Thursday, September 29, 2011

Isotope evidence for the diet of the Neanderthal type specimen.

Isotope evidence for the diet of the Neanderthal type specimen. Introduction We present here the results of a dietary isotopic study (of bonecollagen carbon and nitrogen isotope ratios) of the Neanderthal typespecimen, a second Neanderthal found in 1997/2000, and associated faunafrom the Kleine Feldhofer Grotte site in the Neander Valley, Germany.There is limited faunal material available from this site, and like mostother Middle Palaeolithic sites in Europe, plant remains do not survive,so isotopic analysis is the only method that could be used toreconstruct the diet and subsistence strategies of these twoNeanderthals. The measurements of the stable isotope stable isotopen.An isotope of an element that shows no tendency to undergo radioactive breakdown. ratios of carbon([sup.13]C/[sup.12]C, the [delta][sup.13]C value) and nitrogen([sup.15]N/[sup.14]N, the [delta][sup.15]N value) in bone collagenextracted from human and animal bones has become a well-establishedmethod for determining past diets (Schwarcz & Schoeninger 1991;Ambrose 1993; Katzenberg 2000; Sealy 2001). Through a number of lines ofresearch, including empirical observations on living organisms, it hasbeen determined that the carbon and nitrogen isotope ratios in mammalbone collagen reflect the isotope ratios of dietary protein consumedover the last years of the life of that mammal (Ambrose & Norr1993). Early applications of the method focussed mainly on the[delta][sup.13]C values to determine the amounts of [C.sub.3] vs.[C.sub.4] plants (i.e. the spread of maize agriculture in the Americas)in human diets (Vogel & van def Merwe 1977; van der Merwe &Vogel 1978) or the amounts of marine vs. terrestrial foods in humandiets (Tauber 1981; Chisholm et al. 1982; Schoeninger et al. 1983). One of the more successful later applications of the method hasfocussed mainly on the [delta][sup.15]N values to determine the diets ofNeanderthals in Europe. Nitrogen in diets can only come from dietaryprotein, so mammal bone collagen [delta][sup.15]N values indicate themain sources of dietary protein in a mammal's long-term diet, andis enriched by approximately 2-4 [per thousand] over dietary protein[delta][sup.15]N values (DeNiro & Epstein 1981; Schoeninger &DeNiro 1984). An early application of this method, by Bocherens andcollaborators (Bocherens et al. 1991; Fizet et al. 1995) showed that, incomparison to animals from the same site, the Neanderthals from the siteof Marillac, France (now called Les Pradelles), most resembledcarnivores such as wolves in their isotope values. Therefore, theyconcluded that Neanderthals were top-level carnivores in this ecosystem.Since this early study other isotope studies of Neanderthals have beencarried out at the sites of Scladina, Spy and Engis in Belgium(Bocherens et al. 1999; 2001), Vindija, in Croatia (Richards et al.2000) and Saint-Cesaire (Bocherens et al. 2005), LesRochers-de-Villeneuve (Beauval et al. 2006) and Jonzac (Richards et al.2008) in France. In all of these studies, the authors reached the sameconclusion; that Neanderthals were top-level carnivores in theirecosystems. The existing isotopic studies are on Neanderthals from centralFrance, northern Western Europe Western EuropeThe countries of western Europe, especially those that are allied with the United States and Canada in the North Atlantic Treaty Organization (established 1949 and usually known as NATO). (Belgium) and southern Central Europe Central Europe is the region lying between the variously and vaguely defined areas of Eastern and Western Europe. In addition, Northern, Southern and Southeastern Europe may variously delimit or overlap into Central Europe. (Croatia) and so far there are no isotopic studies on Neanderthals fromwhat is likely to be a much harsher environment in northern CentralEurope. Therefore, we obtained material from a right humerus humerus:see arm. of theNeanderthal type specimen, first discovered at the site of KleineFeldhofer Grotte in the Neander Valley in Germany in 1856 (Schaaffhausen1858; Fuhlrott 1859) as well as a right humerus from a secondNeanderthal from the same context, discovered by Schmitz and colleaguesin 1997/2000 (Schmitz & Thissen 2000a; 2000b; Bolus & Schmitz2006) for isotopic analysis. It is necessary, where possible, to compare the isotope values ofhumans and Neanderthals to those of associated animals, especiallyspecies that are clear herbivores or carnivores. This assists in theinterpretation of the isotope values of omnivores like humans, and isalso necessary as there are variations in isotope values in differentregions related to climate (Heaton et al. 1986), as well as changes inisotope values through time (Richards & Hedges 2003). Here we wereable to sample associated fauna from the site to compare with theNeanderthals. Below we discuss the results of our analysis and comparethe data with those of other Neanderthals from Eurasia. The Kleine Feldhofer Grotte site The two Neanderthals and associated fauna in the study come fromthe site of the Kleine Feldhofer Grotte in the Neander Valley in westernGermany The geographic term Western Germany (German: Westdeutschland) is used to describe a region in the west of Germany. The exact area defined by the term is not constant, but it usually includes North Rhine-Westphalia and Hesse, the . The site was destroyed by quarry workers in 1856 and 16Neanderthal skeletal elements were recovered (Fuhlrott 1859). TheNeanderthal type specimen has been curated at the RheinischesLandesmuseum in Bonn since 1877 and from 1991 has been the subject of aninterdisciplinary research project directed by the museum (Schmitz1995). Excavations undertaken by the Rheinisches Amt furBodendenkmalpflege, Bonn, under the direction of Schmitz and Thissen in1997 and 2000 (Schmitz & Thissen 2000a; 2000b; Schmitz 2006) ofsediments that were removed from the cave in 1856 recovered anadditional 73 human skeletal elements from the Middle Palaeolithic(Smith et al. 2006). In total, the skeletal remains indicate a minimumof two adult individuals and at least one sub-adult (Smith et al. 2006).The two adult Neanderthals date to the Late Middle Palaeolithic, basedon AMS AMS - Andrew Message System radiocarbon ages of Neanderthal 1 (sample Nea 1) of 39 900 [+ or-] 620 BP (ETH-209810) and Neanderthal 2 (sample NN1) of 39 240 [+ or -]670 BP (ETH-19660) (Bonani 2006). Very fragmentary faunal remains (c. 50000 pieces) were recovered from the 1997/2000 excavations, of whichapproximately 20 000 were burnt or charred (Feine et al. 2006). Thesefaunal remains were from both the Upper and Middle Palaeolithic levelsat the site. Feine et al. (2006) identified 14 bones withanthropogenetic alterations (including cut marks) that could beidentified to species from the Middle Palaeolithic level at the site.These were from various species, including horse, reindeer, cervids andprobably mammoth. Stable isotope measurement methods We took small samples of bone powder (c. 40mg) from both of theNeanderthals (both from right humeri) found at this site and wesuccessfully extracted small amounts of collagen from each followingmethods outlined in Richards and Hedges (1999) and Brown et al. (1988).We also sampled 12 animals from the more recent excavations. Only six ofthese samples yielded good quality collagen (based on the C:N ratios,collagen yields and %C and %N in collagen, following DeNiro 1985 andAmbrose 1990). As the remains are so fragmentary, it was not possible toidentify all of the herbivore herbivore:see carnivore. herbivoreAnimal adapted to subsist solely on plant tissues. Herbivores range from insects (e.g., aphids) to large mammals (e.g., elephants), but the term is most often applied to ungulates. faunal samples to individual species. Oneof the bone fragments was identified as red deer Red Deer, city, CanadaRed Deer,city (1991 pop. 58,134), S central Alta., Canada, on the Red Deer River. It developed as a trade and service center for a region of dairying and mixed farming. (Cervus elaphus) whilethe other three were only identified as either roe deer, red deer orreindeer. We were also able to sample red fox (Vulpes vulpes Vulpes vulpessee red fox. ) as well asa fish sample, but we were unable to identify the fish to its specificspecies. We have plotted the data from the two Neanderthals and thewell-preserved animal samples in Figure 1. The animal samples all comefrom the same layer as the Neanderthals with the exception of the fishwhich may have also come from this layer, but also may have come from anUpper Palaeolithic level at the site. Discussion The two Neanderthals have carbon isotope values ([delta][sup.13]C =-21.5 [per thousand], -21.6 [per thousand]) consistent with a diet ofprotein from terrestrial sources (plants or animals) with no evidence ofany measurable consumption of marine protein. The carbon isotope dataalso indicate that the Neanderthals did not consume measurable amountsof [C.sub.4] plants, or animals that consumed [C.sub.4] plants. This isunsurprising as there were almost no edible [C.sub.4] plants inprehistoric Europe This bulk of this article encompasses the time in Europe from c 900,000 years ago to 8th-7th century BCE. Pre-PleistoceneThrough most of Earth's history, various subcontinental land masses such as Baltica and Avalonia that would later be part of Europe moved about the globe . The carbon isotope values of all of the fauna arealso consistent with diets where the protein was from terrestrial([C.sub.3] plant-based) sources. The nitrogen isotope value of consumerbone collagen is 2-4 [per thousand] higher than the [delta][sup.15]Nvalue of dietary protein (i.e. plants or animal meat) (Schoeninger &DeNiro 1984). As can be seen in Figure 1, the herbivores have[delta][sup.15]N values ranging between 1.8 and 5.4 [per thousand]. Wesuspect, based on comparisons with published isotope values of otherherbivores in Europe (Iacumin et al. 2000; Bocherens et al. 2001;Drucker & Gambier gambier:see catechu. 2005; Richards et al. 2008) that the individualwith the highest [delta][sup.13]C (-18.1 [per thousand]) and lowest[delta][sup.15]N (1.8 [per thousand]) values is actually a reindeer,while the other three herbivore bone fragments are more likely to be roeor red deer. Reindeer are believed to have these relatively unusualisotope values due to the consumption of lichens (Beazley et al. 2002;Batts et al. 2004). The red fox is a carnivore carnivore(kär`nəvôr'), term commonly applied to any animal whose diet consists wholly or largely of animal matter. In animal systematics it refers to members of the mammalian order Carnivora (see Chordata). and consequently has ahigher [delta][sup.15]N value of 8.6 [per thousand]. The fox[delta][sup.15]N value is 3.8 [per thousand] higher than the average ofthe three cervid cervida member of the family Cervidae, deer, elk, reindeer, moose, wapiti, muntjacs and sikas. (roe and red deer, and excluding the single possiblereindeer) [delta][sup.15]N values of 4.8 [+ or -] 0.8 [per thousand].Therefore, the fox could have derived the majority of its dietaryprotein from herbivores like these cervids. As the possible reindeer hasa much lower [delta][sup.15]N value of 1.8 [per thousand] it is unlikelythat these species would have been prey for the red fox, as thedifference in [delta][sup.15]N value is too high (7 [per thousand]). Ifwe compare the Neanderthal [delta][sup.15]N values to the other mammal[delta][sup.15]N values we can see that they are most similar to the redfox, with values of 9.0 and 7.9 [per thousand]. Similarly, the cervidsare the most likely source of dietary protein for the Neanderthals, astheir [delta][sup.15]N values are 4.2 [per thousand] and 3.1 [perthousand] higher than the average of the three cervid samples. Thesingle fish sample we were able to measure has very different[delta][sup.13]C and [delta][sup.15]N values than the mammals. This isdue to the different sources of carbon in freshwater ecosystems asopposed to terrestrial systems, as well as reflecting the many trophic trophic/tro��phic/ (tro��fik) (trof��ik) pertaining to nutrition. troph��icadj.Of, relating to, or characterized by nutrition. levels in freshwater ecosystems, where piscivorous piscivorousfisheating; said of birds. fish often have[delta][sup.15]N values of 10 to 12 [per thousand] (Dufour et al. 1999).Based on this limited dataset we argue that the isotope values indicatethat the two Neanderthals were top-level carnivores at this site, andderived the majority of their dietary protein from terrestrial mammalherbivores, and freshwater fish and terrestrial plants were notsignificant dietary sources of protein. [FIGURE 1 OMITTED] Summary and conclusions We presented here the results of a palaeodietary isotopic analysisof two Neanderthals, including the type specimen and associated faunafrom the Kleine Feldhofer Grotte site in the Neander Valley in Germany.Our subsequent isotopic analysis showed that the Neanderthals, whencompared with a single carnivore and a number of herbivores from thesite, were top-level carnivores that derived the majority of theirprotein from herbivores, such as roe and red deer, but not reindeer. Wealso found that freshwater fish were not a source of dietary protein.Due to the history of the site, which was destroyed in themid-nineteenth century, very little faunal material has been recoveredfrom the site, and no plant remains. Therefore, isotopic analysis is theonly means of reconstructing the diet of these Neanderthals. Our isotopic results are remarkably similar to those of otherresearchers who measured the isotope values of other Neanderthals inEurope. These other isotopic studies focussed on Neanderthals fromeither central France (Les Pradelles (Marillac), Saint-Cesaire, LesRochers-de-Villeneuve and Jonzac), northern Western Europe (Spy, Engisand Scladina in Belgium) and southern Central Europe (Vindija, inCroatia). This is the first study of Neanderthals from northern CentralEurope. This was likely to be an area with a harsher climate thancentral France or Southern Central Europe, yet the isotopic studiesshowed that Neanderthals had the same subsistence adaptation in thiscolder environment as they did in areas which may conceivably have hadreadily available plant foods that could have been significant sourcesof protein for Neanderthals (i.e. hazelnuts). Therefore, the isotopicdata produced so far shows a remarkable similarity in Neanderthaladaptation throughout Europe in different time periods and in differentenvironments. In an earlier paper, Richards et al. (2000) argued thatthis meant that Neanderthals had to be sophisticated hunters, and notopportunistic scavengers, to be able to consistently obtain this amountof herbivore meat. This again supports the idea that, contrary toprevious views (e.g. Lieberman 1992; 2007) Neanderthals would have beenable to communicate effectively, most likely through speech, to plan andundertake systematic and frequent hunting. Our new isotopic resultspresented here as well as other recent studies support this contention,and reinforce the idea that Neanderthals were sophisticated hunters withan advanced ability to organise and communicate. Acknowledgements We would like to thank the Rheinisches Landesmuseum Bonn for thepermission to take samples from the Neanderthal and animal bones. Wewould also like to thank Annette Weiske and Stephanie Bosel forassistance with the isotopic measurements and Daniela C. Kalthoff(University of Bonn The University of Bonn (German: Rheinische Friedrich-Wilhelms-Universit?t Bonn) is a public research university located in Bonn, Germany. Founded in 1818 the University of Bonn is nowadays one of the largest universities in Germany. ) and Teresa Steele (Max Planck Noun 1. Max Planck - German physicist whose explanation of blackbody radiation in the context of quantized energy emissions initiated quantum theory (1858-1947)Max Karl Ernst Ludwig Planck, Planck Institute) foridentifying the animal remains. This research was funded by the MaxPlanck Society The Max-Planck-Gesellschaft zur F?rderung der Wissenschaften e. V. (abbreviated MPG, meaning Max Planck Society for the Advancement of Science) is an independent German non-profit research organization funded by the federal and state governments. . Received: 12 September 2007; Accepted: 3 December 2007; Revised: 29January 2008 References AMBROSE, S.H. 1990. 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SMITH, F.H., M.O. SMITH & R.W. SCHMITZ. 2006. Human skeletalremains from the 1997 and 2000 excavations of cave deposits derived fromKleine Feldhofer Grotte in the Neander Valley, Germany, in R.W. Schmitz(ed.) Neanderthal 1856-2006: 187-246. Mainz: Philipp von Zabern. TAUBER, H. 1981. [sup.13]C evidence for dietary habits ofprehistoric man in Denmark. Nature 292: 332-3. VAN DER MERWE, N.J. & J.C. VOGEL. 1978. [sup.13]C content ofhuman collagen as a measure of prehistoric diet in Woodland NorthAmerica. Nature 276: 815-16. VOGEL, J.C. & N. J. VAN DER MERWE. 1977. Isotopic evidence forearly maize cultivation in New York State. American Antiquity 42:238-42. Michael P. Richard (1) & Ralf W. Schmitz (2) (1) Department of Human Evolution, Max Planck Institute forEvolutionary Anthropology The Max Planck Institute for Evolutionary Anthropology is a research institute for evolutionary anthropology based in Leipzig, Germany founded in 1997. It is part of the Max Planck Institute network. The Institute currently employs three-hundred and thirty-four people. , Deutscher Platz 6, D-04103, Leipzig, Germanyand Department of Archaeology, Durham University, South Road Durham, DH13LE, UK (Email: Richards@eva.mpg.ed) (2) Institut fur Ur- und Fruhgeschichte und Archaologie desMittelalters, Abteilung Altere Urgeschichte und Quartarokologie,Universitat Tubingen, Schloss Hohentubingen, D-72070, Tubingen, Germany

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