Exploring the Mesolithic and Neolithic transition in Croatia through isotopic investigations.

Exploring the Mesolithic and Neolithic transition in Croatia through isotopic investigations. [ILLUSTRATION OMITTED] Introduction Isotopic evidence indicates that Mesolithic people in Europe tendedto be heavily reliant on aquatic foods, while Neolithic subsistence wasdominated by agricultural products (Tauber 1981; Richards & Hedges1999; Schulting & Richards 2001; see also papers in Bailey &Spikins 2008 and Price 2000, respectively). The Neolithic'package' consisted of farming and non-farming elements,including domesticated do��mes��ti��cate?tr.v. do��mes��ti��cat��ed, do��mes��ti��cat��ing, do��mes��ti��cates1. To cause to feel comfortable at home; make domestic.2. To adopt or make fit for domestic use or life.3. a. crops, such as emmer wheat emmer wheat:see wheat. and barley;domesticated animals This article or section may contain original research or unverified claims.Please help Wikipedia by adding references. See the for details.This article has been tagged since September 2007.This is a list of animals which have been domesticated by humans. , including sheep, goats, cows, pigs and dogs; theuse of pottery; and the establishment of permanent settlements (Whittle1996). However, this suite of elements was neither adopted in itsentirety nor at the same rate in every region (Eriksson et al. 2008;Bocquet-Appel et al. 2009; Gkiasta et al. 2003; Forenbaher & Miracle2005). Various models have been proposed for the spread of agriculturefrom western Asia, most famously Ammerman and Cavalli-Sforza's(1973) 'wave-of-advance' model. In this theory, agriculturalcommunities have a demographic and dietary advantage over foraginggroups, as farming can support a larger population per land unit(Bocquet-Appe12002; Bellwood & Oxenham 2008). The strict applicationof this model calls for a near-complete population replacementthroughout Europe. At the other end of the spectrum is the argument thatindigenous forager peoples adopted farming with little or no populationmovement (Dennell 1983). Although in some areas one or other of theseextreme models may be accurate, it seems most likely that in others acombination of the two occurred, with interaction between forager andfarmer groups (e.g. Gregg 1988; Zvelebil & Lillie 2000; Robb &Miracle 2007). However, it is becoming increasingly clear that theMesolithic-Neolithic transition in Europe was heterogeneous in both itsmanner (e.g. Tringham 2000) and speed (e.g. Gkiasta et al. 2003;Forenbaher & Miracle 2005; Bocquet-Appel et al. 2009). Thus regionalstudies examining the diversity of responses to incoming subsistenceregimes, technologies or peoples are of growing significance (cf. Lidenet al. 2004). The Balkan Peninsula Balkan Peninsula,southeasternmost peninsula of Europe, c.200,000 sq mi (518,000 sq km), bounded by the Black Sea, Sea of Marmara, Aegean Sea, Mediterranean Sea, Ionian Sea, and Adriatic Sea. is of particular importance sincefarming is thought to have entered Europe via this region. Here we usetemporal and spatial comparisons of carbon and nitrogen stable isotopesfrom bone collagen to investigate the dietary changes that took placeduring this transitional period in coastal areas of Croatia, and soassess the diversity in Neolithic adaptation. The spread of the Neolithic in south-east Europe and Croatia The Balkans have traditionally been perceived as having a lowMesolithic population density, and having been rapidly colonised byagricultural immigrants (e.g. van Andel & Runnels 1995). Morerecently, scholars have argued for the presence of foragers, withdistribution maps suggesting an aquatic adaptation Several animal groups have undergone aquatic adaptation, going from being purely terrestrial animals to living at least part of the time in water. The adaptations in early speciation tend to develop as the animal ventures into water in order to find available food. (Boric bo��ric? also bo��rac��icadj.Of, relating to, derived from, or containing boron.boricAdjectiveof or containing boronAdj. 1. 2005). Modelsfor the spread of the Neolithic in the Balkans have therefore come toinclude a role for indigenous foragers, with a complex,'mosaic' pattern of transformation (Tringham 1971, 2000;Zvelebil & Lillie 2000). Croatia is particularly interesting as there are two main distinctecological regions: the Adriatic coastal zone (Istria and Dalmatia),where agriculture was spread by seafarers For Seafarers International Union and affiliates, see Seafarers International Union of North America. ''Note: This article title may be easily confused with The Seafarer. (Forenbaher & Kaiser2005); and the inland Pannonian basin The Pannonian Basin or Carpathian Basin is a large basin in Central Europe. The basin forms a topographically discrete unit set in the European landscape, surrounded by imposing geographic boundaries that have created a fairly unified cultural area that looks more towards , which received agriculture via anoverland route Overland Route or Overland Trail refers to the following travel routes: The Overland Trail (United States), the roughly parallel routes of the Overland Stage Line and First Transcontinental Railroad (Chapman & Muller 1990). On the Adriatic coast, thebeginning of agriculture is closely paralleled by the spread ofImpressed Ware pottery (e.g. Chapman & Muller 1990) and populationchange seems to have played a large role in the transition to farming(e.g. Biagri 2003). Other models suggest a scenario whereby there is acombination of population movement and indigenous adoption (e.g.Forenbaher & Miracle 2005). The earliest Neolithic in continentalCroatia belongs to the Starcevo culture which appears suddenly in c.6000 cal BC (Minichreiter & Bronic 2006). Agriculture is thought tohave reached this area via an overland route that spread north fromGreece (Gkiasta et al. 2003; Bocquet-Appel et al. 2009). Juric et al.(200 I) argue that as there is no evidence for a gradual development ofagricultural methods, the arrival of farming must be part of acolonisation process. Macro-evidence for diet in coastal and inland Croatia Evidence for Mesolithic diet from the Adriatic coast is limited asno archaeobotanical evidence has been discovered (despite the intensiveflotation programmes of recent projects) to suggest which plant foodsmay have been exploited. Nevertheless, the sites of Pupicina (Miracle2001, 2002), Abri Sebrn (Mirade et al. 2000) and Vela Luka Vela Luka is a town and municipality of Dubrovnik-Neretva County in western Croatia. Vela Luka is located on the western side of the island of Korcula at the bottom of a sharply cut and wide bay indented by many coves. (Cecuk &Radic 2005) provide evidence for game hunting (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. , roe deer, wildboar, fallow deer fallow deera small, 150 lb, fawn deer with white spots and a white spot bordered with black on each buttock. Called also Dama dama. , hare, marten marten,name for carnivorous, largely arboreal mammals (genus Martes) of the weasel family, widely distributed in North America, Europe, and central Asia. Martens are larger, heavier-bodied animals than weasels, with thick fur and bushy tails. and fox), fishing (including deep waterspecies such as tuna and dolphin) and gathering of marine andterrestrial molluscs. More dietary evidence is available from the Neolithic sites alongthe Adriatic. Excavations at Tinj-Podlivade (Chapman et al. 1996),Danilo Binj (Moore 2007), Grapceva Cave (Borojevic et al. 2008) andBukovic-Lastivine (Chapman et al. 1996) have shown that Neolithic dietsincluded domestic crops: einkorn wheat Einkorn wheat (from German Einkorn, literally "one grain") can refer either to the wild species of wheat, Triticum boeoticum (the spelling baeoticum is also common), or to the domesticated form, Triticum monococcum. , emmer wheat, free-threshingwheat, hulled barley, oats, flax and lentil lentil,leguminous Old World annual plant (Lens culinaris) with whitish or pale blue flowers. Its pods contain two greenish-brown or dark-colored seeds, also called lentils, which when fully ripe are ground into meal or used in soups and stews. ; and domestic animals:sheep/goat, cattle and pig. Wild foods were also utilised; acorn,juniper and almond were gathered, and red deer and hare were hunted. Inaddition, marine shells have also been found, but their food value isthought to have been modest (Schwartz 1988; Moore 2007). A sequence of several cultures, the Starcevo, Sopot and Vucedol, isfound in inland Croatia throughout the period of the Neolithic, and allare thought to have been primarily agricultural, growing legumes LegumesA family of plants that bear edible seeds in pods, including beans and peas.Mentioned in: Cholesterol, Highlegumes (l ,einkorn ein��korn?n.A one-seeded wheat (Triticum monococcum) grown in arid regions. Native to southwest Asia, it is one of the first crops to be domesticated by Neolithic peoples. and emmer wheat, and raising sheep, goat, cattle and pigs (Juricet al. 2001; Obelic et al. 2004). The choice of sites for settlementduring the Starcevo period, however, suggests that they were chosen fortheir access to areas for hunting, for gathering fruits and nuts, andrivers for water and fishing, as well as for agricultural potential(Minichreiter 2001). Isotopic evidence for diet Stable carbon and nitrogen isotopic analyses of human remains canbe used in conjunction with the evidence from archaeobotany andzooarchaeology to provide quantitative information about past diets(Lee-Thorp 2008). As the skeleton is made from consumed food and drink,isotopic ratios in the diet are retained in bone collagen. Due to theeffects of metabolic processes, the proportion of 15N increases witheach trophic level trophic leveln.A group of organisms that occupy the same position in a food chain.trophic level, and therefore stable nitrogen isotopic ratiosprovide an assessment of degrees of meat-eating (O'Connell &Hedges 1999). As both freshwater and marine ecosystems tend to have longfoodchains, nitrogen stable isotopes can be used to distinguish betweenaquatic and terrestrial diets (Schoeninger & De Niro Noun 1. De Niro - United States film actor who frequently plays tough characters (born 1943)Robert De Niro 1984). Carbonisotopic ratios allow for discrimination between two types of plants (C3and C4) with different mechanisms of carbon uptake (Vogel & Van derMerwe 1977), and between marine and terrestrial foodwebs (Schoeninger& De Niro 1984). As the isotopic ratios at the base of the foodchain vary through space and rime (Stevens & Hedges 2004), it isnecessary to sample contemporaneous fauna wherever possible. The stable isotope stable isotopen.An isotope of an element that shows no tendency to undergo radioactive breakdown. composition of bone collagen reflects the dietover the majority of adult life, thus indicating an 'average'diet (Hedges et al. 2007). Changes in diet between foodstuffs foodstuffsnpl → comestibles mplfoodstuffsnpl → denr��es fpl alimentairesfoodstuffsfood npl → of similarisotopic composition will not produce a change in consumer bone collagenstable isotope ratios. Furthermore, physiological effects in theconsumer (e.g. starvation: Mekota et al. 2006) and environmentalinfluences on soil or plants can theoretically produce isotopicdifferences in bone collagen (e.g. Farquhar et al. 1982; Heaton 1987).Infants tend to have higher [[delta].sup.15]N values than adults due toa trophic level increase associated with breastfeeding (Fuller et al.2006). In the discussion below, therefore, infants are excluded wherediscussing typical population values. [FIGURE 1 OMITTED] In total, 42 humans and 95 animals were sampled from a range ofsites across Croatia (Figure 1); however it should be noted that it isnot certain that every human sample represents a different individual.The number of human individuals available per site is small, as iscommon in these periods; nevertheless, when combined the samplesrepresent a reasonably sized dataset. The Neolithic human samplespresented here span the whole of the Neolithic and include samples fromtwo sites, Kargadur and Metljka, which are likely to be of Eneolithicdate. The data analysed in this paper can be found in the onlinesupplement which lists the location, period, radiocarbon date (whereapplicable) and number of humans sampled at each site, measurement forthe animal and human samples and the results of statistical tests ofsignificance (1). [FIGURE 2 OMITTED] Faunal results: Mesolithic Faunal isotope data are crucial for our interpretation of the humanresults that follow. Published results are available from Pupicina(Paine et al. 2009) and further data is presented here, largely from thesite of Vela velaplural of velum. Luka-Vela Spilja (Korcula), with four samples (one hare,one roe deer, one wild boar and one fish) from Vela Spilja Losinj. Thesedata are consistent with the values expected for Europe (Figure 2). Ingeneral, omnivores have higher [[delta].sup.15]N values than herbivores.That the wild boar results are similar to those of the deer and two ofthe hares suggests that they are eating a herbivorous herbivorous/her��biv��o��rous/ (her-biv��ah-rus) subsisting upon plants. diet. One haresample (VSF VSF - Virtual Software Factory 032) has an unexpectedly high [[delta].sup.13]C result and islikely to have been misidentified; however there is insufficient boneremaining to reassess the species identification. The fish samples havea lower [[delta].sup.15]N than would usually be expected, suggestingthat these are low trophic trophic/tro��phic/ (tro��fik) (trof��ik) pertaining to nutrition. troph��icadj.Of, relating to, or characterized by nutrition. leve1 fish, although two samples have beententatively identified as seabream, which is carnivorous car��niv��o��rous?adj.1. Of or relating to carnivores.2. Flesh-eating or predatory: a carnivorous bird.3. . With theexception of VSF032, the fish and the dolphin samples are statisticallydifferent from the terrestrial fauna in [[delta].sup.13]C. Faunal results: Neolithic The Neolithic faunal values reported here are derived from threesites, Kargadur, Pupicina and Vela Spilja Losinj. These data areconsistent with the values expected for Europe (Hedges et al. 2008;Figure 3). The sample sizes of the different species were too small toallow for statistical comparison between sites, but these results stillrepresent a good dataset for overall comparison with the human samples.As can be seen from Figure 3, herbivores are generally lower in[[delta].sup.15]N than the foxes and fish. Pigs are statisticallydifferent to sheep/goat, cow and deer in [[delta].sup.15]N and thusappear to be more omnivorous omnivorouseating both plant and animal foods. , suggesting a different management strategyto the other domesticates. There is a clear and statisticallysignificant separation of fish from the other fauna in both carbon andnitrogen isotopic values. This is due to marine-terrestrial isotopicdifferences, and suggests that human consumption offish off��ish?adj.Inclined to be distant and reserved; aloof.offish��ly adv.off should beclearly evident in the isotopic results. The high [[delta].sup.15]Nresults of Neolithic fish are in direct contrast to those from theMesolithic, where the fish [[delta].sup.15]N values are equivalent tothose of the herbivores. Although sample size is limited, this maysuggest that the Mesolithic and Neolithic people who lived at thesesites exploited different marine resources. [FIGURE 3 OMITTED] Human results There is substantial variation in both carbon and nitrogen isotopicvalues of all the humans analysed (with ranges of 3.4[per thousand] in[[delta].sup.13]C and 6.0[per thousand] in [[delta].sup.15]N, or 3.8[perthousand] in [[delta].sup.15]N if infants are excluded), suggesting thatthe individuals represented consumed varied diets. When comparing the human carbon isotopic results in terms of theirtime period and location (Figures 4 & 5), the Mesolithic samples(all of which are from coastal locations) have the highest mean[[delta].sup.13]C (-19.0[per thousand] [+ or -] 0.5). Neolithicindividuals from coastal sites have an intermediate mean[[delta].sup.13]C, with a slightly wider range (-19.6[per thousand] [+or -] 0.7), whilst those Neolithic individuals from inland sites havethe lowest mean [[delta].sup.13]C (-20.3[per thousand] [+ or -] 0.5).The coastal Mesolithic and Neolithic sites are statisticallyindistinguishable, while the inland Neolithic results are different toboth groups of coastal sites. [FIGURE 4 OMITTED] [FIGURE 5 OMITTED] [FIGURE 6 OMITTED] In terms of [[delta].sup.15]N, a different pattern emerges (Figure6). The Mesolithic individuals have a mean [[delta].sup.15]N valuesimilar to that of the inland Neolithic samples (10.0[per thousand] [+or -] 0.9 and 10.3[per thousand] [+ or -] 1.0 respectively), whilst thecoastal Neolithic individuals have a mean [[delta].sup.15]N value whichis somewhat lower (9.3[per thousand] [+ or -] 1.0) and this isstatistically different to the inland Neolithic results on our resultsshowing that Mesolithic fish [[delta].sup.15]N results are lower thanthose from the Neolithic, it seems odd that the Mesolithic humans haveslightly higher [[delta].sup.15]N than the coastal Neolithic humans. Oneexplanation for this may be the consumption of small amounts of marinepredators during the Mesolithic, such as dolphin. Our results indicate that Mesolithic individuals living on thecoast had a mixed diet of both marine and terrestrial protein. For theNeolithic people living inland, data show that they consumed aterrestrial diet with no isotopic evidence for aquatic resources intheir diet. The coastal Neolithic people, however, had a slightly widerdietary range, which were generally of foods of low trophic level andsometimes included marine protein. The Neolithic data cover the entirespan of the Neolithic, including two potentially Eneolithic sites, burwe see no simple trend in isotopic values with time. For all individualsanalysed, the proportion of marine as opposed to terrestrial proteinconsumed on the Adriatic coast during the Mesolithic and Neolithic wasrelatively low and terrestrial foodstuffs would have constituted animportant source of protein. [FIGURE 7 OMITTED] Whilst a direct overall comparison between the two periods isdifficult due to the differing range of site locations, ourinterpretation is borne out by the isotopic results of the two siteswith burials dating to both the Mesolithic and Neolithic. At Vela Luka,a coastal site, there is a clear difference between the Mesolithicindividuals and the single Neolithic individual, an infant, in[[delta].sup.13]C but not in [[delta].sup.15]N (Figure 7). Although thesmall number of samples limits the interpretation, it can be tentativelysuggested that there was a decrease in (low trophic level) marineprotein consumption with the introduction of agriculture. From Pupicina,our data have been combined with results published in Paine et al.(2009) to increase sample size. These results fall into two clustersrelating to relating torelate prep → concernantrelating torelate prep → bez��glich +gen, mit Bezug auf +acctime period with clear differences in both [[delta].sup.13]Cand [[delta].sup.15]N. The sample numbers are again small, and it is notcertain that each sample represents a different individual, neverthelessthe difference between the Mesolithic and Neolithic results suggestsdecreased fish consumption with the introduction of farming to thissite. The range of variation in [[delta].sup.13]C within and betweensites for the Neolithic samples indicates that more diverse subsistencestrategies were followed in the Neolithic than is often assumed (Figure4). While most of the Neolithic individuals sampled have aterrestrial-based diet, individuals from the island sites of Losinj andGrapceva have higher [[delta].sup.13]C, which reflects a small amount ofmarine protein in their diets. At the coastal site of Metaijka, therange in [[delta].sup.13]C values suggests that some individuals wereconsuming marine protein whereas others were eating a terrestrial diet,and at Kargadur only one sample, an infant, has a [[delta].sup.13]Cresult indicative of marine protein consumption. This could suggest thateven within a site, people had a certain degree of dietary choice,however we cannot be sure of the contemporaneity of these individualsdue to a lack of secure dating. Discussion The combination of marine and terrestrial resources can be seenelsewhere in the Mesolithic Mediterranean. At the Spanish site of ElCollado (Guixe et al. 2006), isotope results suggest a diet of 25 percent marine protein with the remainder procured from terrestrialresources. At Grotta dell'Uzzo in Italy, finds of marine molluscsshow that marine resources were utilised, however a clear marine signalis not seen in the human isotope results (Francalacci 1988). Similarly the geographical pattern of Neolithic inland populationsbeing dependent on terrestrial resources whilst Neolithic coastaldwellers consume some marine foods, is also found in the neighbouringareas of Slovenia, Malta and Greece. At Ajdovska jama in Slovenia, aninland Neolithic site, two studies have concluded that individualsconsumed a mixed [C.sub.3]-based diet with no evidence for fishconsumption (Ogrinc & Budja 2005; Bonsall et al. 2007). Results from coastal Neolithic sites are available from Italy,Malta and Greece. Four individuals from Arene Candide cave in Italy have[[delta].sup.13]C values consistent with the terrestrial fauna, despitefinds of marine mollusc molluscmembers of the phylum Mollusca, which comprises about 50,000 species. Includes snails, slugs and the aquatic molluscs��oysters, mussels, clams, cockles, arkshells, scallop, abalone, cuttlefish, squid. shells (Francalacci 1988). Sixteen individualswere analysed from the Brochtorff Circle site in Malta, with isotopicresults indicating no consumption of marine resources (Richards et al.2001; Stoddart et al. 2009). A larger sample size is available fromGreece, where terrestrial diets predominate at all Neolithic sitesanalysed so far, but at two of the four coastal sites there is evidencefor small amounts of marine consumption (Papathanasiou et al. 2000;Papathanasiou 2003; Richards & Hedges 2008). In contrast, at the inland riverine riv��er��ine?adj.1. Relating to or resembling a river.2. Located on or inhabiting the banks of a river; riparian: "Members of a riverine tribe ... sites along the Danube Gorges(Bonsall et al. 1997; Bonsall et al. 2004; Boric et al. 2004), isotopicevidence suggests that the diet of Mesolithic individuals was mainlybased on aquatic resources. Aquatic resource use persisted into theNeolithic, although the diet shifted towards terrestrial protein. The samples obtained for this study were small in number, due tothe scarcity of excavated human remains in this area. Nevertheless, thepatterns seen here provide the first isotopic insights into diet inthese regions during the Mesolithic--Neolithic transition. Overall, ourresults indicate that while the Mesolithic people of the Adriatic coastconsumed some marine protein, terrestrial protein constituted asignificant part of the diet. In the Neolithic, people in the BalkanPeninsula generally consumed terrestrial [C.sub.3]-based diets, likelyto have been produced through farming. The Neolithic individuals fromthe Danube Gorges seem somewhat atypical in their continued substantialutilisation of aquatic resources, but there is also diversity insubsistence strategies at coastal Neolithic sites in both Croatia andGreece. This is in stark contrast to the Mesolithic-Neolithic pattern seenon the Atlantic edge of Europe, where isotopic results suggest that astrong reliance on marine resources gives way to a terrestrial diet inthe Neolithic (Tauber 1981; Lubell et al. 1994; Richards & Mellars1998; Richards & Hedges 1999; Schulting & Richards 2001, 2002;Richards et al. 2003). Guixe et al. (2006) suggest that the differencebetween the Atlantic and Mediterranean Mesolithic diets could be relatedto either a different dietary adaptation or to the lower productivity ofthe Mediterranean compared to the Atlantic. These results must be seenin the light of an ongoing debate about the ability of isotopic evidenceto indicate low levels of marine resource consumption (Hedges 2004;Liden et al. 2004; Milner et al. 2004; Richards & Schutting 2006).But data from this and previous studies suggest that there is adifference in the isotopic pattern between the Atlantic andMediterranean across the Mesolithic-Neolithic transition, withimplications for dietary composition during the transition toagriculture. In general, the isotopic changes between the Mesolithic andNeolithic in the Balkan Peninsula fit with the expected pattern ofdecreasing aquatic resource use as agriculture is introduced. However,the results presented here argue for diversity. In the Adriatic coastalzone, at least one-third of the analysed individuals consumed enoughmarine protein for it to register in their bone collagen isotopicvalues. The coastal individuals who appear to have had a whollyterrestrial diet may have consumed a smaller amount of marine foods, asmarine intake may not be isotopically detectable in diets that containless than 20 per cent marine protein (Hedges 2004; Milner et al. 2004).The intra-population variation at Kargadur and Metaljka shows thatindividuals had some scope for dietary choice, which is more likely tobe linked to economic organisation or personal preference rather thanstatus differences (Forenbaher 2002). It is, as yet, unclear why some people at coastal Neolithic sitesutilised marine resources while others did not. Isotopic analysis cangive us an indication of the structure of the economy rather thandirectly informing us about the process of change. We can speculate,however, that our data may be indicative of population continuity acrossthe Mesolithic--Neolithic transition, with a role for indigenous peoplesin the origins of agriculture in this area. The diversity within sitescould reflect individual movement between communities, as bone collagenreflects a long term dietary signal. It is also possible, however, thatthe utilisation of marine resources reflects local environmentalconditions and resource availability, whether marine or terrestrial. Conclusion This discussion has shown that the Mesolithic-Neolithic transitioncannot be seen as a simple dichotomy between people who used aquaticresources and people who did not. Naturally, some Neolithic coastaldwellers did exploit the sea. There is a growing consensus that we mustgo beyond the oppositional views of foragers versus farmers across theMesolithic--Neolithic transition, since the archaeological evidence isshowing diversity in patterns of behaviour related to material culturewithin and between regions. Isotopic studies are now showing a similarrange in patterns of behaviour related to subsistence and diet. Thisstudy emphasises how crucial it is to examine the patterns derived fromdifferent lines of evidence in order to understand more fully thisimportant process in human history. Acknowledgements The authors are grateful to Catherine Kneale, Mike Hall and Jamesgolfe for their assistance with the isotopic analysis, and to DarkoKomso (Archaeological Museum of Istria), Staso Forenbaher (Institute ofAnthropology, Zagreb) and Dinko Radic (Centre for Culture, Vela Luka)for providing samples. ORAU ORAU Oak Ridge Associated Universities is thanked for providing two extractedcollagen samples from Grapceva. The authors would like to thank MarcVander Linden, Cameron Petrie and Rhiannon Stevens for discussions ofthe manuscript. This work arises out EL's PhD research (Universityof Cambridge), funded by the AHRC AHRC Asian Human Rights CommissionAHRC Arts and Humanities Research Council (UK)AHRC American Homeowners Resource CenterAHRC Army Human Resources CommandAHRC Association for the Help of Retarded Children , and BB's MPhil dissertationsubmitted to the University of Cambridge. Online supplement (seehttp://antiquity.ac.uk/projgall/lightfoot327/) Appendix 1: Table showing site information; Appendix 2: Tablesummarising statistical data; Appendix 3: Table summarising the Mesolithic faunal results;Appendix 4: Table showing human results. Received: 2 December 2009; Accepted: 24 February 2010; Revised: 19July 2010 References AMBROSE, S.H. 1990. Preparation and characterization of bone andtooth collagen for isotopic analysis. 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O'Connell [3] [1] Department of Archaeology, University of Cambridge, DowningStreet, Cambridge, CB2 3DZ, UK [2] Croatian Academy of Sciences and Arts The Croatian Academy of Sciences and Arts (Latin Academia Scientiarum et Artium Croatica, Croatian Hrvatska akademija znanosti i umjetnosti, abbrev. HAZU) is the national academy of Croatia. , Zrinski trg 11, 10000Zagreb, Croatia [3] McDonald Institute for Archaeological Research, University ofCambridge, Downing Street, Cambridge, CB2 3ER, UK * Author for correspondence (Email: ELFL2@cam.ac.uk)