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Historical Tropical Forest Reliance amongst the Wanniyalaeto (Vedda) of Sri Lanka: an Isotopic Perspective

Headland and Bailey (1991) argued in Human Ecology that tropical forests could not support long-term human foraging in the absence of agriculture. Part of their thesis was based on the fact that supposedly isolated 'forest' foragers, such as
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  Historical Tropical Forest Reliance amongst the Wanniyalaeto (Vedda)of Sri Lanka: an Isotopic Perspective Patrick Roberts 1,2 &  Thomas H. Gillingwater 3 &  Marta Mirazon Lahr 4 &  Julia Lee-Thorp 2 &  Malcolm MacCallum 3 & Michael Petraglia 1 &  Oshan Wedage 1,5 &  Uruwaruge Heenbanda 6 &  Uruwaruge Wainnya-laeto 6 # The Author(s) 2018 Abstract Headland and Bailey (1991) argued in  Human Ecology  that tropical forests could not support long-term human foraging in theabsence of agriculture. Part of their thesis was based on the fact that supposedly isolated  ‘ forest  ’  foragers, such as theWanniyalaeto (or Vedda) peoples of Sri Lanka, could be demonstrated to be enmeshed within historical trade networks and relyon crops as part of their overall subsistence. Yet, in the same volume and in the years that followed scholars have presentedethnographicand archaeologicalevidence, including fromSriLanka, thatcounter thisproposition,demonstratingthe occupationand exploitation of tropical rainforest environments back to 38,000 years ago (ka) in this part of the world. However, archaeo-logical and ethnohistorical researchhas yet toquantifythe overallrelianceofhuman foragersontropical forestresources throughtime. Here, we report stable carbon and oxygen isotope data from historical Wanniyalaeto individuals from Sri Lanka, in fullcollaboration with the present-day members of this group, that suggest that while a number of individuals made use of agricul-tural resources in the recent past, others subsisted primarily on tropical forest resources as late as the 1800s. Keywords  Tropicalrainforest  .Hunter-gatherers .Indigenouspeoples .Stablelightisotopes .SriLanka .TheWanniyalaeto Introduction During the 1970s and 1980s, tropical forests were seen as ‘  pristine ’  habitats, home to some of the last groups of hunter-gatherer societies untouched by agriculture and capi-talism anywhere in the world (Stiles 1992). Countering this perception, Headland and Bailey (1991) argued that humanforager habitation of tropical forest environments was virtual-ly impossible without the consistent trade with agriculturalsocieties noted in surveys of ethnographic and historical trop-ical forest societies. Key to their argument was the perceiveddietary constraint imposed by highly spaced resources, sea-sonal flux, and the scarcity of energy-rich wild foods, such asfat-rich animals and carbohydrate-rich tubers (Hart and Hart 1986; Headland 1987; Bailey  et al.  1989). Archaeologistsquickly adopted these views. Since then, tropical forests havetended to be considered as  ‘  barriers ’  to the movement of humans, from their first expansion beyond Africa during theLate Pleistocene onwards (Gamble 1993; Bird  et al.  2005;Boivin  et al.  2013).As many have since pointed out, however, this does not mean that purely foraging lifestyles in tropical forests are im- possible,noworinthepast(Bahuchet  etal. 1991;Balée1994; Brosius 1991; Roberts and Petraglia 2015). In 1991  Human Ecology  published a number of studies from different parts of the world that argued strongly against this thesis. Bahuchet  et al.  (1991) demonstrated that it was possible to gain suffi-cient carbohydrate resources from wild yams and other plant  Electronic supplementary material  The online version of this article(https://doi.org/10.1007/s10745-018-9997-7) contains supplementarymaterial, which is available to authorized users. *  Patrick Robertsroberts@shh.mpg.de 1 Max Planck Institute for the Science of Human History, KahlaischeStr. 10, 07745 Jena, Germany 2 Research Laboratory for Archaeology and the History of Art, Schoolof Archaeology, University of Oxford, Oxford, UK  3 Anatomical Museum, College of Medicine and Veterinary Medicine,University of Edinburgh, Edinburgh, UK  4 Leverhulme Centre for Human Evolutionary Studies, Department of Archaeology & Anthropology, University of Cambridge,Cambridge, UK  5 Department of History and Archaeolpogy, University of SriJayewardenepura, Nugegoda, Sri Lanka 6 Wariga Maha Gedara, Kotabakina, Dambana, Sri Lanka Human Ecologyhttps://doi.org/10.1007/s10745-018-9997-7  foods in the Central African rainforest, and that contemporaryrelationships between hunter-gatherers and farmers in this re-gion distorted the perceived importance of such resources inthe past. Similarly, Brosius (1991) discussed how the Penanhunter-gatherers of Sarawak, Borneo, manipulated the sago palm  Eugeissona utilis  to the extent that it could comfortablymeet their calorific needs, while Dwyer and Minnegal (1991)and Stearman (1991) made comparable arguments for popu-lations living in lowland Papua New Guinea and the BolivianAmazon, respectively.Importantly, archaeological research has definitivelyestablished that humans exploited tropical forest resources inthe past. In the same volume, Endicott and Bellwood (1991)reviewed archaeological evidence at a series of Malaysiancave sites for the exploitation of tropical forest animals, in-cluding gibbons, flying foxes, and pigs, suggesting that for-agers lived independently on these forests resources. Recent archaeological research in the rainforests of Southeast Asiaand Melanesia has established the use of forests and forest edges by prehistoric hunter-gatherers since at least 45 ka(Barker   et al.  2007; Summerhayes  et al.  2010; Roberts andPetraglia 2015). South Asia, and particularly Sri Lanka, hasalsoyieldedanabundanceofevidenceforthehuntingofsemi-arboreal and arboreal mammals from 38 ka to 3 ka in tropicalevergreen and semi-evergreen rainforest habitats (Wijeyapala1997; Roberts  et al.  2015a, 2017). Nevertheless, anthropolog- ical and archaeological studies have done little to test theoverall contribution of wild tropical forest resources to humanforagerdietsrelativetootherhabitatsorsubsistencestrategies.In order to investigate this topic, we carried out stablecarbon and oxygen isotope analysis of human tooth enamelfrom historical Wanniyalaeto (also known as  ‘ Vedda ’ ) in-dividuals from Sri Lanka. The status of these groups asdedicated foragers has been questioned since some of theearliest formulations of the hypothesis that tropical forestsare not productive environments for human foragers. It wasargued that by the 1800s, the Wanniyalaeto were part of trade networks in a globalized, colonial South Asia andmade use of agricultural resources obtained through thesenetworks (Bailey  et al.  1989). However, numerous ethno-graphic accounts from the nineteenth and early twentiethcenturies clearly indicate that despite the historical con-tacts, and indeed the long-standing presence of small-scale farming communities in Sri Lanka, these hunter-gatherers retained a mode of subsistence that relied signif-icantly on forest resources (Seligmann and Seligmann1911). Stable isotope analysis of human tooth enamel hasthe potential to directly test the use of forest resources andthus quantify the potential dietary and cultural effects of contact with farmers and imperial powers, and provide ahistorical dataset to compare to recent work on archaeolog-ical collections of forest foragers dating back to 36 ka(Krigbaum 2003, 2005; Roberts  et al.  2015a, 2017). Forest Foraging amongst the Wanniyalaetoof Sri Lanka Sri Lanka has been a key part of the broader debate regard-ing the suitability of tropical forests for long-term humanforager subsistence. The Wanniyalaeto are a minorityIndigenous group in Sri Lanka, whose language is com-monly called  B Vedda ^ , and are often linked to a pre-Sinhalese and Tamil period of occupation (Seligmann andSeligmann 1911). The term  B Vedda ^  actually derives fromthe Tamil word for hunting, but has become a derogatoryterm in Sri Lankan society for anyone leading a rural, mo- bile way of life (Brow 1978; Boyle 2004). The Wanniyalaeto take pride in tropical forest foraging as atraditional way of life, and many historians and anthropol-ogists (De Silva 1972, 1990; Bandaranayake 1985), as well as more recently geneticists (Ranaweera  et al.  2014), haveseen this as an isolating backdrop to cultural and geneticdistinctiveness in this group relative to their neighbours.Between the eighteenth and the twentieth-first century,British colonialism, the growth of the Sri Lankan stateeducation system, upheaval during the Sri Lankan civilwar, and global capitalism have endangered this group ’ ssurvival, as well as their traditional culture and subsistence practices (Spittel 1961; Wickramasinghe 2016). The primary climatic parameter in Sri Lanka is precipita-tion. The highest annual precipitation within Sri Lanka occursin a Wet Zone at the altitudinal gradient between the south-western coastal plain and the central highlands (Roberts  et al. 2015b),whichreceivesbetween4840and2201mm ofannualrainfall and is home to the island ’ s tropical flora of closed-canopy wet deciduous and tropical evergreen mixed diptero-carp forests (Ashton and Gunatilleke 1987; Gunatilleke  et al. 2005) (Fig. 1). Tropical moist deciduous rainforest and tropi- cal semi-evergreen forestextendintothe IntermediateZoneof the island (Ashton and Gunatilleke 1987; Gunatilleke andGunatilleke 1991), which forms an arc from the centre of itswestern coast to the southern tip, with mean annual rainfall of  between1701and 2200mm. The so-called  ‘ DryZone ’  makesupthemajorityofSriLanka ’ s remaininglandmass,withmeanannual rainfall between 1001 and 1700 mm and is character-ized by large expanses of shrubs and grasslands, with some ‘ monsoon scrub jungle ’  or   ‘ arid zone forest  ’  along the north-ern and southern coasts. Although Wanniyalaeto villages aretoday limited to the open  ‘ Intermediate ’  rainforest and drynorthern monsoonal jungles, during the nineteenth and twen-tieth centuries, particularly prior to British colonialism, theywere widespread across the Wet and Intermediate rainforestsof the island (Seligmann and Seligmann 1911; Knox 1981). During the nineteenth and twentieth centuriesWanniyalaeto starch requirements were met by  Dioscorea yams (Spittel 1924, 1961), wild date palms (  Phoenix pusilla ) and wild breadfruit (Sarasin and Sarasin 1893), Hum Ecol  and the seeds, stems, and rhizomes of various tropical forest  plants (Spittel 1924, 1961). Honey was also reported as a major carbohydrate staple in the Wanniyalaeto diet (Seligmann and Seligmann 1911; Lewis 1915). Animal  protein, including bee grubs, terrapins, tortoise, pangolin(  Manis crassicaudata ), bandicoot rats (  Bandicotabengalensis ), porcupine (  Hystrix indica ), giant squirrel(  Ratufa macroura ), hare (  Lepus nigricollis ), jungle fowl( Gallus lafayetti ), mongoose (  Herpestes  sp.), and freshwa-ter eels and fish, appears to have been the most important source of nutrition (Sarasin and Sarasin 1893). TheWanniyalaeto focused their subsistence in this regard onlarge-bodied monitor lizards ( Varanus bengalensis ), ma-caques (  Macaca sinica ), langurs ( Semnopithecus priamthersites ), pigs ( Sus  sp.), mouse-deer (  Moschiolameminna ), barking deer (  Muntiacus muntjak  ), spotted deer (  Axis axis ), and sambhur (  Rusa unicolor  ) (Bailey 1863;Sarasin and Sarasin 1908; Seligmann and Seligmann1911), although their relative importance varied regionally.The basic method of procuring this larger game was bowand arrow, made entirely from available tree parts (Parker 1909; Seligmann and Seligmann 1911; Lewis 1915). The Wanniyalaeto clearly developed specializedhunting and gathering strategies tuned to the capture of tropical forest prey and other products. Bailey  et al. (1989), however, used historical evidence to argue that the Wanniyalaeto maintained economic contacts with ag-riculturalists as early as the seventeenth century, and that they did not completely rely on tropical forest foraging for their dietary requirements (Seligmann and Seligmann1911; Knox 1981). A number of ethnographers record- ed that the Wanniyalaeto traded forest produce, such ashoney, wax, dried venison, and elephant tusks, with localSinhalese communities for cultivars, such as rice and mil-let, alongside cloth, iron arrow-heads and axes, through-out the nineteenth and twentieth centuries (Sarasin andSarasin 1893; Seligmann and Seligmann 1911; Spittel 1924; Morrison, 2014). Knox (1981) even noted that they served in the armies of Sinhalese kings. Seligmann andSeligmann (1911) also described  ‘ Coast Veddas ’  in certain parts of the island. Nevertheless, while these Indigenous peoples undoubtedly exploited new connections, econom-ic relationships, and resources, this does not necessarilymean that they were culturally and economically divorcedfrom the forest and its dietary and other resources. Indeed,the limited forest use of the Wanniyalaeto today is primar-ily a result of land reorganization and the expansion of state systems, initiated under British rule and continued inthe twentieth and twenty-first centuries (Spittel 1961).Furthermore, the recent diversification of theWanniyalaeto economy does not mean that tropical forest resources were insufficient for subsistence without suchtrade, as documented from growing archaeological evi-dence in the region (Deraniyagala 1992; Perera  et al. 2011; Roberts  et al.  2017). Stable Isotope Analysis as a Direct Testof Human Forest Resource Reliance The differing isotopes of elements such as carbon and oxygenrespond differently to physical and biochemical processes be-cause of their mass differences (Sharp 2006). This fraction-ation leads to different relative isotopic abundances in biolog-icaltissuesthatcanbelinkedtoenvironmentalfactors,suchastemperature, or physiological factors, such as modes of pho-tosynthesis. By convention the results are displayed in parts per thousand as the relative abundance of heavy (less abun-dant) to light (more abundant) isotope relative to an interna-tional standard (McKinney  et al.  1950): δ  ‰ ð Þ ¼  R  sample = R  standard − 1   * 1000 ; where  R  is the ratio of the heavy to light isotope.Because the international standard is a marine limestone, Fig. 1  Map showing the vegetation zones of Sri Lanka after Erdelen(1988) and Roberts  et al.  (2015a) Hum Ecol  which is relatively enriched in  13 C and  18 O, most of the δ -values for biological materials (such as plants, toothenamel) are negative.Differential fractionation during photosynthesis results indistinct non-overlapping  δ 13 C values between C 3  ( − 24 to − 36 ‰  (global mean − 26.5 ‰ )) and C 4  ( − 9 to  − 17 ‰  (globalmean − 12 ‰  (Smith and Epstein 1971)) plants. In a tropicalcontext, this distinction is useful for studying the relative pro- portion of C 4  grassland and C 3  woodland or forest, or therelative proportion of C 4  crops such as millet and C 3  cropssuch as rice, in human diets and, indirectly, their associatedenvironments (Krigbaum 2003, 2005; Roberts  et al.  2015a).CAM plants may either fix atmospheric carbon in the manner of C 3  plants or through a modified, diurnal C 4  sequence that leadsto δ 13 CvaluesthatareeitherwithintherangeofC 3 orC 4  plants, or intermediate between the two (O ’ Leary 1981).However, while CAM plants can be found in tropical forests(Whitmore 1998), they are rare (Krigbaum 2001). Within tropical forests, vegetation growing under a closedforest canopy is strongly depleted in  13 C, due to low light (Farquhar   et al.  1989) and large amounts of respired CO 2  that remains semi-trapped under the canopy (van der Merwe andMedina 1991). As a result of the  ‘ canopy effect  ’ , CO 2 , soils,and plants under a closed canopyhave low  δ 13 C values that arealso reflected in the tissues of animals feeding in the sameenvironments (van der Merwe and Medina 1991; Cerling et al.  2004). In the pre-fossil fuel era, tropical faunal toothenamel with  δ 13 C lower (i.e., more negative) than  − 14 ‰  sug-gests reliance on dense or closed canopy forest, while averagevaluesforherbivoresinopenlandscapeswouldbeabout  − 12 ‰ and 0 ‰  for C 3 - and C 4  –  feeders, respectively (Lee-Thorp  et al. 1989a, b; Levin  et al.  2008; Roberts  et al.  2015a, 2017). Stable oxygen isotope data from human tooth enamel cantheoretically provide additional palaeoecological informationabout water resources and food. In tropical ecosystems, veg-etation δ 18 Oprimarilyreflectsthe sourceandnatureofrainfalland then evaporative potential, which is dependent on relativehumidity (Buchmann  et al.  1997; Buchmann and Ehleringer 1998). The relationship between plant   δ 18 O and evaporative potential can be used to infer levels of evapotranspiration andtherefore, indirectly, canopy density (Roberts  et al.  2017). For obligate drinking mammals such as humans, tooth enamel δ 18 O will reflect a combination of imbibed water, climaticand environmental effects on plants at the base of thefoodchain, physiological factors of the individual and the spe-cies it consumes, and the diet of an individual.Although bone collagen is typically the tissue of choice inhumanpalaeodietaryanalysisbecauseoftheextrainformationabout trophic level that can be obtained from stable nitrogenisotope analysis (Ambrose 1993), it is generally poorly pre-served in tropical contexts (Krigbaum 2005). Tooth enamel ischosen here because it is more resistant to post-mortem deg-radation (Lee-Thorp  et al.  1989b; Lee-Thorp 2008) and represents the  ‘ whole-diet  ’  for the period of enamel formation(Passey etal. 2005).Thisisbetweenonetothreeyearsinmost mammals depending on the tooth (Hillson 1996). Moreover,analysis of this tissue enables the data produced for historicalforagers in Sri Lanka to be compared to a growing stableisotope dataset for human tooth enamelthathas beenaccumu-lated for Late Pleistocene and Holocene Sri Lanka (Roberts et al.  2015a, 2017) and Holocene Southeast Asia (Krigbaum 2001, 2003, 2005). Methods Samples Wesampledteethfromgroupslabeledas ‘ Vedda ’ ( n =14)inthehistorical collections of the Duckworth Laboratory, Universityof Cambridge and the Department of Anatomy, University of Edinburgh (Table 1). All samples were donated to the museuminthelatenineteenthorearlytwentiethcenturiesandarethought to belong to members of Wanniyalaeto culture (Table 1). In the process of repatriation negotiations involving these remains, theWanniyalaeto elders indicated an interest in testing the forest relianceoftheirancestorsgiventherapiddisappearanceofthesesubsistence sources from their diet in the twenty-first century asa result of relocation and the expansion of national education,urban, and agricultural infrastructure into their territories.TheCouncilofWanniyalaetoEldersagreedtominimalsam- pling of tooth enamel for stable isotope analysis prior to repa-triation. This project also forms a smaller part of larger collab-orative research with the Wanniyalaeto that has been grantedethical approval from Friedrich Schiller Universität, Jena,Germany and the University of Jayawardenepura, Colombo,Sri Lanka. Close consultation with Indigenous peoples in thisstudy has enabled a simultaneous scientific and cultural output and enriched interpretation of the results that will also be madeavailable to the  ‘ Vedda Heritage Centre ’  in Dambana, SriLanka, in a multi-lingual poster. When selecting the teeth to besampled,teeththatgrowlateinthelifeofanindividualwere preferredsoastoavoidanypotentialinterferenceofweaninginthe dietary signal. We focused on second or third molar teeththat form during the juvenile and early-adult periods of humanlife (Hillson 1996) (Table 1). Photographs were taken of all individualspriortosamplingandareavailablefromtheauthorson request; these will, nevertheless, only be circulated withexpress permission from the Wanniyalaeto elders. Stable Isotope Analysis The sampled teeth were cleaned using air-abrasion to removeany adhering external material. Enamel powder was obtainedusing gentle abrasion with a diamond-tipped drill along thefull length of the buccal surface in order to maximize the Hum Ecol   period of formation represented by the resulting isotopic anal-ysisforbulksamples.Theresultingenamelpowderswerepre-treated using a protocol to remove organic and secondarycarbonate contaminates. This involved a series of washes in1.5% sodium hypochlorite for 60 min, followed by threerinses in purified H 2 O and centrifuging, before 0.1 M aceticacid was added for 10min,followedbyanother three rinsesin purified H 2 O (as per Lee-Thorp  et al.  2012). Following reac-tion with 100% phosphoric acid, the evolved CO 2  wasanalysed for stable carbon and oxygen isotopic compositionusing a Thermo Gas Bench 2 connected to a Thermo Delta VAdvantage Mass Spectrometer at the Division of Archaeological, Geographic and Environmental Sciences,Bradford University. Carbon and oxygen isotope values werecompared against two International Atomic Energy Agency(NBS 19, CO-8) standards and an in-house standard(MERCK). Replicate analyses of standards suggest that ma-chinemeasurementerroris c. ±0.1 ‰ for  δ 13 Cand±0.2 ‰ for  δ 18 O. Overall measurement precision was studied through themeasurement of repeat extracts from a bovid tooth enamelstandard ( n =20, ± 0.2 ‰  for   δ 13 C and±0.4 ‰  for   δ 18 O). Statistical Analysis ANOVAtestswereperformedonhumanenamel δ 13 Cand δ 18 Oto determine if the  B Vedda ^  populations differed from LatePleistocene and Holocene tropical forest foragers from SriLankanarchaeologicalsitesintheWetZoneoftheislandreport-ed by Roberts  et al.  (2015a, 2017), as well as farmers and for- agersfromLateHoloceneBorneoreportedbyKrigbaum(2003,2005). A linear regression was also performed to test whether human enamel  δ 13 C and  δ 18 O were correlated for the  ‘ Vedda ’ individuals sampled here. All statistical analyses were conduct-ed using the free program R software (R Core Team 2013). Results We made no adjustment of the  δ 13 C and  δ 18 O data for humantoothenamelsamplesanalysedfortheSuesseffectas δ 13 C CO2 in1930 (after the srcins of all of the dataset) differsfrom pre-industrial values by just   c.  0.2 ‰  (Friedli  et al.  1986) (Table 1; Fig. 2). The historical  B Vedda ^  sampled have a wide  δ 13 Crange ( − 14.2 to  − 5.2 ‰ ), indicating varied individual relianceon closed-canopy C 3 , C 3 , and C 4  or marine resources.The majority of individuals (~57%: 8  B Vedda ^ ) have  δ 13 Cvalues between  − 15.0 and − 10.0 ‰ , indicative of a clear die-tary reliance on C 3  resources. The two  B Vedda ^  individualswith  δ 13 C values between  − 15.0 and − 14.0 ‰  clearly docu-ment reliance on closed canopy forest resources (Fig. 2). B Vedda ^  individuals with values between  − 14.0 and  − 10.0 ‰  could reflect more open tropical forest foraging, insettings akin to that of the Intermediate Zone rainforest today,or rice reliance, or varying proportions of the two. Over onethird of the  B Vedda ^  sample ( n =5) has  δ 13 C values between − 8.0 and − 5.0 ‰  (Fig. 3). While heavy reliance on marinefoods could result in tooth enamel values as high as  − 10.0and − 9.0 ‰ ,thehighervaluesreportedhereclearlydocument some contribution of C 4  resources to their diets.The  δ 18 O ranges of all groups are much smaller than thosefor   δ 13 C ( B Vedda ^  = − 9.3 to  − 2.9 ‰ ) and the  δ 18 O values ob-taineddonotcorrelatewith δ 13 C(MultipleR-squared=0.01,  p - Table 1  Stable carbon and oxygen isotope ratios of historical Wanniyalaeto ( B Vedda ^ ) individuals analyzed in this studySample Group Accession number Tooth Source Sex  δ 13 C ( ‰ )(VPDB) δ 18 O ( ‰ )(VPDB)VED1  B Vedda ^  XXI.H.2 Lower left M3 Edinburgh Male  − 10.9  − 4.8VED2  B Vedda ^  XXI.H.4 Lower left M3 Edinburgh Male  − 6.0  − 5.1VED3  B Vedda ^  XXI.H.6 Upper left M2 Edinburgh Male  − 13.6  − 5.8VED4  B Vedda ^  XXI.H.8 Upper right M2 Edinburgh Male  − 12.7  − 2.9VED5  B Vedda ^  XXI.H.1 Lower right M3 Edinburgh Male  − 5.5  − 4.2VED6  B Vedda ^  XXI.H.3 Lower left M3 Edinburgh Male  − 7.3  − 7.3VED7  B Vedda ^  XXI.H.5 Upper left M1 Edinburgh Male  − 5.2  − 5.4VED8  B Vedda ^  XXI.H.7 Upper left PM2 Edinburgh Female  − 14.1  − 6.1VED9  B Vedda ^  XXI.G.18 Lower right M3 Edinburgh Male  − 14.2  − 5.2VED10  B Vedda ^  AS.54.01 Lower left M2 Cambridge Male  − 9.3  − 4.1VED11  B Vedda ^  6101 Lower left M2 Cambridge Male  − 10.0  − 6.2VED12  B Vedda ^  6100 Lower left M2 Cambridge Male  − 6.8  − 4.6VED13  B Vedda ^  1197 Upper right M1 Cambridge -  − 11.1  − 4.3VED14  B Vedda ^  1196 Upper right M2 Cambridge -  − 12.0  − 4.8All samples come from historical collections at the Duckworth Laboratory, University of Cambridge, and the Department of Anatomy, University of Edinburgh Hum Ecol
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