Palaios 2011 Book Review PALAIOS DOI: 10.2110/palo.2011.BR71 Emphasizing the impact of life on Earth’s history

Carnivoran Evolution: New Views on Phylogeny, Form and Function, by Anjali Goswami and Anthony Friscia, editors, 2010, Cambridge University Press, Cambridge, UK, 506 p., ISBN 978-0-521-73586-5, USD59, paperback; ISBN 978-0- 521-51529-0, USD135, hardback.

Carnivoran Evolution brings together an impressive collec- carnivores, the creodonts (hyaenodtonids and oxyaenids). tion of studies addressing the complicated lineage of Carnivora, Chapter 1 concludes with a brief discussion on ecomorphology arguably one of the most popular groups of mammals. This and macroevolutionary patterns, specifically on mammalian lineage is beginning to become clearer thanks to the presented hypercarnivory and its unsustainability, a topic that is addressed works of over 20 international researchers, addressing a variety further in Chapter 7. In Chapter 2, Flynn, Finarelli, and Spauld- of topics from the evolution of carnivoran biogeography to cra- ing provide a very detailed history of carnivoran systematics, nial and postcranial mechanics of extant and fossil taxa. and they discuss recent molecular, morphological, and total The 2006 meeting of the American Society of Mammalogists evidence character analyses. Results of these analyses suggest in Amherst is the true origin of this collection, where the future strong monophyly for nearly all members of carnivoramorpha, editors and authors discussed the fact that there had never been with the exceptions of viverrids (civets, binturongs, and genets) a symposium on the evolution of carnivorans at a Society of and mustelids (weasels), allowing for the diagnosis of a new Vertebrate Paleontology meeting. Such a symposium was also clade, Carnivoraformes, which includes all carnivoramorphans in need, as the previous comprehensive works on carnivoran except early Cenozoic viverravids and miacids. The inclusion evolution, behavior, and ecology (Gittleman 1989, 1996) were of abundant fossil taxa in this study is discussed by Flynn et al., already more than a decade old and did not include updates on noting how crucial fossils are to interpreting evolutionary and new techniques and phylogenies, nor the attention to ecomor- temporal transformations, regardless of the lack of molecular phology included in this text. The following Society of Vertebrate data, which is a welcomed support for total evidence analyses. Paleontology meeting held in Austin, Texas, in 2007, had such a Chapter 3, by Veron, examines the phylogeny of viverrids symposium, where many of the presentations were later adapted (not to be confused with viverravids) and viverrid-like feli- to the chapters included in Carnivoran Evolution. formes, the latter of which includes taxa such as the African The first four chapters deal almost exclusively with car- Palm Civet and the Malagasy carnivores. Here, Veron reevalu- nivoran systematics, reviewing known carnivoran phylogeny and ates the clade of Viverridae, and based on molecular analyses, addressing some problematic issues, such as the phylogenetic Veron excludes African Palm Civets, Malagasy viverrids, and placement of Viverridae (Chapter 3) and Ailuridae (Chapter 4). Asian linsangs from Viverridae. This leads to a new descrip- In Chapter 1, Goswami supplies an excellent introduction to tion of Viverridae, which consists of four subfamilies: Asian the order Carnivora, and outlines their relationship to other pla- (Hemigalinae, Paradonxurinae), African (Genettinae), and centals, going back to the early Cenozoic (Paleocene–Eocene). Afro-Asian (Viverrinae). These results also highlight challenges Included here is a discussion on how advances in molecular of traditional morphological features used to classify feliform phylogenetics have significantly changed our understanding of carnivores, such as apparent convergent or primitive characters carnivoran evolution, as it has for biology and paleontology in of dental and basicranial features. Similarly, Chapter 4 (Morlo general. As Goswami walks us through the two major groups and Peigne) reviews the family Ailuridae and investigates its of carnivorans, Feliformia (e.g., felids, viverrids, hyaenids), and phylogenetic placement, which has previously been problematic. Caniformia (e.g., canids, mustelids, ursids), we see not only Ailuridae is currently represented by only one living genus, simple descriptions of extant groups, but also an overview of Ailurus fulgens (Red Panda). While currently exclusive to Asia, how such fossil taxa as nimravidae (such false saber-toothed ailurids have strong fossil roots in Europe, where they held their cats as Hoplophoneus) and amphicyonidae (such bear-dogs as highest diversity in the Oligocene through Miocene. Morlo and Amphicyon) fit into the lineage, along with the noncarnivoran Peigne conduct phylogenetic analyses, including both molecular

Copyright © 2011, SEPM (Society for Sedimentary Geology) PALAIOS BOok review 2 and morphological data of extant and fossil taxa, yielding results there is little or no overlap in morphospace occupation when that suggest a placement for Ailuridae within Musteloidea in members of either clade have interaction potential; this sug- Arctoidea in Caniformia. The chapter concludes with an histori- gests that the ecological roles of taxa in the two families differs cal overview of Ailurus studies and comparisons to giant panda, between Africa and Asia. Despite this result, understanding the and a highly detailed review of ailurid evolution. processes underlying the pattern is still problematic. The remainder of the book takes a considerable shift in at- In Chapter 10, Morlo, Gunnell, and Nagel provide an tention from phylogenetic analyses to comparative methods and extremely comprehensive ecomorphological analysis of Lau- novel techniques toward an understanding of carnivoran eco- rentian carnivoran guilds from the Eocene through Miocene morphology. Chapter 5 addresses character dependence, a major to investigate the usefulness of carnivoran guild analyses for assumption in many morphology-based phylogenetic analyses, paleoclimate and paleoenvironment estimates. Comparisons of although most studies of modularity and morphologic integra- carnivore guild structures are made based on parameters such tion find significant correlations among phenotypic traits. Here, as body mass, diet class, and locomotor class, and they yield authors Goswami and Polly call the validity of this assumption suggestions on the role climate, environment, and time depth into question, and they use empirically derived data in cranial across faunal turnover have on guild structures throughout the modularity and morphological integration in the carnivoran skull Cenozoic. Furthermore, throughout the Cenozoic, guilds differ to assess the impact of trait correlations on phylogeneic analyses significantly in terms of taxonomic composition and diversity, of Carnivora. By testing whether individual modules differ in although structurally they remain quite similar. their relationships between shape and phylogenetic relatedness Chapter 11, by Friscia and Valkenburgh, takes a slight shift or whether correlated characters significantly misled previous in direction and investigates evidence for direct competition phylogenetic analyses of Carnivora based on morphology, this between carnivorans and creodonts in the Eocene. This study chapter takes a very novel approach to the problem of potential investigates craniodental features, diet, and body size diversity biases in analyses. The utilization of 3D morphometrics and in Eocene carnivores over time to explore the potential for Monte Carlo simulations to investigate the correlations of charac- ecological overlap, and hence competition between creodonts ters also has potential for other avenues in vertebrate paleontol- and carnivorans. The case for competition is strengthened by ogy, where it has not been broadly used or investigated to date. this study, also suggesting that competition may have been In Chapter 6, Benoit discusses the challenges of assigning influential in the extinction of the creodonts, due to the lack of fossil specimens to extant taxa. Notable size differences of the specialization in carnivoran dental plasticity. modern and American lion (Pantera leo and Pantera leo atrox, Chapter 12, by Jones and Goswami, utilizes 3D morphomet- respectively) have led to several misidentifications of distin- ric data to quantitatively assess pinneped cranial morphology guishing features, because the differences between the taxa were to address whether differences in cranial shape correlate with not considered allometrically. This study examines features that phylogenetic relationships among pinnepeds or their ecologi- revealed some of these characteristics are not allometrically cal attributes. The authors also investigate how differences in distinct in the two taxa, while some are. While the results do not cranial ontogeny reflect different reproductive strategies among disprove the hypothesis that American Lion and modern lions pinnepeds. The results show a surprising pattern in the repeated are separate species, the argument is weakened by these results. evolution of mating and feeding specializations toward walrus- Chapters 7 through 13 focus more on carnivoran morpho- like morphology. space and disparity, the latter of which is used here to address Chapters 13 and 14 both discuss postcranial analyses of ecomorphology (evolutionary diversification of form). In Chap- carnivorans. In Chapter 13, Polly addresses faunal ecomorphol- ter 7, Holliday analyzes directionality of the character evolution ogy and living carnivoran locomotor systems in the context of of hypercarnivorous specializations in carnivoran dentition, modern climate, and relying on the calcaneal gear ratio, in which suggesting that the repeated and unsustainable reemergence of high ratios are associated with digigrady in canids and felids of hypercarnivory in Mammalia may be influenced by increasingly arid regions, and low ratios associated with plantigrady among specialized morphospaces, leading to decreases in the long-term ursids and mustelids of temperate forests. Chapter 14 examines success of hypercarnivores, such as nimravids or hyaenids. the postcrania of machairodontine felids, specifically addressing Chapters 8 and 9 discuss ecomorphology and biogeography of scaling patterns in extant felids. The results of this study sug- Carnivora, and focus specifically on herpestids and viverrids gest that contrary to popular belief, saber-toothed cats are not (mongooses and civets). Chapter 8 investigates ecomorpho- scaled-up versions of extant felids. Furthermore, multivariate logical disparity and distribution among extant members of comparisons performed in this study imply different forelimb Carnivora. Authors Werdenlin and Wesley-Hunt report that and hindlimb locomotor function than extant felids. the different families of Carnivora significantly differ in mean In the last chapter, Wroe investigates carnivoran cranial ecomorphological disparity, the lowest being hypercarnivorans mechanics with a novel approach. Chapter 15 utilizes finite ele- (e.g., felidae), compared to high disparity in hyaenids and ursids, ment modeling analysis to reconstruct cranial mechanics and despite no detectable difference in mean disparity of continental predatory behavior in Smilodon fatalis. Finite element modeling differences. Chapter 9 expands upon this analytical method and (FEM) is a relatively new technique, but has previously been focuses on herpestids and viverrids, small tropical-subtropical applied to (and embraced by) vertebrate paleontology (Snively feliformes from Africa and Southeast Asia. Compared to the and Cox, 2008; Snively and Theodor, 2011). The results of results in Chapter 8, Wesley-Hunt, Dehghani, and Werdenlin this comparison concur with previous studies, which suggest a report that even though potential for morphospace overlap exists, head-depressing bite. An interesting comparison is also made PALAIOS BOok review 3 between morphometric bite force and canine shape analysis of at the very least, an excellent reference for not only mammalian hypercarnivorous marsupials and placentals, the marsupial wolf researchers, but vertebrate paleontologists and zoologists in (Thylacinus cynocephalus) and the dingo (Canis lupus dingo) general. respectively. The results of this analysis suggest a significant REFERENCES niche overlap, implying considerable competition as a contribut- Gittleman, J.L., editor, 1989, Carnivore behavior, ecology and ing factor to the ultimate extinction of the Thylacine. evolution, vol. 1: Cornell University Press, New York, 624 p. This volume is very well organized and written. The di- Gittleman, J.L., editor, 1996, Carnivore behavior, ecology and versity of the topics covered by the contributors relieved the evolution, vol. 2: Cornell University Press,New York, 664 p. potential for significant overlap in chapters and content, which Snively, E., and Cox, A., 2008, Structural mechanics of is somewhat common with compiled volumes such as this, pachycephalosaur crania permitted head-butting behavior. especially when they arise out of conference symposia. Many Palaeontologica Electronica, v. 11, no. 1; 3A, 17 p., http:// of the novel and unique approaches to carnivoran evolutionary palaeo-electronica.org/2008_1/140/index.html study, such as FEM, character modules, or integration analysis, Snively, E., and Theodor, J.M., 2011, Common functional cor- have not been as heavily utilized in other fields of vertebrate pa- relates of head strike behavior in the pachycephalosaur leontology, although the potential for such study certainly exists. Stegoceras validum (Ornithischia, Dinosauria) and combat- The figures and tables produced for this book are ive artiodactyls: PLoS ONE, v. 6, no. 6, p. e21422. nicely placed, although the color appendix in middle of book is Joseph E. Peterson somewhat distracting. I highly recommend this text; the organi- Department of Geology zation and content make it an ideal piece for any paleontology University of Wisconsin, Oshkosh or biology professional or graduate student’s library, though it 800 Algoma Blvd. is not off-putting to upper-level undergraduates who may pursue Oshkosh, Wisconsin 54901, USA mammalian paleontology or biology. Carnivoran Evolution is, [email protected]