1988 Bernor et al - Dionysopithecus Miocene of Pakistan

Raymond L. Bernor College of Medicine, Department of Anatomy, Laboratory of Paleobiology,., Howard University, Washington, D.C. 20059, U.S.A.

Lawrence J. Flynn Peabody Museum, Harvard University, Cambridge, MA 02138, U.S.A.

Terry Harrison Department of Anthropology, New York University, New York, NY 10003, U.S.A.

S. Taseer Hussain College of Medicine, Department of Anatomy, Laboratoy of Paleobiology, Howard University, Washington, D.C. 20059, U.S.A.

and Jay Kelley” Division of Biology and Medicine, Box G, Brown University. Providence, RI 02912, U.S.A.

Received 27 August 1987 Revision received 3 February 1988 and accepted 15 February 1988

Publication date June 1988

Keywords: Catarrhine, Miocene, biogeography, biochronology, Pakistan

* Author order alphabetical

Dionysopithecus from southern Pakistan and the biochronology and biogeography of early Eurasian catarrhines

New specimens of a small, advanced catarrhine primate from the Manchar Formation in Sind, southern Pakistan, are referred to Dionysopithecus sp. Their age is biochronologically estimated to be close to the early/middle Miocene boundary. Dionysopithecu is considered closely related to, and possibly congeneric with, Micropithecusfrom the East African early Miocene. The Manchar Dionysopitheczs is among the earliest of Eurasian catarrhines. Catarrhines may have first emigrated from Afro-Arabia around 16.5 Ma, coincident with a major short-term eustatic sea level lowering event, and with the earliest records in South Asia of certain other African mammal groups. The first appearances in Eurasia oflater, more advanced catarrhine lineages also appear to correlate with episodes of global sea level lowering.

Journal of Human Evolution (1988) 17, 339-358

Introduction

Recent geological and paleontological surveys of the Manchar Formation in Sind, Southwest Pakistan, were undertaken at the Gaj River near the town of Dadu and in the hills surrounding the town of Sehwan (Figure 1). In 1975-1976 the Yale Geological Survey of Pakistan Siwalik project (Y-GSP) recovered a diverse vertebrate sample, including a small catarrhine upper canine from their locality S2 (Primates, incertae sedis; Raza et al.,

1984). In 198 1, members of the Howard-Geological Survey of Pakistan project (H-GSP), found a left P4 and a right Mi of a small catarrhine at locality H-GSP 8114. Both localities are low in the Manchar Formation and may represent the same collecting horizon. The three specimens probably represent a single species, Dionyso@thectlr sp.

These small catarrhines are important from a number of standpoints. Biochronologically and biostratigraphically they and their associated fauna correlate with Kamlial Formation faunas in the Potwar Plateau Siwalik Group that are dated magnetostratigraphically at 17 to 16 Ma (megaannum; see Johnson et al., 1985; Barry et al., 1985). They are also approximately contemporaneous with catarrhine-bearing faunas

0047-2484/88/030339 + 20 $03.00/O @ 1988 Academic Press Limited

340 R. L. BERNOR ETAL

PAKISTAN

SHAH

Figure 1. Locations of the Gaj and Sehwan sections of the Manchar Formation, as well as Bugti, Banda Daud Shah (Murree and Chinji Formations), and the Potwar Plateau CPG section. For geographical reference. the Sehwan section is on the east shore of Lake Manchar.

from China, correlated to Orleanian biochron MN 4 (see below), estimated to be ca 17 Ma (see Mein, 1985). They are therefore among the oldest known fossil catarrhines outside of Africa. Biogeographically they appear to represent part of a diverse African mammalian community, some members of which extended their range into southern Asia during the late early Miocene.

Geology and biochronology of the sites

The Manchar Formation is broadly exposed on the borders of Lake Manchar, along the foothills of the Kirthar Mountain Range and in areas between the Kirthar Range and the Indus River (Raza et al., 1984). The Gaj (Dadu) and Sehwan sections have been most extensively studied. The entire Gaj section is several kilometers thick and contains five superimposed formations: Kirthar, Nari, Gaj, Manchar and Dada conglomerate, ranging from Eocene through Pliocene age(?). In contrast, the Sehwan section, where the primate specimens occur, is substantially shorter and contains at its base only the uppermost Kirthar Fm. and short, partial sections of Nari and Gaj Fms. below the Manchar Fm. In both areas the uppermost Gaj Fm. has a near shore estuarine facies with the lowermost Manchar Fm. marking the onset of continental sedimentation. Raza et al. (1984) measured the Manchar Beds near Sehwan (termed Bhagothoro section by them) and reported a vertical thickness of 430 m, in contrast to a vertical thickness of more than 2200 m in the Gaj River section. The H-GSP team measured live stratigraphic sections near Sehwan,

EARLY EURASIAN CATARRHINE PRIMATES 341

Ma

POTWAR BANDA DAUD SHAH,

GAJ

T 1 H-GSP

I I

116

-i- 1 H-GSP 8106

L

I 5 I I I GAJ FM. I

I BUGTI

i

BUGTI

i I I H-GSP 107

GAJ FM.

Figure 2. Correlation ofisolated early and middle Miocene fossil localities to the Potwar CPG sequence (left, chevrons indicate fossil horizons). Imprecision in correlation is indicated by vertical bars, but note that H-GSP 8106 is faunally intermediate between 116 and 8114A, and should not overlap either temporally. Bugti is placed with the two Banda Daud Shah sites to show that it predates both.

SEHWAN

T H-GSP 8224

I , H-GSP 8227

MANCNAR FM.

T ,H-GSP 8114, 52

i H-GSP 8114A

which varied in thickness from 740 m to 280 m. The variable thickness of the different Sehwan sections is due mostly to recent erosion. Raza et al. (1984) argued that attenuation of the Sehwan section relative to the Gaj River section may be due either to major unconformities resulting from periods of nondeposition or to lower sedimentation rates, particularly the latter. Both factors probably contributed to the shorter Sehwan section, but major unconformities may be the dominant cause, based on apparent gaps in the fauna1 sequence (de Bruijn & Hussain, 1984).

Most of the vertebrate localities, including the Dionysopithecus sp. localities S2 and H-GSP 8114 (Figure 2), occur in the lower half of the Manchar Formation (de Bruijn & Hussain, 1984, 1985; Bernor, 1984; Raza et al., 1984). Presently only biochronology provides an estimate for the age of the Manchar levels where the Dionysopithecus specimens occur. De Bruijn & Hussain (1984) analyzed microfaunas from six Manchar Fm. localities, including H-GSP 8114. In Table 1, microfaunas from three of these sites, H-GSP 8106, 8 114A, and 8 I 14, plus H-GSP 116 at the base of the Murree Fm. at Banda Daud Shah, are compared to those from two magnetostratigraphically dated Siwalik micromammal

342 R. L. BERNOR ET AL.

Table 1 Rodents from Potwar localities Y721 and Y592, and from H-GSP localities 116,8 106,8114A and 8114. H-GSP lists are modified from those of de Bruijn & Hussain (1984) and de Brnijn et ~2. (1981). Unique or indeterminate species have been omitted. Potwar cricetids are under study by E. H. Lindsay, University of Arizona. Age decreases from left to right.

Taxa 116 8106 Y721 8114A 8114 Y592

Sciuridae Sayimys minor Sayimys aff. S. minor Sayimys aff. S. sivalensisl Prokanisamys arij Prokanisamys n. sp.2 Prokanisamys betjavuni2 Kanisamys indicus3 Spanocricetodon lii Democricetodon spp, “Me.gacricetodon” spp4 Myocricetodon spp. Dendromurinae5 Thryonomyidae (small)

X X X X X x

X X X X

X X X X X X

X X X X

X X X X X X X X X X X X

X X ? X

X X

1 Forms of Sayimys follow usages of de Bruijn & Hussain (1984); identifications for Y721 and Y592 are provisional.

2 Forms of Prokanisamys are documented in the Potwar record. Their presence at 8106, 8114A and 8114 is inferred from forms designated Kanisamys sp. by de Bruijn & Hussain (1984).

3 The 8114 record was only conferred to K. indicus by de Bruijn & Hussain (1984). 4 Potwar and Sind localities record the same species of this group, but de Bruijn & Hussain (1984) ally these

forms with Myocricetodontinae. 5 We consider Antemus sp. from 8114 (de Bruijn & Hussain, 1984) to be an advanced muroid, perhaps a

dendromurine, and like one from Y592.

localities, Y721 (ca 18 Ma) and Y592 (16.1 Ma), f rom the Potwar Plateau. The proposed tempotal sequence of localities was established primarily according to “stage of evolution” criteria for species of the rodent genera Sayimys, Prokanisamys, and Kanisamys. H-GSP 8114 overlies 8 114A stratigraphically.

H-GSP 116 is early Miocene in age (de Bruijn et al., 1981; de Bruijn & Hussain, 1984) and is certainly younger than Dera Bugti, which has an archaic microfauna Uacobs et al., 1981; Flynn et al., 1986). Bugti has been considered to reflect the regional “Proboscidean Datum” at ca 17.5 Ma (Berggren & Van Couvering, 1974), but workers now concur on a 20-t Ma age (Adams et al., 1983; Bernor, 1983, 1984; Whybrow, 1984; Raza & Meyer, 1984; Raza et al., 1984; Barry et al., 1985; Tassy, 1985). Further issues surrounding the age and significance of the Bugti beds are discussed by Bernor et al. (1987) and Pickford & Rogers ( 1987). Flynn et al. (1986) correlated H-GSP 116 with the base of the Potwar Chita Parwala-Gabhir (CPG) section, dated paleomagnetically at 18.3 Ma (Johnson et al., 1985).

De Bruijn & Hussain (1984) considered H-GSP 8 106 to be somewhat younger than, but close in age to, H-GSP 116, as both sites contain Sayimys minor, Prokanisamys a$, and Spanocricetodon lii. Y72 1, at ca 18 Ma, is younger than either, based on the presence of larger and more advanced species of Sayimys, Prokanisamys, and cricetids. H-GSP 8114A appears to be still younger based on the presence of a small thryonomyid and a Democricetodon similar to D. kohatensis known from Y592 and younger sites. However, the presence of P. ariji at H-GSP 8114A, but not at Y721, is an anomaly. H-GSP 8114 and Y592 correlate well and both are younger than H-GSP 8114A. In addition to Dionysopithecus sp., the two sites share Prokanisamys benjavuni, Kanisamys indicus, advanced myocricetodontines, and perhaps

Table

2

Dim

en

sion

s of

Seh

wan

teeth

refe

rred

to D

ionyso

~it

hecus s

p.,

an

d c

om

pari

son

s w

ith o

ther

small

cata

rrhin

es

(MD

, m

esi

odis

tal

len

gth

; B

L,

buccoli

ngual

bre

adth

; B

HT

, buccal

heig

ht)

C’

P*

Ml

MD

B

L

BH

T

MD

B

L

MD

B

L

N

?C

Ran

ge

N

zi

Ran

ge

N

%

Ran

ge

N

%

Ran

ge

N

x R

ange

N

%

R

ange

N

x

Ran

ge

Dio

nyso

pith

ecus

sp,

Pro

plio

pith

ecus

ch

irob

ates

l

Aeg

ypto

pith

ecus

ze

uxis

l P

liopi

thec

us

antiq

uusz

M

icro

pith

ecus

cl

arki

Li

mno

pith

ecus

le

gete

t3

Lim

nopi

thec

us

evan

si

Den

drop

ithec

us

mac

inne

si3

1 4.

9 4.

3 1

3.8

3.8

1 6.

6 66

1

3.6

3.6

1 5.

6 5.

6 1

5.4

5.4

1 4.

9 4.

9

1 4.

2 4.

2 1

4.2

4.2

2 3.

3 3.

2-3-

3 2

5.9

5.9

7 5.

0 4.

7-5.

1 7

4.8

4.55

.0

3 3.

9 3.

3-4.

5 3

6.9

6.4-

7.1

10

5.9

56-6

.1

10

5.3

4Wj.l

3 6.

9 5G

9.1

3 5.

8 4.

4-7.

3 2

7.5

7.3-

7.7

5 4-

6 4.

54.8

5

7.5

7.2-

8.0

14

6.8

5.9-

7.5

14

5.9

54-6

.4

1 3.

0 3.

0 1

5.0

5.0

7 5.

7 5.

1-6.

2 7

4.6

4G4.

8

4 6.

0 5.

3-6.

3 4

4.9

46-5

.2

7 4.

1 39

-4.3

7

6.0

5.8-

6.3

11

6.0

5.66

.5

12

5.0

4G5.

4

6 6.

3 5.

5-7.

1 6

5.0

4.1-

6.3

2 7.

7 7.

5-7.

9 1

3.5

3.5

1 5.

4 5.

4 11

5.

9 5.

5-6.

4 11

4.

8 4.

3-5.

3

28

8.0

6.6-

10.6

28

5.

7 4.

7-7.

1 12

13.

5 9.

8-16

.9

7 4.

3 4G

4.7

8 7.

0 6.

S7.6

14

6.

6 6.

0-7.

0 15

5.

6 5.

0-6.

3

1 Kay

et

al.

(198

1)

2 H

arri

son

(unp

ublis

hed

data

) 3

Har

riso

n (1

982)

w 5



a b

Figure 4. Comparison of the occlusal plans for the first lower molar in (a) Dionyropithecus sp. (H-GSP 8114/609) and (b) A4icropithecus clarki (KNM-CA 380) (illustration reversed). The black dot represents the mesial aspect of the molar. Scale represents 1 mm.

a dendromurine, none of which is known from presumed earlier sites. The oifferent Sayimys species from the two sites may be significant and might indicate that H-GSP 8114 is actually somewhat older than Y592. Given the 16.1 Ma date ofY592 (references in Barry et al., 1987), we conclude that H-GSP 8114 (and presumably S2) is at least 16 Ma, but certainly not as old as Y721 at 18 Ma. This is consistent with an initial comparison of the large mammal faunas from the Manchar localities and the CPG section fJ. Barry, pers. comm.), and with the results of a preliminary paleomagnetic survey by Khan et al. (1984) of the Manchar Formation in the Gaj River section.

Referring to Figure 2, micromammal sites H-GSP 8114A, 8114,8227, and 8224 all lie in the same measured section, and S2 is near or equivalent to 8114. H-GSP 8227 and 8224, like H-GSP 107 (Wessels et al., 1982)) h ave Antemus chinjiensis, a species restricted to about

14 to 12 Ma. H-GSP 107 and 8224 also have glirids, which first appear at about 13.5 Ma. These data provide consistent upper limits on the age of the Dionysopithecus sites.

We follow de Bruijn & Hussain (1984) who interpreted the base of the Manchar Fm. to be diachronous in the Gaj River and Sehwan sections, a consequence of the previously described hiatuses in the Sehwan section. Similarly, we propose the existence of a significant hiatus between Sehwan localities 8114A and 8114, which are separated by only 4 m of section. Their fauna1 differences suggest a greater time lapse than their stratigraphic proximity might otherwise indicate.

Description and comparison of the primate specimens

The specimens (Figures 3 and 4) are: a left maxillary canine germ (GSP S-76), a left P4 (H-GSP 8114/3690), and a right MI germ (H-GSP 8114/609). The three teeth are compatible in size (see Table 2) and morphology, and can be reasonably referred to a single species.

Figure 3. SEM photomicrographs ofDionysopi&cus sp. from the Manchar Formation, Sind, Pakistan: (a) H-GSP 609, right MI in occlusal (above) and buccal views; (b) GSP S-76, left Cl in lingual view; (c) H-GSP 3690, left P4 in occlusal (above) and distal views. Bar scales equal 1 mm.


H-GSP 8114/609 The specimen is an isolated, unerupted germ of a right Mi. The crown is complete and generally well preserved, although the enamel layer is not yet fully formed and small flakes of enamel have been lost from the tips of the distal cusps by postdepositional abrasion. The crown is quite broad and relatively short, being slightly narrower mesially than it is distally. The principal cusps are low, rounded and voluminous. The protoconid and metaconid are more or less transversely aligned, and are connected by a low, rounded and continuous crest. The mesial fovea is relatively well-defined, being slightly broader than it is long. The hypoconid and entoconid are both well developed and are similar in size to the metaconid. The hypoconulid is the smallest of the cusps and is situated on the distal margin of the crown in the midline of the tooth. The occlusal crests are short and rounded, but well developed. The crests linking the protoconid and hypoconid are interrupted by a line transverse groove which passes lingually from the talonid basin onto the cingular shelf. A short, fine crest links the hypoconid to the hypoconulid. Short crests descend from the apices of the hypoconulid and entoconid and meet midway between the two cusps to define a small, but distinct distal fovea. The talonid basin is quite shallow and is intersected by a system of fine grooves. The buccal cingulum is moderately well developed and forms a narrow, but more or less continuous shelf of enamel around the buccal margin of the tooth.

H- GSP 811413690 This isolated left P4 is only slightly worn but, unfortunately, poorly preserved. The enamel is pitted by weathering, and the base of the crown has been lost buccally and distally by abrasion. The crown is short and relatively broad, and ovoid in occlusal outline. The two cusps are well developed and transversely aligned. The buccal cusp is much more elevated than the lingual cusp with steep and sharply defined mesial and distal crests originating from its apex. The lingual cusp is more rounded with well developed mesial and distal ridges which sweep buccally to become confluent with the marginal ridges. The two cusps are connected by a sharp transverse crest. Mesially, the transverse crest delimits a short, narrow and well defined mesial fovea. The distal basin is much more extensive than the mesial fovea, but is shallow and less well defined. The lingual cingulum is narrow and rounded, and appears to be best developed around the distolingual margin of the lingual cusp.

GSP S-76 This specimen is a left maxillary canine germ preserving an intact crown and approximately 3 mm of the root. The crown enamel is weathered but the specimen appears to have been nearly fully formed. Its most striking feature is its small size (Table 2), perhaps due in part to its not yet being fully developed. S-76 is smaller than all early and middle Miocene small catarrhine canines from East Africa listed by Harrison (1982). There is a deep mesial groove that extends almost to the crown apex. Distal to the groove the lingual face is markedly convex, leading to a sharp distal ridge. There is very little development ofthe lingual cingulum. Lingually, the cervix makes a smooth transition with the root and is not inflated. These latter features may also be due in part to the developmental stage of the tooth.

Comparisons In addition to these specimens, Barry et al. (1987) described an isolated upper first molar (GSP 24307) from locality Y592, in the Kamlial Formation of northern Pakistan, dated


magnetostratigraphically at 16.1 Ma. It is compatible in both size and morphology with the Sindhi specimens, and can be considered to represent the same species.

The Sindhi Mi shows some distinct structural similarities to the corresponding Oligocene catarrhine molars from the Fayum of Egypt that have been assigned to Aegyptopithecus and Propliopithecus. However, there are several derived characters that distinguish the Sindhi specimen from the Fayum primates. The crown is slightly narrower, it has larger and more distinct mesial and distal foveae, a less inflated buccal cingulum and more sharply defined occlusal crests. It appears to be more similar in these respects to the contemporaneous small catarrhine primates from the early to middle Miocene of East Africa. It compares particularly closely to the early Miocene East African form Micropithecus clarki (Fleagle & Simons, 1978)) in having a relatively short and broad crown, with low, rounded occlusal crests, a discontinuous buccal cingulum and a relatively small, but distinct mesial fovea (see Figure 4). In fact, the minor differences between the Asian specimen and M. darki (i.e., in the former the crown is relatively slightly broader, the two mesial cusps are more transversely aligned, and the buccal cingulum is slightly better developed) are probably of insufficient importance, when individual variation is taken into account, to be considered taxonomically significant.

The Sindhi molar is quite distinct from those of the middle to late Miocene European pliopithecids. Pliopithecid lower molars can be distinguished by their combination of largely primitive features and striking apomorphies. They tend to be long and narrow, with high sharp crests, a long and obliquely directed cristid obliqua, a specialized triangle

pliopith&in, at least on M2 and MS, a relatively large mesial fovea, a small hypoconulid, and no true distal foveae.

This single specimen is sufficient to demonstrate that the Sindhi species is derived compared to the Oligocene catarrhines, and to the pliopithecids from the later Miocene of Europe. It has distinct affinities with the more advanced East African early Miocene catarrhines, particularly Micropithecus. The structures of the Sindhi upper premolar and canine, although less useful for determining relationships among early catarrhines, are consistent with this assessment. The canine is closest in size to small, presumably female canines of Limnopithecus legetet and L. evansi (see Harrison, 1982), and to UMP 66-19 from Napak, Uganda, assigned to Micropithecus clurki by Fleagle & Simons ( 1978). No isolated upper canines were assigned to M. clarki by Harrison, but the Sindhi specimen is comparable in size to the base of the crown preserved in the holotype, UMP 64-02. Morphologically, it is most similar to UMP 66-19 and to the larger, possibly male specimen from the same site, UMP 68-03, also assigned to Micropithecus clarki by Fleagle & Simons (1978). It differs from these only in the degree of cingulum development.

A small catarrhine maxillary fragment with MI-3 from the Xiacaowan Fm. at Songlinzhuang (Sihong District, Jiangsu Province), eastern China, also bears a marked similarity, at least in the structure of its molars, to Micropithecus clurki. However, on the basis of morphological, temporal and geographical evidence, Li (1978) assigned it to a new genus and species of primate, Dionysopithecus shuangouensis. The subsequent description of Micropithecus clarki made the distinction between D. shuangouensis and East African taxa less convincing. The upper molars of Dionysopithecus and Micropithecus share a number of distinctive morphological features (see also Etler, in press):

(1) the Ml and Ms are relatively quite narrow, with a convex lingual margin and rounded mesio-lingual and disto-lingual corners; (2) the cusps of the trigon are moderately high and


sharp; (3) the-metacone is at least as voluminous as the protocone on Ml and M*; (4) the occlusal ridges of the trigon are low, but well-defined; (5) the trigon forms an almost perfect equilateral triangle; (6) the mesial fovea is well-defined, but quite restricted; (7) the hypocone on Ml and M* is relatively small, lingually positioned, and is linked to the protocone by a short crest; (8) the lingual cingulum is narrow and forms a C-shaped ledge around the protocone; (9) the distal fovea and cingulum are relatively expansive; and (10) the posterior molar is morphologicaIly and metrically reduced. The minor morphological differences that distinguish the upper molars ofD. shuangouensis from those of‘M. clarki (i.e., the shape and proportions of the M2 and the development of the cingulum) are barely sufficient to warrant even a species distinction between them, and it may prove preferable, when more material is available, to include them in a single genus, Dionysopithecus.

However, synonymy would be premature at this time. Unfortunately, no lower molars attributed to D. shuangouensis have been described. A

number of additional specimens of this species have been recovered from the Xiacaowan Fm. at Songlinzhuang, but these remain unpublished (D. Etler, pers. comm.). Nevertheless, the isolated Mi from Sind is compatable in size and morphology with the holotype of D. shuangouensis. Fending description of the new Songlinzhuang material, and resolution of the question of synonymy of Micropithecus and Dionysopithecus, we provisionally attribute the new specimens from Pakistan to the Asian genus, as Dionysopithecus sp. This taxonomic assessment is supported by the approximate contemporaneity of the Xiacaowan and Manchar Fms. (see Barrv et al., 1987, and below).

Evolutionary, biogeographic and temporal relationships of early Eurasian catarrhines

In addition to D. shuangouensis, four other species of fossil catarrhines have been described from the Xiacaowan Formation. Gu & Lin (1983) re erred five isolated molars from a site f near Songlinzhuang to a new genus and species, Platodontopithecusjianghuaiensis, while Lei (1985) described three additional species from the same site, Hylobates tianganhuemis,

Pliopithecus mongi and Dryopithecus sihongensis, based on single isolated molars. The isolated molars of P.jianghuaiensis are rather worn and poorly preserved, but they do

provide substantive evidence that a second, slightly larger species is represented in the Xiacaowan Fm. The upper molars are broader than those of Dionysopithecus, with a well-developed, shelf-like cingulum bordering the entire lingual aspect of the crown and a prominent cingulum on the buccal margin. The lower molars also have a well-developed buccal cingulum, and the cusps are low and rounded with poorly developed occlusal crests. In general, like Dionysopithecus, Platodontopithecus bears a stronger resemblance to the East African early Miocene taxa than to any of the later Eurasian taxa, such as the Pliopithecidae or Dryopithecus. Etler (in press) finds the strongest resemblance to be with the larger East African genus Proconsul.

The type specimen of H. tianganhuensis, a left Ml (P. 83.3), is slightly larger than the corresponding tooth in the holotype of Dionysopithecus shuangouemis, but judging from the photograph published by Lei (1985), the specimen is morphologically very similar. In fact, it has all of the characteristic features of the upper molars referred to Dionysopithecus and Micropithecus. The minor morphological differences noted by Lei (1985) to distinguish H.

tianganhuensis from D. shuangouensis, and the slight size difference, are in our view rather trivial and do not provide sufficient grounds to recognize a taxonomic separation. We therefore consider H. tianganhuensis to be a junior synonym of D. shuangouensis.


Pliopithecus won@ (Lei, 1985) is represented,by an isolated left lower molar (P. 83.4). The distal narrowing of the crown, distribution of the cusps, and disruption of the distal fovea by a subsidiary crest running from the entoconid, suggest that the specimen is a third molar. The crown is short and elliptical, with low rounded cusps, small but distinct occlusal basins, and a narrow buccal cingulum. Although the tooth cannot be compared directly with D. shuangouensis, as no lower molars are described for the latter, it is an appropriate match for the holotype and compares favorably with some of the isolated third molars of M. clarki. Etler (in press), however, likens it to East African Dendropithecus. It is generally similar to the Sindhi Ml, differing mostly in features that distinguish third from first molars in M. clarki. Although there is no way to be certain of the affinities of P. 83.4 because of the lack of associated material, all the available evidence indicates that P. zoongi may also be a junior synonym of D. shuangouensis.

The third specimen described by Lei (1985), as the holotype of Dryopithecus sihongensis

(P. 83.5), appears to be an Ml rather than an Ma as suggested by Lei. Given its size, it would therefore belong to a relatively large species about the size of Proconsul africanus. As such, it may represent the earliest Eurasian record of a large catarrhine. The crown is quite long and narrow with low, voluminous cusps separated by deeply incised fissures. The major grooves coqform to the basic Dryopithecus Y-5 pattern. In its general morphology, it is most similar to the lower molars ofSimpithecus and Dryopithecus. However, the specimen has a well-developed buccal cingulum. The cingulum appears to be more pronounced thanin later large hominoids, including the middle Miocene species of cf. Siuupithecus from the Vienna Basin and Pasalar (see Kelley & Pilbeam, 1986, regarding the taxonomy of these samples) and Kenyapithecus africanus from Maboko, all of which retain a distinct cingulum (Andrews, 1978a; Andrews & Tobien, 1977; Pickford, 1985aJ). In this respect D. sihongensis is similar to the more conservative genera, Proconsul and Afropithecus, from the early Miocene of East Africa (Andrews, 19783; Leakey & Leakey, 1986). Without further material of D. sihongensis it is difficult to determine its affinity, although the attribution to Dryopithecus can be questioned. As with Platodontopithecus, Etler (in press) sees it as strikingly Proconsul-like.

The age of the Xiacaowan fauna has been estimated indirectly as late early Miocene. Li et al. (1983) correlated it to European mammalian biochrons MN 5 or 6, but this has been revised in two recent more detailed studies. From examination of the Xiacaowan genera Semigenettu and Pseudaelurus, Qiu & Gu (1986) estimated it to be equivalent to MN 4 or 5, while Qiu & Lin (1986) suggested correlation to MN 3 or 4, based on the Sciuridae. The Xiacaowan fauna is older than the Shanwang fauna (Li et al., 1983; Li, pers. comm.; contra Mein & Ginsburg, 1985), which is considered to be MN 4 or 5 (Qiu et al., 1986) and is bracketed by radiometric dates of 18 and 14 Ma (Jin, 1985; see also Etler, in press). Thus, the precise age of the Xiacaowan Dionysopithecus is uncertain, but it may not be any older than earliest middle Miocene, or cu 16 Ma, the age of the oldest reliably (geochronologically) dated Eurasian catarrhine (Barry et al., 1987).

An edentulous mandibular symphysis from Hsi Shui, Taben Buluk, Gansu Province, northern China, provides further evidence of a small catarrhine primate from the early to middle Miocene of Asia. Bohlin ( 1946) ori mally referred the specimen to a new genus, ‘g’ “Kansupithecus”, but, as he failed to provide a species name for the taxon, this does not constitute a valid nomen (see Szalay & Delson, 1979). A second specimen, consisting of a fragment of a lower molar from the neighboring, late Oligocene locality ofyindirte, Taben Buluk (Thomas, 1985), was also referred by Bohlin (1946) to “Kansupithecus”. Casts of

3.50 R. L. BERNOR ET AL.

these specimens indicate that the mandibular fragment is of the appropriate size and morphology to represent a further specimen of Dionysopithecus shuangouensis. The molar fragment, however, is much too incomplete to determine its affinities, but it is likely not even a primate.

Bohlin (1946) considered the Hsi Shui assemblage to be certainly younger than Yindirte, and therefore probably Miocene. The rodent Sayimys obliquidens, although from different sites, was used as evidence of Miocene age deposits in the vicinity. The Hsi Shui locality was placed in the early middle Miocene by Conroy & Bown (1974). Similarly, Li et al.

(1984) correlated it with the Astaracian or latest Orleanean of Europe. Thomas (1985), however, followed Russell & Zhai (1987) who assign it to the early Miocene without discussion. Until material from Taben Buluk is better studied, and until more primate remains are found, the Hsi Shui mandible should be considered indeterminate and not necessarily older than the Xiacaowan and South Asian Dionysopithecus records.

The late early Miocene also witnessed the immigration ofpliopithecids into Europe. The pliopithecids are more conservative than Asian and African early Miocene catarrhines, but more derived than the Fayum primates (Remane, 1965; Groves, 1972, 1974; Delson & Andrews, 1975; Delson, 1977; Ciochon & Corruccini, 1977; Ginsburg & Mein, 1980; Harrison, 1982, 1987; Fleagle, 1984, 1986; Andrews, 1985). This suggests that the pliopithecids were potentially a distinct lineage by the end of the Oligocene and presumably originated in Africa. However, their evolutionary history is unknown until they first appear in Europe during the terminal Burdigalian regression at MN 5 (Ginsburg & Mein, 1980; Bernor, 1983; Ginsburg, 1986; Steininger, 1986). Pliopithecids are represented by at least seven species assignable to five genera or subgenera (Hiirzeler, 1954; Zapfe, 1960, 1961; Bergounioux & Crouzel, 1965; Kretzoi, 1975; Ginsburg & Mein, 1980).

Pliopithecids have also been reported from the Miocene of Asia, although most of these records are based on inadequate material (e.g., Schlosser, 1924; Chopra & Kaul, 1979). Recently, Qiu & Guan ( 1986) referred a left M2 from the Miocene locality of Maerzuizigou, Tongxin, China, to Pliopithecus sp. This specimen appears to resemble European pliopithecids in its proportions, the development of the occlusal crests and buccal cingulum, and in the claimed presence of a triangle pliopithe’cin. The fauna from this site has been considered to be slightly older than the Tung-gur fauna, which appears to be late middle Miocene in age (ca 13-11 Ma; Li et al., 1984; Qiu & Guan, 1986). As noted by Qiu & Guan, the specimen may provide the earliest record of the Pliopithecidae in China. The middle Miocene Paratethys extended continuously from Switzerland to east of the present day Aral Sea, and could have provided a warm, equable environmental corridor for pliopithecids into Asia, north of the rising Himalayan erogenic belt. This corridor would have become progressively diminished during the Sarmatian and Pannonian stages of the Paratethys time scale (ca 13-9 Ma), as marine conditions were replaced by brackish then fresh water conditions, and the Central and Eastern Paratethys became disconnected (R6gl & Steininger, 1983; Steininger, 1986).

The late Miocene site of Shihuiba, Lufeng County, Yunnan Province, southern China, has yielded a large collection of specimens, including a nearly complete skull (Wu & Pan, 1985; Pan, 1988), which were named Laccopithecus robustus Wu & Pan, 1984. The material has not yet been described in great detail, but it seems clear from the dental remains that Laccopithecus is closely related to European Pliopithecidae and should be included in the same family. Possibly, then, true Asian pliopithecids appear relatively late in Asia, being


first represented with certainty at Lufeng, ca 8 Ma or younger (Flynn & Qi, 1982), which also would be the latest known occurrence of the family.

The large, more derived catarrhine primates, cf. Sivapithecus and Dryopithecus, first appear in Eurasia slightly later than the Pliopithecidae. Both groups are presumably of African derivation, but only the thick-enameled cf. Sivapithecus has obvious precursors in the East African middle Miocene. Cf. Sivapithecus is first found in the Vienna Basin by MN 8, ca 13.5 Ma, at the site of Neudorf-Sandberg, and at the middle Miocene site of Pasalar in Turkey. Both sites have traditionally been considered to be very early middle Miocene in age, ca 16-15 Ma (Mein, 1979; Andrews & Tobien, 1977), but Ginsburg & Mein (1980) and Mein (1985, 1986) reassign the Sandberg locality from MN 6 to MN 8 based on a reassessment of certain fauna1 elements. A similar adjustment might be required for Pasalar based on preliminary descriptions of its fauna (Andrews & Tobien, 1977), although Mein (1986) considers it to be definitely older than Sandberg. DrTopithecus is found earliest at La Grive and St. Gaudens in France, and Sant Quirze in Spain, which are also correlated to MN 8, approximately contemporaneous with Neudorf-Sandberg (Mein, 1986).

Neogene eustatic sea-level changes and catarrhine migrations

Early in the Neogene a series of fauna1 migrations between Afro-Arabia and Eurasia commenced. The timing of these episodes of fauna1 migration depended to a large extent on regional and global tectonic and eustatic events. The collision of the Afro-Arabian Plate with Eurasia during the late Oligocene/early Miocene established the potential for a subaerial migratory corridor across the Ethiopian-Yemen Isthmus, through Arabia and into southwestern Asia, probably prior to 20 Ma (Adams et al., 1983; Barry et al., 1985; Bernor, 1983; Raza et al., 1984; Whybrow, 1984; Steininger et al., 1985; Thomas, 1985). Once established, exposure of this corridor was dependent upon both local uplift and regional and global sea level changes. Haq et al. (1987) have recently presented a calibration of global eustatic events. They identify a number of early and middle Miocene short-term sea lowering episodes, which are times of increased likelihood of subaerial connections between Afro-Arabia and Eurasia. These occurred at the following times (Figure 5): a gradual lowering episode beginning at 23 Ma; a major sea level lowstand at 2 1 Ma, followed by gradually rising sea level over the next few million years; a minor sea level drop at 17.5 Ma, followed by successive major drops and sea level lowstands at 16.5 and 15.5 Ma; major lowering episodes at 13.5 Ma and 12.5 Ma, with finally a drastic sea-level lowering at 10 Ma that produced sea levels as low as at any time in the Tertiary.

On a regional scale in the Miocene, Adams et al. (1983) and Whybrow (1984) recognized growing provinciality in the planktonic biota of the Mediterranean and Arabian seas, and apparent interruption in the continuity of marine sedimentation across the Middle East. They argued that the earliest Neogene interval during which a Saudi Arabia-Iran/Iraq subaerial corridor would have been emergent was during the local equivalent of the Aquitanian. This correlates well with the progressive sea level lowering beginning at 23 Ma, culminating with the major lowstand at 21 Ma. The Bugti megafauna, at 20+ Ma, represents the earliest Asian evidence of this connection with the presence of archaic African elements. There is no evidence that primates were involved in this initial interchange between Afro-Arabia and Eurasia, despite the suggestions of Thomas (1985) regarding pliopithecids.

This corridor did eventually allow primates to cross into southwest Asia later in the

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Miocene. Subsequent dispersal into China occurred either across the circum-Indian Ocean arc of southwestern Asia through Indo-Pakistan, or via the Paratethys corridor through central Asia. African primates also entered Europe through Arabia, then dispersed either across the Sub-Alpine Arch of Anatolia and southern Europe (Antunes, 1979; Bernor, 1983), or across the Pannonian and Vienna Basins of eastern Europe (Steininger et al., 1985). On present evidence, both events appear to have occurred ca 17 to 16 Ma, suggesting association with the major sea level lowstand at 16.5 Ma, or possibly with the minor sea level lowering event at 17.5 Ma. Dispersal of catarrhines into Asia was very likely associated with a second episode of African mammal immigration into south Asia sometime between 20 and 16 Ma, when the middle to late Miocene Siwalik chronofauna acquired its characteristic composition (Barry et al., 1985). Important large mammal elements of this immigration were bovids, tragulids and perhaps large giraffoids (Figure 5). From the still somewhat archaic character of ca 18 Ma large and, to a lesser extent, small mammal assemblages from the Potwar Plateau (Barry et al., 1985; J. Barry, pers. comm.) catarrhines and the associated fauna may have entered Asia only toward the end of this interval, and not long before the 16.1 Ma date that marks their current earliest reliably documented occurrence (Barry et al., 1987). More recent collecting in the Potwar suggests that the immigrant mammalian assemblage with which Dionysopithecus is associated seems unlikely to have arrived earlier than about 17 Ma 0. Barry, pers. comm.). The most likely time of immigration therefore seems to be the major sea lowering event at 16.5 Ma.

We must emphasize that the approximate synchroneity of catarrhine immigration into Asia and Europe that now seems apparent is still poorly documented and may be unreal for several reasons. Whereas the South Asian chronology is tied to the established magnetostratigraphic framework of the Potwar Plateau, those of Europe and East Asia depend largely on fauna1 biochronology tied irregularly to interdigitated marine sequences, so that time correlation is both uncertain and imprecise. Despite doubts surrounding the time correlation of European Mammal Ages, the issue of the earliest appearance of European pliopithecids, in biochronological terms, can be addressed more confidently than can the earliest appearance of catarrhines in Asia. Early Miocene faunas of Europe are relatively well known (Mein, 1979) and the likelihood of finding European catarrhines older than MN 5 is low. In contrast, early Miocene faunas of South and East Asia are both sparse and still unreliably dated or correlated. The Xiacaowan fauna could be older than we have suggested, and the possibility of recovering still older catarrhines in East Asia cannot be ruled out. If primates and other immigrants reached China via a Paratethyan route, then the apparent time constraints on immigration by Siwalik faunas could be irrelevant.

The next eustatic sea lowering event, ca 13.5 Ma (Haq et al., 1987), corresponds closely with the apparent first occurrence of Dryopithecus (s.s.) in West-Central Europe and perhaps with the appearance of the thick-enameled hominoids in eastern Europe and Turkey (Figure 5). This correlation is more reliable for Europe than for Turkey, however, because of the uncertainty about the age of Pasalar. The initial immigration of thick-enameled hominoids from Africa to southwest Asia may have been associated with one of the earlier sea lowering episodes. Thick-enameled hominoids first occur in Indo-Pakistan about 12 Ma as part of a minor Siwalik fauna1 immigration episode (Barry et

al., 1985; Barry, 1986). This corresponds with yet another episode of lowered sea level, and a sea level lowstand equal to those of the early middle Miocene (Figure 5).

354 R. L. BERNOR ET AL..

We must be cautious with these interpretations. While global eustatic events certainly influenced catarrhine and other mammal distribution, other factors such as local tectonic structures and ecology also bear on migratory events. Thus, although interprovincial migrations would seem to have been increasingly unlikely between about 2 1 and 17.5 Ma because of gradually rising sea levels accordingly to the Haq et al. (1987) curve, it cannot be assumed that impenetrable marine barriers existed in the tectonically complex region between Afro-Arabia and South Asia during this entire interval. Based on studies of regional stratigraphy and biostratigraphy, Adams et al. (1983) and Whybrow (1984) both argue for an uninterrupted terrestrial connection between Afro-Arabia and southwest Asia after the mid-Burdigalian, cu 19 Ma.

The earliest Neogene deployment of African primates into Eurasia cannot be constrained by simple dispersal hypotheses. Global eustatic sea level lowering, tectonic rearrangements, and shifting paleogeographic features all played crucial roles in these episodes of deployment. As the record of Neogene faunas of the Indian subcontinent matures, it becomes clear that there were numerous periods of interchange with African and East Asian faunas. What had been recognized as one or two early and middle Miocene “migratory pulses” are now best considered as several short intervals of fauna1 exchange.

Summary

Three isolated teeth of a small catarrhine are described from the lower part of the Manchar Formation of southern Pakistan. The specimens, although collected by two field parties, come from the same section, probably the same horizon, and represent a single species. They are compatible in size and morphology with the catarrhine molar from Potwar Plateau locality Y 592, Kamlial Formation, northern Pakistan, described recently by Barry et al. (1987).

The fauna1 assemblages associated with the Y 592 and Manchar specimens show a high degree ofsimilarity (Table l), in contrast to other assemblages in their respective sections. Based mainly on micromammals, the Manchar sites correlate well with Y 592, paleomagnetically dated at 16.1 Ma, although they could be somewhat older. Thus the small catarrhine teeth from both areas are approximately contemporaneous and, although separated by 800 km, may represent the same species.

This small catarrhine is most similar to Dionysopithecus shuangouensis from the Xiacaowan Formation of Jiangsu Province, China. The age of this fauna is not certainly known at present, but could be about the same. It also resembles Micropithecus clurki from the early Miocene of East Africa in several derived features. Analysis of dental traits supports the opinion that these taxa may be accommodated by a single genus Dionysopithecus, although we do not yet formally synonymize the two genera. Unfortunately, there are no homologous teeth for direct comparison of the D. shuungouensis holotype with the Pakistani specimens, and therefore the latter are designated Dionysopithecus sp. until more material is found.

Other catarrhine records from the Xiacaowan Formation include “Hylobutes

tiungunhuensis” and “Pliopithecus wongi”, both probably junior synonyms of Dionysopithecus

shuungouensis. Plutodontopithecus jiunghuuiensis and “Dryopithecus” sihongensis resemble East African early Miocene taxa such as Proconsul and Afropithecus (see also Etler, in press). Thus, the Xiacaowan Formation contains not only a small catarrhine like that of Pakistan, but perhaps also the earliest Eurasian large catarrhines. All three lineages may have


entered eastern Asia from Africa at about the same time, although diversification in Asia from a single immigrant stock cannot be ruled out at present. In contrast, large catarrhines (hominoids) appeared in the Indian subcontinent well after Dionysopithecus, suggesting that if small and large catarrhines entered Asia at the same time, they may have done so by different routes.

The small catarrhine (“Kansupithecus”) from Gansu Province, northern China, is indeterminate and may be no older than the records of Dionyso$thecus.

Pliopithecidae are another primate group that exited Africa in the late early Miocene. They appear in Europe during the terminal Burdigalian regression at MN 5, cu 17 to 15 Ma. The group may be represented at Maerzuizigou, Tongxin, China (Qiu & Guan, 1986), suggesting spread through Asia via the Paratethyan corridor, north of the Himalayan erogenic belt. If this fossil is not a pliopithecid, then it may be that the group did not enter China until the late Miocene, represented by the Lufeng Laccopithecus (but see also Pan, 1988).

Large catarrhines appear in Europe and the Middle East later than the Pliopithecidae. Cf. Sivapithecus occurs at Neudorf-Sandberg in the Vienna Basin by MN 8 (see, for example, Mein, 1986) and at Pasalar, Turkey, at about the same time. Dryopithecus is a late Astaracian (MN 8) European fauna1 element.

Barry et al. ( 1985) h s ow that Siwalik fauna1 history reflects a series of immigrations into south Asia, which are tied strongly to global tectonic and eustatic events. Haq et nE. (1987) have further calibrated global eustatic events, and the spread of primates and other fauna1 elements into southern Asia and Europe correlates to these fluctuations (Figure 5). An early Miocene sea level lowstand corresponds to a phase of emigration from Africa that did not include primates. New African elements appear in south Asia 17 to 16 Ma, including Dionysopithecus. Pliopithecidae appear in Europe at about the same time. East Asian earliest records of small and (perhaps) large catarrhines are about the same age, but the spotty record there does not rule out a still undiscovered earlier appearance of catarrhines in East Asia. Large hominoids enter Europe in a later dispersal, which predates their first occurrence with other African elements in the Indian subcontinent.

Acknowledgements

We thank S. M. Raza, J. C. Barry, D. R. Pilbeam, L. L. Jacobs, J. G. Fleagle, and F. Rijgl for helpful discussions and encouragement during the course of this study. Li C.-K., W. R. Downs, D. Etler, and E. Delson supplied valuable information on Asian localities and Chinese texts. J. Barry compared Manchars and Kamlial/Murree large mammal assemblages. Suggestions by Li, L. Martin, and two anonymous reviewers improved the manuscript. We are grateful to the Geological Survey of Pakistan for support of this project, and thank the Kenya Government and the National Museums of Kenya for permission to study the East African Miocene primates. Support for this study was derived from NSF grants BSR 85-17396 (to Bernor), BNS 84-19703 (to Pilbeam, Barry and Hill), and BSR 85-00145 (to Jacobs, Flynn and Lindsay); Smithsonian Foreign Currency Program Grants to Hussain (41007800) and Pilbeam and Barry (20203700); National Geographic Grant (no. 3494-87, to Hussain); and NATO grant RG 85/0045 (to Bernor). We thank Dr M. Yoder, New York University, and Trisha Rice, Museum of Comparative Zoology, for skilled creation of the SEM photos. This paper is dedicated to Noye Johnson, who so consistently promoted and advanced work in the Siwaliks.


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