Neolithic animal management practices andstable isotope studies in the AdriaticEmily Zavodny1, Sarah B. McClure1, Brendan J. Culleton1, Emil Podrug2,Douglas J. Kennett1

1Department of Anthropology, The Pennsylvania State University, University Park, PA, USA,2Šibenik City Museum, Šibenik, Croatia

We examine bone samples of known domesticates (sheep, goat, cattle and pig) from five open-air villagesites spanning most of the Neolithic period in Dalmatia, Croatia (cal 6000–4700 BC) to characterise dietsof domestic animals and address questions of the origin and development of animal husbandry strategiesin early farming communities. Carbon and nitrogen stable isotope values are analysed as proxies of dietand local environment that may indicate differences in herding and management practices betweendomesticated species. Results are compared to those reported for faunal remains found at other Neolithicsites from coastal Croatia and the wider Adriatic region. We find that isotopic values remain stable forcattle and ovicaprids during most of the Neolithic, suggesting that husbandry of these species remainedfundamentally the same throughout the period in much of the Adriatic. However, temporal differencesidentified among pigs indicate changes in associated management practices through time, and may be aresult of different foddering practices.

Keywords: Neolithic, Adriatic, Animal management, Domestication, Isotopes

IntroductionRecent excavations at key Neolithic sites on the centralDalmatian coast of Croatia provide insights into thedevelopment of animal management practices for themajority of the Neolithic, from early Impresso toearly Hvar cultures (cal 6000–4700 BC; Legge andMoore 2011; McClure 2013; Moore et al. 2007a, b;Podrug 2010). Excavations show that these earlyfarming societies practised a mixed agro-pastoral sub-sistence strategy, settled in dispersed villages, and uti-lised nearby natural caves and rock shelters.However, the ways in which animal husbandry strat-egies formed and changed as domesticated animalswere increasingly incorporated into local economiesremain unclear.Several stable isotope studies have begun to map

and characterise the spread of livestock managementpractices throughout the Adriatic and Balkans (Lelliet al. 2012; Lightfoot et al. 2011). Carbon and nitrogenstable isotopes are especially important for inferringchanges in diet and environment indicative of anthro-pogenic influence. This study uses stable carbon andnitrogen isotope analyses of faunal remains from fiveopen-air Neolithic sites in central Dalmatia toaddress differences in animal diet between species

and through time that may be linked to changes inanimal management practices along the Dalmatiancoast. Comparison of our results with published dataon domesticates from other Adriatic Neolithic sites(see Fig. 1; Lelli et al. 2012; Lightfoot et al. 2011;Ogrinc and Budja 2005) provides a framework forinterpreting management strategies on a regionalscale.

BackgroundStable isotope studies on domesticated faunal remainshave recently advanced our understanding of theorigin and spread of agriculture in theMediterranean world (Lelli et al. 2012; Lightfootet al. 2011). In the Adriatic region, ongoing archaeolo-gical studies focus on the introduction, adoption andadaptation of domesticates by early farming commu-nities and hunter–gatherer groups (Bass 2008; Lelliet al. 2012; Lightfoot et al. 2011; Miracle andForenbaher 2005, 2006). However, the precise timingof these transformative events are oft-debated, partlydue to a dearth of Mesolithic material in theAdriatic region (Biagi 2003; Lelli et al. 2012; Miracleand Forenbaher 2005; Moore et al. 2007a, b), withmost Mesolithic traces being found in caves and onlya few open-air localities identified (Komšo 2006a, b).The elusive Mesolithic occupation in centralDalmatia (Komšo 2006a, b; Miracle and Forenbaher

Correspondence to: Emily Zavodny, Department of Anthropology, ThePennsylvania State University, University Park, PA 16802, USA.Email: ekz5008@psu.edu

© Association for Environmental Archaeology 2014DOI 10.1179/1749631414Y.0000000021 Environmental Archaeology 2014 VOL. 0 NO. 0 1

2005), then, means the timing of theMesolithic–Neolithic transition is still not wellconstrained.Consequently, the advent of farming in Dalmatia

has been variably explained with diffusionist,migratory or native developmental models (e.g., Bass2008; Chapman et al. 1996; Forenbaher and Miracle2005; Marijanovic 2009; Mlekuž 2003; Moore et al.2007a, b). The appearance of a developed Neolithiclifeway at key village sites, such as Pokrovnik, suggeststo some that the spread of farming and husbandry wasrapid and complete (Legge and Moore 2011; Mooreet al. 2007a, b). Others believe that the spread wasmuch more gradual, slowly unfolding along theMediterranean coast and into the Adriatic (Miracleand Forenbaher 2005).Despite this ambiguity, however, current archaeolo-

gical surveys and excavations along the Dalmatiancoast are clarifying the temporal and spatial trajec-tories of the spread of farming and domesticates inthe area (Marijanovic 2009; Miracle and Forenbaher2006; Moore et al. 2007a, b). The earliest knownappearance of the Dalmatian Neolithic package,

which included domestic wheat, barley, sheep, goats,pigs and cattle, has now been pushed back to theearly 6th millennium cal BC (for new radiocarbondates of Dalmatian Neolithic, see Forenbaher andKaiser 2008; Forenbaher et al. 2013; Marijanovic2009; Miracle and Forenbaher 2006; Moore et al.2007a, b; Podrug 2010). As such, the Neolithic chron-ology of coastal Croatia is divided into three sub-periods associated with characteristic pottery styles:Early Neolithic or Impresso (cal 6000–5500 BC),Middle Neolithic or Danilo (cal 5500–4900 BC) andLate Neolithic or Hvar (cal 4900–4000 BC).

Neolithic animal management in centralDalmatiaQuantitative zooarchaeological studies of Neolithicassemblages have demonstrated the predominance ofdomesticated fauna at village sites (Legge andMoore 2011; McClure 2013; Miracle andForenbaher 2005; Moore et al. 2007a, b). At theEarly to Middle Neolithic site of Pokrovnik inDalmatia, for instance, total counts of identifiedbone show that domesticated animals were utilised

Figure 1 Map of Neolithic sites mentioned in the text.

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from the beginning of the site’s occupation (ca. cal6000 BC), and that hunting played a negligible partin Pokrovnik’s overall economy (Legge and Moore2011; Moore et al. 2007b). Similarly, the identifiedanimal bones at the Middle Neolithic site of Daniloare almost exclusively those of domesticated species(Legge and Moore 2011; Moore et al. 2007a).Studies of the seasonality of Neolithic slaughteringand birthing peaks demonstrate that herd size andcomposition were managed differently according tothe products desired (i.e. meat, dairy, wool; Balasseet al. 1997; Miracle and Pugsley 2006). Tooth eruptionand wear patterns are used to determine ages of death,and the resulting slaughter patterns are used as a proxyfor changing herd management strategies. When sheepare managed primarily for milk production, veryyoung lambs are slaughtered at a higher rate thanadults. This is in contrast to a strategy concentratedon meat, where larger numbers of animals survive tobe older juveniles or adults (e.g., Legge and Moore2011; Payne 1973; Vigne and Helmer 2007).However, kill-off models are only an approximationof animal management, since other factors such asaccidental mortality or disease affect mortality profiles(e.g., Legge andMoore 2011, 187) and mixed manage-ment strategies and statistical similarities betweensome strategies may complicate interpretations (e.g.,Bréhard et al. 2010; Brochier 2013; Vigne andHelmer 2007). Sheep age data from Pokrovniksuggest that these early farmers practiced a mixed ormeat-focused strategy (Legge and Moore 2011,187–188).Furthermore, seasonal transhumance is a histori-

cally well-documented pastoral adaptation in theMediterranean and in the case of central Dalmatiahas played a fundamental role in local farming adap-tations well into the 20th century (Moore et al.2007b). In this system, domestic stock is taken tomore distant pastures for part of the year, usually inthe summer months. Animals are able to graze onfresh grass and this allows the area surrounding eachvillage to regenerate from grazing activity. The roleof transhumance in the Neolithic, however, is lessclear. Archaeological data suggest that higherelevation and/or rockier terrain in the Adriaticregion were primarily used by pastoralists during theNeolithic (e.g., Istria, Slovenia; see Miracle 2006;Mlekuž 2003, 2005; see also Dennell 1978; Halstead2006). Elsewhere in the Balkans, sedentary pastoralistshave been identified in Romania (Greenfield andJongsma 2008), but transhumance as a pastoral strat-egy is not documented until the Bronze Age (Arnoldand Greenfield 2006). Recent ethnographic work atthe modern village of Pokrovnik in Dalmatia demon-strates the existence of a transhumant managementstrategy as recently as the past century, with herders

pasturing sheep in the nearby Dinaric Alps duringthe summer months and wintering in the coastalvalley alongside the village (Moore et al. 2007b). Wehypothesise that such seasonal rounds, if they existedin the Neolithic, likely followed the same spatialpattern, and transhumance was limited to sheep andgoats.Some integral aspects of livestock management, e.g.

foddering and grazing, are less visible through tra-ditional archaeological methods. Stable isotopestudies offer a systematic approach to mapping thesedifferent activities both temporally and geographi-cally, and have been applied successfully in a varietyof contexts (e.g., Bocherens et al. 2000, 2001;Makarewicz and Tuross 2006; Pearson et al. 2007).In this study, we use collagen δ13C and δ15N ofcattle, pigs and ovicaprids to determine whether differ-ences in husbandry are visible isotopically. We predictthat there will be isotopic variation between domesti-cated animals during the Neolithic because eachspecies was probably managed differently.

Stable isotopes, diet and managementStable isotopes of carbon and nitrogen in bone col-lagen can be used as proxies of diet, and reflect theaverage dietary protein during the last several yearsof an animal’s life (DeNiro and Epstein 1978, 1981).In addition, local environmental variation in basalproductivity and water availability, as well as meta-bolic factors, can alter the stable isotope compositionof bone and collagen (DeNiro and Epstein 1978,1981; Towers et al. 2011). Animal diet is inferredbased on the isotopic composition of food at thetrophic level of foods consumed. Anticipated differ-ences in stable isotopic values between species andover time can be attributed to changing patterns ofmobility, residence, and, by extension, managementstrategies during the Neolithic as communitiesinvested more time and energy into agriculturallifeways.On the basis of limited archaeological and ethno-

graphic data in Dalmatia, we hypothesise that ovica-prids participated in a seasonal pattern oftranshumance between the coast and mountainswhile cattle and pigs remained in coastal valleys nearpermanent settlements throughout the year (Nimac1940; Perišic 1940; Markovic 1987). Carbon and nitro-gen stable isotope values should reflect this differenceif the practice was indeed implemented during theNeolithic in coastal Croatia (see Fig. 2).Furthermore, as wheat and other C3 pathway plantsformed the bulk of the Dalmatian Neolithic package(Bailey 2000, 139), we expect all sampled domesticanimals to have a C3 carbon signature. Millet, a C4

plant, was presumably not in wide use by Neolithiccommunities in this region (Hunt et al. 2008),

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despite its presence at some sites (Legge and Moore2011; Moore et al. 2007b).

MaterialsThirty-four faunal skeletal elements were chosen fromthe archaeological assemblages of five excavated open-air Neolithic villages along the Dalmatian coast:Konjevrate, Pokrovnik, Danilo Bitinj, Krivace andCista Mala-Velištak (Fig. 1). Bones were identifiedto species and element by McClure at the Penn StateZooarchaeology Laboratory using comparativematerials. Differentiation between sheep and goatwas made according to the criteria set forth by Zederand Lapham (2010). We selected a representativesample of sheep, goat, cows and pigs, and each speci-men was measured, weighed and photographed.At each site, archaeological evidence of the intensive

use of domestic animals and crops suggests thathuman populations were dependent on agricultureand livestock throughout the Neolithic. Of the dom-estic animals associated with livestock herding andmanagement, sheep, goats, cattle and pigs are allpresent in the available faunal assemblages fromPokrovnik and Danilo Bitinj (Legge and Moore2011; McClure 2013; Moore et al. 2007a, b).Preliminary assessments of the material by Leggeand McClure1 indicate that numbers at Konjevrate,Krivace and Cista Mala-Velištak will be consistentwith these observations.Konjevrate is an Early Neolithic village located 4

km from Pokrovnik and is currently under a modernchurchyard (Fig. 1). The site was test excavated inthe 1990s and remains unpublished (for a shortdescription see Menđušic 1998). Although excavationwas not extensive, pottery, stone tools and animal

bones were recovered that are now curated by theŠibenik City Museum. The presence of Impressostyle pottery places the site in the Early Neolithic.Three samples of cow and sheep bone were includedin the analysis.

Pokrovnik, a village roughly 3 ha in size, was occu-pied continuously through the Early and MiddleNeolithic, and is located only a few kilometres fromthe Middle Neolithic site of Danilo Bitinj (Fig. 1;Moore et al. 2007b). Excavations were undertaken in1979 (Brusic 2008), and more recently in 2006(Moore et al. 2007b). Reported radiocarbon datessuggest the site was occupied ca. cal 5900–5100 BC(Legge and Moore 2011), making it one of the earliestdated open-air Neolithic villages in Dalmatia. Recentstudies have identified over 2400 animal bones, withover 90% from domesticates (Legge and Moore2011; Moore et al. 2007b). To take advantage of thetemporal breadth of the Pokrovnik assemblage, ninesamples were taken from Early Neolithic contextsand five samples from Middle Neolithic ones.

Located in a valley several kilometres from theAdriatic coast, the settlement of Danilo Bitinj wasoccupied during the Middle Neolithic, and isthought to have been one of the most extensive sitesof its type in southern Europe (Fig. 1; Moore et al.2007a). Past excavations in 1953, 1955 (Korošec1958, 1964) and 1992 (Menđušic 1998) provided awealth of material, although more recent excavationshave focused especially on the recovery and identifi-cation of over 1600 animal bones, most belonging todomesticates (Legge and Moore 2011; Moore et al.2007a). For this study, two sheep elements fromDanilo Bitinj were sampled.

Krivace is another Middle Neolithic village locatedin the Bribir valley (Fig. 1) that was first excavatedin 1963 (Korošec and Korošec 1974), again duringthe early 2000s (unpublished) and most recently in

Figure 2 Expected stable isotopic shifts according to different management strategies. Assuming free-ranging grazing orfoddering with C3 plants as a baseline, a switch to foddering with C4 plants will enrich δ13C by roughly 10–15‰ (DeNiro andEpstein 1978). Foddering with manured crops, either C3 or C4, will enrich δ15N by 1–3‰ (Bogaard et al. 2013). Animals fed withdomestic refuse will shift to a higher trophic level, enriching δ13C by 1–3‰ and δ15N by 3–5‰ (Schoeninger and DeNiro 1984).Grazing exclusively at higher arid elevations, such as in the Dinaric Alps, will enrich δ15N in relation to lowland grazing individuals(Ambrose 1991).

1Prof. Legge conducted a partial analysis of faunal material from thesesites prior to his untimely death and McClure revisited the assemblagesin 2013 to assess their suitability for future analyses.

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2013 (unpublished). A total of six elements fromexcavations in the 2000s was analysed for this study.Cista Mala-Velištak is currently the only excavated

Late Neolithic village in the region (Fig. 1; Podrug2010). The available radiocarbon dates (cal4900–4700 BC) place it firmly in the early phase ofthe Hvar culture (Podrug 2010). For this study, ninesamples of cattle, pig and sheep were analysed.

MethodsDalmatian samples were analysed using standard pro-cedures for collagen extraction at the Penn StateUniversity Human Palaeoecology Laboratory.Approximately 1700 mg of dry bone were taken fromeach archaeological sample, with compact bone pre-ferentially sampled to maximise collagen yield.Samples were crushed to increase the area of reactivesurface, then washed in NanoPure water and deminer-alised in 0·5 N HCl at 5°C. The demineralisationprocess varied from several days to several weeksdepending upon the sample.Skeletal samples were then prepared for collagen

extraction and purification by the modified Longin(1971) method with ultrafiltration (Brown et al.1988). All isotopic measurements were made on>30 kDa ultrafiltered bone collagen. 30 kDa

Centriprep® ultrafilters were thoroughly cleanedprior to ultrafiltration to remove contaminants (cf.Brock et al. 2007; Higham et al. 2006; Hüls et al.2007) according to protocols detailed in McClureet al. (2010). Carbon and nitrogen concentrationsand stable isotope ratios were analysed at ThePennsylvania State University Light IsotopeLaboratory with a Costech EA (ECS 4010), ThermoFinnigan Conflo IV gas handling device and aThermo Finnigan Delta V analyser. δ13C and δ15Nvalues are reported in standard ‰ notation withrespect to VPDB and atmospheric nitrogen, respect-ively. Sample quality was evaluated with %C, %Nand C:N ratios before further analysis. C:N ratiosfell between 3·0 and 3·26, reflecting good preservationfor isotopic analyses (see Table 1; DeNiro 1985; vanKlinken 1999).

ResultsThe results are presented in Table 1 and Fig. 3. At theoutset of this study, we expected to see isotopic differ-ences between species based on different managementstrategies thought to be practised in Neolithic centralDalmatia: ovicaprids participating in a seasonal patternof transhumance between the coast and mountains,and cattle and pigs remaining in the coastal valleys

Table 1 Results of Dalmatian stable isotope analyses

Site Period Sample # Species δ13C δ15N C:N

Konjevrate Early KON-2 Ovis aries −19·8 4·8 3·24Early KON-4 Ovis aries −19·4 6·4 3·20Early KON-5 Bos taurus −20·3 6·2 3·26

Pokrovnik Early PK-3 Ovis aries −20·6 5·7 3·19Early PK-4 Ovis aries −19·8 5·7 3·16Early PK-5 Ovis aries −20·4 6·0 3·18Early PK-7 Bos taurus −20·4 4·3 3·16Early PK-15 Ovis aries −20·1 5·3 3·17Early PK-19 Ovis aries −19·7 5·6 3·17Early PK-21 Ovis aries −20·4 5·2 3·18Early PK-22 Ovis aries −20·6 5·5 3·17Early PK-37 Bos taurus −20·0 5·3 3·17Middle PK-14 Bos taurus −19·7 5·6 3·23Middle PK-27 Sus scrofa −19·9 6·5 3·28Middle PK-31 Ovis aries −20·5 6·0 3·25Middle PK-36 Bos taurus −18·9 5·5 3·02Middle PK-39 Bos taurus −19·7 5·9 3·20

Danilo Middle DA-6 Ovis aries −19·0 5·7 3·21Middle DA-13 Ovis aries −17·4 5·3 3·21

Krivace Middle KRI-1 Ovicaprid −19·3 6·3 3·18Middle KRI-2 Sus scrofa −20·5 5·7 3·31Middle KRI-3 Bos taurus −20·4 4·0 3·12Middle KRI-9 Ovis aries −21·4 6·1 3·18Middle KRI-10 Bos taurus −19·9 4·6 3·16Middle KRI-11 Bos taurus −20·3 4·9 3·16

Cista Mala-Velistak Late CMV-1 Sus scrofa −20·6 5·4 3·19Late CMV-2 Bos taurus −19·5 6·3 3·19Late CMV-3A Bos taurus −18·8 5·8 3·14Late CMV-4 Sus scrofa −20·4 5·9 3·22Late CMV-5 Ovis aries −20·8 4·8 3·17Late CMV-6 Sus scrofa −19·8 7·0 3·19Late CMV-7 Bos taurus −20·0 4·8 3·15Late CMV-28 Capra hircus −20·0 5·0 3·21Late CMV-38 Capra hircus −19·9 5·0 3·23

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year round. However, carbon and nitrogen values for allspecies appear to reflect similar diets over time, asdemonstrated in Fig. 3. Carbon and nitrogen stableisotope values for each of the three time periodsoverlap closely. Early Neolithic samples have a δ13Crange of −20·6 to −19·4‰ and a δ15N range of4·3–6·4‰. Middle Neolithic samples have a δ13C rangeof −21·4 to −17·4‰ and a δ15N range of 4·0–6·5‰.Late Neolithic faunal samples display a δ13C range of−20·8 to −18·8‰ and a δ15N range of 4·8–7·0‰.When grouped by species (cattle, pigs and ovica-

prids), the overlap between carbon and nitrogenvalues remains with some subtle differences. Pigs inthe assemblage are slightly enriched in nitrogen, con-sistent with an omnivorous diet when compared tothe other domesticates. However, this enrichment isnot statistically significant (t-test assuming unequalvariances, Ruxton 2006; P= 0·08 (vs. cow) and 0·1(vs. ovicaprids)) in contrast to elsewhere on thecoast. In Istria, Lightfoot et al. (2011) report a signifi-cantly higher carbon signature for pigs, presumablybecause of a diet consisting of human waste andfood debris. This is not the case in our sample.Dalmatian pigs may have been foddered differentlythan cattle and ovicaprids, but it is difficult to deter-mine whether the observed variation in isotopicvalues reflects actual feeding differences or simplythe natural isotopic variation in regional vegetation.

Neolithic sites in the wider AdriaticSimilar ecological and environmental landscapesacross the Adriatic region allow us to contextualiseour results for Dalmatia with reported values for dom-esticated animals at other Neolithic sites in Istria(Croatia), Italy and Slovenia. We expect that animalmanagement strategies and their corresponding isoto-pic signatures will remain similar throughout the

circum-Adriatic during this time period. Isotopicvalues of domesticated animals reported from threedifferent human dietary studies are summarised inTable 2, and the background of each site is givenbelow.

IstriaTo see whether management strategies may have dif-fered along the eastern Adriatic coast, we compareour results for domesticated animals from Dalmatiansites to those carbon and nitrogen values reportedfor domesticates at Neolithic sites in the coastalIstrian region, located north of central Dalmatia(Fig. 1; Lightfoot et al. 2011). Lightfoot et al. (2011)focused on human dietary changes during theMesolithic–Neolithic transition, finding that therewas more dietary overlap between these periods thanoriginally thought. Three sites have comparable isoto-pic data for domesticated animals: Kargardur, VelaSpilja-Lošinj and Pupicina.

Kargardur is an open-air village site that is one ofthe most recently excavated of its kind in Istria(Komšo 2006b). Vela Spilja-Lošinj is a prehistoriccave site, located on the island of Lošinj off the coastof Istria. Excavations during the 1950s uncovered aMesolithic occupation (Mirosavljevic 1962, 1968,1974), and more recent studies have also focused onNeolithic components of the site, including the identi-fication of Early Impresso wares (Komšo et al. 2004).

Pupicina Cave was occupied numerous timesthroughout prehistory, and excavations have uncov-ered a significant Neolithic presence (Miracle andForenbaher 2006). Zooarchaeological analysis ofNeolithic animal remains have determined that dom-esticates comprised over 80% of identified taxathroughout the Neolithic (Miracle and Pugsley2006). Ovicaprids form the majority of these

Figure 3 Stable carbon and nitrogen isotope values for Dalmatian samples analyzed in this study.

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domesticated faunal remains, although the presence ofcattle and pigs increased in the Late Neolithic.Calculated survivorship curves for the faunal assem-blage suggest that ovicaprids were the focus ofmilking strategies during the Middle Neolithic, butthat they were exploited more for meat in the LateNeolithic along with cattle and pigs (Miracle andPugsley 2006).

SloveniaAjdovska Jama is an inland cave site in southeasternSlovenia that was infrequently occupied from thePaleolithic until the Middle Ages, although a seriesof archaeological excavations undertaken beginningin 1884 have uncovered a substantial Neolithic com-ponent (Bonsall et al. 2007; Horvat 1989). Earlyradiocarbon testing dated approximately 31 human

Table 2 Summary of stable isotope results for samples included in this study, organised by region

Region Site Period Sample # Species δ13C δ15N C:N References

Italy Basignano Early Bal III Ovis aries −20·0 6·2 3·4 Lelli et al. (2012)Grotta delle

MuraEarly GM 1 Ovicaprid −19·5 6·5 3·4 Lelli et al. (2012)Early GM 2 Ovicaprid −20·2 5·3 3·3Early GM 3 Ovicaprid −20·2 5·3 3·3Early GM 4 Ovicaprid −20·2 7·1 3·4Early GM 5 Ovicaprid −20·8 5·2 3·4Early GM 6 Ovicaprid −21·7 3·8 3·5Early GM 7 Ovicaprid −20·8 4·5 3·5Early GM 8 Ovicaprid −19·8 6·5 3·4

Palata Early Pal B5 Bos taurus −20·1 8·1 3·4 Lelli et al. (2012)Early Pal II fauna Ovicaprid −18·5 6·7 3·4Early Pal US7 Ovicaprid −20·4 6·7 3·4

Portonovo Early P21 Capra hircus −21·0 10·6 3·3 Lelli et al. (2012)Early P7 Ovis aries −24·1 7·8 3·3Early P13 Ovis aries −22·6 10·1 3·2Early P15 Ovis aries −21·2 6·9 3·3Early P18 Ovis aries −22·5 9·1 3·2Early P12 Sus scrofa −21·7 9·4 3·3Early P19 Sus scrofa −21·4 8·5 3·3Early P20 Sus scrofa −21·4 9·0 3·2Early P8 Sus scrofa −22·1 11·0 3·3Early P9 Sus scrofa −21·1 9·4 3·3

Ripa Tetta Early Rp 5 Sus scrofa −20·8 8·0 3·4 Lelli et al. (2012)Early Rp 6 Bos taurus −20·2 6·5 3·5Early Rp 7 Ovicaprid −20·7 8·0 3·3

Istria,Croatia

Kargadur Early BB13 Ovicaprid −20·9 7·1 3·2 Lightfoot et al. (2011)Early BB14 Ovis aries −16·8 8·6 3·1Early BB16 Ovis aries −20·8 5·6 3·2Early BB17 Ovicaprid −20·7 6·6 3·1Early BB18 Ovicaprid −20·5 6·8 3·1

Vela SpiljaLosinj

Early BB26 Sus scrofa −20·5 6·8 3·1 Lightfoot et al. (2011)Early BB29 Ovicaprid −19·2 6·5 3·1Early BB30 Ovicaprid −21·2 5·9 3·1Early BB34 Ovicaprid −20·2 6·1 3·1Early BB36 Ovicaprid −21·5 8·7 3·1Early BB39 Ovicaprid −20·1 9·6 3·1

Pupicina Middle BB50 Ovis aries −20·3 5·1 3·0 Lightfoot et al. (2011)Middle BB51 Ovis aries −20·1 5·1 3·1Middle BB52 Ovis aries −20·3 5·2 3·0Middle BB53 Ovis aries −20·3 5·6 3·0Middle BB54 Ovis aries −20·1 5·5 3·0Middle BB55 Capra hircus −19·0 5·8 3·0Middle BB56 Capra hircus −20·9 4·8 3·1Middle BB57 Bos taurus −21·2 4·8 3·2Middle BB58 Bos taurus −20·0 5·2 3·1Middle BB59 Bos taurus −20·5 5·0 3·1Middle BB62 Sus scrofa −19·6 7·1 3·1Middle BB63 Sus scrofa −19·9 7·0 3·1Middle BB64 Sus scrofa −19·2 7·8 3·0Middle BB65 Sus scrofa −19·3 4·8 3·2Middle BB66 Sus scrofa −19·2 7·9 3·1

Slovenia Ajdovskajama

Late Bos taurus −21·1 5·8 3·4 Ogrinc and Budja (2005)Late Bos taurus −20·4 6·7 3·4Late Bos taurus −21·7 5·8 3·4Late Bos taurus −20·4 5·8 3·2Late Bos taurus −20·0 6·0 3·2Late Ovis aries −19·6 7·0 3·2Late Sus scrofa −20·0 5·7 3·3

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burials to two periods within the Late Neolithic(Ogrinc and Budja 2005), although more recently pub-lished radiocarbon values for these same humanremains cluster between cal 3485 and 3340 BC(Bonsall et al. 2007, 732). Within the cave itself, dom-esticated sheep, goat and cattle bones, many with clearcutmarks, were found in burial contexts alongsidehearths containing carbonised grain. A recent palaeo-dietary study determined that Ajdovska Jama humansate a ‘terrestrial diet’ of mostly domestic animals andC3 plants (Ogrinc and Budja 2005), a conclusion thatwas echoed by Bonsall et al. (2007).

ItalyRecent research suggests that farming practices firstspread along the eastern Adriatic coast, crossing thesea to Italy by the start of the 6th millennium BC(Lelli et al. 2012; Miracle and Forenbaher 2005;Skeates 2000; Starnini 2002). Like central Dalmatia,southeastern Italy also exhibits little evidence of astrong Mesolithic tradition (Biagi 2003; Lelli et al.2012), making it an ideal point of comparison formapping possible introductions and changes inanimal husbandry among first farmers along theAdriatic coast.For this study, we include stable carbon and nitro-

gen isotope values reported for domestic animalsfrom five Early Neolithic village sites by Lelli et al.(2012; see Fig. 1) as part of a human palaeodietarystudy. Radiocarbon dates for four of the villages indi-cate a Neolithic date of these sites Ripa Tetta (cal5860–5600 BC), Palata (cal 5620–5470 BC),Balsignano (cal 5570–5480 BC) and Portonovo (cal5480–5310 BC). Impresso pottery associated withdomesticated fauna are used to date the fifth village,Grotta delle Mura, to the Early Neolithic as well.All of these sites are open-air settlements that were sur-rounded by ditch structures and whose inhabitantsparticipated in an early agricultural economy.Although the majority of the Italian sites cluster inthe southeastern region of the Italian peninsula (seeFig. 1), Portonovo is located along the coast muchfarther north and approximates the same latitudes asthe central Dalmatian sites used in this study.Collectively, these sites offer an opportunity to detectpossible management differences on either side of theAdriatic.

DiscussionFig. 4 illustrates the combined stable carbon and nitro-gen values from our study of central Dalmatia andthose reported from contemporary sites in Italy,Slovenia and Istria.Despite our initial predictions of temporal and

species-specific changes in stable isotope signaturesthroughout the Neolithic period, Fig. 4 demonstrates

that such differences are not actually present overtime or by region. The apparent outliers in theupper left corner of the graph are all early Italian ovi-caprids, cattle and pigs from the northern Italiancoastal site of Portonovo. Samples reported forPortonovo are markedly 15N enriched in comparisonto all other regions, species and time periods. Higherδ15N could signal a unique Neolithic diet forPortonovo’s livestock, such as the inclusion of moremarine foods or plants less able to fix molecularnitrogen (DeNiro and Epstein 1981). This nitrogensignature could also be a result of nursing, as sucklinganimals are enriched over their mothers because theyare feeding at a higher trophic level (Nehlich et al.2009; Richards et al. 2002). The original investigators(Lelli et al. 2012) did not report ages for Portonovospecimens sampled for stable carbon and nitrogenisotopes, so it is possible that Portonovo had ahigher number of young animals than was collectedfrom other Neolithic sites. However, the differencein nitrogen between Portonovo and the rest of thesamples is most likely due to some unknown yetdifferent foddering or management strategy employedby farmers in this village, as proximity to the sea,environment and climate at this site are not thoughtto differ from contemporary coastal sites inDalmatia or Istria.

Disregarding Portonovo samples, then, we see atight cluster of points, suggesting that cattle, ovicapridsand pigs had largely similar diets throughout theNeolithic period regardless of region (Slovenia,Istria, Italy or Dalmatia). A more uniform diet spacedoes not necessarily mean that the spatial patterningof animal management remained the same throughtime or was the same for all species. Very little isknown about the degree of uniformity in vegetationin the region and to what extent coastal valleys mayhave been preferred transhumance routes as opposedto transhumance to inland areas during theNeolithic. Another explanation for similar diet spacebetween species is foddering. In this case farmersprovide animals with much of their subsistence, creat-ing a very different kind of animal management systemthat may well have a similar isotopic signature. Inshort, current stable isotope data support the notionthat animal management was largely stable through-out the Neolithic, but fall short in assessing thedegree of transhumance during this period for ovica-prids and bovines.

A closer look at isotope values reported for pigs,however, does reveal some differences for the speciesacross time and space independent of those individualssampled from Portonovo. Pigs are present in allregions for all time periods (Figs. 4 and 5), and thereappears to be a signal of changing management strat-egies between the Early and Middle–Late Neolithic.

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Early Neolithic pigs from all of Italy, not justPortonovo, and Istria show a markedly differentlinear regression trendline than those for Middle(Istria, Dalmatia) and Late (Slovenia, Dalmatia)Neolithic pigs (Fig. 5). Analysis of covariancebetween the regression lines of Early andMiddle–Late Neolithic pigs indicates a significantdifference in intercepts (P= 0·038) and slopes (P=0·038) (see Fig. 5). Statistical analyses show thatthere is a significant difference in stable carbonvalues between Early and Middle Neolithic pigs(t-test assuming unequal variances, Ruxton 2006;P= 0·0002) and Early and Late Neolithic pigs (P=0·003). Additionally, significant differences in stable

nitrogen values between Early and Middle (P=0·011) and Early and Late Neolithic pigs (P= 0·001)also exist. Carbon and nitrogen values are not signifi-cantly different between Middle and Late Neolithicpigs (P= 0·122 and 0·068, respectively). We suggestthat these differences in stable carbon and nitrogen sig-natures may reflect a shift in the foddering or manage-ment strategy of pigs between Early and Middle–LateNeolithic times throughout the Adriatic.Higher δ15N in pigs may reflect increased manuring

practices in fields (Bogaard et al. 2013; Madgwicket al. 2012) or different climatic and environmental set-tings (Madgwick et al. 2012). However, taking intoaccount the remarkably high 15N enrichment for

Figure 4 Stable carbon and nitrogen isotope values for all samples discussed in this paper (this study; Lelli et al. 2012; Lightfootet al. 2011; Ogrinc and Budja 2005).

Figure 5 Stable carbon and nitrogen isotope values of all pigs discussed in this paper (this study; Lelli et al. 2012; Lightfoot et al.2011; Ogrinc and Budja 2005).

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Portonovo livestock and the prevalence of thesesamples in the Early Neolithic pool, we find that nitro-gen values are not statistically significantly differentbetween Early and Middle–Late Neolithic pigs oncePortonovo values are removed from the sample.Conversely, carbon values for the same time periods

remain statistically significantly different within theAdriatic with or without the Portonovo samples andmay signal a change in livestock strategy. Evidencefrom Neolithic sites indicates that early agriculturalstaples were wheat, primarily einkorn and emmer,barley and legumes – all C3 pathway plants (Bailey2000, 139). A later enrichment in carbon during theNeolithic may have resulted from a shift in diet,from one based primarily on C3 plant foods to oneincorporating small amounts of C4 species. Panicummiliaceum (broomcorn millet) is the likely C4 candi-date, having been recovered from archaeological con-texts in Europe prior to cal 5000 BC (Hunt et al.2008), including Middle Neolithic contexts atPokrovnik in Dalmatia (Legge and Moore 2011;Moore et al. 2007b) and at sites in neighboringSerbia (Gomolava, ca. cal 3700–3600 BC). However,just across the Adriatic, varieties of millet were notknown in northern Italy until the Early Bronze Age(ca. cal 1700–1500 BC, Tafuri et al. 2009; Zoharyand Hopf 2000) or in southern Italy until classicaltimes (Tafuri et al. 2009). While it is possible thatMiddle and Late Neolithic sites in Dalmatia andIstria obtained and utilised domestic millet, eitherfor feeding animals or as part of the human diet, thespread and adoption of this millet species in southeast-ern Europe remains unclear and unequivocal evidencefor the conscious adoption of millet as a cultivar hasnot yet been found. The results of our study canneither support nor refute the clear presence of milletin the Adriatic Neolithic.Statistical tests performed on δ13C and δ15N for

cattle and ovicaprids during each Neolithic periodfound no such difference for either carbon or nitrogenstable isotopes. Thus, we hypothesise that pigs mayhave begun to be treated differently by early farmersas the Neolithic unfolded, whereas sheep, goats andcattle were managed with similar strategies throughoutthe Neolithic.

ConclusionDespite previous assumptions, we find that the isoto-pic values reported in this paper remain largelystable for ovicaprids and cattle over the majority ofthe Neolithic, suggesting that livestock husbandryfor these species remained fundamentally the samethroughout the period in much of the Adriatic.Domesticated pigs, however, may have been foddereddifferently as the Neolithic period progressed, depend-ing on available plant resources. In order to explore

this trend, as well as others suggested in this paper,future studies should focus on expanding samplesizes for known faunal domesticates at both sitesreported here and other Neolithic occupationsthroughout the circum-Adriatic region. Additionally,there is a need for compiled isotopic information onvegetation, either wild or domesticated, that mayhave been used for foddering or grazing by earlyfarmers. Our results demonstrate the utility of isotopicstudies in studying important questions surroundingthe neolithisation of the Adriatic, and highlight theneed to continue quantitative scientific studies in theregion.

AcknowledgementsSpecial thanks to Marko Menđušic, Andrew Mooreand Joško Zaninovic for access to Dalmatiansamples, Karen Trippett and Jennifer Trippett forhelp with sample processing, and two anonymousreviewers for their insightful comments. Funding wasprovided by The Pennsylvania State University.

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