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Carnivoran Evolution: New Views on Phylogeny, Form and Function

Article  in  Systematic Biology · February 2010

DOI: 10.1093/sysbio/syq087

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2011
Syst. Biol. 60(2):241–243, 2011
c The Author(s) 2010. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. All rights reserved.
For Permissions, please email: journals.permissions@oup.com
DOI:10.1093/sysbio/syq087
Advance Access publication on December 24, 2010
Carnivoran Evolution: New Views on Phylogeny,
Form and Function.—Anjali Goswami and Anthony
Friscia, editors. Cambridge, UK: Cambridge Univer-
sity Press, 2010. xiii+492 pp. ISBN 978-0-521-73586-5,
£35, $59 (paperback). ISBN 978-0-521-51529-0, £80, $135
(hardback).
In Carnivoran Evolution, editors Anjali Goswami and
Anthony Friscia bring together more than 20 interna-
tional scientists who present studies focused on fos-
sil and extant mammalian carnivorans. The authors
examine the evolutionary significance of carnivoran
morphology (think flesh-eating felids, clam-eating wal-
ruses, and bamboo-eating pandas) in an ecological con-
text strongly influenced by climate change. The impetus
for the book was a Society of Vertebrate Paleontology
symposium held in Austin, Texas in 2007, and many of
the talks were developed into book chapters. More than
260 species of extant Carnivora are known, and their in-
comparable diversity (i.e., ranging in body size from a
least weasel at 25 gm to the polar bear weighing as much
as 1000 kg) offers an appealing model system. Ques-
tions can be addressed about how such diverse mor-
phology arose and how these traits permitted adapta-
tion to various environmental challenges through time.
To best study patterns and processes of carnivoran evo-
lution, phylogeny-based comparative methods should
be used, as emphasized in this book.
It has been nearly 15 years since publication of the last
comprehensive volumes on the behavior, ecology, and
evolution of mammalian carnivores (Gittleman 1989,
1996). There has been much progress in the field, and
this book incorporates new discoveries as well as new
methods of data analysis. Most importantly, as this
volume well illustrates, carnivoran evolutionary biol-
ogy has become more integrated, benefiting from both
novel and interdisciplinary approaches. Whereas the
Gittleman volumes focused on conservation, this book
reviews the rapidly expanding area of ecomorphology—
emphasizing past and present associations among com-
munities of carnivorans. These associations are based on
BOOK REVIEWS 241
the study of carnivoran feeding and/or locomotor mor-
phology in relation to issues of competition, climate,
and environmental change.
Chapter 1 is a nice introduction to the “players”—
both stem and crown clades. It provides a context for
the book emphasizing that, in addition to their remark-
able diversity, the excellent and near continuous fossil
record of carnivorans during the Cenozoic makes them
an especially interesting group to study. Chapters 2–4
enumerate the many recent advances in carnivoran phy-
logeny that have now yielded a well-resolved and stable
higher level phylogeny based on morphologic, molec-
ular, and combined analyses. That fossils play a crit-
ical role in reconstructing the evolutionary history of
carnivorans is a given, but also emphasized is the of-
ten less-appreciated unique ability of fossils “to break-
up ’long branches’ in phylogenies, . . . preserve
morphologies that can become obscured along these
long branches and also provide a temporal context for
the evolution of living clades” (p. 25). Several contribu-
tions based on more traditional phylogeny and classi-
fication (i.e., Chapters 3 and 4), albeit with new data,
contrast with Chapter 2 by Flynn and colleagues. The
latter provides a broader context for character transfor-
mations revealed through case studies of the evolution
of body size, brain size, and locomotor habitus of early
carnivorans, containing new information from previ-
ously unstudied fossil taxa.
The remainder of the book (more than two-thirds)
builds on a framework rich in comparative methods ap-
plied to various problems in carnivoran ecomorphology
and elucidates broader macroevolutionary patterns.
In Chapter 5, Goswami and Polly use 3D morphomet-
rics in an elegant study of modularity and integration of
the carnivoran cranium together with Monte Carlo sim-
ulations to investigate character correlations. Results
confirm that the basicranium shows the strongest phy-
logenetic signal, a result not unexpected but gratifying,
given that this region has been an area of primary fo-
cus in carnivoran systematics since the time of Flower
(1869). More good news for carnivore tree builders: It
242 SYSTEMATIC BIOLOGY
appears that reliance on dental and basicranial charac-
ters has not significantly misled phylogenetic analyses.
The allometry of fossil and living lions is examined by
Benoit in Chapter 6 using a novel multiphasic regres-
sion analysis of cranial shape and size variables to de-
termine developmental growth phases. Results reveal
misidentification of purported distinguishing features
between the extinct and living lion. For most variables,
the extinct lion shows an allometric shift into the larger
size ranges of the living lion, arguing for population
variation rather than species level separation of these
taxa.
The next 7 chapters (Chapters 7–13) use disparity
to explore carnivoran morphospace. Disparity meth-
ods address lineage diversification in terms of or-
ganismal form (ecomorphology) rather than more
traditional measures of taxonomic richness. Disparity
studies have received recent attention (e.g., Slater et
al. 2010), using methodologies developed by paleon-
tologists and extending them to neontologic studies
of adaptive radiation. In Chapter 7, Holliday evalu-
ates tests for directionality in character evolution or the
case for gain: loss bias by consideration of the effects
of the hypercarnivorous dental specialization on sub-
sequent morphological diversity among carnivorans.
One intriguing explanation for the observed dispar-
ity, which confirms prior results, is that hypercarnivo-
rans (e.g., nimravid feliforms, hyaenids) “ratcheted”
forward into increasingly specialized morphospaces.
This proved to be a strong functional constraint and
likely macroevolutionary pattern, which ultimately
limited their ability to respond to environmental
change.
Chapters 8 and 9 focus on biogeographic patterns—
morphologic disparity explored among carnivoran fam-
ilies and continents in terms of body size and dental
characters. Later (Chapter 10) Morlo and Peign é provide
data suggesting that locomotor pattern is a critically im-
portant additional variable that should be considered
in reconstructing ecomorphologic space. In Chapter 8,
Werdelin and Wesley-Hunt explore morphospace occu-
pation of carnivorans on various continents. They report
high disparity values displayed by euplerids (Malagasy
mongooses), suggesting that this result likely reflects
their being the only carnivorans on Madagascar, thus
occupying niches divided up by several families else-
where. Further work is planned to consider specific en-
vironmental parameters to compare with morphospace
distributions. Chapter 10 presents the most comprehen-
sive investigation to date of carnivoran guild structure
based on body mass, diet, and locomotor patterns sam-
pled over various time slices (Eocene to Recent), which
serves as an excellent model for studies of this kind in
the future.
In Chapter 11, the classic case for competition be-
tween creodonts and carnivores given their temporal
and geographic overlap is investigated based on 2D
craniodental data. More evidence is provided to sup-
port an earlier claim that, in creodonts, their specialized
dentition (i.e., lack of a grinding ability on molars) led
VOL. 60
to their demise, having been outcompeted by carnivo-
rans with more generalized dentitions. Chapter 12, by
Jones and Goswami, is the first rigorous study of shape
variation in the pinniped skull. Results reveal that pho-
cids cover a wider range of morphospace than other
pinniped lineages, perhaps not unexpected given their
greater taxonomic diversity. More interesting, however,
is that both phocids and otariids converge on the wal-
rus morphospace, suggesting morphological constraint
in the rostral region. This may explain why pinniped
ecologic (i.e., mating) specialization is largely limited to
this region (e.g., the large inflated noses of hooded and
elephant seals).
Chapters 13 and 14 (in addition to Chapter 10) de-
part from cranial and tooth shape and diet to exam-
ine morphologic disparity using locomotor characters.
Polly (Chapter 13) investigates limb morphology to
determine whether North American carnivorans are
influenced by various climate and environmental pa-
rameters (i.e., vegetation cover, topography, etc.).
Among various locomotor indices, he found that the cal-
caneal gear ratio performed the best. The high calcaneal
gear ratios of canids and felids reflect digitigrady and
were most often associated with occupants (i.e., mostly
felids) of the desert southwest, whereas the low ra-
tios of ursids and mustelids reflect plantigrady and
were most often observed in the broad-leaf forest re-
gions of the midwest and northeast. This study de-
serves a second step—expansion to other continents.
Lewis and Lague, in Chapter 14, examine machairodon-
tine felid postcrania in light of scaling patterns in extant
felids. Multivariate comparisons reveal that machairo-
dontines are not simply larger versions of extant fe-
lids. Features of the forepaw (e.g., enlarged dew claw)
suggest that they did not use their forelimbs in the
same manner, and their unique femoral morphol-
ogy indicates greater rotary abilities than in extant
felids.
The final chapter (Chapter 15), by Wroe, uses an
engineering technique relatively new to biological ap-
plication (and worthy of more exploration), finite ele-
ment analysis, to reconstruct the cranial mechanics and
predatory behavior of the sabercat, Smilodon fatalis. Re-
sults are consistent with a “bite” activated by head-
depressing musculature. Comparisons are also made
with placental and marsupial wolves and lions. In the
case of the marsupial wolf, Thylacinus cynocephalus, and
dingo, Canis lupus dingo, morphometric bite force and
canine shape analyses suggest both had considerable
niche overlap, increasing the likelihood that competi-
tive exclusion played a role in the demise of the marsu-
pial wolf. Study of the cranial mechanics of marsupial
and placental lions reveals that although the marsupial
lion, Thylacoleo carnifex, was 30% smaller it was able to
generate a bite force 80% that of the placental lion, Pan-
thera leo, suggesting that high bite forces and skull shape
may have allowed the marsupial lion to apply a unique
killing technique.
The book is logically organized and written, and the
editors should be commended for their careful review
2011 BOOK REVIEWS
of contributions, resulting in remarkably little redun-
dancy across chapters. As admonished by several au-
thors, the power of ecomorph comparisons and time se-
ries analyses within the context of a phylogenetic frame-
work might be increased by incorporation of branch
length information or maximum likelihood estimates.
One slight distraction, undoubtedly driven by economic
considerations, is the inclusion of two copies of many
figures, black and white appropriately placed in text and
color versions grouped as an insert in the middle of the
book.
I strongly recommend this book, especially to grad-
uate students. The conceptual breadth of questions,
diversity of approaches, and novel methods of data
analysis aptly reflect the exciting new directions taken
by current researchers. This bookdeserves to be widely
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243
read and serves as an excellent resource for future work
on the paleobiology of carnivorans.
R EFERENCES
Flower W.H. 1869. On the value of the characters of the base of the
cranium in the classification of the order Carnivora, and on the
systematic position of Bassaris and other disputed forms. Proc.
Zool. Soc. Lond. 1869:4–37.
Gittleman J.L., editor. 1989. Carnivore behavior, ecology and evolu-
tion. Volume 1. New York: Cornell University Press.
Gittleman J.L., editor. 1996. Carnivore behavior, ecology and evolu-
tion. Volume 2. New York: Cornell University Press.
Slater G.H., Price S.A., Santini F., Alfaro M.E. 2010. Diversity versus
disparity and the radiation of modern cetaceans. Proc. R. Soc. B.
277:3097–3104.
Annalisa Berta, Department of Biology, San Diego State University, San
Diego, CA 92182, USA; E-mail: aberta@sunstroke.sdsu.edu.