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Home , Ameridelphia, Amphipithecus, Anaptomorphinae, Apterodon, Chriacus, Deltatheridium

Mammalian Evolutionary Morphology Paleobiology and Paleoanthropology Series

Edited by

Eric Delson Vertebrate Paleontology, American Museum of Natural History, New York, NY 10024, USA [email protected]

Ross D. E. MacPhee Vertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA [email protected]

Focal topics for volumes in the series will include systematic paleontology of all (from agnathans to ), phylogeny reconstruction, functional morphology, paleolithic archaeology, taphonomy, geochronology, historical biogeography, and biostratigraphy. Other fi elds (e.g., paleoclimatology, paleoecology, ancient DNA, total organismal community structure) may be considered if the volume theme emphasizes paleobiology (or archaeology). Fields such as modeling of physical processes, genetic methodology, nonvertebrates or neontology are out of our scope.

Volumes in the series may either be monographic treatments (including unpublished but fully revised dissertations) or edited collections, especially those focusing on problem-oriented issues, with multidisciplinary coverage where possible.

Editorial Advisory Board Nicholas Conard (University of Tübingen), John G. Fleagle (Stony Brook University), Jean-Jacques Hublin (Max Planck Institute for Evolutionary Anthropology), Peter Makovicky (The Field Museum), Sally McBrearty (University of Connecticut), Jin Meng (American Museum of Natural, History), Tom Plummer (Queens College/CUNY), Ken Rose (Johns Hopkins University), Eric J. Sargis (Yale University).

Published titles in this series are listed at the end of this volume Mammalian Evolutionary Morphology

A Tribute to Frederick S. Szalay

Edited by

Eric J. Sargis Yale University, Department of Anthropology Peabody Museum of Natural History, Division of Vertebrate Zoology New Haven, CT, USA

and

Marian Dagosto Feinberg School of Medicine Northwestern University, Department of Cell and Molecular Biology, Chicago, IL, USA Library of Congress Control Number: 2008922957

ISBN 978-1-4020-6996-3 (HB) ISBN 978-1-4020-6997-0 (eBook)

Published by Springer, P.O. Box 17, 3300 AA Dordrecht, The Netherlands.

www.springer.com

Cover illustration: Dryomomys szalayi, drawing by Doug M. Boyer.

Printed on acid-free paper

All Rights Reserved © 2008 Springer Science + Business Media B.V. No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfi lming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifi cally for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Justine A. Salton

12 March 1972–28 October 2005 This book is dedicated to the memory of Justine Salton, Fred Szalay’s last student and a friend to many of those who contributed to this volume.

Figure 1. Justine A. Salton (left) with Frederick S. Szalay (right) in New York City after her dissertation defense on August 15, 2005. Preface

Frederick S. Szalay is a commanding figure – one of those Frederick Sigmund Szalay was born in Hungary on peerless inimitable people that leave a lasting impression November 15, 1938. In many ways he was the product of however briefly they are encountered. Passionate and fear- the war-torn years of World War II where as a child he spent less, he approaches his work, as he does everything else in months forced to live in the cellars of Budapest while bombs his life, with great gusto and verve and expects everyone were falling. Towards the end of the war this was followed by around him to do the same. To have worked with him was street combat between the German and Soviet forces, which alternately a terror and a blessing, but was in any case truly he witnessed firsthand when he and other small rascals man- inspirational. Students and colleagues alike were apprehen- aged to sneak upstairs from the cellar. As a 6-year-old at the sive of his much renowned (but in reality rarely displayed and end of 1944, he helped his uncle and some friends coax an usually deserved) critiques, but therefore all the more appre- unexploded 500 lb bomb down the stairs from the third floor ciative of his generously given honest praise and unwavering of the house where they lived. Having to scavenge for food confidence and support. His unbelievable breadth and depth with his beloved grandfather during the winter of 1945–1946 of knowledge of all things mammalian and paleontological stands out as something he will never forget. is due in no small part to his absurdly dense and complete In addition to being a voracious reader of travel and natural library, compiled with the same ravenous collector’s eye that history (and also a student of French and English), most of he applies to souvenirs from foreign locales, abbreviation his high school years were spent playing a variety of sports systems for tooth structures and joint surfaces, and dissect- (swimming, track and field, boxing, and rowing) and shoot- ible road kill carcasses. Those brave readers prepared to work ing photographs, with very little academic effort (but much through the long philosophical preambles and the “very, mischief, and detailed planning with his friends on how to very long sentences and creative grammar constructs” (in the leave the Iron Curtain behind). Having the family background words of one admirer) that distinguish Fred’s insightful work of a Jewish mother and a father from the titled nobility who from more mundane contributions are sure to learn something was a feudal judge in pre-War Hungary nullified any chance valuable from one of the most astute and creative practitioners of his attending university under the communists. Following of evolutionary morphology. the defeat of the 1956 uprising in Hungary, and after a previ- Equally at home with dental, cranial, or postcranial mor- ous attempt at escape which ended in capture, he and a good phology, Fred made major contributions to the literature on friend managed to reach Austria in late November 1956. He mammalian evolutionary morphology, particularly in , never finished his last year in Gymnasium (high school). Archonta, and , as will be detailed below. The esteem Oddly, Fred had no acquaintance with either vertebrate in which he is held by his colleagues can be partly measured by paleontology or while attending col- the number of taxa named after him in honor of his contributions lege in the US. After reaching the United States in December to our knowledge of mammalian evolution. These include the 1956, he worked for nine months at odd jobs. Then a small primates Jemezius szalayi (Beard, 1987), Szalatavus attricuspis Catholic college in Maryland offered him a refugee scholar- (Rosenberger et al., 1991), Tatmanius szalayi (Bown and Rose, ship. Mt. St. Mary’s College offered a straightforward pre- 1991), Dryomomys szalayi (Bloch et al., 2007), and Magnadapis med curriculum with no opportunity for the study of geology. fredi (Godinot, this volume); the Szalinia gracilis His consummate interests in mammalian natural history had (de Muizon and Cifelli, 2001), szalayi (Luo et al., to be satisfied with a biology major and chemistry minor, but 2003), Oklatheridium szalayi (Davis et al., this volume), and his last two summers were spent in near bliss working at the Fredszalaya hunteri (Shockey and Anaya, this volume); and the Catskill Game Farm in New York State. The great variety of multituberculate Ectypodus szalayi (Sloan, 1981). that he worked with while living on the premises

vii viii Preface set the stage for his plans for a Ph.D. at the Evolutionary Morphology University of Massachusetts at Amherst, taking with him an NDEA (National Defense Education Act) Fellowship that he 1. Primate Origins. Fred’s dissertation work on the insecti- won following graduation from college (and naturalization in vore-primate transition set the groundwork for a career-long 1961). While at Amherst Fred took Albert Wood’s year long interest in the subject of primate origins both from a phyloge- course in Vertebrate Paleontology and a seminar with Lincoln netic and ecological/adaptive perspective. His first monograph Brower on Evolution and Ecology at Amherst College. Added on the subject (Szalay, 1969, #10 in Szalay bibliography), the to these pivotal experiences was the reading of George G. publication of his dissertation, concentrated on the dental Simpson’s Meaning of Evolution (followed by Simpson’s evidence for the phylogenetic relationships of the still frus- other books on evolution and systematics), and as he often tratingly difficult to interpret mixodectids and microsyopids. told his students, Simpson’s writings were perhaps the most From this work he developed the hypothesis that the mor- important reasons for shifting his interests from mammalogy phological changes in the dentition that distinguished the to paleontology. A combination of these truly inspirational first primates (plesiadapiforms) from their predecessors was experiences at Amherst led to a quick trip to the American the result of a shift from a primarily insectivorous diet to a Museum of Natural History (AMNH) to interview with more herbivorous one (Szalay, 1968, #6). This work was fol- Malcolm McKenna, followed by a transfer to the Biology lowed by several papers that explored the dental, cranial, and Department at Columbia University, and the AMNH. postcranial evidence linking to Euprimates In addition to McKenna’s much valued tutelage (as Fred and which developed a coherent explanation of the adaptive often stated), and the no less influential atmosphere of the significance of primate synapomorphies. For example, Szalay Vertebrate Paleontology traditions at the AMNH, were the et al. (1975, #52) used postcranial evidence to infer that ple- much treasured associations with fellow graduate students in siadapiforms were arboreal and closely related to euprimates. Biology and Geology at Columbia; professors such as Bobb Although both of these proposals were initially challenged, Schaeffer, Edwin Colbert, and John Imbrie; postdocs like they have subsequently been supported with evidence from Leigh Van Valen and Len Radinsky; and the hosts of perennial new fossils (Bloch and Boyer, 2002; Bloch et al., 2007) and visitors that stream through the AMNH regularly. Yearly field new phylogenetic analyses (Silcox, 2001; Bloch and Boyer, work and field courses in geology rounded out the bases for 2002; Bloch et al., 2007). Primate origins and the evolution- his long continued dedication to understanding mammalian ary morphology of plesiadapiforms are topics addressed in evolutionary history and macroevolutionary dynamics. After this volume by Silcox and Boyer and Bloch. completing his doctorate in 1967, Fred stayed on as an NSF Postdoctoral fellow until taking a job in the Department of 2. Phylogenetic relationships within Primates. Fred Szalay Anthropology at Hunter College, CUNY. He was a Research also worked on the delineation of major taxa within Primates. Associate in the Department of Vertebrate Paleontology of Basicranial evidence was marshaled to understand the rela- the AMNH until 1985, and on the Graduate Faculty of the tionships within Strepsirhini (Szalay and Katz, 1973, #42) City University of New York. He retired from Hunter College and to support the validity of (Szalay, 1975, #58). in 2003, and is now an Adjunct Professor in the Department The latter paper, along with many to follow, argued that the of Biology, University of New Mexico. He is also Professor fundamental division within Primates was Strepsirhini (adap- Emeritus in the Ecology and Evolutionary Biology Doctoral ids + lemuriforms) and Haplorhini (Tarsius + omomyids + Program, City University of New York. anthropoids). These hypotheses, which are the best supported It is an honor and a privilege for all of us to have known today, were defended by Fred against the rival hypotheses him, to have learned from him, and to be able to contribute of “Plesitarsiiformes” (plesiadapiforms + tarsiiforms) and to this volume celebrating his career. This book acknowl- “Simiolemuriformes” (strepsirhines + anthropoids) favored edges and celebrates the contributions of Dr. Frederick S. by other paleontologists. Fred also addressed the origin and Szalay to the field of Mammalian Evolutionary Morphology. phylogenetic relationships of anthropoid primates (Szalay, Fred Szalay has published about 200 articles, 6 mono- 1975, #55; Rosenberger and Szalay, 1980, #75), a theme visited graphs, and 6 books on this subject. His dissertation work in this volume by Rosenberger et al. and Maier. was awarded the Newberry Prize in Vertebrate Zoology. He 3. Major publications. Fred has also described and named has received numerous grants from the National Science numerous and primate taxa, a subject Foundation and the Wenner-Gren Foundation, and was represented here by Godinot and Couette. Fred is the author awarded a Guggenheim Fellowship in 1980. Throughout or editor of several important books and monographs on the his career, Fred has been a strong advocate for biologically subject of primate evolution. These are: and evolutionarily meaningful character analysis. In his 1969: Mixodectidae, Microsyopidae, and the insecti- view, this can be accomplished only through an integrated vore-primate transition. Bulletin of the American Museum strategy of functional, adaptational, and historical analysis. of Natural History 140, 193–330. Using this approach, he has made major contributions to 1975: Approaches to Primate Paleobiology. Contributions the following areas of study: to Primatology, Volume 5. Karger AG, Basel. Preface ix

1975: Phylogeny of the Primates: A Multi-disciplinary logeny, functional morphology, and biogeography in both Approach. Plenum, New York (Luckett, W. P. and F. S. a book (Szalay, 1994, #142) and a monograph (Szalay and Szalay, Eds.). Sargis, 2001, #198). This group is considered by Davis et al. 1976: Systematics of the (, and Kear et al. in this volume. Primates): , Phylogeny, and Adaptations. 3. Other Mammals. In 1990, Szalay, with co-editors M. J. Bulletin of the American Museum of Natural History 156, Novacek and M. C. McKenna, organized an important confer- 157–450. ence on the subject of mammalian phylogeny and evolution, 1979: Evolutionary History of the Primates. Academic, which resulted in the publication of two volumes (Szalay New York (Szalay, F. S. and E. Delson, Eds.). et al., 1993, #130–131). The themes of mammalian systemat- Szalay and Delson (1979, #72) is perhaps the most remarka- ics and paleontology play into several contributions in this ble of all these volumes, as it was a huge undertaking that has volume: e.g., Davis et al., Penkrot et al., Bergqvist, Shockey never been replicated despite enormous interest in primates and Anaya, and O’Sullivan. Fred also published a monograph, and a proliferation of primatologists since the late 1970s. with F. Schrenk in 1998 on “edentates” (#148). This study included an analysis of xenarthrans, a group discussed in this Mammalian Evolutionary Morphology volume by Argot.

1. Archonta. The morphological evidence supporting the supraordinal grouping Archonta, and its adaptive signifi- Theory and Practice of Phylogeny cance. Szalay (1977, #66) provided the first morphological Reconstruction/Adaptive Scenarios support for McKenna’s (1975) revised concept of Gregory’s (1910) Archonta, a that includes Primates, Scandentia, 1. The integration of postcranial evidence into hypotheses Dermoptera, and Chiroptera. Szalay (1977, #66) used tarsal of mammalian systematics. Szalay’s (1977, #66) phylogeny evidence to unite Primates, Scandentia, and Dermoptera, as and classification of mammals were based completely on tarsal well as previously cited similarities to include Chiroptera evidence, which was both novel and controversial at the time as well. Although Archonta (including Chiroptera) has because such studies were typically based on teeth. Most were not been subsequently supported, (excluding critical of this study, but George Gaylord Simpson (1978), Chiroptera) has been strongly supported in molecular stud- probably the best known mammalian systematist in the history ies (e.g., Murphy et al., 2001). In other words, the grouping of the field, was supportive of Szalay’s innovative analysis. In of Primates, Scandentia, and Dermoptera that he originally fact, Fred’s analyses can fairly be seen as building on and refin- recognized based on tarsal evidence has now been supported ing the traditions of “total evidence” practiced by the best of in numerous other studies using different datasets. Szalay and the previous generation of mammalian paleontologists includ- Drawhorn (1980, #73) proposed that Archonta originated and ing Simpson, William K. Gregory, William D. Matthew, and diversified in an arboreal milieu, another hypothesis that has Henry F. Osborne. The hegemony of dental evidence was based been supported in subsequent studies (e.g., Bloch and Boyer, on the assumption that teeth reflected relationships better than 2002; Bloch et al., 2007). Szalay continued to work on this the limb skeleton, which was thought to be more influenced group throughout his career, including the publication of a by functional demands and thus more prone to parallelism. As monograph with S. G. Lucas in 1996 (#145). anticipated by and demonstrated by Szalay, this assumption is faulty at best. Fred was able to use postcranial evidence to 2. Marsupialia. After spending a sabbatical year in Australia support controversial hypotheses on Primates (sensu lato; i.e., in 1980, Szalay (1982, #80) proposed a completely novel including plesiadapiforms), Euarchonta, and Marsupialia (specifi- hypothesis of relationships based on tarsal evi- cally ), as well as many other mammalian groups dence. He hypothesized that the South American Dromiciops such as Glires, , and Mesozoic taxa. The majority of is more closely related to Australasian marsupials than the contributions in this volume build on this aspect of Szalay’s to other South American marsupials. He formalized this work, including those by Kear et al., Argot, Salton and Sargis, by including Dromiciops with Australasian marsupials in Penkrot et al., Bergqvist, Shockey and Anaya, O’Sullivan, Polly, Australidelphia, whereas other South American marsupi- Boyer and Bloch, Dagosto et al., Sargis et al., Harcourt-Smith als were placed in . The classification of et al., and Warshaw. Dromiciops with Australasian taxa in Australidelphia was initially met with strong resistance and was highly criticized, 2. Phylogenetic and adaptational analysis. In the 1970s the but it has subsequently been supported in both morphological trend toward both numerical phenetic and cladistic methods (e.g., Horovitz and Sanchez-Villagra, 2003) and molecular of phylogenetic analysis was rapidly expanding. Fred Szalay (e.g., Amrine-Madsen et al., 2003) analyses. Szalay’s was and is a vocal critic of the superficial character counting, (1982, #80) novel hypothesis of marsupial relationships has distribution-based, algorithm driven solutions to phylogeny major implications for the biogeographic history of this reconstruction advocated by some, particularly cladists. He group. Szalay further developed his ideas on marsupial phy- advocates instead for the primacy of biologically informed x Preface character analysis using functional, developmental, and adap- analyzing adaptation. His 1981 (#78) paper outlined a his- tational criteria to both weight characters and test hypotheses torically informed approach for analyzing adaptations of fossil of homology and polarity (Szalay, 1981, #76). Fred was also organisms. The influence of this point of view is clear in many heavily influenced by the work of Walter Bock, and is a strong of the contributions to this volume. proponent of the logical inseparability of functional-adap- tive and phylogenetic analysis; one is not primary to another, ERIC J. SARGIS they are reciprocally illuminatory (if we might borrow that Department of Anthropology Hennigian phrase) (Szalay, 1981, #78; Szalay and Bock, 1991, Yale University #127; Szalay, 2000, #160). In fact, “The meeting of these two New Haven, CT 06520, USA ‘separate’ disciplines is of course what is usually referred to as [email protected] morphology” (Szalay, 1981, #78, p. 160). This point of view is Division of Vertebrate Zoology represented in his concept of the “transformation series”, a test- Peabody Museum of Natural History able hypothesis of polarity based not on distribution, but on the MARIAN DAGOSTO fossil record and a functionally logical sequence of ancestor- Department of Cell and Molecular Biology descendant states. In Fred’s view, the a-historical approaches, Feinberg School of Medicine those that are phenetic, correlation based, and do not consider Northwestern University the phylogenetic history of the subject organism and its influ- Chicago, IL 60611, USA ence on the likely response to selection, are not adequate for [email protected] Bibliography of Frederick S. Szalay

1965 1970 1. First evidence of tooth replacement in the subclass Allotheria 14. Amphipithecus and the origin of catarrhine primates. Nature (Mammalia). American Museum Novitates 226, 1–12. 227(5256), 355–357.

1966 1971 2. The tarsus of the Paleocene leptictidProdiacodon(, 15. On the cranium of the late Paleocene primate Mammalia). American Museum Novitates 2267, 1–13. tricuspidens. Nature 230(5292), 324–325. 3. (Szalay, F. S. and S. J. Gould) Asiatic Mesonychidae 16. Artiodactyla. Encyclopedia of Science and Technology. (Mammalia, Condylarthra). Bulletin of the American Museum McGraw-Hill, New York, pp. 616–617. of Natural History 132(2), 127–174. 17. Astrapotheria. Encyclopedia of Science and Technology. McGraw-Hill, New York, p. 649. 1967 18. Hyracoidea. Encyclopedia of Science and Technology. McGraw-Hill, 4. The Affinities of Apterodon (Mammalia, Deltatheridia, New York, p. 675–676. ). American Museum Novitates 2293, 1–17. 19. Mammalia. Encyclopedia of Science and Technology. McGraw- 5. (Van Valen, L., P. M. Butler, M. C. McKenna, F. S. Szalay, Hill, New York, pp. 94–96. B. Patterson and A. S. Romer) Galeopithecus Pallas, 1973 20. Monotremata. Encyclopedia of Science and Technology. (Mammalia): proposed validation under the plenary powers McGraw-Hill, New York, pp. 689–690. Z.N. (SP 1792). Bulletin of Zoological Nomenclature 24(3), 21. Perissodactyla. Encyclopedia of Science and Technology. 190–191. McGraw-Hill, New York, pp. 23–25. 22. . Encyclopedia of Science and Technology. 1968 McGraw-Hill, New York, pp. 731–732. 6. The beginnings of primates. Evolution 22(1), 19–36. 23. . Encyclopedia of Science and Technology. McGraw-Hill, 7. The Picrodontidae, a family of early primates. American New York, pp. 396–397. Museum Novitates 2329, 1–55. 24. Tooth (vertebrate). Encyclopedia of Science and Technology. 8. (Mellett, J. S., and F. S. Szalay) Kennatherium shirensis McGraw-Hill, New York, pp. 699–701. (Mammalia, Palaeoryctoidea) a new didymoconid from the 25. (Davis, D. D. and F. S. Szalay) Allotheria. Encyclopedia Eocene of Asia. American Museum Novitates 2342, 1–7. of Science and Technology. McGraw-Hill, New York, pp. 9. Origins of the (Mammalia, Insectivora). 295–296. American Museum Novitates 2352, 1–11. 26. (Davis, D. D. and F. S. Szalay) . Encyclopedia of Science and Technology. McGraw-Hill, New York, p. 136. 1969 27. (Davis, D. D. and F. S. Szalay) Metatheria. Encyclopedia of 10. Mixodectidae, Microsyopidae, and the insectivore-primate Science and Technology. McGraw-Hill, New York, p. 360. transition. Bulletin of the American Museum of Natural 28. (Davis, D. D. and F. S. Szalay) . Encyclopedia of Science History 140(4), 193–330. and Technology. McGraw-Hill, New York, p. 567. 11. The Hapalodectinae and a phylogeny of the Mesonychidae 29. (McKenna, M. C., J. S. Mellett and F. S. Szalay) Affinities of the (Mammalia, Condylarthra). American Museum Novitates . Journal of Paleontology 2361, 1–26. 145(3), 441–442. 12. Uintasoricinae, a new subfamily of early Tertiary mammals 30. (Szalay, F. S. and M. C. McKenna) Beginnings of the Age (Primates). American Museum Novitates 2363, 1–36. of Mammals in Asia: the late Paleocene Gashato fauna, 13. Origin and evolution of function of the feeding Mongolia. Bulletin of the American Museum of Natural History mechanism. Evolution 23(4), 703–720. 144(4), 269–318.

xi xii Bibliography of Frederick S. Szalay

31. The adapid primates Agerina and Pronycticebus. American 1975 Museum Novitates 2466, 1–19. 51. (Szalay, F. S., Ed.) Approaches to Primate Paleobiology. 32. Significance of the basicranium of early Tertiary primates Contributions to Primatology, Volume 5. Karger AG, Basel. for the phylogeny of the . (Abs.) American Journal of 52. (Szalay, F. S., I. Tattersall and R. L. Decker) Phylogenetic Physical Anthropology 35(2), 297. relationships of Plesiadapis – postcranial evidence. In: 33. Biological level of organization of the Chesowanja robust F. S. Szalay (Ed.). Approaches to Primate Paleobiology. australopithecine. Nature 234, 229–230. Contributions to Primatology, 5, 136–166. 34. Relationships of the alleged primate Gesneropithex. Journal 53. Early primates as a source for the taxon Dermoptera. (Abs.); of Mammalogy 52(4), 824–826. American Journal of Physical Anthropology 42(2), 332–333. 54. Where to draw the nonprimate-primate taxonomic bound- 1972 ary. Folia Primatologica 23, 158–163. 35. Cranial morphology of the early Tertiary Phenacolemur 55. Haplorhine relationships and the status of the Anthropoidea. and its bearing on primate phylogeny. American Journal of In: R. H. Tuttle (Ed.). Primate Functional Morphology and Physical Anthropology 36(1), 56–76. Evolution. Mouton Publishers, The Hague, The Netherlands, 36. Paleobiology of the earliest primates. In: R. Tuttle (Ed.). pp.3–22. The Functional and Evolutionary Biology of Primates. Aldine- 56. Hunting-scavenging protohominids: a model for hominid Atherton, Chicago, pp.3–35. origins. Man 10, 420–429. 37. Amphipithecus revisited. Nature 236(5343), 179. 57. (Luckett, W. P. and F. S. Szalay, eds.) Phylogeny of the Primates: 38. Review of “Vertebrate Paleozoology” by E. Olson, American a multi-disciplinary approach. Plenum, New York. Journal of Physical Anthropology 36(3):449–450. 58. Phylogeny of primate higher taxa: the basicranial evidence; 39. (Wilson, R. W. and F. S. Szalay) New paromomyid primate In: W.P. Luckett and F.S. Szalay (Eds.). Phylogeny of the from middle Paleocene beds, Kutz Canyon area, San Juan Primates: A Multidisciplinary Approach. Plenum, New York, Basin, New Mexico. American Museum Novitates 2499, 1–18. pp. 357–404. 59. Phylogeny, adaptations, and dispersal of the tarsiiform 1973 primates; In: W.P. Luckett and F.S. Szalay (Eds.). Phylogeny 40. (Szalay, F. S. and A. Berzi) Cranial anatomy of Oreopithecus. of the Primates: A Multidisciplinary Approach. Plenum, New Science 180,183–185. York, pp. 91–125. 41. New Paleocene primates and a diagnosis of the new suborder Paromomyiformes. Folia Primatologica 19(2–3), 73–87. 1976 42. (Szalay, F. S. and C. C. Katz) Phylogeny of lemurs, galagos, 60. Systematics of the Omomyidae (Tarsiiformes, Primates): tax- and lorises. Folia Primatologica 19(2–3), 88–103. onomy, phylogeny, and adaptations. Bulletin American Museum Natural History 156, 157–450. 1974 61. (Szalay, F. S. and J. A. Wilson) Basicranial morphology of 43. (Szalay, F. S. and R. L. Decker) Origins, evolution and the early Tertiary tarsiiform from Texas. Folia function of the tarsus in late Cretaceous eutherians and Primatologica 25, 88–293. Paleocene primates. In: F.A. Jenkins, Jr. (Ed.). Primate 62. (Wilson, J. A. and F. S. Szalay) New adapid primate of Locomotion. Academic Press, New York, pp. 223–259. European affinities from Texas. Folia Primatologica 25, 44. (Decker, R. L. and F. S. Szalay) Origins and function of the 294–312. pes in the Eocene (Lemuriformes, Primates). In: F.A. Jenkins, Jr. (Ed.). Primate Locomotion. Academic, New 1977 York, pp. 261–291. 63. Constructing primate phylogenies: a search for testable 45. A review of some recent advances in paleoprimatology. hypothesis with maximum empirical content. Journal of American Journal of Physical Anthropology Yearbook (1973), Evolution 6(1), 3–18. 17, 39–64. 64. (Szalay, F. S. and D. Seligsohn) Why did the strepsirrhine tooth 46. Review of “Human evolution”, Vol. II of Symposia of the comb evolve? Folia Primatologica 27(1), 75–82. Society for the Study of Human Biology (Day, M. H., Ed.), 65. Ancestors, descendants, sister groups, and testing of phylogenetic Taylor & Francis/Barnes and Noble Books, London/New York; hypotheses. Systematic Zoology 26(1), 12–18. Evolution 28(3), 507–508. 66. Phylogenetic relationships and a classification of the euth- 47. (Seligsohn, D. and F. S. Szalay) Dental occlusion and the erian Mammalia; In: M.K. Hecht, P.C. Goody, and B.M. Hecht masticatory apparatus on Lemur and Varecia: their bearing (Eds.). Patterns of Vertebrate Evolution. Plenum, New York, on the systematics of living and fossil primates. In: Martin, pp. 315–374. R. D., B.A. Doyle and A.C. Walker (Eds.). Prosimian Biology., 67. (Wilson, J.A. and F. S. Szalay) Mahgarita, a new name for Duckworth, London, pp. 543–561. Margarita Wilson and Szalay, 1976. Journal of Paleontology 48. Comments on “New perspectives on and human evolu- 51(3), 643. tion”, article by A. Kortlandt. Current Anthropology 15(4), 68. (Seligsohn, D. and F. S. Szalay) Relationship between natu- 438–439. ral selection and dental morphology: tooth function and 49. New genera of European Eocene adapid primates. Folia diet in Lepilemur and Hapalemur. In: P. H. Butler, and K. A. Primatologica 22(2–3),116–133. Joysey (Eds.). Development, Function, and Evolution of Teeth. 50. A new and of early Eocene primate from North Academic, London. pp. 289–307. America. Folia Primatologica 22(4), 243–250. Bibliography of Frederick S. Szalay xiii

1978 1984 69. (Luckett, W. P. and F. S. Szalay) versus grades 85. Review of “Mammalian Paleofaunas of the World” by D. in primate phylogeny. In: D. J. Chivers and K. A. Joysey E. Savage and D.E. Russell. American Journal of Physical (Eds.). Recent Advances in Primatology Volume 3: Evolution. Anthropology 64, 205–206. Academic, London. pp. 227–237. 86. Arboreality: is it homologous in metatherian and eutherian mammals? Evolutionary Biology 18, 215–258. 1979 70. The problems of adaptation among Paleogene primates 1985 (Abs.) VIIth Congress of the International Primatological 87. and lagomorph morphotype adaptations, origins, Society, Bangalore, India. and relationships: some postcranial attributes analyzed. In: 71. (Dagosto, M. and F. S. Szalay) The elbow joint of early W. P. Luckett and J.-L. Hartenberger (Eds.). Evolutionary rela- primates (Abs.) American Journal of Physical Anthropology tionships among . NATO ASI Series A: Life Sciences 50(3), 431. Volume. 92, Plenum, New York, pp. 83–132. 72. (Szalay, F. S. and E. Delson) Evolutionary history of the 88. (Szalay, F. S. and J. Langdon) Evolutionary morphology of Primates. Academic, New York. the foot in Oreopithecus; (Abs.) American Journal of Physical Anthropology 66(2), 163. 1980 89. (Delson, E. and F. S. Szalay) Reconstruction of the 1958 73. (Szalay, F. S. and G. Drawhorn) Evolution and diversification cranium of Oreopithecus; (Abs.) American Journal of Physical of the Archonta in an arboreal milieu. In: W.P Luckett (Ed.). Anthropology, 66(2), 163. Comparative Biology and Evolutionary Relationships of Tree 90. Review of “Vertebrate Zoogeography and Evolution in Shrews. Plenum, New York, pp. 133–169. Australasia” by M. Archer and G. Clayton (Eds). American 74. (Szalay, F. S. and M. Dagosto) Locomotor adaptations as Scientist 73,173–174. reflected on the humerus of Paleogene primates. Folia Primatologica (34), 1–45. 1986 75. (Rosenberger, A. L. and F. S. Szalay) On the tarsiiform origins 91. (Szalay, F. S., C.-K. Li, and B.-Y. Wang) Middle Paleocene of the Anthropoidea; In: R. L. Ciochon and A. B. Chiarelli omomyid primate from Anhui Province, China: Decoredon (eds.) Evolutionary Biology of the New World Monkeys and anhuiensis (Xu, 1976), new combination Szalay and Li, and Continental Drift. Plenum, New York, pp. 139–157. the significance of Petrolemur. (Abs.) American Journal of Physical Anthropology 69(2), 269. 1981 92. (Szalay, F. S., and C.-K. Li) Middle Paleocene euprimate 76. Functional analysis and the practice of the phylogenetic from southern China, and the distribution of primates in the method as reflected by some mammalian studies. American Paleogene. Journal of Human Evolution 15, 387–397. Zoologist 21, 37–45. 93. (Szalay, F. S., and J. H. Langdon) The foot of Oreopithecus: 77. Review of “Mesozoic Mammals” by J. A. Lillegraven, Z. an evolutionary assessment. Journal of Human Evolution 15, Kielan-Jaworowska, and W. A. Clemens (Eds.). Journal of 585–621. Mammalogy 62, 443–445. 94. Pedal grasping in primate evolution. (Abs.) American Journal 78. Phylogeny and the problem of adaptive significance: the case of Physical Anthropology 72(2), 260–261. of the earliest primates. Folia Primatologica 36, 157–182. 1987 1982 95. Review of “Major Topics in Primate and Human Evolution” by 79. A critique of some recently proposed Paleogene primate taxa B. Wood, L. Martin, and P. Andrews (Eds.). Quarterly Review and suggested relationships. Folia Primatologica 37, 152–182. of Biology 62, 353. 80. A new appraisal of marsupial phylogeny and classifica- 96. (Szalay, F. S., A. L. Rosenberger, and M. Dagosto) Diagnosis tion; In: M. Archer (Ed.). Carnivorous Marsupials. Australian and differentiation of the order Primates. Yearbook of National University Press, Canberra, pp. 621–640. Physical Anthropology 30, 75–105. 81. (Flannery, T, and F. S. Szalay) Bohra paulae, a new giant fossil tree kangaroo (Marsupialia: Macropodidae) from New South 1988 Wales, Australia. Australian Mammalogy 5, 83–84. 97. A window into evolutionary primate systematics (a review of “Comparative Primate Biology, Volume 1: Systematics, 1983 Evolution, and Anatomy”, by D. R. Swindler and J. Erwin, 82. Phylogenetic relationships of the marsupials; Festschrift for Eds.), American Journal of Primatology, 14, 301–303. R. Hoffstetter, Geobios, Memoire speciale 6, 177–190. 98. (Szalay, F. S. and M. Dagosto) Evolution of hallucial grasp- 83. An eco-ethological reassessment of the living mammals. (A ing in the Primates. Journal of Human Evolution 17(1–2), review of “The Mammalian Radiations” by John E. Eisenberg). 1–33. Evolutionary Theory 6, 219–222. 99. Anaptomorphinae. In: I. Tattersall, E. Delson, and J. Van 84. Review of “Phylogenetic Patterns and Evolutionary Process” Couvering (Eds.). Encyclopedia of Human Evolution and by N. Eldredge and J. Cracraft; American Journal of Physical Prehistory. Garland Publishing, New York/London, pp. Anthropology 61, 509–510. 29–31. xiv Bibliography of Frederick S. Szalay

100. Apatemyidae. In: I. Tattersall, E. Delson, and J. Van 118. Tarsiiformes. In: I. Tattersall, E. Delson, and J. Van Couvering Couvering (Eds.). Encyclopedia of Human Evolution and (Eds.). Encyclopedia of Human Evolution and Prehistory. Prehistory. Garland Publishing, New York/London, p. 39. Garland Publishing, New York/London, pp. 568–570. 101. Archonta. In: I. Tattersall, E. Delson, and J. Van Couvering (Eds.). Encyclopedia of Human Evolution and Prehistory. 1989 Garland Publishing, New York/London, pp. 56–57. 119. Review of “Vertebrates, Phylogeny, and Philosophy: A tribute to 102. Carpolestidae. In: I. Tattersall, E. Delson, and J. Van Couvering George Gaylord Simpson” (K. M. Flanagan and J. A. Lillegraven, (Eds.). Encyclopedia of Human Evolution and Prehistory. Eds.). Journal of Human Evolution; 18(4), 415–416. Garland Publishing, New York/London, pp. 110–111. 120. Review of “The Phylogenetic System. The systematization 103. Donrussellia. In: I. Tattersall, E. Delson, and J. Van Couvering of organisms on the basis of their phylogenesis” by P. Ax; (Eds.). Encyclopedia of Human Evolution and Prehistory. American Journal of Physical Anthropology 78(1), 124–126. Garland Publishing, New York/London, p. 166. 121. On the evolutionary and interpretive frameworks of 104. Euprimates. In: I. Tattersall, E. Delson, and J. Van Couvering hominid lineage paleobiology. In: Evolutionary Biology at (Eds.). Encyclopedia of Human Evolution and Prehistory. the Crossroads (M. K. Hecht, Ed.), Queens College Press, Garland Publishing, New York/London, pp. 184–185. Flushing, New York, pp. 145–148. 105. Evolutionary Morphology. In: I. Tattersall, E. Delson, and J. Van Couvering (Eds.). Encyclopedia of Human Evolution 1990 and Prehistory. Garland Publishing, New York/London, pp. 122. (Szalay, F. S. and R. K. Costello) Evolution of permanent 199–200. estrus displays (PED) in the Hominidae. (Abs.), American 106. Evolutionary Systematics. In: I. Tattersall, E. Delson, and Journal of Physical Anthropology 81(2), 305. J. Van Couvering (Eds.). Encyclopedia of Human Evolution 123. Evolution of the tarsal complex in Mesozoic mammals. and Prehistory. Garland Publishing, New York/London, pp. (Abs.), Journal of Vertebrate Paleontology 9(3), 45A. 200–201. 124. (Szalay, F. S., and S. G. Lucas) Postcranial skeleton of 107. Haplorhini. In: I. Tattersall, E. Delson, and J. Van Couvering Mixodectes and a rediagnosis of the Primates. (Abs.), (Eds.). Encyclopedia of Human Evolution and Prehistory. Journal of Vertebrate Paleontology. 9(3), 45A. Garland Publishing, New York/London, p. 242–243. 108. Microchoerinae. In: I. Tattersall, E. Delson, and J. Van 1991 Couvering (Eds.). Encyclopedia of Human Evolution and 125. Review of “Functional morphology of the evolving hand and Prehistory. Garland Publishing, New York/London, pp. 339– foot” by O. J. Lewis; Journal of Physical Anthropology 84(4), 340. 493–495. 109. Microsyopidae. In: I. Tattersall, E. Delson, and J. Van 126. Causal analysis in historical-narrative explanations; (Abs.), Couvering (Eds.). Encyclopedia of Human Evolution and American Journal of Physical Anthropology Supplement 12, Prehistory. Garland Publishing, New York/London, p. 340. 171. 110. Omomyidae. In: I. Tattersall, E. Delson, and J. Van Couvering 127. (Szalay, F. S. and W. J. Bock) Evolutionary theory and (Eds.). Encyclopedia of Human Evolution and Prehistory. systematics: relationships between process and patterns. Garland Publishing, New York/London, pp. 396–397. Zeitschrift für zoologische Systematik und Evolutionsforschung 111. Paromomyidae. In: I. Tattersall, E. Delson, and J. Van 29, 1–39. Couvering (Eds.). Encyclopedia of Human Evolution and 128. The unresolved world between taxonomy and population Prehistory. Garland Publishing, New York/London, pp. 443– biology: what is, and what is not, macroevolution? (Lead 444. review-essay of Genetics, Paleontology, and Macroevolution by 112. Paromomyoidea. In: I. Tattersall, E. Delson, and J. Van Jeffrey Levinton) Journal of Human Evolution 20, 271–280. Couvering (Eds.). Encyclopedia of Human Evolution and 129. (Szalay, F. S. and R. K Costello) Evolution of permanent Prehistory. Garland Publishing, New York/London, p. 444– estrus displays in hominids. Journal of Human Evolution 20, 445. 439–464. 113. Picrodontidae. In: I. Tattersall, E. Delson, and J. Van Couvering (Eds.). Encyclopedia of Human Evolution and Prehistory. 1993 Garland Publishing, New York/London, p. 452. 130. (Szalay, F. S., M. J. Novacek, and M. C. McKenna, Eds.) 114. . In: I. Tattersall, E. Delson, and J. Van Couvering Mammal Phylogeny: Mesozoic differentiation, , (Eds.). Encyclopedia of Human Evolution and Prehistory. Therians, and Marsupials. Springer, New York, 249 pp. Garland Publishing, New York/London, pp. 464–466. 131. (Szalay, F. S., M. J. Novacek, and M. C. McKenna, Eds.) Mammal 115. Plesiadapiformes. In: I. Tattersall, E. Delson, and J. Van Phylogeny: Placentals; Springer, New York, 341 pp. Couvering (Eds.). Encyclopedia of Human Evolution and 132. Introduction. In: F. S. Szalay, M. J. Novacek, and M. Prehistory. Garland Publishing, New York/London, pp. 466. C. McKenna (Eds.). Mammal Phylogeny: Mesozoic 116. . In: I. Tattersall, E. Delson, and J. Van Differentiation, Monotremes, Therians, and Marsupials. Couvering (Eds.). Encyclopedia of Human Evolution and Springer, New York, pp. 1–3. Prehistory. Garland Publishing, New York/London, p. 466– 133. Pedal in the Mesozoic: tests for 467. taxic relationships. In: F. S. Szalay, M. J. Novacek, and 117. Saxonellidae. In: I. Tattersall, E. Delson, and J. Van Couvering M. C. McKenna (Eds.). Mammal Phylogeny: Mesozoic (Eds.). Encyclopedia of Human Evolution and Prehistory. Differentiation, Monotremes, Therians, and Marsupials. Garland Publishing, New York/London, p. 504. Springer, New York, pp. 108–128. Bibliography of Frederick S. Szalay xv

134. Metatherian taxon phylogeny: evidence and interpreta- 1997 tion from the cranioskeletal system. In: F. S. Szalay, M. J. 147. The phylogenetic affinities of Dromiciops and the osteology Novacek, and M. C. McKenna (Eds.). Mammal Phylogeny: of South American marsupials. Noticiero de Biología (Organo Mesozoic Differentiation, Monotremes, Therians, and oficial de sociedad de biología de Chile) 5(4), 48. Marsupials. Springer, New York, pp. 216–242. 135. Introduction. In: F. S. Szalay, M. J. Novacek, and M. C. 1998 McKenna (Eds.). Mammal Phylogeny: Placentals. Springer, 148. (Szalay, F. S., and Schrenk, F.) The middle Eocene New York, pp. 1–4. Eurotamandua and a Darwinian phylogenetic analysis of 136. Species concepts: the tested, the untestable, and the redun- the “edentates”. Kaupia 7, 97–186. dant. In: W. H. Kimbel, and L. B. Martin (Eds.). Species, Species Concepts, and Primate Evolution. Plenum, New York, 1999 pp. 21–41. 149. Review of Classification of mammals above the species level, 137. (Costello, R. K., C. Dickinson, A. L. Rosenberger, S. Boinski, by M. C. McKenna and S. K. Bell (with contributions from and F. S. Szalay). (Genus Saimiri) tax- G.G. Simpson, R. H. Nichols, R. H. Tedford, K. F. Koopman, onomy: a multidisciplinary study of the biology of species. G. G. Musser, N. A. Neff, J. Shoshani, and D. M. McKenna), In: W. H. Kimbel, and L. B. Martin (Eds.). Species, Species 1997, Columbia University Press, New York, 631 + xii pp., in Concepts, and Primate Evolution. Plenum, New York, pp. Journal of Vertebrate Paleontology 19, 191–195. 177–210. 150. Paleontology and Macroevolution: On the theoretical con- 138. (Szalay, F. S. and S. G. Lucas) Cranioskeletal morphology flict between an expanded synthesis and hierarchic punc- of archontans, and Diagnoses of Chiroptera, Volitantia, tuationism. In: T. G. Bromage, and F. Schrenk (Eds.). African and Archonta. In: R. D. E. MacPhee (Ed.). Primates and their Biogeography, Climate Change, and Human Evolution. Oxford Relatives in Phylogenetic Perspective. Plenum, New York, pp. University Press, New York, pp. 35–56. 187–226. 151. (Szalay, F. S., and Sargis, E. J.) Paleocene marsupial postcrani- 139. (Trofimov, B. A., and F. S. Szalay) New group of Asiatic mar- als from Itaboraí, Brazil: Model-based analysis of adaptations, supials (order Asiadelphia) from the Late Cretaceous of phylogeny, & biogeography. Journal of Vertebrate Paleontology Mongolia. Journal of Vertebrate Paleontology 13(Supplement 19(Supplement 3), 80A. 3), 60A. 2000 1994 152. Review of “Walker’s Mammals of the World. Sixth Edition, 140. (Szalay, F. S., and F. Schrenk) Middle Eocene Eurotamandua Volumes I and II” (Nowak R. M.), 1999, Johns Hopkins and the early differentiation of the Edentatata. Journal of University Press, Baltimore, MD. The Quarterly Review of Vertebrate Paleontology 14 (Supplement 3), 48A. Biology 75(1), 71–72. 141. (Trofimov, B. A., and Szalay, F. S.) New Cretaceous mar- 153. Marsupials. In: R. Singer (Ed.). Encyclopedia of Paleontology. supial from Mongolia and the early radiation of the Fitzroy Dearborn Publishers pp. 703–714. Metatheria. Proceedings of the National Academy of Sciences 154. (Stafford, B. J., and F. S. Szalay). Craniodental functional USA 91, 12569–12573. morphology and taxonomy of dermopterans. Journal of 142. Evolutionary history of the marsupials and an analysis of Mammalogy 81(20), 360–385. osteological characters. Cambridge University Press, New 155. (Szalay, F. S., E. J. Sargis, and B. J. Stafford). Small marsu- York, 481 pp. pial glider from the Paleocene of Itaboraí, Brazil. Journal of Vertebrate Paleontology 20(Supplement 3), 73A. 1995 156. (Schrenk, F., and F. S. Szalay). Enigmatic new mammal 143. (Emry, R. J., S. G. Lucas, F. S. Szalay, and P. A. Tleuberdina) (Dermoptera?) from the Messel middle Eocene, Germany. A new herpetotheriine didelphid (Marsupialia) from the Journal of Vertebrate Paleontology 20(Supplement 3), 68A. of Central Asia. Journal of Vertebrate Paleontology 157. (Warshaw, J., S. C. McFarlin, T. G. Bromage, and F. S. Szalay). 15, 850–854. Some bone microstructure variables in extant therians, 144. Review of Evolution of the cercopithecoid Forelimb – and their relationship to life history, locomotion, and phy- Phylogenetic and Functional Implications from Morphometric logeny. Journal of Vertebrate Paleontology 20(Supplement 3), Analyses, by R. L. Ciochon, 1994, University of California 76A–77A. Publications, Geological Sciences, Volume 138, xxi + 251 pp., 158. Review of “The Atlas of European Mammals” (Mitchell- in Journal of Vertebrate Paleontology 15, 214–215. Jones, A. J. et al., Eds.) Academic, London. The Quarterly Review of Biology 75(4), 472. 1996 159. (Goldman H. M, S. C. McFarlin, J. Warshaw, F. S. Szalay., 145. (Szalay, F. S., and S. G. Lucas) The postcranial morphology and T.G. Bromage.). Application of bone microstructural of Paleocene Chriacus and Mixodectes and the phylogenetic analysis to the comparative study of primate functional relationships of archontan mammals. Bulletin New Mexico adaptation and life history. American Journal of Physical Museum of Natural History 7, 1–47. Anthropology (Supplement 30), 160–161. 146. (Szalay, F. S., and B. A. Trofimov) The Mongolian Late 160. Function and Adaptation in Paleontology and Phylogenetics: Cretaceous Asiatherium, and the early phylogeny and Why Do We Omit Darwin? Palaeontologia Electronica 3(2), paleobiogeography of Metatheria. Journal of Vertebrate 25 pp., 372KB. http://palaeo-electronica.org/2000_2/darwin/ Paleontology 16, 474–509. issue2_00.htm xvi Bibliography of Frederick S. Szalay

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191. Plesiadapiformes. In: E. Delson, I. Tattersall, J. A. Van 2002 Couvering and A. S Brooks (Eds.). Encyclopedia of Human 203. (Warshaw J., T. G. Bromage, C. J. Terranova, F. S. Szalay). Evolution and Prehistory (2nd Edition). Garland Publishing, Diversity in bone collagen fiber orientation patterns New York, pp. 572–573. among primates and other mammals. Journal of Vertebrate 192. Plesiadapoidea. In: E. Delson, I. Tattersall, J. A. Van Paleontology 22(Supplement 3), 117A–118A. Couvering and A. S Brooks (Eds.). Encyclopedia of Human Evolution and Prehistory (2nd Edition). Garland Publishing, 2003 New York, pp. 573–574. 204. (Warshaw J., T. G. Bromage, C. J. Terranova, F.S. Szalay). 193. Saxonellidae. In: E. Delson, I. Tattersall, J. A. Van Couvering Primate bone microstructural variability: relationships to and A. S Brooks (Eds.). Encyclopedia of Human Evolution mechanical and life history adaptation. American Journal of and Prehistory (2nd Edition). Garland Publishing, New York, Physical Anthropology Supplement 36, 220. p. 626. 194. Shoshonius. In: E. Delson, I. Tattersall, J. A. Van Couvering 2004 and A. S Brooks (Eds.). Encyclopedia of Human Evolution 205. (Salton, J. A. and F. S. Szalay). The tarsal complex of and Prehistory (2nd Edition). Garland Publishing, New York, Afro-Malagasy Tenrecoidea: A search for phylogenetically pp. 635–636. meaningful characters. Journal of Mammalian Evolution 11, 195. Tarsiiformes. In: E. Delson, I. Tattersall, J. A. Van Couvering 73–104. and A. S Brooks (Eds.). Encyclopedia of Human Evolution 206. Review of “Late Cretaceous and Cenozoic Mammals of and Prehistory (2nd Edition). Garland Publishing, New York, : Biostratigraphy and Geochronology” (M. O. pp. 691–693. Woodburne, Ed.), 2004, Journal of Mammalian Evolution 11, 196. Tarsioidea. In: E. Delson, I. Tattersall, J. A. Van Couvering 205–206. and A. S Brooks (Eds.). Encyclopedia of Human Evolution and Prehistory (2nd Edition). Garland Publishing, New York, 2005 pp. 693–694. 207. Review of “The Rise of Placental Mammals. Origins and 197. Visual hypothesis. In: E. Delson, I. Tattersall, Relationships of the Major Clades” (Rose, K. D., and J. D. J. A. Van Couvering and A. S Brooks (Eds.). Encyclopedia Archibald, Eds.), 2005, Journal of Mammalian Evolution 12, of Human Evolution and Prehistory (2nd Edition). Garland 533–542. Publishing, New York, pp. 725–726. 2006 2001 208. Review of “Macroevolution-Diversity, Disparity, Contingency: 198. (Szalay, F. S., and E. J. Sargis). Model-based analysis of Essays in Honor of Stephen Jay Gould” (Vrba E., and N. Eldredge, postcranial osteology of marsupials from the Palaeocene Eds.), 2006, Journal of Mammalian Evolution 13, 165–166. of Itaboraí (Brazil), and the phylogenetics and biogeog- 209. (Szalay, F. S., and E. J. Sargis). Cretaceous therian tarsals raphy of Metatheria. Geodiversitas 23(2), 139–302 (con- and the Metatherian-Eutherian dichotomy. Journal of currently published on the Geodiversitas site: http://www. Mammalian Evolution 13, 171–210. mnhn.fr/publication/geodiv/g01n2a1.pdf) 199. (McFarlin, S. C, Warshaw J., Szalay F. S., and Bromage 2007 T. G.). Microstructural features in mammalian bone as 210. Ancestral locomotor modes, placental mammals, and the ori- indicators of organismal life history: a survey of the order gin of euprimates: Lessons from history. In: M. J. Ravosa and Primates. Journal of Vertebrate Paleontology 21(Supplement M. Dagosto (Eds.). Primate Origins: Adaptations and Evolution. 3), 79A. Springer, New York, pp. 457–487. 200. Problems with hominid fossil species taxa and the con- 211. (Chester, S. G. B., Sargis, E. J., Szalay, F. S., Archibald, J. D., struction of taxograms. Ludus Vitalis 9, 143–169. Averianov, A. O.) Functional analysis of mammalian humeri 201. (Szalay, F. S., and F. Schrenk). An enigmatic new mammal from the Late Cretaceous of Uzbekistan. Journal of Vertebrate (Dermoptera?) from the Messel Middle Eocene, Germany. Paleontology 27(Supplement), 58A. Kaupia 11, 153–164. 202. (Warshaw J., S.C. McFarlin, T. G. Bromage, C. J. Terranova, In press F. S. Szalay). The effects of life history, locomotion and 212. Connections between modeling, vertebrate paleobiol- phylogeny on bone microstructural features in mammals. ogy, and systematics. American Zoologist (Walter J. Bock Journal of Morphology 248(3), 298. Festschrift volume). References (not including those listed in Szalay bibliography)

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xix Acknowledgments

We thank Rob Asher, Bert Covert, Darin Croft, John Fleagle, for reviewing multiple manuscripts. We also thank the series Dick Fox, Haviva Goldman, Terry Harrison, Luke Holbrook, editors, Eric Delson and Ross MacPhee, for all their help with Zhexi Luo, Greg McDonald, Mike Plavcan, James Rossie, this volume. Finally, we thank Frederick S. Szalay who served Guillermo Rougier, Tony Tosi, Blaire Van Valkenburgh, as our graduate advisor, and to whom we both owe a huge debt. Robert Walker, John Wible, and Steve Wroe for serving as Although it will be impossible to fully repay that debt, we con- outside reviewers of the chapters in this book, and especially sider this volume to be a small token of our appreciation for all Carl Terranova, Erik Seiffert, Bruce Shockey, and Mary Silcox the support, encouragement, and friendship he has provided.

xxi List of Contributors

Federico Anaya Richard L. Cifelli Facultad de Ingeniería Geológica Sam Noble Oklahoma Museum of Natural History Universidad Autónoma “Tomás Frías” and Department of Zoology Potosí, Bolivia University of Oklahoma 2401 Chautauqua Ave. Christine Argot Norman, OK 73072, USA Département Histoire de la Terre [email protected] Muséum national d’Histoire naturelle 8 rue Buffon Sébastien Couette 75005 Paris, France Ecole Pratique des Hautes Etudes [email protected] UMR 5143 Paléobiodiversité et Paléoenvironnements K. Christopher Beard Case Courrier 38 Section of Vertebrate Paleontology Département d’Histoire de la Terre Carnegie Museum of Natural History Muséum national d’Histoire naturelle 4400 Forbes Avenue 8 rue Buffon Pittsburgh, PA 15213, USA 75005 Paris, France [email protected] [email protected] Lilian P. Bergqvist Marian Dagosto Universidade Federal do Rio de Janeiro Department of Cell and Molecular Biology Departamento de Geologia Feinberg School of Medicine Av. Athos da Silveira Ramos, 274 Northwestern University Rio de Janeiro/RJ 303 E. Chicago Ave. 21941-916 Brasil Chicago, IL 60611, USA [email protected] [email protected] Jonathan I. Bloch Brian M. Davis Florida Museum of Natural History Sam Noble Oklahoma Museum of Natural History and University of Florida Department of Zoology P.O. Box 117800 University of Oklahoma Gainesville, FL 32611, USA 2401 Chautauqua Ave. [email protected] Norman, OK 73072, USA Doug M. Boyer [email protected] Department of Anatomical Sciences Eric Delson Stony Brook University Department of Anthropology, Lehman College/CUNY T8 040 Health Science Center New York Consortium in Evolutionary Primatology Stony Brook, NY 11733, USA (NYCEP) [email protected] Department of Vertebrate Paleontology

xxiii xxiv List of Contributtors

American Museum of Natural History Benjamin P. Kear Central Park West & 79th Street Department of Genetics New York, NY 10024, USA School of Molecular Sciences [email protected] La Trobe University Melbourne, Victoria 3086 Timothy F. Flannery South Australian Museum Division of Environment and Life Sciences North Terrace, Adelaide Macquarie University South Australia 5000, Australia Sydney, 2109, Australia [email protected] Stephen R. Frost Zofia Kielan-Jaworowska Department of Anthropology Instytut Paleobiologii PAN University of Oregon ul. Twarda 51/55 Eugene, OR 97403, USA PL-00-818 Warszawa, Poland [email protected] [email protected] Daniel. L. Gebo Michael S. Y. Lee Department of Anthropology South Australian Museum Northern Illinois University North Terrace, Adelaide DeKalb, IL 60115, USA South Australia 5000, [email protected] Australia Wayne R. Gerdtz Wolfgang Maier School of Ecology and Environment Lehrstuhl Spezielle Zoologie Melbourne Campus Universität Tübingen Deakin University Auf der Morgenstell 28 Burwood, Victoria D-72076 Tübingen, Germany 3125, Australia [email protected] Marc Godinot Xijun Ni Ecole Pratique des Hautes Etudes Department of Vertebrate Paleontology UMR 5143 Paléobiodiversité et American Museum of Natural History Paléoenvironnements Central Park West & 79th Street Case Courrier 38 New York, NY 10024, USA Département d’Histoire de la Terre [email protected] Muséum national d’Histoire naturelle 8 rue Buffon Jay A. O’Sullivan 75005 Paris, France Department of Exercise Science and [email protected] Sport Studies University of Tampa William E. H. Harcourt-Smith 401 West Kennedy Boulevard Department of Vertebrate Paleontology Tampa, FL 33606, USA American Museum of Natural History [email protected] Central Park West & 79th Street New York, NY 10024, USA Tonya A. Penkrot [email protected] Marshall University Department of Biological Sciences Russell Hogg 1 John Marshall Drive Department of Anthropology Huntington, WV 25755, USA The Graduate Center [email protected] City University of New York 365 Fifth Avenue P. David Polly New York, NY 10016, USA Department of Geological Sciences [email protected] Indiana University New York Consortium in Evolutionary 1001 East 10th Street Primatology (NYCEP) Bloomington, IN 47405, USA Hard Tissue Research Unit [email protected] New York University List of Contributtors xxv

Tao Qi Mary T. Silcox Institute of Vertebrate Paleontology and Paleoanthropology University of Winnipeg Chinese Academy of Sciences Department of Anthropology Beijing, 100044, China 515 Portage Ave. Winnipeg, MB R3B 2E9, Canada F. James Rohlf [email protected] Department of Ecology and Evolution Stony Brook University Melissa Tallman Stony Brook, NY 11794, USA Department of Anthropology [email protected] The Graduate Center City University of New York Kenneth D. Rose 365 Fifth Avenue Johns Hopkins University School of Medicine New York, NY 10016, USA Center for Functional Anatomy and Evolution [email protected] 1830 E. Monument St. New York Consortium in Evolutionary Baltimore, MD 21205, USA Primatology (NYCEP) [email protected] Carl J. Terranova Alfred L. Rosenberger Department of Anatomy Department of Anthropology and Archaeology Touro College of Osteopathic Medicine Brooklyn College, CUNY 230 West 125th St. 2900 Bedford Ave. New York, NY 10027, USA Brooklyn, NY 11210, USA [email protected] [email protected] Department of Anthropology Johanna Warshaw The Graduate Center, CUNY Hard Tissue Research Unit New York Consortium in Evolutionary Departments of Biomaterials & Basic Sciences Primatology (NYCEP) Rm 817-S American Museum of Natural History/Mammalogy New York University College of Dentistry NYU Mail Code: 9448 Justine A. Salton 345 East 24th Street Program in Biology New York, NY 10010, USA Bard College [email protected] Annandale-on-Hudson, NY 12504, USA David F. Wiley Eric J. Sargis Institute for Data Analysis and Department of Anthropology Visualization Yale University University of California P.O. Box 208277 Davis, CA 95616, USA New Haven, CT 06520, USA [email protected] [email protected] Division of Vertebrate Zoology Sai Man Wong Peabody Museum of Natural History Department of Anthropology and Archaeology Brooklyn College, CUNY Bruce J. Shockey 2900 Bedford Ave. Biology Department Brooklyn, NY 11210, USA Manhattan College [email protected] Manhattan College Parkway Riverdale, NY 10471, USA Shawn P. Zack [email protected] Marshall University Department of Vertebrate Paleontology Department of Biological Sciences American Museum of Natural History 1 John Marshall Drive Central Park West & 79th Street Huntington, WV 25755, USA New York, NY 10024, USA [email protected] Contents

Section I: Non-primate Mammals 1. Earliest Evidence of (Mammalia: Metatheria) from the Early Cretaceous of North America ...... 3 Brian M. Davis, Richard L. Cifelli, and Zofia Kielan-Jaworowska 2. Evolution of Hind Limb Proportions in Kangaroos (Marsupialia: Macropodoidea) ...... 25 Benjamin P. Kear, Michael S. Y. Lee, Wayne R. Gerdtz, and Timothy F. Flannery 3. Changing Views in Paleontology: The Story of a Giant (Megatherium, Xenarthra) ...... 37 Christine Argot 4. Evolutionary Morphology of the Tenrecoidea (Mammalia) Forelimb Skeleton ...... 51 Justine A. Salton and Eric J. Sargis 5. Postcranial Morphology of Apheliscus and Haplomylus (Condylarthra, Apheliscidae): Evidence for a Paleocene Holarctic Origin of Macroscelidea ...... 73 Tonya A. Penkrot, Shawn P. Zack, Kenneth D. Rose, and Jonathan I. Bloch 6. Postcranial Skeleton of the Upper Paleocene (Itaboraian) “Condylarthra” (Mammalia) of Itaboraí Basin, Brazil ...... 107 Lilian P. Bergqvist 7. Postcranial Osteology of Mammals from Salla, Bolivia (Late Oligocene): Form, Function, and Phylogenetic Implications ...... 135 Bruce J. Shockey and Federico Anaya 8. Evolution of the Proximal Third Phalanx in Oligocene-Miocene Equids, and the Utility of Phalangeal Indices in Phylogeny Reconstruction ...... 159 Jay A. O’Sullivan 9. Adaptive Zones and the Ankle: A Three-Dimensional Quantitative Analysis of Carnivoran Tarsal Evolution ...... 167 P. David Polly

Section II: Primates 10. The Biogeographic Origins of Primates and Euprimates: East, West, North, or South of Eden? ...... 199 Mary T. Silcox 11. Evaluating the Mitten-Gliding Hypothesis for Paromomyidae and Micromomyidae (Mammalia, “Plesiadapiformes”) Using Comparative Functional Morphology of New Paleogene Skeletons ...... 233 Doug M. Boyer and Jonathan I. Bloch

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12. Morphological Diversity in the Skulls of Large Adapines (Primates, ) and Its Systematic Implications ...... 285 Marc Godinot and Sébastien Couette 13. Primate Tibiae from the Middle Eocene Shanghuang Fissure-Fillings of Eastern China ...... 315 Marian Dagosto, Daniel L. Gebo, Xijun Ni, Tao Qi, and K. Christopher Beard 14. Rooneyia, Postorbital Closure, and the Beginnings of the Age of Anthropoidea ...... 325 Alfred L. Rosenberger, Russell Hogg, and Sai Man Wong 15. Epitensoric Position of the Chorda Tympani in Anthropoidea: a New Synapomorphic Character, with Remarks on the Fissura Glaseri in Primates ...... 347 Wolfgang Maier 16. Evolutionary Morphology of the Guenon Postcranium and Its Taxonomic Implications ...... 361 Eric J. Sargis, Carl J. Terranova, and Daniel L. Gebo 17. Analysis of Selected Hominoid Joint Surfaces Using Laser Scanning and Geometric Morphometrics: A Preliminary Report ...... 373 William E. H. Harcourt-Smith, Melissa Tallman, Stephen R. Frost, David F. Wiley, F. James Rohlf, and Eric Delson 18. Comparative Primate Bone Microstructure: Records of Life History, Function, and Phylogeny ...... 385 Johanna Warshaw Taxonomic Index ...... 427 Subject Index ...... 435