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2-1975

Cenozoic Mammals from the Central Great Plains

C. Bertrand Schultz University of Nebraska State Museum

Larry D. Martin University of Kansas Museum of Natural History

R. George Corner Highway Salvage Paleontologist, State of Nebraska Department of Roads

Lloyd G. Tanner University of Nebraska State Museum

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Schultz, C. Bertrand; Martin, Larry D.; Corner, R. George; and Tanner, Lloyd G., "Cenozoic Mammals from the Central Great Plains" (1975). Bulletin of the University of Nebraska State Museum. 106. https://digitalcommons.unl.edu/museumbulletin/106

This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Bulletin of the University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. BULLETIN OF VOLUME 10, NUMBER 1 The University of Nebraska State Museum FEBRUARY, 1975

CENOZOIC MAMMALS FROM THE CENTRAL GREAT PLAINS

Part 1. Middle and Late Cenozoic Tapirs from Nebraska. By C. Bertrand Schultz, Larry D. Martin, and R. George Corner. Part 2. Stratigraphic Occurrences of Teleoceras, with a New Kimballian Species from Nebraska. By Lloyd G. Tanner. Part 3. A New Kimballian Peccary from Nebraska. By C. Bertrand Schultz and Larry D. Martin. Part 4. Bears (Ursidae) from the Late Cenozoic of Nebraska. By C. Bertrand Schultz and Larry D. Martin. Part 5. Scimitar-toothed Cats, Machairodus and Nimra­ vides, from the Pliocene of Kansas and Nebraska. By Larry D. Martin and C. Bertrand Schultz. ACKNOWLEDGMENTS

The editors and authors of these papers gratefully acknowledge the critical comments and valuable suggestions of Craig C. Black, Texas Tech Univer­ sity, and Frank C. Whitmore, Jr., United States Department of the Interior, Geological Survey.

Date submitted: January 21, 1974 Date accepted for publication: April 29, 1974 BULLETIN OF VOLUME 10, NUMBER 1, PART 1 The University of Nebraska State Museum FEBRUARY, 1975

C. Bertrand Schultz Larry D. Martin R. George Corner

Middle and Late Cenozoic Tapirs from Nebraska Frontispiece-Two views of U.N. S.M. Coil. Loc. Ft-40, the locality of the holotypes of ?Tapirus simpsoni, Aphe/ops kimba/lensis, Amebe/odon fricki, Barbourofelis fricki, and new forms of Te/eoceras, Prosthennops, and /ndarctos described in Parts 2, 3, and 4 of the present Bulletin. The upper photograph shows the face of the quarry as it was developed at the time of the discovery of the holotype of ?T. simpsoni in 1947. The "cap rock," which is typical of the upper part of the Kimball Formation in western Nebraska, was being blasted away (when the photograph was taken) in order to expose the fossil-bearing sediments below. The lower photograph shows the cross-bedded channel sands and silts, where so many of the Kimballian fossils are found at Coli. Loc. Ft-40. Some of the "cap rock" is exposed above the channel deposits. Photographs by Schultz. C. Bertrand Schultz Larry D. Martin R. George Corner

CENOZOIC MAMMALS FROM THE CENTRAL GREAT PLAINS

Part 1 Middle and Late Cenozoic Tapirs from Nebraska

BULLETIN OF The University of Nebraska State Museum VOLUME 10, NUMBER 1, PART 1 FEBRUARY, 1975 BULLETIN OF VOLUME 10, NUMBER 1, PART 1 THE UNIVERSITY OF NEBRASKA STATE MUSEUM February, 1975

Pp. 1-21, Tables 1-3 Frontispiece, Figs. 1-13

ABSTRACT

Part 1. Middle and Late Cenozoic Tapirs from Nebraska

C. Bertrand Schultz Larry D. Martin R. George Corner

The distribution and evolution of Late Cenozoic tapirs are discussed and the forms present in Nebraska are reported. Two new species are described from the Ogallala Pliocene of Nebraska, ?Tapirus johnsoni and ?T. simpsoni. Tapirs are known in Nebraska from the Early Oligocene through the Middle Pleistocene. The northern limit of the distribution of the tapirs contracts gradually southward in North America throughout the Tertiary, and even during the Pleistocene interglacials Nebraska must have been near the northern limit of their range.

CONTRIBUTION OF The Department of Geology, College of Arts and Sciences, and the Division of Vertebrate Paleontology of the Museum.

Copyright © 1975 by the Board of Regents of the University of Nebraska Library of Congress Catalog Card Number ISSN 0093-6812 Manufactured in the United States of America Schultz1 Martin2 Corner3

Middle and Late Cenozoic Tapirs from Nebraska

INTRODUCTION contrast with that from South Dakota. This is in part a reflection of the more extensive and more Nebraska has the most complete sequence of productive Early Oligocene (Chadronian) beds late Cenozoic Continental sediments in North present in the latter state. The later Oligocene America, but almost no records of tapirs have beds (Brule Formation) seem to have been de­ been reported from these sediments. Radinsky posited in a less humid environment and tapirs (1963, p. 65) mentions a Nebraska speci men are less abundant in these sediments. Two gen­ housed in the Frick Laboratory of the American era of Oligocene tapirs, C%don and Protapirus, Museum of Natural History, which he identified are known to occur in North America. Only as C%don occidenta/is. He did not give further C%don is known from Nebraska. C%don locality information, identify the elements pres­ belongs to the extinct family Helaletidae which ent, give the stratigraphic horizon, or include the may have given rise to the Tapiridae (Radinsky, occurrence on his map showing the distribution 1963, pp. 94-96.) of Late Eocene and Oligocene tapirs (Radinsky, The earliest tapirid is Protapirus, which may 1963, Fig. 17). Nor was this occurrence plotted have been ancestral to Miotapirus, a genus on Bjork's (1968, Fig. 2) map giving the distribu­ known from the early Miocene of Wyoming, tion of C%don in North America. No additional South Dakota, and Nebraska (Schlaikjer, 1937; published Nebraska records have come to our Macdonald, 1970; Marti n, 1973). Protapirus attention. The scarcity of tapirs in Late Cenozoic robustus has been described from sediments deposits seems to be general in North America, considered to be early Miocene from the John and may be due to lack of extensive subtropical Day Region of Oregon (Sinclair, 1901). The skull forests during this time. of Miotapirus is much more modern than that of The present paper records the distribution of Protapirus. The nasals of Miotapirus are more tapirs in Nebraska from Oligocene to Pleis­ retracted, the diastema is longer, and the palate tocene. The Oligocene record is quite poor in does not extend as far posteriorly. The skull of Miotapirus resembles that of the later genus Tapiravus in having a relatively deep maxilla with a very shallow sulcus on the ascending part of lCurator of Vertebrate Paleontology, University of Nebraska State Museum; and Professor of Geology, Department of the maxillary. No certain records of Hemingfor­ Geology. dian tapirs have been reported, although 2Assistant Curator of Vertebrate Paleontology, University Tapiravus va/idus Marsh may be late Miocene. of Kansas Museum of Natural History; Assistant Professor of Systematics and Ecology, University of Kansas; and Re­ Tapiravus va/idus is known from the type speci­ search Affiliate, Division of Vertebrate Paleontology, Univer­ men from New Jersey and from the Calvert For­ sity of Nebraska State Museum. mation along Chesapeake Bay in Maryland 3Highway Salvage Paleontologist, State of Nebraska De­ partment of Roads; and Division of Vertebrate Paleontology, (Gazin and Collins, 1950, pp. 11-13). The University of Nebraska State Museum. holotype of T. rarus Marsh has never been illus- 2 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM trated. It is from "the Lower Pliocene east of the Referred Specimens.-Partial Palate with Rocky Mountains" (Marsh, 1877, p. 252). Thus ILP3 and partial mandible with P:z-M2, U.N.S.M. the precise geological and geographical prove­ 45113 (Fig. 1B, Tables 1 and 2). nience of the holotype is virtually unknown. Locality of Referred Specimens.-U.N.S.M.5 Tapiravus has also been reported from the Late Coli. Loc. Sx-120, secs. 27-28, T. 34N., R. 57W., Miocene or Early Pliocene of the Bone Valley Sioux County, Nebraska. gravels in Florida, and is recorded in this paper from Nebraska. Tapiravus seems to be restricted Stratigraphic Occurrence.-From a channel to the Late Miocene or Early Pliocene (Valenti­ fill probably equivalent to the Toadstool Channel nian Provincial Age).4 We have referred all the in the Orella Member of the Brule Formation, Clarendon ian and later North American Tertiary White River Group, Oligocene (See Schultz and tapirs discussed in this paper to ?Tapirus. Stout, 1955). Pleistocene records of Tapirus are rare in Ne­ Description.-Upper diastema short; rostrum braska (Fig. 12), suggesting that even during in­ broad; incisive foramina broad but compara­ terglacials this area was near the northern limit tively short; palate narrow anteriorly; incisors of its range (Fig. 13). large, spatulate, and decreasing in size pos­ teriorly; pl small, triangular, with a small antero­ SYSTEMATIC DESCRIPTIONS lingualloph; p2·3 molariform with protocone and hypocone separated, protoloph and metaloph Class: MAMMALIA subparallel, small but distinct parastyle, Order: PERISSODACTYLA metacone elongated posteriorly, trace of a labial cingulum present posteriorly; posterior cing­ Superfamily: TAPIROIDEA ulum well-developed on p2; anterior and pos­ Family HELALETIDAE terior cingula well developed on p3; mandible Genus: C%don Marsh, 1890 relatively short, deep and robust with broad symphysis, roughened and slightly excavated Colodon occidentalis (Leidy), 1868 behind Ia (pathologic?), three small mental foramina present; three incisors present; canine Holotype.-M3, presumably at Academy of abserit; base of crown of p2 indicating small Natural Sciences of Philadelphia (see Radinsky, triangular tooth; p3 subtriangular with promi­ 1968, p. 63, for a detailed account of the occur­ nent paralophid across anterior end and con­ rence of the type of C. occidenta/is). necting with protoconid, metaconid largest Type Locality.-Bad Lands of either South cusp; entoconid large and isolated from Dakota or Nebraska. hypolophid on P3-4 nearly square with re­ duced paralophid and large protoconid; anterior Stratigraphic Occurrence.-? Chadronian. and posterior cingula present on P3-M3 but nearly removed by interstitial wear; lingual and 4Because of the lack of consistency in the use of the term labial cingula only faintly developed on M2-3; mo­ "Pliocene" by various workers, the present writers are con­ lars elongate and increasing in size posteriorly; tinuing with their previous usage in considering the forma­ M3 constricted between trigonid and talonid; tions of the Ogallala Group as Pliocene. However, we realize that the entire Ogallala Group, including the Kimball Forma­ hypolophid of M3 strongly pitched posteriorly tion, may be equal to the uppermost part of the Miocene of toward lingual side; hypoconulid well developed the Mediterranean region of Europe. If the radiometric date on M3. of 2,000,000 ± years ago is considered to be the approximate time for the Quaternary-Neogene ( = Miocene and Pliocene) boundary; and if the radiometric dates are correct, and if five to six million years ago is regarded to be the approximate time for the Miocene-Pliocene boundary in Italy; then the 5Abbreviations used in descriptions: alv., alveolus or al­ Kimballian may be assumed to be equal to very late Miocene. veoli; br., broken; rt., root or roots; U.N.S.M., University of However, in the present Bulletin Parts 1-5, the Valentinian, Nebraska State Museum; C. M., Carnegie Museum; M.C.Z., Clarendonian, Hemphillian, and Kimballian provincial ages Museum of Comparative Zoology, Harvard University; P.U., are all considered to be Pliocene. Princeton University. MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA / 3

Fig. 1-A, ?C%don cingu/atus, U.N.S.M. 45109, partial left ramus (occlusal and labial views), Orella Member, Brule Form­ ation (Oligocene), Sioux County, Nebraska; B, C. occidenta/is, U.N.S.M. 45113, anterior portion of skull (occlusal and labial views, left incisors restored from right side), Toadstool Channel, Orelia Member, Brule Formation (Oligocene), Sioux County, Nebraska; C, U.N.S.M. 45113, partial mandible (occlusal and labial views, P2 alveolus and P4 restored from right side on occlusal view). X 1. 4 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Discussion.-C%don occidentalis appears to shallower and less extensive than in Tapirus; be the most common Oligocene tapir in North mandibular symphysis narrow; cheek-teeth America. Radinsky (1963, p. 63) assigns it a elongate and very brachyodont; 12 at least one­ stratigraphic range of Chadronian through half as large as 11; M2-3 constricted in the middle Whitneyan. The Nebraska record is from an Orel­ with trigonid and talonid separated ,by a deep Ian river channel deposit. Although C%don may groove. have given rise to Protapirus, C. occidenta/is certainly is too specialized to have been the an­ Tapiravus cf. polkensis Olsen, 1960 cestral species. Holotype.-Left p4, Florida Geological Survey, ?Colodon cingulatus Douglass, 1901 V-5941. Paratype.-Lower mandible from same local­ Holotype.-Maxillary with P2(rt.) and P3-M1, C. ity, Florida Geological Survey V-5942. M.722. Type Locality.-Phosphate pit of American Type Locality.-From near Toston, south­ Agricultural Chemical Company, Pierce, Polk western Montana. County, Florida. Stratigraphic Occurrence.-Oligocene (Orel­ Stratigraphic Occurrence.-"Upper Mio­ Ian), Toston Beds. cene-Lower Pliocene" (? Valentinian). Referred Specimen.-Partial left ramus with Amended Diagnosis.-Intermediate in size M2(rt.)-M3, U.N.S.M. 45109, (Fig. 1A). between Tapiravus validus and Tapirus Locality of Referred Specimen.-U.N.S.M. terrestris; cheek teeth brachyodont; molars Coil. Loc. Sx-14, SW. Y4, SW. Y2, NW. Y4 and west elongate; posterior lobes of molars narrow; pro­ edge of SE. %, sec. 7, T. 33N., R. 53W., 11 miles tolophid and metalophid of molars well sepa­ north and about 10 miles west of Crawford, rated. Sioux County, Nebraska. Referred Specimens.-A nearly complete Stratigraphic Occurrence.-Oligocene (Orel­ mandible with complete dentition except for Ia Ian), White River Group, Brule Formation, Orella (alveolus present) U.N.S.M. 45081; partial, Member. juvenile mandible lacking the symphysis with DP2-4 and Ml unerupted but present (exposed by Description.-Ramus massive; M2 relatively dissection on the right ramus), U.N.S.M. 45102; square; M3 lacking prominent cingula, talonid juvenile mandible with DP2-4 and Ml unerupted, only slightly narrower than trigonid, metalophid U.N.S.M. 45103; juvenile mandible with DP2-4 and pitched posteriorly toward lingual side, Ml unerupted, U.N.S.M. 20835; right M3, hypoconulid very prominent. Length of M3, 22.6 U.N.S.M. 20837; left M3, U.N.S.M. 20838 (Figs. 2, mm, and width, 16.6 mm. 4E, 5F, 8, and 9, Tables 2 and 3). Discussion.-?C%don cingu/atus has been Localities of Referred Specimens.-U.N.S.M. previously known only from Chadronian and 45081, 20835, 20837, and 20838 from U.N.S.M. Orellan sediments of Montana (Radinsky, 1963, Coil. Loc. Wt-15A, NW. Y4, sec. 26, T. 1N., R. p. 66), and the Late Eocene of South Dakota 11 W., Webster County, Nebraska; U.N.S.M. (Bjork, 1967). The Nebraska specimen demon­ 45102 and 45103 from U.N.S.M. Coil. Loc. strates that it was sympatric with C%don oc­ Cr-103, from the east side of the Snake River, cidentalis in both Montana and Nebraska in the near center of NE. Y4, sec. 22, T. 32N., R. 30W., Oligocene. Cherry County, Nebraska. Family: TAPIRIDAE Stratigraphic Occurrence.-Pliocene (Valen­ Genus: Tapiravus Marsh, 1877 tinian), Ogallala Group, from sediments equiva­ Diagnosis.-Small tapirs with relatively deep lent to Valentine Formation (U.N.S.M. 45081, maxilla; sulcus on ascending portion of maxilla 20835, 20837, and 20838 from unconsolidated MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA / 5

channel sands eight feet above the Tertiary­ edge of masseteric fossa raised sharply; mas­ Cretaceous unconformity; U.N.S.M. 45102 and seteric fossa not well divided by ridge; margin of 45103 from unconsolidated channel sands 150 ascending ramus directly behind M3 deeply con­ feet below top of cap rock). cave; two grooves present on lingual side of ramus; triangular-shaped lower groove begin­ Description.-!, large and spatulate; 12 spatu­ ning beneath anterior edge of M2 widening pos­ late, two-thirds size of 11; 13 alveolus circular and teriorly and diverging into posterior portion of decidedly smaller than 12 alveolus; IC small, not ramus, upper groove widening anteriorly and as large as paratype of T. polkensis; canine ap­ beginning at about anterior edge of P3 converg­ pressed against 13; IC-P2 diastema shorter than ing into posterior portion of ramus; ascending premolar series; P2 triangular, narrow and more ramus not deep ventrally, ventral border pro­ elongate than in the paratype of T. polkensis; jected medially; deciduous premolars elongate, ectolophid not continuous, metalophid set at 45 lobes narrow; anterior cingula well developed, degree angle to protoconid; P3 subtriangular; posterior cingula crenulated; external reentrant broad paralophids on P3 and P4 and absent in P2; of ectolophid present on DP2 at junction of pro­ molars with trigon ids wider than talonids, tolophid and metalophid. talonids inclined posteriorly, especially in M3; molars narrow waisted; jaw slopi ng continu­ Discussion.-Except for Tapiravus rarus and ously from incisors to beneath P4; mental fora­ the Nebraska records, Tapiravus has been re­ men lying beneath anterior root of P2; mandibu­ ported only from coastal areas. Tapiravus rarus lar foramen ·Iying directly behind M3; anterior Marsh is essentially of unknown geologic age

Fig. 2~Tapiravus cf. polkensis, U.N.S.M. 45081, mandible (labial and occlusal views), from deposits equivalent to lower part of Valentine Formation, Ogallala Group, Webster C04nty,Nebraska. X 3/5. 6 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM and locality, as it was reported by Marsh to be 45110; right M3, U.N.S. M. 45111 (Fig. 5, 0 and G). from the "Lower Pliocene, east of the Rocky Localities.-U.N.S.M. 45110 from U.N.S.M. Mountains" (Marsh, 1877, p. 252). The descrip­ Coil. Loc. Cr-15 (Crookston Bridge Quarry), near tion of it and the genotype of Tapiravus validus W. line sec. 1, T. 32N., R. 30W., Cherry County, (Marsh) are only adequate to show that the ma­ Nebraska; U.N.S.M. 45111 from U.N.S.M. Coil. terials described relate to small tapirs. The inde­ Loc. Cr-103, E. side Snake River, near center NE. terminate nature of these two taxa has also been Y4, sec. 22, T. 32N., R. 30W. pointed out by Sinclair (1901, p. 702) and Simp­ son (1945, p.140). Schlaikjer(1937) reported that Stratigraphic Occurrence.-Pliocene (Valen­ the only generic difference separating Tapiravus tinian), Ogallala Group, Valentine Formation, from the living Tapirus was size, Tapiravus being Crookston Bridge Member. much smaller. Additional material referred to Description.-M1 and M3 brachyodont with Tapiravus cf. validus by Gazin and Collins (1950) protolophid and metalophid deeply separated; from Maryland showed that in addition to its anterior dngula prominent lying against the low smaller size, Tapiravus differed from Tapirus in paralophid; paraconid not distinctly developed; having a relatively deeper maxilla, having a less slight posteriorly directed labial spur off modified lachrymal, and in having a shallower metaconid; posterior cingulum heavy on M1 and more confined sulcus on the lateral face of the light on M3; labial and lingual cingula absent ascending portion of the maxilla. Olsen (1960) from M1 and M3. described a new species, T. po/kensis, from the phosphate beds of Bone Valley, Polk County, Discussion.-These teeth appear to represent Florida, and for the first time figured a portion of a separate lineage of tapirs from that rep­ the mandible of Tapiravus. White (1942) had re­ resented by Tapiravus cf. po/kensis from the ferred a partial ramus with DP2'3 to Tapiravus (?) same horizon. They differ from that species in but did not figure the specimen. The latter having relatively wider and absolutely larger specimen later proved to be the last two perma­ lower molars. nent lower molars and was referred to T. po/ken­ Genus: Tapirus Brisson, 1762 sis by Olsen (1960). The dentition of Tapiravus po/kensis was Diagnosis.-Tapirs of large to medium size shown by Olsen (1960) to be larger than with maxilla shallow; sulcus on ascending pro­ Tapiravus validus but smaller than Tapirus ter­ cess of maxilla deep and extensive; cheek teeth restris. The Nebraska specimen, U.N.S.M. 45081, more quadritubercular than in Tapiravus and is about the same size as the paratype of protolophids not so well separated from Tapiravus po/kensis but larger than the referred metalophids. specimen (M.C.Z. 3808) which includes only M1 Discussion.-The Pliocene tapir material from and M2. Nebraska represents at least in part a single The slope of the ventral margin of the lower lineage. The taxonomic division of so many jaw between the canines and P4 is unbroken, forms which are closely spaced in time presents forming a continuous arc in the specimens from a special problem, and this is especially true for Nebraska. This same character is also seen in a conservative group like the tapirs. Many of the C%don and Pro tapirus , but in the paratype of characters which are used to separate genera Tapiravus po/kensis and other tapirids there is a and species of tapirs occur in the skull and thus noticeable convexity, usually below P2. The are unavailable to us. Considering these prob­ canines of U.N.S.M. 45021 are smaller than in the lems and considering it unwise to create addi­ paratype of T. po/kensis, and the symphysis is tional generic names for our incomplete mate­ narrower. These characters are probably related rial, we have decided to use Tapiravus for the to sexual or individual variation. small very distinct tapir of the Valentinian. For the later forms we use ?Tapirus, indicating a Tapirid, genus and species undetermined genus which may be distinct from but is still Referred Examples.-Left M1, U.N.S.M. close to Tapirus. ;) //?

s:: o r m >z -tJ )) o >-I m ("') ~~=-- 2=:-~' --- - m ~ oZ oN o ~ "'0

~ o"::0 s:: z m tJI ~ en ~ Fig. 3-?Tapirus johnsoni, new species, holotype, U.N.S.M. 1096, mandible (labial and occlusal views), Ash Hollow Formation, Ogallala Group (Pliocene), Cherry County, Nebraska. X 3/5...... 8 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

~ ---/ " -:::-~

~- :: A

B c D

Fig. 4-A, ?Tapirus johnsoni, new species, U.N.S.M. 45104, partial left ramus (labial view), lower or middle part Ash Hollow Formation, Ogallala Group (Pliocene), Sheridan County, Nebraska; U.N.S.M. 45105, partial right ramus (B, labial view), lower part Ash Hollow Formation, Ogallala Group (Pliocene), Garden County, Nebraska; U.N.S.M. 45108, partial right ?OP3 (C, occlusal view), Ash Hollow Formation, Ogallala Group (Pliocene), Cherry County, Nebraska; U.N.S.M. 45107, Right ?p4 (0, occlusal view), Ash Hollow Formation, Ogallala Group (Pliocene), Cherry County, Nebraska; Tapiravus cf. polkensis, U.N.S.M. 45102, mandible (E, labial view), Valentine Formation, Ogallala Group, Cherry County, Nebraska. X 3/5. MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA I 9

Type Locality.-U.N.S.M. Coli. Loc. Cr-104, from the north side of a small canyon on the south side of Steer Creek, near center of SE. 114, sec. 29, T. 32N., R. 30W., 2112 mi. W. and 2 mi. S. of Burge P.O., Cherry County, Nebraska. Stratigraphic Occurrence.-Pliocene (Clar­ endonian), Ogallala Group, Ash Hollow Forma­ tion (in unconsolidated channel sands about 10 feet above cap rock). Referred Specimens.-A partial right ?Dp3, U.N.S.M. 45108; a right ?P4, U.N.S.M. 45107; a partial left ramus with P2-M2 and M3 present but not erupted, U.N.S.M. 45104; a partial left ramus with M1 and M2 beginning to erupt, U.N.S.M. ~.. '.• 'j'i.".,F ~ 45105 (Fig. 4, A-D and Fig. 5, A and C, Table 2). Localities of Referred Specimens.-U.N.S.M. 45108 same as holotype; U.N.S.M. 45107 from the west side of the Snake River in the NE. 114, NE. 114, sec. 28, T. 32N., R. 30W., Cherry County, Ne­ braska; U.N.S.M. 45104 from U.N.S.M. Coli. Loc. Fig. 5-? Tapirusjohnsoni, new species, U.N.S.M. 45104, par­ Sh-7 on E. 112, sec. 25, T. 31 N., R. 41 W., 11 miles tial left ramus (A, occlusal view), lower or middle part Ash south, 1 mile east of Gordon, Sheridan County, Hollow Formation, Ogallala Group (Pliocene), Sheridan Nebraska; U.N.S.M. 45105 from southwest of County, Nebraska, ?Tapirus simpsoni, new species, U.N.S.M. 26132, right P2 (B, labial and occlusal views), U.N.S.M. Coli. Loc. Gd-14, Garden County, Ne­ U.N.S.M. 26134, left Dp1 (E, occlusal view), Band E from braska. Sidney Member, Kimball Formation, Ogallala Group (Pliocene), Frontier County, Nebraska, Tapirus johnsoni, Stratigraphic Occurrences.-U.N.S.M. 45108 U.N.S.M. 45108, partial right ramus (C, occlusal view), Ash same as holotype; U.N.S.M. 45107 Pliocene, Hollow Formation, Ogallala Group (Pliocene), Cherry County, Nebraska, Tapir gen. et sp. indet., U.N.S.M. 45111, Ogallala Group, Ash Hollow Formation (from un­ right M3 (D, occlusal view), U.N.S.M. 45110, left M1 (G, la­ consolidated channel sands which overlie and bial and occlusal views), D and G from Valentine Forma­ cut into the caprock); U.N.S.M. 45104, lower or tion, Ogallala Group (Pliocene), Cherry County, Nebraska, Tapiravus cf.polkensis, U.N.S.M. 45102 (F, unerupted right middle part of the Ash Hollow Formation; M1, labial and occlusal views; G. mandible, occlusal view), U.N.S.M. 45105, middle part of the Ash Hollow Valentine Formation, Ogallala Group, Cherry County, Ne­ Formation. braska. X 3/5. Diagnosis.-Larger than Tapiravus polkensis with more square and hypsodont teeth; teeth smaller and more brachyodont than in Pleis­ ?Tapirus johnsoni6 new species tocene or recent tapirs. Holotype.-Complete mandible with right and Description.-11 large and spatulate; 12 smaller left 11·3, IC, P2-M3; U.N.S.M. 1096; collected by and less procumbent than in Tapiravus; 13 ex­ Frank W. Johnson, PaulO. McGrew, Charles S. tremely reduced as in other Tapirus; dorsal sur­ Osborn, and Grayson E. Meade, 1931 (Fig. 3, face of band antero-dorsal margin of IC showing Table 2). pronounced wear facets; canine incisiform and inclined forwards; P2 short and triangular; ec­ tolophid continuous; metalophid directed pos­ 6Named in honor of Frank Walker Johnson who helped teriorly; P3-4 subtriangular; ectolophid discon­ collect the type and referred material, and who has contrib­ uted much to a better understanding of the Pliocene deposits tinuous on P3-M3; prominent anterior and pos­ of Nebraska. terior cingula on molars decreasing in promi- 10 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM nence anteriorly until almost absent on P3 and slightly smaller and more brachyodont than the absent on P2; external and internal cingula are referred material. This is especially true for absent from the cheek teeth; molars with an­ U.N.S.M. 45104 from Sheridan County, Ne­ terior lophs slightly wider than posterior lophs; braska, which is Hemphillian in age. If beUer ma­ ramus deep; mental foramen beneath anterior terial were available it might be useful to erect a root of P2; almost circular masseteric fossa high new subspecies for the Hemphillian form which on ascending ramus; ascending ramus dished has a thicker and more massive ramus and out below condyles on lingual side and bearing higher crowned teeth. Two isolated upper cheek many pronounced ridges extending from the teeth of ?T. johnsoni are available. However, margin of the angle of the ramus across the de­ their precise placement in the premolar series pression. must be regarded as tentative. We regard U.N.S.M. 45108 as a deciduous tooth, in which Discussion.-There is a general chronocline case it would be either the right Dp3 or DP4. It is for increase in size in the Pliocene tapirs. brachyodont, fairly square in outline and has the ?Tapirus johnsoni is larger than Tapiravus po/­ enamel on the lophs somewhat roughened and kensis from Florida and smaller than the crenulated. The ectoloph between the paracone ?Tapirus from the Kimballian. The holotype of and the metacone dips down to form a shallow ?T. johnsoni, which is Clarendonian in age, is V. There is a well-developed external cingulum

Fig. 6-?Tapirus simpsoni, new species, holotype, U.N.S.M. 45106, partial palate (labial and occlusal views), Kimball Forma­ tion, Ogallala Group (Pliocene), Frontier County, Nebraska. X 3/5. MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA / 11

on the posterior one half of the tooth and a small Referred Specimens from Type Locality. style is present labially between the protoloph -Left DP1, U.N.S.M. 26134; right P2, U.N.S.M. and the metaloph. The paracone is fairly well de­ 26132 (Fig. 5, B and E). veloped. We regard U.N.S.M. 45107 as a perma­ Diagnosis.-Larger than ?Tapirus johnsoni, nent tooth, probably p4 or M1, 20.4 mm. long and and near the size of T. veroensis sellardsi; teeth 20.6 wide. It is also square in outline, brachyo­ more brachyodont and wider in proportion to dont, and has the enamel slightly ridged on the their length than in that species; parastyles large anterior margin of the protoloph. The parastyle on molars, p2 more molariform than in any is large; the ectoloph forms a deep V between Tapirus except T. copei Simpson; diastema rela­ the paracone and the metacone, and just below tively short. this V is a large external style. There is no exter­ nal or internal cingulum. There is a prominent Description.-A tapir about the size of T. internal style between the protocone, hypocone, veroensis; premaxilla with a posterior spur end­ paracone and metacone. These cusps are all bet­ ing just above and anterior to p1; short diastema ter defined in ?T. johnsoni than in living or Pleis­ (15.2 mm.) between 13 and C/; diastema between tocene Tapirus. CI and p1 relatively short (43.0 mm.); infraorbital ?Tapirus johnsoni is also represented by sev­ foramen large; palate relatively elongate and eral fragmentary rami (U.N.S.M. 21158 and narrow (width of maxilla at p4, 126.5 mm.); palate 21159) from U.N.S.M. Coli. Loc. Wt-12 (SE. V4, deeply arched; premaxillaries rounded with a sec. 28, T. 1N., R. 11W., Webster County, Ne­ wide groove situated above and paralleling the braska) from Lower Ash Hollow (Clarendonian) incisive foramina; maxillary deep; zygomatics deposits (Turner, 1972). The cheek teeth are massive; infraorbital foramen large and oval, nearly the same size as the holotype of ?Tapirus leading posteriorly on to large concave shelf johnsoni although the depth of the ramus is formed by maxillary at ventral border of orbit; much shallower in the Webster County speci­ shelf bordered by pits; foramen across from mens. posterior border of anterior root of zygomatic premaxillary portion of palate with paired ?Tapirus simpsonF grooves extending into incisive foramina; inci­ sive foramina large extending from just anterior to C/ to level of posterior border of p1; palatal Holotype.-Palate with 11(alv.), C/, P2_M3; surface of diastema flat; lateral grooves on pal­ U.N.S.M. 45106 (Fig. 6, Table 1). ate deepening posteriorly; lateral wings of palatines heavy, rounded and well separated Type Locality.-U.N.S.M. Coli. Loc. Ft-40, from maxillaries by grooves; palatines terminat­ = "Amebe/odon fricki Quarry," (E. V2, SW. 1/4, SE. ing anteriorly across from anterior border of M1; 1/4, sec. 15, T. 5N., R. 26W.), 8 mi. N. and 5 mi. W. distinct interstitial wear facet on posterior bor­ of Cambridge in Frontier County, Nebraska. The der of tooth alveolus of j1 indicating a tooth history of U.N.S.M. Coli. Lac. Ft-40 and a pre­ about the size of 12; crown of 12 semicircular and liminary list of the Kimballian fauna can be found basin-like; anterior edge of 12 maintaining a in Schultz, Schultz, and Martin (1970). sharp edge; 13 caniniform and heavily worn pos­ Stratigraphic Occurrence.-Upper Pliocene teriorly; CI small and triangular; p1 triangular (Kimball ian), Ogallala Group, Kimball Formation and submolariform with the posterior width gre­ (see Schultz, Schultz, and Martin, 1970, Fig. 8), ater than anteroposterior length; internal cing­ from channel deposits which rest on the upper ulum on anterior portion of p1; all cingula of p1 part of the Ash Hollow Formation. on posterior one-half of tooth, protocone large and relatively far forwards as in T. copei, pro­ toloph fully developed; molars all with large 7Named in honor of George Gaylord Simpson who has parastyles; DP1 lower crowned than p1 with dis­ contributed much to our understanding of fossil tapirs, and who aided the senior writer in so many ways during the prep­ tinct parastyle running into thin low protoloph, aration of the revision of the oreodonts. protoloph attached to prominent metaloph, la- 12 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM bial cingulum developed mostly posteriorly; DP1, Type Locality.-About a mile southwest of 19.5 mm. antero-posterior and 17.5 mm. wide; pl Enon, Missouri, on Moreau Creek, Moniteau lacking parastyle; P2 elongate with faint lingual County. but with no labial cingulum; paraconid of P2 Stratigraphic Occurrence.-Late Pleistocene. connected to protoconid by blade-like paralophid, paraconid and protoconid about . Referred Specimens.-Partial right ramus equal sized but with protoconid higher, with M2 and M3, U.N.S.M. 45114 (Fig. 7A), col­ metaconid small and almost completely joined lected by George Meginnis; left P3, U.N.S.M. to protoconid; ectolopid of P2 at junction of pro­ 39420 (Fig. 7C); left p2, U.N.S.M. 39421 (Fig. 78). toconid and metaconid with a labial groove at this point; hypoconid of P2 large and joined to entoconid by metalophid; p34 with smaller parastyles and p2 with a very small parastyle; la­ bial styles present as slight swellings between the protoloph and the metaloph on molars; mo­ lars and p4 with very slight indication of cristae on ectoloph; protoloph and metaloph well sepa­ rated on P2-M3; ectoloph slightly depressed be­ tween paracone and metacone, but not as deeply as in Tapirus terrestris. Discussion.-?Tapirus simpsoni differs from ?T. johnsoni in being larger and in having P2_M3 much wider than long. The latter proportion along with the molariform pl distinguishes it from T. veroensis which it resembles in having a short diastema. The length of the diastema of ?T. simpsoni falls just within the lower range given for this measurement on T. terrestris by Simpson Fig. 7-Tapirus ct. excelsus, U.N.S.M, 45114, partial right (1945, p. 49, Table 2) and is muc'h less than the ramus (A, occlusal and labial views), U.N,S.M. 39421, left p2 (B, labial and occlusal views), U.N.S.M. 39420, left P3 (C, length he gives for the diastema of T. exce/sus occlusal and labial views), Band C from ?Middle to Late (52 mm.), Simpson (1945, p. 75, Table 11). ?T. Pleistocene, Cherry County, Nebraska. X 3/5. simpsoni most closely resembles T. copei from Port Kennedy Cave which also has a sub­ Localities of Referred Specimens.-U,N.S.M. molariform pl. It differs from T. copei in having 45114 from gravel pit south of Fremont, Dodge the length of the pl about equal to the width and County, Nebraska; U.N.S.M. 39420 and 39421 in being slightly smaller. ?T. simpsoni is distictly from U.N.S.M.Coll. Loc. Cr-10, Pit 3, NW. V4, sec. smaller than T. merriami Frick. The molariform 18, T. 25N., R. 33W., north of Mullen, Cherry p12 separates ?T. simpsoni from Tapirus tapirella County, Nebraska. bairdii, T. indicus and T. roulini. The Dpl of ?T. simpsoni has a constricted anterior portion Stratigraphic Occurrence.-U.N.S.M. 45114 which makes it a much less molariform tooth from Late Pleistocene; U.N.S.M. 39420 and than pl. It is also more brachyodont with a better 39421 from Middle to Late Pleistocene. developed protoloph and metaloph. Description.-Posterior and anterior cingula of M2 and M3 of U.N.S.M. 45114 prominent; Tapirus cf. excelsus Simpson, 1945 paralophids large; hypolophid of M3 small, not pitched far lingually; labial and lingual cingula Holotype.-Skull and jaws and numerous not well developed; U.N.S.M. 39420 (P3) sub­ skeletal elements of a juvenile; A.M.N.H. 39406 triangular; paralophid extremely large; anterior (Simpson, 1945, p. 70). and posterior cingula present; posterior cin- MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA I 13 gulum interrupted; small tubercle developed lin­ gually between metaconid and entoconid; U.N.S.M. 39421 (P2) anterior and posterior cin­ gula well developed; small tubercle present lin­ gually between protocone and hypocone; large A tubercle at antero-Iabial base of paracone.

Tapirus Sp.

Referred Specimen.-Partial left ramus with roots of M2'3, U.N.S.M. 45115 collected by K. A. B Richey. Locality.-U.N.S.M. CoiL Loc. Tm-102, Mallory Sand and Gravel Pit near Thedford, Thomas County, Nebraska. c Stratigraphic Occurrence.-Pleistocene. Description.-Very large tapir with robust ramus and elongate teeth. D Discussion.-This fragmentary specimen rep­ resents a very large species, possibly T. copei. Fig. 8-Lower left P2'S: A, ?Tapirus simpsoni, new species, Some Early Pleistocene fossils have been col­ U.N. S.M. 26132 (Kimballian); B, ?Tapirus johnsoni, new lected from the same pit but the exact horizon species, U.N.S.M. 45104 (Hemphillian); C, ?Tapirus johnsoni, new species, holotype, U.N.S.M. 1096 (Clarendo­ from which the tapir came is uncertain. nian); D, Tapiravus cf. polkensis, U.N.S.M. 45081 (Valenti­ nian). SUMMARY AND CONCLUSIONS C%don in having the nasal notch narrow with The earliest known tapirid, Protapirus, has its its lower (maxillary) border parallel to the nasals. first known occurrence in the early Oligocene of Miotapirus of the early Miocene has a more Europe (Radinsky, 1963, p. 95). The earliest modern aspect to its skull with the dorsal bor­ North American occurrence is from the Pro­ ders of the premaxillary and maxillary smoothly toceras beds (Whitneyan?) of South Dakota inclined towards the nasal notch. In Protapirus (Wortman and Earle, 1893, p. 168). Protapirus the incisive foramina are relatively small and do has non molariform premolars, and resembles not extend posteriorly past the canine. They are C%don among the helaletid tapiroids in having elongated posteriorly in all later tapirs and are retracted nasals. It must have branched off from elongated to the anterior end of p2 in? Tapirus this lineage by the late Eocene because of its simpsoni. The lower canine of Protapirus is mi­ retention of a small canine and nonmolariform nute and it becomes progressively larger upper premolars. The premolars become through Miotapirus and Tapirus while the upper molariform in C%don and the canines are ves­ canine remains relatively small (Fig. 9). The func­ tigial or lost Radinsky (1963, p. 69) suggests that tion of the upper canine is eventually taken over C%don? hancocki from the late Eocene of by a caniniform !3 which shows a progressive Montana would make a good intermediate be­ increase in size. The upper incisors of Protapirus tween He/a/etes and Protapirus. Throughout the and Miotapirus are almost uniform in size. In North American Tertiary there is an increase in Protapirus 13 is already reduced in size and along molarization of the premolars of the tapirs cou­ with 12 maintains about the same relative size in pled with some increase in hypsodonty. Much of most lineages. 11 becomes progressively larger, the molarization (especially in terms of P1) oc­ procumbent, and spatulate (Fig. 9). There also curs during the Oligocene. Protapirus resembles appears to be a tendency for the postero-lingual 14 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

crease in size is well reflected in the P2'S from the Valentinian through the Kimballian (Fig. 8). There is an interesting correspondence be­ A tween the southward retreat of the tapir's range in North America and the southward shrinkage of the subtropical flora as given by Dorf (1957). This same pattern can also be seen in the Ter­ tiary distributions of the giant land tortoises (Fig. 13). B ACKNOWLEDGMENTS The writers wish to thank the following for their aid in making this research project possi­ ble: The late Mr. Alex Keith for the discovery of Kimballian faunal locality (U.N.S.M. Coil. Loc. Ft-40) in 1927; the late Dr. Erwin H. Barbour for his interest in and encouragement of late Ter­ c tiary and Quaternary field work; the late Profes­ sor E. F. Schramm and Mr. Childs Frick for pro­ viding funds for field work; Professors T. M. Stout, A. L. Lugn, Lloyd G. Tanner, and Vincent Dreeszen, Dr. John Boellstorff and Messrs. King Richey, W. D. Frankforter, and Paul Edwards for o assistance and advice. The members of the Uni­ versity of Nebraska State Museum's 1947 field crew, who worked under the direction of W. D. Frankforter and C. Bertrand Schultz, deserve special thanks for their discoveries of various fossils described in Parts 1-5 of this Bulletin. E Members of this crew were Allen Graffham (party leader), James Allen, William H. Berninghausen, Fig. 9-Lower left incisors and canines: A, Tapirus cf. exce/sus, C. M. 159 (Late Pleistocene); B, ?Tapirus Kenneth Harding, J. Knox Jones, Richard johnsoni, U.N.S.M. 1096 (Clarendonian); C, Tapiravus cf. Loomis, Richard Lugn, Neal McClymond, polkensis, U.N.S.M. 45081 (Valentinian); D, Miotapirus Maurice Mendenhall, George Scheffert, Robert harrisonensis, M.C.Z. 2949 (Arikareean), from Schlaikjer, 1937, p. 235, Fig. 1; E, Protapirus va/idus, P.U. 10,900 Truxell, and Olin Webb. The late Florence Erford (Whitneyan), from Hatcher, 1896,2 plates, 4 figures. Sunderland contributed many weeks of volun­ teer time to the field party, and also provided corner of the M3 to become directed further financial aid to help make the 1947 excavations a posteriorly. The width of the talonid of M3 more success. The manuscript was typed by Mrs. Mary nearly approaches that of the trigonid in later Tanner and Miss Rhonda Carlsten. The illustra­ tapirids, thus the M3 appears somewhat squarish tions for Part I of this Bulletin were made by the in shape. Present evidence indicates that North following artists: Mrs. Mary Tanner (Figs. 1, 2, 3, American tapirids have always been browsing 7,8, and 9), Mr. Jerry Tanner (Figs. 4 and 5), Mr. animals, restricted to forest and forest parkland. Robert Miller (Fig. 6), and Mr. Thomas H. They remained about the size of the domestic Swearingen (maps, Figs. 10-13). Special thanks pig (Sus scrofa) from the Oligocene through the are due to Mr. Cecil Williams, owner of the fossil Valentinian. From the Clarendonian through the quarry (Ft-40), and to Mr. Delbert Lewis, former Kimballian there was a rapid increase until they owner, and Mr. Gary Myers, present owner of the reached their present size. This cline for in- fossil locality (Wt-15a). MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA I 15

103 100 97 I

24·!5848 Fig. 10-Map of tapiroid occurrences in Nebraska. 16 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

120 110 100 90 80 70

40

30 30

o 200 400 ~~ Scale in Miles

110 100 90 80

Fig. 11-Map of North American middle and late Tertiary tapiroid occurrences. MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA I 17

120 110 100 90 80 70

40

30 30

200 400 oL=: ' '-=-4 Scale in Miles

110 100 90 80

Fig. 12-Map of Pleistocene tapir occurrences in the United States and Canada (Data from the following: Alexander, 1963; Davis, 1969; Gazin, 1950; Gray and Cramer, 1961; Guilday, 1962; Gut and Ray, 1963; Hibbard and Dalquest, 1966; Holman, 1959; Lance, 1959; Leffler, 1964; Lundelius, 1960, 1967; Neill, 1953; Oesch, 1967; Ray, 1964; Simpson, 1945; Slaughter, 1966; Slaughter et a/., 1962; Strain, 1959; and Schultz, Martin, and Corner, present paper.) 18 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

80 SCJ 60

+ + 60

x Late .. 1- ,; Eocene· .::.". \ ~I '-'

\ / I ;~~ / ir-':...... II. I 21 I -n I { -.--'--1_. I '-\i 40 V""'--:-~-...." Late Pli6'cene Modern

30 30 f--

20 20

o • + (7• '"o

10 10 Q)OO 500 1000 Miles

120 100 eo

Fig_ 13-Northern limit of tapir and giant tortoise distributions respectively: Early Oligocene (1, 1 '); Late Miocene (2, 2'); Early Pleistocene (3, 3'); Middle Pleistocene (4, 4'). The so lid lines give northern limit for the subtropical flora over a simi­ lar time-span (from Dort, 1957). MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA I 19

TABLE 1 C%don, ?Tapirus COMPARATIVE MEASUREMENTS OF SKULLS

C%don occidentalis ?Tapirus simpsoni n.sp. Referred Holotype SKULLS Sx-120 Ft-40 U.N.S.M.45113 U.N. S.M. 45106 pI greatest anteroposterior diameter ...... 8.2 19.7 pI greatest transverse diameter ...... 9.8 20.0 p2 greatest anteroposterior diameter ...... 10.8 20.6 p2 TA', Tp2 ...... 13.6, 14.7 23.8, 25.9 p3 greatest anteroposterior diameter ...... 12.4 21.1 p3 TA, TP ...... 16.2, 16.5 27.8, 27.6 P" greatest anteroposterior diameter ...... 21.8 P"TA, TP ...... 29.5, (28.3)3 M' greatest anteroposterior diameter ...... 22.2 M' TA, TP ...... 27.0, 24.0 M2 greatest anteroposterior diameter ...... 26.3 M2 TA, TP ...... 30.5, 26.5 M3 greatest anteroposterior diamter ...... 26.2 M3 TA, TP ...... 28.7, 23.7 pl_p4 (incl.) ...... 84.9 M'-M3 (incl.) ...... 73.8 PI_M3 (incl.) ...... 159.5 Length of diastema C/_pl ...... 33.9 'greatest transverse diameter of anterior lobe 2greatest transverse diameter of posterior lobe 3( ) = approximate

TABLE 2 C%don, Tapiravus, and ?Tapirus COMPARATIVE MEASUREMENTS OF MANDIBULAR RAMI

C%don occidentalis ?Tapirus johnsoni n.sp. ?Tapiru8 johnsoni n.sp. Referred Tapiravus cf. po/kensis Holotype Referred MANDIBULAR RAMI Sx-120 Wt-15 Cr-104 Sh-7 U.N.S.M.45113 U.N.S.M. 45081 U.N.S.M. 1096 U.N.S.M. 45104 P2 greatest anteroposterior diameter .. 19.1 21.1 21.0 P2 TP' ...... 11.6 15.5 14.9 P3 greatest anteroposterior diameter .. 12.2 17.6 19.7 19.8 P3 TA2, TP ...... 10.1, 11.5 13.0, 14.3 16.2, 19.0 16.4, 18.2 P4 greatest anteroposterior diameter .. 12.6 18.3 19.8 21.0 P4 TA, TP ...... 11.5, 12.2 14.6, 15.1 17.5, 19.0 18.0, 18.1 MI greatest anteroposterior diameter . 15.7 18.1 20.5 22.2 M, TA, TP ...... 12.4, 12.4 14.5, 13.4 17.3, 17.0 18.3, 17.2 M2 greatest anteroposterior diameter . 16.2 19.8 23.4 23.7 M2TA, TP ...... 14.0, 13.3 15.0, 14.2 18.8, 18.2 19.5, 18.3 M3 greatest anteroposterior diameter. 20.6 20.5 23.7 unerupted M3 TA, TP ...... 15.3, 13.1 15.1, 13.8 18.7, 16.9 P2-P4 (incl.) ...... 54.9 60.5 62.4 M,-M3 (incl.) ...... 53.7 59.5 67.8 P2-M3 (incl.) ...... 115.7 65.5 Length of diastema IC-P2 ...... 39.2 41.5 Width across canines ...... 52.3 Depth of jaw below anterior edge M3 . 131.3 'greatest transverse diameter of posterior lobe 2greatest transverse diameter of anterior lobe 20 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

TABLE 3 Tapiravus cf. po/kensis COMPARATIVE MEASUREMENTS OF IMMATURE MANDIBULAR RAMI

Cr-103 Cr-103 Wt-15 IMMATURE MANDIBULAR RAMI U.N.S.M. 45102 U.N.S.M. 45103 U.N.S.M.20835 DP2 greatest anteroposterior diameter ...... 21.2 22.1 DP2 WP' ...... 12.5 12.7 DP3 greatest anteroposterior diameter ...... 18.3 19.0 18.1 DP3 WA2, WP ...... 12.7, 12.6 13.3, 13.1 12.5, 12.0 DP4 greatest anteroposterior diameter ...... 18.9 (20.5) 18.5 DP4 WA, WP ...... 14.1, 13.6 13.5, 12.6 M, greatest anteroposterior diameter ...... 20.5 unerupted unerupted M, WA, WP ...... 14.9, 13.0 DP2-DP4 (incl.) ...... 59.8 60.4 'greatest transverse diameter of posterior lobe 2greatest transverse diameter of anterior lobe MIDDLE AND LATE CENOZOIC TAPIRS FROM NEBRASKA I 21

REFERENCES Miocene Gering Formation of western Nebraska and the early evolution of the North American Cricetidae. Unpub­ Alexander, Herbert L. 1963. The Levi Site: a Paleo-Indian lished Ph.D. thesis, Univ. Kansas. campsite in central Texas. Amer. Antiquity 28(4): 510-528, Neill, Wilfred T. 1953. Notes on the supposed association of Figs. 1-5, 2 tables. artifacts and extinct vertebrates in Flagler County, Florida. Bjork, Philip R. 1967. Latest Eocene vertebrates from north­ Am. Antiquity 19(2): 170-171, 1 figure. western South Dakota. Jou r. Paleo. 41 (1): 227-236, 2 Oesch, R. D. 1967. A preliminary investigation of a Pleis­ plates, 4 tables. tocene vertebrate fauna from Crankshaft Pit, Jefferson --. 1968. New Records of Helaletid Tapiroids from the County, Missouri. Nat. Spel. Soc. Bull. 29(4): 163-185, 13 Oligocene of South Dakota. Papers Michigan Acad. Sci., figures, 8 tables. Arts and Letters 53: 73-78, 2 figures. Olsen, Stanley J. 1960. Age and faunal relationship of Davis, L. C. 1969. The biostratigraphy of Peccary Cave, New­ Tapiravus remains from Florida. Jour. Paleo. 34(1): ton County, Arkansas. Arkansas Acad. Sci. Proc. 43: 164-167, Figs. 1-2. 192-196. Radinsky, Leonard. 1963. Origin and early evolution of North Dort, E. 1957. The Earths changing climates. Weatherwise American Tapiroidea. Bull. Peabody Mus. Nat. Hist. 10: 54-59. 17:1-106, Figs. 1-21, Pis. 1-4, 14 tables. Gazin, C. Lewis. 1950. Annotated list of fossil Mammalia as­ Ray, Clayton E. 1964. Tapirus copei in the Pleistocene of sociated with human remains at Melbourne, Florida. Jour. Florida. Quart. Jour. Florida Acad. Sci. 27(1): 59-66, PI. 1, Washington Acad. Sci. 40(12): 397-404. Tables 1-2. Gazin, C. Lewis, and R. Lee Collins. 1950. Remains of land Schlaikjer, Erich M. 1937. A new tapir from the Lower mammals from the Miocene of the Chesapeake Bay Re­ Miocene of Wyoming. Bull. Mus. Compo Zool. 80(4): gion. Smithsonian Misc. Coil. 116(2): 1-21, Figs. 1-7. 231-251, Figs. 1-5. Gray, Stephen W., and Howard R. Cromer. 1961. A Tapir Schultz, C. Bertrand, Marian R. Schultz, and Larry D. Martin. mandible from a northwest Georgia Cave. Bull. Georgia 1970. A new tribe of saber-toothed cats (Barbourofelini) Acad. Sci. 19(4): 1-8, Figs. 1-4. from the Pliocene of North America. Bull. Univ. Nebr. State Guilday, John E. 1962. Notes on Pleistocene vertebrates from Mus. 9(1): 1-31, Frontispiece, Figs. 1-13, 2 tables. Wythe County, Virginia. Ann. Carnegie Mus. Art. 8:77-86. Schultz, C. Bertrand, and Thompson M. Stout. 1948. Pleis­ Gut, H. James, and Clayton E. Ray. 1963. The Pleistocene tocene mammals and terraces in the Great Plains. Bull. vertebrate fauna of Reddick, Florida. Quart. Jour. Florida Geol. Soc. Amer. 59: 553-558, Figs. 1-4, 1 plate. Acad. Sci. 26(4): 315-328. --. 1955. Classifications of Oligocene sediments in Ne­ Hibbard, Claude W., and Walter W. Dalquest. 1966. Fossils braska, a guide for the stratigraphic collecting of fossil from the Seymour Formation of Knox and Baylor Counties, mammals. Bull. Univ. Nebraska State Mus. 4(2): 17-52, Texas and their bearing on the Late Kansan climate of that Figs. 1-12,2 tables. region. Contr. Mus. Paleo. Univ. Michigan 21(1): 1-66, Figs. Simpson, George Gaylord. 1945. Notes on Pleistocene and 1-8, Pis. 1-5. Recent tapirs. Bull. Amer. Mus. Nat. Hist. 86(2): 33-82, Figs. Holman, J. Alan. 1959. Birds and mammals from the Pleis­ 1-11, Pis. 5-10,18 tables. tocene of Williston, Florida. Bull. Florida State Mus. 5(1): Sinclair, W. J. 1901. The discovery of a new fossil tapir in 1-24, Pis. 1-2, 11 tables. Oregon. Jour. Geol. 9(8): 702-707, 1 figure. Lance, John F. 1959. Faunal remains from the Lehner Mam­ Slaughter, Bob H. 1966. The Moore Pit Local Fauna. Pleis­ moth Site. In the Lehner Mammoth Site. Haury and others. tocene of Texas. Jour. Paleo. 40(1): 78-91, Figs. 1-6. Amer. Antiquity 25(1): 2-42. Slaughter, Bob H., Wilson W. Crook, Jr., R. K. Harris, D. C. Leffler, Sanford R. 1964. Fossil mammals from the Elk River Allen, and Martin Seifert. 1962. The Hill-Shuler local faunas Formation, Cape Blanco, Oregon. Jour. Mammal. 45(1): of the Upper Trinity River, Dallas and Denton Counties, 53-61, Figs. 1-2, 3 tables. Texas. Bureau Econ. Geol. Univ. Texas Rept. Inv. 48: 1-73, Lundelius, Ernest L. 1960. Mylohyus nasutus long-nosed Figs. 1-16, PI. 1, 14 tables. peccary of the Texas Pleistocene. Bull. Texas Mem. Mus. Strain, W. W. 1959. Blancan mammalian fauna from Rio (1): 9-40, Figs. 1-8, Pis. 1-3,6 tables. Grande Valley, Hudspeth County, Texas. Geol. Soc. Amer. --.1967. Late Pleistocene and Holocene faunal history of 70(3): 375-377, Fig. 1. central Texas in Pleistocene Extinctions the Search for a Turner, Mary Ann. 1972. A faunal assemblage from the Lower Cause (ed. by P. A. Martin and H. E. Wright, Jr.): 287-319, Ash Hollow formation (Neogene) of southern Nebraska. Figs. 1-11, 1 table. Unpublished M.Sc. thesis, Univ. Nebraska Dept. Geol. Macdonald, J. R. 1970. Review of the Miocene Wounded White, Theodore E. 1942. Additions to the fauna of the Knee Faunas of southwestern South Dakota. Bull. Los Florida Phosphates. Proc. New England Zool. Club. 21: Angeles Co. Mus. Nat. Hist. (8): 1-82, Figs. 1-32,52 tables. 87-91, 3 plates. Marsh, O. C. 1877. Notice of some new vertebrate fossils. Wortman, J. L., and Charles Earle. 1893. Ancestors of the Amer. Jour. Sci. 14: 249-256. tapir from the Lower Miocene of Dakota. Bull. Amer. Mus. Martin, Larry D. 1973. The mammalian fauna of the Lower Nat. Hist. 5(21): 159-180, Figs. 1-7. BULLETIN OF VOLUME 10, NUMBER 1, PART 2 The University of Nebraska State of Museum FEBRUARY, 1975

Lloyd G. Tanner

Stratigraphic Occurrences of Teleoceras with a New Kimballian Species from Nebraska Frontispiece-View of U.N.S.M. Coil. Loc. Ft-40, Kimball Formation, Ogallala Group (Pliocene), Frontier County, Nebraska. The photograph shows the quarry when the overburden was being removed in 1947. The letters A, B, and C show approxi­ mate locations where important specimens were found. A shows the approximate place where the holotype of Amebelodon fricki Barbour was found in 1927. The holotypes of Te/eoceras schultzi Tanner, new species; Aphelops kimballensis Tanner; and ?Tapirus simpsoni Schultz, Martin, and Corner were also from this area of the quarry. B shows the approxi­ mate spot where the holotype of Barbourofelis fricki Schultz, Schultz, and Martin was discovered. This latter specimen was found in a small local channel, which had been cut into the very fossiliferous cross-bedded channel sands. C is at the approximate site where the holotype of a new species of Prosthennops (Macrogens) was discovered (described in Pt. 3 of this Bulletin). 0 shows Terrace 2B fill , the site of U.N. S.M. Coli. Loc. Ft-41 , the Early Man Lime Creek Site (see Schultz, Lueninghoener, and Frankforter, 1948; Schultz and Frankforter, 1948; Davis, E. Mott, 1962). Photograph by C. Bertrand Schultz. Lloyd G. Tanner

CENOZOIC MAMMALS FROM THE CENTRAL GREAT PLAINS

Part 2 Stratigraphic Occurrences of Teleoceras, with a New Kimballian Species from Nebraska

BULLETIN OF The University of Nebraska State Museum VOLUME 10, NUMBER 1, PART 2 FEBRUARY, 1975 BULLETIN OF VOLUME 10, NUMBER 1, PART 2 THE UNIVERSITY OF NEBRASKA STATE MUSEUM FEBRUARY, 1975

Pp. 23-33, Tables 1-4 Frontispiece, Figs. 1-6

ABSTRACT

Part 2. Stratigraphic Occurrences of Teleoceras, with a New Kimballian Species from Nebraska

Lloyd G. Tanner

Study of Te/eoceras remains in the University of Nebraska State Museum indicates that this specialized, short-limbed rhinoceros inhabited the Central Great Plains from Early through Late Pliocene. Previously thought to have become extinct at the end of the middle Pliocene, this genus is now known from the very latest Pliocene. A new species, Te/eoceras schultz;, is de­ scribed from the Kimball Formation, Ogallala Group, Frontier County, Nebraska.

CONTRIBUTION OF The Department of Geology, College of Arts and Sciences, and the Division of Vertebrate Paleontology of the Museum. Tanner1

Stratigraphic Occurrences of Teleoceras, with a New Kimballian Species from Nebraska2

INTRODUCTION evolution is considered to be a new species, Te/eoceras schultzi (Fig. 1, A). Study of the evidence regarding the short­ Skinner, Skinner, and Gooris (1968, p. 431) limbed rhinoceros, Teleoceras, from Upper Ter­ clarified a misconception of some researchers tiary deposits of the Great Plains Area indicates regarding the genotype material for Teleoceras. that this fossil rhinoceros evolved from a rela­ They pointed out that Teleoceras major Hatcher tively small animal to a large, robust form. The had priority over Teleoceras fossiger (Cope) and smallest, most primitive Teleoceras has been also presented evidence that the two species found in the Valentinian and Lower Ash Hollow were collected from different stratigraphic sediments (Skinner, Skinner, Gooris, 1968, p. levels. Teleoceras major was collected from 431) and has not been adequately studied; there­ Clarendonian, Lower Ash Hollow, deposits and fore only the generic identifications are given. Teleoceras fossiger was recovered from the The largest of the species is recorded from the upper portion of the Middle Ash Hollow (Hem­ Kimballian (see Schultz, Schultz, and Martin, phillian). Both Teleoceras and Aphelops are 1970, pp. 123-128 for discussion of the use of usually found in most Late Cenozoic Quarries in this Provincial Age name). The Teleoceras, as did the Great Plains and adjacent areas, especially the other rhinoceroses, became extinct on the from the Hemphillian. North American continent near the close of the Present understanding of the stratigraphie oc­ Kimballian. This final stage of Teleoceras currences of the Late Cenozoic succession of species for these genera is shown below:

1Coordinator of Systematic Collections, University of Ne­ Kimballian Aphelops kimballensis Teleoceras schultzi n. sp. braska State Museum, Research Associate in Vertebrate U. Hemphillian Aphe/ops longinaris Teleoceras hicksi Paleontology, and Assistant Professor of Geology. Hemphillian Aphelops mutilus Teloceras fossiger Clarendonian Aphelops malacorhinus Teleoceras major 21n the present paper "Pliocene" includes the Valentini an, Valentinian Aphelops sp. Teleoceras sp., Clarendonian, Hemphillian, and Kimballian provincial ages, Skinner 1968 although the writers realize that the Kimballian may be equiv­ alent to the very late Miocene of Europe. See Part 1, p. 2 of The interpretation used in this chart regarding the present Bulletin for further remarks on this subject. the stratigraphic occurrences of Teleoceras is 24 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM based mainly on the fossil evidence observed in composite hind foot from the Edson Quarry (U. the University of Nebraska State Museum fossil K.3817). vertebrate collections. Teleoceras hicksi, from the upper Hemphillian Some objection may be raised as to the use of deposits of Wray County, Colorado, is consid­ Hemphillian for the occurrence of Teleoceras ered to be specifically different from Teleoceras fossiger. Matthew (1932, p. 435) states that fossiger. Teleoceras hicksi appears to be the an­ Teleoceras is not recorded at the Coffee Ranch cestral form for Teleoceras schultzi. Two Ne­ Locality but further indicates that it is found as­ braska specimens, a skull, U.N.S.M. 62095, and sociated with Aphelops in other Pliocene de­ lower jaw, U.N.S.M. 62099 (Fig. 1, B) from posits (Higgins Quarry B). Skinner, Skinner, and U.N.S.M. Coil. Loc. Gd-104, have been referred Gooris (1968, p. 431) concur that Teleoceras fos­ to Teleoceras hicksi. siger came from deposits comparable to middle Hemphillian. SYSTEMATIC DESCRIPTIONS The Edson Quarries of Sherman County, Kan­ sas, have yielded Teleoceras remains which are Class: MAMMALIA comparable to T. fossiger. Hibbard (1934, p. 247). Order: PERISSODACTYLA in his faunal assemblage from the Edson Quar­ ries of Sherman County, Kansas, lists Aphelops Family: RHINOCEROTIDAE cf. A. mutilus Matthew as coming from the de­ Genus: Teleoceras posits which were dated by M. K. Elias as middle Pliocene in age. More recently Frye, Leonard, Te/eoceras schultzi,3 new species and Swineford (1956, p. 30) discussed the Edson Quarry Local Fauna and its relation to the Holotype.-U.N.S.M. 5800, left mandibular stratigraphy of the Ogallala Formation of north­ ramus with I, P3-M3 (Fig. 1A, Table 1). Lacks por­ ern Kansas. They placed the stratigraphic hori­ tion of ascending ramus and condyle. zon of the Edson Local Fauna in the upper part Type Locality.-U.N.S.M. Coil. Loc. Ft-40 = of the Ash Hollow Member of the Ogallala For­ "Amebelodon fricki Quarry," (E. 1/2, SW. 114, SE. mation. They state that the Kimball-Ash Hollow 114, Sec. 15, T. 5N., R. 26W.), 8 mi. N. and 5 mi. W. contact was not present at this locality, but infer­ of Cambridge in Frontier County, Nebraska. red that this contact " ... may be only a few feet above the position of the fauna." Stratigraphic Occurrence.-Upper Pliocene, The faunal lists from the Edson Fossil Locality Ogallala Group, Kimball Formation, Sidney of Sherman County, Kansas, prepared by Hib­ Member from channel deposits which rest on bard (1935, p. 247) and reviewed by Robert W. upper part of Ash Hollow Formation. Wilson for Frye, Leonard, and Swineford (1956, Paratypes.-Skull, partial, lacking frontal­ p. 30), when compared to the faunal list prepared nasal area on both sides and dental series an­ by Matthew and Sti rton (1930a, p. 172) from the terior to upper molar two, U.N.S.M. 62103; skull Coffee Ranch Quarry, "Locality 20," indicated fragment, occipital portion with posterior half of that there are eleven genera and species in left zygomatic arch, U.N.S.M. 62098. common between these two localities. Matthew and Stirton (1930, p. 366) make the Locality of Paratype Specimens.-U.N.S.M. comparison of the Hemphill County, Texas, 62103 and U.N.S.M. 62098 are from U.N.S.M. "Locality 20" local fauna with the Snake Creek Coil. Loc. Ft-40 (E. 1/2, SW. 114, SE. 114, Sec. 15, T. assemblage, and demonstrate that there is a dis­ 5N., R. 26W.), 8 mi. N. and 5 mi. W. of Cambridge tinct similarity in the vertebrate fauna from these in Frontier County, Nebraska. two localities. The composite hind foot illustrated by 3Named in honor of Professor C. Bertrand Schultz, Curator Matthew (1932, p. 429) from the Coffee Ranch of Vertebrate Paleontology, University of Nebraska State Locality 20 is nearly identical in size to a Museum, and Director of the Museum, 1941-1973. STRATIGRAPHIC OCCURRENCES OF TELEOCERAS, WITH A NEW KIMSALLIAN SPECIES / 25

~~. ...~~

-- c---~----. - ~ ----.------

Fig. 1-Mandibular rami (occlusal and labial views): A, Teleoceras schultzi, new species, holotype, U.N.S.M. 5800, Kimball Formation, Ogallala Group (Pliocene), Frontier County,Nebraska; S, Teleoceras hicksi, referred, U.N.S.M. 62099, left mandibular ramus, upper part of Ash Hollow Formation, Ogallala Group (Pliocene), Garden County, Nebraska. X 1/4. 26 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

TABLE 1 Teleoceras COMPARATIVE MEASUREMENTS' OF MANDIBULAR RAMI

T. schultzi T. hicksi n. sp. Holotype MANDIBULAR RAMI Holotype Denver Musem Nat. U.N.S.M. 5800 History 304 Total length of lower dental series ...... 238.0' 258.0' Total length of lower molar series ...... 165.0' 173.0' Total length of diastema in front of P3 ...... 55.0' 56.0' Depth of jaw below M2 ...... 145.0 98.0 Depth of jaw below P3 ...... 128.0 78.0 P3 greatest anteroposterior diameter ...... 35.8 P3 greatest transverse diameter ...... 23.0 P4 greatest anteroposterior diameter ...... 44.5 P4 greatest transverse diameter ...... 37.0 M, greatest anteroposterior diameter ...... 48.7 M, greatest transverse diameter ...... 38.4 M2 greatest anteroposterior diameter ...... 60.0 M2 greatest transverse diameter ...... 37.0 M3 greatest anteroposterior diameter ...... 59.0 M3 greatest transverse diameter ...... 40.0 'The measurements are taken to the nearest millimeter except on dentition where they are measured to the nearest one-tenth of a millimeter. 'These diameters vary considerably with maturity, as interstitial wear from crowding is considerable.

Diagnosis.-A species of Teleoceras slightly teristic, as crowding and interstitial wear vary larger than T. hicksi Cook; upper dentition more with the stage of maturity. The tooth pattern is complicated; skull more brachycephalic and relatively simple and the cingula, as in all Late having a relatively wider basio-occipital region. Pliocene Teleoceras, are very weak to absent. The holotype ramus, U.N.S.M. 5800, is more The lateral incisor is worn through attrition with robust than Teleoceras hicksi. The length of the the opposing upper incisor (see Fig. 3). The most lower dental series is not a diagnostic charac- diagnostic characteristics for the species are the

TABLE 2 Teleoceras COMPARATIVE MEASUREMENTS OF SELECTED POST-CRANIAL MATERIAL

.... IX) ..... 0 0 N IX) IX) Q;IX) 0> 0> 0 ;; ~O '-!:IN0 '-!:IN0 .N '-!:IN0 '- !:IN0 ~N N UJ UJ UJ -to as -to "5~ "5~ as => . :; => . as ci tO as " ~~ .~; " ..c::::2: :0 :0 '8::2: 0::2: " '8::2: .0 '8::2: :;::; ~::!E :;::; POST-CRANIAL MATERIAL ",(J). € ;;::: "'00 ~ "'00 ~ '"o . ~ ~cri "'00 "'00 t--oo ""::i ""::i ~Z ""::i ""::i ....:~ :i =i =i ....:::j =i ::::> ::::> ::::>

Length (maximum) ...... 270 295 245 268 203 250 247 Length (articular) ...... 245 267 228 255 224 225 218 Transverse diameter, proximal end (maximum) ...... 104 115 90 133 47 126 118 Transverse diameter, center of shaft ...... 60 65 41 56 30 56 54 Transverse diameter, minimum ...... 55 61 37 56 30 56 54 Transverse diameter, distal end (maximum) ...... 122 105 86 95 58 97 91 Anteroposterior diameter, proximal end ...... 55 72 53 138 59 128 117 Anteroposterior diameter, center ...... 36 40 32 67 22 52 55 Anteroposterior diameter, distal end ...... 59 54 55 72 69 70 69 STRATIGRAPHIC OCCURRENCES OF TELEOCERAS, WITH A NEW KIMBALL/AN SPECIES 1 27

B

G

Fig. 2-Cheek teeth (occlusal views): A, Teleoceras schultzi, new species U.N.S.M. 62092, right partial maxillary, P4_M3, Kim­ ball Formation, Ogallala Group (Pliocene), Cheyenne County, Nebraska; B, T. schultzi, referred, U.N.S.M. 61220, right P4_M', Kimball Formation, Cheyenne County, Nebraska; C, T. schultzi, referred, U.N.S.M. 5817, right P4_M2, Kimball Forma­ tion, Frontier County, Nebraska; D, T. schultzi, referred, U. N.S.M. 5820, left M3, Kimball Formation, Frontier County, Ne­ braska; E, T. schultzi, referred, U.N.S.M. 62093, right M3, Kimball Formation, Garden County, Nebraska; F, T. major, referred, U.N.S.M. 62097, right P3_M3, basal part of Ash Hollow Formation, Brown County, Nebraska; G. T. hicksi, refer­ red, U.N.S.M. 62095, right maxillary with P2_M3, Ash Hollow Formation, Garden County, Nebraska. X 1/2. 28 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

c

F G H

Fig. 3-Lateral incisors: A-C, Te/eoceras schultzi, referred, upper lateral incisors, U.N.S.M. 62080, 5477, 62083, Kimball mation, Frontier County, Nebraska; D-E, T. fossiger, referred, upper lateral incisors, U.N.S.M. 62081, 62082, middle part of Ash Hollow Formation, Banner County, Nebraska; F, Aphe/ops kimballensis, referred, lower lateral incisor, U.N.S.M. 5809, Kimball Formation, Frontier County, Nebraska; G-H, T. schultzi, referred, lower lateral incisors, U.N.S.M. 62079, 62078, Kimball Formation, Frontier County, Nebraska; I, Aphe/ops kimballensis, referred, lower lateral incisor, U.N.S.M. 62078, Kimball Formation, Frontier County, Nebraska. X 1/2. STRATIGRAPHIC OCCURRENCES OF TELEOCERAS, WITH A NEW KIMBALLIAN SPECIES / 29

Fig. 4-Radii (anterior views): A, Teleoceras schultzi, new species, U.N.S.M. 62087, Kimball Formation, Ogallala Group, (Pliocene), Frontier County, Nebraska; B, T. schultzi, new species, U.N.S.M. 62088, Kimball Formation, Frontier County, Nebraska; C, T. fossiger, referred, U.N.S.M. 62084, middle part of Ash Hollow Formation, Ogallala Group, Ban­ ner County, Nebraska. X 1/3. large, complicated upper molars shown in Fig. 2, Study and comparison of Teleoceras post-cra­ A. This partial maxillary from the uppermost nial elements (Figs. 4, 5, 6) which have been col­ Pliocene depOSits shows the fossette in the area lected from the Ogallala deposits indicates that of the protocone, and the upper dentition is rela­ the bones of T. schultzi are large and become tively more hypsodont. relatively more massive. Table 2, see p. 26, is Teleoceras schultzi is a more specialized form presented as evidence. and is larger than T. hicksi in most dimensions. The radii, U.N.S.M. 62087 and 62088 (Fig. The occipital portion of a skull with one zygoma­ 4A-B), from the Kimballian deposits of U.N.S.M. tic arch attached of Teleoceras schultzi, Coil. Loc. Ft-40, are distinctly more robust and U.N.S.M. 62098, provides some information re­ massive when compared to the radius U.N.S.M. garding the width of the skull of this species. The 62084 (Fig. 4C) which was removed from de­ greatest transverse dimension at the zygomatic posits herein considered to be Hemphillian or arches would be approximately 396 mm. In com­ near equivalent at U.N.S.M. Coil. Loc. Bn-10, parison, the like dimension of the holotype skull Banner County, Nebraska. Like evidence can be of T. hicksi (Cook, 1927) is 334 mm. (see Table 4). observed regarding the larger size range of the The type skull of T. fossiger measures 339 mm. tibiae of Teleoceras schultzi from the Kimballian across the zygomatic region. Further measure­ (see Fig. 6). U.N.S.M. 62102 and the articulated ments taken on the OCCipital region of U.N.S.M. tibia-fibula U.N.S.M. 62090 are from U.N.S.M. 62098 and U.N.S.M. 62103 indicate that this Coil. Loc. Ft-40. The selected tibia (listed in species is more brachycephalic than T. hicksi. Kent, 1963, p. 108), U.N.S.M. 61331 from 30 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

U.N.S.M. Coil. Loc. Cn-105 (Cheyenne County) is that these forms became extinct during the mid­ slightly smaller in most dimensions, even though dle portion of the Pliocene. the deposits yielding this specimen are also con­ Further study of this group is anticipated in an sidered to be uppermost Pliocene (Kimballian) in effort to clarify the differences in relative sizes of age. Other skeletal elements of Teleoceras from certain of the Kimballian forms. Perhaps a dwarf this locality, when compared to those from tribe of Teleocerines may have been living at that U.N.S.M. Ft-40 (Frontier County) are rather time. robust, but seem to indicate the possibility that a smaller tribe of Teleoceras may have lived at the ACKNOWLEDGMENTS same time as T. schultzi. Perhaps the sediments of the Cheyenne County locality are slightly Special credit should be given to Professors C. younger than the deposits in Frontier County. Bertrand Schultz and T. M. Stout of the Univer­ sity of Nebraska State Museum for their advice. SUMMARY Professor Larry Martin of the University of Kan­ sas and Mr. George Corner, University of Ne­ The fossil remains of Teleoceras are now braska State Museum, also contributed aid to the known to occur in deposits from Valentini an preparation of this report. All figures were drawn through Kimballian or throughout the Pliocene by Mary Tanner. Rhonda Carlsten typed the Epoch. Previous published records proposed manuscript.

A c

Fig. 5-Metacarpals (posterior and proximal views): A, Te/eoceras major, referred, U.N.S.M. 62074, lower part of Ash Hol­ low Formation, Ogallala Group (Pliocene), Cherry County, Nebraska; B, T. fossiger, referred, U.N.S.M. 62077, middle part of Ash Hollow Formation, Ogallala Group, Banner County, Nebraska; C, T. schultzi, new species, U.N. S.M. 62076, Kimball Formation, Ogallala Group, Frontier County, Nebraska. X 1/2. STRATIGRAPHIC OCCURRENCES OF TELEOCERAS, WITH A NEW KIMBALLIAN SPECIES / 31

Fig. 6-A, Teleoceras schultz;, new species, U.N.S.M. 62090, tibia and fibula articulated, Kimball Formation, Ogallala Group (Pliocene), Frontier County, Nebraska; B, Teleoceras sp., U.N.S.M. 62101, tibia, Kimball Formation, Ogallala Group, Mor­ rill County, Nebraska; C, T. schultz;, referred, U.N.S.M. 62102, tibia, Kimball Formation, Frontier County, Nebraska: D, Teleoceras sp., U.N.S.M. 61331, tibia, Kimball Formatio n, Cheyenne County, Nebraska. X 1/3.

TABLE 3 Teleoceras COMPARATIVE MEASUREMENTS OF METACARPALS

T. schultzi T. fossiger ref. T. major ref. METACARPALS U.N.S.M. 62076 U.N.S.M. 62077 U.N.S.M. 62074 Metacarpal III Metacarpal III Metacarpal III Length (maximum) ...... " ...... , ...... 126.5 134.3 115.5 Length (articular) ...... 116.5 122.5 114.5 Transverse diameter, proximal end (maximum) ...... 76 64.2 67 Transverse diameter, center of shaft ...... 62 53.5 46.2 Transverse diameter, minimum ...... 62 53.5 44.8 Transverse diameter, distal end (maximum) ...... 72 64 56 Anteroposterior diameter, proximal end ...... 51 64.2 44.2 Anteroposterior diameter, center ...... 26 23.5 20.8 Anteroposterior diameter, keel (maximum) ...... (45)1 36:7 37.4 Distance from minimum diameter to distal end (maximum) ...... " .. . 64.3 78.2 54

1( ) = approximate 32 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

TABLE 4 Tefeoceras COMPARATIVE MEASUREMENTS OF PARTIAL SKULLS

T. schu/tzi T. schu/tzi T. hicksi SKULLS T. schu/tzi ref. Paratype Paratype Denver Mus. U.N.S.M.62092 U.N.S.M.62103 U.N.S.M.62098 Nat. Hist. 304 Transverse diameter of palate at M2, labial edges ...... 230 Transverse diameter at zygomatic arch ...... 387 (396)' 334 Transverse diameter at mastoids ...... 263 294 M2 anteroposterior diameter ...... 55.7 M2 transverse diameter ...... 72.5 M3 anteroposterior diameter ...... (46.5)2 53.2 M3 transverse diameter ...... (40.0) 64.5 p4 anteroposterior diameter ...... 52.5 p4 transverse diameter ...... 70.8 M' Anteroposterior diameter...... 63.2 M' transverse diameter ...... (79) M2 anteroposterior diameter...... 70.2 M2 transverse diameter...... (71.2) '( ) = approximate 'Erupting STRATIGRAPHIC OCCURRENCES OF TELEOCERAS, WITH A NEW KIMBALLIAN SPECIES / 33

REFERENCES Osborn, Henry Fairfield. 1936. Proboscidea: A monograph of the discovery, evolution, migration and extinction of the Cope, E. D., and W. D. Matthew. 1915. Hitherto unpublished mastodonts and elephants of the World. Vol. plates of Tertiary mammalia and Permian Vertebrata. Mon. 1-Moeritheroidea, Deinotherioidea, Mastodontoidea. Amer. Mus. Nat. Hist. Ser. 2: Pis. 125--144b. New York, Amer. Mus. Nat. Hist.: i-xl, 1-802, Figs. 1-680. Davis, E. Mott. 1962. Archeology of the Lime Creek Site in Schultz, C. Bertrand, and W. D. Frankforter. 1948. Prelimi­ southwestern Nebraska. Special Publ. Univ. Nebraska nary report on the Lime Creek Sites: New evidence of Early State Mus. (3): 1-106, Figs. 1-38. Man in southwestern Nebraska. Bull. Univ. Nebraska State Davis, E. Mott, and C. Bertrand Schultz. 1952. The archeolog­ Mus. 3(4), Pt. 2: 43-62, Figs. 1-13. ical and paleontological salvage program at the Medicine Schultz, C. Bertand, Gilbert C. Lueninghoener, and W. D. Creek Reservoir, Frontier County, Nebraska. Science 115 Frankforter. 1948. Preliminary geomorphological studies (2985): 288-290. of the Lime Creek area. Ibid. 3(4), Pt. 1: 31-42, Figs. 1-6. Eisele, C. Robert. 1964. Salvaging fossils in Nebraska. Univ. Schultz, C. Bertrand, and Thompson M. Stout. 1939. Practi­ Nebraska State Mus. Museum Notes (24): 1-8, Figs. 1-6. cal application of paleoecology in the study of Cenozoic Frye, John C., A. Byron Leonard, and Ada Swineford. 1956. mammals. (Abstract) Bull. Geol. Soc. Amer. 50(12): 1967. Stratigraphy of the Ogallala formation (Neogene) of north­ ---.1948. Pleistocene mammals and terraces in the Great ern Kansas. Bull. Kansas Geol. Surv. (118): 1-92, Figs. 1-5, Plains. Bull. Geol. Soc. Amer. 59(6): 553-558, Figs. 1-4, PI. Pis. 1-9. 1,3 tables. Galbreath, E. C. 1953. A contribution to the Tertiary Geology --. 1961. Field conference on the Tertiary and Pleis­ and Paleontology of northeastern Colorado. Kansas Univ. tocene of western Nebraska (with contributions by Charles Paleo. Contr. (4): 1-20, Figs. 1-26, Pis. ~-2. H. Falkenbach and Lloyd G. Tanner and field conference Hesse, Curtis J. 1935. A vertebrate fauna from the type local­ assistance from Harold J. Cook and A. L. Lugn) (Guide ity of the Ogallala formation. Bull. Univ. Kansas 36(8) and Book for the Ninth Field Conference of the Society of Ver­ Univ. Kansas Sci. Bull. 22(5): 79--101,8 plates. tebrate Paleontology). Special Publ. Univ. Nebraska State ---.1940. A Pliocene vertebrate fauna from Higgins, lips­ Mus. (2): 1-55, Figs. 1-47,2 charts, 1 map. comb County, Texas. Publ. Univ. Texas (3945): 671-698. Schultz, C. Bertrand, Marian R. Schultz, and Larry D. Martin. Hibbard, Claude W. 1934. Two new genera of felidae from the 1970. A new tribe of saber-toothed cats (Barbourofelini) middle Pliocene of Kansas. Trans. Kansas Acad. of Sci. 37: from the Pliocene of North America. Bull. Univ. Nebraska 239--255, Pis. 4-6, Tables 1-2. State Mus. 9(1): 1-31, Tables 1-2. Kent, Douglas C. 1963. A late Pliocene faunal assemblage Simpson, George Gaylord. 1945. The principles of classifica­ from Cheyenne County, Nebraska. Unpublished M.Sc. tion and a classification of mammals. Bull. Amer. Mus. Nat. thesis, Univ. Nebraska Dept. Geol.: 1-143, Figs. 1-41, and Hist. 85: 1-350. Appendices A, B, C. Skinner, Morris F., Shirley M. Skinner, and Raymond J. Lane, H. H. 1927. A new Rhinoceros from Kansas. Bull. Univ. Gooris. 1969. Cenozoic rocks and faunas of Turtle Butte, Kansas Sci. 17(2): 297-311, Pis. 22-25. South-central South Dakota. Bull. Amer. Mus. Nat. Hist. Lugn, Alvin L. 1939. Classification of the Tertiary system in 138 art. 7: 379-436, Figs. 1-16, Pis. 20-25, Tables 1-7. Nebraska. Geol. Soc. Amer. (50): 1245--1276, 1 plate. Stout, T. M., H. M. DeGraw, L. G. Tanner, K. O. Stanley, W. J. Matthew, W. D. 1932. A review of the rhinoceroses with a Wayne, and J. B. Swinehart. 1971. Guidebook to the Late description of the Aphelops material from the Pliocene of Pliocene and Early Pleistocene of Nebraska. Conservation Texas. Bull. Univ. California Dept. Geol. Sci. 20(12): and Survey Division (Nebraska Geol. Survey) Univ. 411-480, Figs. 1-12, Pis. 61-69. Nebraska-Lincoln: i-v, 1-109, Figs. 1-11, Pis. 1-2, table. Matthew, W. D. and R. A. Stirton. 1930a. Osteology and af­ Tanner, Lloyd G. 1960. Rhinoceroses of the past. Univ. Ne­ finities of Boraphagus. Bull. Univ. California Dept. Geol. braska State Mus. Museum Notes (11): 1-4,6 illustrations. Sci. 19(7): 171-216,2 figures, Pis. 21-34. ---.1967. A new Species of rhinoceros. Aphelops kimba/­ --. 1930b. Equidae from the Pliocene of Texas. Bull. lensis from the latest Pliocene of Nebraska. Bull. Univ. Univ. Calfornia Dept. Geol. Sci. 19(17): 349--396, Pis. 45--58. Nebraska State Mus. 6(1): 1-16, Fig. 1, Pis. 1-5, Tables 1-2. Osborn, Henry Fairfield. 1904. New Miocene rhinoceroses with revisions of known species. Bull. Amer. Mus. Nat. Hist. 20(27): 307-367, Figs. 1-21. BULLETIN OF VOLUME 10, NUMBER 1, PART 3 The University of Nebraska State Museum FEBRUARY, 1975

C. Bertrand Schultz Larry D. Martin

A New Kimballian Peccary from Nebraska Frontispiece -Prosthennops (Macrogens) graffhami, new species, holotype, U.N.S.M. 76051, skull, from U.N.S.M. Coil. Loc. Ft-40, Kimball Formation, Ogallala Group (Pliocene), Frontier County, Nebraska. X 1/2. C. Bertrand Schultz Larry D. Martin

CENOZOIC MAMMALS FROM THE CENTRAL GREAT PLAINS

Part 3 ANew Kimballian Peccary from Nebraska

BULLETIN OF The University of Nebraska State Museum VOLUME 10, NUMBER 1, PART 3 FEBRUARY, 1975 BULLETIN OF VOLUME 10, NUMBER 1, PART 3 THE UNIVERSITY OF NEBRASKA STATE MUSEUM FEBRUARY, 1975

Pp. 35-46, Tables 1-3 Frontispiece, Figs. 1-5

ABSTRACT

Part 3. ANew Kimballian Peccary from Nebraska

c. Bertrand Schultz Larry D. Martin

A new species of Pliocene peccary, Prosthennops (Macrogens) graffhami; is described from the Kimball Formation, Ogallala Group, Frontier County, Nebraska. This new species is the latest in geologic age and most advanced in the genus.

CONTRIBUTION OF The Department of Geology, College of Arts and Sciences, and the Divison of Vertebrate Paleontology of the Museum. Schultz1 Martin2

ANew Kimballian Peccary from Nebraska

INTRODUCTION Peccaries are relatively common in Ogallala faunas, although few good skulls have been de­ The remains of fossil vertebrates are gener­ scribed. This is especially unfortunate because ally rare in the upper part of the Ogallala group, the teeth tend to be conservative while the skulls and some have maintained that Hemphillian are often highly mod ified (Colbert, 1938, p. 263). faunas such as Coffee Ranch and Smith County, At least two distinct lineages of peccaries occur Kansas, represent the latest Ogallala faunas. in the Ogallala Pliocene of the Central Great Later faunas than are typically considered as Plains. One of these, which includes Hemphillian, containing more advanced forms, Prosthennops (Macrogens), has large, triangular occur in the Kimball Formation and deposits of malar tuberosities, while the other lineage, equivalent age. Known faunas which may be Prosthennops (Prosthennops), lacks this fea­ close to Kimballian are rare outside of Nebraska, ture. The new peccary described in this paper is although part of the Rattlesnake fauna in the latest and most advanced known species of Oregon, the Axtel fauna in Texas, and the Prosthennops (Macrogens). Guymon fauna in Oklahoma contain forms only SYSTEMATIC DESCRIPTIONS somewhat more primitive than forms considered Class: MAMMALIA to be early Kimballian in age. The fauna of the Order: ARTIODACTYLA lower part of the Kimball Formation in Nebraska Family: TAYASSUIDAE is rapidly becoming the best known Pliocene Subfamily: Tayassuinae fauna from North America, if the Blancan faunas are considered to be Early Pleistocene in age. Prosthennops Gidley, 1904 Much Kimballian fauna has been recovered from U.N.S.M. Coil. Loc. Ft-40 (= Amebe/odon fricki Prosthennops (Macrogens) Matthew, 1924 Locality). (See Schultz, Schultz, and Martin, 1970, for discussion of the faunal and strati­ Prosthennops (Macrogens) graffhamj3 graphic position.) new species

1Curator of Vertebrate Paleontology, University of Ne­ Holotype.-Skull (essentially uncrushed and braska State Museum; and Professor of Geology, Depart­ ment of Geology. complete except for the tip of the snout with the 2Assistant Curator of Vertebrate Paleontology, University of Kansas Museum of Natural History; Assistant Professor of Systematics and Ecology, University of Kansas; and Re­ 3Named in honor of Mr. Allen Graffham, field party leader at search Affiliate, Division of Vertebrate Paleontology, Univer­ U.N.S.M. Coil. Loc. Ft-40 in 1947, when the holotype of this sity of Nebraska State Museum. species was discovered. 36 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM incisors) with C, P2_M3, U.N.S.M. 76051 (Figs. supraorbital foramina on each side, large lateral 1-2). foramen and small medial foramen connected by a small branch of supraorbital canal; infraorbital Referred Specimens from Type foramen large, oval, and above pH; buccinator Locality.-Partial left maxillary with Dp2_M1, fossa shallow, extending back to M1, canine­ U.N.S.M. 76055 (Fig. 2C); partial left maxillary premolar diastema short; canine buttresses with Dp3_M1, U.N.S.M. 76056; fragment of left strongly developed; skull domed as in My/ohyus; maxillary with p3_p4 (br.), U.N.S.M. 76057; left orbits posteriorily and dorsally situated with an­ DP4, U.N.S.M. 76070; partial mandible with terior margins above middle of glenoid fossa; l1(br.), 12-3, IC(alv.), P2-M3, U.N.S.M. 76052 (Fig. 3, lacrymal process small and distinct; large lac­ Table 2); anterior portion of mandible with 11-/C rymal.foramen near anteroventral border of eye; (Fig. 3); fragment of anterior portion of mandible prominent groove on posterointernal wall of with 11-13 alveoli, IC(br.); partial left ramus with orbit; posterior opening of infraorbital foramen P2, P3(alv.), P4, M1-3(br.), U.N.S.M. 76059; right P4, sm~II;. cranium short; sagittal crest prominent; U.N.S.M. 76068; 4 upper canines, U.N.S.M. occIpital region less inclined than in P/atygonus; 76062, 76072, 76073, 76083; 5 lower canines, ventral borders of occipital condyle, glenoid U.N.S.M. 76060, 76061, 76063, 76067, 76084; fossa and maxillary bone at about the same right hind limb, essentially complete from femur level; malar tuberosities massive, triangular and to distal phalanges, U.N.S.M. 76071 (Fig 4); right wing-like; origin of M. masseter /ateralis profun­ femur (lacking proximal end), U.N.S.M. 76065; dus large, short and oval; origins of M. depressor proximal end of left femur, U.N.S.M. 76064. rostri and M. dilator naris /atera/es long and Referred specimen from U.N.S.M. Coil. Loc. deep; orbital processes of the zygoma and fron­ Gd-10 (Oshkosh Loc. A, Q. 1, Pit 1).-Partial tals relatively long; height of zygomatic process mandible with 11-/C, P2-M3, U.N.S.M. 76054. much less posterior to postorbital process than Type Locality.-U.N.S.M. Coil. Loc. Ft-40 = anterior; prominent postzygomatic crest extend­ "Amebe/odon fricki Quarry," (E. 1/2, SW. 114, SE. ing from zygomatic arch above glenoid fossa in­ 114, sec. 15, T. 5N., R. 26W.), 9 miles north and 5 cluding external auditory meatus but not con­ miles west of Cambridge in Frontier County, tinuous with occipital crest; external auditory Nebraska. meatus slightly dorsal to ventral border of orbits; palate narrow and rugose with a low median keel Stratigraphie Occurrence.-Upper Pliocene and lateral palatine grooves extending from (Kimballian),4 Ogallala Group, Kimball Forma­ canine buttresses to foramen behind M3; palate tion, from channel deposits which rest on upper bec~ming narrower and shallower posteriorly, part of Ash Hollow Formation. leading into a deep, wide groove (beginning Diagnosis.-A peccary larger than across from M2); palatine-maxillary groove di­ Prosthennops crassigens with p4 and P4 sub­ rectly behind posterior lingual portion of M3; pal­ molariform; skull with zygomatic arch expanded ate termi nati ng posteriorily across from pos­ and forehead domed. terior margin of glenoid fossae; glenoid fossae oval and shallow; opening of internal nares Description.-Skull massive; rostrum not about at level of orbits and communicating with elongate as in My/ohyus; and expanded anterior larynx through relatively small heart-shaped to infraorbital foramen; dorsal surface of nasals opening lying anterior to basisphenoid and be­ flat; supraorbital canals extend from nasal notch tween auditory bullae; auditory bullae small along nasals and up lateral sides of maxillaries, kidney-shaped and filled with cancellous bone: over frontals and into supraorbital foramina; two anteromedial border of bullae blade-shaped and pointed medially; bullae inclined anteriorly with 41n the present paper "Pliocene" includes the Valentinian lateral sides enclosed by posterior roots of Clarendonian, ~emphilli~n, and Kimballian provincial ages: zygomatics which curve around to join wings of although the wnters realize that the Kimballian may be equi­ valent to the latest Miocene of Europe. See Part 1, p. 2 of the pterygoids; distinct groove anterodorsal to bulla present Bulletin for further remarks on this subiect. and joining groove leading into eustachian {:::=>

:> z , m , ~ , '"~ III :> r r 5>z "'0m () () :> JJ -< -n oJJ :s::: z m Fig. 1-Prosthennops (Macrogens) graffhami, new species: A, holotype, U.N. S.M. 76051, skull (lateral view); B, referred, U.N.S.M. 76052, partial mandible III (lateral view), from the Kimball Formation, Frontier County, Nebraska. X 3/5. ~ en ~

...... w -..j -~~--=~~--~~-­--===--==

" .d..71iiTfft11it7iil(((1fi,{~ -, "-

>

Fig. 2-Prosthennops (Macrogens) graffhami, new species, holotype: A-B, U. N.S.M. 76051, skull (A, dorsal and B, palatal views); C, referred, U.N.S.M. 76055, left juvenile maxillary with Dp2_Ml, from the Kimball Formation, Frontier County, Nebraska. X 3/5. A NEW KIMBALLIAN PECCARY FROM NEBRASKA I 39 canal; posterior lacerate foramen large and head at same level (greater trochanter more dis­ separate from condyloid foramen; paramastoid tal in My/ohyus and P/atygonus); large nutrient process a low ridge extending from base of con­ foramen centered about one-fourth of way down dyles and wrapping around posterior lacerate anterior face of shaft; distal end narrower than in foramen; basisphenoid short and tilted dorsally My/ohyus; intercondyloid fossa narrower than in with two prominent basilar eminences for origin My/ohyus; lesser trochanter on posterior surface of M. rectus capitis ventralis major; basicranial of shaft; deep sulcus in base of postero-medial region sharply tilted upwardly from occipital surface of greater trochanter; attachment for condyles; foramen magnum small and shield­ round-ligament shallow and posteriorly situated; shaped; expansion of occipital region above three prominent ridges for muscle attachment foramen magnum; large heart-shaped depres­ on posterior face of shaft and digital extensor sion above expansion; upper canine blade-like, fossa not deeply excavated; rotular groove nar­ sharp posteriorly and with thin line of wear on row; patella triangular with a double articulating anterior edge; p2 small and circular with two facet; tibia same length as femur (tibia longer major anterior cusps and a broad, flat posterior than femur in My/ohyus); bowed outwards and area; p3-4 square and submolariform with pro­ similar in proportions to My/ohyus except with tocone, paracone and metacone all rounded narrower proximal end; cnemial crest short (25% cusps of about equal size, hypocone small and of length of tibia as compared to 34% in low (p3-4 less molariform in P. edensis); molars My/ohyus (Lundelius, 1960, p. 22); three crests with transverse lophs of about equal height on posterior surface of shaft; fibula slender, ex­ formed by paracone-protocone and metacone­ panded on both ends, and with an abrupt bend hypocone; no labial or lingual cingula on cheek on the proximal end; calcaneum more elongate teeth except for lingual cingulum on M1; anterior than in My/ohyus and facet for the posterior sur­ and posterior cingula developed on all cheek face of astragalus more anterior; astragalus long teeth and M3 with a small slightly crenulate pos­ and narrow with dorsal and ventral portions of terior basin. Deciduous upper premolars worn about equal width; tarsals about as in My/ohyus; on all examples preserved; Dp2 elongate, and metatarsals III and IV unfused and fairly long, triangular with a large anterior and two posterior metatarsal V small but present; toes not espe­ cusps; DP3 more rectangular with transverse ciallyelongated. lophs formed by protocone-paracone and metacone-hypocone; DP4 submolariform; and SUMMARY AND CONCLUSIONS deciduous premolars with distinct lingual cin­ gula. The origins of Prosthennops may be traced Mandible robust, thickening ventrally with back to the Valentinian species P. (M.) nio­ strong lateral rugosities and a flat ventral mar­ brarensis (Colbert, 1935, 1938). There appears to gin; symphysis long, fused, with an anteroventral have been a split into two lineages at about that median groove and a foramen on each side of it; time. Prosthennops (Macrogens) graffhami from dorsal surface of symphysis anterior to canine the "Amebe/odon fricki Quarry," (Barbour, 1927; roughened; two small pits for genio-hyoids; two 1929), represents the culmination of one of these large and two small mental foramina ventral to lineages. Prosthennops edensis (Frick, 1921) diastema; diastema short and broad; anterior may represent a late form of the other lineage, border of ascending ramus across from heel of although this cannot be clearly shown without a M3; 11 small, procumbent, spatulate and ventrally skull. located; 12 vestigial; wear on incisors flat and According to Guilday (1971, p. 289) the de­ confined to tips; lower canine large with a trian­ velopment of the canine buttresses and suborbi­ gular cross-section; P2 small and triangular; P3 tal zygomatic flares are variable in the flat­ also small but rectangular with four major cusps; headed peccary, P/atygonus. He was not able to P4 molariform; molars rectangular and M3 not establish if these or other characters might be especially elongated. Femur straight and elon­ assigned to sexual and age variation. In the small gate; proximal ends of greater trochanter and sam pie we have of Prosthennops, some size 40 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Fig. 3-Prosthennops (Macrogens) graffhami, new species, referred, partial mandibles: A, U.N.S.M. 76053 (dorsal and lateral views), possible male; B, U.N.S.M. 76052 (dorsal view), from the Kimball Formation, Frontier County, Nebraska; C, U.N.S.M. 76054 (dorsal and lateral views, reversed except for IC), Kimball Formation, Garden County, Nebraska. X 3/5. ~ > .-~-.:-:~~~--=--=--=---=~-

--- ==='5"=-==-~- ca

-::-~~~o::-:-==-~~c"'~§="~_'"

'- ('\

.~--~ ------~~_~__~_-_~_~.~-~~-==-"S-~ -~~~--~ -

r- .."

-'~.~'=- ~=~ ~--~ »­ ~ .'.~- ~- G>. __7''-' • z ." ';:'; ~

(;') "3: r~ r ,.""..,. 0 ); -==~~-~~,? Z mo" z :c~ ,.. -< o ." o:c s:: z m m

~ Fig. 4-Right hind limb, Prosthennops (Macrogens) graffhami, new species, U.M.S.M. 76071, referred: A-E, femur (anterior, lateral, posterior, proximal, and ~ distal views); F-G, patella (anterior and posterior views); H-K, tibia (anterior, lateral, posterior, and distal views); L, fibula (posterior view); M, calcaneum (medial view); N-O, astragalus (anterior and posterior views); P-R, rear pes (medial, anterior, and posterior views), Kimball Formation, Frontier County, Nebraska. X 1/2. ~ 42 1 BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

, ~ .,------­

D

Fig. 5-Possible phylogeny of Prosthennops (Macrogens): A, P. (M.) graffhami, Kimball Formation, Kimballian; B, P. (M.) crassigens, ?Clarendonian (from Gidley, 1904, Fig. 15); C, P. (M.) niobrarensis, Burge Member of Valentine Formation, Valentinian (from Colbert, 1935, Fig. 199); D, Dyseohyus stirtoni, Sheep Creek Formation, late Hemingfordian (from Woodburne, 1969, PI. 48, Fig. 1). X 1/2.

bimodality is present which we ascribe to sexual ness of the cranium, and expanded zygomatic variation (Fig. 3). The larger individuals are also flares. Guilday (1971) associates these features more robust and have larger canines. with an open country habitat, and a savannah The orbits of Prosthennops are located far to parkland might not be a bad estimate of the the rear and are high on the skull, resembling habitat of Prosthennops. The central axis of the My/ohyus in these respects. While not so ex­ orbit is directed anteriorly when the snout is tremely modified, the skull of Prosthennops pointed ventrally. This would help to permit use­ resembles that of the wart hog Phacochoerus ful vision while the animal was eating and further aethiopieus in the position of the orbit, short- suggests the presence of a habitat where free A NEW KIMBALLIAN PECCARY FROM NEBRASKA I 43

TABLE 1 Prosthennops (Macrogens) graffhami, new species MEASUREMENTS' OF SKULL

P. (M.) graffhami n. sp. SKULL Holotype U.N.S.M. 26051

Length (max., including supraoccipital crest and incisors) ...... (362.00)2 Length (ant. of canines to posterior of condyles) ...... 256.5 Width (max., across zygomatic arches) ...... 207.7 Width (max., across supraorbital processes) ...... 123.5 Width (min., across lacrimal tuberosities) ...... 105.0 Distance from anterior rim of orbit to anterior portion of canine buttress ...... 73.0 Distance from anterior rim of orbit to supraoccipital crest ...... 130.0 Distance from ventral portion of occipital condyles to top of supraoccipital ...... 118.0 Width across condyles (max.) ...... 53.3 Breadth across canines (max.) ...... 84.5 Width of palate between fourth superior premolars ...... 29.0 Width of palate between canines ...... 49.0 Length of dental series (C-M3 inclusive) ...... 153.0 Length of diastema between C and p2 ...... 39.0 Length of superior premolar series (P2_p4) ...... 37.5 Length of superior molar series (M'-M3) ...... 56.0 C/3 ...... 21.2 C/4 ...... 12.8 p2 ...... 10.8 p2 ...... 10.7 p3 ...... 13.5 p3 ...... 13.3 p4 14.7 p4 ...... 15.0 M' ...... 16.6 M' ...... 14.9 M2 ...... 20.2 M2 ...... 17.1 M3 ...... 20.5 M3 ...... 16.0 lThe measurements are taken to the nearest millimeter except on dentition where they are measured to the nearest one-tenth of a millimeter. 2( ) = approximate. 3First measurement, anterior-posterior length. 'Second measurement, max, width. 44 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

TABLE 2 Prosthennops (Macrogens) graffhami, new species COMPARATIVE MEASUREMENTS' OF MANDIBULAR RAMI

P. M.) graffhami, P. (M.) graffhami, n. sp. n. sp. MANDIBULAR RAMI Referred Referred U.N.S.M.76052 U.N.S.M. 76504

Depth below anterior of third inferior molar ...... 45.1 Length of dental series (lC-M3 inclusive) ...... 160.4 150.4 Lenth of inferior molar-premolar series ...... 91.6 92.7 Length of inferior premolar series (max.) ...... 36.0 36.5 Length of inferior molar series (max.) ...... 53.5 56.7 h ...... 10.6 h ...... 5.1 12 ...... 12.7 11.7 12 ...... 7.0 5.8 b ...... 4.7 b ...... 2.6 ~ ...... (18.4) 16.5 IC ...... (14.6) 12.5 ~ ...... 11.0 10.3 P2 ...... 7.6 6.8 P3 ...... 11.8 12.7 P3 9.7 9.6 P4 ...... 13.2 14.7 P4 ...... 12.4 12.0 M, ...... 15.3 13.7 M, ...... 13.2 (12.6) M2 ...... 16.8 18.6 M2 ...... 14.4 14.5 M3 ...... 24.5 25.8 M3 ...... 14.9 15.0 Post-canine diastema ...... , ...... 51.5 49.9 Pre-canine diastema ...... 7.5 6.5 Width symphysis at narrowest point ...... 36.9 33.3 'See footnotes for Table ,. A NEW KIMBALLIAN PECCARY FROM NEBRASKA I 45

TABLE 3 Prosthennops (Macrogens) graffhami, new species COMPARATIVE MEASUREMENTS OF LIMBS

P. (M.) graffhami n. sp. Referred LIMBS U.N. S.M. 76071 and 76065

Length of femur (articular) ...... 196.5 Length of tibia (articular) ...... , " ...... , ...... , ...... 192.0 Length of tibia (max.) ...... 204.0 Length of calcaneum (max.) ...... 76.0 Length of fibula (max.) ...... 185.5 Length of metatarsal II ...... 69.3 Length of metatarsal III ...... 94.5 Length of metatarsal IV ...... 99.0 Length of astragalus (max.) ...... 42.0 Proximal width of astragalus (art.) ...... 21.5 Distal width of astragalus (art.) ...... 22.0 Calcaneum, anteroposterior-length distal end (max.) ...... 32.0 Proximal phalanx, lateral toe, right, max. length...... 40.0 Medial phalanx, lateral toe, max. length...... 25.5 Distal phalanx, lateral toe, max. length...... 29.0

vIsion was important. The legs were elongate ble: the late Professor E. F. Schramm and Mr. and suggest a cursorial existence, although Childs Frick for providing funds for field work; there was not as much toe reduction on the hind and Professors T. M. Stout and Lloyd G. Tanner foot as in the Pleistocene genera (Fig. 4). for assistance and advice. The members of the According to Woodburne (1969, p. 331), University of Nebraska State Museum's 1947 and Oyseohyus stirtoni from the late Miocene of Ne­ 1948 field crews at U.N.S.M. Call. Loc. Ft-40 in braska is either ancestral or is near the lineage Frontier Cou nty, Nebraska, deserve special that produced Prosthennops niobrarensis. thanks for the discovery of the fossils listed in Oyseohyus stirtoni differs from Prosthennops in this paper (see Acknowledgments in Part 1 of having a larger and more anterior eye socket, a this Bulletin for names of the collectors). The nearly straight dorsal skull outline (forehead not 1970 field party, which collected U.N.S.M. domed), and no expanded zygomatic flares as in specimens numbered 76055 and 76056, con­ Prosthennops. An increase in size, development sisted of Messrs. Robert Buschow, John Knapp, of a domed cranium and expanded zygomatic Ben Obitz, and William Rovnak. The U.N.S.M. flares, and the posterior movement of the eye are 76054 specimen from Garden County was col­ all shown in the possible evolutionary series: lected by Mr. Tom Middleswart and the late Oyseohyus stirtoni~ Prosthennops (Macrogens) Messrs. William Chalupka and S. R. Sweet. Mr. niobrarensis~ P. (M.) crassigens~ P. (M.) graf­ George Corner has been very helpful in the fhami (Fig 5). Prothennops (Macrogens) became cataloging and curating of the specimens and in extinct at the end of the Kimballian, and the the preparation and editing of the manuscript. North American Pleistocene peccaries must be Mr. Robert Miller was responsible for preparing derived from other lineages. the drawings on Fig. 1, A and B, on Fig. 2, and A and B on Fig. 3. Mrs. Mary Tanner illustrated ACKNOWLEDGMENTS Figs. 2C, 3C, 4, and 5, and also helped with edit­ The writers are indebted to the following for ing of the manuscript. The manuscript was typed making the publication of this manuscript possi- by Mrs. Mary Tanner and Miss Rhonda Carlsten. 46 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

REFERENCES sociated fauna, Kentucky Pleistocene. Annals of Carnegie Mus. 43(9): 249-320, Figs. 1-31,8 tables. Lundelius, Ernest L. 1960. Mylohyus nasutus. Long-nosed Barbour, Erwin Hinckley. 1925. Prosthennops xiphodonticus, peccary of the Texas Pleistocene. BUll. Texas Mem. Mus. sp. nov. A new fossil peccary from Nebraska. Bull. [Univ.J (1): 1-40, Figs. 1-8, Pis. 1-4, Tables 1-6. Nebraska State Mus. 1(3): 25-32, Figs. 12-13. Matthew, W. D. 1924. Third contribution to the Snake Creek ---. 1927. Preliminary notice of a new proboscidean Fauna. Bull. Amer. Mus. Nat. Hist. 50(2): 59-210, Figs. 1-63. Amebelodon fricki, gen. et sp. nov. Ibid., 1(13): 131-134, Matthew, W. D., and J. W. Gidley. 1904. New or little known Figs. 89-91. mammals from the Miocene of South Dakota. American ---.1929. The mandible of Amebelodon fricki. Ibid., 1(15): Museum Expedition of 1903. BUll. Amer. Mus. Nat. Hist. 20 139-146, Figs. 93-97. (22): 241-268, Figs. 1-15. Colbert, Edwin H. 1935. A new fossil peccary, Prosthennops Schultz, C. Bertrand, Marian R. Schultz, and Larry D. Martin. niobrarensis, from Brown County, Nebraska. Bul!. Univ. 1970. A new tribe of saber-toothed cats (Barbourofelini) Nebraska State Mus. 1(44): 419-430, Fig. 198-199. from the Pliocene of North America. Bull. Univ. Nebraska ---. 1938. Pliocene peccaries from the Pacific Coast Re­ State Mus. 9 (1): 1-31, Figs. 1-13, Frontispiece, Tables 1-2. gion of North America. Carnegie Inst. Washington Pub!. Tanner, Lloyd G. 1967. A new species of rhinoceros, (487): 241-269, Figs. 1-4, Pis. 1-6. Aphelops kimballensis from the latest Pliocene of Ne­ Frick, Childs. 1921. Extinct vertebrate faunas of the badlands braska. Bull. Univ. Nebraska State Mus. 6 (1): 1-16, 4 of Bautista Creek and San Timoteo Canon, southern plates. California. Univ. California Pub!. Bull. Dept. Geo!. 12(5): Wood burne, Mic~hael O. 1969. Systematics, biogeography, 277-424, Figs. 10-65, Pis. 43-50. and evolution of Cynorca and Dyseohyus (Tayassuidae). Guilday, John E., Harold W. Hamilton, and Allan D. McCady. BUll. Amer. Mus. Nat. Hist. 141 (2): 271-356, Figs. 1-14, Pis. 1971. The Welsh Cave Peccaries (Platygonus) and as- 41-51, Tables 1-17. BULLETIN OF VOLUME 10, NUMBER 1, PART 4 The University of Nebraska State Museum FEBRUARY, 1975

c. Bertrand Schultz Larry D. Martin

Bears (Ursidae) from the Late Cenozoic of Nebraska Frontispiece-Photograph showing Allen Graffham and Alex Keith (left to right) at U.N.S.M. Coli. Loc. Ft-40, Frontier County, Nebraska. Mr. Keith discovered the Kimballian fossil locality in 1927 when he uncovered the holotype of Amebelodon fricki Barbour, a shovel-tusked mastodont, at approximately the spot where the two men are standing. Mr. Graffham was the field leader in 1947, when the following holotypes were discov­ ered: Indarctos keithi Schultz and Martin, new species; Prosthennops (Macrogens) graffhami Schultz and Martin (described in Pt. 3 of this Bulletin); Teleoceras schultzi Tanner (described in Pt. 2 of this Bulletin); Aphelops kimballensis Tanner; ?Tapirus simpsoni Schultz, Martin, and Corner (described in Pt. 1 of this Bul­ letin); Barbourofelis fricki Schultz, Schultz, and Martin; and Proagriocharis kimballensis Martin and Tate. Photograph by Schultz, 1947. c. Bertrand Schultz Larry D. Martin

CENOZOIC MAMMALS FROM THE CENTRAL GREAT PLAINS

Part 4 Bears (U rsidae) from the Late Cenozoic of Nebraska

BULLETIN OF The University of Nebraska State Museum VOLUME 10, NUMBER 1, PART 4 FEBRUARY, 1975 BULLETIN OF VOLUME 10, NUMBER 1, PART 4 THE UNIVERSITY OF NEBRASKA STATE MUSEUM FEBRUARY, 1975

Pp. 47-54, Tables Frontispiece, Figs. 1-3

ABSTRACT

Part 4. Bears (Ursidae) from the Late Cenozoic of Nebraska

c. Bertrand Schultz Larry D. Martin

A ramus and partial premaxilla establish the presence of a new subspecies of Indarctos in the upper Pliocene (Kimball Formation, Ogallala Group) of Frontier County, Nebraska. An extremely large species of Agriotherium is represented by fragmentary remains from the middle Pliocene (middle part of Ash Hollow Formation, Ogallala Group) of Sherman County, Nebraska.

CONTRIBUTION OF The Department of Geology, College of Arts and Sciences, and the Division of Vertebrate Paleontology of the Museum. Schultz1 Martin2

Bears (Ursidae) from the Late Cenozoic of Nebraska

INTRODUCTION SYSTEMATIC DESCRIPTIONS

This study is part of a series of papers dealing Class: MAMMALIA primarily with the fauna of the Kimball formation Order: CARNIVORA in Nebraska (Barbour 1927, 1929; Barbour and Schultz, 1941; Schultz and Stout, 1948, 1961; Family: URSIDAE Kent 1963, 1967; Tanner, 1967; Short, 1969; Mar­ tin and Tate, 1970; Schultz, Schultz, and Martin, Indarctos oregonensis keithi, new subspecies 1970). The Kimballian fauna from Nebraska dif­ fers from the Hemphillian fauna in that most of Holotype.-Left ramus with 11-13 alv., IC, Pl, the known forms are markedly more advanced, P2-P3 alv., P4-M2, M3 (alv.), U.N.S.M. 76009 (Figs. but definitely pre-Blancan (Early Pleistocene). 1-2). A large bear, /ndarctos oregonensis keithi, Referred Specimen from the Type new subspecies, from U.N.S.M. Coli. Loc. Ft-40 is Locality.-Left premaxilla with 11_1 3 alv., CI, included in the Kimballian fauna from Nebraska. U.N.S.M. 76010 (Fig. 2) This specimen appears to An edentulous ramus and a radius of be from the same individual as the holotype, and Agriotherium brought in by Frank Garvel of Ash­ therefore should be considered to be part of the ton, Nebraska, from near U.N.S.M. Coil. Loc. holotype. Sm-101 in Sherman County, Nebraska, is also reported. This latter locality is Hemphillian in age Type Locality.-U.N.S.M. Coil. Loc. Ft-40 (= (Ash Hollow Formation) and has produced "Amebe/odon fricki Quarry"), E. 1/2, SW. 114, SE. Machairodus as well as some rhinoceros mate­ 114, sec. 15, T. 5N., R. 26W., 8 miles North and 5 rial (personal communication from Lloyd Tan­ miles West of Cambridge, Frontier County, Ne­ ner). braska. Stratigraphie Occurrence.-Upper Pliocene (Kimballian),3 Ogallala Group, Kimball Forma­ tion, from channel deposits which rest on the upper part of the Ash Hollow Formation. lCurator of Vertebrate Paleontology, University of Ne­ braska State Museum; and Professor of Geology, Depart­ ment of Geology. 2Assistant Curator of Vertebrate Paleontology, University of 31n the present paper "Pliocene" includes the Valentinian, Kansas Museum of Natural History; Assistant Professor of Clarendonian, Hemphillian, and Kimballian Provincial ages, Systematics and Ecology, University of Kansas; and Re­ although the writers realize that the Kimballian may be equiv­ search Affiliate, Division of Vertebrate Paleontology, Univer­ alent to the latest Miocene of Europe. See Part 1, p. 2 of the sity of Nebraska State Museum. present Bulletin for further remarks on this subject. .j>. co

cOJ Q)D o"~ r r m :::::lz ~ J/-Cpp// o ~.,;Y--:-'"7?~/"~ "T1 >' - /")' --f /~' ~//A :::c m c z <: m :D CJ) =i -< o "T1 z m OJ :D» CJ)

~ CJ)

~ --fm s:: c CJ) m c s::

Fig. 1-lndarctos oregonensis keithi, new subspecies, holotype, U.N. S.M. 76009, left ramus (occlusal, labial, and lingual views), from the lower part of the Kimball Formation, Frontier County, Nebraska. X 1/2. BEARS (URSIDAE) FROM THE LATE CENOZOIC OF NEBRASKA I 49

seteric fossa roughened and ridged; angular process large and continuing as a strongly rugose masseteric line under masseteric fossa; coronoid process large (upper portion broken off), and mandibular condyle enormous.

Discussion.-Indarctos oregonensis keithi is the first record of this genus from Nebraska. It differs from Agriotherium in the shape of the M2, as the talonid and trigonid are about equal in size. In Agriotherium the trigonid is usually larger than the talonid. This tooth is also more elongate than the M2 of Agriotherium. The M3 is represented by a broken alveolus which indi­ cates that the tooth was single rooted. The shape of the ramus is different from that found in Agriotherium and is tapered in its forward por­ Fig. 2-lndarctos oregonensis keithi, new subspecies, refer­ red, U.N.S.M. 76009, left canine, from same quarry as tion, making it similar to the shape of the ramus holotype. X 1/2. in Ursus. There is no premasseteric fossa, which is often present in Agriotherium. The Nebraska species of Indarctos resembles Diagnosis.-Mandible very large and elon­ I. arctoides from the upper Pliocene of Europe, gate; premasseteric fossa absent; mandibular but it is much larger in size. I. o. keithi has condyle large and close to the angular process; marked similarities to I. maraghanus from the dental formula 11-3, IC, P1-4, M1-3; incisors Pliocene sediments of the Maragha in Persia, dif­ crowded with 12 largest and posterior to hand 13; fering from that species in the shape of the M1 IC large and robust; P1-2 small and single rooted; and having slightly smaller dentition. Indarctos P3-4 double rooted; M1 large, narrow, and elon­ oregonensis keithi differs from I. nevadensis gate; trigonid of M1 narrower than talonid; from the Hemphillian of Nevada and from I. at­ trigonid and talonid about equal on M2. ticus from the upper Pliocene (Pontian) of the Premaxilla massive with crowded 11-3 and a large Island of Pikermi (in the Aegean Sea), both in its robust C/; P and 12 approximately same size; 13 large size and the presence of a double-rooted very large. P3 (see Thenius, 1959). It might be noted here Description.-Dentition worn; alveoli indicat­ that I. nevadensis and I. atticus are very similar in ing small laterally flattened incisors; canine most respects. It is unfortunate that the C/, IC, short and heavy with a wear facet on the antero­ and M2'S are too worn and mutilated to have lingual margin; P1 peg-like (worn down to the much diagnostic value in the new subspecies. root); single alveolus for P2; short diastema (18.6 These teeth do agree quite closely in size and mm.) between P2 and P3; double alveolus for P3; proportions with I. oregonensis, but the upper P4 inclined posteriorly against M1; accessory canine of the Nebraska specimen is definitely cusps on P4 not noticeably developed; cingulum larger and more robust. The measurements of barely evident on cheek teeth; M1 large and the left CI of the holotype of I. o. oregonensis, elongate, with trigonid narrow, talonid wide and U.C. 22362, are as follows: anteroposterior rounded, paraconid fairly large; M2 elongate diameter at base of enamel, approximately 29.8 with trigonid and talonid about equal in size; M3 mm.; and the transverse diameter at base of large and single rooted (based on alveolus); two enamel, 20.2 mm. (Merriam, Stock, and Moody, large mental foramina (under diastema between 1916, p. 582). The corresponding measurements P2 and P3 and under P4) posterior to two small of the CI of I. o. keithi are approximately 39 mm. mental foramina (under IC); masseteric fossa and 28 mm. large and posteriorly situated; interior of mas- The specimens from the Hemphillian Coffee 50 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Ranch Local Fauna reported by Dalquest (1969, unusually large; strong rugosity for muscle at­ pp. 9-10) as Indarctos oregonensis4 include a M2 tachment along inner side of ventral border of with the trigonid much wider then the talonid. jaw; anterior portion of premasseteric fossa lies The carnassial Dalquest (1969, Fig. 4) illustrates beneath M3; right radius U.N.S.M. 26202 is large is very similar in size and configuration to the and massive; bicipital tuberosity well developed, one illustrated as Hyaenarctos, species, by Frick as is styloid process- general configuration of (1926, Fig. 29). Hyaenarctos is a junior synonym radius very similar to Ursus (concavity above of Agriotherium, and it is probably to the latter facet which articulates with ulna much larger, genus that the Coffee Ranch specimens should and all of ridges for muscle attachments strongly be referred. They would belong to one of the developed); about 10% larger than radius of largest species of Agriotherium and are possibly Arctotherium simum from Potter Creek Cave conspecific with the undescribed form from near (Merriam and Stock, 1925). Sm-101 reported in this paper. Discussion.-This material is notable for its large size, and the relative shortness of the Agriotherium sp. ramus. It contrasts sharply with the elongate, Material.-Right ramus with 12 (alv.), 13 (rt.), IC more dog-like ramus of Indarctos (Fig. 1) in this (rt.), P3-P4 alv., M1 (rt.), M2-M3 alv., U.N.S.M. respect. The ramus resembles most closely that 76011 (Fig. 3B); and right radius, U.N.S.M. 76013 of Agriotherium schneideri Sellards, but differs (Fig. 3A), which is probably from the same indi­ from that species in the lack of P1, the presence vidual. of P3, and the largeness of the ramus. Agriotherium sivalensis is also similar, but is not Locality.-From the same drainage as so large, and has a P1 as well as a P2. U.N.S.M. Coil. Loc. Sm-101, 7 mi. north and 1 mi. Agriotherium gregoryi was also slightly smaller. east of Ashton, Sherman County, Nebraska. The Nebraska material probably represents an Stratigraphie Occurrence.-Middle Pliocene undescribed form, but no name is proposed be­ (?Hemphillian), Ogallala Group, Ash Hollow cause of the fragmentary nature of the material. Formation (probably middle). SUMMARY AND CONCLUSION Description.-The right ramus of this large ursid has a dental formula of II?, IC, P3-4, M1·3; Although Agriotherium has been designated incisors probably reduced in number with only a an index fossil for the Hemphillian age (Wood, et trace of alveolus and root for very small 12, and ai, 1941, p. 12), both it and Indarctos are ex­ no signs alveolus for b; 11 moderate in size; tremely rare fossil mammals. Because of this it canine very large with oval base; P3 very small seems worth while to report the ramus of and peg-like with a single root; P4 unusually Agriotherium from Sherman County, Nebraska, large, double rooted and inclined posteriorly; M1 although it is badly damaged. It differs from and M2 large; M3 small and apparently single other described species of Agriotherium in being rooted; M2 and M3 placed high on ascending somewhat larger and more massive. A premas­ ramus (alveolar measurements are given for the seteric fossa comparable to the one in teeth, and except for P4, are all probably some­ Arctotherium and even better developed than what smaller than actual measurements of the the premasseteric fossa of Agriotherium greg­ teeth would have been); interalveolar border oryi (Frick) (Frick, 1926, Fig. 36) may have been strongly roughened symphysial surface large present. Agriotherium and Arctotherium are and squarish in outline; three prominent mental both short-faced bears with massive limbs (Mer­ foramina present and posterior one (under P4) rian and Stock, 1925). Most species of Agriotherium however have lost too many of the anterior premolars to be regarded as ancestral to 4Apparently the reference of this material to Indarctos Arctotherium. The development of a premas­ nevadensis in table 7 (Oalquest, 1969) is a lapsus, although on page 10 of the same paper he seems to suggest identity seteric fossa might be related to the develop­ with I. nevadensis. ment of a short and deep skull and relatively BEARS (URSIDAE) FROM THE LATE CENOZOIC OF NEBRASKA I 51 deep jaws. It is found in Agriotherium, Arctodus 879) is much closer to the Asian forms than is I. and Tremarctos. However, it is also found in the keithi. The exact relationship of I. oregonensis rather dog-like genus Hemicyon. keithi to I. oregonensis oregonensis cannot be Both Indarctos and Agriotherium probably determined until additional material of both sub­ originated in Eurasia and migrated to North species is discovered and described. America during the Hemphillian or perhaps as early as the Clarendonian. The large Middle ACKNOWLEDGMENTS Miocene carnivores were primarily am­ phicyonids in North America. The diversity of The two Sherman County Indarctos these amphicyonids was very reduced before the specimens, U.N.S.M. 76011 and 76013, were col­ appearance of these giant bears. Indarctos and lected in 1950 and donated by Mr. Frank Garvel Agriotherium achieved an essentially holarctic of Ashton, Nebraska. The holotype of Indarctos distribution during the Ogallala Pliocene, essen­ oregonensis keithi, U.N.S.M. 76009 and 76010, tially similar to that of Machairodus. were collected in 1947 by the University of Ne­ The presence of Indarctos in the Kimballian of braska State Museum field party (see Acknowl­ Nebraska is interesting as I. keithi is one of the edgments in Part 1 of this Bulletin for names of largest and most advanced known species of the the collectors). Figs. 1 and 3 were illustrated by genus. Indarctos nevadensis Macdonald has Mrs. Mary Tanner, and Fig. 2 by Mr. Jerry Tanner. teeth almost as large but has a much smaller Mr. George Corner and Mrs. Mary Tanner were ramus, and as described by Macdonald (1959, p. very helpful in the preparation of the manuscript. 52 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

A 8

Fig. 3-Agriotherium sp., A, U.N.S.M. 76013, right radius (medial and lateral views), B, U.N.S.M. 76011, edentulous right ramus (labial and lingual views), from the Ash Hollow Formation (?middle), Sherman County, Nebraska. X 1/3. BEARS (URSIDAE) FROM THE LATE CENOZOIC OF NEBRASKA I 53

TABLE 1 Indarctos COMPARATIVE MEASUREMENTS OF MANDIBULAR RAMI

1 oreogonensis keithi n. subsp. MANDIBULAR RAMI Holotype I. nevadensis (Macdonald) U.N.S.M. 76009 U.C. Mus. Pal. 38629 I. arctoides (Deperet)' I. atticus (Dames)' Total length of the ramus ...... 369 (253)2 (248) Depth of ramus behind canine ...... 70 (52) Depth of ramus posterior to M, ...... 92 (64) 49.5 61 IC anteroposterior ...... 35 26.5 18.4 IC transverse ...... 24 20.5 13.8 p, anteroposterior ...... 7 12.1 8.5 (7.0) p, transverse ...... 7 7.5 6.8 (6.8) P2 anteroposterior ...... (7.5) 11.9 9.7 (7.0) P2 transverse ...... 6 8.1 6.1 (6.4) P3 anteroposterior ...... (11.2) 12.7 11.0 (6.8) P3 transverse ...... 6 9.1 5.5 (6.4) P4 anteroposterior ...... 19.5 23.5 15.8 21.0 P4 transverse ...... 12.4 14.7 9.6 13.2 M, anteroposterior ...... 39.8 40.5 30.8 39.3 M, transverse ...... (21) 20.6 15.8 20.4 M2 anteroposterior ...... 30.4 31.4 24.1 28.7 M2 transverse ...... 20.7 16.5 20.8 M3 anteroposterior ...... ? 17.0 M3 transverse ...... ? 16.4

'Measurements from Macdonald, 1959. 2( ) = alveloar or approximate measurement.

TABLE 2 Agriotherium sp. COMPARATIVE MEASUREMENTS' OF MANDIBULAR RAMUS AND RADIUS

MANDIBULAR RAMUS U.N.S.M.76011

Depth of ramus at posterior margin of the canine...... 69 Depth of ramus at posterior margin of M, ...... 100 IC greatest anteroposterior diameter...... (39)2 IC greatest transverse diameter...... (23) P3 greatest anteroposterior diameter...... (3.5) P3 greatest transverse diameter...... (4) P4 greatest anteroposterior diameter...... (27) P4 greatest transverse diameter ...... (14) M, greatest anteroposterior diameter ...... (45) M, greatest transverse diameter ...... (15) M2 greatest anteroposterior diameter ...... (36) M2 greatest transverse diameter...... (11) M3 greatest anteroposterior diameter ...... (11) RADIUS U.N.S.M.76013

Greatest length...... 421 Greatest diameter of proximal extremity...... 80 Width of shaft at middle...... 43 Thickness of shaft at middle...... 33 Greatest diameter of distal extremity...... 58

'The measurements are taken to the nearest millimeter except on dentition where they are measured to the nearest one-tenth of a millimeter. 2( ) = alveolar. 54 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

REFERENCES Sellards, E. H. 1916. Fossil vertebrates from Florida: A new Miocene fauna; new Pliocene species; the Pleistocene Barbour, Erwin Hinckley. 1927. Preliminary notice of a new fauna. Ann. Rept. Florida State Geol. Surv.: 77-119, Pis. proboscidean Amebe/odon fricki, gen. et sp. nov. Bull. 10-14. Univ. Nebraska State Mus. 1 (13): 131-134, Figs. 89-91. Schultz, C. Bertrand, and W. D. Frankforter. 1948. Prelimi­ --.1929. The mandible of Amebelodon fricki. Ibid. 1 (15): nary report on the Lime Creek Sites: New evidence of Early 139-146, Figs. 93-97. Man in southwestern Nebraska. Bull. Univ. Nebraska State Barbour, Erwin H., and C. Bertrand Schultz. 1941. A new Mus. 3 (4), Pt. 2: 43-62, Figs. 1-13. species of Sphenopha/os from the upper Ogallala of Ne­ Schultz, C. Bertrand, Gilbert C. Lueninghoener, and W. D. braska. Bull. Univ. Nebraska State Mus. 2 (6): 59-62, Fig. Frankforter. 1948. Preliminary geomorphological studies 23. of the Lime Creek area. Ibid. 3 (4). Pt. 1: 31-42, Figs. 1-6. Dalquest, Walter W. 1969. Pliocene carnivores of the Coffee Schultz, C. Bertrand, Marian R. Schultz, and Larry D. Martin. Ranch. Texas Mem. Museum Bull. 15: 1-44, Figs. 1-11, 1970. A new tribe of saber-toothed cats (Barbourofelini) Tables 1-31. from the Pliocene of North America. Ibid. 9 (1): 1-31, Fron­ Frick, Childs. 1921. Extinct vertebrate faunas of the badlands tispiece, Figs. 1-13. of Bautista Creek and San Timoteo Canon, southern Schultz, C. Bertrand, and Thompson M. Stout. 1948. Pleis­ California. Univ. California Publ. Bull. Dept. Geol. (12): tocene mammals and terraces in the Great Plains. Bull. 341-347, Figs. 1-165, Pis. 43-50. Geol. Soc. Amer. 59 (6): 553-588, Figs. 1-4, PI. 1, --. 1926. The Hemicyoninae and an American Tertiary (Also:Discussion, Ibid.: 623-625) bear. Bull. Amer. Mus. Nat. Hist. 56 (1): 1-119, Figs. 1-63, --. 1961. Field Conference on the Tertiary and Pleis­ Frontispiece. tocene of Western Nebraska (Guide book for the Ninth Kent, Douglas C. 1963. A late Pliocene faunal assemblage Field Conference of the Society of Vertebrate Paleontol­ from Cheyenne County, Nebraska. Unpublished M.Sc. ogy). Special Publ. Univ. Nebraska State Mus. (2): 1-55, thesis, Univ. Nebraska Dept. Geol.: 1-143, Figs. 1-41, Ap­ Figs. 1-47, 2 charts, 1 map. pendices A, B, and C. Short, Lester L., Jr. 1969. A new genus and species of ---. 1967. Citel/us kimbal/ensis, a new Late Pliocene gooselike swan from the Pliocene of Nebraska. Amer. Mus. ground squirrel. Bull. Univ. Nebraska State Mus. 6(2): Novit. (2369): 1-7, Fig. 1. 17-26, Figs. 1-3. Simpson, George Gaylord. 1945. The principles of classifica­ Lugn, A. L. 1939. Classification of the Teritary system in Ne­ tion and a classification of mammals. Bull. Amer. Mus. Nat. braska. Bull. Geol. Soc. Amer. 50: 1245-1276, 1 plate. Hist. 85: 1-350. Macdonald, J. R. 1959. The Middle Pliocene mammalian Tanner, Lloyd G. 1967. A new species of rhinoceros, fauna from Smiths Valley, Nevada. Jour. Paleont. 33 (5): Aphe/ops kimbal/ensis, from the latest Pliocene of Ne­ 877-880, Figs. 1-3. braska. Bull. Univ. Nebraska State Mus. 6 (1): 1-16, 4 Martin, Larry D., and James Tate, Jr. 1970. A new turkey from plates. the Pliocene of Nebraska. Wilson Bull. 82 (2): 214-218, Fig. Thenius, Erich. 1959. Indarctos arctoides (Carnivora, Mam­ 1. malia) aus dem Pliozan Osterreichs nebst einer Revision Mecquenem, R. de. 1925. Contribution a I'etude des fossiles der Gattung. Neues Jb. Geol. u. Palaont., Abh. 108 (3): de Maragha. Ann. Paleont., 13-14, 135-160 u. 1-36, Paris. 270-295, Figs. 1-8. Merriam, John C., and Chester Stock. 1925. Relationships WOOd, Horace E., 2nd, Ralph W. Chaney, John Clark, Edwin and structure of the short-faced bear, Archtotherium, from H. Colbert, Glenn L. Jepson, John B. Reeside, Jr., and the Pleistocene of California. Pub. Carnegie Inst. 347: 1-35. Chester Stock. 1941. Nomenclature and correlation of the --. 1927. A hyaenarctid bear from the Later Tertiary of North American continental Tertiary. Bull. Geol. Soc. Amer. the John Day basin of Oregon. Ibid. 346: 39-44, 1 fig. 52: 1-48, 1 plate. Merriam, John C., Chester Stock, and Clarence L. Moody. Zdansky, Otto. 1924. Jungtertiare Carnivoren Chinas. 1916. An American Pliocene bear. Univ. California Publ. Paleont. Sinica 2: 1-155, Figs. 1-24, Pis. 1-33. Bull. Dept. Geol. 10:87-109. BULLETIN OF VOLUME 10, NUMBER 1, PART 5 The University of Nebraska State Museum FEBRUARY, 1975

Larry D. Martin C. Bertrand Schultz

Scimitar-toothed Cats, Machairodus and Nimravides, from the Pliocene of Kansas and Nebraska Frontispiece-Restoration of the head of Machairodus c%radensis tanneri from the Pliocene of Kansas and Nebraska. Restoration by Mary Tanner. X 1/4. Larry D. Martin C. Bertrand Schultz

CENOZOIC MAMMALS FROM THE CENTRAL GREAT PLAINS

Part 5 Scimitar-toothed Cats, Machairodus and Nimravides, from the Pliocene of Kansas and Nebraska

BULLETIN OF The University of Nebraska State Museum VOLUME 10, NUMBER 1, PART 5 FEBRUARY, 1975 BULLETIN OF VOLUME 10, NUMBER 1, PART 5 THE UNIVERSITY OF NEBRASKA STATE MUSEUM FEBRUARY, 1975

PP.55-63, Frontispiece, Figs. 1-5

ABSTRACT

Part 5. Scimitar-toothed Cats, Machairodus and Nimravides, from the Pliocene of Kansas and Nebraska

Larry D. Martin C. Bertrand Schultz

"Machairodus catocopis Cope" is shown to be a pseudaelurin cat belonging to the genus Nimravides Kitts. Nimravides thinobates (Macdonald) is a possible synonym of N. catocopis (Cope). Nimravides is compared with the Eurasian Machairodus-like cat, Dinofelis. Machairodus (HeterofeJis) coloradensis is reported from the Kimball Formation, upper Pliocene (Kimballian) of Cheyenne County, Nebraska, and from the upper part of the Ash Hollow Formation, Pliocene (Hemphillian) of Sherman County, Nebraska. The Kimballian form is described as a new sub­ species, Machairodus coloradensis tanneri.

CONTRIBUTION OF The Department of Geology, College of Arts and Sciences, and the Division of Vertebrate Paleontology of the Museum. Martin1, Schultz2

Scimitar-toothed Cats, Machairodus and Nimravides, from the Pliocene of Nebraska and Kansas3

INTRODUCTION coloradensis in synonomy with M. catocopis and this concept has generally been followed. New The genus Machairodus has long been as­ material from Smith County, Kansas, which is sociated with the Hemphillian of North America near the type locality and horizon of M. and the Pontian of Eurasia. It was first reported Catocopis, permits a reevaluation of that species from North America when Cope, in 1887, de­ and shows that it is a pseudaelurin cat, scribed a partial symphysis of a mandible from Nimravides. Machairodus coloradensis is re­ the Pliocene of Kansas as Machairodus ported from the Upper Ash Hollow deposits catocopis. He did not illustrate this specimen, (Hemphillian) of Sherman County, Nebraska, and it has largely been ignored (see Fig. 3, C and and is compared to a new subspecies of E in the present paper). The concept of the Machairodus described in this paper from the species has rather been based on abundant Kimball (Kimballian) Formation of Cheyenne material from Ogallala deposits (Hemphillian) of County, Nebraska. This new subspecies demon­ Yuma County, Colorado, described by Cook strates evolutionary trends which seem to be (1922) as M. (Heterofelis) coloradensis; later Burt leading to Ischyrosmilus from the lower Quater­ (1931) published additional records from the nary (Blancan). We do not regard the pseudaelu­ Hemphillian of Texas. Matthew (1924) placed M. rin cats to be of subfamily rank, although they probably should be placed in a separate tribe, Pseudaelurini, which represents primitive 'Assistant Curator of Vertebrate Paleontology, University felines. of Kansas Museum of Natural History, and Assistant Profes­ sor of Systematics and Ecology, University of Kansas, Law­ rence; Research Affiliate, University of Nebraska State 31n the present paper "Pliocene" includes the Valentinian, Museum. Clarendonian, Hemphillian, and Kimballian provincial ages, 2Curator of Vertebrate Paleontology, University of Ne­ although the writers realize that only the Kimballian may be braska State Museum, and Professor of Geology, Depart­ equivalent to the latest Miocene of Europe. See Part 1, p. 2, ment of Geology. of the present Bulletin for further remarks on this subject. 56 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

SYSTEMATIC DESCRIPTIONS

Class: MAMMALIA Order: CARNIVORA Family: FELIDAE Gray, 1821 Subfamily: Machairodontinae Gill, 1872 Genus: Machairodus Kaup, 1833

Machairodus aphanista (Kaup)

Geographic Distribution.-North America, Eurasia, and Africa. Amended Diagnosis.-Scimitar-toothed cats about the size of a lion (Panthera leo); skull with a relatively elongate face; upper canines large, blade-like, and coarsely serrated; lower canine greatly reduced and incisiform; upper and lower incisors evenly spaced and arranged in a semicircle; upper carnassial with protocone re­ duced; metaconid very reduced or lost on lower carnassial; P2 usually absent; coronoid process on ramus low; limbs elongate; and feet digiti­ grade.

Machairodus cf. coloradensis Cook, 1922

Machairodus catocopis Matthew, 1924; Burt, 1931; Savage, 1941; Mawby, 1965; Shotwell, 1956; and Dalquest, 1969.

Type Locality.-Near Wray, Yuma County, Colorado. Fig. 1 -A, Machairodus cf. c%radensis, U.N.S.M. 25510, right ramus (crown and labial views), and B, right canine Stratigraphic Occurrence.-Middle Pliocene, (labial view) from the upper part of the Ash Hollow Forma­ Ogallala Group, upper part of Ash Hollow Forma­ tion, Sherman County, Nebraska; C, Nimravides catocopis, tion. U.N.S.M. 21819, distal end of humerus, from the upper part of the Ash Hollow Formation, Smith County, Kansas. X 1/2. Referred Specimen.-Right CJ; right ramus with !1-2(alv.), 13, IC, P3-4(alv.), Ml, U.N.S.M. 25510 (Fig. 1, A and B). Amended Diagnosis.-Scimitar-toothed cat intermediate in size between jaguar and lion; Locality.-U.N.S.M. Coil. Loc. Sm-101, SV2, dorsal surface of skull rounded and occiput in­ SW. V4, sec. 13, T. 16N., R. 13W. East of Ashton, clined as in feline cats; upper canine flattened Sherman County, Nebraska. and crenulated on anterior and posterior edges; Stratigraphic Occurrence.-Middle Pliocene p3 double-rooted and relatively large with a large (Hemphill ian), Ogallala Group, Ash Hollow For­ posteriorly directed paracone, well-developed mation. parastyle and metacone of about equal size; p4 SCIMITAR-TOOTHED CATS, MACHA/ROD US AND N/MRAVIDES, FROM THE PLIOCENE / 57

Fig.2-Machairodus c%radensis tanneri, new subspecies, holotype, U.N.S.M. 26086, right ramus (labial and lingual views), from the Kimball Formation, Cheyenne County, Nebraska. large with a prominent anterior cusp on the of Agriotherium. The skeleton of the North parastyle (absent or very small in Nimravides), American Machairodus has been described by protocone well developed, metacone elongate Dalquest (1969) and shows the long-legged and blade-like; M' small (absent in Nimravides); form, typical of scimitar-toothed cats. The lower ramus with anterior margin inclined; coronoid canine is in place in U.N.S.M. 25510 and has the process low (higher in Nimravides); anterior form illustrated by Burt (1931, pI. 45) contrary mental foramen larger than in Nimravides; in­ to Dalquest (1969, p. 16). The species of cisors graded in size becoming larger from 11-3, Machairodus need a thorough revision, and a recurved and serrated on both edges (crowded good series of the North American forms should in Nimravides); lower canine reduced in size, in­ be studied. The four rami of M. coloradensis cisiform, and serrated anteriorly and posteriorly; reported by Cook (1922) show considerable vari­ P3-4 large and double-rooted; M, larger than in ation in size and morphology. The smaller Nimravides and more posteriorly located, ramus, No. 207, that he illustrates (Cook, 1922, p. paraconid almost as large as protoconid; 24) resembles U.N.S.M. 25510 in the lack of a metaconid very small or absent. dependent flange on the ramus, the lack of de­ velopment of the metaconid on M" and in its Discussion.-The diagnosis is based in part relatively small size. The other rami illustrated on figures in Cook (1922) and Burt (1931). are much larger and appear to have prominent Cook's figures are of poor quality and it is dif­ metaconids on M,. ficult to ascertain the nature of the metaconid on M, which he mentions. It is absent from all other specimens of Machairodus that have been de­ Machairodus c%radensis tanneri,4 new sub­ scribed from North America (Mawby, 1965). The species upper canine of U.N.S.M. 25510 has large crenu­ lations on its anterior and posterior margins. It Holotype.-Right ramus with 11-3 alv., IC rt., seems well adapted for slashing and contrasts P3-M, broken, U.N.S.M. 26086; Figs. 2, 3A. with the stabbing saber with fine crenulations found in Barbourofelis. U.N.S.M. 25510 was 4Named in honor of Lloyd G. Tanner who has contributed much to a found in association with undetermined better understanding of the Kimballian fauna and the Kimball Forma­ rhinoceros remains and an undescribed species tion. 58 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

E

~------~------

Fig. 3 -A, Machairodus c%radensis tanneri, new subspecies, holotype, U.N.S.M. 26086, ramal symphysis (anterior view), from the Kimball Formation, Cheyenne County, Nebras ka; B, D, F, G, Nimravides catocopis referred: B, U.N.S.M. 21818, mandibular symphysis (anterior view), from Smith Cou nty Kansas; D, U.N.S.M. 21817, partial left maxillary (labial view), from Smith County, Kansas; F, U.N.S.M. 21818, partial left ramus (labial view), P3-4 and the side of the ramus below the cheek teeth restored from the left side of the mandibl e, from Smith County, Kansas; G, U.N.S.M. 45116, left ramus (labial view), from Frontier County, Nebraska; C, E, Nimravides catocopis, holotype, A.M.N.H. 8318, from Phillips County, Kansas; C. mandibular symphysis (anterior view); E, mandibular symphysis, (labial view). X 1/2. SCIMITAR-TOOTHED CATS, MACHAIRODUS AND NIMRAVIDES, FROM THE PLIOCENE I 59

A c

Fig. 4 -Outlines of right incisor and canine alveolae: A, Machairodus cf. coloradensis, U.N.S.M. 25501, from the upper part of the Ash Hollow Formation, Sherman County, Nebraska; Band C, Nimravides catocopis, B, holotype, A.M.N.H. 8318, from Phillips County, Kansas; C, referred, U.N.S.M. 21818, from Smith County, Kansas. X 1.

Type Locality_-U.N.S.M. Coil. Loc. Cn-101, Colorad 0 (Cook, 1922), and from the Coffee NW. Y4, sec. 23, T. 15N., R. 49W., 41/2 miles south Ranch Local Fauna at Hemphill, Texas (Burt, and 31/2 miles east of Gurley, Cheyenne County, 1931; Dalquest, 1969). It seems to have a better Nebraska. developed dependent flange on the ramus and to differ from M. c. coloradensis in lacking the Stratigraphie Occurrence.-Upper Pliocene metaconid on Ml. It may also have a lower (Kimballian), Ogallala Group, Kimball Formation. coronoid process, although damage to U.N.S.M. Diagnosis.-A scimitar-toothed felid with a 26086 makes it impossible to determine this with well-developed, dependent flange on the ramus certainty. and two roots on P3. Subfamily: Felinae Trouessart, 1885 Description.-Mandible very elongate; coronoid process low; masseteric fossa large, deep, and extending under middle of Ml; angu­ Nimravides Kitts, 1958 lar process an elongated knob; flange excavated Type Species.-Nimravides thinobates for saber; contour of dependent flange similar to (Macdonald), 1942. that in Ischyrosmilus; anterior border of sym­ physis square with two large foramina as in Geographic Distribution.-California, Texas, Ischyrosmilus; alveolae show incisors slightly Oklahoma, Kansas, and Nebraska. crowded; IC root flattened laterally; diastema Amended Diagnosis.-Very large pseudaelu­ relatively long; P3 with two roots; P4 slightly rin cats; anterior buccal side of ramus flattened tilted posteriorly; Ml moderately large. and excavated in a manner analogous to the Discussion.-Machairodus coloradensis tan­ condition in Machairodus; ventral symphysial neri is highly advanced in terms of the develop­ extension projecting further ventrally than in ment of a distinct dependent flange on the Pseudaelurus; p4 with elongate (blade-like) ramus. It mig ht make a good ancestor for metacone, and large parastyle; upper and lower Ischyrosmilus, from which it differs in having P3 carnassial notches more shallow than in much less reduced and the mandible more elon­ Pseudaelurus but not as shallow as in gate. It would have been contemporaneous with Machairodus; lower carnassial having the giant dirk-toothed cat Barbourofelis fricki, metaconid but usually lacking talonid and Machairodus coloradensis tanneri might be (metaconid absent or extremely vestigial in represented by some undescribed skeletal ele­ North American Machairodus); lower carnassial ments from U.N.S.M. Coil. Loc. Ft-40, the type relatively smaller than in Machairodus; lower in­ locality for Barbourofelis fricki. cisors small and crowded as in Pseudaelurus In size U.N.S.M. 26086 falls well within the (larger and more evenly spaced in Machairodus); range given for Machairodus from Yuma County, anterior margin of ramus less prognathous than 60 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Machairodus sp.: Burt, 1931. Machairodus coloradensis: Savage, 1941.

Holotype.-Symphysis of a mandible with 11-3(alv.), C (br.); A.M.N.H. 8318 (Fig. 3, C and E). Type Locality.-Republican River beds, Phil­ lips County, Kansas. Stratigraphie Occurrence.-Pliocene (?Hem­ phillian), Ogallala Group, Ash Hollow Formation. Referred Specimens.-Partial maxilla with p3·4, U.N.S.M. 21817 (Fig. 3,0); partial mandible with 11·3(alv.), IC(br.), P2(alv.)-Ml, U.N.S.M. 21818 (Fig. 3, B, F; 4, C); partial right ramus with 11·3(alv.), IC(alv.) and P3(alv.), U.N.S.M. 21816; dis­ tal end of humerus, U.N.S.M. 21819, all from Smith County, Kansas; and right ramus with h·3(alv.),/C, P3(alv.), P4, Ml(alv.), U.N.S.M. 45116 (Fig. 3, G), from upper part of the Ash Hollow Formation, Frontier County, Nebraska. Localities.-U.N.S.M. Coil. Loc. Kan-1, Smith County, Kansas, and U.N.S.M. Coil. Loc. Ft-47, upper part of Ash Hollow Formation, Frontier County, Nebraska (more precise locality and stratigraphic data is available from the locality files of the University of Nebraska State Museum). Stratigraphic Occurrence.-Pliocene (Hem­ c phillian), Ogallala Group, upper part of Ash Hol­ low Formation. Diagnosis.-Not distinguished from ·generic diagnosis. The only other described species, Fig. 5 -Restoration of outlines of skulls: A, Machairodus coloradensis (based on Cook, 1922); B, Nimravides thino­ Nimravides thinobates (Macdonald), is a proba­ bates (based on Kitts, 1958, PI. 1 and Macdonald, 1948); C, ble synonym. Dinofelis diastemata (after Hemmer, 1965, Fig. 36). Approx­ imately X 1/4. Discussion.-The holotype of Nimravides catocopis has the anterior-buccal margin to the in Machairodus; coronoid process high as in ramus less excavated, the ventral margin of the Pseudaelurus but with broader base and more symphysis less angular, the canine larger, and recurved posteriorly than in Machairodus; mas­ the incisors smaller and more crowded than in seteric fossa larger and with a thinner ventral Machairodus. It resembles Nimravides in all of border than in Pseudaelurus. these features. The partial mandible, U.N.S.M. 21818, is as close to being a topotype of N. Nimravides catocopis (Cope), 1887, new combi­ catocopis as we can reasonably hope to find nation with the data available. It agrees with the holotype on almost all visible features. It demon­ Machairodus catocopis: Cope, 1887; Hesse, strates that the carnassial was smaller and not as 1940. posteriorly placed as in Machairodus coloraden- SCIMITAR-TOOTHED CATS, MACHA/ROD US AND N/MRAV/DES, FROM THE PLIOCENE I 61 sis. The lower canine of Nimravides does not expressions of a persistent tendency in feline have the extensive anterior and posterior serra­ cats to revert to the saber-toothed condition. The tions found in Machairodus (Burt, 1931, PI. 45). living cloud leopard (Neofelis nebulosus) The P2 was present in U.N.S.M. 21818 but was represents another example. In North America, absent in U.N.S.M. 21816 from the same locality. Machairodus may have given rise to The P3 is a large tooth with a well developed Ischyrosmi1us, a genus which would then be paraconid, protoconid, and talonid. The valid and should not be put into synonomy with paraconid is very large on P4 while M1 has a Homotherium as has been suggested (Mawby, small metaconid and no talonid. p3 has a large 1965; Churcher, 1968).The P2 of Nimravides was parastyle, paracone, metacone, and a well­ a small Single-rooted peg-like tooth which seems developed posterior cingular cusp as described to have been present in a fairly high percentage in Kitts (1958, p. 373). The upper carnassial is of the population, and also often occurs in badly damaged, but seems to have been similar Pseudaelurus. to that illustrated by Macdonald (1948, Fig. 14). Kitts (1958) includes a substantial amount of The M1 is unfortunately not preserved. size variation in his concept of N. thinobates. If The lower carnassial referred by Savage (1941, this amount of variation is accepted, N. thino­ p. 697) to Machairodus (Heterofelis) coloraden­ bates (Macdonald) may be a synonym of N. sis on the basis of its prominent metaconid may catocopis (Cope). All of the specimens from for that same reason be better placed in Smith County, Kansas, are near the size of the Nimravides catocopis, while the specimen lack­ holotype of N. catocopis and the smaller speci­ ing the Metaconid from Optima may be a men from Arnett, but the specimen from Machairodus. U.N.S.M. Coil. Loc. Ft-47 is near the size of the larger specimen from Arnett. CONCLUSION Kitts did not compare Nimravides with the es­ sentially similar machairodontoid pseudaelurin The typical scimitar-toothed cat of the cat of Eurasia, Dinofelis. Dinofelis does not usu­ Pliocene is Machairodus. It apparently origi­ ally have a well-developed metaconid on the nated in Eurasia from the Miocene scimitar­ lower carnassial as does Nimravides. It does toothed cat Nimravus and spread through most have the anterior margin of the symphysis more of the Northern Hemisphere. The time of its arri­ inclined. Dinofelis barlowi has a large metaconid val in North America is presently uncertain, and on M1. Both genera have large lower canines, we are not aware of any records older than Hem­ high coronoid processes, and generally similar phillian. It seems probable that it migrated to skull configuarations (Fig. 5). It appears that North America at the same time as the Nimravides, like Dinofelis, lacks serrations on postorbital-bar cats, the Barbourofelini. The lat­ the upper canine, while Machairodus and ter first appear in North America during the Ischyrosmilus have coarsely serrated upper Clarendon ian (Schultz, Schultz, and Martin, canines. Dinofe/is has the auditory bulla sepa­ 1970). Although several species of North Ameri­ rated into two parts, as do other pseudaelurin can felids have been referred to Machairodus, cats (Piveteau, 1948). The auditory bulla of only M. coloradensis Cook seems to be valid. Nimravides has not been described although we Machairodus hesperus Gazin from the Early would expect it to be similar to that found in Pleistocene of Idaho has been shown to be a Dinofelis. It is possible that these cats may be Megantereon (Schultz and Martin, 1970), closely related although they are widely sepa­ Machairodus niobrarensis (Thorpe) is a bar­ rated in the phylogenetic tree published by bourofelin cat,S and M. catocopis Cope belongs Thenius (1967, Fig. 1). Kurten (1972) has shown to the genus Nimravides Kitts. that Panthera palaeoonca Meade is a species of Nimravides is closely similar to the Eurasian Dinofelis. The latter genus is now known from machairodontoid cat Dinofelis. Both genera are the Blancan of North America and the Villafran­ chian of Eurasia and Africa. It is possible that it SMartin, manuscript in preparation. was derived from Nimravides in North America 62 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

TABLE 1 Machairodus and Nimravides COMPARATIVE MEASUREMENTS' OF MANDIBULAR RAMI

Machairodus coloradensis Nimravides MANDIBULAR RAMI tanneri, n. subsp. Nimravides catocopis Holotype M. coloradensis catocopis U.N.S.M.45116 U.N.S.M. 26086 U.N.S.M.25510 U.N.S.M.21818 U.N.S.M.21817

Distance between alveoli for IC and P3 ...... 39 29.5 26.3 40.2 Depth of ramus posterior to M, ...... 44.1 39.2 37.6 49.3 Thickness of ramus below M, ...... 19.2 18.2 18.7 23 Height from inferior border of angle to summit of condyle ...... 40.7 42.3 (45)2 Height from inferior border of angle to summit of coronoid process ...... (73.6) (103.6) Transverse width of condyle ...... 42 h greatest anteroposterior diameter ...... (12.5) (9.6) I, greatest transverse diameter ...... (4.7) (3.5) 12 greatest anteroposterior diameter ...... (13.8) (9.6) 12 greatest transverse diameter ...... (7.4) (5.2) b greatest anteroposterior diameter ...... 9.5 b greatest transverse diameter ...... 9.1 IC greatest anteroposterior diameter at base of enamel ...... (19.2) 15.5 20.5 20 IC greatest transverse diameter ...... 12.8 13.2 12 12.1 Length P3-M, ...... 78.8 77.6 65.8 P2 anteroposterior diameter ...... 3.6 P2 greatest transverse diameter ...... 3.0 P3 anteroposterior diameter ...... (19.6) (20) 17.1 (19.2) P3 greatest transverse diameter ...... (7.9) (7.8) 8.3 P4 anteroposterior diameter ...... (26.3) (26.4) 21.9 23.9 P4 greatest transverse diameter ...... (10.5) (10.2) 9.4 10.7 M, anteroposterior diameter ...... (32.0) 31.9 25.4 (28.6) M, greatest transverse diameter ...... (12.6) 14 11.6 M, length of protoconid blade ...... 13.5 12.4 UPPER DENTITION CI greatest anteroposterior diameter ...... 38.4 CI greatest transverse diameter ...... 15.8 p3 anteroposterior diameter'...... 21.4 p3 greatest transverse diameter ...... ' ... . 10.0 . p4 anteroposterior diameter ...... : ...... 31.3 p4 greatest transverse diameter ...... 13.6

'The measurements are taken to the nearest millimeter except on dentition where they are measured to the nearest one tenth of a millimeter 2( ) = alveolar or approximate measurement SCIMITAR-TOOTHED CATS, MACHA/RODUS AND N/MRAVIDES, FROM THE PLIOCENE I 63 and then migrated to Eurasia during the Blancan Cope, E. D. 1887. A saber-tooth tiger from the Loup Fork. as did the early lynxes, Amer. Nat. 21: 1019-1020. Dalquest, W. W. 1969. Pliocene Carnivores of the Coffee Ranch (Type Hemphill) Local Fauna. Bull. Texas Memorial ACKNOWLEDGMENTS Mus. (15): 1-43, Figs. 1-11, 1 plate, 31 tables. Hemmer, H. 1965. Zur Nomenklatur und Verbreitung des The writers are grateful to Messrs. Dick Her­ Genus Dinofelis Zdansky, 1924 (Therailurus Piveteau, man and Leonard Egging for the discovery of the 1948). Paleont. Africana 9: 78-89, 4 figures. holotype of Machairodus coloradensis tanneri, Hesse, Curtis J. 1940. A Pliocene vertebrate fauna from Hig­ U.N.S.M. 26086, and for reporting the fossil gins, Lipscomb County, Texas. Univ. Texas Publ. 3945: 671-698,8 figures. quarry, U.N.S.M. Coli. Loc. Cn-101, from Kitts, D. B. 1958. Nimravides, a new genus of felidae from the Cheyenne County to the Museum's Division of Pliocene of California, Texas and Oklahoma. Jour. Mam­ Vertebrate Paleontology. They are also thankful mal. 39: 368-375 to Mr. Dale Anderstrom for the donation of the Kurten, B. 1972. The genus Dinofe/is (Carnivora, Mammalia) Machairodus specimen, U.N.S,M. 25510, from in the Blancan of North America. Univ. Texas Bull., Pearce-Sellards Ser. (19); 1-7, 1 figure, 1 table. Sherman County. The Smith County (Kansas) Macdonald, J. R. 1948. The Pliocene carnivores of the Black specimens, U.N.S.M. 21816, 21817, 21818, and Hawk Ranch. Univ. California Publ. Geol. Sci. 28: 53-80, 15 21819, were collected in 1931 by a University of figures. Nebraska State Museum field party consisting of Matthew, W. D. 1924. Third contribution to the Snake Creek Messrs. Emery L. Blue, Frank Crabill, Eugene Fauna. BUll. Amer. Mus. Nat. Hist. 50 (2): 59-210, 63 fig­ ures. Vanderpool, and C. Bertrand Schultz; and the Mawby, J. E. 1965. Machairodonts from the Late Cenozoic of Frontier County (Nebraska) specimen, U.N.S.M. the Panhandle of Texas. Jour. Mammal. 46(4): 573-587, 45116, was collected in 1948 by a U.N.S.M. field Figs. 1-5. party which was under the direction of Messrs. Piveteau, J. 1948. Un Felide du Pliocene de Rousillon. Ann. W. D. Frankforter and C. Bertrand Schultz. Mr. Paleont. 34: 97-124. Savage, D. E. 1941. Two new middle Pliocene Carnivores Jerry Tanner prepared the illustrations for Figs. from Oklahoma, with notes on the Optima fauna. Amer. 1, 2, 3A, and 3D, and Mrs. Mary Tanner was re­ Midland Nat. 25(3): 672-710, Figs. 1-40,4 plates. sponsible for the drawings in Figs. 4 and 5, and Schultz, C. B., and L. D. Martin. 1970. Machairodont cats also 3B, 3C, 3E, 3F, and 3G. The writers are from the Early Pleistocene Broadwater and Lisco Local grateful to Mrs. Mary Tanner and Mr. George faunas. Bull. Univ. Nebraska State Mus. 9(2): 1-36, Figs. 1-2, Frontispiece. Corner for their help in the preparation of the Schultz, C. B., M. R. Schultz, and L. D. Martin. 1970. A new manuscript. tribe of saber-toothed cats (Barbourofelini) from the Pliocene of North America. Bull. Univ. Nebraska State Mus. REFERENCES 9(1): 1-31, Figs. 1-13, Frontispiece. Shotwell, J. A. 1956. Hemphillian mammalian assemblage Burt, W. H. 1931. Machaerodus catocopis Cope from the from northeastern Oregon. Bull. Geol. Soc. Amer. 67: Pliocene of Texas. Univ. California Publ., Bull. Dept. Geol. 717-738, Figs. 1-7. Sci. 20(7): 261-292, 8 plates. Thenius, E. 1967. Zur Phylogenie der Feliden (Carnivora, Churcher, C. S. 1966. The affinities of Dinobastis serus Cope Mamm.). Zeitschrift Zool. Syst. Evolutionsforschung 5(2): 1893. Quaternaria (8): 263-275. 129-143, 1 fig ure. Cook, H. J. 1922. A Pliocene fauna from Yuma County, Col­ orado, with notes on the closely related Snake Creek Beds of Nebraska. Proc. Colo. Mus. Nat. Hist. 4(2): 3-15, 14 fig­ ures. THE BOARD OF REGENTS

Robert R. Koefoot, M.D., chairman Mrs. J. G. Elliott Kermit Hansen James H. Moylan Robert J. Prokop, M.D. Robert L. Raun Edward Schwartzkopf Kermit Wagner Ralph H. Bradley, corporation secretary

THE PRESIDENT OF THE UNIVERSITY OF NEBRASKA

D. B. Varner

THE CHANCELLOR OF THE UNIVERSITY OF NEBRASKA-LINCOLN

James H. Zumberge

THE COMMITTEE ON SCHOLARLY PUBLICATIONS

Frederick M. Link, chairman Ned S. Hedges Warren W. Caldwell C. Bertrand Schultz Virginia L. Faulkner Gerald E. Thompson