The youngest occurrence of Hexaprotodon Falconer and Cautley, 1836 (Hippopotamidae, Mammalia) from South Asia with a discussion on its extinction

Accepted Manuscript The youngest occurrence of Hexaprotodon Falconer and Cautley, 1836 (Hippopotamidae, Mammalia) from South Asia with a discussion on its extinction Advait M. Jukar, Rajeev Patnaik, Parth R. Chauhan, Hong-Chun Li, Jih-Pai Lin PII:

S1040-6182(18)31008-5

DOI:

https://doi.org/10.1016/j.quaint.2019.01.005

Reference:

JQI 7701

To appear in:

Quaternary International

Received Date: 29 August 2018 Revised Date:

13 December 2018

Accepted Date: 3 January 2019

Please cite this article as: Jukar, A.M., Patnaik, R., Chauhan, P.R., Li, H.-C., Lin, J.-P., The youngest occurrence of Hexaprotodon Falconer and Cautley, 1836 (Hippopotamidae, Mammalia) from South Asia with a discussion on its extinction, Quaternary International (2019), doi: https://doi.org/10.1016/ j.quaint.2019.01.005. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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The youngest occurrence of Hexaprotodon Falconer and Cautley, 1836 (Hippopotamidae,

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Mammalia) from South Asia with a discussion on its extinction Advait M. Jukar1, Rajeev Patnaik2, Parth R. Chauhan3, Hong-Chun Li4, Jih-Pai Lin4

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Washington DC 20013, USA

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Center for Advanced Study in Geology, Panjab University, Chandigarh 160014, India

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Department of Humanities and Social Sciences, Indian Institute of Science Education and

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Research, Mohali 140306, India 4

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Department of Geosciences, National Taiwan University, No. 1, Sec. 4, Roosevelt Road, Taipei,

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Corresponding author: Advait M. Jukar

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Email: [email protected]

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Department of Paleobiology, National Museum of Natural History, Smithsonian Institution,

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Abstract

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The earliest hippopotamid fossils from the Indian Subcontinent come from the Miocene of the

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Siwalik Group. South Asian hippopotamidae are represented by the genus Hexaprotodon, and

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remains of these hippos are commonly found in Neogene and Quaternary sites. Here we report

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on the first directly dated specimen of Hexaprotodon sp. from the Narmada Valley of Central

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India and its associated paleoecological implications. The specimen, an upper right canine

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fragment, was dated to 16,467-15,660 cal BP using accelerator mass spectrometry. This

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individual lived during Heinrich Event 1, a particularly arid period. Isotopes from dental enamel

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revealed this animal to have lived in a savanna environment, and likely experienced a shortage of

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water. Using other dated remains of Hexaprotodon from the Indian Subcontinent, we developed

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a tentative extinction chronology, which showed that Hexaprotodon likely survived into the

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Early Holocene. We hypothesize that a combination of climatic stress and anthropogenic impacts

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would have caused the species’ eventual extinction.

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Keywords: India, Quaternary, Narmada, Megafauna, AMS 14C dating

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1. Introduction Hippopotamids first disperse into Eurasia from Africa during the Upper Miocene and are represented by a single genus, Hexaprotodon Falconer and Cautley, 1836 (Geraads, 2010;

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Patnaik, 2013). Hexaprotodon diversifies in South and Southeast Asia during the Late Neogene

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and Quaternary (Deraniyagala, 1944; Hooijer, 1950; van den Bergh et al., 2001; Louys et al.,

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2007; Chauhan, 2008; Patnaik, 2013). While widespread during the Early-Middle Pleistocene,

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Hexaprotodon is found only in South Asia during the Late Pleistocene (Dennell, 2005; van den

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Bergh et al., 2001; Louys et al., 2007; Chauhan, 2008; Patnaik, 2013). Here, we present the

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youngest directly dated specimen of Hexaprotodon from the Narmada Valley in Central India

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and discuss possible causes for its extinction in South Asia.

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The earliest records of Hexaprotodon come from the Siwaliks of India and Pakistan

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(Barry et al., 1982; Patnaik, 2013). Hexaprotodon sivalensis Falconer and Cautley, 1836 is

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known from the Late Miocene to the Middle Pleistocene of the Indian Subcontinent (Barry et al.,

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1982; Patnaik, 2013). Hexaprotodon iravaticus Falconer and Cautley, 1847, a smaller species, is

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known from the Plio-Pleistocene Irrawaddy Beds of Myanmar (Colbert, 1938; Takai and

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Saegusa, 2006). While this species has been reported from the Early Pleistocene of the Pakistan

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Siwaliks (Akhtar and Khurshid, 1997), recent re-evaluations have shown that the specimen

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cannot be distinguished from H. sivalensis (de Visser, 2008). Another species, H. dhokwazirensis

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Akhtar and Bakr, 1995 from the Early Pleistocene of Pakistan has been synonymized with H.

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sivalensis (de Visser, 2008). By the Early Pleistocene, an insular form, Hexaprotodon simplex, is

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found in the Satir fauna on Java (van den Bergh et al., 2001). This species is succeeded by

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Hexaprotodon sivalensis, which persists in insular and mainland Southeast Asia until the late

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Middle Pleistocene (Hooijer, 1950; Peacock, 1958; van den Bergh et al., 2001; Louys et al.,

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2007). Two species are known from Middle-Late Pleistocene sites in India, H. namadicus

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Falconer and Cautley, 1847, and H. palaeindicus Falconer and Cautley, 1847. A third species,

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Hippopotamus deccanensis Anantharaman, Dassarma and Kumar, 2005 was also described from

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India, but the crushed nature of the skull makes it difficult to determine whether this is a valid

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taxon or not. A fourth species, Hexaprotodon sinhaleyus Deraniyagala, 1944 was described from

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the Ratnapura Beds in Sri Lanka. These beds have not been dated, but it has been suggested that

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this species is Middle-Late Pleistocene in age (Deraniyagala, 1955). There is also some

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contention as to whether the two better known Indian species represent one sexually dimorphic

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species, a chronospecies, or two distinct taxa (Salahuddin, 1989). However, a detailed taxonomic

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review is beyond the scope of this study.

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Hexaprotodon is thought to have gone extinct in India at the end of the Pleistocene (Chauhan, 2008), although two disputed specimens have been reported from Holocene sites in

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the Ganges Plain (Alur, 1980; Joglekar et al., 2003). Hexaprotodon is one of several genera of

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large mammals that go extinct in the Late Quaternary (Koch and Barnosky, 2006). This

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extinction event is unique in the Cenozoic because of its size-biased nature whereby large

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species preferentially go extinct (Smith et al., 2018), and several hypotheses have been

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developed to explain this extinction including human overexploitation (Martin, 1973, 1984),

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climate change and subsequent community change (Graham and Lundelius Jr, 1984), synergies

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between climate change and human pressure (Lorenzen et al., 2011; Cooper et al., 2015; Metcalf

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et al., 2016), removal of keystone herbivores (Owen-Smith, 1987), and a hyperdisease (MacPhee,

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1997). While weakening monsoons and changing fluvial regimes have been implicated in the

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extinction of Hexaprotodon sivalensis in the Early-Middle Pleistocene (Dennell, 2005; Jablonski,

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2004; Louys et al., 2007), the extinction of Late Pleistocene Hexaprotodon has seldom been

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investigated.

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In this study, we 1) use accelerator mass spectrometry (AMS) radiocarbon dating method to date a Hexaprotodon specimen from the Narmada Valley, 2) analyze carbon and oxygen

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isotopes from dental enamel from the specimen to determine the environment it lived in, 3)

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develop an extinction chronology for Hexaprotodon using dated occurrences from the Indian

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Subcontinent to estimate the probable last appearance, and 4) correlate the chronology with

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environmental and anthropogenic events in an effort to develop hypotheses for the extinction.

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2. Regional Setting and Geology

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The specimen BNF2-1 was recovered from a fossiliferous silt horizon of the Baneta Formation in the Upper Narmada alluvium exposed at Baneta (Fig. 1A and 1B). Fossils from the

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Narmada alluvium have been known since the mid 19th century (Prinsep, 1834). The alluvium

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has been divided stratigraphically into seven formations, loosely spanning the entire Quaternary

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(Tiwari and Bhai, 1997). Of the seven, Hexaprotodon along with a diverse mammalian herbivore

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fauna and lithics can be found in the Surajkund Formation—roughly correlated with the Middle-

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Late Pleistocene, and in the overlying Baneta Formation—correlated with the Latest Pleistocene

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(Badam et al., 1986; Biswas, 1997; Patnaik et al., 2009; Sonakia and Biswas, 2011). The

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Surajkund Formation at the type locality Hathnora exhibits as a ~4 m section of boulder

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conglomerate and cross-bedded sandy-pebbly layers containing fossils, and Acheulian and early

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Middle Palaeolithic tools (Tiwari and Bhai, 1997; Patnaik et al., 2009). The Baneta Formation, in

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contrast, comprises of cross-bedded gravelly sand followed by fine-grained silts and

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carbonaceous clay (Fig. 1B). Pollen assemblages suggest that the Surajkund Formation was

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deposited under warm, humid conditions, whereas palynologically, the Late Pleistocene Baneta

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Formation indicates cool, dry conditions similar to the Last Glacial Maximum (Patnaik et al.,

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2009). Other Late Pleistocene occurrences of Hexaprotodon also come from similar fluvial

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deposits in peninsular India and the Indo-Gangetic Plain, and are often also spatially or

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stratigraphically associated with lithics (Fig. 1A; Chauhan, 2008).

3. Materials and Methods

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3.1. Specimen

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The specimen, BNF-2-1, is an upper right canine (Fig. 2A and 2B). The cross section

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shows that the specimen is a little fragmented and distorted (Fig. 2B), however, the specimen

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exhibits a deep posterior longitudinal groove (Fig. 2A and 2B). A deep posterior groove on the

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upper canine is considered diagnostic to Hexaprotodon (Coryndon, 1977; Harris, 1991; Harrison,

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1997; Boisserie, 2005). However, some variation does exist within the genus. For example,

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Hexaprotodon bruneti from Middle Awash, Ethiopia shows both shallow and wide grooves

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(Boisserie and White, 2004). Hippopotamus upper canines, in contrast, are characterised by

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having narrow and shallow posterior longitudinal grooves (Boisserie, 2005). Since there are no

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known upper canine characters that distinguish the better-known Middle-Late Pleistocene

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species, Hexaprotodon namadicus and Hexaprotodon palaeindicus, we assign the BNF-2-1 to

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Hexaprotodon sp.

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3.2. Radiocarbon Dating

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Sample BNF-2-1 was sent to the NTUAMS lab in the Department of Geosciences, National Taiwan University for AMS 14C dating. The sample was physically cleaned with a

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metal brush and deionized water to remove any detritus on the surface. Bone fractions were

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separated from the bulk sample, then grinded into about 3 mm size grains. The ground bone

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sample was first pretreated by acid-base-acid (ABA) method to remove any carbonates and

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dissolvable humic acids on the surface (Brock et al., 2010; Zhao et al., 2017). Then, the

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pretreated sample was placed in 0.5 N HCl and heated at 80oC overnight for collagen extraction.

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Under this condition, any carbonates inside the sample would also be removed. Normally,

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collagens would be extracted from the bone. However, we did not get a visible amount of

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collagen from the sample, probably due to small size of the bone sample and very low collagen

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concentration of the bone. Regardless, organic matters including collagens would be leached out

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with dissolved phosphates throughout this procedure (Brock et al., 2010). The solution was

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neuturalized with 0.1 N NaOH. The phosphates together with any organic components including

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collagens would be precipitated out. During this process, pH does not exceed 7, so that

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absorption of the atmospheric CO2 is negligible. The precipitates were separated, cleaned and

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dried. The dried phosphate is pure white in color and contains small amount of organic carbon.

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The sample phosphate was placed in a 9 mm quartz tube with pre-combusted CuO and a piece of silver. The tube was placed on a vacuum line and sealed under vacuum of 1e-5 mbar,

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then put into a Muffle furnace at 850oC for 6h. The CO2 oxidized from the sample was purified

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and converted into graphite following the method of Xu et al. (2007). The graphite target was

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measured with a HVE 1.0 MV Tandetron Model 4110 BO-accelerator mass spectrometer (AMS)

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together with at least three international standards (OXII, 4900C), three backgrounds (BKG) and

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two inter-comparison samples (IRIs). The measured 14C/12C and 13C/12C ratios were used to

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calculate ∆14C values and conventional 14C ages with a 14C half-life of 5568 years after

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correction for carbon isotopic fractionation using the δ13C values of the samples. We used the

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IntCal13 calibration curve (Reimer et al., 2013) in the OxCal Online Radiocarbon Calibration

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program (Ramsey, 2001) to calibrate the AMS date.

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3.3. Stable Isotope Analyses

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A cross-section of the canine fragment was prepared using a Beuhler Precision Saw, Isomet 1000. Stable isotope analysis on the canine enamel was performed using conventional

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H3PO4 digestion at SIRFER Stable Isotope Ratio Facility for Environmental Research at the

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University of Utah. Approximately 5 milligrams of powdered enamel was treated to remove

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carbonates and analysed on a Finnigan Delta V plus IRMS (isotope ratio mass spectrometer)

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equipped with a Finnigan TC/EA through Thermo Finnigan Conflo IV.

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3.4. Extinction Chronology

We constructed an extinction chronology using other available dates for Hexaprotodon

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from the Indian Peninsula (Table 1). Available radiocarbon dates were ranked using the scale

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developed by Barnosky and Lindsey (2010) (Supplementary Table S1). Because of the

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incomplete fossil record, the youngest occurrence of a taxon oftentimes predates the actual

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extinction time because the last individual of a species is almost never preserved (Wang and

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Marshall, 2016). In order to account for this, we used the Gaussian-Resampled Inverse Weighted

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McInerny et al. (GRIWM) method developed by Bradshaw et al. (2012) to analytically estimate

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the last appearance datum (LAD) for Hexaprotodon. GRIWM progressively up-weights the

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temporal gaps between dates closer to the time the taxon ceases to appear in the fossil record

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(Bradshaw et al., 2012). For each site, we included either the only available date, or the oldest

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and youngest dates in the GRIWM analysis. Radiocarbon dates were calibrated using the

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IntCal13 calibration curve (Reimer et al., 2013) in the OxCal Online Radiocarbon Calibration

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program (Ramsey, 2001) before they were included. We used code developed by Saltré et al.

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(2015) to perform the analyses. This chronology is then compared with the history of human

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activities in the Indian Subcontinent as well as global and regional climates (Fig. 3). We used the

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GICC05 Greenland Ice Core oxygen isotope curve (Andersen et al., 2006; Rasmussen et al.,

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2006; Steffensen et al., 2008; Seierstad et al., 2014) as a proxy for ice volume, and therefore

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global temperature. For regional monsoon data, we used a composite curve of oxygen isotope

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data from speleothems from Bittoo Cave in Himachal Pradesh, Northern India (Kathayat et al.,

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2016), and Mawmluh Cave, Meghalaya, Northeastern India (Berkelhammer et al., 2012; Dutt et

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al., 2015).

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4. Results and Discussion

The Baneta Hexaprotodon specimen is dated to 13,344 ± 135 BP (16,467-15,660 cal BP,

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Table 2). This is the only directly dated Hexaprotodon from the Indian Subcontinent, and

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represents the youngest unambiguously known specimen. This date also conforms to the

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chronostratigraphic framework of the Baneta section where a carbonaceous clay layer from

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~4.0m above this specimen was dated to 8,740 ± 540 BP (Fig. 1B; Patnaik et al., 2009).

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The Baneta Hexaprotodon lived during a particularly dry period in the Late Quaternary known as

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Heinrich Event 1. This period is characterized by a catastrophic drought in South Asia caused by

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an extremely weak Indian Monsoon forced by high latitude cooling (Stager et al., 2011;

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Denniston et al., 2013; Deplazes et al., 2014; Dutt et al., 2015; Tierney et al., 2015; Zhou et al.,

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2016). The enamel isotope analyses support the paleoclimatic characterization. The δ18O enamel

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isotope value of -2.9‰ indicates dry conditions (negative δ18O values around -8.0‰ suggest wet

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conditions, whereas less negative or positive values are indicative of water stress or dry

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conditions [Kohn, 1996; Kohn and Cerling, 2002]). Previous palynological studies on terminal

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Pleistocene sediments from the Narmada Valley have shown the presence of savannas in the

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surrounding region (Patnaik et al., 2009; Verma et al., 2009). The δ13C value from the

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Hexaprotodon specimen of 0.1‰ suggests a C4-dominated diet, further supporting the

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paleoenvironmental interpretation since C4 vegetation tends to flourish in arid, seasonal climates

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(Bond, 2008).

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The GRIWM analysis (Fig. 3) estimated the most probable LAD at 9.001 ka (95% CI =

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10.160-8.173 ka) suggesting that Hexaprotodon survived into the early Holocene. The probable

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extinction time depends on the quality of the dates used in the GRIWM analysis. We used a

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ranking protocol developed by Barnosky and Lindsey (2010) to assess their quality. All previous

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dates ranked below 12 largely because they 1) were obtained using conventional radiocarbon

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dating procedures, and 2) the samples used were freshwater bivalve shells instead of actual

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Hexaprotodon specimens (Table 1). The ‘hard water’ reservoir effect, and diagenesis of the outer

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layers of the shell can affect the age produced from these samples (Preece et al., 1983; Yates,

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2016; Wright, 2017). According to standard protocols used for radiocarbon dating in India, dilute

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hydrochloric acid was typically used to remove carbonates from the surface of the samples

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(Kusumgar et al., 1963; Agrawal and Kusumgar, 1974), thus reducing the effect of diagenesis.

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However, since no information on reservoir effects was available, we cannot rule out that some

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of these sites/specimens are several hundreds of years younger than the current available dates

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suggest they are. This effect would only serve to concentrate the chronology closer to the

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terminal date obtained in this study, and is unlikely to affect the overall pattern of extinction. It

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would however constrain the GRIWM extinction estimate closer to the terminal date since the

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analysis up-weights the gaps between the younger dates to estimate true extinction time

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(Bradshaw et al., 2012). Therefore, we present this chronology tentatively until further samples

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are dated. At present, there is no definitive evidence of Hexaprotodon from Holocene sites in the

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Subcontinent (Pandey, 1990; Thomas and Joglekar, 1994). While possible hippopotamid remains

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have been reported from the Mesolithic oxbow lake sites of Sarai Nahar Rai and Mahadaha

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(Alur, 1980; Joglekar et al., 2003), these occurrences have been disputed by other researchers

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as misidentifications of Rhinoceros remains (Pandey, 1990). Additionally, subsequent studies

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on the fauna at these two sites have not recovered or identified any remains of Hexaprotodon

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(Chattopadhyaya, 1996, 2008). Photographic evidence of the fossils does not strongly suggest

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that the specimens belong to Hexaprotodon (J-R Boisserie, pers comm. 2018), therefore, we

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take a conservative approach and do not consider these occurrences as valid until the fossils

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have been thoroughly reassessed. Nevertheless, if Hexaprotodon persisted into the Holocene

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when monsoon intensity was high (Kale, 2007), it shows that peninsular Hexaprotodon survived

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several abrupt changes in the monsoon (Dutt et al., 2015), periods of river aggradation (Rajaguru

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and Kale, 1985; Kale and Rajaguru, 1987; Mishra et al., 2003; Williams et al., 2006; Roy et al.,

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2012), and aridity during the Late Pleistocene (Blinkhorn and Petraglia, 2017). Therefore, other

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factors likely played a role in its extinction.

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Anthropogenic impacts such as overhunting, and habitat alteration are also thought to

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have played a dominant role in the Late Quaternary extinctions (Koch and Barnosky, 2006). The

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unusual large-size bias of the megafaunal extinction is strong evidence for the dominant role of

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humans as an extinction driver (Smith et al., 2018). Homo sapiens co-occurred with

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Hexaprotodon for several thousand years before it went extinct (Bae et al., 2017). Its extinction

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is predicted to have taken place in the early Holocene, after cultures have become more

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sophisticated and human populations experienced exponential growth (Fig. 3; Atkinson et al.,

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2008; Petraglia et al., 2009). However, while evidence for hippopotamid butchery and processing

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for consumption is known from Quaternary sites in Africa and Europe (Säve-Söderberg, 1953;

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Hill, 1983; Bunn, 1994; Horwitz and Monchot, 2002; Fiore et al., 2004; Schrire, 2014;

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Landeck and Garriga, 2016; Altamura et al., 2018), there is at present no clearly demonstrated

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evidence of comparable processing in the Indian Subcontinent despite the long co-occurrence

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with humans and the association of Hexaprotodon remains with lithics at Pleistocene

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localities (Chauhan, 2008). The lack of kill-sites doesn’t necessarily indicate that

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Hexaprotodon was not hunted. The paucity of paleontological sites in the Subcontinent that

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preserve large mammal remains indicates that potential butchery sites are likely under-

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sampled. It is also possible that historically collected remains have not been thoroughly

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inspected for signs of processing or taphonomic biases prevented the preservation of these

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sites or specimens.

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An alternative explanation to direct anthropogenic impacts or environmental

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deterioration is a combination of the two factors. Hexaprotodon is thought to be semi-aquatic

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like the extant Hippopotamus amphibious (Boisserie et al., 2011) which would make it

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particularly sensitive to changes in the monsoon and the amount of water available in rain-fed

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rivers and lakes. Reduction of the availability of water in rivers and lakes during Heinrich Event

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1 and other comparable dry periods would have resulted in a loss of favorable habitat. Such

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changes are known to fragment ranges, and weaken meta-population dynamics (Lorenzen et al.,

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2011; Mays et al., 2018). In fact, populations of modern hippos under drought conditions are

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known to produce fewer pregnant females because of overcrowding, and a lack of food sources

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and shelter (Smuts and Whyte, 1981). Declining populations can show genetic abnormalities and

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allee effects, further putting them at risk (Brook et al., 2008; Rogers and Slatkin, 2017). Thus,

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the gradual fragmentation of Hexaprotodon populations and associated declines in genetic

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diversity through the Late Pleistocene, and repeated drought stress may have made the species

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more vulnerable to extinction. Low levels of human exploitation occurring concurrently might

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have been devastating to these already stressed populations. Indeed, simulation studies on North

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American megafauna have shown that low levels of human hunting can produce the magnitude

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of extinction seen in the fossil record (Alroy, 2001).

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5. Conclusion

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Indian Hexaprotodon were Eurasia’s last surviving species of hippopotamid. Here we provide the first direct date for this species in the Indian Subcontinent, and using the extinction

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chronology, we hypothesize that a combination of climate stress and growing human populations

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would have caused its extinction. Fluctuating environments may have fragmented populations

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and eroded genetic diversity making the species more susceptible to even low-levels of

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anthropogenic pressure—either from hunting or from competition for habitat, as has been seen in

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the case of the Sumatran rhino (Mays et al., 2018). The evidence at present does not allow us to

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test the hypothesis of population reduction and range fragmentation caused by climate change.

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Ancient DNA can provide insights into whether this species was declining gradually and whether

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the extinction occurs during a recovery period, which would more strongly implicate humans as

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a major contributor. Furthermore, additional directly dated Hexaprotodon specimens will help

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refine the extinction chronology, and provide more robust estimates for when the species

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eventually went extinct.

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Data Availability

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The specimen BNF2-1 is accessioned in the Department of Geology, Panjab University,

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Chandigarh, India.

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Acknowledgements

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We would like to thank Dr. T. Cerling and Dr. K. Uno for their kind help in the stable isotope

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analysis, Dr. F. Saltré for assistance with the GRIWM analysis, and Ms. C.-Y. Chou at the

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NTUAMS Lab for AMS 14C dating. We wish to thanks the editorial team, and two anonymous

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reviewers for their feedback that helped improve this manuscript. This work was supported in

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part by the Ministry of Earth Sciences, Government of India (MoES/P.O.(Geosci)/46/2015),

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Promotion of University Research and Scientific Excellence (PURSE) from the Department of

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Science and Technology, Government of India, and the Ministry of Science and Technology,

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Taiwan, R.O.C. (MOST 106-2116-M-002-018 and MOST 107-2116-M-002-007).

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Table 1. Dates from sites where Hexaprotodon remains have been recovered in the Indian Subcontinent. Rank refers to the quality of dates assessed using the scale developed by Barnosky and Lindsey, 2010. Raw date ± SD (BP)

Median age (Cal BP)

Freshwater Shells

20135 ± 220

24222

Freshwater Shells

26250 ± 420

30431

Sigma (Cal BP)

From (Cal BP)

To (Cal BP)

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24900

23680

6

Jones and Pal, 2009

31111

29537

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Jones and Pal, 2009

Material Dated

14

Beta 4791

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Beta 4793

Baghor-Son River

TL

Alpha 898

26100 ± 5400

Jones and Pal, 2009

Baghor-Son River

IRSL

BN 2

24000 ± 3000

Williams et al., 2006

Baghor-Son River

IRSL

BN 3

39000 ± 9000

Williams et al., 2006

Baneta

AMS

NTUAMS 4285

BNF2-1

Inamgaon

14

TF 1177

Freshwater Shells

Inamgaon

14

Kalpi

IRSL

Kalpi

IRSL

C

C

TF 1003

SC

421

M AN U

TE D

Baghor-Son River

C

EP

Baghor-Son River

Method

RI PT

Lab Number

Site

AC C

299 300

Freshwater Shells

Rank Reference

13344 ± 135

16050

198

16467

15660

19290 ± 360

23255

420

24072

22464

Agrawal and 9 Kusumgar, 1975a

24769

Agrawal and 7 Kusumgar, 1975a

21725 ± 605

26062

651

27392

12 This study

45000 ± 9000

Tewari et al., 2002

43000 ± 7000

Tewari et al.,

15

ACCEPTED MANUSCRIPT

2002 TF 1245

Freshwater Shells

19175 ± 340

23126

14

PRL 86

Freshwater Shells

25790 ± 830

29951

14

BS 163

Freshwater Shells

27410 ± 425

14

TF 967

Freshwater Shells

33700 ± 1723

Tadula

14

BS 562

Freshwater Shells

34470 ± 2070

Wangdari

14

BS 561

Freshwater Shells

26820 ± 750

Rati Karar (Devakachar)

C

C

C C

818

23925

31435

22418

Agrawal and 10 Kusumgar, 1975b

28275

10 Thapar, 1979

429

32474

30780

Mishra, 1995; 9 Rajagopalan et al., 1982

38297

1983

42218

34799

Agrawal and 9 Kusumgar, 1975a

39217

2424

44221

34881

6 Sathe, 1989

30935

820

32790

29403

6 Sathe, 1989

SC

Nandur Madhmeshwar

C

395

31364

M AN U

Mahagara Gravels III

C

RI PT

14

TE D

Mahagara Gravels III

301

14

302

luminescence; cal BP = calibrated years before present. Raw dates for 14C and AMS dates refer to uncalibrated ages.

304 305

EP

AC C

303

C = traditional radiocarbon dates; AMS = accelerator mass spectrometry; TL = thermoluminescence; IRSL = infrared stimulated

16

ACCEPTED MANUSCRIPT

Table 2. Results of the AMS dating procedure on the Baneta Hexaprotodon sp. specimen, BNF2-1 C12

C14

current

counts statistical

(A) NTUAMS- 7.30E-06

11219

C14

pMC

δ14C (‰

14

(% ±

± 2σ)

(BP ± 2σ)

Calibrated age (cal BP)

error (%)

2σ)

0.94

18.99 ±

-810.1 ±

0.19

8.2

4285

C Age

RI PT

Lab Code

13,344 ±

SC

306 307

135

16,46715,660

pMC = % of modern 14C activity; BP = uncalibrated years before present; cal BP = calibrated

309

years before present

310

Figure Captions

311

Fig. 1. Location of dated localities where Hexaprotodon has been recovered from the Indian

312

peninsula and the Gangetic Plain (A), and stratigraphic column of the Baneta section where

313

BNF-2-1 was recovered (B). The yellow star in (A) shows the location of the Baneta section in

314

the Narmada Valley. Black circles represent other dated localities that were included in the

315

analyses. (B) shows the position in the stratigraphic section where BNF-2-1 was found.

TE D

EP

AC C

316

M AN U

308

317

Fig. 2. Upper Right Canine of Hexaprotodon sp. (BNF-2-1). (A), posterior view showing the

318

longitudinal posterior groove. (B), cross section of the canine showing the damage and the

319

diagnostic deep and wide longitudinal posterior groove.

320

17

ACCEPTED MANUSCRIPT

Fig. 3. Extinction chronology for Hexaprotodon in the Indian Subcontinent. Black circles are the

322

median calibrated 14C dates with 95% confidence intervals (CI) ranked <12. The red diamond is

323

the median calibrated 14C date ranked >12 from this study. Grey circles are the non-radiocarbon

324

dates with 95% CI. Each vertical bar represents a site where a particular species was found;

325

multiple dots on a site represent the youngest and oldest dates and error bars. The orange hollow

326

circle the median expected extinction date estimated using GRIWM with 95% CI. The grey bars

327

represent archaeological phases. The Middle Palaeolithic and Late Palaeolithic, which is

328

characterized by microliths (James and Petraglia, 2005) that overlap with each other. The δ18O

329

record from the GICC05 Greenland ice core as a proxy for global temperature and a composite

330

record of δ18O from Bittoo and Mawmluh cave speleothems as a proxy for monsoonal intensity

331

in the Subcontinent. Horizontal blue bars represent the last glacial maximum (LGM), Heinrich

332

Event 1 (H1), and Younger Dryas (YD) cold periods. MIS 1-3 represents the marine isotope

333

stages 1 through 3.

TE D

M AN U

SC

RI PT

321

334

Supplementary Table S1. Detailed site description and ranking protocol. Dates in bold were

336

used in the GRIWM analysis.

338

AC C

337

EP

335

339

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