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Research Exploring the Mesolithic and Neolithic transition in Croatia through isotopic investigations E. Lightfoot 1, B. Boneva 1 , P.T. Miracle 1 , M. ˇ Slaus 2 & T.C. O’Connell 3 The generalised picture of Mesolithic marine diet giving way to a Neolithic terrestrial diet, as derived from isotope measurements, has been both championed and challenged in this journal. Here new results from the Balkans offer a preliminary picture of a diversity of food strategy, both before and after the great transition. Keywords: Balkans, Croatia, Mesolithic, Neolithic, diet Introduction Isotopic evidence indicates that Mesolithic people in Europe tended to be heavily reliant on aquatic foods, while Neolithic subsistence was dominated by agricultural products (Tauber 1981; Richards & Hedges 1999; 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 crops, such as emmer wheat and barley; domesticated animals, including sheep, goats, cows, pigs and dogs; the use of pottery; and the establishment of permanent settlements (Whittle 1996). However, this suite of elements 1 Department of Archaeology, University of Cambridge, Downing Street, Cambridge, CB2 3DZ, UK 2 Croatian Academy of Sciences and Arts, Zrinski trg 11, 10000 Zagreb, Croatia 3 McDonald Institute for Archaeological Research, University of Cambridge, Downing Street, Cambridge, CB2 3ER, UK Author for correspondence (Email: [email protected]) Received: 2 December 2009; Accepted: 24 February 2010; Revised: 19 July 2010 ANTIQUITY 85 (2011): 73–86 http://antiquity.ac.uk/ant/085/ant0850073.htm 73
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Exploring the Mesolithic and Neolithictransition in Croatia through isotopicinvestigationsE. Lightfoot1∗, B. Boneva1, P.T. Miracle1, M. Slaus2

& T.C. O’Connell3

The generalised picture of Mesolithic marinediet giving way to a Neolithic terrestrial diet,as derived from isotope measurements, hasbeen both championed and challenged in thisjournal. Here new results from the Balkansoffer a preliminary picture of a diversity offood strategy, both before and after the greattransition.

Keywords: Balkans, Croatia, Mesolithic, Neolithic, diet

IntroductionIsotopic evidence indicates that Mesolithic people in Europe tended to be heavily reliant onaquatic foods, while Neolithic subsistence was dominated by agricultural products (Tauber1981; Richards & Hedges 1999; Schulting & Richards 2001; see also papers in Bailey &Spikins 2008 and Price 2000, respectively). The Neolithic ‘package’ consisted of farmingand non-farming elements, including domesticated crops, such as emmer wheat and barley;domesticated animals, including sheep, goats, cows, pigs and dogs; the use of pottery; andthe establishment of permanent settlements (Whittle 1996). However, this suite of elements

1 Department of Archaeology, University of Cambridge, Downing Street, Cambridge, CB2 3DZ, UK2 Croatian Academy of Sciences and Arts, Zrinski trg 11, 10000 Zagreb, Croatia3 McDonald Institute for Archaeological Research, University of Cambridge, Downing Street, Cambridge, CB2

3ER, UK∗ Author for correspondence (Email: [email protected])

Received: 2 December 2009; Accepted: 24 February 2010; Revised: 19 July 2010

ANTIQUITY 85 (2011): 73–86 http://antiquity.ac.uk/ant/085/ant0850073.htm

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was neither adopted in its entirety nor at the same rate in every region (Eriksson et al. 2008;Bocquet-Appel et al. 2009; Gkiasta et al. 2003; Forenbaher & Miracle 2005).

Various models have been proposed for the spread of agriculture from western Asia, mostfamously Ammerman and Cavalli-Sforza’s (1973) ‘wave-of-advance’ model. In this theory,agricultural communities have a demographic and dietary advantage over foraging groups,as farming can support a larger population per land unit (Bocquet-Appel 2002; Bellwood &Oxenham 2008). The strict application of this model calls for a near-complete populationreplacement throughout Europe. At the other end of the spectrum is the argument thatindigenous forager peoples adopted farming with little or no population movement (Dennell1983). Although in some areas one or other of these extreme models may be accurate, itseems most likely that in others a combination of the two occurred, with interaction betweenforager and farmer groups (e.g. Gregg 1988; Zvelebil & Lillie 2000; Robb & Miracle 2007).

However, it is becoming increasingly clear that the Mesolithic–Neolithic transition inEurope was heterogeneous in both its manner (e.g. Tringham 2000) and speed (e.g. Gkiastaet al. 2003; Forenbaher & Miracle 2005; Bocquet-Appel et al. 2009). Thus regional studiesexamining the diversity of responses to incoming subsistence regimes, technologies or peoplesare of growing significance (cf. Liden et al. 2004). The Balkan Peninsula is of particularimportance since farming is thought to have entered Europe via this region. Here we usetemporal and spatial comparisons of carbon and nitrogen stable isotopes from bone collagento investigate the dietary changes that took place during this transitional period in coastalareas of Croatia, and so assess the diversity in Neolithic adaptation.

The spread of the Neolithic in south-east Europe and CroatiaThe Balkans have traditionally been perceived as having a low Mesolithic population density,and having been rapidly colonised by agricultural immigrants (e.g. van Andel & Runnels1995). More recently, scholars have argued for the presence of foragers, with distributionmaps suggesting an aquatic adaptation (Boric 2005). Models for the spread of the Neolithicin the Balkans have therefore come to include 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 distinct ecological regions:the Adriatic coastal zone (Istria and Dalmatia), where agriculture was spread by seafarers(Forenbaher & Kaiser 2005); and the inland Pannonian basin, which received agriculturevia an overland route (Chapman & Muller 1990). On the Adriatic coast, the beginning ofagriculture is closely paralleled by the spread of Impressed Ware pottery (e.g. Chapman &Muller 1990) and population change seems to have played a large role in the transition tofarming (e.g. Biagri 2003). Other models suggest a scenario whereby there is a combinationof population movement and indigenous adoption (e.g. Forenbaher & Miracle 2005). Theearliest Neolithic in continental Croatia belongs to the Starcevo culture which appearssuddenly in c. 6000 cal BC (Minichreiter & Bronic 2006). Agriculture is thought to havereached this area via an overland route that spread north from Greece (Gkiasta et al. 2003;Bocquet-Appel et al. 2009). Juric et al. (2001) argue that as there is no evidence for a gradualdevelopment of agricultural methods, the arrival of farming must be part of a colonisationprocess.

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Macro-evidence for diet in coastal and inland CroatiaEvidence for Mesolithic diet from the Adriatic coast is limited as no archaeobotanicalevidence has been discovered (despite the intensive flotation programmes of recent projects)to suggest which plant foods may have been exploited. Nevertheless, the sites of Pupicina(Miracle 2001, 2002), Abri Sebrn (Miracle et al. 2000) and Vela Luka (Cecuk & Radic 2005)provide evidence for game hunting (red deer, roe deer, wild boar, fallow deer, hare, martenand fox), fishing (including deep water species such as tuna and dolphin) and gathering ofmarine and terrestrial molluscs.

More dietary evidence is available from the Neolithic sites along the Adriatic. Excavationsat Tinj-Podlivade (Chapman et al. 1996), Danilo Binj (Moore 2007), Grapceva Cave(Borojevic et al. 2008) and Bukovic-Lastivine (Chapman et al. 1996) have shown thatNeolithic diets included domestic crops: einkorn wheat, emmer wheat, free-threshing wheat,hulled barley, oats, flax and lentil; and domestic animals: sheep/goat, cattle and pig. Wildfoods were also utilised; acorn, juniper and almond were gathered, and red deer and harewere hunted. In addition, marine shells have also been found, but their food value is thoughtto have been modest (Schwartz 1988; Moore 2007).

A sequence of several cultures, the Starcevo, Sopot and Vucedol, is found in inlandCroatia throughout the period of the Neolithic, and all are thought to have been primarilyagricultural, growing legumes, einkorn and emmer wheat, and raising sheep, goat, cattleand pigs (Juric et al. 2001; Obelic et al. 2004). The choice of sites for settlement duringthe Starcevo period, however, suggests that they were chosen for their access to areas forhunting, for gathering fruits and nuts, and rivers for water and fishing, as well as foragricultural potential (Minichreiter 2001).

Isotopic evidence for dietStable carbon and nitrogen isotopic analyses of human remains can be used in conjunctionwith the evidence from archaeobotany and zooarchaeology to provide quantitativeinformation about past diets (Lee-Thorp 2008). As the skeleton is made from consumedfood and drink, isotopic ratios in the diet are retained in bone collagen. Due to the effects ofmetabolic processes, the proportion of 15N increases with each trophic level, and thereforestable nitrogen isotopic ratios provide an assessment of degrees of meat-eating (O’Connell& Hedges 1999). As both freshwater and marine ecosystems tend to have long foodchains,nitrogen stable isotopes can be used to distinguish between aquatic and terrestrial diets(Schoeninger & De Niro 1984). Carbon isotopic ratios allow for discrimination betweentwo types of plants (C3 and C4) with different mechanisms of carbon uptake (Vogel &Van der Merwe 1977), and between marine and terrestrial foodwebs (Schoeninger & DeNiro 1984). As the isotopic ratios at the base of the food chain vary through space andtime (Stevens & Hedges 2004), it is necessary to sample contemporaneous fauna whereverpossible.

The stable isotope composition of bone collagen reflects the diet over the majority of adultlife, thus indicating an ‘average’ diet (Hedges et al. 2007). Changes in diet between foodstuffsof similar isotopic composition will not produce a change in consumer bone collagen stable

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Figure 1. Map of Croatia showing analysed sites.

isotope ratios. Furthermore, physiological effects in the consumer (e.g. starvation: Mekota etal. 2006) and environmental influences on soil or plants can theoretically produce isotopicdifferences in bone collagen (e.g. Farquhar et al. 1982; Heaton 1987). Infants tend to havehigher δ15N values than adults due to a trophic level increase associated with breastfeeding(Fuller et al. 2006). In the discussion below, therefore, infants are excluded where discussingtypical population values.

In total, 42 humans and 95 animals were sampled from a range of sites across Croatia(Figure 1); however it should be noted that it is not certain that every human samplerepresents a different individual. The number of human individuals available per site issmall, as is common in these periods; nevertheless, when combined the samples represent areasonably sized dataset. The Neolithic human samples presented here span the whole of theNeolithic and include samples from two sites, Kargadur and Metljka, which are likely to beof Eneolithic date. The data analysed in this paper can be found in the online supplementwhich lists the location, period, radiocarbon date (where applicable) and number of humans

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Figure 2. Scatter graph of Mesolithic faunal carbon and nitrogen isotopic values.

sampled at each site, measurement for the animal and human samples and the results ofstatistical tests of significance1.

Faunal results: MesolithicFaunal isotope data are crucial for our interpretation of the human results that follow.Published results are available from Pupicina (Paine et al. 2009) and further data is presentedhere, largely from the site of Vela Luka-Vela Spilja (Korcula), with four samples (one hare,one roe deer, one wild boar and one fish) from Vela Spilja Losinj. These data are consistentwith the values expected for Europe (Figure 2). In general, omnivores have higher δ15Nvalues than herbivores. That the wild boar results are similar to those of the deer and twoof the hares suggests that they are eating a herbivorous diet. One hare sample (VSF032) hasan unexpectedly high δ13C result and is likely to have been misidentified; however there isinsufficient bone remaining to reassess the species identification. The fish samples have alower δ15N than would usually be expected, suggesting that these are low trophic level fish,although two samples have been tentatively identified as seabream, which is carnivorous.With the exception of VSF032, the fish and the dolphin samples are statistically differentfrom the terrestrial fauna in δ13C.

Faunal results: NeolithicThe Neolithic faunal values reported here are derived from three sites, Kargadur, Pupicinaand Vela Spilja Losinj. These data are consistent with the values expected for Europe (Hedgeset al. 2008; Figure 3). The sample sizes of the different species were too small to allow for

1 http://antiquity.ac.uk/projgall/lightfoot327/

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Figure 3. Scatter graph of Neolithic faunal carbon and nitrogen isotopic values.

statistical comparison between sites, but these results still represent a good dataset for overallcomparison with the human samples. As can be seen from Figure 3, herbivores are generallylower in δ15N than the foxes and fish. Pigs are statistically different to sheep/goat, cow anddeer in δ15N and thus appear to be more omnivorous, suggesting a different managementstrategy to the other domesticates. There is a clear and statistically significant separationof fish from the other fauna in both carbon and nitrogen isotopic values. This is due tomarine-terrestrial isotopic differences, and suggests that human consumption of fish shouldbe clearly evident in the isotopic results. The high δ15N results of Neolithic fish are in directcontrast to those from the Mesolithic, where the fish δ15N values are equivalent to thoseof the herbivores. Although sample size is limited, this may suggest that the Mesolithic andNeolithic people who lived at these sites exploited different marine resources.

Human resultsThere is substantial variation in both carbon and nitrogen isotopic values of all the humansanalysed (with ranges of 3.4‰ in δ13C and 6.0‰ in δ15N, or 3.8‰ in δ15N if infants areexcluded), suggesting that the individuals represented consumed varied diets.

When comparing the human carbon isotopic results in terms of their time period andlocation (Figures 4 & 5), the Mesolithic samples (all of which are from coastal locations)have the highest mean δ13C (−19.0‰ +− 0.5). Neolithic individuals from coastal sites havean intermediate mean δ13C, with a slightly wider range (−19.6‰ +− 0.7), whilst thoseNeolithic individuals from inland sites have the lowest mean δ13C (−20.3‰ +− 0.5). Thecoastal Mesolithic and Neolithic sites are statistically indistinguishable, while the inlandNeolithic results are different to both groups of coastal sites.

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Figure 4. Scatter graph showing human carbon and nitrogen isotopic results, separated by site.

Figure 5. Boxplot comparing human carbon isotopic results by period and location.

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Figure 6. Boxplot comparing human nitrogen isotopic results by period and location, excluding infants.

In terms of δ15N, a different pattern emerges (Figure 6). The Mesolithic individualshave a mean δ15N value similar to that of the inland Neolithic samples (10.0‰ +− 0.9and 10.3‰ +− 1.0 respectively), whilst the coastal Neolithic individuals have a mean δ15Nvalue which is somewhat lower (9.3‰ +− 1.0) and this is statistically different to the inlandNeolithic results only (P = 0.016). This pattern holds if infants are excluded. In the light ofour results showing that Mesolithic fish δ15N results are lower than those from the Neolithic,it seems odd that the Mesolithic humans have slightly higher δ15N than the coastal Neolithichumans. One explanation 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 the coast had a mixed dietof both marine and terrestrial protein. For the Neolithic people living inland, data showthat they consumed a terrestrial diet with no isotopic evidence for aquatic resources intheir diet. The coastal Neolithic people, however, had a slightly wider dietary range, whichwere generally of foods of low trophic level and sometimes included marine protein. TheNeolithic data cover the entire span of the Neolithic, including two potentially Eneolithicsites, but we see no simple trend in isotopic values with time. For all individuals analysed,the proportion of marine as opposed to terrestrial protein consumed on the Adriatic coastduring the Mesolithic and Neolithic was relatively low and terrestrial foodstuffs would haveconstituted an important source of protein.

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Figure 7. Scatter graph showing Vela Luka and Pupicina carbon and nitrogen isotopic results.

Whilst a direct overall comparison between the two periods is difficult due to the differingrange of site locations, our interpretation 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, thereis a clear difference between the Mesolithic individuals and the single Neolithic individual,an infant, in δ13C but not in δ15N (Figure 7). Although the small number of samples limitsthe interpretation, it can be tentatively suggested that there was a decrease in (low trophiclevel) marine protein consumption with the introduction of agriculture. From Pupicina, ourdata have been combined with results published in Paine et al. (2009) to increase samplesize. These results fall into two clusters relating to time period with clear differences inboth δ13C and δ15N. The sample numbers are again small, and it is not certain that eachsample represents a different individual, nevertheless the difference between the Mesolithicand Neolithic results suggests decreased fish consumption with the introduction of farmingto this site.

The range of variation in δ13C within and between sites for the Neolithic samples indicatesthat more diverse subsistence strategies were followed in the Neolithic than is often assumed(Figure 4). While most of the Neolithic individuals sampled have a terrestrial-based diet,individuals from the island sites of Losinj and Grapceva have higher δ13C, which reflects asmall amount of marine protein in their diets. At the coastal site of Metaljka, the range inδ13C values suggests that some individuals were consuming marine protein whereas otherswere eating a terrestrial diet, and at Kargadur only one sample, an infant, has a δ13C resultindicative of marine protein consumption. This could suggest that even within a site, peoplehad a certain degree of dietary choice, however we cannot be sure of the contemporaneityof these individuals due to a lack of secure dating.

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Discussion

The combination of marine and terrestrial resources can be seen elsewhere in the MesolithicMediterranean. At the Spanish site of El Collado (Guixe et al. 2006), isotope results suggest adiet of 25 per cent marine protein with the remainder procured from terrestrial resources. AtGrotta dell’Uzzo in Italy, finds of marine molluscs show that marine resources were utilised,however a clear marine signal is not seen in the human isotope results (Francalacci 1988).

Similarly the geographical pattern of Neolithic inland populations being dependent onterrestrial resources whilst Neolithic coastal dwellers consume some marine foods, is alsofound in the neighbouring areas of Slovenia, Malta and Greece. At Ajdovska jama in Slovenia,an inland Neolithic site, two studies have concluded that individuals consumed a mixedC3-based diet with no evidence for fish consumption (Ogrinc & Budja 2005; Bonsall et al.2007).

Results from coastal Neolithic sites are available from Italy, Malta and Greece. Fourindividuals from Arene Candide cave in Italy have δ13C values consistent with the terrestrialfauna, despite finds of marine mollusc shells (Francalacci 1988). Sixteen individuals wereanalysed from the Brochtorff Circle site in Malta, with isotopic results indicating noconsumption of marine resources (Richards et al. 2001; Stoddart et al. 2009). A largersample size is available from Greece, where terrestrial diets predominate at all Neolithic sitesanalysed so far, but at two of the four coastal sites there is evidence for small amounts ofmarine consumption (Papathanasiou et al. 2000; Papathanasiou 2003; Richards & Hedges2008).

In contrast, at the inland riverine sites along the Danube Gorges (Bonsall et al. 1997;Bonsall et al. 2004; Boric et al. 2004), isotopic evidence suggests that the diet of Mesolithicindividuals was mainly based 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 to the scarcity of excavatedhuman remains in this area. Nevertheless, the patterns seen here provide the first isotopicinsights into diet in these regions during the Mesolithic–Neolithic transition. Overall, ourresults indicate that while the Mesolithic people of the Adriatic coast consumed somemarine protein, terrestrial protein constituted a significant part of the diet. In the Neolithic,people in the Balkan Peninsula generally consumed terrestrial C3-based diets, likely to havebeen produced through farming. The Neolithic individuals from the Danube Gorges seemsomewhat atypical in their continued substantial utilisation of aquatic resources, but there isalso diversity in subsistence strategies at coastal Neolithic sites in both Croatia and Greece.

This is in stark contrast to the Mesolithic–Neolithic pattern seen on the Atlantic edge ofEurope, where isotopic results suggest that a strong reliance on marine resources gives way toa terrestrial diet in the Neolithic (Tauber 1981; Lubell et al. 1994; Richards & Mellars 1998;Richards & Hedges 1999; Schulting & Richards 2001, 2002; Richards et al. 2003). Guixeet al. (2006) suggest that the difference between the Atlantic and Mediterranean Mesolithicdiets could be related to either a different dietary adaptation or to the lower productivity ofthe Mediterranean compared to the Atlantic. These results must be seen in the light of anongoing debate about the ability of isotopic evidence to indicate low levels of marine resourceconsumption (Hedges 2004; Liden et al. 2004; Milner et al. 2004; Richards & Schulting

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2006). But data from this and previous studies suggest that there is a difference in theisotopic pattern between the Atlantic and Mediterranean across the Mesolithic–Neolithictransition, with implications for dietary composition during the transition to agriculture.

In general, the isotopic changes between the Mesolithic and Neolithic in the BalkanPeninsula fit with the expected pattern of decreasing aquatic resource use as agriculture isintroduced. However, the results presented here argue for diversity. In the Adriatic coastalzone, at least one-third of the analysed individuals consumed enough marine protein for itto register in their bone collagen isotopic values. The coastal individuals who appear to havehad a wholly terrestrial diet may have consumed a smaller amount of marine foods, as marineintake may not be isotopically detectable in diets that contain less than 20 per cent marineprotein (Hedges 2004; Milner et al. 2004). The intra-population variation at Kargadur andMetaljka shows that individuals had some scope for dietary choice, which is more likelyto be linked to economic organisation or personal preference rather than status differences(Forenbaher 2002).

It is, as yet, unclear why some people at coastal Neolithic sites utilised marine resourceswhile others did not. Isotopic analysis can give us an indication of the structure of theeconomy rather than directly informing us about the process of change. We can speculate,however, that our data may be indicative of population continuity across the Mesolithic–Neolithic transition, with a role for indigenous peoples in the origins of agriculture in thisarea. The diversity within sites could reflect individual movement between communities,as bone collagen reflects a long term dietary signal. It is also possible, however, thatthe utilisation of marine resources reflects local environmental conditions and resourceavailability, whether marine or terrestrial.

ConclusionThis discussion has shown that the Mesolithic–Neolithic transition cannot be seen as a simpledichotomy between people who used aquatic resources and people who did not. Naturally,some Neolithic coastal dwellers did exploit the sea. There is a growing consensus that wemust go beyond the oppositional views of foragers versus farmers across the Mesolithic–Neolithic transition, since the archaeological evidence is showing diversity in patterns ofbehaviour related to material culture within and between regions. Isotopic studies are nowshowing a similar range in patterns of behaviour related to subsistence and diet. This studyemphasises how crucial it is to examine the patterns derived from different lines of evidencein order to understand more fully this important process in human history.

AcknowledgementsThe authors are grateful to Catherine Kneale, Mike Hall and James Rolfe for their assistance with the isotopicanalysis, and to Darko Komso (Archaeological Museum of Istria), Staso Forenbaher (Institute of Anthropology,Zagreb) and Dinko Radic (Centre for Culture, Vela Luka) for providing samples. ORAU is thanked for providingtwo extracted collagen samples from Grapceva. The authors would like to thank Marc Vander Linden, CameronPetrie and Rhiannon Stevens for discussions of the manuscript. This work arises out EL’s PhD research (Universityof Cambridge), funded by the AHRC, and BB’s MPhil dissertation submitted to the University of Cambridge.

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Online supplement (see http://antiquity.ac.uk/projgall/lightfoot327/)Appendix 1: Table showing site information; Appendix 2: Table summarising statistical data;

Appendix 3: Table summarising the Mesolithic faunal results; Appendix 4: Table showing humanresults.

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