Mammalian Evolutionary Morphology: A Tribute to Frederick S. Szalay

452 Pages·2008·12.35 MB·English

by  Brian M. Devis

Mammalian Evolutionary Morphology
Vertebrate 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 vertebrates (from agnathans to humans),
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Volumes in the series may either be monographic treatments (including unpublished but fully revised dissertations) or edited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 (e-book)
Published by Springer,
P.O. Box 17, 3300 AA Dordrecht, The Netherlands.
www.springer.com
Cover illustration:
Dryomomys szalayi, drawing by Doug M. Boyer.
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No part of this work may be reproduced, stored in a retrieval system, or transmitted
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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 Primates, never finished his last year in Gymnasium (high school).
Archonta, and Metatheria, 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 evolutionary biology 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
primatesJemezius 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 marsupials Szalinia gracilis His consummate interests in mammalian natural history had
(de Muizon and Cifelli, 2001), Sinodelphys 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
multituberculateEctypodus szalayi (Sloan, 1981). mammals that he worked with while living on the premises
vii
viii Preface
set the stage for his plans for a Mammalogy Ph.D. at the Primate 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 Plesiadapiformes 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 Haplorhini (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 Paleocene and Eocene 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 Omomyidae (Tarsiiformes, and Kear et al. in this volume.
Primates): Taxonomy, 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
Theory and Practice of Phylogeny
supraordinal grouping Archonta, and its adaptive signifi-
Reconstruction/Adaptive Scenarios
cance. Szalay (1977, #66) provided the first morphological
support for McKenna’s (1975) revised concept of Gregory’s
(1910) Archonta, a clade 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, Euarchonta (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 marsupial relationships based on tarsal evi-
cally Australidelphia), as well as many other mammalian groups
dence. He hypothesized that the South American Dromiciops
such as Glires, Xenarthra, 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 Ameridelphia. 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
Department of Anthropology
they are reciprocally illuminatory (if we might borrow that
Yale University
Hennigian phrase) (Szalay, 1981, #78; Szalay and Bock, 1991,
New Haven, CT 06520, USA
#127; Szalay, 2000, #160). In fact, “The meeting of these two
[email protected]
‘separate’ disciplines is of course what is usually referred to as
Division of Vertebrate Zoology
morphology” (Szalay, 1981, #78, p. 160). This point of view is
Peabody Museum of Natural History
represented in his concept of the “transformation series”, a test-
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. Amphipithecusand 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(Insectivora, 15. On the cranium of the late Paleocene primate Plesiadapis
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.
Hyaenodontidae). 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. Proboscidea. Encyclopedia of Science and Technology.
1968 McGraw-Hill, New York, pp. 731–732.
6. The beginnings of primates. Evolution 22(1), 19–36. 23. Sirenia.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 Apatemyidae (Mammalia, Insectivora). 295–296.
American Museum Novitates 2352, 1–11. 26. ( Davis, D. D. and F. S. Szalay) Eutheria. 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) Theria.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 Cretaceous mammalDeltatheridium. 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 mesonychid feeding Mongolia. Bulletin of the American Museum of Natural History
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xi