A Cretaceous Hoofed Mammal from India G. V. R. Prasad, 1 * O. Verma, 1 A. Sahni, 2 V. Parmar, 1 A. Khosla 2 U ngulate or hoofed mammals represent the most important herbivores of Ceno- zoic land mammal communities. Condy- larths or archaic ungulates are a paraphyletic group regarded as an ancestral or sister taxon to living ungulate mammals (1) and are known by fossils from the Early Paleocene of North America and South America, the Late Paleocene and Eocene of Europe, and the Eocene of Africa and possibly Australia. The earliest definitive ungulates represented by Protungulatum, Oxy- primus, Baioconodon, and Mimatuta (1, 2) come from the Early Paleocene (Puercan) of north- eastern Montana. No Cre- taceous condylarth has yet been documented. Here, we describe an isolated lower molar (m 1 or m 2 ) of a condylarth mammal from the Late Cretaceous (Maastrichtian) lacustrine rocks interbedded with Deccan volcanic flows of Central India [Supporting Online Material (SOM) text and fig. S1]. The tooth is cataloged as VPL/JU/IM/31 and deposited in the Verte- brate Paleontology Labora- tory of Jammu University. Class: Mammalia Lin- naeus, 1758. Order: Con- dylarthra Cope, 1881. Family: Incertae Sedis. Kharmerungulatum vanvaleni genus et species nova. Holotype. VPL/JU/ IM/31, isolated right lower molar. Generic Diagnosis. Asymmetrical trigonid wider and longer than talonid, bul- bous cusps, paraconid slight- ly labial to the lingual margin, voluminous proto- conid twice as large as meta- conid and closely appressed to it, metaconid slightly posterior to protoconid with the posterior trigonid wall slightly oblique to the long axis, entoconid smaller than hypoconulid and basally conjoined to it forming an oblique posterolingual crest, hypocon- id is the most voluminous talonid cusp, talonid basin partially closed lingually, obliquely trans- verse talonid groove, size smaller as compared to other archaic ungulates (SOM text). Etymology. Genus named after the nearby Kharmer River; species named in honor of Leigh Van Valen. Specific Diagnosis. As for genus. Basal expansion and sidewall convexity of the lower molars are conspicuous differences between archaic ungulates and Cretaceous eutherians such as Cimolestes , Procerberus , Batodon, Gypsonictops , and Deccanolestes . Archaic ungulates developed the ability to crush and grind food through reduced height difference between the trigonid and talon- id, cusp bunodonty, the possession of a large hypoconid, and characteristic abrasion causing beveling of cusp apices (1). The morphology of Kharmerungulatum (Fig. 1, A to F) is like that of the lower molar morphology of archaic ungulates, but the latter are relatively larger and have more bunodont cusps. Kharmerungulatum retains many plesiomorphic characters that occur variably in Protungulatum, Oxyprimus, Baioconodon, and Mimatuta. These include trigonid moderately taller than the talonid; paraconid slightly labial to the lingual margin and well separated from the metaconid, as in Mimatuta, Protungulatum gor- gun, and Baioconodon middletoni; obliquely oriented paracristid, as in Protungulatum and Oxyprimus; obliquely oriented posterior trigonid wall with a slightly posteriorly developed meta- conid and asymmetrical trigonid, as in Protungu- latum donnae; short entocristid partially closing the talonid lingually, as in Protungulatum, Oxy- primus, and Baiconodon; and a cristid obliqua joining the posterior trigonid wall below the posterolabial base of the metaconid. However, in relative dimensions of talonid cusps and closely appressed entoconid and hypoconulid forming a posterolingual crest, Kharmerungulatum is also comparable to South American Molinodus (3). Among all species of archaic ungulates, Kharmerungulatum is closest to P. gorgun in cusp morphology except that it lacks cingulids [figure 3F of (4)] and to Baioconodon in having a talonid shorter and narrower than the trigonid (SOM text). We consider Kharmerungulatum to represent an early stage in the evolution of ungulates. The presence of an archaic ungulate in the latest Cretaceous of India may reflect that (i) archaic ungulates had a pan-Gondwanan distri- bution, and their absence in other landmasses may be an artifact of limited field investigations; (ii) Kharmerungulatum immigrated to India from Western Asia, which had a diversified assem- blage of zhelestids (85 million years ago) re- garded by some as ancestral to archaic ungulates (5), although any connection is disputed (6); and (iii) the drifting Indian subcontinent may have served as a center of origin for many mammalian orders and other vertebrate and plant groups. References and Notes 1. J. D. Archibald, Univ. Calif. Publ. Geol. Sci. 122, 1 (1982). 2. L. Van Valen, Evol. Theory 4, 45 (1978). 3. C. de Muizon, R. L. Cifelli, Geodiversitas 22, 47 (2000). 4. J. J. Eberle, J. A. Lillegraven, Rocky Mount. Geol. 33, 49 (1998). 5. J. D. Archibald, Science 272, 1150 (1996). 6. J. R. Wible, G. W. Rougier, M. J. Novacek, R. J. Asher, Nature 447, 1003 (2007). 7. G.V.R.P. acknowledges support from the Department of Science and Technology (DST), New Delhi (grant no. SR/S4/ES/24/2002). A.S. thanks the Indian National Science Academy for grant no. SP/SS/2006/2841, and O.V. and A.K. are thankful to the Council of Scientific and Industrial Research, New Delhi and DST, respectively. Supporting Online Material www.sciencemag.org/cgi/content/full/318/5852/937/DC1 SOM Text Fig. S1 References 15 August 2007; accepted 27 September 2007 10.1126/science.1149267 BREVIA 1 Department of Geology, University of Jammu, Jammu 180 006, India. 2 Centre of Advanced Study in Geology, Panjab University, Chandigarh 160 014, India. *To whom correspondence should be addressed. E-mail: guntupalli.vrprasad@gmail.com Fig. 1. Holotype of Kharmerungulatum vanvaleni genus et species nova, isolated right lower molar (m 1 or m 2 , VPL/JU/IM/31). End, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; oc, cristid obliqua; pad, paraconid; pcc, precingulid cuspule; and prd, protoconid. (A) Occlusal view. (B) Line drawing of (A). (C) Posterior view. (D) Lingual view. (E) Anterior view. (F) Labial view. www.sciencemag.org SCIENCE VOL 318 9 NOVEMBER 2007 937 on November 9, 2007 www.sciencemag.org Downloaded from