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Novltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICANt MUSEUM Novltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2984, 60 pp., 18 figs., 5 tables, 2 appendices October 23, 1990 Systematic Revision of Chinese Hipparion Species Described by Sefve, 1927 RAYMOND L. BERNOR,1 ZHANXIANG QIU,2 AND LEE-ANN C. HAYEK3 ABSTRACT Sefve's type series of Chinese Neogene hippar- refer it here to "H." forstenae (sensu Zhegallo, ionine horses, in the Palaeontologiska Museet, 1971). We recommend that four of Sefve's (1927) Uppsala, is studied, along with relevant specimens original taxa be considered nomina dubia: Hip- housed by the American Museum ofNatural His- parion kreugeri, Hipparion plocodus, Hipparion tory Frick Laboratories. Morphological and sta- tylodus, and Hipparion parvum. The phylogenetic tistical analyses have been made of skull and di- relationships of the valid species are discussed, rectly associated mandibular material. The and these species are found to represent parts of morphological analysis, using multiple states for other known and some new lineages. The species 40 characters, identifies 11 hipparion groups in "Hipparion" platyodus, "Hipparion" coelophyes the assemblage. The statistical analysis documents (s.l.), and "Hipparion" hippidiodus are found to range of variability in these morphological group- represent a morphoclinal series in China, but the ings and compares range of variability and size to coexistence of"H." coelophyes (s.l.) and "H." hip- the Howenegg (late Miocene, Hegau, Germany) pidiodus at the same localities and the later strati- quarry sample. These analyses guide us in vali- graphic occurrence of"H." platyodus provide evi- dating six of Sefve's (1927) original species: Hip- dence which rejects the hypothesis of their parion platyodus, Hipparion coelophyes, Hippar- chronospecific relationship. "Hipparion" platyo- ion hippidiodus, Hipparion ptychodus, Hipparion dus, "Hipparion" ptychodus, and Proboscidippa- dermatorhinum, and Proboscidipparion sinense. rion sinense are cited as being potentially related We follow Zhegallo's (1971) recommendation that to the "Sivalhippus Complex" which has a geo- Sefve's (1927) species Hipparion richthofeni be graphic distribution including mostly eastern and recognized as a taxon distinct from Koken's orig- southern Asia and Africa. "Hipparion" dermator- inal material ofH. richthofeni from Mongolia; we hinum is believed potentially to be a monospe- ' Raymond L. Bemor, College of Medicine, Department of Anatomy, Laboratory of Paleobiology, Howard Uni- versity, Washington D.C. 20059; Department of Paleobiology, Smithsonian Institution, Washington D.C. 20010. 2 Zhanxiang Qiu, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing 10004, China. 3Lee-Ann C. Hayek, Department of Mathematical Statistics, Smithsonian Institution, Washington D.C. 20010. Copyright © American Museum of Natural History 1990 ISSN 0003-0082 / Price $5.50 2 AMERICAN MUSEUM NOVITATES NO. 2984 cific lineage unique to northern Asia. "Hipparion" tribution including China, western U.S.S.R., forstenae is found to be a member of the Cre- Greece, and Iran. mohipparion lineage, which had a geographic dis- INTRODUCTION Hipparionine horses have long been rec- revised Sefve's (1927) initial work on the ognized as valuable index fossils for age dis- Uppsala assemblage. Because this collection crimination of late Neogene Old World con- is of key importance for all subsequent sys- tinental deposits. Recently there has been a tematic work on Chinese hipparionines, and renewed interest in this group, and a glance because there are accurate locality data as- over the contemporary literature suffices to sociated with some of it, we believe that it is show how much progress has been made in timely to revise the alpha systematics of the understanding this group's evolutionary his- Uppsala collection and, building on the work tory. New discoveries have been made, es- by Qiu et al. (1987), to further develop hy- pecially as regards Old World early late Mio- potheses of these species' evolutionary rela- cene forms (Vallesian age of European tionships. chronology). Many of the old museum col- Three other major assemblages of Chinese lections from classical localities such as Sa- hipparionines currently are housed at the mos, Pikermi, Maragheh, and the Siwalik American Museum of Natural History, New Hills, Pakistan, have been restudied and re- York; the Institute of Vertebrate Paleontol- vised (for a review see Bernor and Hussain, ogy and Paleoanthropology, Beijing; and the 1985; Bemor et al., in press; Woodbume, Tianjin Museum, Tianjin. Currently, ourjoint 1989). Since publications by Skinner and project is to prepare and describe the Amer- MacFadden (1977) and Woodbume and Ber- ican Museum Chinese hipparion collections. nor (1980), a number of the American pa- We provide here descriptions ofthose AMNH leontologists have become increasingly in- specimens that have recently been prepared, volved in formulating a superspecific and are referable to species of the Uppsala classification ofhipparionine horses. With the type series. notable exception ofwork by Qiu et al. (1987), hipparionine research on Chinese assem- blages has remained comparatively dormant. ABBREVIATIONS AND DEFINITIONS We believe that one of the principal reasons lies in difficulties of extending the existing Institutional nomenclature of the Uppsala type collection AMNH Department of Vertebrate Paleontology, to other, newer Chinese discoveries. American Museum of Natural History, Sefve's (1927) work is the first and only New York detailed description of the Uppsala hippar- F:AM Frick fossil mammal collection, AMNH IVPP Institute of Vertebrate Paleontology and ionine collection. Sefve described 44 skulls, Paleoanthropology, Beijing, China some associated with lowerjaws, and a num- PMU Palaeontologiska Museet, Uppsala, Swe- ber ofisolated jaws and limb bones. Whereas den the main portion ofthis assemblage was from Baode, there were some specimens from the Anatomical Wuxiang-Yushe and Qingyang areas (see fig. MC III metacarpal III 1 and table 1). Altogether Sefve (1927) de- MT III metatarsal III scribed 11 new species and one new genus of POB preorbital bar hipparionine horses. While Sefve's work was POF preorbital fossa a milestone in the systematic organization of the assemblage, it contains critical flaws. Sefve Hipparionine or hipparion -Any horse with designated no type specimens, made no an isolated protocone of the maxillary pre- species diagnoses, paid little or no attention molar and molar teeth, and, as far as known, to ontogenetic variation (especially of cheek tridactyl feet, including species ofthe follow- teeth), and entirely ignored the assemblage's ing genera: Hipparion, Neohipparion, Nan- geological age. No author has yet properly nippus, Cormohipparion, Hippotherium, 1990 BERNOR, QIU, HAYEK: CHINESE HIPPARION SPECIES 3 Fig. 1. Map of principal collecting areas of Uppsala Chinese hipparionines. A. Gansu (= Kansu) Province; Qingyang (= King-Yang) County; Locality 115. B. Shanxi (= Shansi) Province; Baode (= Pao- Te) County and Hequ (= Hochii) County; Localities 30, 31, 43, 44, 49, 52, 109, 110, and 114. C. Shanxi Province (= Shansi); Yushe-Wuxiang (= Yii-she-Wu-Hsiang) County; Localities 70 and 73. D. Henan (= Honan) Province; Xin-an (= Hsin-An) County; Localities 12 and 39; E. Shaanxi (= Shensi) Province; Fugu (= Fu-ku) County; Locality 51. Proboscidipparion, "Sivalhippus," Pseudhip- on the face. The posterior pocket becomes parion, Stylohipparion, and Cremohipparion. reduced and eventually lost, and confluent Recent characterizations ofthese taxa can be with the adjacent facial surface (includes found in MacFadden (1984), Bemor and Group 3 of Woodburne and Bernor, 1980). Hussain (1985), Webb and Hulbert (1986), We differ with some of the previous authors Hulbert (1987), Qui et al. (1987), Bemor et in the specific referrals ofHipparion s.s.; this al. (in press) and Woodburne (1989). issue has most recently been discussed in de- Hipparion s.s.-We follow MacFadden tail by Bemor et al. (in press). (1980, 1984), Woodburne and Bemor (1980), "Hipparion"-Following Woodburne and Woodburne et al. (1981), McFadden and Bemor (1980), MacFadden and Woodburne Woodburne (1982), Bemor and Hussain (1982), Bemor and Hussain (1985), Bemor (1985), Bernor (1985), Bernor et al. (1987), (1985), Bemor et al. (1988), and Bemor et al. Bernor et al. (in press), and Woodburne (1989) (in press): Old World hipparion horses with in restricting this nomen to a specific lineage facial morphologies that differ from Hippar- of horses that have a fossa positioned high ion s.s. (includes superspecific taxa listed 4 AMERICAN MUSEUM NOVITATES NO. 2984 TABLE 1 Locality Distribution of the Uppsala Chinese Hipparionine Species Geo- Sam- Locality graphic pling numbera locationh method, Taxa present (this report)d 12 D 2 "H." plocodus 30 B 1 "H." forstenae, "H." dermatorhinum, "H." ?dermatorhinum, hipparionine gen. and sp. indet. 31 B hipparionine gen. and sp. indet. 39 D 2 Proboscidipparion sinense 43 B "H." coelophyes, "H." aff. coelophyes, "H." hippidiodus, "H. plocodus," hipparion- ine gen. and sp. indet. 44 B "H." coelophyes, "H." hippidiodus 49 B "H. plocodus" 51 E 2 "H." ?dermatorhinum 52 B "H. kreugeri," "H." l hippidiodus 70 C 2 "H." platyodus 73 C 2 "H." ptychodus, "H. tylodus" B 109 2 "H." aff. coelophyes 110 B "H." 2 ?derpratorhinum 114 B "H." ptychodus 115 A "H." hippidiodus A "H. parvum" a Localities as used by Sefve and referred
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