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SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION
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Smithsonian Contributions to the Marine Sciences
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Lawrence M. Small
Secretary
Smithsonian Institution
Cenozoic Mammals of Land and Sea:
Tributes to the Career
of Clayton E. Ray
Robert J. Emry
Editor
Smithsonian Institution Press
Washington, D.C.
2002
ABSTRACT
Emry, Robert J., editor. Cenozoic Mammals of Land and Sea: Tributes to the Career of Clay¬
ton E. Ray. Smithsonian Contributions to Paleobiology, 93, 372 pages, 197 figures, 11 plates,
46 tables, 2002.—This is a volume of collected papers published to honor the career of Clayton
E. Ray, now Curator Emeritus in the Department of Paleobiology, National Museum of Natural
History, Smithsonian Institution, and Curator of Late Cenozoic Mammals and of Fossil Marine
Mammals in the same department for more than 30 years before his retirement in 1994. The
volume includes a preface, a biography and bibliography of Clayton E. Ray, and 19 papers
devoted principally to Pleistocene mammals and to fossil marine mammals. Gary Morgan
describes late Pleistocene mammalian faunas from several sites in southernmost Florida and
discusses the Neotropical influence in Florida’s Pleistocene faunas. Richard H. Tedford
describes the basicranium of the Pleistocene giant wombat Phascolonus gigas Owen and dis¬
cusses its significance in marsupial phylogenetic reconstruction. Gerardo De luliis and A. Gor¬
don Edmund describe Vassallia maxima Castellanos, the only pre-Pleistocene pampathere
known in which a skull and mandible are associated with osteoderms; the range of osteoderm
variation in one associated individual allows them to synonymize other taxa that had been
based on osteoderm differences. Paul W. Parmalee and Russell Wm. Graham report additional
records of the giant beaver, Castoroides, from the mid-South. Frederick Grady, Joaquin
Arroyo-Cabrales, and E. Ray Garton report the northernmost known occurrence of vampire
bats in the Pleistocene of eastern North America. H. Gregory McDonald reports the second
known occurrence of the badger Taxidea taxus in the Pleistocene of Kentucky and discusses
the paleoecological implications of the occurrence. Jerry N. McDonald and George E. Lam-
mers describe Bison antiquus from Ontario and discuss the evolution of bison in the Holocene
of North America. Daryl P. Domning presents a new analysis and interpretation of the terres¬
trial posture in desmostylians. Thomas A. Demere and Annalisa Berta describe new material
and present a phylogenetic analysis of the Miocene pinniped Desmatophoca oregonensis from
Oregon. Irina A. Koretsky and Dan Grigorescu describe and evaluate the systematic position of
the fossil monk seal Pontophoca sarmatica from the Miocene of eastern Europe. Irina A.
Koretsky and Peter Holec describe a new, primitive, phocid pinniped from the early middle
Miocene of Slovakia and discuss its bearing on the phylogeny and classification of pinnipeds.
Irina A. Koretsky and Albert E. Sanders report remains of the oldest known phocid pinniped
from the late Oligocene of South Carolina. R. Ewan Fordyce describes and discusses a bizarre
archaic Oligocene dolphin from the eastern North Pacific, on which he bases a new species,
genus, and subfamily. Christian de Muizon, Daryl P. Domning, and Darlene R. Ketten describe
and discuss the paleobiology and behavior of an unusual walrus-convergent delphinoid ceta¬
cean from the early Pliocene of Peru. Susan D. Dawson and Michael D. Gottfried report paleo-
pathologic conditions in a Miocene odontocete cetacean. Albert E. Sanders and Lawrence G.
Barnes contribute two papers, both describing and analyzing new, primitive, cetotheriid mys-
ticete cetaceans from the late Oligocene of South Carolina. James W. Westgate and Frank C.
Whitmore, Jr., describe a new species of bowhead whale from the Pliocene Yorktown Forma¬
tion in Virginia. James G. Mead and Rosemary G. Dagit present an account of the search for
the 1880s manuscript of J.A. Allen’s unpublished monograph on the mammalian orders Cete
and Sirenia; the manuscript was not found but the 12 plates that were prepared for it are pub¬
lished herein.
Official publication date is handstamped in a limited number of initial copies and is
recorded in the Institution’s annual report. Annals of the Smithsonian Institution. SERIES COVER
DESIGN: The trilobite Phacops rana Green.
Library of Congress Cataloging-in-Publication Data
Cenozoic mammals of land and sea : tributes to the career of Clayton E. Ray / Robert .1, Emry
p. cm.—(Smithsonian contributions to paleobiology ; no. 93)
1. Mammals, Fossil. 2. Paleontology—Cenozoic. 1. Emry, Robert J. 11. Series.
QE881 .C46 2002 569-dc21 2002030521
© The paper used in this publication meets the minimum requirements of the American
National Standard for Permanence of Paper for Printed Library Materials Z39.48_1984
Contents
Page
Preface. i
Biography and Bibliography of Clayton Edward Ray
Ralph E. Eshelman, Robert J. Emry, Daryl R Damning, and David J. Bohaska.1
Late Rancholabrean Mammals from Southernmost Florida, and the Neotropical In¬
fluence in Florida Pleistocene Faunas
Gary S. Morgan .15
The Basicranium of the Giant Wombat Phascolonus gigas Owen (Vombatidae: Mar-
supialia) and Its Significance in Phytogeny
Richard H. Tedford .39
Vassallia maxima Castellanos, 1946 (Mammalia: Xenarthra: Pampatheriidae), from
Puerta del Corral Quemado (Late Miocene to Early Pliocene), Catamarca
Province, Argentina
Gerardo De Iidiis and A. Gordon Edmund.49
Additional Records of the Giant Beaver, Castoroides, from the Mid-South; Alabama,
Tennessee, and South Carolina
Paul W. Parmalee and Russell Wm. Graham.65
The Northernmost Occurrence of the Pleistocene Vampire Bat Desmodus stocki Jones
(Chiroptera: Phyllostomatidae: Desmodontinae) in Eastern North America
Frederick Grady, Joaquin Arroyo-Cabrales, and E. Ray Garton .73
Second Record of the Badger Taxidea taxus (Schreber) from the Pleistocene of Ken-
mcky and Its Paleoecological Implications
H. Gregory McDonald .77
Bison antiquus from Kenora, Ontario, and Notes on the Evolution of North Ameri¬
can Holocene Bison
Jerry N. McDonald and George E. Lammers.83
The Terrestrial Posture of Desmostylians
Daryl P Damning.99
The Miocene Pinniped Desmatophoca oregonensis Condon, 1906 (Mammalia; Car¬
nivora), from the Astoria Formation, Oregon
Thomas A. Demere and Annalisa Berta.113
The Fossil Monk Seal Pontophoca sarmatica (Alekseev) (Mammalia: Phocidae:
Monachinae) from the Miocene of Eastern Europe
Irina A. Koretsky and Dan Grigorescu.149
A Primitive Seal (Mammalia: Phocidae) from the Early Middle Miocene of Central
Paratethys
Irina A. Koretsky and Peter Holec .163
Paleontology of the Late Oligocene Ashley and Chandler Bridge Formations of
South Carolina, 1: Paleogene Pinniped Remains; The Oldest Known Seal
(Carnivora: Phocidae)
Irina A. Koretsky and Albert E. Sanders.179
Simocetus rayi (Odontoceti: Simocetidae, New Family): A Bizarre New Archaic Oli¬
gocene Dolphin from the Eastern North Pacific
R. Ewan Fordyce. 185
Odobenocetops peruvianas, the Walrus-Convergent Delphinoid (Mammalia: Ceta¬
cea) from the Early Pliocene of Peru
Christian de Muizon, Daryl P. Damning, and Darlene R. Ketten .223
iii
IV SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY
Paleopathology in a Miocene Kentriodontid Dolphin (Cetacea: Odontoceti)
Susan D. Dawson and Michael D. Gottfried .263
Paleontology of the Late Oligocene Ashley and Chandler Bridge Formations of
South Carolina, 2: Micromysticetus rothauseni, a Primitive Cetotheriid Mys-
ticete (Mammalia: Cetacea)
Albert E. Sanders and Lawrence G. Barnes.271
Balaena ricei, a New Species of Bowhead Whale from the Yorktown Formation
(Pliocene) of Hampton, Virginia
James W. Westgate and Frank C. Whitmore, Jr..295
Paleontology of the Late Oligocene Ashley and Chandler Bridge Formations of
South Carolina, 3: Eomysticetidae, a New Family of Primitive Mysticetes
(Mammalia: Cetacea)
Albert E. Sanders and Lawrence G. Barnes .313
Plates for Joel Asaph Allen’s Unpublished Monograph on the Mammalian Orders
Cete and Sirenia and a Record of the Search for the Manuscript
James G. Mead and Rosemary G. Dagit.357
Preface
When Clayton Ray retired from the National Museum of Natural History at the end of
1994, those of us who had been closely associated with him for many years, who had sat
down to lunch with him several times each week, who had talked with him almost daily,
were left with the feeling that we should do something to recognize his career and his con¬
tributions to his profession. Clayton, typically, had discouraged any kind of social event,
such as a department party or luncheon, to celebrate the occasion of his retirement. So,
mindful of what Clayton’s advice would have been had we asked for it, we opted to do
something useful instead: we invited some of his colleagues, mainly those who had
worked closely with him on certain projects, who had been his coauthors, or who had
studied groups of special interest to him, to contribute papers to be assembled as a
festschrift. The process of completing this volume has gone on longer than intended (al¬
though not so much longer than expected), but this result surely makes the wait worth¬
while. Although the “Contents” might suggest that a rather disparate range of topics is rep¬
resented, in fact the contributions are all representative of Clayton’s diverse interests
during his career—late Cenozoic mammals, Caribbean/Neotropical mammalian biogeog¬
raphy, marine mammals.
There is hardly a paper in this volume to which Clayton hasn’t contributed in some way,
in some cases quite directly, and many of these papers would simply not have been possi¬
ble without some kind of aiding and abetting by him. When we began to assemble this vol¬
ume, some of the papers now included herein were in fact being prepared as individual
Contributions to Paleobiology, with Clayton’s encouragement, and in at least one in¬
stance, with his coauthorship. The authors of these contributions were eager to have them
included in this tribute, even though some of the papers would undoubtedly have been
published sooner as individual contributions. As editor, I take this opportunity to express
my gratitude to all of the contributors for their patience in seeing this through the long pro¬
cess from conception to publication. I also want especially to thank David J. Bohaska,
Irina A. Koretsky, and Robert W. Purdy, without whose ideas, encouragement, and assis¬
tance that long process might have been immeasurably longer. Irina A. Koretsky and Mary
A. Parrish both stepped into the breach to help with illustration revisions during the final
stage of production.
Early in the following biography, I mention that Clayton was bom a Hoosier, which I
am confident that readers will interpret in its usual meaning: the nickname for a native or
resident of Indiana. The term sometimes also connotes a hayseed, a bumpkin, an unskilled
mstic. Clayton’s well-known sense of humor sometimes includes self-deprecation, so
whereas he might not reject this meaning outright, it is hardly appropriate for a naturalized
Virginian gentleman farmer. But there is a particular meaning of the term that applies to
Clayton. The several office dictionaries that I consulted gave no etymology for hoosier,
but the Webster s Third New International Dictionary, Unabridged does have this proba¬
ble etymology: “alter, of E dial, hoozer anything large of its kind.” So the word does fit.
Although Clayton is not large in physical stature, he must have carried some weight; the
breadth and depth of the imprints he has left through the course of his professional career
are unmistakable evidence of “a large one of his kind,” not the Hoosier by biogeographical
chance, but the self-made Hoosier.
Robert J. Emry
V
Biography and Bibliography
of Clayton Edward Ray
Ralph E. Eshelman, Robert J. Emry,
Daryl R Domning, and David J. Bohaska
With this volume we pay tribute to our mentor and colleague, fondly the rural life he experienced around his boyhood home
Clayton E. Ray. Scientist, historian, ardent supporter of ama¬ in Indiana, especially the summers spent on his grandparents’
teur and professional paleontologists alike, Clayton is most es¬ farm. He attended high school in Indianapolis, ranked third in
pecially a sound advisor and a valued friend. One of the things his class at graduation, and was awarded a full scholarship to
we miss most as a result of his retirement is the benefit of his Harvard College. He married his high-school sweetheart,
full-time good counsel; we counted on his instinct for the clear¬ Donna, and together they raised four daughters.
headed, no-nonsense approach to problems, minor or major, Clayton enrolled at Harvard College, Cambridge, Massachu¬
and we relied upon him as a source for thoughtful, good advice, setts, in 1951 and received his A.B. degree in geology in 1955.
always strengthened with encouragement. In all of his personal He was elected to Phi Beta Cappa. His advisor as an undergrad¬
interactions, Clayton is invariably perceptive, stimulating,
uate was Alfred S. Romer, one of the most prominent and
helpful, patient, and kind. All of these qualities, combined with
highly respected vertebrate paleontologists of his time. Toward
his quick wit and keen sense of humor, have enriched our long
the end of his senior year, as Clayton met with Romer to dis¬
associations with him. We all hope that some of his attributes
cuss graduate school possibilities, Romer revealed to him, in
have been communicable—that something might have rubbed
confidence, that Brian Patterson would be coming to Harvard
off on all of us to make us more like him.
and that Clayton might want to discuss graduate school possi¬
When we began to organize this tribute, our intention was to
bilities with Patterson when he came to visit. Clayton believes
publish it as a surprise to Clayton, but the process has now
that he was the first “civilian” (as he puts it) to know that Brian
taken so long that the bigger surprise will be if Clayton does
Patterson would be moving from Chicago to Harvard. Clayton
not find out what is afoot. One disadvantage of intending this
followed Romer’s advice, met with Patterson, and decided to
as a surprise is that it inhibited our grilling him for biographical
remain at Harvard for advanced degrees with Patterson as his
information, and thus limited our biography essentially to what
advisor. Clayton does stress, however, that, in the real, practical
is known or could be researched by all of us—the professional,
sense, it was Professor Ernest Williams who served as his grad¬
public aspect. Consequently, this cannot be expected to be an
uate advisor and had the greatest impact on his career. It was
in-depth biography, but a rather superficial review of the gen¬
Williams, for example, who steered Clayton’s interests into
eral trends in Clayton’s professional career.
Quaternary mammals of the Neotropics, and the Caribbean re¬
Clayton’s wife, Donna, supplied us with a few of the basic
gion in particular. Clayton was awarded his A.M. degree in
facts of his pre-university days. Clayton was bom a Hoosier, in
Henry County, Indiana, to Lloyd and Ruth Ray, on 6 February 1958 and his Ph.D. in 1962 from Harvard, both in geology.
In September 1959 Clayton moved to the University of Flor¬
1933 and grew up in central Indiana. Clayton remembers
ida, Gainesville, where he accepted the positions of Interim As¬
sistant Professor in the Department of Biology and Interim As¬
Ralph E. Eshelman and Darryl P. Domning, Research Associates, De¬ sistant Curator at the Florida State Museum (now the Florida
partment of Paleobiology, National Museum of Natural History,
Museum of Natural History ). Upon receiving his doctorate
Smithsonian Institution, Washington, D.C. 20560-0121; Robert J.
from Harvard, he was promoted to Assistant Professor and As¬
Emry and David J. Bohaska, Department of Paleobiology, National
sistant Curator, positions he held until he moved to the Smith¬
Museum of Natural History, Smithsonian Insttitution, Washington,
sonian Institution in 1963.
D.C. 20560-0121.
1
2 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY
Fieldwork and Research
Among his duties in Florida was, of course, fielding ques¬
tions about fossils from visitors. One day he was manning the
Clayton’s earliest publications were on modem mammals,
museum’s front office in the old Seagle Building on Gaines¬
including remains of mammals recovered from archeological
ville’s main street when a high school student from Biloxi,
sites, on Pleistocene mammals, and even on an osteological pe¬
Mississippi, named Daryl Domning walked in, his long-suffer¬
culiarity of the extant gopher tortoise (see bibliography, follow¬
ing parents in tow, seeking publications and tips on where to
ing). In 1960 he cofounded, with two archeologists at the Uni¬
collect fossils in the area. Clayton patiently gave them direc¬
versity of Florida, the first formally constituted museum pro¬
tions to Paynes Prairie, an area on the edge of town where spoil
gram in archaeozoology. Elizabeth Wing, an early associate in
piles from interstate highway construction offered bones of the program, has gone on to become a leading figure in the
Pleistocene critters to the casual collector; afterward, he helped field, and similar formal programs in archaeozoology are now
them identify their finds. This brief encounter was followed by common in other museums and universities. Clayton’s interests
countless more in subsequent years, as Clayton came to play a in archaeozoology have continued throughout his career, even
major role in fostering Domning’s career as a vertebrate pale¬ as his research evolved to focus more on Quaternary mammals
ontologist. The two have since coauthored nearly a dozen pub¬ and eventually on fossil marine mammals. The latter, however,
lications on marine mammals. had been prominent among his interests almost from the start.
Early in the 1960s, the National Museum of Natural History, When Remington Kellogg, the preeminent curator of fossil ma¬
Smithsonian Institution, began planning a major new exhibition rine mammals at the Smithsonian, passed away, Clayton’s at¬
hall devoted to Pleistocene mammals. When the museum de¬ tention shifted progressively to marine mammals, and by the
cided in 1963 to hire a new curator to be in charge of developing late 1970s this assumed priority over his work on Quaternary
these new exhibits, Clayton was the choice. Clayton’s letter ac¬ paleontology. The collections and research data on Quaternary
cepting the new position was typical of him in its thoughtfulness mammals that Clayton had brought together were largely be¬
queathed to others. For example, bats were turned over to Gary
and clarity; he outlined prior obligations that he had to meet and
Morgan, ungulates to Jerry McDonald, Caribbean oryzomyine
asked to make his move to Washington, D.C., between school
rodents to Mike Carleton, and faunal studies to Ralph Eshel-
semesters to lessen the disruption to his children’s education.
man. In concentrating on marine mammal fossils at the Smith¬
On 18 December 1963, Clayton became Associate Curator of
sonian, Clayton continued a tradition that goes back almost as
Later Cenozoic Mammals at the museum, where he remained
far as the Institution itself In fact, the first vertebrate fossil cat¬
until his retirement more than 30 years later.
aloged in the Smithsonian’s collections was a marine mammal
It is probably not generally known that when research scien¬
bone, and work on fossil marine mammals has been conducted
tists (i.e., curators) are hired at the National Museum, they do
by scientists such as F.A. Lucas, Frederick True, Gerrit S.
not immediately become federal civil service employees, but
Miller, Jr., Remington Kellogg, and Frank C. Whitmore, Jr. Be¬
must first complete a conditional, term appointment of at least
cause of this long-standing emphasis, the fossil marine mam¬
one year. If all is well at the end of that term, the appointment is
mal collection at the Smithsonian is the largest in the United
then converted to Career Civil Service status. C.L. (Lew)
States and probably in the world; certainly it is one of the two
Gazin, senior vertebrate paleontologist at the Smithsonian at
most important fossil marine mammal collections in the world.
the time Clayton arrived, wrote the following evaluation of
Clayton’s contributions have added substantially to this already
Clayton’s first year:
strong collection; most important, he was the agent directly re¬
I have found him to be a highly intelligent and industrious worker. During the sponsible for the Smithsonian’s acquiring the Douglas Emlong
rather short time since receiving his doctorate at Harvard he has had published collection of Tertiary marine mammals from the West Coast,
or has completed for publication a surprising number of well-written and care¬ and for nearly three decades he oversaw the collecting of the
fully thought out papers. Much of the time of his first year here had been de¬
huge volume of material from the late Tertiary Lee Creek Mine
voted, as anticipated, to planning with the exhibits staff renovation of Hall 6 for
in North Carolina. In his own research, Clayton has taken a
a new series of exhibits which are to depict the Pleistocene epoch in vertebrate
lively interest in nearly every group of aquatic mammals, but
paleontology. Nevertheless, this has not prevented him from carrying on field
pinnipeds most of all; among these, thanks largely to the gar¬
work in Mexico during the early part of the summer and in initiating at other
times several short trips of profit to the museum. Moreover, in addition to gantuan Lee Creek project, he has focused mainly on the pho-
keeping abreast of a rather considerable amount of examination and report cid seals and on walruses (the subject of his first marine mam¬
work he has managed to squeeze in an appreciable amount of research and give mal paper, way back in 1960).
a few lectures. Dr. Ray is a personable young man who gets along well with his
In the 1970s, Clayton and Charles Handley, Curator of Mam¬
colleagues and makes a good impression on outsiders. It gives me pleasure to
mals in the Division of Mammalogy of the Department of Ver¬
report that we are entirely satisfied with his work.
tebrate Zoology, attempted to form a marine mammal study
So began Clayton’s career of more than three decades as a
center for both fossil and modern marine mammals at the
Smithsonian Curator. Clayton officially retired at the end of
Smithsonian. An important objective was to bring the separate
September 1994 and is now a Curator Emeritus in the Depart¬ collections together both administratively and, especially,
ment of Paleobiology. physically. As anyone who has studied whale skeletons knows.
