First introduction and early exploitation of the Persian fallow deer in Cyprus (8,000-6,000 cal. Bc)








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First introduction and early exploitation of the Persian fallow deer in Cyprus (8,000-6,000 cal. BC)
Vigne Jean-Denis, Daujat Julie and Monchot Hervé
Supporting information
Appendix S1: Brief summary of Cyprus Prehistory – Shillourokambos and Khirokitia

The earliest evidence for the presence of humans on Cyprus comes from the upper layer of the small rock shelter of Akrotiri-Aetokremnos. This site, dated to c. 10,500 cal. BC, is contemporaneous with the Late Natufian or the Khiamian in the Levant. It is characterized by a rough lithic industry, associated with fish and shellfish remains, bird bones and a small wild boar (Sus scrofa ssp.) (Simmons, 1999; Vigne et al., 2009). There are no data for the 10th millennium. The open-air sites at Aya Varvara-Asprokremnos and Ayios Tychonas-Klimonas indicate that human groups belonging to the PPNA koine were living on Cyprus during the first half of the 9th millennium cal. BC (Guilaine & Briois, 2007; Manning et al., 2010; Vigne et al., 2011b ). They exploited small wild boar, the only large mammal present on the island at that time, as well as a dog (Vigne et al., 2011a,b). Archaeological and archaeozoological data become more frequent from 8,400-8,300 cal. BC onwards, with the two wells of Kissonerga-Mylouthkia (Peltenburgh, 2003) and, especially, the large open-air site of Parekklisha-Shillourokambos, where an area of over 5,000 m2 was excavated.

The long lasting occupation of Area 1 of Shillourokambos can be divided into four main phases, for which archaeological and archaeozoological analyses have recently been published in detail (Guilaine, 2003; Guilaine & Briois, 2007; Guilaine et al., 2011; Vigne et al., 2011a ), and can be summarized as follows:

  1. Early A (8,400-8,100 cal. BC), contemporaneous with the Early PPNB in Southeast Anatolia and the Northern Levant; cultivation of cereals and legumes is attested; the rather small sample of animal bones (NISP=282) is characterized by the prevalence of the small wild boar – already attested during the preceding periods, as well as the presence of small dogs and the first occurrence of the domestic goat (Capra aegagrus/hircus) and cattle (Bos taurus);

  2. After a short abandonment of the site – or a gap in the deposits due to erosion, the Early phases B and C (7,900-7,600 cal. BC) are contemporaneous with the Middle PPNB; here again, agriculture is attested; archaeozoological samples (NISP=1586+763) indicate a decrease in the predominance of suids, and the first occurrence of domestic sheep (Ovis aries), red fox (Vulpes vulpes) and Pfd; animal supply relied upon sheep husbandry (i.e., milk and meat), pig and cattle, and hunting of feral goats, wild boars and Pfd; dogs rapidly became very rare;

  3. Middle phases (7,600-7,400 cal. BC) are divided into at least two (A and B) sub-phases; multiple changes in the material culture and in the economic system are evidenced; after the collapse of sheep herding, new sheep lineages were introduced for meat production and the Cypriot feral goats were locally domesticated (Vigne et al., 2015 ); cattle drastically decreased in importance whilst pig breeding became more important (NISP=4707+942);

  4. Late phase (7,400-7,000 cal. BC) is contemporaneous with the Late PPNB on the continental mainland; archaeozoological data (NISP=471) indicate a specialization in sheep and goat herding, the former for meat and the latter for milk.


Numerous 7th and early 6th millennium Cypriot villages were contemporaneous with the PPNC/pre-Halaf/Halaf cultures on the continental mainland. They belong to the Aceramic culture of Khirokitia (Le Brun & Daune-Le Brun, 2003, 2009). The latter is a large village containing hundreds of thousands of animal bones, dominated by domestic sheep, goat and pig, as well as Pfd; cattle were absent and dogs remained very rare (Davis, 1984, 1994, 2003). The following periods (Pottery Neolithic of the Sotira period, Chalcolithic and Bronze Age) are not taken into consideration in this paper; research by one of the authors (JD) into the study of the Pfd during these periods is ongoing.
Method S1: Geographic distribution of the Pfd during the PPN (material and methods)

The geographic distribution of Dama in the northeast Mediterranean area and the Near East during the Early Holocene presented in this paper (Figure 1) and supporting information (see below Figure S1 and Tables S1 & S2), is based on fallow deer remains from sites listed in Conolly et al. (2011). These data were extracted from the OSSK database, to which 14 sites were added (n. 20, 23, 27, 29, 35, 66, 97, 98, 123 to 129). This represents a total of 129 sites, 198 contexts and more than 16,796 Dama remains NISP. B. Stopp extracted Dama data from the database, which are listed according to three categories of identification: D. d. dama, D. d. mesopotamica and D. d. ssp. D. dama ssp. was attributed to one of the two subspecies according to location. Attributions to one subspecies or another were corrected for several sites as follows: D. d. dama into D. d. mesopotamica for sites n. 20, 25, 43, 48, 99, 101 and 102; D. d. mesopotamica into D. d. dama for sites n. 10, 24, 35 and 123.

According to the cultural (after Aurenche & Kozlowski, 1999) and geographic regions, 11 areas were delimitated: 1, Greece; 2, Western and Central Anatolia; 3, Zagros; 4, Djezireh; 5, Middle Euphrates and “High valleys”; 6, Northern Levant and Adana Plain; 7, Central Levant; 8, Southern Levant; 9, Sinai; 10, Syro-Jordanian desert; 11, Cyprus. On Cyprus, contexts prior to the introduction of Dama, i.e., Akrotiri, Klimonas, Asprokremnos, Mylouthkia well 116 and Shillourokambos Early A phase, were taken into consideration.
Method S2: Protocol for the analysis of the morphology, sex ratios, evolution of size and comparisons

Mixture analysis in PAST© version 2.12 (Hammer et al., 2001) were used in this study. It is based on the Expectation-maximization (EM) algorithm of Dempster et al. (1977). The procedure is automatically run 20 times, each time with new, random starting positions for the mean. The Akaike Information Criterion (AIC) is calculated with a small sample correction (Akaike, 1974). The minimum value of the AIC is often obtained for a high number of groups. Mixture analyses with the lowest AIC values – or at least close to the lowest values, were considered in this study.

Each measurement with sufficient data was analysed as follows – slightly improved protocol with reference to Vigne (2011a):

  1. The normality of the entire distribution of each measurement is tested using the Shapiro-Wilk test, in order to detect – and eliminate – small datasets and/or variables that are not or poorly influenced by sexual dimorphism;

  2. When a sample is large enough and the Shapiro-Wilk test is negative (p<0.9), a repeated series of mixture analysis (bi-normal distribution) is computed and the most accurate model is retained based on both the congruence with the Kernel distribution of density of the data (Figure 2B), and the lowest value of AIC – at this step variables which produced unstable or models with two (or more) entirely overlapping Gaussian profiles are excluded;

  3. When the bimodal partition is stable and the structure of the AIC low enough, the mean and standard deviation (std) are estimated for the two Gaussian profiles – i.e., both sexes (Figure 2B);

  4. For each measurement with more than 20 data, the sex ratio is estimated based on the probability computed for each of the two Gaussian profiles and compared to the null hypothesis (males=females) using a z test,

  5. Sexual dimorphism is estimated using the following index (ISD): for each i measurement ISDi=(XiMales-XiFemales)/X(XiMales+XiFemales), where X is the mean;

  6. The mean size of each sex is compared separately between phases or sites using a z test – sample size for both males and females is calculated as follows: for each i measurement Ni=ni pi where N is the sample size, n is the number of data and p the probability given by the mixture analysis.


Method S3: Age profiles and slaughtering strategies (method & detail profiles by phase)

Lower cheek teeth are mostly used for the construction and statistical processing of mortality profiles. Upper molars are only exceptionally used if it was not possible to match them with any mandibles – in terms of minimal individuals. Isolated first and second molars are separated based on their measurements – breadth and length. Slaughter profiles are estimated based either on the dental MNI (Early B and C, Middle B phases) or the total number of teeth (isolated or in cheek row; Middle A phase). Unlike in Vigne (2011: 992-998), frequencies were not corrected, except for the construction of density histograms, following Brochier (2013). Survival profiles are deduced from non-corrected age frequencies starting at a 100% population, and postulating that humans were the only cause of mortality – in the absence of any other large predator on Cyprus, apart from dogs. Survival profiles obtained for Shillourokambos are compared with the survival profile of small red deer from the Scottish Isle of Rùm, in the Inner Hebrides (Clutton-Brock et al., 1982).

Age profiles of the Early B phase at Shillourokambos (Figure S4A)

Age profiles of the Early B phase are based on 51 dental data representing 13 individuals, and on 417 epiphyseal fusion data. The latter also include the Early C phase. Individuals less than two years old are nearly absent from dental records. This may be partly due to the bad preservation of young teeth, since 14% of animals were culled at less than two years old according to the epiphyseal fusion profile. Slaughtering targeted mostly young 2-3-year-old adults (28 and 14% from dental and epiphyseal fusion data, respectively), and secondarily adults between 3 and 9 years old (53% of dental ages). Apart from the relative deficit in young animals, these profiles are similar to those for the population living on the Isle of Rùm. Combined with the well-balanced sex ratio evidenced above, this indicates a low level of selection by the Shillourokambos inhabitants.

Age profiles of the Middle A2 phase at Shillourokambos (Figure S4B)

Bone preservation is far better than for the Early B phase, especially for young teeth. The sample comes from a large pit filled over a short period of time c. 7,500 cal. BC. This makes this profile more reliable than the previous one, with 196 teeth representing 27 MNI.

The dental age profile indicates that more than 27% of individuals were slaughtered between 6 and 12 months old. The remaining animals were killed without any clear selection between 1 and 9 years of age. This cannot correspond to any sustainable exploitation of a deer population. Such a strategy would not only yield a very low return – each 6-12 month animal would have supplied less than 20kg of meat, based on the weight growth of the Rùm deer (after Clutton-Brock et al., 1982) and adapted to Pfd weights (after Tajbaksh & Jamali, 1995; Vigne, 2011a: 992, figure 16), but would also have rapidly entailed the collapse of the population. This profile is obviously truncated, i.e., most of the sub-adult and adult teeth are missing. This is probably due to the fact that heads were left at the kill site. Indeed, the age profile based on epiphyseal fusion data (n=571) is drastically different, and is probably more representative of the culling strategy. Fawns represent only 5%. Most of the slaughtering focused on young adults (28% between 1-3 years), and adults (67%). The discrepancy between dental and epiphyseal fusion profiles demonstrates that animals between 6-12 months old (and less than 40kg) were brought to the village whole. Conversely, older animals were butchered outside the dwelling area, and their heads were often left there. On the other hand, the epiphyseal fusion profile does not differ from the Early phase profile nor from the survival profile of modern red deer on the Isle of Rùm – apart from the bias in class [3-4 years], which can mostly be considered as artefactual (Vigne, 1984, 2011a). Again, combined with the well-balanced sex ratio evidenced earlier for this phase, it suggests a poor selective slaughtering strategy.
References cited only in supporting information (Appendix S1, Methods S1-3)

NB: for references present in the main text and also cited in supporting information (excluding references for Method S1), refer to main text references.

Akaike H. 1974. A new look at the statistical model identification. IEEE Transactions on Automatic Control 19: 716-723.

Aurenche O, Kozlowski SK. 1999. La naissance du Néolithique au Proche-Orient. Errance: Paris.

Brochier JE. 2013. The use and abuse of culling profiles in recent zooarchaeological studies: some methodological comments on “frequency correction” and its consequences. Journal of Archaeological Science 40: 1416-1420.

Davis SJM. 1984. Khirokitia and its mammal remains. A Neolithic Noah's ark. In Fouilles récentes à Khirokitia (Chypre), 1977-1981, Le Brun A (ed.). Recherche sur les civilisations: Paris; 147-162.

Dempster AP, Laird NM, Rubin DB. 1977. Maximum likelihood from incomplete data via the EM algorithm. Journal of the Royal Statistical Society, Series B 39: 1-38. DOI: 10.1.1.133.4884

Guilaine J. 2003. Parekklisha-Shillourokambos. Périodisation et aménagements domestiques In Le Néolithique de Chypre. Actes du Colloque organisé par l’École Framçaise d’Athènes et le Département des Antiquités de Chypre, Nicosie, 17-19 mai 2011, Guilaine J, Le Brun A (eds). BCH Suppl. 43, École Française d'Athènes: Athènes; 3-14.

Guilaine J, Briois F. 2007. Shillourokambos and the neolithization of Cyprus: some reflections. Eurasian Prahistory 4: 159-175.

Hammer Ø, Harper DAT, Ryan PD. 2001. PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4(1): 9pp.

Le Brun A, Daune-Le Brun O. 2003. Deux aspects du Néolithique pré-céramique récent de Chypre: Khirokitia et Cap Andreas-Kastros. In Le Néolithique de Chypre. Actes du Colloque organisé par l’École Framçaise d’Athènes et le Département des Antiquités de Chypre, Nicosie, 17-19 mai 2011, Guilaine J, Le Brun A (eds). BCH Suppl. 43, École Française d'Athènes: Athènes; 45-59.

Le Brun A, Daune-Le Brun O. 2009. Khirokitia (Chypre) : La taille et les pulsations de l'établissement néolithique pré-céramique, nouvelles données. Paléorient 35(2): 67-76.

Manning SW, McCartney C, Kromer B, Stewart ST. 2010. The earlier Neolithic in Cyprus: recognition and dating of a Pre-Pottery Neolithic A occupation. Antiquity 84: 693-706.

Peltenburg E (ed.). 2003. The colonisation and settlement of Cyprus. Investigations at Kissonerga-Mylouthkia, 1976-1996. SIMA 70(4)/Lemba Archaeological Project, Cyprus, Vol. III.1, Paul Åströms: Savedalen.

Simmons A (ed.). 1999. Faunal extinction in an island society. Pygmy hippopotamus hunters of Cyprus. Kluwer Academy: New York, Boston; Plenum Publisher: London, Moscow, Dordrecht.

Vigne J-D. 1984. Premières données sur les débuts de l'élevage du Mouton, de la Chèvre et du Porc dans le sud de la Corse (France). In Animals and Archaeology 3. Early Herders and their Flocks. 4th International Council for Archaeozoology, London, 1982. Clutton-Brock J, Grigson C (eds). BAR International Series 202: London; 47-65.

Vigne J-D, Helmer D. 2007. Was milk a “secondary product” in the Old World Neolithisation process? Its role in the domestication of cattle, sheep and goats. Anthropozoologica
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