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Ostrich expansion into India during the Late Pleistocene: Implications forcontinental dispersal corridors

James Blinkhorn, Hema Achyuthan, Michael D. Petraglia

PII: S0031-0182(14)00537-9DOI: doi: 10.1016/j.palaeo.2014.10.026Reference: PALAEO 7063

To appear in: Palaeogeography, Palaeoclimatology, Palaeoecology

Received date: 2 July 2014Revised date: 14 October 2014Accepted date: 21 October 2014

Please cite this article as: Blinkhorn, James, Achyuthan, Hema, Petraglia, Michael D.,Ostrich expansion into India during the Late Pleistocene: Implications for continen-tal dispersal corridors, Palaeogeography, Palaeoclimatology, Palaeoecology (2014), doi:10.1016/j.palaeo.2014.10.026

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Ostrich expansion into India during the Late Pleistocene: Implications for continental dispersal

corridors

Blinkhorn, James1, Achyuthan, Hema

2, Petraglia, Michael D.

3

1UMR5199 PACEA, Université de Bordeaux, Avenues de Facultes, Cedex Pessac, France

2Department of Geology, Anna University, Chennai 600 025, India

3School of Archaeology, Research Laboratory for Archaeology and the History of Art, University of

Oxford, Oxford OX1 3QY, United Kingdom

Correspondance: J. Blinkhorn, UMR5199 PACEA, Université de Bordeaux, Avenues de Facultes,

Cedex Pessac, France; Email: j.blinkhorn@pacea.u-bordeaux1.fr Phone : 00 33 5 40 00 25 45

ABSTRACT

New evidence is presented for the earliest occurrence of ostrich (Struthio sp.) in India during the Late

Pleistocene along with a synthesis on the evidence for ostrich populations in the subcontinent. Direct

dating of ostrich eggshell using Accelerator Mass Spectrometry (AMS) radiocarbon methods on

excavated samples from Katoati, Rajasthan, India, are supported by Optically Stimulated

Luminesence (OSL) dating of associated sediments to demonstrate the arrival of ostrich in India

before 60 thousand years ago (ka). In addition, the first stable isotope studies on ostrich eggshell from

India have been conducted, yielding a new form of palaeoenvironmental proxy data for the Late

Pleistocene. The geographic expansion of ostrich into India corresponds with the distribution of

Sahel-like environments, bordering but not substantially colonising endemic Indian vegetation zones.

The dispersal of ostrich into India marks a rare introduction of megafauna into the subcontinent

during the Late Pleistocene, the longevity of which spans a period more than 40 ka. The timespan and

range of this colonisation indicates the availability and exploitation of suitable habitats in India. The

continental dispersal of ostrich into India during the Late Pleistocene offers useful insights into the

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debate surrounding the dispersal of modern humans, and contrasts with the hypothesized coastal

movement of Homo sapiens into India.

Keywords: Dispersal; India; Katoati; Late Pleistocene; stable isotope; Struthio

Highlights

Oldest directly dated evidence for Late Pleistocene expansion of Struthio into India

First stable isotope study of Late Pleistocene ostrich eggshell in India

Synthesis of existing evidence for Struthio populations in South Asia

Struthio expansion associated with extension of Sahel-like habitats

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1. Introduction

Broad continuity in the presence of megafauna can be observed in the Indian subcontinent

throughout the Middle Pleistocene and extending into the Late Pleistocene (Roberts et al., 2014). The

Pakistani Siwalik formations yield repeated evidence for the presence of Struthiodiae (family of

flightless ratite birds), spanning the Middle Miocene to Middle Pleistocene (Dennell, 2004; Patnaik et

al., 2009b; Stern et al., 1994). However, a significant change in the distribution of Struthiodiae is

evident in the Late Pleistocene, becoming absent in the Himalayan forelands and instead appearing in

western and central India for the first time (Andrews, 1911; Badam, 2005; Bidwell, 1910). The timing

of this significant range expansion is poorly constrained and has yet to be set within the broader

context of Late Pleistocene faunal dynamics and their relationship with palaeoenvironmental changes.

The only osteological remains of Struthiodiae from South Asia were collected from unspecified

Siwalik strata and described as a new species, Struthio asiaticus, by Milne-Edwards (1871) on the

basis of size differences with the extant African ostrich, S. camelus. A metric study of these

specimens noted that despite presenting a more robust neck, the Siwalik specimens were comparable

in form and size with a large male specimen of S. camelus (Davies, 1880). Further description by

Lydekker (1884) suggested that the stouter neck of the Siwalik specimen should be considered as

within the realm of individual variation, rather than at a species level, thus suggesting S. asiaticus be

treated as a ‘preliminary’ taxon. Since this time, a number of osteological remains and, more

problematically, eggshell specimens, have been attributed to S. asiaticus spanning from the Pliocene

to the Late Pleistocene and ranging from South Africa to East Asia (Kurochkin et al., 2010; Manegold

et al., 2013; Mikhailov, 1991; Mourer-Chauviré and Geraads, 2008), making it the most widespread

species in the genus Struthio. Osteological studies of specimens attributed to S. asiaticus are noted to

be between 20-50% more robust than contemporary African ostrich, although not necessarily taller

(Mourer-Chauvire and Geraads 2008).

In contrast to the paucity of osteological remains, ostrich eggshells (OES) have been recovered

from a number of contexts in the Siwalik Range (Dennell, 2005; Stern et al., 1994) and in peninsula

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India (Badam 2005). Ratite eggshells with aepyornithoid-type pore patterns are reported from the

mid-Miocene Dhok Pathan formations near Hasnot (Patnaik et al., 2009b; Sauer, 1972), and a range

of sites in the Siwalik formations dating between 11.35-1.25 millon years ago (Ma.) (Stern et al.,

1994). Eggshells with struthionid-type pore patterns are found in younger Siwalik deposits ranging in

age from 2.24–0.5 Ma. (Stern et al., 1994) and at a number of sites in the Indian states of Rajasthan,

Uttar Pradesh, Madhya Pradesh and Maharashtra dating between >43–18 thousand years ago (ka)

(Badam 2005). More detailed description of eggshell morphology and putative taxonomic affiliations

of these OES samples have been restricted to the Late Pleistocene samples (Sahni et al., 1990). The

earliest samples examined, from Ken River (Uttar Pradesh), were considered as “practically

indistinguishable” from egg-shell of modern Somali ostrich (S. camelus molydophanes)(Andrews

1911; Bidwell 1910), though noted as thicker than contemporary examples, with more recent studies

reaching the same conclusions based upon wider sampling from peninsula India (Sahni et al., 1990).

The extinction of Late Pleistocene Struthio sp. from India has been briefly discussed (e.g. Badam

2005) yet the timing and potential causes for their expansion into western and central India has not be

addressed. Pertinent to this are patterns of climatic amelioration during the Late Pleistocene that may

have permitted the extension of favourable habitats for ostrich from their endemic regions into India.

Here, evidence is presented for the earliest dated Late Pleistocene Struthio samples from western

India, which is placed in its palaeoenvironmental context along with all known samples of

Struthiodiae from South Asia. Factors that may have affected this Late Pleistocene expansion are

discussed

2. Materials and Methods

Ostrich eggshell has been recovered from both surface and excavated contexts at Katoati,

Rajasthan, India. OES fragments have been recovered from six sites, labelled KAT1–5 and JFL2. The

excavation of a 4.5 m trench at KAT1 yielded two pieces of OES. The two pieces were recovered in

10mm screens from sediment excavated at a depth of 3.4 m from surface in horizon S4 (Fig.

1)(Blinkhorn et al., 2013). At KAT2 a cluster of OES was observed eroding from an exposed

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sediment sequence below the contact between two distinct geological units, homologous with S2 and

S3 at KAT1. Shallow excavation (ca 10 cm) identified three discrete clusters of OES fragments

embedded in the lower sediment deposits (S3) (Fig. 1), overlying a thin, laterally extensive pebble

horizon. Further exploration at exposures of this pebble horizon resulted in the recovery of three OES

samples from S3 deposits at sites KAT3, KAT4 and KAT5. Inspection of an exposed sediment

sequence yielded a single piece of OES from site JFL2, which was recovered from an aeolian deposit

between a thin, upper and thick, lower gravel horizon (Blinkhorn et al., 2013). Individual fragment

thickness of OES from all sites were measured using digital calipers.

Fig. 1: Excavated sediment sequences at KAT1 (left) and KAT2 (right; not to scale) illustrating the

sediment context for horizons bearing OES and associated OSL dates.

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A total of nine OES samples were submitted and processed for AMS radiocarbon dating.

Eight samples were processed in the Oxford Radiocarbon Accelerator Unit (ORAU). These samples

were sandblasted in aluminium oxide powder, etched in ~0.2 M HCl for 2 minutes, rinsed in distilled

water and dried prior to acid hydrolysis for producing CO2 for graphitisation for AMS dating and

mass spectrometer analysis (see Bronk-Ramsey et al., 2002, 2004a, 2004b for further details). One of

four samples recovered from KAT2 was analysed by Prof. Jay Quade, University of Arizona, Tucson.

Additional dating has been undertaken at KAT2 using Optically Stimulated Luminescence (OSL).

OSL samples were recovered from sediment deposits ca 5 cm above and below the excavated OES

deposits at KAT2, relating to units S2 (above) and S3 (below) from KAT1, and submitted to the

Wadia Institute of Himalayan Geology for analysis (see Supplementary Information for

methodology).

Mass spectrometer analysis of 12 OES samples (including seven samples analysed by ORAU)

was undertaken without physical or chemical abrasion of the surface. Subsamples of OES pieces were

selected and rinsed in ethanol to remove any adhering sediments prior to crushing in an agate pestle

and mortar and drying at 40°C. Oxygen and carbon stable isotopic results were obtained using a VG

Isogas Prism II mass spectrometer with an on-line VG Isocarb common acid bath preparation system.

Each sample was reacted with purified phosphoric acid (H3PO4) at 90°C with the liberated carbon

dioxide cryogenically distilled prior to admission to the mass spectrometer. Both oxygen and carbon

isotopic ratios are reported relative to the VPDB international standard. Calibration was against the in-

house NOCZ Carrara Marble standard with a reproducibility of better than 0.2%. Stable isotope ratios

are expressed using the notation as difference in parts per thousand (permil, ‰) relative to the

standard, calculated as δ‰([Rsample/Rstandard]-1)x1000, where R = 13C/12C or 18O/16O.

3. Results

3.1 Ostrich Eggshell Thickness and Form

Thirty-six measurements of OES thickness indicate a mean thickness of 2.27 mm, with a

range between 2.13 and 2.38 mm. This range matches closely with the reported thicknesses of

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samples from the Indian peninsula, including from Ken River (2.4 mm) (Andrews 1910) and from the

sites of Ledkheri, Chandresal and Anjar (2.1 to 2.2 mm) (Sahni et al., 1990). Collectively, these

samples are all notably larger than a single measurement of 1.7 mm from a sample from Pakistan

(Grellet-Tinner, 2006), but substantially smaller than aepyornithoid-type OES from the Dhok Pathan

formation at Hasnot with a mean of 2.76 mm (range=2.5 to 2.9 mm)(Patnaik et al., 2009). Pores are

concentrated in spherical to sub-spherical pits (Fig. 2) matching descriptions of previously studied

Late Pleistocene OES from peninsular India (Sahni et al., 1990, 1989), and appear analogous in form

to the pore structures of S. camelus molydophanes (Sahni et al., 1990, 1989) and descriptions of S.

asiaticus (Moure-Chevrire and Geraard 2008).

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Fig. 2: Photographs of OES from Katoati, matching descriptions of other OES from India, Struthio

camelus molydophanes (Sahni et al., 1990; 1989) and some descriptions of Struthio asiaticus (Moure-

Chevrire and Geraard 2008) showing: a) the exterior surface exhibiting pore structures concentrated

into pits; b) a close up of a single pore pit; c) the interior surface morphology.

3.2 Dating

The results of direct dating of OES samples from Katoati are presented in Table 1, together

with a compilation of all previous dates for OES in the Late Pleistocene of South Asia. Our results

show that the application of AMS radiocarbon dating methods have returned the oldest directly dated

evidence for the presence of Struthio in Late Pleistocene India. The minimum ages returned for two of

the oldest samples from KAT1 occur below OSL ages indicating an age of change in depositional

environment at Katoati at ca 60 ka (Blinkhorn et al., 2013). This indicates that the OES from KAT1

predates this change in depositional environment and are therefore older than 60 ka. In addition, they

overly a horizon dated to 77±18 ka, restricting the likely maximum age of these specimens to late

MIS 5 or MIS 4.

Table 1: The results of AMS radiocarbon dating of OES samples from Katoati, Rajasthan,

synthesized with other dated OES samples from India.

Site Lab no Method Material Date (+/-) Ref

Hathnora 1 FT42 ESR Equid tooth 62,800 9000 Patnaik et al., 2009

Hathnora 1 FT39 ESR Bovid tooth 61,600 900 Patnaik et al., 2009

KAT1-23b OxA-25898 AMS C14

OES >62,000 Blinkhorn et al., 2013

KAT2-a OxA-25899 AMS C14

OES >59,900 This article

KAT1-23a OxA-25897 AMS C14

OES >57,900 Blinkhorn et al., 2013

KAT3 OxA-25902 AMS C14

OES 52,100 1100 This article

KAT2-e OxA-25901 AMS C14

OES 49,500 800 This article

JFL2 OxA-19407 AMS C14

OES 45,350 650 Blinkhorn et al., 2013

KAT4 OxA-25903 AMS C14

OES 45,100 500 This article

KAT2 AA 96726 AMS C14

OES >45,000 This article

Mehtakheri AA 8463 AMS C14

OES >41,900 Mishra 1995

Chandresal Grn 10638 C14

OES 38,000 700 Mishra 1995

Chandresal Grn 10639 C14

OES 36,550 600 Mishra 1995

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KAT2-c OxA-25900 AMS C14

OES 35,210 220 This article

Nagda PRL-854 C14

OES >31,000 Agrawal et al., 1991

Ramgada (Ramnagar) PRL-1196 C14

OES >31,000 Kumar et al., 1990

Bori AA 8461 AMS C14

OES 30,000 420 Mishra et al., 2003

Patne Grn 7200 C14

OES 25,000 200 Sali, 1985

Morgaon AA 8846 AMS C14

OES 22,485 320/310 Mishra et al., 2003

Khapardkhera A 9446 C14

Charcoal 15,680 440/415 Mishra et al., 2003

The chronometric evidence provides robust support for the earlier age estimate from KAT2

(>59.9 ka), and for the overall antiquity of all samples analysed, which occur at the limit of AMS

radiocarbon range. The ages of the four samples from KAT2 either predate or fall within the error

range of the OSL date associated with S3 sediments at the site (38±6 ka), which corroborates the

antiquity of the samples, clearly indicating that they are not related with the younger (21±3 ka) S2

sediments (see Table 2). The dating results from KAT 3 (52.1±1.1 ka) and KAT4 (45.1±0.5 ka), also

derived from the upper part of S3 sediment deposits, yielding similar age estimates. The single sample

collected from JFL2, occurring in a slightly different sediment sequence, returned an age of 45.3±0.6

ka.

Table 2: OSL ages and supporting De and dose rate data for the sediment samples from KAT2.

Lab

Sample

Field

Sample

U

(ppm)

Th

(ppm)

K

%

Mean

De

(Gy)

Least

De

(Gy)

Wt.

mean

De

(Gy)

Dose

Rate

(Gy/ka)

Mean

Age

(ka)

Least

Age

(ka)

Wt.

Mean

Age

(ka)

LD-

1099

KAT2 –

Above

1.5 11.6 1.3 48±5 42±1 46±5 2.2±0.1 21±3 19±1 21±3

LD-

1100

KAT2 –

Below

1.9 13.5 1.3 98±19 79±7 93±13 2.4±0.2 40±8 32±4 38±6

We present the eight oldest directly dated samples of OES from India. Our findings support

similar evidence for the antiquity of OES from gravel deposits of the Surajkund Formation at

Hathnora 1 (Patnaik et al., 2009a). At Hathnora 1 samples of OES pieces were found in association

with an equid and a bovid tooth, which were ESR dated to 61.6 ka and 62.8 ka respectively; a third,

reworked tooth from the same sediment context was dated to 131 ka. As a result, the maximum age of

the sediment context, and therefore the OES, is ca 62 ka. This corroborates the antiquity of previously

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dated samples for which only minimum age estimates could be returned, including at Mehtakheri

(Mishra, 1995), Nagda (Agrawal et al., 1991) and Ramgada (Ramnagar)(Kumar et al., 1990).

Elsewhere, OES samples have been dated between >42 to 22 ka, with the exception of Khapardkhera,

where charcoal associated with an archaeological horizon containing OES beads post-dates the Last

Glacial Maximum (LGM). However, it may be anticipated that numerous other previously dated

samples would return older dates if subject to the same pre-treatment protocols and AMS dating

methods that have been employed at Katoati.

3.3 Stable Isotopes

The results of the first stable isotope analyses of Late Pleistocene OES in India are presented

in Table 3 and Fig. 3. The isotopic composition of OES is characterized as a “snap-shot” of diet,

averaged over 3–5 days, with breeding occurring after the rainy season when sufficient nutrient stores

have been acquired (Johnson et al., 1998). As non-obligate drinkers, a range of fractionation processes

within either surface or plant water sources as well as physiological responses of ostrich to humidity

may affect oxygen isotope data preserved in OES, suggesting these results may be best used only as a

qualitative measure of palaeoclimate (Johnson et al., 1998). δ18

O results from the Katoati sites can be

readily separated into three groups. The majority of samples range between -0.5‰ to 0.5‰, whereas

the two samples from the KAT1 excavated sequence show a large step in enrichment of δ18

O,

returning results of ~6‰, and a similarly large step is observed with KAT3a, returning results of

~12‰. As a result, it can be suggested that the majority of samples relate to eggs produced during

more humid conditions, with those from KAT1 and KAT3a indicating increasing aridity.

Table 3: Results of stable isotope studies of OES samples from Rajasthan; a carbon isotope results

reported from physically and chemically abraded samples produced during AMS radiocarbon dating

at University of Oxford; bstable isotopes results reported from sample dated at University of Arizona.

Sample δ13C‰ δ

13C‰

a δ

18O‰ Age (BP)

KAT1-23a -3.277 -5.1 6.057 >57,900

KAT1-23b -3.413 -4.1 6.391 >62,000

KAT2-a -2.902 -3.9 0.173 >59,900

KAT2-b -2.948 -0.007

KAT2-c -2.969 -2.8 0.139 35,210±220

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KAT2-d -3.095 -0.062

KAT2-e -2.655 -2.8 -0.538 49,500±800

KAT2-f -2.904 0.338

KAT2b -2.57 -0.086 >45,000

KAT3-a -9.917 -9.9 11.937 52,100±1,100

KAT3-b -3.311 0.218

KAT4 -3.015 -2.6 0.038 45,100±500

KAT5 -2.921 0.058

Fig. 3: Stable oxygen and carbon isotope results of un-abraded Late Pleistocene OES samples from

KAT1–KAT5 compared with Pliocene to Middle Pleistocene samples from Siwalik sites illustrating

(top) the relationship between isotope values and (bottom) stable isotope variability through time. The

carbon isotope results from Katoati overlap with other Pleistocene Siwalik samples, indicating a

greater focus on C4 plant resources, whereas oxygen isotope results from Katoati show greater overlap

with some Miocene samples, indicating greater aridity.

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Carbon isotope results from OES appear to directly reflect the isotopic composition of the

diet, with an enrichment of ~16‰ (Johnson et al., 1998; Ségalen et al., 2006) such that a pure C3 diet

would produce an average δ13C of -9.5‰ and a pure C4 diet would return results of 4.2‰. Ostriches

show a slight preference for C3 shrubs and select against some vegetation (Johnson et al., 1998),

which could relate to requirement for plant water intake where surface water supplies are not

available. Overall, the results from most samples indicate that the estimated ostrich diets at Katoati

show relatively equal proportions of C3 and C4 input with δ13C of -2 to -3.5‰ relating to a slight bias

toward C3 consumption forming 51.1 to 55.6% of consumed vegetation. The results from KAT3a are

substantially different, indicating a pure C3 diet. The carbon isotope results of samples that were

physically and chemically abraded match well, although not exactly, with un-abraded samples

suggesting potential contamination from secondary carbonate sources have been effectively limited.

Overall, no clear diachronic trends in isotopic composition are evident in this Late Pleistocene

sample.

When compared to isotope results from earlier Siwalik Struthiodiae (Stern et al., 1994), it is

evident that the majority of results from Katoati occupy a tight range within that observed in the

Quaternary period (Fig. 2). Oxygen isotope results from Katoati, which indicated increased aridity

(KAT 23a; KAT 23b; KAT 3a), extend beyond the typical range observed in Indian ratite populations.

Instead, these match results from a limited number of aepyornithoid-type Pliocene specimens. This

may represent aridity experienced at an individual, rather than population-wide, scale. Similarly, only

the carbon isotope results from KAT3a extend beyond the variability observed in ratite δ13

C‰ from

the Siwalik Quaternary sequence, but within the range of Miocene samples, again potentially the

result of individual scale variation.

3.4 Geographic Distribution

The distribution of sites with OES in South Asia is presented in Fig. 4. This illustrates a clear

spatial distinction between the range of Late Pleistocene sites in peninsular India and the earlier

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Siwalik sites. The majority of Late Pleistocene occurrences of OES in India occur in the Chambal and

Narmada valleys, associated with the Malwa Plateau of west-central India. Fewer sites are reported

from the Tapi valley and between the headwaters of the Krishna and Bhima Rivers on the Deccan

Plateau in south-central India. A small number of sites, including Katoati, are notable in that they not

located in the immediate vicinity of one of the subcontinents large drainage basins. Nevertheless, the

landscape at Katoati indicates that the excavated sites occur at the edge of a braided stream network

that may have ultimately fed into the Luni River. Three sites located in Kachchh, Gujarat, are located

close to smaller fluvial regimes in near-coastal locations. Sites bearing OES are reported in both in

lowland or wide valley floor contexts, as well as within regions displaying more relief, occurring at

the edge of different watersheds. In contrast, Pliocene to Middle Pleistocene OES sites are restricted

to the foothills of the Himalaya and the Siwalik ranges and occur in the upper reaches of the Indus

drainage system.

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Fig. 4: Geographic distribution of OES sites in South Asia.

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Despite the occurrence of Late Pleistocene OES in a wide variety of landscape contexts, their

distribution appears geographically limited, predominately associated with the Malwa Plateau and

associated drainage systems. Occupation of the Deccan Plateau is significantly more limited to a

cluster of sites close to Pune, Mahrashtra, and a single site at the watershed of the Tapi River.

Combined, these sites present a SW–NE trend, although Katoati, the Kachchh sites, Ken River and

Shindi are clear exceptions to this. The earliest dated examples of OES in India, dating between >60–

35 ka at Katoati and Hathnora, appear in river systems that flow to the west/south-west. The

concentration of sites on the Malwa Plateau that are associated with eastward flowing drainage

networks mostly date between >40 to >30ka, with the exception of Khapardkhera, which dates to

16ka. The dated sites from the Deccan Plateau, also relating to eastward flowing river systems, are

younger than both of these groups, dating to 30–22 ka.

4. Discussion

The presence of ostrich in Late Pleistocene India appears to be restricted both temporally and

spatially, and marks a significant change in geographic range compared to the Miocene to Middle

Pleistocene. The earliest evidence for ostrich in Late Pleistocene India occurs in contexts dating to

>60 ka, in MIS 4, at the margins of the Thar Desert and the westward draining Narmada basin. While

ostrich are present at Katoati and the Narmada basin during MIS 3, the range of ostrich expands in

this period to include the Malwa Plateau. With the onset of MIS 2, evidence for ostrich is only found

on the north western Deccan Plateau. Only a single example of ostrich, at Khapardkhera, on the

Malwa Plateau, is known from India post-dating the LGM. In order to further assess the timing and

nature of ostrich expansion into India, it is necessary to situate these findings in broader context.

4.1 Chronology

The presence of ostrich in India currently appears restricted from MIS 4 to MIS 2 (>60–16

ka). The earliest dated evidence for ostrich, occurring before 60 ka (MIS 4), stretches beyond the

range of traditional radiocarbon methods, although earlier dates from Katoati are suggested by OSL

dating. Therefore the use of dating methods, such as OSL, is necessary to corroborate the antiquity of

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the samples. As this has typically not been the case in South Asia, expansion of ostrich into western

and central India prior to MIS 4 cannot be precluded. Nevertheless, both environmental and

ecological evidence reviewed below may further support the appearance of ostrich in western and

central India during MIS 4.

Khaparkeda presents the only evidence for the survival of ostrich in India <20 ka. The

regional extinction of ostrich from India is suggested to have occurred across the LGM, perhaps

resulting from extreme climatic pressure and increased competition and human predation (see below).

The presence of numerous OES samples dating to before the LGM and the scarcity of similar samples

dating after the LGM supports the suggestion of significant collapse of ostrich populations at this

time, which ultimately leads to their regional extinction. Based on examples of Struthio extirpation in

North Asia (Janz et al., 2009) and numerous megafaunal taxa at a global scale (Koch and Barnosky,

2006), an alternative hypothesis may suggest the regional extinction of ostrich around the

Pleistocene–Holocene boundary with the return to interglacial climates. However, as yet there is no

evidence to support continuity of ostrich populations in India into the Holocene, and the unique,

mosaic nature of Indian habitats may indicate that the global collapse of megafaunal diversity ca 10

ka is not a suitable analogy (Roberts et al., 2014).

4.2 Taxonomy

The Late Pleistocene ostrich population represented by OES in peninsula India can be most

securely referred to as Struthio sp. In their comparison of OES samples from late Miocene and

Pliocene of East Africa and Namibia, Harrison and Msuya (2005) identify a pattern of decreasing

shell thickness and pore diameter and increasing pore density through time between S. karingarbensis

(shell thickness 2.9–3.2; pore density 2.2/cm2; pore diameter 2.7 mm; age range - 6.5–4.2 Ma.), S.

kakesiensis (Shell thickness 2.5–4.4 mm, mean=3.2; pore density=4.6/cm2; pore diameter =2.5 mm;

age range 4.5–3.6 Ma.), S. daberasensis (Shell thickness 1.7–2.6 mm; mean=2.3; pore density=4–

5/cm2; pore diameter=2.2 mm; age range 6.5–3 Ma.) and S. camelus (Shell thickness 1.5–2.8 mm;

mean = 2.1; pore density 10.8/cm2; pore diameter=1 mm; age range 3.8 Ma. to present). It is worth

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noting that dated OES from Siwalik deposits bearing struthionid-type morphotypes only overlap

chronologically with S. camelus. OES comparable to S. camelus molydophanes associated with

osteological remains identified as S. asiaticus in Late Pliocene deposits at Ahl al Oughlam, Morocco,

exhibit a shell thickness (2.3 to 2.7 mm; mean=2.54 mm) that overlaps the range of S. daberasensis

but a greater pore density (10–20/cm2) comparable to S. camelus, thus fitting the broad trend (Moure-

Chevrire and Geraard 2008). The decreased shell thickness in the Late Pleistocene ostrich population

in India with high pore density continues this African trend. However, while the thickness range

overlaps with S. asiaticus samples from Ahl al Oughlam, the Indian samples are notably thinner.

The occurrence of OES in Central and East Asia share some similarities with South Asia.

The recovery of aepyornithoid-type eggshells from Miocene deposits followed by struthionid forms

appearing in the Pliocene deposits in Mongolia and reported as Struthiolithus (Sauer 1972) parallel

the records from the Siwalik Range. Notably, the samples, originally reported as Struthiolithus,

exhibit thinner shells than their African counterparts, ranging between 1.8–2.4 mm. In addition, the

struthionid-type eggshells exhibit needle point pores rather than pore-pits observed in the

contemporary African sample. A rich collection of Upper Pleistocene specimens dated using C14

methods are reported from China, Mongolia and Transbaikal Russia (Janz et al., 2009; Kurochkin et

al., 2010). Further descriptions of their morphology (i.e. shell thickness, pore structure and density)

are required before meaningful comparison can be made with the South Asian sample.

The direct ancestry and taxonomic affiliation of Late Pleistocene Struthio in western and

central India is difficult to assess based on ootaxonomy alone, particularly as a number of potential

source populations exist. They may represent descendants of the Siwalik populations, a more

widespread population of S. asiaticus that extends from the Siwalik Range to Morocco and potentially

including South Africa, more recent Struthio populations found within this range, such as those

known from Arabia (Potts, 1999) including Late Pleistocene samples from the Mundafan depression

(Crassard et al., 2013), or of shared source population with Central and East Asian Struthio

populations. While it may not be possible to identify which source population, and therefore which

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taxon, expanded into western and central India in the Late Pleistocene, it is possible to explore why

this expansion occurred.

4.3 Neogene and Quaternary habitats of Struthiodiae in South Asia

Significant changes to the geographic and ecological make up of South Asia have occurred

since the earliest known appearance of ostrich, and particularly in the mountainous northern region

which has yielded all evidence for ostrich prior to the Late Pleistocene. Himalayan orogeny and

associated changes to Eurasian climate systems, particularly the evolution of the Asian monsoons, are

likely to have had significant impacts upon ostrich populations. Studies of Himalayan exhumation

rates indicate a decrease in monsoonal intensity after 10 Ma., following the mid Miocene climatic

optimum, followed by an enhanced monsoon system after 4 Ma. (Clift et al., 2008). Isotopic studies of

pedogenic carbonates have indicated a sharp change to C4 dominated ecologies in the Late Miocene

(Cerling et al., 1997; Quade and Cerling 1995), but more recent studies have suggested lateral

variability in floral gradients prior to C4 expansions and greater heterogeneity in floral change

(Behrensmeyer et al., 2007; Morgan et al., 2009). Similarly the expansion of C4 flora has a notable but

complex effect on faunal isotopic signatures (Barry et al., 2002; Morgan et al., 1994).

Two distinct phases in Siwalik OES isotope chemistry can clearly be identified. Mid-Miocene

samples older than ca 9 Ma., exclusively comprising specimens with aepyornithoid-type pore patterns,

present a strong C3 signature associated with relatively humid conditions. Pleistocene samples,

predominately comprising specimens with struthionid-type pore patterns, present a strong C4

signature associated with relatively humid conditions. Limited sample size prevents a clear

characterisation of Late Miocene and Pliocene OES isotope chemistry, although the carbon isotope

signature is intermediate between the range of the mid-Miocene and Pleistocene groups whereas the

oxygen isotope signature indicates increased aridity in contrast to both.

These results broadly corroborate existing trends identified in other isotope archives,

particularly highlighting the appearance of C4 plants in the Siwaliks and their exploitation by

Pleistocene ostrich populations. The apparent synchronicity of the appearance of a C4 signature in

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OES isotope chemistry and the appearance of struthionid-type pore patterns may indicate that the

appearance of new ostrich populations in the Siwalik sequence was related to wide ranging ecological

reconfiguration. However, the continuity of specimens with aepyornithoid-type pore patterns that

substantially overlaps the appearance of C4 flora and OES with struthionid-type pore patterns suggests

that existing ostrich populations were not rapidly replaced. In this context, the Late Pleistocene

expansion of ostrich into western and central India exhibits the exploitation of a similar resource base

with earlier Pleistocene ostrich populations, focused upon C4 habitats.

Extirpation of Siwalik ostrich populations may have been the result of changing relief,

environments, impacts of climate change on seasonality or a combination of these factors, and

identifying which factor may have been dominant in this process is beyond resolution of data

available. However, these Siwalik habitats differed significantly in their geographic structure and

climatic seasonality from the regions of western and central India into which ostrich populations

expanded during the Late Pleistocene.

4.4 Geographic and Palaeoenvironmental context of Late Pleistocene ostrich expansion in western

and central India

Given the known distribution of Struthiodiae spanning the Miocene–Pleistocene, and the

geographical and ecological makeup of north India and Pakistan, the Late Pleistocene expansion of

ostrich into western and central India is most likely to have been the result of dispersals from the

north-west of the subcontinent. Indeed, the NW–SE cline noted in the dated occurrences of OES in

western and central India appears to support this. Geographic and environmental conditions in the

Thar Desert, marking the most north-westerly extremity of the Late Pleistocene ostrich range in India,

are therefore critical to understanding the timing and nature of ostrich expansions.

An expansion into the Thar Desert, whether eastwards from Africa and Arabia or southeast-

wards from the Siwalik Range, requires a crossing of the Indus River. A GIS model of eastward

dispersal routes across southern Asia, based on least cost surface models, indicates the Indus as a

major barrier, indicating that any expansions would likely be directed in to the interior of the Thar

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Desert to be capable of traversing these river systems (Field et al., 2007). Increased humidity

experienced across southern Asia during MIS 5 would have both provided an important context both

for population expansion and an increased potential for river systems to prevent major changes to

geographic ranges. While the Indus may have been a major physical barrier to dispersals, it is likely to

have also acted as a refugium for flora and fauna alike during periods of increased aridity, such as

MIS 4, delivering a constant supply of water from the Himalaya through landscapes lacking direct

precipitation. In this context, the Indus corridor provides the most likely direct source for ostrich

expansions into western and central India. Indeed, during glacial conditions such as MIS 4, changes to

river channel morphology in the mid and lower Indus valley modulated by varying flow and sediment

supply may have provided the most suitable conditions for the crossing of this physical barrier. This

provides the immediate context for the appearance of Late Pleistocene ostrich in India.

The Thar Desert provides the best recorded evidence for Late Pleistocene environmental

change in South Asia and suggests humid conditions were present for much of MIS 3, with evidence

for faltering fluvial systems and the onset of dune mobility occurring by 35 ka (see Blinkhorn 2014).

During this period of faltering humidity in MIS 3, ostrich appear to successfully expand to the

majority of its known range in India, encompassing all dated sites within and to the north of the

Narmada Valley. The occurrence of ostrich in Arabia in this time period has also been recently

reported, dating to 49.8 ka and >50 ka (Crassard et al., 2013). Increasing aridity occurs during leading

up to the LGM, evident in the onset of sand dune mobility commencing in the central Thar Region

and extending to the mouth of the Narmada. It is during the transition between MIS 3–2 and prior to

the LGM that the geographic range of ostrich in India reaches its southernmost extent on the north-

west Deccan Plateau. Only the site of Kharpakeda indicates the limited continuity of ostrich

populations on the Marwa plateau across the LGM.

Fig. 5 presents the distribution of OES sites in India plotted with modelled rainfall during

peak humidity in MIS 5 (which may also serve as an appropriate analogy for early MIS 3 given the

presence of terrestrial proxies directly indicating humid conditions) and during the LGM (a suitable,

but more arid analogue for MIS 4). The earliest evidence for Late Pleistocene ostrich in India, at

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Katoati, occurs when increased aridity in the Thar Desert would have presented arid environments

that may have depressed other faunal populations. Increasing aridity between MIS 3 and 2 may have

extended arid and semi-arid habitats to the east of the Thar Desert, while eventually prohibiting

occupation of the expanding extreme arid desert. The majority of OES sites in India are located in

regions that have experienced arid to semi-arid climates between MIS 4–2, with little indication for

the colonization of sub-humid to humid regions of the subcontinent. Similarly, Janz et al., (2009) have

suggested the Late Pleistocene expansion of Struthio populations in Central Asia may reflect a

strengthening summer monsoon.

Fig. 5: (Left) Distribution of OES sites in India plotted with modeled annual rainfall during the last

interglacial (MIS 5) (Otto-Bliesner et al., 2006) and (right) distribution of OES sites in India plotted

with modeled annual rainfall during the LGM (Braconnot et al., 2007).

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Recent models of vegetation changes across southern Asia (Boivin et al., 2013) offer a useful

context in which to situate the dispersal of ostrich into South Asia. Fig. 6 illustrates the known

distribution of ostrich in India plotted on modelled vegetation from MIS 5 (which may also serve as

an analogy for humid phases of MIS 3) and MIS 4 (which may be analogous to periods of increasing

aridity in MIS 3 and 2). On the enhanced humidity model, a corridor of Sahel-like vegetation is

evident across southern Asia and encircling the arid core of the Thar Desert. Katoati, alongside five

other undated sites occur at the interface between this Sahel corridor and a mosaic of tropical

savannah/woodland grass. The Marwa Plateau sites also appear predominately at a vegetative

boundary, between tropical savannah/woodland grass mosaics and dry tropical woodland habitats.

However, on the heightened aridity model, these sites also cluster at the boundary between Sahel and

tropical savannah/woodland grass mosaics.

Fig. 6: Distribution of OES sites in India plotted with modeled vegetation in (left) MIS 5 and (right)

MIS 4, analogous to humid and arid phases in MIS 3–2 respectively (following Boivin et al., 2013).

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The heterogeneous habitats of the Indian subcontinent appear to have been critical to the

continuity of large faunal taxa in India. Faunal records from the Billasurgum caves, south India,

suggest 20 of 21 large animal species have remained present in the Indian subcontinent over the Late

Pleistocene, with the disappearance of Theropithecus marking the only example of extinction (Roberts

et al., 2014). Nevertheless, a number of species, such as wild ass and Indian rhinoceros, now only

appear in restricted distributions within the Indian subcontinent (Roberts et al., 2014). Fragmentation

and spatial redistribution of India’s habitat mosaic appears to have played a critical role in faunal

mobility and long term continuity in the region (Roberts et al., 2014). The arrival of ostrich in western

and central India before 60 ka appears to have occurred during such a period of habitat redistribution.

The high levels of mobility of the ostrich populations may have been critical to effectively exploit

changing habitat boundaries and compete with Indian fauna, particularly with the extension of Sahel-

like vegetation in the region. Nevertheless, there is little evidence to suggest the successful ostrich

colonization of endemic Indian habitats. It remains unclear what caused the regional extinction of

ostrich from India, although the lack of an endemic source population, competition for marginal

resources and increased predation from Indian fauna over the LGM, as well as exploitation by rapidly

growing modern human populations may all have played some role.

4.5 The importance of ostrich expansion into India for the human dispersal debate

The ostrich is one of two megafaunal species known to have expanded into western and

central India during the Late Pleistocene, the other being Homo sapiens. The dispersal of ostrich into

western and central India before 60 ka offers an important opportunity to assess a number of aspects

of the modern human dispersal debate. Meanwhile, the demographic expansion of human populations

within India may be significant in explaining the regional extinction of ostrich, either through

predation or indirect impacts upon regional ecology.

The appearance of ostrich in Late Pleistocene India prior to 60 ka occurs before the time

frame suggested by some for the appearance of modern humans in India, between 60 to 50 ka (e.g.

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Mellars et al., 2013). The argument for a dispersal of modern humans after 60 ka favours a rapid

coastal dispersal in to India on account of advanced technological and social adaptations, with a

timeframe based on mtDNA evidence (Mellars et al., 2013). Ostrich expansions into India during the

Late Pleistocene indicate continental routes of dispersal into India were also possible. Predation of

ostrich and use of OES by contemporary and prehistoric human populations indicate significant

geographic overlap of humans and ostriches in Sahel-like and savannah habitats (Cooper et al., 2009).

This suggests that the continental routes of expansion exploited by ostrich populations prior to 60ka,

focused upon Sahel-like and savannah habitats, may also have been habitable to human populations

and offer an alternative dispersal route to those proposed by coastal models.

Ostrich beads and symbolic pieces play a role in the identification of behavioural modernity

in the archaeological record, both in Africa (e.g. d’Errico and Stringer, 2011) and in South Asia

(James and Petraglia, 2005; Mellars et al., 2013). The presence of OES beads, and fragments incised

with hatched patterns, such as at Patne (Sali, 1985), is suggested to support a dispersal of human

populations associated with these material behaviours at the onset of MIS 3 (Mellars et al., 2013).

Yet, alternate models argue for earlier expansions of modern humans into South Asia during MIS 5

(see Blinkhorn and Petraglia 2014), possibly preceding the arrival of ostrich. Resolving the timeframe

of the earliest expansion of ostrich into western and central India is therefore critical to assess whether

OES may have been available to human populations throughout MIS 5. An earlier arrival of human

populations would preclude the use of OES, and the appearance of symbolic material culture

produced using OES may indicate regional innovations in response to local demographic and

environmental pressures (Petraglia et al., 2009) rather than an innate feature of modern human

behaviour. Nevertheless, ostrich appear to have been recorded, albeit rarely, in the rich corpus of rock

art known from South Asia, such as at the site of Firengi, south of Bhopal (Badam, 2005). Evidence

for ostrich regional extinction prior to the Holocene supports the suggestion that rock art was being

created in India during the late phases of the Late Pleistocene (Taçon et al., 2010).

If the extension of Sahel-like vegetation across South Asia was a key factor in the dispersal of

both ostrich and humans, potential dispersals of both taxa during MIS 5 should be considered based

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on available palaeoenvironmental reconstructions. However, modern humans appear to have been

uniquely successful in expanding beyond these Sahel-like habitats, to which ostrich appear restricted.

The successful adaptation of modern humans to new habitats and changing environments may have

also impacted upon the regional extinction of ostrich. Overlapping factors of environmental

deterioration and demographic expansion appear to have provided a context for technological

innovation around the MIS 3–2 boundary, associated with the emergence of more efficient hunting

technologies and the earliest evidence for human-modified OES (Petraglia et al., 2009). Whether

through direct predation of ostrich and their eggs, or more indirect ecological impacts on increasingly

fragmented habitats, the growing population of modern humans in South Asia is likely to have been a

factor contributing to the regional extinction of ostrich from India.

5. Summary:

Our results of direct dating of OES samples from Katoati, along with dating of their sedimentary

context, present the first robust evidence to indicate a significant Late Pleistocene range expansion of

ostrich in India before 60 ka. By synthesising these results with existing reports and placing them

within their palaeoenvironmental and ecological context, a restricted timespan for the occurrence of

ostrich expansion into peninsula India, limited to MIS 3–2, has been demonstrated. The range of Late

Pleistocene ostrich in India now appears to track the margin of Sahel-like habitats, with limited

evidence to suggest any extensive colonisation of regions with endemic Indian vegetation. The

dispersal of ostrich into India during the Late Pleistocene is notable in the context of long-term

stability in the diversity of mega-fauna in the region, as well as potential analogies for the more

successful dispersals of modern humans.

Acknowledgements

The fieldwork was supported by the Emslie Horniman Scholarship (Royal Anthropological Institute,

London) awarded to JB. AMS radiocarbon dating at ORAU was funded by the NERC Radiocarbon

Facility (2011/2/16) awarded to MP and JB. JB is supported by a Fondation Fyssen Post-Doctoral

Fellowship. MP is supported by a grant from the European Research Council (no. 295719). We thank

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Jay Quade (University of Arizona) for AMS dating an OES sample from KAT2, Peter Ditchfield

(University of Oxford) for facilitating the stable isotope analysis, and N. Suresh (Wadia Institute of

Himalayan Studies, Dheradun) for dating the OSL samples from KAT2 and providing the

methodology and figures reported in SI. JB and HA thank the Anna University for their support and

facilities. We thank Francesco d’Errico for Fig. 2.

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SUPPORTING INFORMATION

Additional Supporting Information may be found in the online version of this article:

Appendix S1: Quartz Optically Stimulated Luminescence (OSL dating)

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Ostrich expansion into India during the Late Pleistocene: Implications for continental dispersal

corridors

Highlights

Oldest directly dated evidence for Late Pleistocene expansion of Struthio into India

First stable isotope study of Late Pleistocene ostrich eggshell in India

Synthesis of existing evidence for Struthio populations in South Asia

Struthio expansion associated with extension of Sahel-like habitats