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Oligocene and Early Miocene Ruminants (Mammalia, Artiodactyla) from Pakistan and Uganda

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Oligocene and Early Miocene Ruminants (Mammalia, Artiodactyla) from Pakistan and Uganda Palaeontologia Electronica http://palaeo-electronica.org OLIGOCENE AND EARLY MIOCENE RUMINANTS (MAMMALIA, ARTIODACTYLA) FROM PAKISTAN AND UGANDA John C. Barry, Susanne Cote, Laura MacLatchy, Everett H. Lindsay, Robert Kityo, and A. Rahim Rajpar ABSTRACT Late Oligocene and Early Miocene fossil ruminants from Napak and Moroto in Uganda and the Zinda Pir sequence in Pakistan comprise at least eight taxa, including an indeterminate lophiomerycid, ?Gelocus gajensis, Walangania africanus, Bugtimeryx pilgrimi, an indeterminate large cervoid, Progiraffa exigua, Palaeohypsodontus zinen- sis, and an unnamed bovid. The fossils range in age between 25 and 16 million years old and together with species of tragulids provide evidence for the existence of diverse latest Oligocene and earliest Miocene ruminant faunas in sub-Saharan Africa and southern Asia. The fossils from Pakistan may also document the first appearance and subsequent radiation of giraffes and bovids, two groups that dominate later Neogene and modern herbivore faunas. John C. Barry. Peabody Museum and Department of Anthropology, Harvard University, 11 Divinity Ave., Cambridge, Massachusetts 02138 USA [email protected] Susanne Cote. Department of Anthropology, Harvard University, 11 Divinity Ave., Cambridge, Massachu- setts 02138 USA Laura MacLatchy. Department of Anthopology, University of Michigan, 550 East University Ave., Ann Arbor, MI 48109-1092 USA Everett H. Lindsay. Department of Geosciences, University of Arizona, Tucson, AZ 85721, USA Robert Kityo. Zoology Department, Makerere University, Kampala, Uganda A. Rahim Rajpar. Pakistan Museum of Natural History, Islamabad, Pakistan KEY WORDS: Ruminants; Miocene, Early; Oligocene; Pakistan; East Africa PE Article Number: 8.1.22A Copyright: Society of Vertebrate Paleontology May 2005. Submission: 15 November 2004. Acceptance: 5 March 2005 INTRODUCTION world and within the abstract geographies of the past and future. He wanted to go everywhere and Our friend Will Downs liked to travel; both pretty much did. Precambrian river gorges, Triassic within the natural and cultural geographies of this Barry, John C., Cote, Susanne, MacLatchy, Laura, Lindsay, Everett H., Kityo, Robert, and Rajpar, A. Rahim, 2005.Oligocene and Early Miocene Ruminants (Mammalia, Artiodactyla) from Pakistan and Uganda, Palaeontologia Electronica, Vol. 8, Issue 1; 22A:29p, 885MB; http://palaeo-electronica.org/paleo/2005_1/barry22/issue1_05.htm BARRY ET AL.: OLIGOCENE MIOCENE RUMINANTS FROM PAKISTAN AND UGANDA and in most cases represents species that are poorly known. The fossils represent at least eight species belonging to six major groups. Seven spe- cies come from Pakistan and one from Uganda. The scope of the study is limited in two ways. First, we only discuss new material and do not attempt to reanalyze older collections, which in East Africa are quite extensive. Such an undertaking would be too much for the limited time available and is only significant in the case of the East African record. Secondly, with the exception of one species, the taxa discussed in this paper are all members of the Pecora. Tragulids are also very well represented in these collections, perhaps with as many as seven additional species in two genera from the Oli- gocene and Early Miocene. The material, however, requires a thorough study of the very large and species rich Middle and Late Miocene collections of tragulids from the Potwar Plateau and East Africa. Tragulids are thus not considered here. METHODS Figure 1. Geographic positions of localities: 1) fossil Molar and premolar terminology in this paper sites in Uganda; 2) fossil sites in Pakistan. follows that of Janis and Scott (1987) and Gentry (1994), although we use “prehypocristid” to refer to what is otherwise called the cristid obliqua. Other tribal deserts, Miocene tropical volcanoes, and modifications are noted in the text. Measurements even densely urban, modern civilizations were all are in millimeters or centimeters, with lengths Will’s habitats. being the maximum dimension parallel to the This paper uses our memories of Will as an mesiodistal axis of the tooth, trigonid width the opportunity to bring together descriptions of fossil maximum through the protoconid and metaconid material from just three of the places where he perpendicular to the mesiodistal axis, and talonid worked: Uganda and the Potwar Plateau and width the maximum through the hypoconid and Sulaiman Ranges of Pakistan. While these fossils entoconid perpendicular to the mesiodistal axis. would ordinarily not have been described together, General locations for the fossil sites are they do have a commonality. They are early repre- shown in Figure 1.1-1.2.The ages of the sites are sentatives of one of the great mammalian radia- based on the published literature in the cases of tions, the Oligocene to Early Miocene radiation of Uganda and the Potwar Plateau, and the accom- the ruminants. The ruminant radiation gave rise to panying paper by Lindsay et al. (this issue) for the six modern families with nearly 200 extant species, Zinda Pir sites in the Sulaiman ranges. The age and it also produced an abundance of extinct spe- estimates for East African sites are primarily radio- cies and higher level taxa. However, most of what metric, while those for the Potwar Plateau are we know of this radiation is based on taxa from the based on paleomagnetic sequences. The Lindsay temperate Oligocene and Miocene of North Amer- et al. (this issue) paper presents new interpreta- ica, Europe, and central Asia, with little being tions of the stratigraphy and paleomagnetic record known of contemporaneous ruminants from tropi- of the Zinda Pir Dome area. In citing ages from it cal southern Asia or Africa, which are now the sites we have used their second interpretation (see fig- of the greatest species diversity. In this paper we ure 6b in Lindsay et al. this issue), which suggests describe fossil material from the Early Miocene of a Late Oligocene through Early Miocene age for all Uganda and the Oligocene through Early Miocene of the Chitarwata Formation. The alternative, Inter- of southern Asia, and, particularly in the case of pretation A (Lindsay et al. this issue, Figure 6A), Pakistan, this adds a new element to our under- suggests an Early Oligocene age for the lower part standing of the evolutionary history of early rumi- of the Chitarwata Formation, and a Late Oligocene nants. to Early Miocene age for the upper part. Both inter- The material we discuss was collected during pretations favor an Early Miocene age for the lower fieldwork by several of us during the past 15 years, 2 BARRY ET AL.: OLIGOCENE MIOCENE RUMINANTS FROM PAKISTAN AND UGANDA Table 1. Length and Widths of Lower Teeth. p3 p4 dp4 m1 m2 m3 indet. m (mm) (mm) (mm) (mm) (mm) (mm) (mm) Lophiomerycidae, gen. et sp. indet. Z 2278 Length: 8.4 Width: 4.7 ?Gelocus gajensis Z 271 Length: -- Width: 9.1 Z 2298 Length: 22.7 Width: 10.0 Walangania africanus BUMP 721 Length: 9.8 9.9 11.1 Width: 5.5 6.8 7.7 BUMP 722 Length: 8.7 8.6 9.9 Width: 4.5 5.3 6.4 BUMP 584 Length: 10.8 Width: 7.2 BUMP 552 Length: 16.0 Width: 6.9 BUMP 450 Length: 9.1 Width: 4.5 Bugtimeryx pilgrimi Z 269 Length: 8.7 11.0 Width: 6.1 7.5 Z 270 Length: 18.0 Width: 8.1 Z 2024 Length: 12.2 Width: 7.8 Z 2030 Length: 12.2 Width: 8.2 Z 2032 Length: -- Width: 8.0 Z 2068 Length: 8.1 -- Width: 4.9 5.9 ?Cervoidea, gen. et sp. indet. Z 2031 Length: 17.8 Width: 11.2 Vihowa Formation. Interpretation A of Lindsay et University Project; GSP-Y: Geological Survey of al. (this issue) is the same as an age correlation Pakistan - Harvard University Project; GSP-DGK: suggested by Welcomme et al. (2001) for the Chi- Geological Survey of Pakistan - D.G. Khan collec- tarwata Formation in Bugti Hills 300 km south of tion; GSP-S: Geological Survey of Pakistan - Sind the Zinda Pir area. collection; PMNH-Z: Pakistan Museum of Natural Measurements for the lower and upper teeth History - Zinda Pir collection. and astragalus are in Tables 1, 2, and 3. SYSTEMATIC PALEONTOLOGY Abbreviations BUMP: Boston University Uganda Museum Order Artiodactyla Makerere University Paleontology Expedition; Suborder Ruminantia GSP-H: Geological Survey of Pakistan - Howard Infraorder Tragulina 3 BARRY ET AL.: OLIGOCENE MIOCENE RUMINANTS FROM PAKISTAN AND UGANDA Table 1. (continued). p3 p4 dp4 m1 m2 m3 indet. m (mm) (mm) (mm) (mm) (mm) (mm) (mm) Progiraffa exigua S 88 Length: 18.6 Width: 14.3 S 412 Length: 17.7 Width: 9.2 S 305 Length: 21.3 Width: 13.9 H 208 Length: 32.1 Width: 14.5 Y 31676 Length: 20.2 Width: 14.8 Y 41662 Length: 21.2 Width: 14.4 Palaeohypsodontus zinensis Z 241 Length: 12.0 Width: 7.2 Z 2075 Length: 9.6 12.1 13.8 Width: -- -- -- Table 2. Length and Widths of Upper Teeth. P3 P4 dP4 M1 M2 M3 Indet. M (mm) (mm) (mm) (mm) (mm) (mm) (mm) Walangania africanus BUMP 23 Length: 10.0 Width: 12.0 BUMP 274 Length: 9.0 9.1 10.9 11.2 11.5 Width: 7.8 9.8 11.5 12.1 11.9 BUMP 390 Length: 10.1 Width: 12.1 Bugtimeryx pilgrimi Z 2027 Length: 9.7 Width: 10.0 Z 2028 Length: 10.3 Width: -- Z 2029 Length: 9.7 Width: 11.6 Progiraffa exigua H 664 Length: 26.7 Width: 29.1 H 312 Length: 18.5 10.8 22.3 24.0 24.3 Width: 15.1 17.5 21.7 25.9 23.7 Lophiomerycidae Janis, 1987 fragment of a right p3; Z 2278, left mandible with Genus and Species Indeterminate m1-m2; Z 2299, right astragalus. Material Locality and Age Locality Z 108: Z 760, fragment of a left P3; Z 761, Known from only one Zinda Pir site, Locality Z 108 fragment of a very worn upper right molar; Z 2277, in the lower part of the Chitarwata Formation.
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