RISE OF AMPHIBIANS: 365 MILLION YEARS OF EVOLUTION, by Robert L. Carroll, 2009.
The John Hopkins University Press, Baltimore, 392 pp. US$65.00, ISBN: 978-0-801-89140-3.
The evolution of amphibians (‘non-amniotic tetrapods’) is
a fascinating subject for which a large part of the scientific
community has shown a new, tremendous interest. Fossil amphi-
bians (or stegocephalians) lead to all living amphibians (lissam-
phibians), reptiles, mammals, and birds, which are represented by
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approximately 21,100 modern species. Studying stegocephalians
and their terrestrialization involves many fields of earth and life
sciences, far beyond vertebrate paleontology, including sedimen-
tary geology, paleogeography, biomechanics, and developmental
genetics.
For several decades, numerous research teams from all over
the world have been working intensively in original and mul-
tidisciplinary programs, producing major results. Yesterday,
Ichthyostega and Acanthostega were the only known Devonian
tetrapods, and Eusthenopteron was the only ichthyan sarcoptery-
gian traditionally used to depict the fish–tetrapod transition. To-
day, we have more than 10 Devonian tetrapods from all over
the world (e.g., Clack, 2006), without forgetting of course the re-
cent discovery of Tiktaalik from the Devonian of Canada, the
sarcopterygian closest to tetrapods, which helps better explain
the great transition (Daeschler et al., 2006). We know also that
most Devonian tetrapods were not terrestrial, but aquatic or even
marine, with odd polydactylous limbs, and that this polydactyly
could be linked with a life (and a development) in salt water
(Steyer, 2009). We know also, based on newly discovered track-
ways in Poland, that the tetrapod clade must have originated be-
fore the Late Devonian, somewhere during the early Middle De-
vonian (Niedzwiedzki et al., 2010) if not during the Early Devo-
nian (Georges and Blieck, 2011).
If there is a taxonomic group in paleontology that underwent
a ‘revolution,’ with new discoveries, new ideas, and new sce-
narios, the amphibians are number one, perhaps even ahead of
the dinosaurs. But the amphibians, as fascinating they are, re-
main also a very difficult and puzzling scientific group: today, the
origin of lissamphibians and, to a lesser extent that of amniotes,
remain highly debated because there is no consensus about stego-
cephalian phylogeny—the dark side of the amphibians, one might
say.
The Rise of Amphibians is a thick book divided almost chrono-
logically into 14 chapters. Chapters 1 and 2 correspond to a very
precise and updated review of the evolution of the Earth and of
life (entitled ‘History of Earth and Life’ and ‘Advanced Meta-
zoans and the Ancestry of Vertebrates,’ respectively), starting
from the formation of the planet and the origin of life, and ending
with the sarcopterygian Eusthenopteron. Chapter 3, ‘The Origin
of Amphibians,’ starts with Tiktaalik but deals mostly with De-
vonian tetrapods. In these two chapters, the author added inte-
resting sections dealing with genetics, hox genes, and ‘evo-devo,’
pointing out the importance of the neural crest in the origin and
evolution of chordates (Chapter 2) and the essential role of the
famous ‘AER’ (Apical Ectodermic Ridge) in fin or limb develop-
ment. Chapter 4, ‘The Radiation of Carboniferous Amphibians,’
is by far the longest (83 pages), and provides an exhaustive
state-of-the-art coverage of Carboniferous paleoherpetology. All
of the tetrapod groups, with or without limbs, are described and
figured in detail, including enigmatic and aberrant forms such
as Crassigyrinus and Diplocaulus. A special emphasis is given to
1406
BOOK REVIEW
important Carboniferous localities from Scotland, Gilmerton,
Cowdenbeath, and East Kirkton, which have yielded numerous
key stegocephalians of this age. The author also mentions the
main difficulty encountered by those paleontologists working on
fossil amphibians (and that is also probably the main reason why
there is still no agreed phylogeny), the famous ‘Romer’s gap,’
a 30 million year interval within the Lower Carboniferous that
is almost devoid of fossils. Chapter 5, ‘Adaptation, Radiation,
and Relationships,’ is a very personal but constructive section
dealing with the ‘problem’ of phylogeny and proposing another
alternative classification. Chapter 6, ‘The Zenith of Amphibian
Diversity,’ portrays most of the Late Carboniferous and Permian
amphibians with a special focus on my favorite group, the
temnospondyls, whereas Chapter 7, ‘The Origin of Amniotes,’
describes Carboniferous and Permian reptiles without forgetting
their amphibian relatives such as Diadectes. The author ends this
fantastic travel through the stegocephalian world with Chapter 8,
entitled ‘Stereospondyls’ (a clade of advanced temnospondyls),
and by briefly discussing the Permian-Triassic mass extinction.
The following chapters deal with lissamphibians, both extinct
and extant: Chapter 9, ‘The Enigma of Modern Amphibian
Origins,’ points out again, as in Chapter 5, the absence of an
agreed tetrapod phylogeny. It also presents in detail the anatomy
of each lissamphibian group based on living anurans (frogs and
toads), caecilians (gymnophionans), and urodeles (salamanders).
Chapters 10–12 beautifully describe fossil representatives of each
modern group such as the famous Triadobatrachus from the Tri-
assic of Madagascar, the earliest lissamphibian. The reader could
even learn that this key fossil was initially found on the island in a
chief’s hut! In these chapters, the author also attempts to link the
modern groups with Paleozoic taxa he considers as ‘ancestors.’
Chapter 13, ‘The Success of Modern Amphibians,’ deals with
the interrelationships within each modern group (totaling 5948
recognized species today) and their geographic distribution.
And last, in Chapter 14, ‘The Future of Amphibians,’ the author
recognizes the current decline in amphibian biodiversity but
considers predictions for the future as difficult to make (and
therefore not precise), because the amphibians respond too
randomly to environmental changes. Finally, Bob Carroll stays
optimistic and questions the scenario of the soon-to-be extinction
by arguing that the invasive species are all but extinct and that
the amphibians, through their evolutionary history, have already
crossed many rivers, and will cross many others with or without
Homo sapiens.
There is no need to present Robert L. Carroll, worldwide ex-
pert, author, and editor of numerous articles and books dealing
with amphibians and, to a wider extent, with vertebrate evolu-
tion. His books have often been considered as references by many
generations of students, as is the case of Vertebrate Paleontology
and Evolution (Carroll, 1988), which accompanied me through
my university education (even if it needs today a second edition).
Carroll’s books are always masterpieces of comparative anatomy,
and The Rise of Amphibians does not escape this rule: Bob of-
fers us a very precise and comprehensive review on amphibian
anatomy and osteology, and a huge and powerful bibliographic
synthesis that is partly based on his own publications (e.g.,
Carroll, 2007).
 BOOK REVIEWS
The illustrations, consisting mostly of black-and-white figures,
are very precise and of good quality because they are taken
from the literature. But it is unfortunately not the case with the
colored plates that attempt to reconstruct key paleoenvironments
such as Horton Bluff from the Carboniferous of Canada or the
Texas Permian: they are neither realistic nor aesthetic, the an-
imals included are ecologically too numerous, too visible, and,
with other elements such as trees, they just seem to have been
‘cut and pasted’ into landscapes that owe too much to (if not
badly copied from) previous paleo-artworks. It is clear, in com-
parison with other recent books related to the subject, that these
illustrations were not a priority for the editor.
The text is readable, forceful, and very didactic, and it is easy to
recognize a pedagogic talent mixed with the professorial tone of
a teacher who benefits from long experience in science commu-
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nication at the university level. Speaking about the teacher, the
dedication Bob made to the “many students [he has] supervised
[. . .] who have assisted [him] during [his] research and continue to
contribute to [his] education,” well portrays the man, definitively
modest and modern.
As mentioned above, the book is organized almost chronologi-
cally, but the chapter division is sometimes not logical. To render
the reading more fluent, the definition of the choanates should
appear before Chapter 3, and this is also true for Tiktaalik, which
should be described in Chapter 2, together with other ichthyan
sarcopterygians such as Eusthenopteron, not in Chapter 3. This
would show Tiktaalik’s real importance as the closest taxon to the
tetrapod clade, an importance that is sometimes not made clear in
this book. When dealing with the Devonian fossils, Bob does not
mention the tetrapod trackways discovered in the Middle Devo-
nian of Poland (see above) and which push the origin of the tetra-
pod clade far beyond 365 million years, a number he retained in
the book title. Interestingly, however, he seems to question the
assignment of the Devonian Sinostega from China to Tetrapoda
(“another possible tetrapod jaw,” p. 58), although this specimen
has been described as a true tetrapod (Zhu et al., 2002).
The main criticism I have of The Rise of Amphibians deals
with its phylogenetic discussion, which well illustrates Bob’s
personal conception of phylogeny and cladistics. Indeed, his
final result presented at the end of the book (Fig. 12.11,
p. 304) is not testable and does not involve a hypothetico-
deductive approach, as is the case with cladistics. As men-
tioned above, there is no agreed tetrapod phylogeny in the
literature. Three major phylogenies are currently in competi-
tion to explain the origin(s) of lissamphibians: the best sup-
ported (in term of article numbers) proposes temnospondyls
as the sister taxon to the monophyletic lissamphibians (e.g.,
Ruta et al., 2003), whereas a second hypothesis suggests lepo-
spondyls as the lissamphibian sister group (e.g., Laurin, 1998).
The third (and less widespread) suggests that lissamphibians are
polyphyletic; frogs and salamanders (batrachians) being the sis-
ter taxon to temnospondyls, with caecilians allied to lepospondyls
(e.g., Anderson et al., 2008). The latter idea is developed in The
Rise of Amphibians but with a pure stratophenetic approach, and
surprisingly Bob does not cite the work of his ex-Ph.D. student,
Jason Anderson (Anderson et al., 2008), who based this hypothe-
sis upon a cladistic analysis. After using computer-assisted cladis-
tics himself for a while (Carroll 1995, 2007; or Fig. 5.1.A and D,
p. 146 in this book), Bob returned to the use of a stratophe-
netic approach in which fossils must be considered as ‘ancestors’
of living taxa, together linked by a priori relationships based on
stratigraphy. “Another way to investigate the nature of relation-
ships [. . .] is by discussing each of the individual lineages [. . .]
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1407
in a temporal sequence based on the time of first occurrence of
each taxon in the fossil record [. . .]” (pp. 63–64). He explains this
‘return to stratophenetics’ by noting that “The primary lesson to
be learned from these [incongruent] cladograms is that phyloge-
netic analyses using PAUP have failed to provide consistent or
informative answers to the probable interrelationships of the ma-
jor clades of Paleozoic tetrapods” (p. 147), or more amusingly (p.
231): “Even a three-year-old child should be able to differenti-
ate frogs, salamanders, and caecilians. Why was this not possible
using PAUP?”
This very personal conception of phylogeny unfortunately
gives to this book an ‘old school’ style, with gradualist or teleolo-
gist expressions (“sequence of changes” p. 40, “step in evolution”
p. 60, “intermediate forms” p. 144, “directionally selective forces”
p. 145, etc.) and figures directly inherited from Romer (cf. the
trees in Fig. 1.2—the first of the book—and Fig. 9.1). But despite
these phylogenetic criticisms, it can be said that Bob reaches the
same global conclusions admitted by all the ‘fellowship of the
amphibians’: we lack fossils presenting a mosaic of characters and
linking stegocephalians with modern taxa. Finally, with talent,
modesty, and humor (I particularly appreciated his nod to Star
Wars on p. 13), Bob Carroll is telling us, in a modern and very
readable style, the history of the amphibians, with an osteological
precision nobody else has. In fact, with this new book, he shows
us he stays the same, our ‘Master Yoda of the amphibians,’
phylogenetically a bit old school but anatomically very powerful!
J. SÉBASTIEN STEYER
CNRS–Muséum national d’Histoire naturelle
Département Histoire de la Terre
CP38, 8 rue Buffon
75005 Paris, France
LITERATURE CITED
Anderson, J., R. R. Reisz, D. Scott, N. Fröbisch, and S. Sumida. 2008. A
stem batrachian from the Early Permian of Texas and the origin of
frogs and salamanders. Nature 453:515–518.
Carroll, R. L. 1988. Vertebrate Paleontology and Evolution. W. H. Free-
man and Company, New York, 698 pp.
Carroll, R. L. 1995. Problems of the phylogenetic analysis of Paleo-
zoic choanates. Bulletin du Muséum national d’Histoire naturelle
4(17C):389–445.
Carroll, R. L. 2007. The Palaeozoic ancestry of salamanders, frogs
and caecilians. Zoological Journal of the Linnean Society
150(Suppl.1):1–140.
Clack, J. A. 2006. The emergence of early tetrapods. Palaeogeography,
Palaeoclimatology, Palaeoecology 232:167–189.
Daeschler, E. B., N. H. Shubin, and F. A. Jenkins. 2006. A Devonian
tetrapod-like fish and the evolution of the tetrapod body plan. Na-
ture 440:757–763.
Georges, D., and A. Blieck. 2011. Rise of the Earliest Tetrapods: an Early
Devonian Origin from Marine Environment. PlosOne 6(7):1–7.
Laurin, M. 1998. The importance of global parsimony and historical bias
in understanding tetrapod evolution. Part I. Systematics, middle ear
evolution and jaw suspension. Annales des Sciences Naturelles, Zo-
ologie, Paris 13(19):1–42.
Niedzwiedzki, G., P. Szrek, K. Narkiewicz, M. Narkiewicz, and P. E.
Ahlberg. 2010. Tetrapod trackways from the early Middle Devonian
period of Poland. Nature 463:43–48.
Ruta, M., M. I. Coates, and D. L. Quicke. 2003. Early tetrapod relation-
ships revisited. Biological Reviews 78:251–345.
Steyer, J. S. 2009. La Terre avant les dinosaures. Eds Belin, Paris, 203 pp.
Zhu, M., P. E. Ahlberg, W. Zhao, and L. Jia. 2002. First Devonian tetra-
pod from Asia. Nature 420:760–761.