evolution in India 729

India at the cross-roads of

R PATNAIKa,* and P CHAUHANb aCentre of Advanced Studies in Geology, Panjab University, Chandigarh 160 014, India bThe Institute and CRAFT Research Center (Indiana University), 1392 W Dittemore Road, Gosport, IN 47433, USA *Corresponding author (Email, [email protected])

The Indian palaeoanthropological record, although patchy at the moment, is improving rapidly with every new fi nd. This broad review attempts to provide an account of (a) the Late and their gradual disappearance due to ecological shift from forest dominated to grassland dominated ecosystem around 9–8 Ma ago, (b) the immigration/evolution of possible hominids and associated fauna, (c) the record of fossil hominins, associated fauna and artifacts, and (d) the Holocene time of permanent settlements and the genetic data from various human cultural groups within India. Around 13 Ma ago (late Middle Miocene) Siwalik forests saw the emergence of an -like . By 8 Ma, this disappeared from the Siwalik region as its habitat started shrinking due to increased aridity infl uenced by global cooling and intensifi cation. A contemporary and a close relative of Sivapithecus, (Indopithecus), the largest that ever-lived, made its fi rst appearance at around 9 Ma. Other smaller that were pene-contemporaneous with these apes were Pliopithecus (Dendropithecus), Indraloris, Sivaladapis and Palaeotupia. The Late Pliocene and Early Pleistocene witnessed northern hemisphere glaciations, followed by the spread of arid conditions on a global scale, setting the stage for hominids to explore “Savanahastan”. With the prominent expansion of grassland environments from East to China and Indonesia in the Pliocene, monkeys and baboons dispersed into the Indian subcontinent from Africa along with other . Though debated, there are several claims of the presence of early hominins in this part of the world during the Late Pliocene, based primarily on the recovery of Palaeolithic tools. of our own ancestor and one of the fi rst globe-trotters, early erectus, has been documented from the Early Pleistocene of East Africa, Western Asia and Southeast Asia, thus indirectly pointing towards Indian subcontinent as a possible migration corridor between these regions. The only defi nite pre- Homo sapiens fossil hominin remains come from the Central Narmada Valley and are thought to be of Middle to age, and the cranium has been shown to be closely linked to archaic Homo sapiens/H. heidelbergensis of Europe. Around ~74,000 yrs ago, a super volcanic eruption in Sumatra caused the deposition of Youngest Toba Tephra, that covered large parts of the Indian peninsula. Just around this time anatomically-and-behaviorally modern or Homo sapiens possibly arrived into India as evidenced by the so called Middle and Upper Palaeolithic assemblages and associated symbolic evidence. The available genetic data reveals that the gene pool to which modern Indians races belong was extremely diverse and had variable mixed links with both European and Asian populations.

[Patnaik R and Chauhan P 2009 India at the cross-roads of human evolution; J. Biosci. 5 729–747] DOI 10.1007/s12038-009-0056-9

1. Introduction the early primates, the order to which, we humans belong. In fact, recent fi nds of anthropoid primates from the Early According to the “Out of India” hypothesis, several Eocene lignite mines situated in Eastern Gujarat, India groups of modern Asian organisms had their roots in the (Bajpai et al. 2008) and Oligocene of Pakistan (Marivuax northwardly moving Indian plate (Bossuyt and Milinkovich et al. 2005), may hold clues to the origin and dispersal of 2001; Karanth 2006). One of these groups could have been our earliest ancestors. These recent fi nds have lent support

Keywords. Genetics; human evolution; Indian subcontinent; palaeoclimate; phylogeny; primate http://www.ias.ac.in/jbiosci J. Biosci. 34(5), November 2009, 729–747, © IndianJ. Biosci. Academy 34(5), of November Sciences 2009 729 730 R Patnaik and P Chauhan to some of the long-standing assertions that Asia was the and Ramapithecines were the African Dryopithecines, centre of anthropoid origins (Ciochon 1985; Culotta 1995; which may have arrived in southern Asia during the Early Beard et al. 1996). Miocene (Begun et al. 2003). Our aim here is to broadly review the palaeoanthropo- The Middle Miocene was the time of warm and humid logical records of India (with a few examples from Pakistan) climate and evergreen to deciduous tropical forests covered from the Miocene onwards, which essentially begins with a large part of the northwestern Indian subcontinent. These the emergence and disappearance of fossil apes, followed by ecological conditions are very well refl ected in the fossil the possible presence of early hominins in the late Pliocene as well as sediment record (Ashton and Gunatilleke 1987; and Early Pleistocene in the context of changing climate Nanda and Sehgal 1993; Prasad 1993; Thomas et al. 2002). and ecology. We also briefl y touch upon the Holocene Sivapithecus fossils have been found from the Potwar by reviewing data related to the gene fl ow of modern Plateau of Pakistan in the west and from in the east human races into and out of India. Finally, we conclude by from 13.5-8.4 Ma (Barry et al. 2002; Nelson 2003; Patnaik evaluating the lacunae in the fossil and technological record, et al. 2005). There were at least three of Sivapithecus and suggesting possible future directions of research to fi ll that occurred in the Siwaliks, namely Sivapithecus indicus, S. these gaps in our knowledge of human evolution in India sivalensis and S. parvada. Sivapithecus indicus ranged from (e.g. Chauhan 2006; Petraglia and Allchin 2007). 12.5 to 10.3 Ma and S. sivalensis ranged between 9.8 to 8.4 Ma. S. parvada comes from a site dated to 10 Ma (Berggren 2. The Miocene et al. 1985; Kappelman et al. 1991; Flynn et al. 1995; Barry et al. 2002). S. sivalensis shows , with Though short-lived, India had its share of fame associated males averaging around 45 kg whereas females weighed with Ramapithecus, the Miocene ape from the Siwaliks. around 20 kg. S. parvada males were as large as modern Lewis (1934) fi rst recognized and named Ramapithecus orangutan males (Kelley 1986). Low crowned, thick brevirostris, the Rama’s ‘short-faced ape’ and placed it in the enameled molars of Sivapithecus indicate a tough diet, , the family to which all bipedal such as nuts and fruits with tough rinds (Kay 1981). Dental and we belong. At the time of its discovery and soon after, microwear studies of their occlusal surfaces have been Ramapithecus was widely known to be the ancestor of found to indicate a diet similar to those of modern fruit- , the “ape of the South”, which in turn gave eating apes, with some hard-object feeding (Nelson 2003). rise to our own genus Homo. This idea of Ramapithecus being Kelley and Pilbeam (1986) have shown that the cranio-facial a hominid gained general acceptance (Simons and Pilbeam morphology of Sivapithecus is heavily buttressed, pointing 1972) and researchers started assigning Miocene specimens towards an adaptation to either withstand prolonged from Kenya (Leakey 1962; Andrews and Walker 1979); cyclical loading or to generate high occlusal loads. As far as China (Woo and Wu 1984), Nepal (West 1984); Turkey locomotion is concerned, Sivapithecus postcranials indicate (Andrews 1982); Hungary (Kretzoi 1975; Fleagle 1988) and that it was more like a pronograde quadruped that walked Greece (Pilbeam et al. 1977) to the Ramapithecines. In fact, above the branches very similar to most of other Miocene von Koenigswald (1976) went further, by proposing that hominoids, but might have also been an active climber (Rose Ramapithecus should be the right candidate to be at the root 1986). Sivapithecus does not show any specializations for of hominid evolution that gave rise to Australopithecus as extensive terrestriality (Kelley and Pilbeam 1986). Kelley Ramapithecus bearing Siwaliks are located geographically (1997), based on enamel growth lines of a Sivapithecus between fossil hominid yielding Africa and Southeast Asia. parvada juvenile, found that the life history pattern with However, with the discovery of key facial and postcranial a prolonged growth and maturation period of Sivapithecus remains of Sivapithecus in the early nineteen eighties, was similar to that of modern great apes. Morphology of Ramapithecus lost its place in Hominidae (Andrews 1982; Sivapithecus and its life history pattern, suggest that this Andrews and Cronin 1982; Pilbeam 1983). The advent of large-bodied frugivore must have been vulnerable to periods analysing molecular clocks, which indicated ~5 Ma as the of ripe fruit shortages and may have relied heavily upon time of divergence between humans and apes, ultimately different fallback foods, such as hard seeds and nuts (Nelson made this contention more convincing (Sarich and Wilson 2005). 1967). Nevertheless, Ramapithecus still appears to hold an With the advent of Late Miocene global cooling and spread important place in primate evolution, as it belongs to the of arid conditions, these forests began to shrink (Kennett and pool of Late Miocene hominoids one of which gave rise to Hodell 1986; Scott et al. 1999). In Nepal it has been found later hominids (Ciochon and Fleagle 1985). This change in (Hoorn et al. 2000) that from the late Middle Miocene to the phylogenetic position of Ramapithecus (fi gure 1) has early Late Miocene (~11.5–8 Ma), the Himalayan foothills been well illustrated by Kennedy (2003: 119). Possible and the Gangetic fl oodplain were forested with subtropical to ancestors to these Middle to Late Miocene Sivapithecines temperate broad-leafed and tropical forest taxa, which were

J. Biosci. 34(5), November 2009 Human evolution in India 731

Figure 1. Phylogenetic placement of Ramapithecus, (A) in the 1970’s and (B) today (modifi ed after Kennedy 2003). then replaced by grasslands between the early to late Late (mainly bushes and trees) prior to 8 Ma, but by 7 Ma the C4 Miocene (~8–6.5 Ma). Intensifi cation of the Asian monsoon grasses dominated the Siwalik fl oodplain biomass (Quade et and disturbance of the vegetation on Himalayan slopes due al. 1989, 1995; Cerling et al. 1997). Palaeosol deposits from to their uplift might have facilitated such a change (Hoorn various Siwalik sequences also indicate marked seasonality et al. 2000). Around this time, a change in the sedimentary in rainfall (Retallack 1991, 1995). Recently, Barry et al. regime also takes place. The large emergent river system of (2002) observed three very brief periods of high Siwalik Nagri Formation gave way to an inter-fan river system of faunal turnover at 10.3, 7.8, and 7.37–7.04 Ma. Latest the Dhok Pathan Formation at around 10.1 Ma (Barry et Miocene faunal turnover at 7.37 and 7.04 Ma, particularly, al. 2002). There are other independent observations that has been correlated with expansion of C4 grasses, the strengthen this view further. In numerous deep-sea cores, oxygen isotope and sedimentological evidence indicating it was observed (Prell et al. 1992) that an upwelling of an increasingly drier and more seasonal climate (Barry et al. endemic foraminifers and radiolarians fauna took place at 2002). A dramatic change in the diversity of muroid ~8 Ma. As the Tibeto-Himalayan zone started rising around (from cricetid-dominated to murid-dominated) at ~9–8 Ma 12–9 Ma ago (Amano and Taira 1992; Harrison et al. 1993), has also been attributed to an intensifi cation of the the heat budget of the region probably changed drastically (Patnaik 2003) (fi gure 2). leading to the intensifi cation of the monsoon climate system A combination of Late Miocene change in climate, in South Asia (Ruddiman and Kutzbach 1989; Raymo and tectonics and sedimentary environments may have Ruddiman 1992; Kutzbach et al. 1993; Hay 1996; Ramstein resulted in the shrinking of forests that were occupied et al. 1997). A rather rapid uplift began at ~10 Ma and ended by Sivapithecus, leading to their eventual disappearance at ~5 Ma (Prell and Kutzbach 1992). The diatom record from the Siwaliks. Around this time (~9 Ma) the largest of the Indian Ocean also points towards an intensifi cation ape that ever lived on this planet and a close relative of of regional monsoons between 11 and 7 Ma (Schrader Sivapithecus appears: Gigantopithecus. It co-existed with 1974; Burkle 1989). In the terrestrial Siwalik deposits, soil Sivapithecus in the Late Miocene and and carbonate isotopic studies indicate a C3 dominant vegetation Pongo pygmaeus in the Pleistocene (Ciochon et al. 1996).

J. Biosci. 34(5), November 2009 732 R Patnaik and P Chauhan

Figure 2. (A) Time series of carbon (crosses) and oxygen (circles) isotopes in soil carbonates in Pakistan as an index of a shift in regional climate and to C4 vegetation which is interpreted to indicate a more arid, highly seasonal environment, after Quade et al. (1989) (redrawn from Prell and Kutzbach 1992, fi gure 4d). (B) Time series of endemic upwelling species of plankton: % foraminifera, Globigerina bulloides (dotted line), and qualitative abundance of radiolarian Actinomma spp. As indices of upwelling induced by monsoon winds (redrawn from Prell and Kutzbach 1992, fi gure 4c). (C) Percentage of grassland (redrawn from Hoorn et al. 2000, fi gure 4). (D) Late Miocene to present Phytogeography based on palaeosol data (redrawin from Retallack 1991, fi gure 6.12). (E) Muroid diversity (data from Patnaik 2003). (F) Age ranges of Siwalik apes (Barry et al. 2002; Patnaik 2008). (Modifi ed after Patnaik and Cameron 2007.)

For example, this genus goes on to thrive until the Late key locality for Miocene apes in India is Haritalyanagar Pleistocene (100,000 yrs BP) in China and Vietnam but not (fi gure 3) which is still yielding fossil ape specimens in the Indian subcontinent. Gigantopithecus bilaspurensis/ beside other faunal elements (fi gure 4). The other important giganteus from the late Miocene Siwaliks (Simons and hominoid locality is the Lower Siwalik Ramnagar locality Chopra 1969; Simons and Ettel 1970; Simons and Pilbeam exposed near Jammu and has yielded several fossil ape 1972) has recently been reassigned to Indopithecus (von specimens (see Nanda and Sehgal 1993; Sehgal and Nanda Koenigswald 1950) based on the distinct differences in 2002). mandibular and dental morphologies between the Indian and Chinese Gigantopithecus specimens (Cameron 2001, 2003). However, Miller et al. (2008) argue against assigning the 3. The Pliocene Miocene form to a separate genus. Ciochon et al. (1990) found opal on the teeth Discovery of tchadensis from (Brunet enamel of Gigantopithecus, which indicates a varied diet et al. 1995) may indicate that by the late Miocene, early comprising of , grasses and fruits. One of the hominids had already occupied areas beyond the East Gigantopithecus (Indopithecus) shows a large caries, African Rift valley. Dennell and Roebroeks (2005) have which also suggests a diet of grasses and fruits as compared argued that “If Australopithecus bahrelghazali, 2,500 km to a predominantly frugivorous diet of Sivapithecus (Patnaik west of the Rift Valley, implies that by 3.5 Myr ago hominids 2008). Their inferred giant size (based only on fossil were distributed throughout the woodland and savannah belt ) ranging from 300-150 kg and diet suggest from the Atlantic Ocean across the Sahel through eastern terrestrial locomotion. Accurately reconstructing the actual Africa to the Cape of Good Hope” (Brunet et al. 1995), why size and shape of Gigantopithecus has been very diffi cult, could they not have done the same across the grasslands of since it is known only by isolated teeth and a few . The western, southern and central Asia? By ~3 Ma ago grasslands

J. Biosci. 34(4), October 2009 Human evolution in India 733

Figure 3. Geographical and chronological placement of the some of the important fossil vertebrates at Haritalyangar (modifi ed after Patnaik et al. 2005). (A) (1) the Dangar I locality; (2) the Gigantopithecus (Indopithecus) locality; (3) Gigantopithecus (Indopithecus) molar site Hari Devi I; (4) ostrich-like eggshell locality, Dharamsala; triangles denote other major hominoid localities. (B) Magnetic polarity stratigraphy near Haritalyangar showing the levels of important fossil fi ndings (modifi ed after Pillans et al. 2005): HTA10-S.sivalensis (Sankhyan, 1985), IG-Indopithecus giganteus (Gigantopithecus mandible site), CK-Chob Ka-nala Svalhippus site HD- Hari Devi- Gigantopithecus site, D1-Dangar Sivapithecus site HT-1-Sivaladapis site. Eggshells- Cf. Struthiolithus. existed between East Africa and India (fi gure 5). Currently, Africa, Israel, Spain, Italy (Delson 1993; Delson et al. there is no evidence for the presence of Australopithecines in 1993; Gibert et al.1995; Belmaker 2002; Rook et al. 2004). the Siwalik Hills region. However, artifacts possibly dated to The sabre- felid and the large hyena ~2.0 Ma from Riwat and to 2.0 – 1.0 Ma from the Pabbi Hills which occur in the Pinjor Formation may (both in northern Pakistan) (Rendell et al. 1987; Dennell et also have used this corridor to disperse from Europe during al. 1988) indicate that early Homo possibly had made its the Early Pleistocene (Turner 1992; Rook et al. 2004). way to South Asia during the Plio-Pleistocene boundary. Beside large mammals, small mammals such as murine The northern hemisphere glaciation at ~2.5 Ma ago rodents are common in Late Pliocene deposits of Ethiopia, brought about a major change in the global climate Kenya, Tanzania and Indo-Pakistan (Patnaik 2000, 2001; (Shackleton et al. 1984), an event possibly refl ected in Wynn et al. 2006) and may indicate the absence of any the faunal turnover at the Tatrot-Pinjor boundary in the physical barrier between the Indian subcontinent and East Siwaliks (Patnaik 2003). With the spread of grasslands, the Africa to prevent faunal migrations. The large- fi rst primates to enter Siwaliks ~2.5 Ma ago were colobines Plio-Pleistocene taxa from East African such as Oryx, such as Presbytis sivalensis and Cercopithecines for ex. and Crocuta crocuta (Tchernov 1992) have Macaca palaeindica, Procynocephalus subhimalayanus. also been recorded in the Siwaliks. Siwalik mammals such Theropithecus delsoni probably entered a bit later at around as , gazelle and Hippopotamus have also been found 1 Ma (Delson 1993). Theropithecus, which has invariably at the North African Early Pleistocene hominid site of Ain been globally associated with early Homo is also known Hanech in Algeria (see Dennell 2003 for a review). It has from Plio-Pleistocene deposits of south, east and north been observed that the Upper Irrawaddy fauna of Myanmar 734 R Patnaik and P Chauhan

Figure 4. Select fossil mammals from the hominid interval at Haritalyangar (modifi ed after Patnaik 2008). (a) occlusal view of Gigantopithecus (Indopithecus) mandible (CYP359/68 -Simons and Chopra 1969). (b, c, d) Occlusal, labial and lingual views of Gigantopithecus (Indopithecus) M2 (VPL/HD I-1, Patnaik et al. 2005). (e, f) Lingual and labial views of Sivapithecus I1(VPL/HD I-2). (g, h, i) Occlusal, lingual and labial views of P3 Sivapithecus (VPL/RP-H1, Patnaik and Cameron 1997). (j, k) Cf. Struthiolithus eggshells. (l, m, n) Brachyrhizomys choristos ,dorsal lateral and ventral views of the (VPL/BS/PU 101). (o, p) Labial and occlusal view of a mandible (VPL/BS/PU 102, Flynn et al. 1990). (q, r, s) Lingual, labial and occlusal view of Sivaladapis mandible (P11AS, Gingerich and Sahni 1979). Bar scales represent 1 cm. has several taxa, such as Potamochoerus, Merycopotamus known archaeological evidence for such an early dispersal dissimilis, palaeindicus, sp., Hemibos into South Asia is ambiguous and not clearly established triquetricornis, Rhinoceras sivalensis, insignis (discussed below). This refl ects major palaeontological and and Elephas hysudricus, which have also been recorded archaeological gaps prior to the Middle Pleistocene, some in the Pinjors (Takai et al. 2006). The faunal remains from of which may be due to the lack of absolute dates for known Xiaochangliang, China, like Cervus and Gazella (Zhu et al. artifact assemblages and possibly due to the discontinuous 2001) are also found in the Pinjor deposits of India. Other occupation of this region prior to the early Middle common forms occurring in China as well as in Pinjors are Pleistocene (Dennell 2003). The only-known pre-modern Pachycrocuta brevirostris, Equus, Rhinoceras, Stegodon, hominin fossil in the subcontinent may be contemporary Coelodonta, Potamochoerus and Sus (Colbert 1940; Tang with the Late Acheulean or early Middle Palaeolithic 1980; Han 1987; He 1997; Zhu et al 2003). The Pliocene phase(s) and has been recovered from Hathnora (fi gure 6) in monkey from China, Procynocephalus wimani (Schlosser the central Narmada Valley (Sonakia 1984; Kennedy 2001). 1924), has been found to be very similar to Procynocephalus This partial calvarium (fi gure 6 inset) was supplemented by subhimalayanus of Pinjor (Verma 1969). It appears that the possibly-associated clavicles and a rib fragment (Sankhyan Pliocene conditions in India during the Pinjor times were 1997a,b; 2005). The calvarium (possibly that of a female) conducive to possibly allow dispersion of early hominin was originally identifi ed as an ‘advanced’ Homo erectus (de through the region. Lumley and Sonakia 1985) and later re-classifi ed as an archaic or early form of H. sapiens (Kennedy et al. 1991). A detailed 4. The Pleistocene phylogenetic analysis of the Narmada calvarium reveals that it falls between the Steinheim (H. heidelbergensis) and H. The re-dating of Javan H. erectus by Swisher et al. (1994) neanderthalensis (fi gure 7), which in turn strongly supports to ~1.8 Ma hints that the Indian subcontinent may have an European connection for the Narmada hominin as been utilized as a geographic corridor between East opposed to a strictly Asian origin (Cameron et al. 2004). The Africa and Southeast Asia. Unfortunately, the currently Narmada hominin calvarium has recently been classifi ed as

J. Biosci. 34(5), November 2009 Human evolution in India 735

Figure 5. The extent of grasslands ca. 3 Ma: note the absence of a Saharo-Arabian desert barrier between Africa and Asia at this time. (Source: Dowsett et al. 1994.)

Homo sp. indet. by Athreya (2007). Mammals such as diverse ecological environments throughout the entire region, Stegodon namadicus, Equus namadicus, Hippopotamus and which must have collectively affected early hominin namadicus, Sus namadicus, and Cuon alpinus have been ecological adaptations, subsistence strategies and seasonal found to be stratigraphically associated with the calvarium dispersal patterns within India. Indeed, the South Asian from Hathnora. This mammalian assemblage, belonging to faunal evidence offers valuable opportunities for comparative the Surajkund Formation, broadly suggests a warm climate taxonomic studies, isotope analyses, palaeoenvironmental with intermittent arid phases in the late middle Pleistocene. reconstructions, biochronological correlations and inter- In general in the Narmada Valley, it has been found that the regional faunal migration links (with West Asia and Southeast early to middle Pleistocene vertebrate assemblages, including Asia, respectively). cyprinid fi shes, crocodiles, Hippopotamus palaeindicus, and Elephas hysudricus suggest a warm climate, whereas 4.1 The archaeological evidence presence of wild ass (Equus hemionus khur) and ostrich (Struthio camelus) may indicate drier conditions in the In contrast to the meager hominin fossil record, the stone terminal part of the upper Pleistocene (Patnaik et al. 2009). tool record of the Indian subcontinent is overwhelming and Although Pleistocene vertebrate faunal assemblages have lithic assemblages have been recovered in diverse contexts been recovered from throughout the subcontinent, including from various ecological regions (Sankalia 1974). Although a the Siwalik Hills, central and peninsular India and Sri Lanka robust chronological framework is lacking for most of this (often in stratigraphic or spatial association with stone evidence, a complete sequence of Palaeolithic occupation tools), no convincing evidence of butchery has been clearly from at least the early Middle Pleistocene has been clearly demonstrated (Chauhan 2008a). Nonetheless, this rich established and separated into the traditional triparate Lower, vertebrate palaeontological evidence (Badam 2002) points to Middle and Upper Palaeolithic phases (for latest respective

J. Biosci. 34(5), November 2009 736 R Patnaik and P Chauhan

Figure 6. Narmada hominin locality, Hathnora and the hominin cranium shown in inset (courtesy Arun Sonakia). reviews, see Settar and Korisettar 2002). Although broad stratifi ed or surface association with fi ne-grained fl uvial ‘transitions’ between these techno-chronological phases and lacustrine sediments, ferricretes, laterites, and gravel have been presumed based on evidence from other regions or conglomerate deposits. Most of the Indian localities have of the Old World, a clear understanding of the precise been directly dated through the Uranium-Thorium (234Th- of the South Asian cultural shifts remains to be properly 230U) and thermoluminescence (TL) methods and include worked out. This current drawback is collectively due to a predominance of Acheulean sites (Mishra 1995). Ages the dearth of absolute dates, few excavations of extensive for other occurrences such as Riwat, Dina, Jalapur, Pabbi cultural sequences and prominent sterile horizons at Hills, Morgaon, and Satpati Hill have been estimated using culturally-continuous sites (Chauhan 2009d). In any case, palaeomagnetism and geostratigraphic correlations (Chauhan it is clear that the South Asian Palaeolithic record preserves 2009a). At Teggihalli, Chirki-Nevasa, and Yedurwadi, the typo-technological attributes shared by other regions (e.g. 234Th-230U ages for the Acheulean extend beyond 350 Ka (or Acheulean) as well as assemblages resulting from indigenous 390 at Didwana), the maximum limit of the dating methods, mechanisms of cultural evolution, particularly from the early an assessment partly supported by lithic typology. With the Middle Palaeolithic onwards (Petraglia 2008). possible exceptions of the Satpati Hill site in Nepal and The South Asian Lower Palaeolithic has been Morgaon and Chirki-on-Pravara in Maharashtra, there is traditionally divided into core-and-fl ake and Acheulean no unequivocal evidence of Acheulean occupation prior to lithic industries that occur independently as well as in the Middle Pleistocene in the subcontinent. Though the site shared geographic and geomorphologic contexts (Jayaswal of Isampur in the Hunsgi Valley has been dated to c. 1.27 1982; Petraglia 1998; Gaillard and Mishra 2001; Chauhan Ma using electron spin resonance (ESR) on teeth 2009a). These assemblages are frequently found in associated with the cultural horizons (Paddayya et al. 2002),

J. Biosci. 34(5), November 2009 Human evolution in India 737

Figure 7. Cladogram showing place of the Narmada hominin in the broad phylogeny of the Pleistocene hominins (after Cameron et al. 2004). this estimate is preliminary and requires corroboration. In abundance issues. Beyond this evidence, no unequivocal sum, the South Asian Acheulean evidence chronologically evidence of pre-Acheulean assemblages is known from the ranges from the early Middle Pleistocene to the early Late entire subcontinent. Systematic surveys in the Siwalik Hills Pleistocene. of northern India and other regions in central and peninsular One of the fi rst claims of pre-Acheulean evidence in the India may eventually yield Oldowan assemblages - but only subcontinent was made through the Soanian industry by if the region was a corridor for early Homo from East Africa de Terra and Patterson (1939). Later work by the British to Southeast Asia (Chauhan 2009b). Given the abundance Archaeological Mission to Pakistan (BAMP) resulted in a of stone raw material sources, diverse faunal populations, major revision of de Terra and Paterson’s interpretations. a lengthy monsoon regime and conducive environmental Subsequently, multiple lines of evidence, including a conditions during the South Asian Plio-Pleistocene, it is comparison of Soanian and Acheulean technology, landscape likely that the region witnessed pre-Acheulean hominin geoarcheology, surveys of dated geological features, and a occupations. comparative morphometric analysis, clearly revealed that The Acheulean evidence is much more prominent and the majority of Soanian assemblages, if not all, comprise better studied: with the exception of northeast India and Levallois elements and probably postdate the Acheulean parts of Konkan Maharashtra, western Kerala, south of the (Chauhan 2009a). Subsequent claims for a pre-Acheulean Cauvery River in Tamil Nadu and Sri Lanka, Acheulean occupation have come from central India and the Siwalik assemblages are found throughout most of the Indian Hills in Pakistan and India (Chauhan 2009b, c). The most subcontinent (Misra 1989; Pappu 2001a, b; Petraglia systematically studied of all these pre-Acheulean claims 2006). The South Asian Acheulean is generally divided come from Riwat and the Pabbi Hills in the Siwalik deposits into Early or Late developmental phases, based primarily of northern Pakistan (Dennell et al. 1988), and is also the on typo-technological features, assemblage compositions, most controversial because of contextual and artifact- comparative stratigraphy and associated metrical analyses

J. Biosci. 34(5), November 2009 738 R Patnaik and P Chauhan

(Paddayya 1984). The behavioural record is particularly systematically justifi ed. Early Acheulean assemblages continuous from the early Middle Pleistocene and comprises are known to comprise handaxes (fi gure 8), choppers, a rich and diverse array of technological, structural and polyhedrons, and spheroids, usually a lower number of symbolic evidence. As more absolute dates and detailed cleavers (but not always) and fl ake tools, the predominant metrical data become available, current classifi cations of use of the stone-hammer technique, and a marked absence of many assemblages are likely to change. While the term the Levallois technique (Misra 1987). The Early Acheulean ‘Middle Acheulean’ has been occasionally applied to bifaces are often asymmetrical, large with thick butts or mid- ‘transitional’ assemblages, such a facies have never been sections and possess large, bold and irregular fl ake scars,

Figure 8. Select stone tools from Narmada Valley. (a) Early Pleistocene chopper from Dhansi Formation. (b) Early Acheulean handaxe from Pilikarar. (c) Late Acheulean handaxe from Surankund Formation. (d) Late Acheulean miniature hand axe from Surajkund. (e) Late Acheulean cleaver from Surajkund Formation (Patnaik et al. 2009).

J. Biosci. 34(5), November 2009 Human evolution in India 739 indicative of hard-hammer percussion. In contrast, Late a Middle Palaeolithic tradition with a prominent heavy-duty Acheulean assemblages are defi ned by the low proportion tool component instead of representing a Mode 1 technology of bifaces, the high ratio of cleavers to hand axes, the very thought to precede the Acheulean (see Chauhan 2007; 2008a, high ratio of fl ake tools such as scrapers, and the extensive b for reviews, research results and related citations). employment of the soft-hammer technique and the Levallois The super volcanic eruption of Toba (northern Sumatra) and discoid-core techniques (Misra 1987). These bifaces ~74 ka ago led to the deposition of a blanket of volcanic ash are also generally smaller, thinner, and morphologically all over India (William and Royce 1982; Rose and Chesner more refi ned, with a signifi cant increase in the degree of 1987, 1990). Terrestrial deposits of this Tephra have been retouching and controlled bifacial thinning/fl aking (fi gure recorded in several river Valleys throughout India (Acharyya 8). Early Acheulean characteristics have been identifi ed at and Basu 1993; Shane et al. 1995; Westgate et al. 1998; such sites as Chirki-Nevasa, Morgaon, Pilikarar, Singi Talav Jones 2007; Raj 2008). Researchers have referred this ~74 and Satpati Hill while Late Acheulean characters have been ka eruption to be a ‘super eruption’ with a DRE (Dense Rock recognized at Attirampakkam, Bhimbetka, Raisen District, Equivalent) estimate of ~2800 km3 (Knight et al. 1986; Rose Hunsgi-Baichbal Valleys, the Kaldgi Basin and Gadari and Chesner 1987, 1990; Rampino 2002; Mason et al. 2004) (among many others) (Pappu 2001a, b; Petraglia 2006). and ~800 km3 of ash (Viseras and Fernandez 1995). This huge The gradual shift in characterization from ‘‘Middle eruption has been estimated to be much larger in magnitude Stone Age’’ to the ‘‘Middle Palaeolithic’’ for the South compared to those of very recent times such as1815 Tambora Asian evidence is thought to be due to the Mousterian and and 1991 Pinatubo eruptions (Stothers 1984; McCormick et Levallois affi nities between assemblages in the northwestern al. 1995). It has been proposed (Rampino et al. 1988) that region of the subcontinent and other penecontemporaneous the after affects of Toba Supervolcano can be equated with occurrences in Central Asia, northern Africa, and Europe a nuclear winter as modeled by Turco et al. (1983; 1990). (Kennedy 2003). Separating the Middle Palaeolithic horizons Further, the volcanic winter could have decreased the land from the Late Acheulian ones, however, has proved to be a temperatures between 30-700 Latitude by ~5-150C (Rampino recurrent methodological problem (Mishra 1995) because and Self 1992). This super eruption has been found to the Levallois technique and other forms of prepared-core coincide with a major shift in global climatic conditions technology are also present in the Late Acheulean phase from interglacial marine isotopic stage (MIS) 5 to glacial of the subcontinent. Additionally, the Middle Palaeolithic MIS 4 (Ninkovich et al. 1978; Kale et al. 2004). Regarded sites often overlap geographically with the Late Acheulean as the largest volcanic eruption in the last two million , occurrences and indicate successful adaptations and this eruption must have caused signifi cant change in the exploitation of a range of ecological and topographic global climate leading to widespread biotic and a settings. In comparison with the South Asian Acheulean, severe reduction in the human population (Ninkovich et al. the four features that distinguish Middle Palaeolithic 1978; Ambrose 1998). Although a recent study claims that assemblages are: (i) a decrease in size of the artifacts, (ii) early humans survived this devastating event (Petraglia et a noticeable shift from large Acheulian bifaces to more al. 2007), only long-term and large-scale projects at multiple smaller, specialized tools (iii) an increase in the prepared- locations in India will reveal the extent of the YTT impact(s) core technique, and (iv) a preference for fi ne-grained raw on human populations material (such as quartz, fi ne-grained quartzite, chert, The South Asian Upper Palaeolithic is not as clearly jasper, chalcedony, fl int, agate, crypto-crystalline silica, defi ned (James and Petraglia 2005) as the regional lydianite and bloodstone (Kennedy 2003). In some regions Acheulean or the South Asian Middle Palaeolithic nor such as Rajasthan, parts of Andhra Pradesh, parts of coastal well understood; as a result, it still requires extensive Maharashtra, and the Narmada Valley, quartzite continues multidisciplinary research at a large scale (Chauhan 2009d). to be used. Some of the new types that either fi rst appear or The dominating and defi ning features of South Asian Upper become prominent in the South Asian Middle Palaeolithic Palaeolithic assemblage compositions include a notable are prepared-cores, discoids, fl akes, fl ake-scrapers, borers, increase in the production of more specialized tools such awls, blades, and points. Since choppers and diminutive as blades, burins, and borers. Although the production of handaxes are often found in certain Middle Palaeolithic blades is known from Late Acheulean levels at a few sites contexts, it may be suitable to arbitrarily divide South Asian (e.g. Bhimbetka), this behavior became highly prominent, Middle Palaeolithic assemblages into two separate groups: prolifi c, and technologically consistent and standardized light-duty assemblages and heavy-duty assemblages. The only during the Upper Palaeolithic. Additional tool types factors for such variation in assemblage composition may during this technochronological period include fl akes, include function, raw material variability, ecology, culture, knives, awls, scrapers, cores including cylindrical types, style, and/or natural post-depositional formation processes. choppers, and bone tools. The techniques of making many The Soanian evidence is also now increasingly classifi ed as of these lithic and nonlithic tools also changed from the

J. Biosci. 34(5), November 2009 740 R Patnaik and P Chauhan preceding technochronological phases. For example, the use (e.g., steep, convex, convergent) and backed blades (Misra of pressure fl aking and the soft hammer technique for fl ake 2001). Figure 9 illustrates the key Palaeolithic localities in detachment appears to increase signifi cantly as compared the Indian Subcontinent, some of which have been discussed with the South Asian Middle Palaeolithic and the later briefl y in this paper. Acheulean. The degree or intensity of retouch also appears In the terminal Pleistocene cool and dry conditions to increase considerably compared with Lower and Middle corresponding to the Last Glacial Maxima (LGM) prevailed Palaeolithic assemblages in general. Compared with earlier in the Indian subcontinent. Conditions between ~18,000 to Palaeolithic technology, the South Asian Upper Palaeolithic ~13,000 yrs. have been referred to as hyper-arid (Pant 2003). shows a greater degree of regional typo-technological When compared to present day climate, such dry conditions variation as well as an increase in different types of scrapers were probably due to low precipitation of summer monsoon

Figure 9. Locations of key occurrences in the Indian Subcontinent.

J. Biosci. 34(5), November 2009 Human evolution in India 741 and a higher winter precipitation (Singh et al. 1990). Nilgiri ethnic populations of India and that some of the earliest Hills of southern India also experienced LGM climate continuous occupiers of the region may have been Austro- (Sukumar et al. 1993). Under such conditions predominance Asiatic tribal populations although the Dravidians have also of tropical grass vegetation during 20-16 ka clearly been ancestral contenders. Modern tribal groups (461) make indicates a very arid phase as this type of vegetation grows up over 8% of the Indian population (Census of India 1991) favourably under low aridity and low soil moisture. This and the majority is separated geographically, linguistically also points to a period of weak southwest summer monsoon and culturally and many groups, up to varying levels, have during LGM. It has been noticed that the change in climate retained traditional or primitive modes of subsistence; and vegetation has adversely infl uenced the structure and though not as extensive as some other regions in the Old composition of the montane ecosystem (Sukumar et al. World. For example, the long-term geographical isolation 1995). Similar conditions of LGM have also been identifi ed of the Andaman Islanders is signifi cant and mitochondrial in the Central Narmada Valley (Patnaik et al. 2009). sequences obtained from museum specimens revealed that some of their physical attributes converge with African pygmoid groups representing a southern movement from 5. The Holocene Africa (Endicott et al. 2003); although Indians are genetically classed as Caucasoid (Kivisild 2000). It should also be noted The terminal Pleistocene cool and dry conditions of LGM that some of these early tribal populations have retained some were followed by warm and humid conditions of early of their genotype by not mixing with later groups migrating Holocene time. Around ~7–10 ka the northwestern part into the subcontinent and at the same time, Indian tribal of India was very warm and wet (Goodbred and Kuhel and caste populations derive predominantly from a shared 2000; Fleitmann et al. 2003), which would have facilitated gene source in southern and western Asia (Kivisild et al. the hunter-gatherers to domesticate plants and , 2003). Studying the variation in their DNA will help explain followed by the start of agriculture (Gupta 2004). Eventually, biological affi nities as well as gene fl ow between peninsular agriculture led to the permanent settlement of populations. Indian populations and Southeast Asian populations. For Invasion of Aryans into India is still a hotly debated issue. example, it has been demonstrated by Reddy and Stoneking It has been hypothesized that Harappan settlers (6000-2000 (1999) that Australian Aboriginals are very distinct and BC) were basically the modern humans that came from closely resembles certain Indian groups (also see Kumar Africa around 60,000 yrs ago, and were different from and Reddy 2003). Such affi nities or independent origins of the Aryans that arrived via Europe after the LGM (Vahia Indian populations has been studied by understanding the 2004). The main difference between these two populations 9-bp depletion between COII/tRNA and comparing it with hinges on the idea that the Harappan sites lack any evidence African and eastern Asian populations: Koya and Chenchu of ‘Horses’, whereas, they were integral part of ritualistic tribal groups differ by seven HVS-I mutations, inheriting a customs mentioned in the Rigvedic literature supposed to subset of African mtDNA lineages (see Kivisild et al. 2003 have been written between 4000-2000 BC (Thapar 2003). for explanations). Furthermore, the frequency of haplogroup M and its diversity are highest in India (Edwin et al. 2002) 5.1 Changing human populations in South Asia: the suggesting that the subcontinent was settled soon following genetic evidence the archaic Homo dispersal from Africa, neither showing complete nor a replacement of the initial hominin In recent years, there has been increasing interest in the groups (Kivisilid et al. 2003). Though the archaeological genetic background of the human population diversity of the record is rich and continuous, such vital information as Indian subcontinent. In fact, physical differences in South radiometric dates, hominin fossils, and high-resolution Asian populations have been a subject of study since the palaeoenvironmental data is currently inadequate to answer early British occupation of India. More recently, a number questions regarding dispersals and population movements of studies have been conducted to explain the biological within South Asia and its periphery zones. diversity as well as subsequent historical developments Recently Eswaran (2002) and Eswaran et al. (2005) of caste and regional group origins (Bamshad et al. 2001; make an important argument about gene fl ow rather Cordaux et al. 2004). Though not generally utilized to than direct population movement outside of Africa. understand hominin dispersal patterns, such studies have However, it is currently diffi cult to test such inferences important implications for understanding the dispersal of from the archaeological record and the fossil record is too archaic Homo (Quintana-Murci et al. 1999) into South Asia fragmentary in key regions (western China, India, Russia, rather than for older Homo species. For example, Majumder etc.) to reveal any patterns of gene fl ow at such an early (2001) has presented evidence that mitochondrial DNA stage in hominin dispersals. In fact, hominin groups appear haplotypes (based on RFLPs) are remarkably similar across to have adapted to different palaeoenvironments at regional

J. Biosci. 34(5), November 2009 742 R Patnaik and P Chauhan levels over a long period of time and in relative isolation. For for ancestors of Miocene hominoids in the Oligocene example, experimental studies involving physical exercises deposits; (3) recovering potential hominid ancestors among between tropical (Malaya and Indian) and temperate the Miocene hominoids; (4) what did Gigantopithecus (Chinese) subjects were conducted by Duncan and Horvath exactly look like?; (5) whether hominids were present in (1988), which showed differential adaptation to varying heat the Pliocene?; (6) Who were the Early Pleistocene hominins stress levels. Another phenotypic example is shovel-shaped (if they were present in the region at that time)?; and (7) of extant East Asian populations also found in the evidence concerning the arrival/evolution of modern human fossil hominin record from the same region. The lack of groups into the region. Current and future multidisciplinary this feature in South Asians is an example of evolution in research projects within the Indian subcontinent are bound to geographical isolation, following a certain demographic reveal exciting new information and changing interpretations saturation point. In other words, there may have been regarding the global role and impact of the Indian landmass relatively minimal gene fl ow between peninsular Indian on primate and human evolution. and East/Southeast Asian groups during earlier Pleistocene times. This is also tentatively supported by the work of Acknowledgements Bamshad et al. (2001) who demonstrate, through mtDNA and y-chromosome variation, that there was minimal or We would like to thank Profs. Sunil Bajpai and Ashok Sahni marginal movement in the northeastern corridor of the for inviting us to contribute to this volume. We are grateful to subcontinent. Additional evidence collected by Kivisild et Claire Gaillard for offering valuable and helpful comments al. (2000) led them to consider India as a part of the source on an earlier draft of this paper. RP thanks Wenner-Gren gene pool ancestral to maternal lineages in Europe. Most Foundation (Grant No. Gr.ICRG-43) and Department of recently, a new genetics study (Reich et al. 2009) reveals that Science and Technology, New Delhi for the fi nancial support northern Indian and southern Indian populations come from (SR/S4/ES-171/2005). We would like to thank National geographically-separate ancestral lineages which disporsed Geographic Society (Grant 7386-02 to RP) and Wenner-Gren into the subcontinent at different times in the past. Foundation for Anthropological Research (Grant 7541 to PC) for funding the Narmada Basin Palaeoanthropology Project. 6. Discussion and conclusions References There is every possibility that the collision of India with Asian landmass in the early Eocene led to the initial Acharyya S K and Basu P K 1993 Toba ash on the Indian dispersal of anthropoid primates into Eurasia. Beside the subcontinent and its implications for correlation of late records from Eocene and Oligocene, there is almost no Pleistocene alluvium; Quat. Res. 40 10–19 record of anthropoid primates until ~13 Ma ago. The Late Amano K and Taira A 1992 Two-phase uplift of higher Himalayas Miocene ape Sivapithecus was primarily arboreal, but since 17 Ma; Geology 20 391–394 some terrestrial locomotion by this ape cannot be ruled Ambrose S N 1998 Late Pleistocene human population bottlenecks, out. Gigantopithecus was defi nitely a terrestrial ape that volcanic winter, and differentiation of modern humans; J. Hum. lived on grasses, fruits and possibly bamboos. Again there Evol. 34 623–651 Andrews P 1982 Hominoid evolution; Nature (London) 295 exists a lengthy gap until the Middle Pleistocene, where we 185–86 have defi nite Acheulean artifacts produced most probably Andrews P and Cronin J 1982 The relationships of Sivapithecus by H. erectus or H. heidelbergensis populations. Then, and Ramapithecus and the evolution of the orangutan; Nature there is the only defi nite hominin fossil, the Narmada (London) 297 541–546 hominin which has been assigned to archaic H. sapiens/H. Andrews P and Walker A 1979 The primate an other fauna erectus H. heidelbergensis/Homo sp. by various specialists. from Fort Ternan, Kenya; in Human origins (eds) G Isaac Genetic evidence demonstrates that the gene pool to which and E McCown (Menlo Park, California: WA Benjamin) pp Indian populations belong is highly diverse and comprises 279–306 a large number of tribal groups, many of which are linked Ashton P S and Gunatilleke C V S 1987 New light on the plant at different levels with both European and Asian lineages. geography of Ceylon, Historical plant geography; J. Biogeogr. Though, the palaeoanthropological data so far appears to 14 249–285 Athreya S 2007 Was in South Asia? be discontinuous, India with appropriate deposits ranging A test using the Narmada fossil from central India; in The in age from the Palaeocene to the Pleistocene and covering evolution and history of human populations in South Asia (eds) a vast geographic area, offers great potential for further M D Petraglia and B Allchin (New York: Springer Press) pp palaeoanthropological exploration and investigations. The 137–170 critical areas of research include: (1) searching for the earliest Badam G L 2002 Vertebrate in India: anthropoids in Palaeocene and Eocene deposits; (2) searching Fifty Years of Research; in Indian archeology in retrospect-

J. Biosci. 34(5), November 2009 Human evolution in India 743

Archaeology and interactive disciplines Vol. 3 (eds) S Settar and Chauhan P R 2008a Large mammal fossil occurrences and R Korisettar (Delhi: Manohar and Indian Council of Historical associated archaeological evidence in Pleistocene contexts of Research) pp 209–246 peninsular India and Sri Lanka; Quat. Int. 192 21–42 Bajpai S, Kay R F, Williams B A, Das D P, Kapur V V and Tiwari Chauhan P R 2008b Soanian lithic occurrences and raw material B N 2008 The oldest Asian record of Anthropoidea; Proc. Natl. exploitation in the Siwalik Frontal zone, northern India: a Acad. Sci. USA 105 11093–11098 geoarchaeological approach; J. Hum. Evol. 54 591–614 Bamshad M, Kivisild T, Watkins W S, Disxon M E, Ricker C E, Chauhan P R 2009a The Lower Paleolithic of the Indian Rao B B, Naidu J M, Prasad B V R, Reddy P G, Rasanayagam Subcontinent; Evol. Anthropol. 18 62–78 A, Papiha S S, Villems R, Redd A, Hammer M F, Nguyen S V, Chauhan P R 2009b The Indian subcontinent and ‘Out of Africa I’; Carroll M L, Batzer M A and L B Jorde 2001 Genetic Evidence in Out of Africa I: The fi rst Hominins of Eurasia (eds) J Fleagle, on the Origins of Indian Caste Populations; Genome Res. 11 J Shea and R Leakey (Kluwer Academic Press. Paleobiology 994–1004 and Series) (in press) Barry J C, Morgan M E, Flynn L J, Pilbeam D, Behrensmeyer A Chauhan P R 2009c Core-and-fl ake assemblages of India; in K, Raza S M, Khan I A, Badgley C, Hicks J and Kelley J 2002 Asian Paleoanthropology: From Africa to China and beyond Faunal and environmental change in the latest Miocene Siwaliks (Book Series: Vertebrate paleobiology and paleoanthropology) of northern Pakistan; Paleobiol. Memoirs 28 1–72 (eds) C J Norton and D Braun (Kluwer Academic Press) Beard K C, Tong Y, Dawson M R, Wang J and Huang X 1996 Earliest (in press) complete of an anthropoid primate from the late middle Chauhan P R 2009d The South Asian Paleolithic record and its Eocene of Shanxi Province, China; Science 272 82–85 potential for transitions studies; in Sourcebook of paleolithic Belmaker M 2002 First evidence of the presence of Theropithecus transitions: Methods, theories and interpretations (eds) M sp. in the Southern Levant; Israel J. Zool. 48 165 Camps and P R Chauhan (Springer Press) pp 121–139 Berggren W A, Kent D V and van Couvering J A 1985 Ciochon R L 1985 Fossil ancestors of Burma; Nat. Hist. 94 26–36 geochronology and chronostratigraphy; in The chronology of Ciochon R L and Fleagle J G 1985 Ramapithecus and human the geological record (ed.) N J Snelling (Oxford: Blackwell origins; in Primate evolution and origins (eds) R L Ciochon Scientifi c) pp 211–260 and J G Fleagle (Menlo Park, California: W A Benjamin) pp Begun D R, Güleç E and Geraads D 2003 Dispersal patterns of 207–208 Eurasian hominoids: Implications from Turkey; in Distribution Ciochon R L, Piperno D R and Thompson R G 1990 Opal and migration of tertiary mammals in Eurasia (eds) J W F phytoliths found on the teeth of the extinct ape Gigantopithecus Reumer and Wessels W Deinsea (Rotterdam: Deisea) vol 10, blacki: implications for paleodietary studies; Proc. Natl. Acad. pp 23–39 Sci. USA 87 8120–8124 Burkle L H 1989 Distribution of diatoms in sediments of the Ciochon R L, Long V T, Roy L, Gonzalez L, Grun R, De Vos J, northern Indian ocean: Relationship to physical oceanography; Younge C, Taylor L, Yoshida H and Reagan M 1996 Dated Marine Micropaleontol. 15 53–65 co-occurrence of Homo erectus and Gigantopithecus from Bossuyt F and Milinkovich M C 2001 Amphibians as indicators of Tham Khuyen Cave, Vietnam; Proc. Natl. Acad. Sci. USA 93 Early Tertiary “Out –of-India” dispersal of vertebrates; Science 3016–3020 292 93–95 Colbert E H 1940 Pleistocene mammals from the Ma Kai valley of Brunet M, Beauvilain A, Coppens Y, Heintz E, Aladji H E Moutaye Northern Yunnan, China; Am. Mus. Novitates 1099 1–10 and David P 1995 The fi rst 2,500 kilometres Cordaux R, Aunger R, Bentley G, Nasidze I, Sirajuddin S M and M west of the Rift Valley (Chad); Nature (London) 378 273–275 Stoneking 2004 Independent origins of Indian caste and tribal Cameron D W 2001 The taxonomic status of the Siwalik Late paternal lineages; Curr. Biol. 14 231–235 Miocene hominid Gigantopithecus (‘Indopithecus’); Himalayan Culotta F 1995 Asian anthropoids strike back; Science 270 918 Geol. 22 29–34 Delson E 1993 Theropithecus fossils from Africa and India and the Cameron D W 2003 A functional and phylogenetic interpretation of of the Genus; in Theropithecus: The rise and fall of the late Miocene Siwalik hominid Indopithecus and the Chinese a primate genus (ed.) N G Jablonski (Cambridge: Cambridge Pleistocene hominid Gigantopithecus; Himalayan Geol. 24 University Press) pp 157–189 19–28 Delson E, Eck G G, Leakey M G and Jablonski N G 1993 A partial Cameron D, Patnaik R and Sahni A 2004 The phylogenetic catalogue of fossil remains of Theropithecus; in Theropithecus: signifi cance of the Middle Pleistocene Narmada Hominin; Int. The rise and fall of a primate genus (ed.) N G Jablonski J. Osteoarchaeol. 14 419–447 (Cambridge: Cambridge University Press) pp 499–525 Cerling T E, Harris J M, MacFadden M G, Leakey M G and Quade de Lumley H and Sonakia A 1985 Contexte stratigraphique et J 1997 Global vegetation change through the Miocene/Pliocene archéologique de l’homme de la Narmada, Hathnora, Madhya boundary; Nature (London) 389 153–158 Pradesh, Inde ; L’ Anthropologie 89 3–12 Chauhan P R 2006 Human Origins Studies in India: Position, Dennell R W 2003 Dispersal and colonisation, long and short Problems, and Prospects. Assemblage 9. Available at: www.asse chronologies: how continuous is the Early Pleistocene record mblage.group.shef.ac.uk/issue9/chauhan.html for hominids outside East Africa?; J. Hum. Evol. 45 421–440 Chauhan P R 2007 Soanian cores and core-tools from Toka, Dennell R W and Roebroeks 2005 Out of Africa: An Asian northern India: towards a new typo-technological organization; perspective on early human dispersal from Africa; Nature J. Anthropol. Archaeol. 26 412–441 (London) 438 1099–1104

J. Biosci. 34(5), November 2009 744 R Patnaik and P Chauhan

Dennell R W, Rendell H M and Hailwood E 1988 Early tool Tibetan uplift, denudation, and climatic histories in two molasse making in Asia: Two Million old artifacts in Pakistan; deposits; J. Geol. 101 157–175 Antiquity 62 98–106 Hay W W 1996 Tectonics and climate; Geol. Rundsch. 85 409–437 De Terra H and Patterson T T 1939 Studies on the ice age in India He Z 1997 Yuanmou hominoid fauna (Kunming, China: Yunana and associated human culture (Carnegie Instit. Washington Science and Technology Press) pp 270 Publ.) pp 493 Han D 1987 Artiodactyla fossils from Liucheng Gigantopithecus Dowsett H, Thompson R, Barron J, Cronin T, Fleming F, Ishman Cave in ; Mem. Inst. Vert. Palaeontol. Palaeoanthropol. S, Poore R, Willard D and Holtz T J 1994 Joint investigations 18 135–208 of the Middle Pliocene climate I: PRISM palaeoenvironmental Hoorn C, Ojha T and Quade J 2000 Palynological evidence reconstructions; Global Planetary Change 9 169–195 for vegetation development and climatic change in the sub- Duncan M T and Horvath S M 1988 Physiological adaptations to Himalayan zone (Neogene, Central Nepal); Palaeogeogr. thermal stress in tropical Asians; Eur. J. Appl. Physiol. Occupat. Palaeoclimatol. Palaeoecol. 163 133–161 Physiol. 57 540–544 James H V A and Petraglia M D 2005 Modern human origins and Edwin D, Vishawanathan H, Roy S, Usha Rani, M, Majumder the evolution of behavior in the later Pleistocene record of South P 2002 Mitochondrial DNA diversity among fi ve tribal Asia; Curr. Anthropol. 46 S3–S27 populations of southern India; Curr. Sci. 83 158–16 Jayaswal V 1982 Chopper-chopping component of Palaeolithic Endicott P, Gilbert M T P, Stringer C, Lalueza-Fox C, Willersley India (New Delhi: Agam Kala Prakashan) E, Hansen A J and A J Cooper 2003The Genetic Origins of the Jones S C 2007 The Toba supervolcanic eruption: Tephra-fall Andaman Islanders; Am. J. Hum. Genet. 72 178–184 deposits in India and Paleoanthropological implications; in The Eswaran V 2002 A Diffusion Wave Out of Africa: The mechanism evolution and history of human populations in South Asia (eds.) of the modern human revolution?; Curr. Anthropol. 43 M D Petraglia and B Allchin (New York: Springer Press) pp 749–774 137–170 Eswaran V, Harpending H and Rogers A R 2005 Genomics refutes Kale V S, Joshi V U and Hire A S 2004 Palaeohydrological an exclusively African origin of humans; J. Hum. Evol. 49 reconstructions based on analysis of a palaeochannel and Toba- 1–18 Ash associated alluvial sediments in the Deccan Trap region, Fleagle J G 1988 Primate Adaptation and evolution (San Diego: India; J. Geol. Soc. India 64 403–417 Academic Press) Kappelman J, Kelley J, Pilbeam D R, Sheikh K A, Ward S, Fleitmann D Burns S J, Mudelsee M, Neff U, Kramers J, Mangini Anwar M, Barry J C, Brown B, Hake P, Johnson N M, Raza A and Matter A 2003 Holocene forcing of the Indian monsoon S M, and Imbrahim Shah S M 1991 The earliest occurrence recorded in a stalagmite from Southern Oman; Science 300 of Sivapithecus from the middle Miocene Chinji Formation of 1737–1739 Pakistan; J. Hum. Evol. 21 61–73 Flynn L J, Sahni A, Jaeger J J, Singh B and Bhatia S B 1990 Karanth K P 2006 Out-of-India Gondwana origin of some tropical Additional fossil rodents from the Siwalik Beds of India; Proc. Asian biota; Curr. Sci. 90 789–792 K. Ned. Akad. Wet. 93 7–20 Kay R 1981 The nut-crackers—a new theory of the adaptations of Flynn L J, Barry J C, Morgan M E, Pilbeam D R, Jacobs L L, the Ramapithecinae; Am. J. Phys. Anthropol. 55 141–151 and Lindsay E H 1995 Neogene Siwalik mammalian lineages: Kelley J 1986 Paleobiology of Miocene Hominoids, PhD species longevities, rates of change, and modes of speciation; dissertation, Yale University, New Haven, USA Palaeogeogr. Palaeoclimatol. Palaeoecol. 115 249–264 Kelley J 1997 Paleobiological and phylogenetic signifi cance of Gaillard C and Mishra S 2001 The lower Palaeolithic in South like history in Miocene hominoids; in Function, phylogeny and Asia; in Origin of settlements and chronology of the palaeolithic fossils: Miocene Hominoid origins (eds) D Begun, C Ward and cultures in Southeast Asia. Colloque International de la Fondation M Rose (New York: Plenum Press) pp 173–208 Singer Polignac (Under the direction of F Sémah, C Falgueres, D Kelley J and Pilbeam D 1986 The Dryopithecines: taxonomy, Grimaud-Herve, and A Sémah). Semenanjung: Paris comparative anatomy, and phylogeny of Miocene large Gibert J, Ribot F, Gibert L, Leakey M, Arribas A and Martinez- hominoids; in Systematics, evolution, and anatomy (eds) D Navarro B, 1995 Presence of the cercopithecid genus Swindler and D J Erwin (New York: Alan R Liss) pp 361–411 Theropithecus in Cueva Victoria (Murcia, Spain); J. Hum. Evol. Kennedy K A R 2003 God - apes and fossil men: Palaeoanthropology 28 487–493 of South Asia (Ann Arbor: University of Michigan Press) Gingerich P D and Sahni A 1979 Indraloris and Sivaladapis: Kennedy K A R 2001 Middle and Late Pleistocene Hominids of Miocene adapid primates from the Siwaliks of India and South Asia; in Humanity from African Naissance to coming Pakistan; Nature (London) 279 415–416 Millennia (University of Firenze, Firenze) pp 167–174 Goodbred S L and Kuehl S A 2000 Enormous Ganges–Brahmaputra Kennedy K A R, Sonakia A, Chiment J and Verma K K 1991 Is sediment discharge during strengthened early Holocene the Narmada hominid an Indian Homo erectus?; Am. J. Phys. monsoon; Geology 28 1083–1086 Anthropol. 86 475–496 Gupta A K 2004 Origin of agriculture and domestication of plants Kennett J P and Hodell D A 1986 Major events in Neogene oxygen and animals linked to early Holocene climate amelioration; isotopic records; S. Afr. J. Sci. 82 497–498 Curr. Sci. 87 54–59 Kivisild T 2000 The origins of southern and western Eurasian Harrison T M, Copeland P, Hall S A, Quade J, Burner S, Ojha populations: an mtDNA study, PhD Thesis, Tartu University, T P and Kidd W S F1993 Isotopic preservation of Himalaya- Estonia

J. Biosci. 34(5), November 2009 Human evolution in India 745

Kivisild T, Papiha S S, Rootsi S, Parik J, Kaldma K, Reidla M, Nelson S 2003 The extinction of Sivapithecus: Faunal and Laos S, Metspalu M, et al. 2000 An Indian ancestry: a key environmental changes in the Siwaliks of Pakistan (Boston: for understanding human diversity in Europe and beyond; in Brill Academic Publishers) Archaeogenetics: DNA and the population of Europe Nelson S 2005 Paleoseasonality inferred from equid teeth and (eds) C Renfrew and K Boyle (Cambridge: McDonald Institute intra-tooth isotopic Variability; Palaeogeogr. Palaeoclimatol. for Archaeological Research) pp 267–279 Palaeoecol. 222 122–144 Kivisild T, Rootsi S, Metspalu M, Mastana S, Kaldma K, Parik Ninkovich D, Shackleton N J, Abdel-Monem A A, Obradovich J D J, Metspalu E, Adojaan, M, et al. 2003 The Genetic Heritage and Izett G K 1978 K-Ar age of the late Pleistocene eruption of of the Earliest Settlers Persists Both in Indian Tribal and Caste Toba, north Sumatra; Nature (London) 276 574–577 Populations; Am. J. Hum. Genet. 72 313–332 Paddayya K 1984 Stone Age India; in Neue Forschungen zur Knight M D, Walker G P L, Ellwood B B and Diehl J F 1986 Altsteinseit (ed.) H Muller-Karp (Munich: C H Beck Verlag) Stratigraphy, palaeomagnetism and magnetic fabric of the Toba pp 345–403 tuffs: constraints on the sources and eruptive styles; J. Geophys. Paddayya K, Blackwell B A B, Jhaldiyal R, Petraglia M D, Fevrier Res. B 91 10,355–10,382 S, Chaderton II D A , Blickstein J I B, Skinner A R 2002 Recent Kretzoi M 1975 New ramapithecines and Pliopithecus from the fi ndings on the Acheulian of the Hunsgi and Baichbal valleys, lower Pliocene of Rudabanya in northeastern Hungary; Nature Karnataka, with special reference to the Isampur excavation and (London) 257 578–581 its dating; Curr. Sci. 83 641–647 Kumar V and B M Reddy 2003 Status of Austro-Asiatic groups Pant G B 2003 Long-Term climate variability and change over in the peopling of India: An exploratory study based on the monsoon Asia; J. Indian Geophys. Uni. 7 125–134 available prehistoric, linguistic and biological evidences; J. Pappu S 2001a A Re-Examination of the Palaeolithic Archaeological Biosci. 28 507–522 Record of Northern Tamil Nadu, South India (British. Archaeol. Kutzbach J E, Prell W L and Ruddiman W F 1993 Sensitivity of Rep. Int. Series 1003) Pappu R S 2001b Acheulian culture in Peninsular India (New Eurasian climate to surface uplift of the Tibetan Plateau; J. Geol. Delhi: D K Printworld (P) Ltd.) 101 177–190 Patnaik R 2000 Siwalik murid rodents: Origin and dispersal; Leakey L S B 1962 A new lower Pliocene fossil primate from Him. Geol. 21 145–151 Kenya; Ann. Mag. Nat. Hist. Ser. 13 689–696 Patnaik R 2001 Late Pliocene micromammals from Tatrot Lewis G E 1934 Preliminary notice of man-like apes from India; Formation (Upper Siwalik) exposed near village Saketi, Am. J. Sci. 27 161–179 Himalchal Pradesh, India; Palaeontographica 261 55–81 Majumder P P 2001 Ethnic populations of India as seen from an Patnaik R 2008 Revisiting Haritalyangar, the Late Miocene ape evolutionary perspective; J. Biosci. 26 533–545 locality of India; in Elwyn L Simons: A search for origins (eds) J Marivaux L, Antoine P-O, Rafi qul H B S, Benammi M, Chaimanee Fleagle and C Gilbert (New York: Springer) pp 197–210 Y, Crochet J-Y, de Franceschi D, Iqbal N, et al. 2005 Anthropoid Patnaik R and Cameron D W 1997 New Miocene fossil ape primates from the Oligocene of Pakistan (Bugti Hills): Data on locality, Dangar, Hari-Talyangar region, Siwaliks, northern early anthropoid evolution and biogeography; Proc. Natl. Acad. India; J. Hum. Evol. 32 93–97 Sci. USA 102 8436–8441 Patnaik R, Cameron D, Sharma J C and Hogarth J 2005 Extinction Mason B G, Pyle D M and Oppenheimer C 2004 The size and of Siwalik fossil apes: a review based on a new fossil tooth frequency of the largest explosive eruption on the earth; Bull. and on palaeoecological and palaeoclimatological evidence; Volcanol. 66 735–748 Anthropol. Sci. 113 65–72 McCormick M P, Thomason L W, Trepte C R 1995 Atmospheric Patnaik R 2003 Reconstruction of Upper Siwalik palaeoecology effects of Mt. Pinatubo eruption; Nature (London) 373 and palaeoclimatology using microfossil palaeocommunities; 399–404 Palaeogeogr. Palaeoecol. Palaeoclimatol. 197 133–150 Miller S, F Jessica W and Russel L C 2008 Assesing mandi- Patnaik R and David C 2007 On new dates and ecology of recent bular Shape Variation within Gigantopithecus Using a Hominoid fi nds from Haritalyangar area, Himachal Pradesh; Geometric Morphometric approach; Am. J. Phys. Anthropol. in Human origins, genome and people of India: genomic, 137 201–212 palaeontological and archaeological perspectives (eds) A R Mishra S 1995 Chronology of the Indian Stone Age: The impact Sankhyan and V R Rao (New Delhi: Allied Publishers) pp of recent absolute and relative dating attempts; Man Environ. 91–111 20 11–16 Patnaik R, Chauhan P R, Rao M R, Blackwell B A B, Skinner Misra V N 1987 Middle Pleistocene Adaptations in India; in The A R, Sahni A, Chauhan M S and Khan H S 2009 New Pleistocene Old World-Regional perspectives (ed.) O Soffer geochronological, palaeoclimatological and Palaeolithic data (New York: Plenum Press) pp 99–114 from the Narmada Valley hominin locality, central India; J. Misra V N 1989 Stone Age India: An Ecological Perspective; Man Hum. Evol. 56 114–133 Environ. 14 17–64 Patnaik R, Sahni A, Cameron D, Pillans B, Chatrath P, Simons E, Misra V N 2001 Prehistoric human colonization of India; J. Biosci. Williams M and Bibi F 2009 Ostrich-like eggshells from a 10.1 26 491–531 million year old Miocene ape locality, Haritalyangar (Himachal Nanda A C and Sehgal R K 1993 Siwalik Mammalian faunas Pradesh) India; Curr. Sci. 96 1485–1495 from Ramnagar (J. & K.) and Nurpur (H.P.) and Lower limit of Petraglia M D 1998 The Lower Palaeolithic of India and its bearing hipparion; J. Geol. Soc. India 42 115–134 on the Asian record; in Early human behavior in global context:

J. Biosci. 34(5), November 2009 746 R Patnaik and P Chauhan

The rise and diversity of the Lower Palaeolithic record (eds) M D Reddy A J and M Stoneking 1999 Peopling of Sahul: mtDNA Petraglia and R Korisettar (New York: Routledge) pp 343–390 variation in Aborignal Australian and Papua New Guinean Petraglia M D 2006 The Indian Acheulean in Global Perspective; in populations; Am. J. Hum. Genet. 65 808–828 Axe age: Acheulean tool-making from quarry to discard (eds) N Reich D, Thangaraj K, Patterson N, Price A L and Singh L 2009 Goren-Inbar and G Sharon (London: Equinox) pp 389–414 Reconstructing Indian population history; Nature (London) 461 Petraglia M D 2008 the Gap: Factoring the Arabian peninsula 489–495 and the Indian subcontinent into out of Africa models; in The Rendell H M, Hailwood E A and Dennell R W 1987 Magnetic human revolution revisited (eds) P Mellars, O Bar-Yosef, polarity stratigraphy of Upper Siwalik Sub-Group, Soan Valley, K Boyle and C Stringer (Cambridge: McDonald Institute Pakistan: implications for early human occupation of Asia; Archaeological Publications) pp 383–394 Earth Planet. Sci. Lett. 85 488–496 Petraglia M D and Allchin B 2007 The evolution and history of Retallack G J 1991 Miocene Paleosols and ape Habitats of human populations in South Asia; in Inter-disciplinary studies Pakistan and Kenya (New York: Oxford University Press) in archaeology, biological anthropology, linguistics and Retallack G J 1995 Palaeosols of the Siwalik Group as a 15 Mys genetics (New York: Springer) pp 1–464 record of South Asian Palaeoclimate; Mem. Geol. Soc. India 32 Petraglia M, Korisettar R, Boivin N, Clarkson C, Ditchfi eld P, 36–51 Jones S, Koshy J, Lahr M M, et al. 2007 Rook L, Martý´nez-Navarro B and Howell F C 2004 Occurrence assemblages from the Indian Subcontinent before and after the of Theropithecus sp. in the Late Villafranchian of Southern Italy Toba Super-Eruption; Science 317 114–116 and implication for Early Pleistocene “out of Africa” dispersals; Pillans B, Williams M, Cameron D, Patnaik R, Hogarth J, Sahni A, J. Hum. Evol. 47 267–277 Sharma J C, Williams F and Bernor R 2005 Revised correlation Rose M 1986 Further hominoid postcranial specimens from the of the Haritalyangar magnetostratigraphy, Indian Siwaliks: Late Miocene Nagri formation of Pakistan; J. Hum. Evol. 15 implications for the age of the Miocene hominids Indopithecaus 333–368 and Sivapithecus, with a note on a new hominid tooth; J. Hum. Rose W I and Chesner C A 1987 Dispersal of ash in the great Toba Evol. 48 507–515 eruption; Geology 15 913–917 Prell W L, Murray D W, Clemens S C and Anderson D M Rose W I and Chesner C A 1990 Worldwide dispersal of ash and 1992 Evolution and variability of the Indian Ocean summer gases from the earth’s largest known eruption: Toba Sumatra, monsoon: evidence from the western Arabian Sea drilling 75ka; Palaeogeogr. Palaeoclimatol. Palaeoecol. (Global and program; in The Indian ocean: A synthesis of results from Planetary Change Section) 89 269–275 the ocean drilling program (ed.) R A Duncan (Geophysical Ruddiman W F and Kutzbach J E 1989 Forcing of late Cenozoic Monograph, No. 70, American Geophysical Union, Washington northern hemisphere climate by plateau uplift in Southern Asia DC) pp 447–469 and the American West; J. Geophys. Res. 94 409–418 Quade J T, Cerling T E and Bowman J R 1989 Development of the Sankalia H D 1974 The prehistory and protohistory of India and Asian monsoon revealed by marked ecological shift in the latest Pakistan (Pune: Deccan College and Postgraduate Research Miocene of northern Pakistan; Nature (London) 342 163–166 Institute) Quade J, Cater J M, Ojha T P, Adam J and Harrison T M 1995 Sankhyan A R 1985 Late occurrence of Sivapithecus in Indian Late Miocene environmental change in Nepal and the northern Siwaliks; J. Hum. Evol. 14 573–578 Indian subcontinent: stable isotopic evidence from paleosols; Sankhyan A R 1997a A new human fossil from the Central Narmada Geol. Soc. Am. Bull. 107 1381–1397 Basin and its chronology; Curr. Sci. 73 1110–1111 Quintana-Murci L, Semino O, Bandelt H J, Passarino G, Sankhyan A R 1997b Fossil clavicle of a middle Pleistocene McElreavey K and Santachiar-Benerecetti A S 1999 Genetic hominid from the central Narmada Valley, India; J. Hum. Evol. evidence of an early exit of Homo sapiens sapiens from Africa 32 3–16 through eastern Africa; Nat. Genet. 23 437–441 Sankhyan A R 2005 New fossils of Early Stone Age man from Raj R 2008 Occurrence of volcanic ash in the Quaternary alluvial Central Narmada Valley; Curr. Sci. 88 704–707 deposits, lower Narmada basin, western India; J. Earth Syst. Sci. Sarich V M and Wilson A C 1967 Immunological time scale for 117 41–48 hominid evolution; Science 158 1200–1203 Rampino M R 2002 Supereruptions as a threat to civilisations on Schrader H J 1974 Cenozoic marine planktonic diatom stratigraphy Earth-like Planets; Icarus 156 562–569 of the tropical Indian Ocean; Initial Rep. Deep Sea Drill. Proj. Rampino M R and Self S 1992 Volcanic winter and accelerated 24 887–968 glaciation following the Toba super-eruption; Nature (London) Schlosser M 1924 Fossil Primates of China; Palaeontol. Sinica 359 50–52 Ser. D1 1–16 Rampino M R, Self S and Stothers R B 1988 Volcanic winters; Scott R S, J Kappelman and Kelly J 1999 The paleoenvironment of Annu. Rev. Earth Planet Sci. 16 73–99 Sivapithecus parvada; J. Hum. Evol. 36 245–274 Ramstein G, Fluteau F, Besse J and Joussaume S 1997 Effect of Sehgal R K and Nanda A C 2002 Palaeoenvironment and orogeny, plate motion and land-sea distribution on Eurasian palaeoecology of the Lower and middle Siwalik Subgroups of a climate change over the past 30 million years; Nature (London) part of Northwestern Himalaya; J. Geol. Soc.India 59 517–529 386 788–795 Settar S and Korisettar R 2002 Indian archeology in retrospect - Raymo M E and Ruddiman W F 1992 Tectonic forcing of late Archaeology and interactive disciplines Volume 1 (New Delhi: Cenozoic climate; Nature (London) 359 117–122 Manohar and Indian Council of Historical Research)

J. Biosci. 34(5), November 2009 Human evolution in India 747

Shackleton N J, Backman H, Zimmerman D V, Kent M A, Hall D Turco R P, Toon O B, Ackerman T P, Pollock J B, Sagan C, 1983 G, Roberts D, Schitker J, Baldauf A, et al.1984 Oxygen isotope Nuclear winter: Global Consequences of Multiple nuclear calibration of the onset of ice-rafting and history of glaciation in explosions; Science 222 1283–1292 the North Atlantic region; Nature (London) 307 620–623 Turco R P, Toon O B, Ackerman T P, Pollock J B and Sagan C 1990 Shane P, Westgate J, Williams M and Korisettar R 1995 New Climate and Smoke: an appraisal of nuclear winter; Science 247 Geochemical evidence for the youngest Toba Tuff in India; 166–176 Quat. Res. 44 200–204 Turner A 1992 Large carnivores and earliest European hominids: Simons E L and Chopra S R K 1969 Gigantopithecus (Pongidae, changing determinants of resource availability during the Lower Hominoidea): A new species from Northern India; Postilla 138 and Middle Pleistocene; J. Hum. Evol. 22 109–126 1–8 Vahia M 2004 The harappan question. Available at: www.tifr.res.in/ Simons E L and Ettel P C 1970 Gigantopithecus; Sci. Am. 222 ~vahia/harappa.pdf 77–85 Verma B C 1969 Procynocephalus pinjori sp. nov. a new fossil Simons E L and Pilbeam D 1972 Hominid palaeobiology; in The primate from the Pinjor beds (Lower Pleistocene) east of Function and Evolutionary Biology of Primates (ed.) E R Tuttle Chandigarh; J. Palaeontol. Soc. India 13 53–57 (Chicago:Aldine-Atherton) pp 36–62 Viseras C and Fernandez J 1995 The role of erosion and deposition Singh G, Wasson R J and Agrawal D P 1990 Vegetational and in the construction of alluvial fan sequences in the Guadix seasonal climate changes since the last full glacial in the Thar Formation (SE Spain); Geol. Mijnbouw 21–33 desert, northwestern India; Rev. Paleobot. Palynol. 64 351–358 von Koenigswald G H R 1950 Bemerkungen zu Dryopi- 13 Sukumar R, Ramesh R, Pant R K and Rajagopalan G 1993 A C thecus giganteus Pilgrim; Eclogae Geologicae Helvetiae 42 record of late Quaternary climate change from tropical peats in 515–519 southern India; Nature(London) 364 703–706 von Koenigswald G H R 1976 The evolution of man (Ann Arbor: Sukumar R, Suresh H S R and Ramesh R 1995 Climate change and The University of Michigan Press) its impact on tropical montane ecosystem in southern India; J. Westgate J, Shane P, Pearce N, Perkins W, Korisettar R, Chesner C, Biogeogr. 22 533–536 Williams M and Acharyya S K 1998 All Toba tephra occurrences Sonakia A 1984 The skull cap of early man and associated mammalian across peninsular India belong to the 75,000 years B.P. eruption; fauna from Narmada Valley alluvium, Hoshangabad area, Madhya Quat. Res. 50 107–112 Pradesh, India; Rec. Geol. Surv. India 113 159–172 William M A J and Royce K 1982 Quaternary geology of the middle Stothers R B 1984 The great Tambora eruption in 1815 and its Son valley, north central India: implications for prehistoric aftermath; Science 224 1191–1198 Archaeology; Palaeogeogr. Palaeclimatol. Palaeoecol. 38 Swisher C C III, Curtis G H, Jacob T, Getty A G, Suprijo A and Widiasmoro 1994 Age of earliest known hominids in , 139–162 Indonesia; Science 263 1118–1121 West R M 1984 Siwalik fauna from Nepal: palaeoecologic and Takai Masanaru, Saegusa H, Thaung-Htike, Zin-Maung-Maung- palaeoclimatic implications; in The evolution of the East Asian Thein 2006 Neogene mammalian fauna in Myanmar; Asian environment (ed.) R O Whyte (: University of Hong Paleoprimatol. 4 143–172 Kong) vol. 2, pp 724–744 Tang Y 1980 An Early Pleistocene mammalian fauna from Yuxian, Woo J K and Wu X 1984 Hominid fossils from China an their Hebei Province and its signifi cance towards stratigraphic relation to those of neighbouring regions; in The evolution of subdivision; Vert. PalAsiatica 18 256–267 the East Asian environment (ed.) R O Whyte (Hong Kong: Tchernov E 1992Eurasian-African biotic exchanges through the University of Hong Kong) vol. 2, pp 787–795 Levantine corridor during the Neogene and Quaternary; in Wynn J G, Alemseged Z, René B, Denis G, Denné R and Diana C Mammalian migration and dispersal events in the European R 2006 Geological and palaeontological context of a Pliocene quaternary (eds) W von Koenigswald and L Werdelin L juvenile hominin at Dikika, Ethiopia; Nature (London) 443 (Senckenberg, Frankfurt Courier Forschungsinstitut) vol. 153, 332–336 pp 103–123 Zhu R X, Hoffman K A, Potts R, Deng C L, Y X, Guo B, Thapar R 2003 Early India: from the origins to AD 1300 Shi C D, Guo, Z T, Hou Y M and Huang W W 2001 Earliest (California: University of California Press) presence of humans in northeast Asia; Nature (London) 413 Thomas J V, Parkash B and Mohindra R 2002 Lithofacies and 413–417 palaeosol analysis of the Middle and Upper Siwalik Groups Zhu R X, An Z, Pots R and Hoffman K A 2003 Magnetostrati- (Plio–Pleistocene), Haripur-Kolar section, Himachal Pradesh; graphic dating of early humans in China; Earth Sci. Rev. 61 India Sediment. Geol. 150 343–366 341–359

ePublication: 26 October 2009

J. Biosci. 34(5), November 2009