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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2883, pp. 1-36, figs. 1-13, tables 1-3 June 26, 1987 The Occurrence of Hadromys (Rodentia: Muridae) in Early Pleistocene Siwalik Strata in Northern Pakistan and Its Bearing on Biogeographic Affinities Between Indian and Northeaster African Murine Faunas GUY G. MUSSER' ABSTRACT Three upper molars collected from an early lunda, another Indian native, and because that Pleistocene site in the Soan Formation of the Up- genus has also been identified in Pliocene sedi- per Siwaliks outcropping in the Pabbi Hills of ments of Ethiopia, morphologies of specimens in northern Pakistan are described as a new species samples of Recent and fossil Golunda are con- of Hadromys. That genus was known only by the trasted. Golunda remains an Indian region endem- living H. humei, which is found in northeastern ic; the Ethiopian species is not a member of that India. Compositions of the murine faunas of genus and is known only from northeastem Africa. northeastem India, northern Pakistan, and Pleis- Biogeographical relationships between the Indian tocene Pakistan are compared. Because the new subcontinent and northeastern Africa as indicated Hadromys is associated with fossil species of Go- by Recent and fossil murine faunas are discussed. INTRODUCTION With the publication in 1978 of a report posits in northern Pakistan, Dr. Louis L. Ja- on fossil Muridae from Neogene Siwalik de- cobs opened a window on past species di- ' Archbold Curator, Department of Mammalogy, American Museum of Natural History. Copyright © American Museum of Natural History 1987 ISSN 0003-0082 / Price $3.50 2 AMERICAN MUSEUM NOVITATES NO. 2883 versity ofmurine rodents in central Asia. The (Sabatier, 1982; Wesselman, 1984, for ex- view he provided has significantly influenced ample). the ways other glirologists are looking at pos- As a student ofliving murines, particularly sible evolutionary histories that resulted in those native to Asia and the Indo-Australian the modem murine fauna now composed of region, I turn often to morphologies and tem- more than 450 species (Carleton and Musser, poral distributions of fossil murines, seeking 1984). The Siwalik sediments containing mu- insights into origins of patterns of phyloge- rine fossils were deposited between 14 and netic relationships and geographic distribu- 1.6 million years ago (Barry et al., 1985; Ja- tions reflected among contemporary murine cobs et al., in press; Lowrie and Alvarez, 1981) species. Asian fossils, particularly those from and yielded samples of isolated molars and the Pliocene to Recent time period, are es- incisors representing 8 genera and at least 11 pecially important to the understanding of species. The deposits contain Antemus chin- links between ancient and modern faunas. jiensis, the oldest known murine, which ac- However, the uncritical use ofthe genus Rat- cording to Jacobs (1978) possessed a mor- tus as a catchall for many species of fossil phology from which all other murines could murines presents a major hinderance. My re- be derived. Miocene species of Progonomys, search has demonstrated that many contem- Parapodemus, and Mus comprised one pri- porary species conventionally assigned to mary branch that Jacobs derived from basal Rattus (Ellerman, 1941, for example) are only Antemus-like stock; Miocene species ofKar- distantly related to Rattus proper, and I have nimata and Parapelomys formed another accordingly reclassified them (Musser, 1981, branch emerging from the Antemus trunk. 1982; Musser and Newcomb, 1983, for ex- Pleistocene samples of Golunda and another ample). Similarly, I have encountered diffi- form that Jacobs tentatively assigned to Rat- culties with certain samples of Pleistocene tus represented twigs originating from the taxa; for example, fossils from Pleistocene Karnimata-Parapelomys stock. Living species Chinese outcrops originally identified as Rat- of Apodemus, Mus, Acomys, Praomys, and tus (or its synonym Epimys) but which proved their allies form the crown ofthe Progonomys to be examples of Niviventer and Leopold- branch; from the Karnimata branch has leafed amys (Musser, 1981). the contemporary species ofRattus, Arvican- Jacobs' (1978) identification of three iso- this, Pelomys, Mylomys, Golunda, and their lated molars from early Pleistocene Siwalik relatives. From Miocene beginnings, as in- sediments as cf. Rattus sp. suggested the pos- terpreted by Jacobs, the evolutionary diver- sibility of another species erroneously re- sity of murines flourished over Asia, Africa, ferred to that genus. Dr. Jacobs was very gen- and Europe. erous and did not hesitate to send me the Results ofJacobs' study have been used as three teeth. I studied them, took measure- a frame ofreference by others studying Neo- ments, subjected them to strong microscope gene assemblages of fossil murines. Subse- lamps and electron blasts from the scanning quent to publication of his descriptions and electron microscope, managed not to drop comparisons, additional murine fossils from any, and did not lose a single tooth. My iden- northern Pakistan have been described and tification of those small, isolated upper mo- their significance discussed by Cheema et al. lars as well as their significance in under- (1983) and Wessels et al. (1982). The diver- standing the present and past distributions of sity of fossil rats and mice in central Asia Asian murines form the substance ofthis re- suggested by the Pakistan samples has been port. increased by discovery of new species from The fossils are a sample ofan extinct species Miocene and Pliocene sediments in Afghan- ofHadromys.2 Its closest relative is the living istan (Brandy, 1979; Brandy et al., 1980; Sen H. humei, a native of northeastern India. To et al., 1979; Sen, 1983); the interplay of past faunas between Asia and Africa has been 2 Another view has recently been provided by Gaur touched on not only by Jacobs (1978) but also (1986: 543), who in reporting on a new fossil murine by those reporting on compositions of Plio- from the Upper Siwaliks of India states that "cf. Rattus cene murine faunas from northeastern Africa ofJacobs is more related to Nesokia, with which it shares 1 987 MUSSER: HADROMYS 3 place description and comparisons ofthe fos- BM British Museum (Nat. Hist.), sils in proper context, I begin the report with London a redescription ofH. humei that is meant not FMNH Field Museum of Natural Histo- as a definitive account but as an introduction ry, Chicago to this poorly known member of the Indian fauna. My knowledge of the species comes from study of skins, skulls, and dentitions METHODS along with the scanty ecological and distri- Specimens I examined are in collections of butional data available in published ac- the institutions listed above. Casts of fossils counts. This information sets the stage for are in the American Museum ofNatural His- the next section describing the fossil species tory. Specimens requiring identification are of Hadromys and comparing it with H. hu- noted in text, legends to figures, and tables. mei. The part that follows discusses relation- Values I obtained from measuring certain ships between the two species of Hadromys, specimens are listed in either table 2 or leg- between Hadromys and Rattus, and between ends to figures; limits of the measurements Hadromys and genera in the arvicanthine are explained in the legend to table 2. group, particularly Golunda. Faunal and hab- Much ofthe report focuses on comparisons itat associations are then provided which ofdental traits among taxa offossil and mod- contrast the Pleistocene and modern murine ern species. The nomenclature I use for cusps assemblages of northern Pakistan with the (also called cones or tubercles by other au- modern fauna native to northeastern India. thors) and cusplets (sometimes referred to as A next-to-last section discusses biogeo- cuspids or cuspules) is diagrammed in figure graphic relationships of the faunas in north- 1. The terminology brings together those pro- ern Pakistan and northeastern India at the posed by Miller (1912) and van de Weerd level of species and then genera. I focus on (1976) and is a combination I have used in Golunda, which is a constituent ofboth living my previous publications on muroid rodents. and early Pleistocene murine associations in Other systems of names and notations have northern Pakistan, has been identified from been proposed (see table 3 in Jacobs, 1978), Pliocene sediments in northeastern Africa, including a descriptive terminology suggest- and is currently considered to be a close rel- ed by Jacobs (1978) in his report on Siwalik ative of the living African Mylomys. I com- rodents. The nomenclature he applies to most pare samples ofAsian Golunda with African parts ofmolars reflects homologies with den- Mylomys and then with the African Pliocene tal structures in other groups of muroid ro- samples of Golunda. My interpretations of dents and even mammals that are not ro- phylogenetic and biogeographic relationships dents. For that reason his arrangement is derived from morphologies in those samples better than most ofthe others but I still have contrast with current views. Finally, I discuss problems with homologies of certain struc- generic limits in Hadromys and interconti- tures in his scheme and until I find solutions nental affinities between Asian and African I will stick to the terms shown in figure 1. murines. Another item of terminology needs expla- nation. When Jacobs (1978) wrote of Neo- gene Siwalik Muridae, he was referring to a ABBREVIATIONS group of more than 450 species of rats and DP Dartmouth College-Peshawar mice that are native to the Old World. His University Pakistan Project usage excluded hamsters (Cricetidae), voles AMNH American Museum of Natural (Arvicolidae), gerbils (Gerbillidae), and other History, New York groups of muroid rodents from the Muridae. Throughout this report I refer to Jacobs' Mu- ridae as murines, which reflects the arrange- ment ofCarleton and Musser (1984) in which many morphological characters and differs only in the the Murinae and 14 other Recent groups of absence of posterostyle, than to any Rattus species (J.
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