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
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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
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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)
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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).
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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.
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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).
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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.
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Table 16.2
Mammals from the Vihowa Formation (Late Early to
Middle Miocene) of the Sulaiman Province
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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.
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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.
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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.
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410
SOUTH AND SOUTHEAST ASIA
Rodentia
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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
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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).
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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-
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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
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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
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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).
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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
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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).
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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).
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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)
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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
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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.
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