Mammalian Evolutionary Morphology: A Tribute to Frederick S. Szalay Eric J. Sargis and Marian Dagosto (eds.) Dordrect, The Netherlands: Springer, 2008, 439 pp. (hardback), $139.00/€94.95. ISBN-13: 9781402069963.

Reviewed by KRIS KOVAROVIC Department of Anthropology, University College London, 14 Taviton Street, London WC1H 0BW, UNITED KINGDOM; [email protected]

ammalian Evolutionary Morphology is the most recent maries or short concluding sections demystified some of Mpublication in Springer’s Vertebrate Paleobiology the material during my own reading of chapters regard- and Paleoanthropology Series edited by Eric Delson and ing taxa with which I was not very familiar. Regardless of Ross MacPhee. This volume, like others in the series, is an this minor point, all readers will recognise the merit of both extremely professional and well-produced book present- sections for the methodological and analytical issues un- ing up-to-date and cutting-edge research. However, in one der consideration and the thought provoking discussions respect this book stands out from the others; it is a tribute of how skeletal and dental characters can be appropriately to the long and ongoing career of one scientist—Frederick used in the reconstruction of phylogenetic relationships S. Szalay, evolutionary morphologist par excellence. and evolutionary trends. Szalay was once unique amongst To say that Szalay has had a successful career and sig- his peers for pioneering phylogenetic analyses based on nificant influence on many areas of evolutionary morphol- postcranial material—tarsals, in particular—when the ma- ogy is something of an understatement. Eric Sargis and jority of workers believed wholeheartedly that dental char- Marion Dagosto, who edited the book, know firsthand of acters more accurately reflected relatedness. Many of the his influence, having completed their Ph.D.s under his tute- papers in this book reflect his interest in tarsal functional lage. They carefully document Szalay’s impact on the field morphology, as well as his insistence upon the inseparabil- in their own editorial comments and in the organization of ity of phylogeny and function in any analysis and a his- the volume, which includes thoughtful and thought-pro- torical approach to understanding adaptation. These con- voking contributions from many who have been taught or tributions also make it clear that recent fossil discoveries, inspired by him during his more than forty-year career. additional morphological analyses and genetics have often Sargis and Dagosto first provide a sometimes poignant borne out Szalay’s sometimes controversial conclusions re- account of Szalay’s entry into American academics after garding systematics. growing up in his native Hungary and the intellectual path Section I (Non- ) begins with Davis et he followed as a young researcher. They point out that his al.’s detailed dental descriptions of two represent- influence is evidenced by the ten mammalian species that ing the earliest known North American deltatheroidans, have been named after him before detailing the research considered to be a stem group of Metatheria, including a areas in which he has made major contributions, including new species herein named Oklatheridum szalayi in honour of primate evolutionary morphology, mammalian evolution- Szalay’s contributions to our understanding of metatherian ary morphology, and theory and practice of phylogeny re- radiations. Their reassessment of deltatheroidan material construction and adaptive scenarios. A seven page bibliog- indicates a previously unacknowledged level of morpho- raphy of his publications spanning from his first in 1965 to logical diversity during the Early and, as these one article in press at the time of the book’s printing, bears species predate Asian taxa, suggests that a North American further testament to the breadth and depth of his knowl- origin for the group might have been possible. Kear et al. edge. follow with a short paper detailing a functional analysis of The book is divided into two sections. While the au- the hindlimb of extant and fossil macropodoids, investigat- dience of this journal may be most familiar with Szalay’s ing the relationship between limb bone length, locomotor work on , to which the second half is dedicated, habits, and phylogeny. By viewing morphological trends his vast compendium of research also includes major con- over time and comparing analyses that excluded and in- tributions to our understanding of other groups cluded fossil species, they demonstrate that a range of including marsupials and Archonta. Naturally, the two metapodial proportions was established early in kangaroo halves of this book will interest a different subset of evolu- evolution and an increase in metatarsal length is not corre- tionary morphologists. Those interested in the evolution of lated to an increase in aridity during the late Tertiary. The primates may not find the first half of the book as relevant third contribution, from Argot, stands out for providing an or engaging, and vice versa for those involved in mamma- account of the changing interpretations of the giant ground lian evolutionary research of other families. For this reason sloth, Megatherium americanum. Beginning with Cuvier’s my only criticism of the book as a whole is that readers interpretations of the species’ behavior based on its skel- might have found the chapters easier to navigate had they etal anatomy and tracing historical shifts in the portrayal provided abstracts at the outset; certainly those with sum- of this over the years, Argot ties these various in-

PaleoAnthropology 2009: 166−168. © 2009 PaleoAnthropology Society. All rights reserved. ISSN 1545-0031 BOOK REVIEW • 167 terpretations to our limited modern understanding of the primitive primate and euprimate taxa in Asia than previ- species’ behavior. She points out that this famous large- ously acknowledged, she concludes that the scenario is bodied mammal has never been subjected to a systematic possible but certainly not proven. The chapter outlines this functional analysis and, although one might wish that the mounting evidence and presents a cladistic analysis of rel- chapter provided something more along these lines, it is evant taxa, as well as two lengthy but useful appendices nonetheless an interesting story. The fourth chapter, by of data and character definitions. Boyer and Bloch follow Salton and Sargis, outlines a comprehensive functional with a discussion of the mitten-gliding hypothesis for paro- analysis of the tenrecoid forelimb skeleton. Summaries of momyid and micromomyid plesiadapiforms, focussing on the morphologies of the species typifying different locomo- functional interpretations of the postcrania, the phalanges tor patterns are provided and it is one of many chapters in particular. Their analysis, which includes new material, where extremely useful, clear black and white photos are suggests that these taxa do not share the suite of adaptive in abundance. A lengthy fifth chapter from Penkot et al. is mitten-gliding characteristics proposed in an earlier study the first of two focussing on condylarths. It provides de- and are thus more likely to be allied with primates than tailed descriptions of postcranial remains definitively as- with dermopterans, a belief also championed by Szalay. signed to Apheliscus and Haplomylus, two North American The third chapter, by Godinot and Couette, comments on species of apheliscid which were previously best known the morphology and phylogeny of large adapines. They from dental material. Their analysis links them to early identify and compare the cranial material of two genera, macroscelideans, the first non-African taxa which affiliate Leptadapis and their newly named Magnadapis, which com- in this way. The sixth chapter, by Bergqvist, presents a re- prise a total of seven species including Magnadapis fredi, in assessment of condylarth postcrania from the Itaboraí Ba- honour of Szalay. Dagosto et al. provide the fourth contri- sin, Brazil. The morphologies of each element are described bution in a brief and straightforward description of tibiae and assigned to a species already known from the locale from Middle sediments at Shanghuang, China. This (and Bergqvist here supports the somewhat contentious builds on previous work in which analyses of the tarsal position of Victorlemoinea prototypica within Condylarthra). and dental remains suggested the presence of five different A brief functional interpretation of the material infers that small-bodied primate groups. The tibiae, however, appear all of the species were generalised terrestrial mammals, to belong entirely to two haplorhine groups, one with fused although there is some suggestion in the astragalus and tibiafibulae and one displaying the unfused condition, but forelimb long bones that cursorial locomotion may have both bearing functional hallmarks of a leaping primate. been possible. Shockey and Anaya follow with a contribu- Rosenberger et al. follow with an investigation of the “in- tion containing another honorific species announcement. cipient” form of postorbital closure found in the late Eocene Fredszalaya hunteri is named as a new species of carnivo- Rooneyia viejaensis noted by Szalay in the mid-1970’s. This rous marsupial in a chapter that presents a straightforward feature is a major hallmark of Anthropoidea, but research- description of the fauna from Salla, Bolivia, during the late ers argue as to whether or not it links anthropoids and tar- Oligocene when species richness and diversity was particu- siers or if it is a convergent feature of the two groups. This larly high. The authors emphasize anatomical descriptions study is a detailed look at the craniofacial architecture as- of hindlimb morphology, tarsals in particular, and suggest sociated with primate orbital morphology using 3D laser phylogenetic histories for the families represented in the scans. The complexities of placing Rooneyia in the context of fauna. The penultimate chapter of Section I is a very short the major primate clades create difficulties in interpreting piece investigating the use of third proximal phalanx in- the meaning of this feature, but ultimately the authors con- dices in resolving equid systematics in the Oligocene and clude that the genus is more closely related to anthropoids Miocene of Florida. The introduction is an excellent sum- and that the postorbital closure observed in both tarsiers mary of the topic but the relevant text and figures of the and anthropoids arose independently. The sixth contribu- chapter comprise approximately five pages and the intro- tion is Maier’s study of the inner ear anatomy of primates, duction accounts for half of this. Although the results are situated in the broader context of comparative mammalian communicated succinctly—phalangeal data improves phy- anatomy. Histological serial sections allow for an investiga- logenetic analyses compared to analyses based entirely on tion of the chorda tympani nerve; the position of the nerve dental characters—realistically the analysis and discussion above the tensor tympani muscle is shown to be a synapo- require further explanation. The section concludes with an morphy defining Anthropoidea. Sargis et al. then examine engaging and thorough chapter by Polly on the evolution guenon postcrania and locomotor strategies, which range of the pinniped calcaneum and astragalus, addressing is- from arboreal to semi-terrestrial to completely terrestrial. sues of both locomotor function and phylogeny. He clearly Full terrestriality is unique to only three guenon groups, but situates his work within a wider body of research, includ- this analysis suggests that it evolved only once. However, ing Szalay’s interests, assessing the relationship between these three groups of species have diversified substantially tarsal morphology, behavior and relatedness. in terms of their substrate use and postcranial morphology, Section II (Primates) begins with Silcox’s excellent re- inferring that further research in this area may provide a view of evidence in support of the suggestion that the order new avenue for distinguishing between more subtle differ- Primates has an Asian origin. Based on an increasing aware- ences in locomotor behavior in extant and fossil primates. ness of a more diverse fossil record of plesiadapiforms and Harcourt-Smith et al. provide the eighth chapter in this sec- 168 • PaleoAnthropology 2009 tion, wherein the authors report briefly on the initial results Mammalian Evolutionary Morphology is a highly recom- from a 3D morphometric analysis of hominoid tibiae and mended advanced volume that will prove to be a relevant talus joint surface congruence. This will ultimately aid in resource for professionals and postgraduate students in the matching of unarticulated fossil elements from single many evolutionary and palaeontological disciplines. Any- localities and in the identification of fossil genera and spe- one with an interest in mammalian evolution, primate or cies. The book concludes with a lengthy contribution from otherwise, should be familiar with the name Fred Szalay Warshaw in which she discusses the variation in mid-shaft and certainly will recognise that this volume is more than a bone tissue microstructure in extant primates. This novel handy academic textbook. It is a meaningful tribute to the approach can be used to tackle various questions about breadth and depth of Szalay’s long career; the work of his function and phylogeny in living primates and, possibly in students and colleagues herein provide a sincere and grate- the future, similar kinds of questions about fossil taxa. This ful testament to the influence of one person on the develop- detailed report notes a dominance of slow growing tissues ment of a unique branch of science. It is the sort of tribute in all primates, but also identifies variation between taxa as that all academics should aspire towards, but which only a well as between hind and forelimb bones, highlighting the handful of the most worthy will achieve. need for further research in this unique area.