Chapter 16 Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan PIERRE- OLIVIER ANTOINE, GREGOIRE MÉTAIS, MAEVA J. ORLIAC, J.- Y. CROCHET, LAWRENCE J. FLYNN, LAURENT MARIVAUX, ABDUL RAHIM RAJPAR, G. ROOHI, AND JEAN- LOUP WELCOMME he Sulaiman Range is a north–south-trending band of rugged mountains rising 1000–3400 m above sea level that dei nes the modern political boundary between Balochistan and Punjab provinces and extends northward into North-West Frontier Province (igure 16.1). Late Mesozoic and Cenozoic sedimentary rocks here are primarily marine and accumulated in the Tethys Sea in what is now the Indus Basin, bounded to the northwest by the Axial Belt (Shah 1977), or Bela–Waziristan Ophiolite Zone (Bannert et al. 1992), and to the southeast by the Indo–Pakistani Subcontinent. he east side of the Sulaiman Range is of par ticu lar interest since a relatively thick, well-exposed mid-Cenozoic sequence registers the transition from marine shelf to terrestrial deposition episodes related to the uplit and erosion of the orogenic Himalayan highlands and the related closure of the Tethys Sea. Deposition of these thick detrital units resulted from the collision between the Indian and Asian plates, which started near the Paleocene/Eocene transition, ~55 Ma (Beck et al. 1995; Clit et al. 2001). STRATIGRAPHIC AND HISTORICAL CONTEXT -1— 0— +1— he fossiliferous outcrops were recognized early in the nineteenth century (Vickary 1846) and were initially described in detail by Blanford (1883), who assigned the terrestrial conglomerates and sandstones overlying the Eocene “Nummulitic Limestone” to the Upper Nari Formation. hese sediments—especially in the southern 49502_1P_03_Wang15012_text14-31.indd 400 Sulaiman Range, roughly corresponding to the Bugti Hills—soon became famous for their fossil vertebrate faunas (Lydekker 1884; Pilgrim 1907, 1908, 1910, 1912; Forster-Cooper 1913, 1915, 1924), rich in large terrestrial mammals of Miocene to Pleistocene age. he faunas, as a whole, have long been considered as terrestrial, as they provide clear evidence that the environment of deposition was primarily luvial. he stratigraphic classiication of the sediments of the Sulaiman foothills has evolved with the growth of knowledge, and many diferent names have been applied, both generally and locally (Hemphill and Kidwai 1973; Shah 1977; Welcomme et al. 2001). he intensive ieldwork on the Siwalik series of the Potwar Plateau greatly inluenced biostratigraphical interpretation of the entire Himalayan foothills of Pakistan (Pilgrim 1912; Raza and Meyer 1984). he exceptionally well-exposed succession in the Potwar Plateau area of northern Pakistan provides the best standard succession for almost the entire Neogene, and its biostratigraphy is successfully dated by paleomagnetic methods (Johnson et al. 1985; Barry et al. 2002). he period encompassing the Oligocene and earliest Miocene has long been considered as lacking in the entire Sulaiman Geological Province (e.g., Raza and Meyer 1984). Traditionally, the depositional history of preSiwalik continental strata is very poorly known, from the point of view of age, environment, or burial structure. he Murrees sensu lato, which unconformably overlies the Eocene marine rocks in some areas of the Potwar region, may represent this missing sedimentary record. 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN 401 Figure 16.1 Geological map of the Sulaiman Province, Middle Indus Basin, Pakistan. Mammal-yielding localities discussed in the text are located in the Zinda Pir Dome and in the Bugti Hills. Modified after Raza et al. (2002). However, the Murrees deposits are very poor in fossil remains, and correlations are still delicate to establish with pre-Siwalik deposits of other areas, including those of the Sulaiman Range (Najman and Garzanti 2000; Kumar and Kad 2003). In the last decades, the continental series of the Sulaiman Range has been investigated as a southern extension 49502_1P_03_Wang15012_text14-31.indd 401 of the important framework made in the Potwar Plateau area (Barry, Lindsay, and Jacobs 1982; Barry, Flynn, and Pilbeam1990; Barry et al. 2002) by a Yale University– GSP team since the early 1980s. Important data came with the irst collection of small mammals in the Sulaiman foothills (Jacobs et al. 1981; Flynn, Jacobs, and Cheema 1986). Renewed reconnaissance of the Sulaiman foothills —-1 —0 —+1 6/27/12 12:09 AM 402 SOUTH AND SOUTHEAST ASIA started in Dalana, in the Zinda Pir Dome area (Raza et al. 2002). he i rst attempt to place the fossils in a stratigraphic framework began there (Downing et al. 1993) in coordination with sediment sampling for magnetic stratigraphy (Friedman et al. 1992). Independently, a French expedition led by Jean-Loup Welcomme and the late Leonard Ginsburg was initiated in the famous Bugti Hills in 1995, with the main objective of establishing a proper lithostratigraphic framework in order to determine the stratigraphic provenance of spectacular fossil collections made by Pilgrim and Forster-Cooper nearly a century before. he i rst survey of the Dera Bugti Syncline produced a detailed section of the Early Miocene deposits that, at some points, lie unconformably on the Eocene marine limestones of the Kirthar Formation (Welcomme et al. 1997) and provided the i rst unambiguous evidence of (fossiliferous) Oligocene deposits in the Sulaiman Lobe (Welcomme and Ginsburg 1997). hese critical results (summarized in Welcomme et al. 1999; Welcomme et al. 2001), led to a reexamination of the age of the Zinda Pir sequence (Lindsay et al. 2005) and had important biostratigraphic implications for the entire Sulaiman Province, the chronology of deposition of exhumed material, and the history of West Himalayan foreland basins, especially the Indus paleodrainage and its tributaries (Downing and Lindsay 2005; Métais et al. 2009). So far, the pre-Siwalik deposits exposed in the Sulaiman Range constitute one of the best terrestrial and fossiliferous archives for the eroded materials from the rising Himalaya. Formally, the units documenting Neogene time in the Sulaiman Province are as follows, from the base to the top. Chitarwata Formation, Upper Member (Earliest Miocene) -1— 0— +1— In the Zinda Pir area, the Chitarwata Formation is up to 480 m thick (Raza et al. 2002; Lindsay et al. 2005) while it is much more condensed in the Bugti Hills (120–260 m thick; Métais et al. 2009; new data). he lower and middle units recognized in the Zinda Pir area (Lindsay et al. 2005) and the Bugti Member of the Bugti Hills (Métais et al. 2009) are considered similar faunally and temporally and, being Oligocene in age, are discussed only in reference to younger sites. he upper part of the Chitarwata Formation in both areas is referred to the earliest Miocene (i.e., roughly corresponding to the Aquitanian marine stage or Agenian European Land Mammal Age; Antoine et al. 2010). he thickness of the upper member 49502_1P_03_Wang15012_text14-31.indd 402 varies laterally, but it reaches about 200 m near Dalana (Zinda Pir). he measured thickness is 50 m (at Kumbi, most condensed section) up to 130 m (igure 16.2 [Habib Rahi]; unpublished data) in the Bugti Hills, with two distinct and successive fossiliferous levels (igure 16.3 [level M = 3; Q = level]; Welcomme et al. 2001; Métais et al. 2009). Diferences in thickness of the Chitarwata Formation between the Bugti Hills and Zinda Pir are discussed in Métais et al. (2009). In the Bugti Hills, the upper member of the Chitarwata Formation is the richest stratigraphic interval in terms of fossil vertebrates (table 16.1). It has been widely investigated in the past, and it yields the most diversiied Bugti faunas (levels 3bis and 4; Welcomme et al. 2001). However, this interval is only documented by mediumsize and large mammals in the Bugti area: no small mammal was recovered. Near Dalana in the Zinda Pir area, ive small mammal localities were recovered within this interval (see igure 16.3 [Z113, Z139, Z150, Z135, and Z126, from base to top]; Lindsay et al. 2005). Large mammal remains are scarce, although they occur in various localities (Raza et al. 2002; Barry et al. 2005; Lindsay et al. 2005). Vihowa Formation (Late Early Miocene–Middle Miocene) he Vihowa Formation is 720 m thick in the Zinda Pir Dome (Raza et al. 2002). Only the lower part was studied thoroughly (E. H. Lindsay, personal com., 2010). Vihowa thickness is likely to reach only ca. 100–200 m in the Bugti Hills (see igure 16.3), but this is far from being well constrained. Again, only the lower part was sampled densely. Various sites from three successive levels (levels 5 [=T], 6, and 6sup [=Bugti sup]) document the lowermost part of the Vihowa Formation in the Bugti Hills (late Early Miocene), while a single locality (see igure 16.2 [Lundo W]) is referable to the middle part (Middle Miocene, i.e., considered similar to the Chinji fauna of Potwar Plateau; Welcomme et al. 2001; Métais et al. 2009; Orliac et al. 2009; Antoine et al. 2010). he levels 6 and 6sup yielded both large mammal and micromammal faunas (table 16.2; Welcomme et al. 1997; Welcomme et al. 2001; Métais et al. 2009). For Zinda Pir, three small mammal localities were recovered within this interval (see igure 16.3 [Z124, Z122, and Z120, from base to top]; Lindsay et al. 2005). Large mammal remains are rather scarce, but they occur in various localities (Raza et al. 2002; Barry et al. 2005; Lindsay et al. 2005). 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN 403 Figure 16.2 Location of main Cenozoic vertebrate-yielding sections from the Bugti Hills, Balochistan, Pakistan. Litra Formation (Late Miocene) he Litra Formation, in the Zinda Pir area, is 1700 m thick (Raza et al. 2002); no concentrated survey was done in that interval (E. H. Lindsay, personal com., 2010). It is most likely much thinner in the Bugti Hills, but this hypothesis is not constrained by formally measured sections, except in the Habib Rahi area (ca. 150 m; see igure 16.2; unpublished data). Two localities referable to this stratigraphic interval are recognized in the Bugti Hills (see igures 16.2 [Djigani] and 16.3 [levels 7 and 7sup]; Welcomme et al. 1997; Welcomme et al. 2001; Antoine, Duranthon, and Welmomme 2003b). hey have yielded a large mammal fauna similar to what is observed in the lower part of the Middle Siwaliks in the Potwar Plateau, including the First Local Appearance (FLA) of Hipparion sensu lato (table 16.3; Antoine, Duranthon, and Welmomme 2003b). 49502_1P_03_Wang15012_text14-31.indd 403 Post-Miocene Formations (Chaudhwan Formation, Pliocene–Pleistocene) he Chaudhwan Formation is ca. 1500 m thick in Zinda Pir (Raza et al. 2002). In the Bugti area, its thickness is not constrained by measured sections, but it appears as highly variable depending on local tectonic context: the corresponding deposits are much deformed and consist essentially of boulder conglomerates and luvial terraces, mainly observable in the Dera Bugti Syncline and in Habib Rahi (see igures 16.1 and 16.2). Welcomme et al. (1997) mention only egg shells of an unidentiied struthioniform (ostrich). Crochet et al. (2009) report prehistoric rock paintings in the vicinity of Lundo (see igures 16.1 and 16.2), with anthropomorphic, geometric, and zoomorphic (e.g., cervid and felid) sketches attesting to favorable climatic conditions in the area around the Last Glacial Maximum and during subsequent periods. —-1 —0 —+1 6/27/12 12:09 AM -1— 0— +1— Figure 16.3 Stratigraphic sections of post- Eocene deposits of the Sulaiman Province (Zinda Pir Dome, to the left; Bugti Hills, to the right), with First Local Appearances (FLA) and Last Local Appearances (LLA) of index mammals and tentative correlation to the GPTS. Zinda Pir localities appear as “Z numbers” in the left column. Completed and modified after Marivaux et al. (1999, 2001, 2005, 2006), Welcomme et al. (2001), Raza et al. (2002), Antoine et al. (2003b, 2010), Barry et al. (2005), Gradstein et al. (2005), Lindsay et al. (2005), Métais et al. (2009a, 2009b), and Orliac et al. (2009, 2010). 49502_1P_03_Wang15012_text14-31.indd 404 6/27/12 12:09 AM Table 16.1 Mammals from the Upper Member of the Chitarwata Formation (Earliest Miocene) of the Sulaiman Province Soricidae genus and species indet. ZP Chiroptera genus and species indet. ZP Tupaiidae genus and species indet. ZP Lorisidae genus and species indet. ZP Sciuridae genus and species indet. ZP Petauristinae gen et sp. indet. ZP Cricetidae Eumyarion kowalskii ZP ?Eumyarion sp. ZP Democricetodon sp. ZP Democricetodon sp. A ZP Spanocricetodon khani ZP Spanocricetodon sulaimani ZP Spanocricetodon sp. ZP Primus sp. ZP Rhizomyidae Prokanisamys arii ZP Prokanisamys sp. ZP Ctenodactylidae genus and species indet. ZP “Z135 Sayimys sp.” ZP Prosayimys lynni ZP Prosayimys sp. ZP ?baluchimyine indet. ZP Diatomyidae Marymus dalanae ZP Carnivora genus and species indet. ZP Amphicyonidae Amphicyon shahbazi BH genus and species indet. ZP Proboscidea genus and species indet. ZP Deinotheriidae Prodeinotherium sp. ZP, BH P. pentapotamiae BH Elephantoidea genus and species indet. ZP, BH Gomphotherium sp. ZP, BH Table 16.1 (continued ) Sanitheriidae ?Sanitherium jef reysi ZP Sanitherium jef reysi BH Palaeochoeridae Pecarichoerus sp. BH Suidae Listriodon ainis BH Anthracotheriidae Parabrachyodus sp. ZP P. hyopotamoides ZP Microbunodon silistrense ZP Sivameryx palaeindicus ZP, BH Hemimeryx cf. blanfordi BH Telmatodon sp. BH ?Masritherium sp. ZP genus and species indet. ZP Tragulidae genus and species indet. ZP Dorcatherium sp. ZP, BH Giraidae Progirafa exigua ZP Bovidae Eotragus sp. BH Ruminantia incertae sedis Bugtimeryx pilgrimi ZP, BH Chalicotheriidae Phyllotillon naricus BH “Chalicotherium” pilgrimi BH genus and species indet. ZP Rhinocerotidae Protaceratherium sp. ZP, BH Plesiaceratherium naricum BH Mesaceratherium welcommi ZP, BH Pleuroceros blanfordi ZP, BH Brachypotherium gajense ZP, BH B. fatehjangense ZP, BH Prosantorhinus shahbazi BH Gaindatherium cf. browni BH ?cf. Rhinoceros sp. BH Bugtirhinus praecursor BH genus and species indet. ZP Note: ZP, Zinda Pir; BH, Bugti Hills Sources: Completed and modiied ater Welcomme et al. (2001), Raza et al. (2002), Ginsburg and Welcomme (2002), Antoine et al. (2004, 2010), Barry et al. (2005), Gradstein et al. (2004), Lindsay et al. (2005), Métais et al. (2009a, 2009b), Orliac et al. (2009, 2010) and references therein, and new data. —-1 —0 —+1 49502_1P_03_Wang15012_text14-31.indd 405 6/27/12 12:09 AM Table 16.2 Mammals from the Vihowa Formation (Late Early to Middle Miocene) of the Sulaiman Province -1— 0— +1— Erinaceidae Amphechinus sp. ZP genus and species indet. ZP Soricidae genus and species indet. ZP, BH Chiroptera genus and species indet. ZP Tupaiidae genus and species indet. ZP Sivaladapidae Guangxilemur sp. ZP Sciuridae Petauristinae gen. et sp. indet. ZP genus and species indet. ZP Cricetidae Democricetodon sp. B ZP Democricetodon sp. X ZP Spanocricetodon khani BH Spanocricetodon sulaimani ZP Spanocricetodon sp. BH Megacricetodon ?daamsi BH Megacricetodon sp. ZP ?Megacricetodon sp. ZP Myocricetodon sivalensis ZP Myocricetodon sp. 1 ZP Myocricetodon sp. 2 ZP Myocricetodon sp., large ZP Rhizomyidae Prokanisamys arii ZP Prokanisamys cf. benjavuni BH Prokanisamys benjavuni BH Prokanisamys sp. ZP Ctenodactylidae genus and species indet. ZP Sayimys intermedius BH Sayimys cf. intermedius ZP Sayimys sp. ZP Diatomyidae Diatomys sp. Creodonta Hyanailouros sulzeri ZP Pterodon bugtiensis BH genus and species indet. ZP Carnivora Megamphicyon giganteus BH Amphicyon shahbazi BH genus and species indet. ZP 49502_1P_03_Wang15012_text14-31.indd 406 Table 16.2 (continued ) Deinotheriidae Deinotherium sp. ZP Prodeinotherium pentapotamiae BH Elephantoidea Gomphotherium browni BH Gomphotherium cooperi BH Choerolophodon corrugatus ZP, BH cf. Protanancus chinjiensis ZP genus and species indet. ZP Sanitheriidae genus and species indet. ZP Suidae Listriodon guptai ZP, BH ?Listriodon ainis ZP Anthracotheriidae Parabrachyodus hyopotamoides BH Microbunodon silistrense ZP Sivameryx palaeindicus ZP, BH ?Sivameryx palaeindicus ZP genus and species indet. Tragulidae Dorcatherium sp. ZP Dorcatherium cf. parvum BH genus and species indet. ZP Giraidae Progirafa exigua ZP, BH ?Girafokeryx sp. ZP Bovidae Eotragus noyei ZP Eotragus sp. ZP cf. Elachistocerus sp. ZP genus and species indet. ZP Chalicotheriidae Anisodon sp. BH genus and species indet. ZP Rhinocerotidae ?Plesiaceratherium naricum BH Mesaceratherium welcommi BH Pleuroceros blanfordi BH Brachypotherium fatehjangense BH Brachypotherium gajense BH Brachypotherium perimense ZP, BH cf. Rhinoceros sp. ZP, BH Bugtirhinus praecursor ZP genus and species indet. ZP Note: ZP = Zinda Pir; BH = Bugti Hills. Sources: Completed and modiied ater Welcomme et al. (2001); Ginsburg and Welcomme (2002); Raza et al. (2002); Barry et al. (2005); Gradstein et al. (2004); Lindsay et al. (2005); Métais et al. (2009a, 2009b); Orliac et al. (2009); Antoine et al. (2010), references therein; and new data. 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN Table 16.3 Mammals from the Litra Formation (Late Miocene) of the Sulaiman Province Deinotheriidae Deinotherium sp. BH Elephantoidea Gomphotherium sp. ZP Choerolophodon corrugatus ZP, BH genus and species indet. BH Suidae Listriodon sp. ZP, BH ?Propotamochoerus sp. ZP Parachleuastochoerus sp. BH Giraidae ?Bramatherium perimense BH ?Bramatherium sp. ZP genus and species indet. ZP Bovidae Prostrepsiceros vinyaki ZP Hispanodorcas terrubiae ZP Reduncini gen. et sp. indet. ZP genus and species indet. BH Equidae Cormohipparion (Sivalhippus) theobaldi ZP Hippotherium sp. ZP, BH Rhinocerotidae Alicornops complanatum BH Brachypotherium perimense BH Rhinoceros af . sivalensis BH genus and species indet. ZP Note: ZP = Zinda Pir = BH, Bugti Hills. Sources: Completed and modiied ater Welcomme et al. (2001), Raza et al. (2002), Antoine et al. (2003b, 2010), Zouhri and Ginsburg (2003), Gradstein et al. (2004) and references therein, and new data. MAMMALIAN FAUNAS Fossil mammals can be split into two categories, roughly corresponding to the collection methods used in the ield: prospecting (middle-sized and large mammals) and screening/washing (small mammals). Small Mammals Neogene micromammal stes were much more investigated in the Zinda Pir area than in the Bugti Hills (Wel- 49502_1P_03_Wang15012_text14-31.indd 407 407 comme et al. 2001; Lindsay et al. 2005; igure 16.4a). he Zinda Pir Chitarwata and Vihowa formations notably yielded diversiied rodent assemblages (Lindsay et al. 2005). By contrast, no micromammal locality is available in Neogene deposits of the Bugti Hills, with the exception of two successive levels—documenting the same assemblage—in the base of the Vihowa Formation (levels 6 and 6sup, late Early Miocene; Welcomme et al. 1997; Métais et al. 2009). Interestingly, a wide array of micromammals (marsupials, insectivores, bats, dermopterans, primates, and rodents) was recovered in the Bugti Member of the Chitarwata Formation in the Bugti Hills (Paali C2, Early Oligocene; Marivaux et al. 2001; Marivaux et al. 2001 2005). To our knowledge, neither marsupial nor lagomorphs nor dermopteran remains have been found in the Neogene of the Sulaiman Province so far. Lipotyphla and Chiroptera Unidentiied soricid remains were recovered from the upper member of the Chitarwata Formation in Zinda Pir (Z113 and Z150) and from the lower part of the Vihowa Formation in both the Bugti Hills (level 6; Welcomme et al. 1997) and Zinda Pir (Z122; Lindsay et al. 2005). Amphechinus sp. and an unidentiied hedgehog are mentioned in the lower part of the Vihowa Formation (Z122; Lindsay et al., 2005). Bats of uncertain ai nities occur in Z113 (upper member of the Chitarwata Formation, earliest Miocene) and in Z124–Z122 (lower part of the Vihowa Formation, late Early Miocene; Lindsay et al., 2005). Euarchontes By contrast with the rich and diversiied primate and dermopteran fauna unearthed in the Bugti member of the Chitarwata Formation in the Bugti area (Paali C2, Early Oligocene; Marivaux et al. 2001; Marivaux et al. 2005; Marivaux et al. 2006), Neogene deposits of the Sulaiman Province have yielded only scarce euarchontan remains from Zinda Pir, consisting of unidentiied Tupaiidae (Z113, upper Chitarwata Formation; Z122, lower Vihowa Formation) and Lorisidae (Z135, upper Chitarwata Formation; Lindsay et al. 2005). he sivaladapid adapiform Guangxilemur sp. occurs in the lower part of the Vihowa Formation (Z122; Lindsay et al. 2005). he corresponding stratigraphic interval is restricted to the Early Miocene period (see tables 16.1 and 16.2 and igure 16.4a). No Neogene euarchontan is identiied in the Bugti Hills. —-1 —0 —+1 6/27/12 12:09 AM Figure 16.4 (a) Biostratigraphical range of small mammals from the Neogene of the Sulaiman Province, Pakistan. Taxa restricted to the Bugti Hills appear in gray. Taxa from Zinda Pir (most of them correlated to the GPTS) and/or from the whole Sulaiman Province (Zinda Pir + Bugti Hills) appear in black. Uncertain ranges are indicated by dashed lines. Taxa are sorted by order and/or family, and then by order of appearance. Completed and modified after Welcomme et al. (2001), Raza et al. (2002), Gradstein et al. (2005), Lindsay et al. (2005), Métais et al. (2009), and references therein. (b) Biostratigraphical range of large mammals from the Neogene of the Sulaiman Province, Pakistan. Taxa restricted to the Bugti Hills appear in gray. Taxa from Zinda Pir (most of them correlated to the GPTS) and/or from the whole Sulaiman Province (Zinda Pir + Bugti Hills) appear in black. Uncertain ranges are indicated by dashed lines. Taxa are sorted by order and/or family, and then by order of appearance. Completed and modified after Welcomme et al. (2001), Ginsburg and Welcomme (2002), Raza et al. (2002), Antoine et al. (2003b, 2010), Zouhri and Ginsburg (2003), Gradstein et al. (2005), Lindsay et al. (2005), Métais et al. (2009a, 2009b), Orliac et al. (2009), references therein, and new data. -1— 0— +1— 49502_1P_03_Wang15012_text14-31.indd 408 6/27/12 12:09 AM —-1 —0 —+1 49502_1P_03_Wang15012_text14-31.indd 409 6/27/12 12:09 AM 410 SOUTH AND SOUTHEAST ASIA Rodentia -1— 0— +1— In the Sulaiman Province, Neogene rodents are essentially known by micromammal localities from the Zinda Pir area, encompassing the upper member of the Chitarwata Formation (earliest Miocene; see table 16.1) and the lower part of the Vihowa Formation (late Early Miocene; see table 16.2; Lindsay et al. 2005). Only two levels from the lower part of the Chitarwata have yielded rodents in the Bugti Hills (see igures 16.2 [Dera Bugti] and 16.3 [levels 6 and 6sup]), but no micromammal is known from the upper member of the Chitarwata Formation (Welcomme et al. 2001). Correlation based on rodents within the Sulaiman Province is therefore restricted to the late Early Miocene (see igure 16.3). Sciuridae are documented by unidentiied remains from the upper member of the Chitarwata Formation (Z113, Z139, and Z135, from base to top) and from the lower part of the Vihowa Formation (Z122; Lindsay et al. 2005). A petauristine of uncertain ai nities occurs in the same interval (Z113 and Z122; Lindsay et al. 2005), and a ratui ne is found at Z135. Cricetidae are by far the most diversiied rodents of this interval, with at least 16 species (Welcomme et al. 2001; Lindsay et al. 2005). Eumyarion kowalskii (Z113), E. sp. (Z139), Spanocricetodon sp. and Spanocricetodon khani (Z113–Z135), and Primus sp. are restricted to the upper member of the Chitarwata Formation. h is part of the section records also the First Local Appearance (FLA) of Democricetodon (D. sp. and D. sp. A, in Z150) and Spanocricetodon sulaimani (Z113), while a couple of other species referable to Democricetodon appear in the lower part of the Vihowa Formation (D. sp. X, Z124; D. sp. B, Z122; Lindsay et al. 2005). Megacricetodon sp. and Myocricetodon (My. sp. 1, My. sivalensis, My. sp. 2, and My. sp., large) have their FAD in the base of the Vihowa Formation in the Zinda Pir Dome (Z124; Lindsay et al. 2005). In Zinda Pir, the rhizomyids Prokanisamys sp. and P. arii occur in the upper part of the Chitarwata Formation (Z150) and persist in the lower part of the overlying Vihowa Formation (up to Z122; Lindsay et al. 2005). P. cf. benjavuni is recorded in the levels 6 and 6sup of the Bugti Hills (lower Vihowa Formation, late Early Miocene; Métais et al. 2009). Ctenodactylidae are documented by various species in Zinda Pir (see tables 16.1 and 16.2 and igure 16.4). Prosayimys lynni (Z113) and P. sp. (Z113 and Z150) are restricted to the upper part of the Chitarwata Formation (Lindsay et al. 2005). he FLA of Sayimys sp. is recorded in the same interval, but dei nitive Sayimys (S. cf. intermedius) appears in the lower part of the Vihowa Formation 49502_1P_03_Wang15012_text14-31.indd 410 (Z122; Lindsay et al. 2005), which coincides with the occurrence of S. intermedius in the Bugti Hills (level 6sup; Welcomme et al. 2001). h ree enigmatic teeth were identiied as Baluchimys sp. and Zindapiria quadricollis in a single locality of the upper member of the Chitarwata Formation (Z113; Lindsay et al. 2005), but these records are possibly poorly preserved muroids. Diatomyidae are documented by Marymus dalanae (Z113, Z139, and Z150) in the upper member of the Chitarwata Formation (earliest Miocene) and by Diatomys sp. in the lower part of the Vihowa Formation in the Zinda Pir Dome (Z124 and Z122, late Early Miocene; Lindsay et al. 2005; Flynn 2007). he absence of any micromammal site from the upper part of the Chitarwata Formation (earliest Miocene) in the Bugti Hills prevents testing the rodent i rst appearances as observed in Zinda Pir for the corresponding interval. However, the rodent assemblages from the lower part of the Vihowa Formation are totally homotaxic in both areas (see table 16.2), at the genus or species level, with notably Megacricetodon and Sayimys appearing in the same interval in the Bugti Hills (level 6 and/or level 6sup; Welcomme et al. 2001; Métais et al. 2009) and in Zinda Pir (Z124, base of the Vihowa Formation; Lindsay et al. 2005). Large Mammals Ferae Pending a revision of a few medium-size specimens unearthed in the upper part of the Chitarwata Formation in the Bugti Hills (Samane, level 4, earliest Miocene; see igure 16.1), and due to the sampling method used in the ield—surface collection favoring large specimens— creodonts and carnivore remains are either unidentiied to genus and species or referred as hyaeonodontid creodonts and amphicyonid carnivores (i.e., large or gigantic taxa). heir fossil record is so far restricted to the Early Miocene in the Sulaiman Province (see igure 16.4b). he hyaenodontids Pterodon and Hyanailouros are documented in the lower part of the Vihowa Formation. Corresponding remains are referred to P. bugtiensis in the Bugti Hills (levels 6 and 6sup, late Early Miocene; Ginsburg and Welcomme 2002; Métais et al. 2009) and to H. sulzeri in coeval deposits of Zinda Pir (Z120; Lindsay et al. 2005). In the Bugti Hills, the amphicyonid Amphicyon shahbazi occurs in several localities encompassing the top of 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN the Chitarwata Formation (level 4, earliest Miocene) and the lower part of the Vihowa Formation (see igure 16.3 [levels 5 –6 sup, late Early Miocene]), while its close ally Megamphicyon giganteus occurs only in the lower part of the Vihowa Formation (levels 5– 6sup, late Early Miocene; Ginsburg and Welcomme 2002; Métais et al. 2009). On the other hand, only unidentiied carnivore and amphicyonid remains have been unearthed in coeval Zinda Pir deposits (Lindsay et al. 2005). Proboscidea Until recently, the Paleogene history of proboscideans was restricted to Africa, and elephants and their close relatives were supposed to disperse toward Eurasia during the late Early Miocene (Shoshani and Tassy 1996). h is dispersal was given strong biostratigraphic signiicance and named Proboscidean Datum Event (PDE) by Madden and Van Couvering (1976). he recognition of a diagnostic elephantoid incisor from the Bugti Member of the Chitarwata Formation in the Bugti Hills (locality DB-J1, late Oligocene; Antoine, Welcomme, et al. 2003), the discovery of another tusk lower in the same stratigraphic interval in 2004 (level 2=F; Bugti Member, Chitarwata Formation; unpublished data), and the subsequent mention of an unidentiied elephantoid in the lower Chitarwata at Zinda Pir (Oligocene Z108; Lindsay et al. 2005; now rejected as dei nitive) seriously challenge the concept of PDE as a single event occurring at ca. 17.5 Ma (late Early Miocene; Madden and Van Couvering 1976), and indicate rather that proboscideans dispersed several times from Africa to adjacent plates in Middle Cenozoic times, as suspected by Tassy (Barry, Flynn, and Pilbeam). Deinotheriidae have their FLA in the upper member of the Chitarwata Formation (earliest Miocene), with Prodeinotherium sp. (level 3bis; tusk and dental fragments) and P. pentapotamiae (level 4; dental remains) in the Bugti Hills (see igure 16.3 and table 16.1). Scarce remains are reported from the same interval in Zinda Pir (Z129 and upward; Raza et al. 2002; Lindsay et al. 2005). hese small deinotheres occur also throughout the Vihowa Formation (late early to middle Miocene) throughout the Sulaiman Province, and in coeval deposits of the Potwar Plateau (Kamlial and Chinji formations; Welcomme et al., 2001; pers. obs. POA) and Europe (e.g., Antoine et al. 1997). A larger deinotheriid referable to Deinotherium sp. occurs in the level 7sup of the Bugti Hills (Djigani, middle late Miocene; new data). In the Sulaiman Province, the earliest elephantoid remains identiied at genus level are referred to Gomphoth- 49502_1P_03_Wang15012_text14-31.indd 411 411 erium sp. hey originate from the upper part of the Chitarwata Formation in the Bugti Hills (see igure 16.3 [level 4, earliest Miocene]; Welcomme et al. 2001; Métais et al. 2009). he elephantoid assemblage is much more diversiied in the subsequent stratigraphic interval—that is, in the lower part of the Vihowa Formation in the Bugti Hills (levels 6 and 6 sup.; late Early Miocene)—with Gomphotherium browni, G. cooperi, Zygolophodon metachinjiensis, Choerolophodon cf. corrugatus, and Protanancus chinjiensis (Welcomme et al. 2001; Métais et al. 2009). he two latter taxa are also recognized in the middle part of the Vihowa Formation (Z205; Lindsay et al. 2005). C. corrugatus persists until the middle Late Miocene in the Bugti Hills (level 7sup at Djigani; new data), as in Potwar Plateau (Dhok Pathan Formation; Pilbeam et al. 1979, 1996). Cetartiodactyla Odontoceti An unidentiied odontocete is mentioned in the lower part of the Vihowa Formation (level 6), in the Bugti Hills (Welcomme et al. 1997). he available tooth belongs probably to a river dolphin, given the luvial depositional environment of the whole formation (Welcomme et al. 2001; Métais et al. 2009). Ruminantia Initially, Pilgrim (1908) referred dental remains to a primitive giraid, Progirafa exigua, with the mention “Dera Bugti Area,” which was later rei ned as “upper Nari” of “Dakko Nala” (Pilgrim 1911). Pilgrim (1912) described two new species of ruminants from the base of the “Gaj” in the Bugti Area, based on sparse dental material; the species were doubtfully assigned as Gelocus(?) gajensis and Prodremotherium(?) beatrix. At that time, both genera were common ruminants restricted to the Oligocene of Europe. Despite this weak biochronological evidence of Oligocene age, Pilgrim (1912), probably inluenced by the sequence observed in the Sind Province to the south, revised his former opinion concerning the probable occurrence of Oligocene deposits in the Bugti Hills (Pilgrim 1908) and argued for an early Miocene age for the whole Bugti fauna. Later on, there was no further extensive study of the ruminants from the Bugti Hills, principally due to the uncertain geographic and stratigraphic provenance of the described material, and the lack of ieldwork in the Sulaiman Lobe until the early 1980s and late 1990s (Raza and Meyer 1984; Welcomme et al. 1997). —-1 —0 —+1 6/27/12 12:09 AM 412 -1— 0— +1— SOUTH AND SOUTHEAST ASIA Recent washing and screening in the lower Chitarwata Formation near Dera Bugti led to the discovery of new ruminant material with typically Oligocene taxa (Métais, Welcomme, and Ducrocq 2009). he upper Chitarwata has also yielded ruminant remains whose afi nities remain somewhat obscure. Most eforts made by MPFB in the past decade have focused on the lower Chitarwata, and further investigation is necessary to better determine the diversity and stratigraphic ranges of Neogene ruminants of the Bugti Hills. Ginsburg, Morales, and Soria (2001) reassessed the systematics of ruminants collected by Pilgrim (1908, 1911, 1912) and ForsterCooper (1915), in light of new material collected in 1997– 1998 from the early Miocene strata of the Bugti Hills (essentially the levels 4, uppermost Chitarwata, and 6, basal Vihowa), allowing them to describe six taxa, referred to Giraidae, Tragulidae, or Bovidae. Even if Ginsburg, Morales, and Soria (2001) did not indicate the geographic location of their material, and thus tended to perpetuate errors made by ancient authors, we were able to reassess the stratigraphic allocation of the concerned specimens thanks to their inventory number. Bugtimeryx pilgrimi was erected by Ginsburg, Morales, and Soria (2001), based on mandibular, dental, and postcranial material collected from level 4 in the Bugti Hills (uppermost Chitarwata; see igures 16.3 and 16.4b and table 16.1). In the same article, Gelocus (?) gajensis (Pilgrim, 1912) and Prodremotherium (?) beatrix (Pilgrim 1912) were tentatively referred to Bugtimeryx, and this new genus was included within Giraidae, without further discussion. Barry et al. (2005) and Lindsay et al. (2005) mention B. pilgrimi in several localities referred to the upper Chitarwata (Z151–Z133; see igure 16.4b), but rather consider it as a representative of “Pecora, incertae sedis.” Progirafa exigua occurs in the uppermost Chitarwata (Z126) and the lower Vihowa in the Zinda Pir (Z124–Z205; see igure 16.4b; Barry et al. 2005; Lindsay et al. 2005) and only in the lower Vihowa deposits in the Bugti Hills (level 6; Ginsburg, Morales, and Soria 2001). he type specimen of P. exigua was until recently all that was known of the taxon, but the new collections show that the species is rather common in the Early Miocene deposits of Zinda Pir (see igure 16.4b; Barry et al. 2005). Even if Ginsburg, Morales, and Soria (2001) also consider it as a giraid closely related to Canthumeryx from Gebel Zelten (Libya, late Early Miocene), this taxon is of uncertain ai nities within Ruminantia, following Barry et al. (2005). he earliest Tragulidae from the Bugti Hills occur in the Bugti Member of the Chitarwata Formation at Paali C2 (Early Oligocene; see igure 16.3; Marivaux et al. 49502_1P_03_Wang15012_text14-31.indd 412 2001; Métais, Welcomme, and Ducrocq 2009). Higher in the series, Ginsburg, Morales, and Soria (2001) mention a supposedly primitive species of Siamotragulus (S. bugtiensis) based on mandibular, dental, and postcranial material from the uppermost Chitarwata (see igure 16.3 [level 4, earliest Miocene]). Small mandibular, dental, and postcranial remains from the base of the Vihowa Formation (see igure 16.3 [level 6, late Early Miocene]) are referred to Dorcabune welcommi by Ginsburg, Morales, and Soria (2001). Dorcabune was initially described by Pilgrim (1910, 1915) in the base of the lower Manchar Formation of Sind—that is, in levels considered as coeval to lower Vihowa deposits from the Sulaiman Province (late Early Miocene; for correlation, see igure 16.4b). In Zinda Pir, Barry et al. (2005) and Lindsay et al. (2005) mention unidentiied tragulids in the upper Chitarwata (Z150–Z126, earliest Miocene) and Dorcatherium sp. in the lower Vihowa (Z127–Z116, late Early Miocene; see igure 16.4b). A large taxon documented only by postcranials from the uppermost Chitarwata and the lowermost Vihowa formations in the Bugti area was tentatively referred by Ginsburg, Morales, and Soria (2001) to Tragulidae, under the name “?Siamotragulus indicus (Forster-Cooper 1915).” he corresponding remains most probably document an unidentiied tragulid, since the type specimen (NHM M15421) of “Gelocus indicus Forster-Cooper 1915,” for which the stratigraphic provenance is unknown, rather shows close ai nities with the lophiomerycid Nalameryx sulaimani Métais, Welcomme, and Ducrocq 2009 from the Early Oligocene locality of Paali C2 in the Bugti Hills (Bugti Mb., lower Chitarwata). h is family is restricted to Eurasian Paleogene localities (Métais, Welcomme, and Ducrocq 2009). Interestingly, several Oligocene localities of Zinda Pir have yielded scarce remains of ruminants (Barry et al. 2005). Locality Z108, stratigraphically situated in the lower part of the Chitarwata Formation, has produced an indeterminate lophiomerycid (considered as conspeciic to Nalameryx sulaimani by Métais, Welcomme, and Ducrocq 2009) and ?Gelocus gajensis. he generic status and familial ai nities of ?Gelocus gajensis are still debated, but either of these taxa can be useful for biochronology. We consider as highly dubious the referral of isolated postcranial remains with a tragulid pattern from the upper Chitarwata Formation and the lower Vihowa Formation in the Bugti Hills (levels 4 to 6, early Miocene; Ginsburg, Morales, and Soria 2001) to the probable Oligocene lophiomerycid “Gelocus indicus Forster-Cooper 1915” (see igure 16.3). At locality Z142, stratigraphically equivalent to Z108 (lower member, Chitarwata Formation, Oligocene; Lind- 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN say et al. 2005), Barry et al. (2005) recognized the bovidlike ruminant Palaeohypsodontus zinensis, which is also known in the late Oligocene locality of Lundo Chur J2 in the Bugti area (Métais et al. 2003). here is no mention of this taxon in Miocene deposits of the Bugti area, but it is worth noting that P. zinensis is also reported from the locality Z121, situated in the late Early Miocene Vihowa Formation (Lindsay et al. 2005). h is occurrence may correspond to the LAD of this puzzling genus. Bovids are well represented throughout the Early Miocene of the Sulaiman Province. he earliest undisputable remains (horn core, tooth fragments, and postcranials were unearthed in the uppermost Chitarwata Formation (see igures 16.3 [level 4, earliest Miocene]and 16.4b; reinterpreted ater Ginsburg, Morales, and Soria 2001) and in the lowermost Vihowa (level 6, late Early Miocene) in the Bugti area. All of them are referred to the minute species Eotragus minus (Ginsburg, Morales, and Soria 2001). he occurrence of “Bovidae indet.” in the lower Vihowa (see igures 16.3 [Z110–Z120, late Early Miocene] and 16.4b) dates back to ca. 19.5 Ma among Zinda Pir occurrences (Lindsay et al. 2005). Higher in the Zinda Pir series, Raza et al. (2002) mention Eotragus noyei and cf. Elachistoceras sp. (Vihowa Formation, middle Miocene), while the Litra Formation (Late Miocene) records the occurrence of more advanced forms, such as Prostrepsiceros vinayaki, Hispanodorcas terrubiae, and an unidentiied representative of Reduncini. In the Potwar Plateau, Elachistoceras khauristanensis occurs only in the Late Miocene, with a 11.5–7.4 Ma range, but its inferred FLA is estimated at ca. 14.0 Ma (Barry et al. 2002). In the same area, cf. Prostrepsiceros vinayaki has a short inferred interval (8.6–7.4 Ma) that slightly exceeds its observed time range (8.3–7.9 Ma; Barry et al. 2002). he Reduncini include Recent African reedbucks (Kobus) and waterbucks (Redunca) and their extinct relatives from the Old World, with a late Miocene to early Pleistocene Asian range (McKenna and Bell 1997). No fossil cervid is known from the Neogene of the Bugti Hills and Zinda Pir (Welcomme et al. 2001; Raza et al. 2002; Lindsay et al. 2005), even though an undisputable cervid is portrayed with dichotomic branches in the pre-Holocene cave paintings of Lundo (Crochet et al. 2009). Anthracotheriidae Historically, the classic “Bugti Bone Beds” yielded abundant remains of anthracotheres, which were described in detail by Pilgrim (1912) and Forster- Cooper (1913, 1924). he generic and speciic diversity of anthracothe- 49502_1P_03_Wang15012_text14-31.indd 413 413 riids from Bugti was certainly overestimated by these authors, and upon revision of ancient collections (without any lithostratigraphic control), Pickford (1987) recognized eight species belonging to six genera. Validity of these taxa and implied stratigraphic ranges of the species they would represent in the Bugti Hills remain unclear, making it diicult to use anthracotheriids for biochronologic, paleogeographic, and paleoenvironmental purposes. Anthracothere remains are well represented throughout the Chitarwata Formation in the Bugti Hills, but they tend to be taxonomically more diverse in the upper Chitarwata (Métais et al. 2009). Anthracotherium cf. bugtiense and another large anthracotheriine, as well as the bothriodontine Elomeryx sp., have been found in basal beds of the Chitarwata Formation near Bugti (Early and Late Oligocene; see igure 16.3; Métais et al. 2009). Dental remains of anthracotheres are rare at Paali C2 (basal Chitarwata, Early Oligocene; see igures 16.2 and 16.3), but several postcranial elements suggest the presence of a small species we refer to Microbunodon cf. silistrense. Anthracotherium cf. bugtiense ranges up to duricrust J2 (Late Oligocene) but is absent from the overlying levels M and Q (earliest Miocene; see igure 16.3). Microbunodon is neither registered in the upper Chitarwata (earliest Miocene) nor in the Vihowa Formation (late Early to Middle Miocene) in the Bugti area, while Microbunodon silistrense ranges from the upper Chitarwata (Z151, Z139, Z209) up to the lower Vihowa (Z121, Z120) at Zinda Pir (Lindsay et al. 2005). In the Bugti area, the upper Chitarwata Formation, referred to the earliest Miocene, has yielded an entirely diferent assemblage of anthracotheres, mostly documented by fossils from ferruginous duricrust 4 or Q (see igure 16.3), with dental and postcranial remains of Sivameryx palaeindicus, Hemimeryx cf. blanfordi, Parabrachyodus cf. hyopotamoides, and possibly Telmatodon sp. (see igure 16.4b; Welcomme et al. 2001; Métais et al. 2009). In Zinda Pir, the anthracotheriid assemblage of the upper Chitarwata is somewhat distinct in the absence of Hemimeryx and possibly Telmatodon, and the occurrence of Microbunodon, a genus recorded from the lower Chitarwata in the Bugti area (Lindsay et al. 2005; Métais et al. 2009). Hemimeryx cf. blanfordi and Parabrachyodus cf. hyopotamoides are also recorded in the base of the Vihowa Formation (level 6, late Early Miocene) in the Bugti area. Anthracotheres from Zinda Pir have not been described yet, with the exception of Elomeryx cf. borbonicus from the Oligocene locality Z108 (Ducrocq and Lihoreau 2006). Only a broad revision of the anthracotheriid material from the Sulaiman Province, encompassing both Zinda Pir and Bugti samples, is likely to provide a —-1 —0 —+1 6/27/12 12:09 AM 414 SOUTH AND SOUTHEAST ASIA satisfactory explanation to such a taxonomic/stratigraphic discrepancy. Downing et al. (1993) and Raza et al. (2002) reported the presence of an indeterminate anthracotheriid and remains questionably referred to Masritherium from the Chitarwata Formation, as well as the occurrence of an unidentiied anthracotheriid along with ?Hyoboops sp. (=Sivameryx) in the Vihowa Formation. To our knowledge, no hippopotamid has been mentioned for Neogene deposits of the Sulaiman Province. Suoidea -1— 0— +1— Even though Suoidea are not abundant in terms of number of specimens, all three Eurasian families—that is, Sanitheriidae, Palaeochoeridae (“Old World peccaries”), and Suidae—are represented in mid-Cenozoic deposits of the Sulaiman Province. Remains referable to Sanitheriidae are scarce, but they occur throughout the Chitarwata and Vihowa formations (Early Oligocene–Middle Miocene; see igure 16.4b; Lindsay et al. 2005; Orliac et al. 2010). A right P3 from Paali C2 (Bugti member, Chitarwata Formation, Early Oligocene; see igure 16.3), strongly reminiscent of the holotype of “Hyotherium(?) jef reysi Forster-Cooper 1913” but with a simpler structure, is identiied provisionally as Sanitherium sp. (Hyotherium is a suid, and in our opinion, Sanitheriidae are monogeneric, Diamantohyus Stromer 1926 being a junior synonym of Sanitherium Von Meyer 1866; Orliac et al. 2010). A sanitheriid referred to as “?Diamantohyus jefreysi”—here, “?S. jef reysi”—is recorded in both the lower member (Z144, Oligocene) and the upper member of the Chitarwata Formation in Zinda Pir (Z156, earliest Miocene; Lindsay et al. 2005). S. jeffreysi occurs in coeval deposits from the Bugti Hills, based on a mandibular fragment with m1–3 found at Samane (see igure 16.3 [level 4, earliest Miocene); unpublished data). An unidentiied sanitheriid is also listed in the late Early Miocene of Zinda Pir (Z120, lower Vihowa; Lindsay et al. 2005). Palaeochoeridae (sensu van der Made 1997) are documented unquestionably by a single specimen, an M3 referable to Pecarichoerus sp., from the upper member of the Chitarwata Formation at Samane in the Bugti Hills (see igure 16.3 [level 4, earliest Miocene]; unpublished data). h is specimen strongly recalls the European Taucanamo grandaevum, from the late Early Miocene of France (MN 4; Orliac, Antoine, and Duranthon 2006). h is family has not been mentioned in Zinda Pir so far (Lindsay et al. 2005). he Late Oligocene locality of Lundo J2, in the Bugti Hills (Bugti Member, Chitarwata 49502_1P_03_Wang15012_text14-31.indd 414 Formation; see igure 16.3), yielded recently a mandibular fragment with m2–3 with morphology and measurements strongly comparable to the holotype of “Microbunodon sminthos” igured and described by Forster-Cooper (1913:ig. 5). Pickford (1987) subsequently reassigned this holotype, initially considered as belonging to an anthracotheriid “from the Upper Oligocene deposits of Dera Bugti” (Forster-Cooper 1913:514), to the palaeochoere genus Pecarichoerus Colbert 1933, otherwise known from Middle Miocene localities of the Potwar Plateau (Chinji Formation; Colbert 1933), hailand, and China (Pickford et al. 2004). To date, no specimen documenting this puzzling palaeochoere has been unearthed in Neogene deposits of the Sulaiman Province. Suidae occur throughout the continental series in the Bugti Hills—that is, from the Bugti member of the Chitarwata Formation at Paali C2 (Early Oligocene) up to the level 7 (Litra Formation, Late Miocene; see igure 16.3 and table 16.3). A few specimens from Paali C2, referrable to the hyotheriine Hyotherium, remain unidentiied at species level (Orliac et al. 2010). h is subfamily occurs only in the base of the Chitarwata Formation so far. Higher in the series, Suidae are only represented by Listriodontinae, Tetraconodontinae, and Suinae. Listriodontinae have their FLA in the upper member of the Chitarwata Formation in the Bugti Hills, with Listriodon ainis (see igure 16.3 [level 4, earliest Miocene]; Orliac et al. 2009); this species is also mentioned from the lower part of the Vihowa Formation in Zinda Pir (Z205, late Early Miocene; Lindsay et al. 2005). L. guptai occurs in the base of the Vihowa Formation in both the Bugti Hills (see igures 16.3 [level 6] and 16.4b) and Zinda Pir (Z124, Z210, Z120, and Z205; “L. gupti” of Lindsay et al. 2005). he sublophodont listriodont L. guptai is very close morphologically to L. akatikubas from the late Early Miocene of Maboko, Kenya (ca. 16.5 Ma; Orliac et al. 2009). Fully lophodont listriodonts are found higher in the Vihowa Formation and referred to as either Listriodon pentapotamiae in the Bugti Hills (see igure 16.3 [level W, Middle Miocene]; Welcomme et al. 2001) or Listriodon sp. in the Vihowa Formation (late Early to Middle Miocene) and the Litra Formation (Late Miocene; Raza et al. 2002). he inferred stratigraphic range of L. pentapotamiae in the Potwar Plateau is 14.0–10.3 Ma (Barry et al. 2002:69). As such, the Litra specimens from the Sulaiman Province count among the few attested co-occurrences of listriodontine suids with hipparionine equids and may constitute the Last Local Appearance (LLA) for both Listriodon and Listriodontini (Barry et al. 2002; Raza et al. 2002). 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN Tetraconodontinae are only represented by an m3 from the Litra Formation of the Bugti Hills, identiied as Parachleuastochoerus sp. (see igures 16.3 [level 7, early to middle Late Miocene] and 16.4b and table 16.3; Antoine, Welcomme, et al. 2003). h is genus is known from the late Middle Miocene of Europe (MN 7–8; Golpe Posse 1972) and the early Late Miocene of China (correlated with MN10; Pickford and Liu 2001) Suinae are likely to occur in the Litra Formation (Late Miocene) of Zinda Pir, with ?Propotamochoerus sp. (Raza et al. 2002). he only species referred to this suine genus to be documented in the Potwar Plateau is P. hysudricus, with a Late Miocene inferred stratigraphic range (10.2– 6.5 Ma; Barry et al. 2002:69). Perissodactyla Perissodactyls are particularly abundant in the Neogene of the Bugti Hills, and they provide critical information for the age assessment (Welcomme et al. 2001; Antoine, Ducrocq, et al. 2003; Antoine, Duranthon, and Welmomme 2010; Métais et al. 2009). No equoid is recorded in Chitarwata and Vihowa deposits: Hipparionine equids occur only in the top of the Miocene series—that is, in the Litra Formation (Zinda Pir) and in levels 7 and 7sup (Sartaaf = Djigani; Bugti Hills)—in deposits probably equivalent in age to the Dhok Pathan Formation of the Potwar Plateau (see igure 16.3; DB7; Welcomme et al. 1997; Antoine, Duranthon, and Welmomme 2003; Zouhri and Ginsburg 2003). he hipparionine teeth from Djigani recall those of Hippotherium nagriense (Nagri Formation; early Late Miocene) and Cormohipparion (Sivalhippus) theobaldi from the lower part of the Dhok Pathan Formation of Potwar Plateau, which indicates an age earlier than middle Turolian (Zouhri and Ginsburg 2003). Hipparion sensu lato in Zinda Pir is recorded at Z168—that is, 1500 m above the base of the Chitarwata Formation, at ca. 10.5 Myr (see igure 16.3; Raza et al. 2002). Raza et al. (2002) mention C. (S.) theobaldi in the same formation, without further precision concerning its stratigraphic range. Ancylopoda are uncommon but present throughout the post-Eocene series in the Bugti Hills (Métais et al. 2009) and in the Zinda Pir Dome (Raza et al. 2002; Lindsay et al. 2005). In the latter area, all the specimens remain unidentiied. In the Bugti Hills, the large schizotheriine Phyllotillon naricus is documented by dental and postcranial remains from the upper part of the Bugti member of the Chitarwata Formation (Oligocene in age) up to the top of the formation (level 4 (=Q ); earliest Miocene), while the smaller chalicotheriine “Chalicothe- 49502_1P_03_Wang15012_text14-31.indd 415 415 rium” pilgrimi seems to be restricted to the upper member, in levels 3bis (=M) and 4 (Métais et al. 2009). Higher in the series, a few postcranial and dental specimens unearthed from the middle part of the Vihowa Formation in the Bugti Hills (level W; Middle Miocene) are similar to specimens from Sansan in France and thus referable to Anisodon sp. (Anquetin, Antoine, and Tassy 2007). In the Sulaiman Province, the bulk of Neogene perissodactyls is constituted by rhinocerotids, for which nine or ten species are recognized in the upper part of the Chitarwata Formation, seven or eight species in the Vihowa Formation, and at least three in Litra Formation or coeval deposits (see tables 16.1–16.3 and igure 16.4; Welcomme et al. 1997; Raza et al. 2002; Lindsay et al. 2005; Antoine et al. 2010). Most rhinocerotid suprageneric groups recognized in the Old World occur in the Neogene of the area (see igure 16.4). Rhinocerotinae are much diversiied throughout the concerned period, with 12 species referred to Rhinocerotina, Aceratheriina, Teleoceratina, and Rhinocerotinae incertae sedis; on the other hand, Elasmotheriinae are represented by a single species (Antoine, Duranthon, and Welmomme 2003; Antoine et al. 2010). Four basal ofshoots of the Rhinocerotinae were unearthed in the upper Chitarwata Formation of both the Bugti Hills and Zinda Pir (earliest Miocene): Protaceratherium sp., Pleuroceros blanfordi, Mesaceratherium welcommi, and Plesiaceratherium naricum. he latter may also occur in the basal Vihowa Formation in the Bugti Hills (see table 16.2). P. naricum is the earliest representative of a well-known Eurasian genus, so far restricted to the late Early–early Middle Miocene interval (Yan and Heissig 1986; Antoine et al. 2000). Pleuroceros blanfordi and Mesaceratherium welcommi occur in the upper member of the Chitarwata Formation in the Bugti Hills and the Zinda Pir Dome and in the basal Vihowa Formation in the Bugti Hills (Early Miocene; see tables 16.1 and 16.2; Welcomme et al. 2001; Antoine et al. 2010). Both species are endemic to the Sulaiman Province, but they are sister taxa of the European P. pleuroceros and M. paulhiacense, respectively, from the earliest Miocene of Western Europe (Antoine et al. 2006, 2010). Rhinocerotina include all ive living rhino species; their fossil record is restricted to the Old World. he earliest representatives of the clade are restricted to Pakistan until the late Early Miocene (Gaindatherium; Antoine et al. 2000; Antoine et al. 2010). Gaindatherium cf. browni, recognized in the upper member of the Chitarwata Formation in the Bugti Hills (earliest Miocene), widely predates the FLA of other representatives of this onehorned genus elsewhere (Heissig 1972). An early putative representative of the extant genus Rhinoceros is —-1 —0 —+1 6/27/12 12:09 AM 416 -1— 0— +1— SOUTH AND SOUTHEAST ASIA recognized in the lower part of the Vihowa Formation (late Early Miocene, the Bugti Hills and Zinda Pir; Antoine et al. 2010). Undescribed specimens recovered from the base of the Kamlial Formation in the Potwar Plateau and referable to the same taxon (obs. by POA) help for correlating both intervals (see igure 16.3). Higher in the series, a partial maxilla with M1–3 unearthed in the level 7 (Djigani, the Bugti Hills; see igure 16.3) is referred to Rhinoceros af. sivalensis. Undescribed specimens from the lower Dhok Pathan Formation in the Potwar Plateau document the same taxon. hus, a middle Late Miocene age can be hypothesized for Djigani locality (Barry et al. 2002). Teleoceratina are hippo-like extinct rhinos, with shortened limb bones adapted to swamps and riversides. Prosantorhinus shahbazi is restricted to the earliest Miocene deposits of the Bugti Hills (see table 16.1; Antoine et al. 2010). Prior to the latter recognition, the genus was only recorded in the late Early and early Middle Miocene of Western Europe (e.g., Antoine et al. 2000). h ree representatives of the large and robust genus Brachypotherium are known. B. fatehjangense and B. gajense are recorded from the upper Chitarwata Formation and the lowest Vihowa Formation in both the Bugti Hills and Zinda Pir (Early Miocene; see table 16.1; Welcomme et al. 2001; Antoine et al. 2010). B. gajense is restricted to this stratigraphic interval, while B. fatehjangense persists until the Late Miocene in the Potwar Plateau (middle Dhok Pathan Formation; new data). In the Sulaiman Province, B. perimense ranges from the base of the Vihowa Formation in Zinda Pir (late Early Miocene; see table 16.1) up to level 7 in the Bugti Hills (middle Late Miocene; Antoine, Duranthon, and Welmomme 2003). New data from the Potwar Plateau provide similar ranges (LAD at ca. 7.1 Ma; Barry et al. 2002; POA, personal observation). Bugti and Zinda Pir remains document the earliest occurrences of Prosantorhinus and Brachypotherium at Eurasian and Old World scales, respectively. Aceratheriina are extinct hornless rhinos, widespread in the Miocene of North America and Eurasia and in the Miocene of Africa. he only aceratheriine sensu stricto recognized in the Sulaiman Province is Alicornops complanatum, which occurs in level 7 of the Bugti Hills (Litra Formation, middle Late Miocene; Antoine, Duranthon, and Welmomme 2003). h is taxon is abundant throughout the Late Miocene Dhok Pathan Formation in the Potwar Plateau (Colbert 1935; Heissig 1972; new data). he Elasmotheriinae are the sister group of Rhinocerotinae. hey are well represented in the Neogene of Eurasia and the Miocene of Africa (Antoine 2002). Bugtirhi- 49502_1P_03_Wang15012_text14-31.indd 416 nus praecursor is the earliest elasmotheriine known so far. h is primitive species is restricted to the upper Chitarwata deposits of the Bugti Hills (levels 3bis and 4, earliest Miocene; see table 16.1) and to the base of the overlying Vihowa Formation of Zinda Pir (Z116, late Early Miocene; see table 16.2 and igure 16.4; POA, personal obs.). NEOGENE FAUNAL SUCCESSION IN THE SULAIMAN PROVINCE AND BIOSTRATIGRAPHICAL CORRELATION In the present work, biostratigraphical correlation between the Bugti and Zinda Pir areas is mostly based on First Local Appearances (FLA) and observed ranges of hoofed mammals (rhinocerotids, proboscideans, artiodactyls, and hipparionine equids) for the whole OligoceneMiocene series, as well as rodents for the early Miocene period (see igures 16.3 and 16.4). hese assemblages are widely homotaxic, at the generic and/or species level (see tables 16.1–16.3). In the Sulaiman Province, the best-documented stratigraphic interval spans the Chitarwata Formation and overlying Vihowa Formation (Oligocene to Middle Miocene), while the Potwar Plateau in northern Pakistan yields essentially Neogene deposits, among which Middle and Late Miocene faunas (Chinji, Nagri, and Dhok Pathan formations) are far better known than Early Miocene faunas (Murree and Kamlial formations; Pilbeam et al. 1979; Barry, Lindsay, and Jacobs 1982; Bary et al. 2002). Such a situation does not facilitate correlating the concerned areas. he tentative correlation to GPTS for Neogene deposits of the Sulaiman Province is primarily based on the “Interpretation B” of Lindsay et al. (2005:ig. 6B), with revised ages for chrons 6C to 5C (Gradstein et al. 2004). h is interpretation is by far the most satisfactory for concerned mammal assemblages (i.e., without hiatus between the top of the Chitarwata Formation and the base of the Vihowa Formation) within C5En.2n, at ca. 19.4 Ma (“C6n” in Lindsay et al. 2005). As a result, four successive faunal assemblages (A to D from old to young) are recognized in the Neogene of the Sulaiman Province, mainly constrained by perissodactyls (rhinocerotids and hipparionine equids) and rodents (cricetids, rhizomyids, and ctenodactylids) and, to a lesser degree, by deinotheriid proboscideans, sanitheriid and listriodontine suoids, anthracotheriids, and most ruminant groups (tragulids, bovids, and Pecora incertae sedis; see igure 16.4). 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN 417 Assemblage A (Upper Chitarwata Formation Assemblage: Earliest Miocene) Assemblage B (Lower Vihowa Formation Assemblage: Late Early Miocene) A major turnover is observed within the Chitarwata Formation in the Sulaiman Province. Lower in the series— that is, in the lower Member (Zinda Pir) and Bugti Member (Bugti Hills)—occurs the Last Local Appearance (LLA) of many groups and genera of Oligocene ainities (see igure 16.3): the rodent Downsimys and most baluchimyines (Lindsaya, Lophibaluchia, Hodsahibia, and Asterratus; Lindsay et al. 2005), Entelodontidae, the anthracotheriids Anthracotherium, Bugtitherium, and Elomeryx, the chalicotheriid Schizotherium, Hyracodontidae (Paraceratherium bugtiense), Amynodontidae (Cadurcotherium indicum), and early rhinocerotids such as Epiaceratherium cf. magnum, Aprotodon smithwoodwardi, “Dicerorhinus” abeli, and a close ally of Diceratherium (Antoine, Duranthon, and Welmomme 2003; Antoine et al. 2004; Lindsay et al. 2005; Métais et al. 2009). h is assemblage is assumed to predate the Oligocene-Miocene transition. Assemblage A coincides with the appearance of a totally renewed assemblage in the upper Member of the Chitarwata Formation (Z113 and Z139 and localities above in Zinda Pir; levels 3bis and 4 in the Bugti area; Lindsay et al. 2005; Métais et al. 2009; Antoine et al. 2010). he rodent fauna is broadly renewed, with FLAs of Eumyarion, Copemyinae (Democricetodon, Spanocricetodon, and Primus), Rhizomyinae (Prokanisamys), and Ctenodactylinae (Prosayimys). Large mammals having their FLA in this assemblage are the large carnivore Amphicyon, Deinotheriidae (Prodeinotherium), Gomphotherium, Listriodontini suids (Listriodon), the anthracotheriids Telmatodon and Hemimeryx, the tragulid Dorcatherium, Bovidae (Eotragus), the ruminant Bugtimeryx, the chalicotheriids “Chalicotherium” and Phyllotillon, and the rhinocerotids Protaceratherium, Mesaceratherium, Pleuroceros, Plesiaceratherium, Brachypotherium, Prosantorhinus, Gaindatherium, and Bugtirhinus (see i gure 16.3). h is stratigraphic interval resembles the Gaj Formation in Sind (Métais et al. 2009) and perhaps the poorly documented base of the Murree Formation in the Potwar Plateau (Barry et al. 2002). Given the faunal content and favored correlation to GPTS (see igure 16.4), this assemblage is tentatively correlated to the earliest Neogene standard age, the Aquitanian, and roughly correlated with the Xiejian Chinese Land Mammal Age (CLMA) and the Agenian European Land Mammal Age (ELMA), MN 1–2. he Chitarwata-Vihowa formation transition is marked by the LLA of the cricetids Eumyarion and Primus, the ctenodactylid Prosayimys, the sanitheriid suoid Sanitherium, the ruminant Bugtimeryx, and several rhinocerotids, such as Protaceratherium sp., Plesiaceratherium naricum, Prosantorhinus shahbazi, and Gaindatherium cf. browni, which only occur in the Chitarwata Formation (see igures 16.3 and 16.4). h is transition is supposedly coeval to both the Aquitanian-Burdigalian transition (marine standard scale) and the Agenian-Orleanian European Land Mammal Ages transition. Assemblage B (lower Vihowa Formation) documents the FLA of key taxa such as the derived muroids Megacricetodon and Myocricetodon, the ctenodactylid Sayimys intermedius, the diatomyid Diatomys, the creodonts Pterodon and Hyanailouros, the carnivore Megamphicyon, the elephantoids Protanancus and Choerolophodon, Listriodon guptai, the tragulid Dorcabune, the “girafoid” Progirafa, and the rhinocerotids cf. Rhinoceros sp. and Brachypotherium perimense. Based on rhinocerotids and early bovids, this stratigraphic interval resembles the lower Manchar Formation in Sind (Raza et al. 1984) and the Kamlial Formation in the Potwar Plateau (Barry et al. 2002). Given the faunal content and favored correlation to GPTS (see igure 16.4), this assemblage is considered late Early Neogene standard age, the Burdigalian, and roughly correlated with the Shanwangian CLMA and the Orleanian ELMA (MN 3–5). 49502_1P_03_Wang15012_text14-31.indd 417 Assemblage C (upper Vihowa Formation assemblage: Middle Miocene) Assemblage C is only documented by a few large mammals, and as such, it is not well constrained in terms of biostratigraphy. he concerned interval yields the FLA of Deinotherium, Listriodon pentapotamiae, the bovid cf. Elachistoceras, and the chalicotheriid Anisodon (see i gure 16.3). h is stratigraphic interval resembles the upper Manchar Formation in Sind and the Chinji Formation in the Potwar Plateau (Barry et al. 2002). Given the faunal content and favored correlation to GPTS (see i gure 16.4b), this assemblage is tentatively considered early Middle Miocene, late Langhian– early Serravallian standard ages, and roughly correlated with the Tunggurian CLMA and the Astaracian ELMA (MN 6). —-1 —0 —+1 6/27/12 12:09 AM 418 SOUTH AND SOUTHEAST ASIA Assemblage D (Litra Formation Assemblage: Late Miocene) In stratigraphic terms, the Vihowa–Litra formation transition is not well constrained, due to the scarcity of available localities and remains across the whole Sulaiman Province (Welcomme et al. 2001; Raza et al. 2002; Antoine, Welcomme, et al. 2003; Zouhri and Ginsburg 2003). h is assemblage consists only of large mammals (see igures 16.3and 16.4b), with the FLA of tetraconodontine and suine suids (Parachleuastochoerus and ?Propotamochoerus, respectively), of giraids sensu stricto (?Bramatherium), of advanced bovids (Hispanodorcas, Prostrepsiceros, and an unidentiied reduncine), of hipparionine equids (Cormohipparion (Sivalhippus) theobaldi and Hippotherium sp.), and of the rhinocerotids Alicornops complanatum and Rhinoceros af. sivalensis and the LLA of listriodontines (Listriodon sp.). he concerned fauna strongly resembles the magnetostratigraphically constrained assemblages recognized in the upper part of the Nagri Formation and the lower part of the Dhok Pathan Formation in the Potwar Plateau (Heissig 1972; Pilbeam et al. 1979; Barry et al. 2002). As such, this assemblage might be early Late Miocene standard age (late Tortonian) and is tentatively correlated with the late Bahean CLMA interval and the late Vallesian– early Turolian ELMA interval (MN 10–12). FIRST LOCAL APPEARANCES AS “DATUMS” Several mammalian taxa have their earliest occurrences in mid-Cenozoic deposits of the Sulaiman Range, either at tribe, family, or even at order level. In this section, we have chosen to focus on key taxa used at a large scale for biochronology and dispersal events, and well represented in the Neogene of the Sulaiman Province, such as proboscideans, suoids, bovids, and rhinocerotids. Proboscidean Datum Event(s) (Bugti Hills/Zinda Pir) -1— 0— +1— Early specimens referrable to elephantoids discovered in the Bugti Hills originate from the Lundo section (Welcomme et al. 2001; Antoine, Welcomme, et al. 2003; Métais et al. 2009). he locality DB-J1 where the i rst tusk was found (Antoine, Welcomme, et al. 2003) is located ca. 40 m below the classical Lundo locality (level J2 = “Chur Lando” of Pilgrim 1908, 1910; Forster-Cooper 1924, 1934), and about 55 m below the Chitarwata–Vihowa formation 49502_1P_03_Wang15012_text14-31.indd 418 transition (see igure 16.3). Tabbutt, Sheikh, and Johnson (1997) provided a ission track date of 22.6 ± 2.9 Myr for the yellow sands of “Chur Lando” (i.e., level J2). Locality DB-J1 is necessarily older than 19.7 Myr (and may date back to 25.4 Myr). he second tusk was found in a still older locality, referred to the level F, ca. 20 m lower in the same section (see igure 16.3). In the Zinda Pir area, Lindsay et al. (2005) mention both “Elephantoidea Indet. genus, indet. species” and “Proboscidea Indet. genus, indet. species” in locality Z108, located in the lower member of the Chitarwata Formation, but we now think these specimens are not dei nitive. A younger specimen at locality Z154 is clearly an elephantoid and found midway in a signiicant normal magnetochron, currently considered chron C6Br, and therefore likely somewhat younger than 23 Ma (Flynn et al., chapter 14, this volume). hese early occurrences alter considerably the concept of Proboscidean Datum Event as documenting a single dispersal of proboscideans out of Africa in the late Early Miocene (ca. 17.5 Ma; Madden and Van Couvering 1976). Proboscideans rather dispersed several times from Africa to Eurasia during the Oligocene and the Early Miocene, as already argued by Tassy (1990). he earliest “diversiied proboscidean fauna” from the Sulaiman Province occurs in the upper Chitarwata Formation, with the FLA of both Deinotheriidae (see igure 16.3 [levels 3bis and 4 in the Bugti Hills, Z129 in Zinda Pir]) and Gomphotherium sp. (see igure 16.3 [level 4 in the Bugti Hills]; Welcomme et al. 2001; Métais et al. 2009). Lindsay et al. (2005) correlate Z129 with C6Bn (ca. 22.5 Ma; Gradstein et al. 2005) and the level 4 may date back to ca. 21 Ma (see igure 16.4b). Sanitheriidae he sanitheriid Sanitherium sp. was recently recognized in the Bugti member of the Chitarwata Formation at Paali C2 (Early Oligocene, ca. 28 Ma; see igure 16.3; Orliac et al. 2010). Sanitherium jef reysi is documented in the upper Member of the Chitarwata Formation at Samane 4 (Early Miocene, ca. 21 Ma; see igure 16.3; Orliac et al. 2010). “?Diamantohyus jef reysi” is mentioned from coeval deposits of the lower Chitarwata Formation in Zinda Pir (Z144; Lindsay et al. 2005). Z144 is located in the same chron as Z108, but higher (C7n or C10n.2n; Lindsay et al. 2005), dating to 24.6 Ma or even 28.6 Ma (see section “Proboscidean Datum Event(s)”). Both Oligocene occurrences widely predate the previous worldwide FAD of the family, so far considered as 6/27/12 12:09 AM MAMMALIAN NEOGENE BIOSTRATIGRAPHY OF THE SULAIMAN PROVINCE, PAKISTAN occurring during the Early Miocene in Africa (Sperrgebiet, Namibia, ~21–19 Ma; Pickford and Senut 2000, 2002). Such an early settlement of sanitheres in the Indian Subcontinent strongly supports the provocative hypothesis of a late Oligocene–earliest Miocene dispersal event from South Asia toward Africa, rather than in the opposite direction (Orliac et al. 2010). Bovidae he lowest bovid-yielding locality in Zinda Pir is Z120 (lowermost Vihowa), which is correlated with C5En (Lindsay et al. 2005)—that is, estimated at ca. 18.4 Myr (see igure 16.4b; Gradstein et al. 2005). he horncore referred to Eotragus minus in the Bugti area (Ginsburg, Morales, and Soria 2001) was recovered from coeval deposits (level 6, lowermost Vihowa Formation; see igure 16.4b). Both occurrences predate the base of the Kamlial Formation in the Siwalik Group (~18 Ma), where Solounias et al. (1995) have described Eotragus noyei, the oldest representative of the family then known. Furthermore, the uppermost Chitarwata Formation in the Bugti Hills (level 4) yields several unambiguous bovid postcranials (Ginsburg, Morales, and Soria 2001). Given the favored correlation hypothesis between Zinda Pir and the Bugti area, these remains may date back to ca. 21 Ma (see section “Proboscidean Datum Event(s)” and igure 16.4b). Early Miocene Rhinocerotidae and the “African Rhinocerotid Datum” he rhinocerotid fauna from assemblage A (upper Chitarwata Formation, earliest Miocene: 23–19.4 Ma; see igure 16.4b) is exceptionally diversiied, with nine cooccurring species in the Bugti Hills: the early elasmotheriine Bugtirhinus praecursor and the hornless rhinocerotines Protaceratherium sp. and Plesiaceratherium naricum, the basal rhinocerotines Pleuroceros blanfordi and Mesaceratherium welcommi, the teleoceratines Brachypotherium gajense, B. fatehjangense, and Prosantorhinus shahbazi, and the rhinocerotine Gaindatherium cf. browni (Antoine and Welcomme 2000; Antoine et al. 2010). Coeval homotaxic rhinocerotid assemblages (at genus level) are recorded from the Agenian ELMA of France (MN 1 to MN 2: 23–20 Ma interval; Gradstein et al. 2005), with Protaceratherium minutum, Plesiaceratherium aquitanicum, Pleuroceros pleuroceros, and Mesaceratherium paulhiacense (de Bonis 1973; Antoine et al. 2006; Antoine et al. 2010). 49502_1P_03_Wang15012_text14-31.indd 419 419 Interestingly, the earliest representatives of Rhinocerotidae in Africa are strongly comparable to those of assemblage A, which persist into assemblage B (see igure 16.4b): Brachypotherium heinzelini and Aceratherium acutirostratum, recognized in Napak II and Songhor (Hooijer 1966, 1973; Hooijer and Patterson 1972), as well as Ougandatherium napakense from Napak I (Guérin and Pickford 2003), are strongly comparable to the teleoceratine Brachypotherium fatehjangense, the acerathere Mesaceratherium welcommi, and the earliest elasmotheriine Bugtirhinus praecursor, respectively (Antoine et al. 2010). he radiometric age of Songhor is ~19.5 Ma (Pickford 1986; Cote et al. 2007) and Napak might be slightly older (Tassy 1986; Cote et al. 2007), which coincides with the assemblage A–assemblage B transition in terms of age (ca. 19.4 Ma; see igure 16.4b). he close ai nities of South Asian, African, and European rhinocerotid assemblages coni rm both the presence of land bridges and the absence of ecological barriers between these continental areas during the Early Miocene. AC KNOW LEDG MENTS he authors warmly thank Xiaoming Wang, Deng Tao, Li Xiang, Mikael Fortelius, and all the Organizing Committee of the symposium “Neogene Terrestrial Mammalian Biostratigraphy and Chronology in Asia” held in Beijing in June 2009. Everett H. Lindsay provided highly valuable comments on a previous version of the manuscript. We are grateful to Francis Duranthon, Mouloud Benammi, Jean-Jacques Jaeger, Yaowalak Chaimanee, and Dario De Franceschi for their participation in ieldwork, and to Michèle E. Morgan, John C. Barry, David Pilbeam, Everett H. Lindsay, and Iqbal U. Cheema for their valuable help and discussion. h is article is dedicated to the memory of Will Downs, Nawab M. A. K. Bugti, and Léonard Ginsburg. h is research was supported by the French ANR-PALASIAFRICA Program (ANR- 08-JCJC- 0017 - ANR-ERC). MPFB Publication no. 39. REFERENCES Anquetin, J., P.- O. Antoine, and P. Tassy. 2007. 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