Human evolution in India
729
India at the cross-roads of human evolution
R PATNAIKa,* and P CHAUHANb
a
Centre of Advanced Studies in Geology, Panjab University, Chandigarh 160 014, India
The Stone Age Institute and CRAFT Research Center (Indiana University), 1392 W Dittemore Road,
Gosport, IN 47433, USA
b
*Corresponding author (Email, [email protected])
The Indian palaeoanthropological record, although patchy at the moment, is improving rapidly with every new find.
This broad review attempts to provide an account of (a) the Late Miocene fossil apes and their gradual disappearance
due to ecological shift from forest dominated to grassland dominated ecosystem around 9–8 Ma ago, (b) the Pliocene
immigration/evolution of possible hominids and associated fauna, (c) the Pleistocene 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 orangutan-like primate
Sivapithecus. By 8 Ma, this genus disappeared from the Siwalik region as its habitat started shrinking due to increased
aridity influenced by global cooling and monsoon intensification. A contemporary and a close relative of Sivapithecus,
Gigantopithecus (Indopithecus), the largest ape that ever-lived, made its first appearance at around 9 Ma. Other smaller
primates 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 Africa to China and Indonesia in the Pliocene, monkeys
and baboons dispersed into the Indian subcontinent from Africa along with other mammals. 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. Fossils of our own ancestor and one of the first globe-trotters, early Homo 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 definite preHomo sapiens fossil hominin remains come from the Central Narmada Valley and are thought to be of Middle to late
Pleistocene 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 humans
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
According to the “Out of India” hypothesis, several
groups of modern Asian organisms had their roots in the
northwardly moving Indian plate (Bossuyt and Milinkovich
2001; Karanth 2006). One of these groups could have been
Keywords.
the early primates, the order to which, we humans belong.
In fact, recent finds of anthropoid primates from the Early
Eocene lignite mines situated in Eastern Gujarat, India
(Bajpai et al. 2008) and Oligocene of Pakistan (Marivuax
et al. 2005), may hold clues to the origin and dispersal of
our earliest ancestors. These recent finds have lent support
Genetics; human evolution; Indian subcontinent; palaeoclimate; phylogeny; primate
http://www.ias.ac.in/jbiosci
J. Biosci. 34(5), November 2009, 729–747, © Indian
Academy
Sciences 2009
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34(5),ofNovember
R Patnaik and P Chauhan
730
to some of the long-standing assertions that Asia was the
centre of anthropoid origins (Ciochon 1985; Culotta 1995;
Beard et al. 1996).
Our aim here is to broadly review the palaeoanthropological records of India (with a few examples from Pakistan)
from the Miocene onwards, which essentially begins with
the emergence and disappearance of fossil apes, followed by
the possible presence of early hominins in the late Pliocene
and Early Pleistocene in the context of changing climate
and ecology. We also briefly touch upon the Holocene
by reviewing data related to the gene flow of modern
human races into and out of India. Finally, we conclude by
evaluating the lacunae in the fossil and technological record,
and suggesting possible future directions of research to fill
these gaps in our knowledge of human evolution in India
(e.g. Chauhan 2006; Petraglia and Allchin 2007).
2. The Miocene
Though short-lived, India had its share of fame associated
with Ramapithecus, the Miocene ape from the Siwaliks.
Lewis (1934) first recognized and named Ramapithecus
brevirostris, the Rama’s ‘short-faced ape’ and placed it in the
Hominidae, the family to which all bipedal Australopithecines
and we belong. At the time of its discovery and soon after,
Ramapithecus was widely known to be the ancestor of
Australopithecus, the “ape of the South”, which in turn gave
rise to our own genus Homo. This idea of Ramapithecus being
a hominid gained general acceptance (Simons and Pilbeam
1972) and researchers started assigning Miocene specimens
from Kenya (Leakey 1962; Andrews and Walker 1979);
China (Woo and Wu 1984), Nepal (West 1984); Turkey
(Andrews 1982); Hungary (Kretzoi 1975; Fleagle 1988) and
Greece (Pilbeam et al. 1977) to the Ramapithecines. In fact,
von Koenigswald (1976) went further, by proposing that
Ramapithecus should be the right candidate to be at the root
of hominid evolution that gave rise to Australopithecus as
Ramapithecus bearing Siwaliks are located geographically
between fossil hominid yielding Africa and Southeast Asia.
However, with the discovery of key facial and postcranial
remains of Sivapithecus in the early nineteen eighties,
Ramapithecus lost its place in Hominidae (Andrews 1982;
Andrews and Cronin 1982; Pilbeam 1983). The advent of
analysing molecular clocks, which indicated ~5 Ma as the
time of divergence between humans and apes, ultimately
made this contention more convincing (Sarich and Wilson
1967). Nevertheless, Ramapithecus still appears to hold an
important place in primate evolution, as it belongs to the
pool of Late Miocene hominoids one of which gave rise to
later hominids (Ciochon and Fleagle 1985). This change in
the phylogenetic position of Ramapithecus (figure 1) has
been well illustrated by Kennedy (2003: 119). Possible
ancestors to these Middle to Late Miocene Sivapithecines
J. Biosci. 34(5), November 2009
and Ramapithecines were the African Dryopithecines,
which may have arrived in southern Asia during the Early
Miocene (Begun et al. 2003).
The Middle Miocene was the time of warm and humid
climate and evergreen to deciduous tropical forests covered
a large part of the northwestern Indian subcontinent. These
ecological conditions are very well reflected in the fossil
as well as sediment record (Ashton and Gunatilleke 1987;
Nanda and Sehgal 1993; Prasad 1993; Thomas et al. 2002).
Sivapithecus fossils have been found from the Potwar
Plateau of Pakistan in the west and from Nepal in the east
from 13.5-8.4 Ma (Barry et al. 2002; Nelson 2003; Patnaik
et al. 2005). There were at least three species of Sivapithecus
that occurred in the Siwaliks, namely Sivapithecus indicus, S.
sivalensis and S. parvada. Sivapithecus indicus ranged from
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
et al. 1985; Kappelman et al. 1991; Flynn et al. 1995; Barry
et al. 2002). S. sivalensis shows sexual dimorphism, with
males averaging around 45 kg whereas females weighed
around 20 kg. S. parvada males were as large as modern
orangutan males (Kelley 1986). Low crowned, thick
enameled molars of Sivapithecus indicate a tough diet,
such as nuts and fruits with tough rinds (Kay 1981). Dental
microwear studies of their occlusal surfaces have been
found to indicate a diet similar to those of modern fruiteating apes, with some hard-object feeding (Nelson 2003).
Kelley and Pilbeam (1986) have shown that the cranio-facial
morphology of Sivapithecus is heavily buttressed, pointing
towards an adaptation to either withstand prolonged
cyclical loading or to generate high occlusal loads. As far as
locomotion is concerned, Sivapithecus postcranials indicate
that it was more like a pronograde quadruped that walked
above the branches very similar to most of other Miocene
hominoids, but might have also been an active climber (Rose
1986). Sivapithecus does not show any specializations for
extensive terrestriality (Kelley and Pilbeam 1986). Kelley
(1997), based on enamel growth lines of a Sivapithecus
parvada juvenile, found that the life history pattern with
a prolonged growth and maturation period of Sivapithecus
was similar to that of modern great apes. Morphology of
Sivapithecus and its life history pattern, suggest that this
large-bodied frugivore must have been vulnerable to periods
of ripe fruit shortages and may have relied heavily upon
different fallback foods, such as hard seeds and nuts (Nelson
2005).
With the advent of Late Miocene global cooling and spread
of arid conditions, these forests began to shrink (Kennett and
Hodell 1986; Scott et al. 1999). In Nepal it has been found
(Hoorn et al. 2000) that from the late Middle Miocene to
early Late Miocene (~11.5–8 Ma), the Himalayan foothills
and the Gangetic floodplain were forested with subtropical to
temperate broad-leafed and tropical forest taxa, which were
Human evolution in India
Figure 1.
731
Phylogenetic placement of Ramapithecus, (A) in the 1970’s and (B) today (modified after Kennedy 2003).
then replaced by grasslands between the early to late Late
Miocene (~8–6.5 Ma). Intensification of the Asian monsoon
and disturbance of the vegetation on Himalayan slopes due
to their uplift might have facilitated such a change (Hoorn
et al. 2000). Around this time, a change in the sedimentary
regime also takes place. The large emergent river system of
Nagri Formation gave way to an inter-fan river system of
the Dhok Pathan Formation at around 10.1 Ma (Barry et
al. 2002). There are other independent observations that
strengthen this view further. In numerous deep-sea cores,
it was observed (Prell et al. 1992) that an upwelling of
endemic foraminifers and radiolarians fauna took place at
~8 Ma. As the Tibeto-Himalayan zone started rising around
12–9 Ma ago (Amano and Taira 1992; Harrison et al. 1993),
the heat budget of the region probably changed drastically
leading to the intensification of the monsoon climate system
in South Asia (Ruddiman and Kutzbach 1989; Raymo and
Ruddiman 1992; Kutzbach et al. 1993; Hay 1996; Ramstein
et al. 1997). A rather rapid uplift began at ~10 Ma and ended
at ~5 Ma (Prell and Kutzbach 1992). The diatom record
of the Indian Ocean also points towards an intensification
of regional monsoons between 11 and 7 Ma (Schrader
1974; Burkle 1989). In the terrestrial Siwalik deposits, soil
carbonate isotopic studies indicate a C3 dominant vegetation
(mainly bushes and trees) prior to 8 Ma, but by 7 Ma the C4
grasses dominated the Siwalik floodplain biomass (Quade et
al. 1989, 1995; Cerling et al. 1997). Palaeosol deposits from
various Siwalik sequences also indicate marked seasonality
in rainfall (Retallack 1991, 1995). Recently, Barry et al.
(2002) observed three very brief periods of high Siwalik
faunal turnover at 10.3, 7.8, and 7.37–7.04 Ma. Latest
Miocene faunal turnover at 7.37 and 7.04 Ma, particularly,
has been correlated with expansion of C4 grasses, the
oxygen isotope and sedimentological evidence indicating
an increasingly drier and more seasonal climate (Barry et al.
2002). A dramatic change in the diversity of muroid rodents
(from cricetid-dominated to murid-dominated) at ~9–8 Ma
has also been attributed to an intensification of the monsoons
(Patnaik 2003) (figure 2).
A combination of Late Miocene change in climate,
tectonics and sedimentary environments may have
resulted in the shrinking of forests that were occupied
by Sivapithecus, leading to their eventual disappearance
from the Siwaliks. Around this time (~9 Ma) the largest
ape that ever lived on this planet and a close relative of
Sivapithecus appears: Gigantopithecus. It co-existed with
Sivapithecus in the Late Miocene and Homo erectus and
Pongo pygmaeus in the Pleistocene (Ciochon et al. 1996).
J. Biosci. 34(5), November 2009
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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, figure 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, figure 4c). (C) Percentage of grassland (redrawn from Hoorn et al. 2000, figure 4). (D) Late Miocene to present
Phytogeography based on palaeosol data (redrawin from Retallack 1991, figure 6.12). (E) Muroid rodent diversity (data from Patnaik
2003). (F) Age ranges of Siwalik apes (Barry et al. 2002; Patnaik 2008). (Modified after Patnaik and Cameron 2007.)
For example, this genus goes on to thrive until the Late
Pleistocene (100,000 yrs BP) in China and Vietnam but not
in the Indian subcontinent. Gigantopithecus bilaspurensis/
giganteus from the late Miocene Siwaliks (Simons and
Chopra 1969; Simons and Ettel 1970; Simons and Pilbeam
1972) has recently been reassigned to Indopithecus (von
Koenigswald 1950) based on the distinct differences in
mandibular and dental morphologies between the Indian and
Chinese Gigantopithecus specimens (Cameron 2001, 2003).
However, Miller et al. (2008) argue against assigning the
Miocene form to a separate genus.
Ciochon et al. (1990) found opal phytoliths on the teeth
enamel of Gigantopithecus, which indicates a varied diet
comprising of bamboos, grasses and fruits. One of the
Gigantopithecus (Indopithecus) molar shows a large caries,
which also suggests a diet of grasses and fruits as compared
to a predominantly frugivorous diet of Sivapithecus (Patnaik
2008). Their inferred giant size (based only on fossil
mandibles) ranging from 300-150 kg and diet suggest
terrestrial locomotion. Accurately reconstructing the actual
size and shape of Gigantopithecus has been very difficult,
since it is known only by isolated teeth and a few jaws. The
J. Biosci. 34(4), October 2009
key locality for Miocene apes in India is Haritalyanagar
(figure 3) which is still yielding fossil ape specimens
beside other faunal elements (figure 4). The other important
hominoid locality is the Lower Siwalik Ramnagar locality
exposed near Jammu and has yielded several fossil ape
specimens (see Nanda and Sehgal 1993; Sehgal and Nanda
2002).
3. The Pliocene
Discovery of Sahelanthropus tchadensis from Chad (Brunet
et al. 1995) may indicate that by the late Miocene, early
hominids had already occupied areas beyond the East
African Rift valley. Dennell and Roebroeks (2005) have
argued that “If Australopithecus bahrelghazali, 2,500 km
west of the Rift Valley, implies that by 3.5 Myr ago hominids
were distributed throughout the woodland and savannah belt
from the Atlantic Ocean across the Sahel through eastern
Africa to the Cape of Good Hope” (Brunet et al. 1995), why
could they not have done the same across the grasslands of
western, southern and central Asia? By ~3 Ma ago grasslands
Human evolution in India
733
Figure 3. Geographical and chronological placement of the some of the important fossil vertebrates at Haritalyangar (modified
after Patnaik et al. 2005). (A) (1) the Dangar I locality; (2) the Gigantopithecus (Indopithecus) mandible 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 findings (modified after Pillans et al. 2005):
HTA10-S.sivalensis (Sankhyan, 1985), IG-Indopithecus giganteus (Gigantopithecus mandible site), CK-Chob Ka-nala Svalhippus site HDHari Devi- Gigantopithecus site, D1-Dangar Sivapithecus site HT-1-Sivaladapis site. Eggshells- Cf. Struthiolithus.
existed between East Africa and India (figure 5). Currently,
there is no evidence for the presence of Australopithecines in
the Siwalik Hills region. However, artifacts possibly dated to
~2.0 Ma from Riwat and to 2.0 – 1.0 Ma from the Pabbi Hills
(both in northern Pakistan) (Rendell et al. 1987; Dennell et
al. 1988) indicate that early Homo possibly had made its
way to South Asia during the Plio-Pleistocene boundary.
The northern hemisphere glaciation at ~2.5 Ma ago
brought about a major change in the global climate
(Shackleton et al. 1984), an event possibly reflected in
the faunal turnover at the Tatrot-Pinjor boundary in the
Siwaliks (Patnaik 2003). With the spread of grasslands, the
first primates to enter Siwaliks ~2.5 Ma ago were colobines
such as Presbytis sivalensis and Cercopithecines for ex.
Macaca palaeindica, Procynocephalus subhimalayanus.
Theropithecus delsoni probably entered a bit later at around
1 Ma (Delson 1993). Theropithecus, which has invariably
been globally associated with early Homo is also known
from Plio-Pleistocene deposits of south, east and north
Africa, Israel, Spain, Italy (Delson 1993; Delson et al.
1993; Gibert et al.1995; Belmaker 2002; Rook et al. 2004).
The sabre-tooth felid Megantereon and the large hyena
Pachycrocuta which occur in the Pinjor Formation may
also have used this corridor to disperse from Europe during
the Early Pleistocene (Turner 1992; Rook et al. 2004).
Beside large mammals, small mammals such as murine
rodents are common in Late Pliocene deposits of Ethiopia,
Kenya, Tanzania and Indo-Pakistan (Patnaik 2000, 2001;
Wynn et al. 2006) and may indicate the absence of any
physical barrier between the Indian subcontinent and East
Africa to prevent faunal migrations. The large-mammal
Plio-Pleistocene taxa from East African such as Oryx,
Hippopotamus and Crocuta crocuta (Tchernov 1992) have
also been recorded in the Siwaliks. Siwalik mammals such
as Equus, gazelle and Hippopotamus have also been found
at the North African Early Pleistocene hominid site of Ain
Hanech in Algeria (see Dennell 2003 for a review). It has
been observed that the Upper Irrawaddy fauna of Myanmar
R Patnaik and P Chauhan
734
Figure 4. Select fossil mammals from the hominid interval at Haritalyangar (modified 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 skull (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
dissimilis, Hexaprotodon palaeindicus, Cervus sp., Hemibos
triquetricornis, Rhinoceras sivalensis, Stegodon insignis
and Elephas hysudricus, which have also been recorded
in the Pinjors (Takai et al. 2006). The faunal remains from
Xiaochangliang, China, like Cervus and Gazella (Zhu et al.
2001) are also found in the Pinjor deposits of India. Other
common forms occurring in China as well as in Pinjors are
Pachycrocuta brevirostris, Equus, Rhinoceras, Stegodon,
Coelodonta, Potamochoerus and Sus (Colbert 1940; Tang
1980; Han 1987; He 1997; Zhu et al 2003). The Pliocene
monkey from China, Procynocephalus wimani (Schlosser
1924), has been found to be very similar to Procynocephalus
subhimalayanus of Pinjor (Verma 1969). It appears that the
Pliocene conditions in India during the Pinjor times were
conducive to possibly allow dispersion of early hominin
through the region.
4. The Pleistocene
The re-dating of Javan H. erectus by Swisher et al. (1994)
to ~1.8 Ma hints that the Indian subcontinent may have
been utilized as a geographic corridor between East
Africa and Southeast Asia. Unfortunately, the currently
J. Biosci. 34(5), November 2009
known archaeological evidence for such an early dispersal
into South Asia is ambiguous and not clearly established
(discussed below). This reflects major palaeontological and
archaeological gaps prior to the Middle Pleistocene, some
of which may be due to the lack of absolute dates for known
artifact assemblages and possibly due to the discontinuous
occupation of this region prior to the early Middle
Pleistocene (Dennell 2003). The only-known pre-modern
hominin fossil in the subcontinent may be contemporary
with the Late Acheulean or early Middle Palaeolithic
phase(s) and has been recovered from Hathnora (figure 6) in
the central Narmada Valley (Sonakia 1984; Kennedy 2001).
This partial calvarium (figure 6 inset) was supplemented by
possibly-associated clavicles and a rib fragment (Sankhyan
1997a,b; 2005). The calvarium (possibly that of a female)
was originally identified as an ‘advanced’ Homo erectus (de
Lumley and Sonakia 1985) and later re-classified as an archaic
or early form of H. sapiens (Kennedy et al. 1991). A detailed
phylogenetic analysis of the Narmada calvarium reveals that
it falls between the Steinheim (H. heidelbergensis) and H.
neanderthalensis (figure 7), which in turn strongly supports
an European connection for the Narmada hominin as
opposed to a strictly Asian origin (Cameron et al. 2004). The
Narmada hominin calvarium has recently been classified as
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
Stegodon namadicus, Equus namadicus, Hippopotamus
namadicus, Sus namadicus, and Cuon alpinus have been
found to be stratigraphically associated with the calvarium
from Hathnora. This mammalian assemblage, belonging to
the Surajkund Formation, broadly suggests a warm climate
with intermittent arid phases in the late middle Pleistocene.
In general in the Narmada Valley, it has been found that the
early to middle Pleistocene vertebrate assemblages, including
cyprinid fishes, crocodiles, Hippopotamus palaeindicus,
and Elephas hysudricus suggest a warm climate, whereas
presence of wild ass (Equus hemionus khur) and ostrich
(Struthio camelus) may indicate drier conditions in the
terminal part of the upper Pleistocene (Patnaik et al. 2009).
Although Pleistocene vertebrate faunal assemblages have
been recovered from throughout the subcontinent, including
the Siwalik Hills, central and peninsular India and Sri Lanka
(often in stratigraphic or spatial association with stone
tools), no convincing evidence of butchery has been clearly
demonstrated (Chauhan 2008a). Nonetheless, this rich
vertebrate palaeontological evidence (Badam 2002) points to
diverse ecological environments throughout the entire region,
and which must have collectively affected early hominin
ecological adaptations, subsistence strategies and seasonal
dispersal patterns within India. Indeed, the South Asian
faunal evidence offers valuable opportunities for comparative
taxonomic studies, isotope analyses, palaeoenvironmental
reconstructions, biochronological correlations and interregional faunal migration links (with West Asia and Southeast
Asia, respectively).
4.1
The archaeological evidence
In contrast to the meager hominin fossil record, the stone
tool record of the Indian subcontinent is overwhelming and
lithic assemblages have been recovered in diverse contexts
from various ecological regions (Sankalia 1974). Although a
robust chronological framework is lacking for most of this
evidence, a complete sequence of Palaeolithic occupation
from at least the early Middle Pleistocene has been clearly
established and separated into the traditional triparate Lower,
Middle and Upper Palaeolithic phases (for latest respective
J. Biosci. 34(5), November 2009
R Patnaik and P Chauhan
736
Figure 6.
Narmada hominin locality, Hathnora and the hominin cranium shown in inset (courtesy Arun Sonakia).
reviews, see Settar and Korisettar 2002). Although broad
‘transitions’ between these techno-chronological phases
have been presumed based on evidence from other regions
of the Old World, a clear understanding of the precise nature
of the South Asian cultural shifts remains to be properly
worked out. This current drawback is collectively due to
the dearth of absolute dates, few excavations of extensive
cultural sequences and prominent sterile horizons at
culturally-continuous sites (Chauhan 2009d). In any case,
it is clear that the South Asian Palaeolithic record preserves
typo-technological attributes shared by other regions (e.g.
Acheulean) as well as assemblages resulting from indigenous
mechanisms of cultural evolution, particularly from the early
Middle Palaeolithic onwards (Petraglia 2008).
The South Asian Lower Palaeolithic has been
traditionally divided into core-and-flake and Acheulean
lithic industries that occur independently as well as in
shared geographic and geomorphologic contexts (Jayaswal
1982; Petraglia 1998; Gaillard and Mishra 2001; Chauhan
2009a). These assemblages are frequently found in
J. Biosci. 34(5), November 2009
stratified or surface association with fine-grained fluvial
and lacustrine sediments, ferricretes, laterites, and gravel
or conglomerate deposits. Most of the Indian localities have
been directly dated through the Uranium-Thorium (234Th230
U) and thermoluminescence (TL) methods and include
a predominance of Acheulean sites (Mishra 1995). Ages
for other occurrences such as Riwat, Dina, Jalapur, Pabbi
Hills, Morgaon, and Satpati Hill have been estimated using
palaeomagnetism and geostratigraphic correlations (Chauhan
2009a). At Teggihalli, Chirki-Nevasa, and Yedurwadi, the
234
Th-230U ages for the Acheulean extend beyond 350 Ka (or
390 at Didwana), the maximum limit of the dating methods,
an assessment partly supported by lithic typology. With the
possible exceptions of the Satpati Hill site in Nepal and
Morgaon and Chirki-on-Pravara in Maharashtra, there is
no unequivocal evidence of Acheulean occupation prior to
the Middle Pleistocene in the subcontinent. Though the site
of Isampur in the Hunsgi Valley has been dated to c. 1.27
Ma using electron spin resonance (ESR) on herbivore teeth
associated with the cultural horizons (Paddayya et al. 2002),
Human evolution in India
Figure 7.
2004).
737
Cladogram showing place of the Narmada hominin in the broad phylogeny of the Pleistocene hominins (after Cameron et al.
this estimate is preliminary and requires corroboration. In
sum, the South Asian Acheulean evidence chronologically
ranges from the early Middle Pleistocene to the early Late
Pleistocene.
One of the first claims of pre-Acheulean evidence in the
subcontinent was made through the Soanian industry by
de Terra and Patterson (1939). Later work by the British
Archaeological Mission to Pakistan (BAMP) resulted in a
major revision of de Terra and Paterson’s interpretations.
Subsequently, multiple lines of evidence, including a
comparison of Soanian and Acheulean technology, landscape
geoarcheology, surveys of dated geological features, and a
comparative morphometric analysis, clearly revealed that
the majority of Soanian assemblages, if not all, comprise
Levallois elements and probably postdate the Acheulean
(Chauhan 2009a). Subsequent claims for a pre-Acheulean
occupation have come from central India and the Siwalik
Hills in Pakistan and India (Chauhan 2009b, c). The most
systematically studied of all these pre-Acheulean claims
come from Riwat and the Pabbi Hills in the Siwalik deposits
of northern Pakistan (Dennell et al. 1988), and is also the
most controversial because of contextual and artifact-
abundance issues. Beyond this evidence, no unequivocal
evidence of pre-Acheulean assemblages is known from the
entire subcontinent. Systematic surveys in the Siwalik Hills
of northern India and other regions in central and peninsular
India may eventually yield Oldowan assemblages - but only
if the region was a corridor for early Homo from East Africa
to Southeast Asia (Chauhan 2009b). Given the abundance
of stone raw material sources, diverse faunal populations,
a lengthy monsoon regime and conducive environmental
conditions during the South Asian Plio-Pleistocene, it is
likely that the region witnessed pre-Acheulean hominin
occupations.
The Acheulean evidence is much more prominent and
better studied: with the exception of northeast India and
parts of Konkan Maharashtra, western Kerala, south of the
Cauvery River in Tamil Nadu and Sri Lanka, Acheulean
assemblages are found throughout most of the Indian
subcontinent (Misra 1989; Pappu 2001a, b; Petraglia
2006). The South Asian Acheulean is generally divided
into Early or Late developmental phases, based primarily
on typo-technological features, assemblage compositions,
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
continuous from the early Middle Pleistocene and comprises
a rich and diverse array of technological, structural and
symbolic evidence. As more absolute dates and detailed
metrical data become available, current classifications of
many assemblages are likely to change. While the term
‘Middle Acheulean’ has been occasionally applied to
‘transitional’ assemblages, such a facies have never been
systematically justified. Early Acheulean assemblages
are known to comprise handaxes (figure 8), choppers,
polyhedrons, and spheroids, usually a lower number of
cleavers (but not always) and flake tools, the predominant
use of the stone-hammer technique, and a marked absence of
the Levallois technique (Misra 1987). The Early Acheulean
bifaces are often asymmetrical, large with thick butts or midsections and possess large, bold and irregular flake 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
indicative of hard-hammer percussion. In contrast, Late
Acheulean assemblages are defined by the low proportion
of bifaces, the high ratio of cleavers to hand axes, the very
high ratio of flake tools such as scrapers, and the extensive
employment of the soft-hammer technique and the Levallois
and discoid-core techniques (Misra 1987). These bifaces
are also generally smaller, thinner, and morphologically
more refined, with a significant increase in the degree of
retouching and controlled bifacial thinning/flaking (figure
8). Early Acheulean characteristics have been identified at
such sites as Chirki-Nevasa, Morgaon, Pilikarar, Singi Talav
and Satpati Hill while Late Acheulean characters have been
recognized at Attirampakkam, Bhimbetka, Raisen District,
Hunsgi-Baichbal Valleys, the Kaldgi Basin and Gadari
(among many others) (Pappu 2001a, b; Petraglia 2006).
The gradual shift in characterization from ‘‘Middle
Stone Age’’ to the ‘‘Middle Palaeolithic’’ for the South
Asian evidence is thought to be due to the Mousterian and
Levallois affinities between assemblages in the northwestern
region of the subcontinent and other penecontemporaneous
occurrences in Central Asia, northern Africa, and Europe
(Kennedy 2003). Separating the Middle Palaeolithic horizons
from the Late Acheulian ones, however, has proved to be a
recurrent methodological problem (Mishra 1995) because
the Levallois technique and other forms of prepared-core
technology are also present in the Late Acheulean phase
of the subcontinent. Additionally, the Middle Palaeolithic
sites often overlap geographically with the Late Acheulean
occurrences and indicate successful adaptations and
exploitation of a range of ecological and topographic
settings. In comparison with the South Asian Acheulean,
the four features that distinguish Middle Palaeolithic
assemblages are: (i) a decrease in size of the artifacts, (ii)
a noticeable shift from large Acheulian bifaces to more
smaller, specialized tools (iii) an increase in the preparedcore technique, and (iv) a preference for fine-grained raw
material (such as quartz, fine-grained quartzite, chert,
jasper, chalcedony, flint, agate, crypto-crystalline silica,
lydianite and bloodstone (Kennedy 2003). In some regions
such as Rajasthan, parts of Andhra Pradesh, parts of coastal
Maharashtra, and the Narmada Valley, quartzite continues
to be used. Some of the new types that either first appear or
become prominent in the South Asian Middle Palaeolithic
are prepared-cores, discoids, flakes, flake-scrapers, borers,
awls, blades, and points. Since choppers and diminutive
handaxes are often found in certain Middle Palaeolithic
contexts, it may be suitable to arbitrarily divide South Asian
Middle Palaeolithic assemblages into two separate groups:
light-duty assemblages and heavy-duty assemblages. The
factors for such variation in assemblage composition may
include function, raw material variability, ecology, culture,
style, and/or natural post-depositional formation processes.
The Soanian evidence is also now increasingly classified as
739
a Middle Palaeolithic tradition with a prominent heavy-duty
tool component instead of representing a Mode 1 technology
thought to precede the Acheulean (see Chauhan 2007; 2008a,
b for reviews, research results and related citations).
The super volcanic eruption of Toba (northern Sumatra)
~74 ka ago led to the deposition of a blanket of volcanic ash
all over India (William and Royce 1982; Rose and Chesner
1987, 1990). Terrestrial deposits of this Tephra have been
recorded in several river Valleys throughout India (Acharyya
and Basu 1993; Shane et al. 1995; Westgate et al. 1998;
Jones 2007; Raj 2008). Researchers have referred this ~74
ka eruption to be a ‘super eruption’ with a DRE (Dense Rock
Equivalent) estimate of ~2800 km3 (Knight et al. 1986; Rose
and Chesner 1987, 1990; Rampino 2002; Mason et al. 2004)
and ~800 km3 of ash (Viseras and Fernandez 1995). This huge
eruption has been estimated to be much larger in magnitude
compared to those of very recent times such as1815 Tambora
and 1991 Pinatubo eruptions (Stothers 1984; McCormick et
al. 1995). It has been proposed (Rampino et al. 1988) that
the after affects of Toba Supervolcano can be equated with
a nuclear winter as modeled by Turco et al. (1983; 1990).
Further, the volcanic winter could have decreased the land
temperatures between 30-700 Latitude by ~5-150C (Rampino
and Self 1992). This super eruption has been found to
coincide with a major shift in global climatic conditions
from interglacial marine isotopic stage (MIS) 5 to glacial
MIS 4 (Ninkovich et al. 1978; Kale et al. 2004). Regarded
as the largest volcanic eruption in the last two million years,
this eruption must have caused significant change in the
global climate leading to widespread biotic extinctions and a
severe reduction in the human population (Ninkovich et al.
1978; Ambrose 1998). Although a recent study claims that
early humans survived this devastating event (Petraglia et
al. 2007), only long-term and large-scale projects at multiple
locations in India will reveal the extent of the YTT impact(s)
on human populations
The South Asian Upper Palaeolithic is not as clearly
defined (James and Petraglia 2005) as the regional
Acheulean or the South Asian Middle Palaeolithic nor
well understood; as a result, it still requires extensive
multidisciplinary research at a large scale (Chauhan 2009d).
The dominating and defining features of South Asian Upper
Palaeolithic assemblage compositions include a notable
increase in the production of more specialized tools such
as blades, burins, and borers. Although the production of
blades is known from Late Acheulean levels at a few sites
(e.g. Bhimbetka), this behavior became highly prominent,
prolific, and technologically consistent and standardized
only during the Upper Palaeolithic. Additional tool types
during this technochronological period include flakes,
knives, awls, scrapers, cores including cylindrical types,
choppers, and bone tools. The techniques of making many
of these lithic and nonlithic tools also changed from the
J. Biosci. 34(5), November 2009
R Patnaik and P Chauhan
740
preceding technochronological phases. For example, the use
of pressure flaking and the soft hammer technique for flake
detachment appears to increase significantly as compared
with the South Asian Middle Palaeolithic and the later
Acheulean. The degree or intensity of retouch also appears
to increase considerably compared with Lower and Middle
Palaeolithic assemblages in general. Compared with earlier
Palaeolithic technology, the South Asian Upper Palaeolithic
shows a greater degree of regional typo-technological
variation as well as an increase in different types of scrapers
Figure 9.
(e.g., steep, convex, convergent) and backed blades (Misra
2001). Figure 9 illustrates the key Palaeolithic localities in
the Indian Subcontinent, some of which have been discussed
briefly in this paper.
In the terminal Pleistocene cool and dry conditions
corresponding to the Last Glacial Maxima (LGM) prevailed
in the Indian subcontinent. Conditions between ~18,000 to
~13,000 yrs. have been referred to as hyper-arid (Pant 2003).
When compared to present day climate, such dry conditions
were probably due to low precipitation of summer monsoon
Locations of key Paleolithic occurrences in the Indian Subcontinent.
J. Biosci. 34(5), November 2009
Human evolution in India
and a higher winter precipitation (Singh et al. 1990). Nilgiri
Hills of southern India also experienced LGM climate
(Sukumar et al. 1993). Under such conditions predominance
of tropical grass type vegetation during 20-16 ka clearly
indicates a very arid phase as this type of vegetation grows
favourably under low aridity and low soil moisture. This
also points to a period of weak southwest summer monsoon
during LGM. It has been noticed that the change in climate
and vegetation has adversely influenced the structure and
composition of the montane ecosystem (Sukumar et al.
1995). Similar conditions of LGM have also been identified
in the Central Narmada Valley (Patnaik et al. 2009).
5. The Holocene
The terminal Pleistocene cool and dry conditions of LGM
were followed by warm and humid conditions of early
Holocene time. Around ~7–10 ka the northwestern part
of India was very warm and wet (Goodbred and Kuhel
2000; Fleitmann et al. 2003), which would have facilitated
the hunter-gatherers to domesticate plants and animals,
followed by the start of agriculture (Gupta 2004). Eventually,
agriculture led to the permanent settlement of populations.
Invasion of Aryans into India is still a hotly debated issue.
It has been hypothesized that Harappan settlers (6000-2000
BC) were basically the modern humans that came from
Africa around 60,000 yrs ago, and were different from
the Aryans that arrived via Europe after the LGM (Vahia
2004). The main difference between these two populations
hinges on the idea that the Harappan sites lack any evidence
of ‘Horses’, whereas, they were integral part of ritualistic
customs mentioned in the Rigvedic literature supposed to
have been written between 4000-2000 BC (Thapar 2003).
5.1
Changing human populations in South Asia: the
genetic evidence
In recent years, there has been increasing interest in the
genetic background of the human population diversity of the
Indian subcontinent. In fact, physical differences in South
Asian populations have been a subject of study since the
early British occupation of India. More recently, a number
of studies have been conducted to explain the biological
diversity as well as subsequent historical developments
of caste and regional group origins (Bamshad et al. 2001;
Cordaux et al. 2004). Though not generally utilized to
understand hominin dispersal patterns, such studies have
important implications for understanding the dispersal of
archaic Homo (Quintana-Murci et al. 1999) into South Asia
rather than for older Homo species. For example, Majumder
(2001) has presented evidence that mitochondrial DNA
haplotypes (based on RFLPs) are remarkably similar across
741
ethnic populations of India and that some of the earliest
continuous occupiers of the region may have been AustroAsiatic tribal populations although the Dravidians have also
been ancestral contenders. Modern tribal groups (461) make
up over 8% of the Indian population (Census of India 1991)
and the majority is separated geographically, linguistically
and culturally and many groups, up to varying levels, have
retained traditional or primitive modes of subsistence;
though not as extensive as some other regions in the Old
World. For example, the long-term geographical isolation
of the Andaman Islanders is significant and mitochondrial
sequences obtained from museum specimens revealed that
some of their physical attributes converge with African
pygmoid groups representing a southern movement from
Africa (Endicott et al. 2003); although Indians are genetically
classed as Caucasoid (Kivisild 2000). It should also be noted
that some of these early tribal populations have retained some
of their genotype by not mixing with later groups migrating
into the subcontinent and at the same time, Indian tribal
and caste populations derive predominantly from a shared
gene source in southern and western Asia (Kivisild et al.
2003). Studying the variation in their DNA will help explain
biological affinities as well as gene flow between peninsular
Indian populations and Southeast Asian populations. For
example, it has been demonstrated by Reddy and Stoneking
(1999) that Australian Aboriginals are very distinct and
closely resembles certain Indian groups (also see Kumar
and Reddy 2003). Such affinities or independent origins of
Indian populations has been studied by understanding the
9-bp depletion between COII/tRNA and comparing it with
African and eastern Asian populations: Koya and Chenchu
tribal groups differ by seven HVS-I mutations, inheriting a
subset of African mtDNA lineages (see Kivisild et al. 2003
for explanations). Furthermore, the frequency of haplogroup
M and its diversity are highest in India (Edwin et al. 2002)
suggesting that the subcontinent was settled soon following
the archaic Homo dispersal from Africa, neither showing
complete extinction nor a replacement of the initial hominin
groups (Kivisilid et al. 2003). Though the archaeological
record is rich and continuous, such vital information as
radiometric dates, hominin fossils, and high-resolution
palaeoenvironmental data is currently inadequate to answer
questions regarding dispersals and population movements
within South Asia and its periphery zones.
Recently Eswaran (2002) and Eswaran et al. (2005)
make an important argument about gene flow rather
than direct population movement outside of Africa.
However, it is currently difficult to test such inferences
from the archaeological record and the fossil record is too
fragmentary in key regions (western China, India, Russia,
etc.) to reveal any patterns of gene flow at such an early
stage in hominin dispersals. In fact, hominin groups appear
to have adapted to different palaeoenvironments at regional
J. Biosci. 34(5), November 2009
R Patnaik and P Chauhan
742
levels over a long period of time and in relative isolation. For
example, experimental studies involving physical exercises
between tropical (Malaya and Indian) and temperate
(Chinese) subjects were conducted by Duncan and Horvath
(1988), which showed differential adaptation to varying heat
stress levels. Another phenotypic example is shovel-shaped
incisors of extant East Asian populations also found in the
fossil hominin record from the same region. The lack of
this feature in South Asians is an example of evolution in
geographical isolation, following a certain demographic
saturation point. In other words, there may have been
relatively minimal gene flow between peninsular Indian
and East/Southeast Asian groups during earlier Pleistocene
times. This is also tentatively supported by the work of
Bamshad et al. (2001) who demonstrate, through mtDNA
and y-chromosome variation, that there was minimal or
marginal movement in the northeastern corridor of the
subcontinent. Additional evidence collected by Kivisild et
al. (2000) led them to consider India as a part of the source
gene pool ancestral to maternal lineages in Europe. Most
recently, a new genetics study (Reich et al. 2009) reveals that
northern Indian and southern Indian populations come from
geographically-separate ancestral lineages which disporsed
into the subcontinent at different times in the past.
for ancestors of Miocene hominoids in the Oligocene
deposits; (3) recovering potential hominid ancestors among
the Miocene hominoids; (4) what did Gigantopithecus
exactly look like?; (5) whether hominids were present in
the Pliocene?; (6) Who were the Early Pleistocene hominins
(if they were present in the region at that time)?; and (7)
evidence concerning the arrival/evolution of modern human
groups into the region. Current and future multidisciplinary
research projects within the Indian subcontinent are bound to
reveal exciting new information and changing interpretations
regarding the global role and impact of the Indian landmass
on primate and human evolution.
Acknowledgements
We would like to thank Profs. Sunil Bajpai and Ashok Sahni
for inviting us to contribute to this volume. We are grateful to
Claire Gaillard for offering valuable and helpful comments
on an earlier draft of this paper. RP thanks Wenner-Gren
Foundation (Grant No. Gr.ICRG-43) and Department of
Science and Technology, New Delhi for the financial support
(SR/S4/ES-171/2005). We would like to thank National
Geographic Society (Grant 7386-02 to RP) and Wenner-Gren
Foundation for Anthropological Research (Grant 7541 to PC)
for funding the Narmada Basin Palaeoanthropology Project.
6. Discussion and conclusions
There is every possibility that the collision of India with
Asian landmass in the early Eocene led to the initial
dispersal of anthropoid primates into Eurasia. Beside the
records from Eocene and Oligocene, there is almost no
record of anthropoid primates until ~13 Ma ago. The Late
Miocene ape Sivapithecus was primarily arboreal, but
some terrestrial locomotion by this ape cannot be ruled
out. Gigantopithecus was definitely a terrestrial ape that
lived on grasses, fruits and possibly bamboos. Again there
exists a lengthy gap until the Middle Pleistocene, where we
have definite Acheulean artifacts produced most probably
by H. erectus or H. heidelbergensis populations. Then,
there is the only definite hominin fossil, the Narmada
hominin which has been assigned to archaic H. sapiens/H.
erectus H. heidelbergensis/Homo sp. by various specialists.
Genetic evidence demonstrates that the gene pool to which
Indian populations belong is highly diverse and comprises
a large number of tribal groups, many of which are linked
at different levels with both European and Asian lineages.
Though, the palaeoanthropological data so far appears to
be discontinuous, India with appropriate deposits ranging
in age from the Palaeocene to the Pleistocene and covering
a vast geographic area, offers great potential for further
palaeoanthropological exploration and investigations. The
critical areas of research include: (1) searching for the earliest
anthropoids in Palaeocene and Eocene deposits; (2) searching
J. Biosci. 34(5), November 2009
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