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Distinguishing Wild Boar from Domestic Pigs in Prehistory: A Review of Approaches and Recent Results Peter Rowley-Conwy, Umberto Albarella & Keith Dobney

Journal of World Prehistory ISSN 0892-7537 J World Prehist DOI 10.1007/s10963-012-9055-0

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Author's personal copy J World Prehist DOI 10.1007/s10963-012-9055-0 ORIGINAL PAPER

Distinguishing Wild Boar from Domestic Pigs in Prehistory: A Review of Approaches and Recent Results Peter Rowley-Conwy • Umberto Albarella • Keith Dobney

 Springer Science+Business Media, LLC 2012

Abstract New methods permit archaeologists to distinguish between wild boar and domestic pigs with greater confidence than has been hitherto possible. Metrical methods are the most commonly used; these are reviewed. Assemblages containing a wider range of measurements (as measured by the coefficient of variation [V]) than is found in one population suggest that two populations of different-sized pigs were present, probably indicating separate wild and domestic populations with little or no interbreeding. These assemblages, sometimes taken to indicate animals ‘intermediate’ between wild and domestic, are clear evidence of full domestication. Other traditional means of diagnosing domestication based on age at death (the killing of many young animals) and biogeography (the export of the animal beyond its natural range, especially to islands) are particularly problematic when applied to pigs. New methods include the frequency of Linear Enamel Hypoplasia, which may increase due to domestication-induced stress, the study of diet through isotopes and dental microwear, and the examination of population histories through ancient and modern DNA and geometric morphometrics. These are all promising but should not be considered in isolation: many problems remain. Keywords

Wild boar  Domestic pig  Domestication  Methods  Old World

Introduction The domestication of the major food mammals is one of the most crucial developments in the history of humankind. If the success of a domestic food animal can be measured by its P. Rowley-Conwy (&) Department of Archaeology, Durham University, South Road, Durham DH1 3LE, UK e-mail: [email protected] U. Albarella Department of Archaeology, University of Sheffield, Sheffield, UK K. Dobney Department of Archaeology, University of Aberdeen, Aberdeen, UK

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geographical dispersal in farming systems, a case can be made that the pig is the most successful of all. Its success places it alongside the commensal rat and the multipurpose dog as the most widely exported species on the planet. Pigs are, however, the most difficult of the major domestic animals to identify as ‘wild’ or ‘domestic’ in the zooarchaeological record. This contribution is an outcome of the ‘Durham Pig Project’, which has examined the nature and identification of domestication by a variety of means. In this contribution, we will examine the various ways in which domestication may be detected, and indeed what the term ‘domestication’ means when applied to a species as versatile as the pig: can the full range of possible human–pig relationships be simply divided into ‘wild’ and ‘domestic’? Wild boar were named Sus scrofa by Linnaeus in 1758, while domestic pigs were initially placed in a different species, Sus domesticus, by Erxleben in 1777 (Gentry et al. 2004). The practice of placing them in a single species, however, soon became common. For example, the Swedish zoologist Nilsson (1847, pp. 451–455) referred to wild boar as Sus scrofa ferus and domestic pig as Sus scrofa domesticus. Subsequent terminology has been variable but has generally placed wild and domestic in the same species. Since they are so closely related, the zooarchaeological distinction between them has always been a problem; feral populations, groups possibly intermediate between ‘wild’ and ‘domestic’, and crosses between all these categories add further difficulties. Many methods have been used. Metrical methods are the most common, and also have the longest history. They are easy to employ, because they are ‘low technology’, and because measurements are routinely taken. The first part of this paper will therefore consider metrical methods. The second part will discuss non-metrical methods, some (but not all) of which are quite new, and more ‘high technology’. Discussion will concentrate on Europe and the Near East, but will also consider some recent advances made by East Asian workers. Archaeological methods are developing all the time. If with the advantage of hindsight we detect errors in the work of our predecessors, we must remember that no conclusions are absolute: future zooarchaeologists will re-evaluate our work, and no doubt demonstrate that it too contains errors. Many questions remain difficult to answer. As an example, the Neolithic pigs from the Swedish island of Gotland (see Fig. 1) will be considered at various places below. These assemblages have been studied in detail over the years—but still cannot be securely identified as ‘wild’, ‘domestic’, or any other category. Maybe this shows that our categories need rethinking; or maybe it just shows that our current methods need refining.

Metrical Methods One of Europe’s earliest identifications of pig was of a canine tooth from a megalithic grave at Kvistofta in Sweden, excavated in June 1819. Nilsson (1822, p. 296) identified this as wild, because of its large size. Some years later the Danish naturalist Japetus Steenstrup (1851, p. 21) identified two third molars from Krabbesholm as wild because of their large size. However, it was a Swiss researcher who first formalised a metrical method of study. From Ru¨timeyer (1860) to Boessneck et al. (1963) Ludwig Ru¨timeyer examined animal bones from the Swiss lake dwellings of Moosseedorf, Wauwyl and Robenhausen, where organic items were well preserved in peat. Jaws were common at these sites. Ru¨timeyer concentrated on complete adult tooth rows when

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Fig. 1 Map of western Eurasia showing the sites and regions mentioned in this chapter. The southern limit of wild boar in Northwest Africa follows Oliver et al. (1993, fig. 10)

available, and measured mainly upper and lower M3s. Ru¨timeyer (1860, pp. 32–36) identified three types of prehistoric pig: – wild boar, Sus scrofa ferus – domestic pig, Sus scrofa domesticus – the torfschwein or ‘peat pig’, Sus scrofa palustris, a new taxon. Ferus was fairly common, while domesticus was rare; palustris was very common on the archaeological sites, but had no living representatives. Although Ru¨timeyer was, of course, aware that wild boar and domestic pigs could interbreed, he did not doubt that each of the three types were was either ‘wild’ or ‘domestic’, and he considered them in those terms. He concluded that palustris was a wild form, possibly of Asian origin, that lived in Europe alongside ferus wild boar. Palustris was later domesticated, giving rise to the domestic pigs of Europe. Not all agreed with this taxonomy: Steenstrup (1861) argued that palustris was merely the female form of ferus. Ru¨timeyer (1862) disproved this by illustrating male and female maxillae of both ferus and palustris (see Fig. 2); ferus and palustris were thus clearly different, however this was interpreted. In 1862 Ru¨timeyer published his first measurements. Based on a large archaeological sample, he gave the following length ranges for lower M3: prehistoric ferus prehistoric palustris

40–53 mm 33–39 mm.

His modern comparatives were few, and somewhat problematic. Three modern domestic M3s were close to palustris (Fig. 2 top). Ru¨timeyer listed only two modern wild

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Fig. 2 Ru¨timeyer’s distinction between Sus scrofa ferus and Sus scrofa palustris. a Chart of lower M3 length, plotting Ru¨timeyer’s five modern comparatives and the ranges of palustris and ferus (from Ru¨timeyer 1862, pp. 36, 122, 185). b and c Drawings of female and male maxillae, showing the distinction between palustris and ferus (reproduced from Ru¨timeyer 1862, tab. VI, figs. 4, 5, 6 and 7). Ru¨timeyer’s drawings were probably at 1:1 scale; in the original, the male ferus fragment was 16 cm in length

boar lower M3s, one from Germany of 40 mm, and one from Algeria of 37 mm. The Algerian specimen was smaller than all his prehistoric ferus wild boar (Fig. 2). Wild boar are now known to be smaller in warmer climates (see below), but this was not known to Ru¨timeyer; had he examined a larger sample of European wild boar, he might have fixed the division between palustris and ferus at around 40 mm with more confidence. Discussion continued (see Pira 1909, pp. 375–403 for a summary). Ru¨timeyer soon concluded that all palustris were in fact domestic, being crosses between European wild boar and Indian domestic pigs (Ru¨timeyer 1864, p.159). Nehring (1889, p. 366) argued that palustris could not be separated from later domestic pigs, and that palustris was part of Sus scrofa domesticus, and this became generally accepted. Ru¨timeyer had thus successfully

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Fig. 3 Measurements of lower M3 from Denmark. a Ertebølle, Aalborg and other sites, values from Winge (1900). b Dyrholm II and Bundsø, values from Degerbøl (1939, table 9 and 1942, table 2 respectively)

distinguished between wild and domestic pigs. His invention of the term palustris has led to prehistoric domestic pigs sometimes being called Sus scrofa palustris. By the early twentieth century, wild boar and domestic pigs were routinely being separated using lower M3 measurements. Figure 3 presents two examples from Denmark. Winge (1900) presented results from shell middens, some Mesolithic (including the famous site of Ertebølle) and some Neolithic (most from Aalborg). The Neolithic teeth were much shorter than the Mesolithic ones, though there was an overlap around 38–41 mm (Fig. 3 top). Winge (1900, pp. 158–159) identified the Mesolithic specimens as wild, and the Neolithic ones as domestic—identical with Ru¨timeyer’s palustris, though he termed them Sus scrofa domesticus (palustris). Degerbøl listed both length and breadth of lower M3s from Mesolithic Dyrholm II (Degerbøl 1942) and Neolithic Bundsø (Degerbøl 1939). Figure 3 (bottom) shows how two measurements can assist interpretation. The two

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scatters show little overlap, except for three large Neolithic teeth that fall in the Mesolithic range. Degerbøl interpreted these as wild boar, hunted by Neolithic farmers who kept domestic pigs (Degerbøl 1939, pp. 94–97). Degerbøl used the term Sus scrofa palustris for his domestic pigs, stressing that they were identical to those presented by Winge and Ru¨timeyer (Degerbøl 1939, p. 134). As the twentieth century progressed, more measurements were taken from more parts of the skeleton in order to separate the conventional wild and domestic categories. This methodological development took place particularly in Germany, and the standard measurement guide now used internationally was produced there (von den Driesch 1976). The Neolithic site of Seeberg Burga¨schisee-Su¨d in Switzerland is a good example, where the pigs were examined by Boessneck et al. (1963). Boessneck stated that there were 1855 wild boar fragments from at least 62 individuals, and 243 domestic pig fragments from at least 23 individuals, while a further 706 bones from at least 10 individuals could not be determined (Boessneck et al. 1963, tab. 2). For lower M3 (Fig. 4a), Boessneck concluded that all the specimens were wild, since none was below 40 mm. In upper M3, however, some smaller domestic specimens were present (Fig. 4b), and the same was true for distal humerus, astragalus and scapula (indicated in Fig. 4c–e). This showed the importance of considering a variety of skeletal elements: had Boessneck relied only on lower M3, he would have concluded that all the animals were wild. These methods have not solved all the problems, however. The Middle Neolithic pigs from Gotland are a good example. Graves at Ire, Grausne and Ajvide (Fig. 1) each contained many pig jaws: Ire, 38 half mandibles from 19 pigs; Grausne, 46 half mandibles from 34 pigs, and Ajvide, 46 half mandibles from 30 pigs (Rowley-Conwy and Dobney 2007, fig. 7.8). The methods described above do not clearly reveal whether they are wild or domestic; and there is the possibility (discussed below) that the conventional division into just ‘wild’ and domestic’ may be an oversimplification. The graves contain only mandibles, so no other skeletal element can be considered (the post-cranial material from the associated settlements is still under study). Most of the jaws are juvenile, and do not contain M3. The 15 lower M3s present are plotted in Fig. 5, compared with domestic specimens from Troldebjerg and wild boar from Ringkloster (both in Denmark). The Gotland specimens fall along the bottom of the Ringkloster scatter. They are larger than Neolithic domestic pigs, which led Ekman (1974) to argue that they were wild boar. They are small for wild boar, however, and Jonsson (1986) argued that they were crosses between wild boar and domestic pigs. Lindqvist and Possnert (1997) suggested that they were crosses between domestic and feral pigs. Metrical methods clearly do not resolve the Gotland problem, which will be returned to below. Geographical Variability in Wild Boar All the studies described above were undertaken in Central Europe (Fig. 1), where wild boar were everywhere of similar size. But when work began in other areas, it became apparent that wild boar size varied for climatic reasons. Davis demonstrated that wild boar in colder climates were larger, while those in warmer climates were smaller (Davis 1981, fig. 3; 1987, fig. 3.10)—and see Ru¨timeyer’s Algerian specimen in Fig. 2. Mesolithic wild boar in southern Portugal show a progressive size decline to the south, on the basis of astragalus because insufficient M3s were present (Fig. 6). Also plotted are wild boar from Zambujal (Copper Age), where domestic pigs were more common (von den Driesch and Boessneck 1976). Zambujal’s environment is relatively cool and moist, while the other sites are in progressively hotter and drier areas less suitable for pigs. Size decreases markedly from

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Fig. 4 Selected measurements from Seeberg Burga¨schisee-Su¨d. a Lower M3; b upper M3; c distal humerus; d astragalus; e scapula (values listed in Boessneck et al. 1963, pp. 60, 61, 64, 67–68 and 63–64 respectively). The measurements correspond with those defined by von den Driesch (1976)

Zambujal to Fiais. The southernmost limit of wild boar is North Africa (Fig. 1); southern Portugal is the closest part of Europe to this. The wild boar from Copper Age Leceia, in a similar environment to Zambujal, are similarly large (Albarella, Davis et al. 2005). Zambujal

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Fig. 5 Lower M3s from Neolithic burials at Ire, Grausne and Ajvide on Gotland, compared with Mesolithic specimens from Ringkloster and Neolithic specimens from Troldebjerg, both in Denmark (Rowley-Conwy unpublished)

and Leceia may, however, raise a further complexity: wild boar size sometimes increases in Neolithic contexts (Albarella, Davis et al. 2005; Albarella, Dobney and Rowley-Conwy 2009; Albarella, Tagliacozzo et al. 2006). We must clearly be cautious before comparing between sites even within Spain and Portugal (Rowley-Conwy 1995); and the Zambujal wild boar were themselves much smaller than those from Ringkloster in Denmark (Fig. 6). Work in Japan and the Far East has also highlighted geographically related size variation, and in addition the effects of small islands (Hongo et al. 2007, pp. 124–126). Prehistoric wild boar on Hokkaido (Fig. 7) were larger than those on Honshu (Yamazaki et al. 2005, p. 163). Those on the Izu Islands were smaller than those on Honshu; this could be due to climatic factors, but insular dwarfing and/or human intervention might also have played a part (Yamazaki et al. 2005, pp. 169–170). Prehistoric wild boar on Okinawa were also smaller than those on Honshu (Matsui et al. 2005). Wild boar on Taiwan are however larger than those on Okinawa, and similar in size to those on Honshu, which suggests that climate is not the only factor involved in determining size on Okinawa (Yamazaki et al. 2005, p. 169). The largest wild boar in eastern Eurasia are Sus scrofa ussuricus from the Russian Far East (Albarella et al. 2009). Climate-related size variation is important with regard to domestication. Since domestic pigs are smaller than wild boar, domestication should appear as a size decrease. But climatic warming at the end of the last glacial should also cause wild boar to decrease in size. In a major survey of pigs from many sites in Israel, Davis identified two size decreases: the first at the end of the last glacial, attributed to climatic change; and the second during the Neolithic, attributed to domestication (Davis 1981, fig. 9; 1987, figs. 6.9, 6.10; see also Ducos and Horwitz 1998). Domestication and ‘Semi-Domestication’ Work on initial domestication in the Near East began only in the mid-twentieth century (see Flannery 1983 for a review). Data were initially very scarce. The first task was to establish the metrical parameters of Near Eastern wild boar. Figure 8 (top) shows lower

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Fig. 6 Astragalus lengths from sites in Portugal, compared with those from Ringkloster in Denmark. Ringkloster from Rowley-Conwy (unpublished); Zambujal from von den Driesch and Boessneck (1976, tabelle 22); Muge, Sado and Fiais from Rowley-Conwy (1995, fig. 8)

M3 length of modern Near Eastern specimens. They are somewhat smaller than their European counterparts, presumably for climatic reasons (see above). Pig bones were usually rare on archaeological sites, so measurable samples were small. Figure 8 shows Flannery’s pioneering results from Jarmo in Iraq; the Pottery Neolithic specimens might be smaller than the Pre-Pottery Neolithic ones, but samples are very small indeed. From a series of small samples, Flannery concluded that pig domestication in the Jarmo region occurred near the start of the Pottery Neolithic. Subsequent work has produced larger samples. Further west, pigs were apparently domesticated during the Pre-Pottery Neolithic. At Gu¨rcu¨tepe in southeastern Turkey (late PPNB), the pigs are smaller than at earlier sites in the region (Peters et al. 2000, 2005, fig. 10). At Atlit Yam in Israel (PPNC), wild and domestic pigs can be distinguished (Horwitz et al. 2000). At Hagoshrim, also in Israel, late PPN and early Pottery Neolithic

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Fig. 7 Map of eastern Eurasia and Japan, showing the sites and regions mentioned in this paper

layers produced large wild boar. The later Pottery Neolithic, however, showed a marked drop in size, identified as the result of domestication, accompanied by an increase in the killing of younger animals (Haber and Dayan 2004; Haber et al. 2005). The clearest evidence for domestication during the PPN comes from C¸ayo¨nu¨ Tepesi in southeastern Turkey (Fig. 8). The sequence shows a continuous reduction of M3 length, starting in the Pre-Pottery Neolithic, which must be the result of domestication (Ervynck et al. 2002). The discussion so far has assumed that pigs can be divided into straightforward ‘wild’ and ‘domestic’ categories. In the late 1960s theoretical discussions of domestication, coming initially mainly from the Cambridge University ‘bone room’ group of researchers, suggested that the situation was more complex (e.g. Higgs and Jarman 1969; Jarman and Wilkinson 1972). Intensive close herding on the one hand, and pure hunting on the other, are extremes on a wide spectrum of behaviour, with various forms of extensive management and loose herding in between. In medieval England, for example, some pigs were intensively managed and fed on crops grown for them (Biddick 1984); others were herded into woodlands in the autumn to eat acorns (‘pannage’), or foraged for themselves in towns (Grigson 1982). In some English towns this continued until the twentieth century (Malcolmson and Mastoris 1998). Very extensive strategies may be employed in more rural settings. On Corsica and Sardinia some pigs live far away from their owners and are visited just two or three times per year, and interbreed with wild or feral animals (Albarella, Manconi et al. 2007). With such a range of management practices, can we think just in terms of ‘wild’ and ‘domestic’?

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Fig. 8 Length of lower M3 teeth from the Near East. Modern specimens are those listed by Flannery (1983, table 3), plus five individuals from Payne and Bull (1988, table 1a), two from Hongo and Meadow (1998, table 4) and those from Dobney et al. (2007); Jarmo from Flannery (1983, tables 6 and 7); C¸ayo¨nu¨ from Ervynck et al. (2002, fig. 13); Hallan C¸emi from Redding and Rosenberg (1998, table 1)

Jarman (1971, 1976) approached this archaeologically at the Neolithic site of Molino Casarotto in Italy. He questioned the osteological separation of wild and domestic pigs, and whether prehistoric pigs were necessarily hunted just because their teeth are the same size as those of modern wild boar. Figure 9 plots Jarman’s measurements (1976, Table 4). Comparing them with sites such as Seeberg Burga¨schisee-Su¨d, Jarman stated that ‘the pigs bridge the accepted size ranges of wild pigs and Neolithic domestic pigs’ (1976, p. 528).

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Fig. 9 Lengths of upper and lower M3 from Molino Casarotto, Italy. Data from Jarman (1976, table 4)

Comparison with Fig. 4a, b confirms this: at Molino Casarotto lower M3 lengths fall each side of 40 mm, and upper M3 lengths fall each side of 36 mm, the measurements where Boessneck divided wild and domestic at Seeberg Burga¨schisee-Su¨d. Jarman offered three possible explanations for the Molino Casarotto pigs: 1. Two populations were present, but were so close in size that no separation can be seen in the measurements. 2. Two populations were present, but the separation was blurred by the presence of feral, hybrid or proto-domestic forms (Jarman did not define the term ‘proto-domestic’). 3. Only one population was present. Jarman favoured the third option, because there was no evidence to suggest that two different modes of pig exploitation were operating (1971, p. 257; 1976, p. 528). He argued that ‘human economic behaviour may have partially caused this [metrically] anomalous population … there was a human exploitation pattern that relied on a closer degree of control over the pig herds than formerly’ (1976, p. 530). Jarman was therefore suggesting that because the pigs were metrically intermediate between wild and domestic, they were also behaviourally intermediate. To examine whether this suggestion is valid, we can consider Dwyer’s (1996) ethnographic study of pig husbandry in New Guinea. In addition to hunting, he defines three forms of human–pig relationship that he terms ‘domestication’: 1. All domestic pigs are captured from the wild as piglets. Males are castrated, and females kept close to the settlement so that they cannot breed with wild males. Dwyer (1996, pp. 485–487) terms this ‘reproductive alienation’, because humans do not breed from their domestic pigs. 2. Domestic females are kept, but no breeding males. All domestic breeding is with wild males, encountered by the domestic females in the bush around the settlements. Females give birth within the settlement, but all male piglets are castrated. Dwyer (1996, pp. 486–488) terms this ‘female breeding’, since humans control only the females. The behavioural gulf between wild and domestic is substantial. Among the Maring, ‘young pigs are treated as pets …. The young animal receives a great deal of loving attention—it

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is petted, talked to, and fed choice morsels’ (Rappaport 1968, p. 58). It lives in the women’s house until released at 4–5 months of age, after which it forages in the village, returning home each evening. Among the Etoro, piglets live inside the houses and are ‘fed and fondled for three to six months so that they will develop a permanent attachment to their owners’ (Kelly 1988, p. 115), after which they are released, but whenever encountered by its owner ‘a pig is invariably called by name, stroked, scratched, and fed bits of food …’ (1988, p. 116). Both the Maring and the Etoro castrate all males in domestic litters, and also hunt the wild pigs that father their domestic litters (Rappaport 1968, pp. 13, 70, 78; Kelly 1988, pp. 115–116). 3. Both domestic males and females are kept, so that all domestic litters are the offspring of two domestic parents. Dwyer (1996) terms this ‘male and female breeding’. The Wola are a good example, keeping both domestic females and breeding males, as well as castrated males (Sillitoe 2007). The archaeological outcomes of these strategies are crucial. No size change can occur in options 1 or 2, because of the continuous input of genes from the wild males. No ‘intermediate’ size grouping of the kind argued for by Jarman (above) is thus likely to emerge. Only in option 3, where both males and females are under close human control, could a size divergence occur, because of the separation of the wild and domestic gene pools. Jarman’s argument that intermediate size indicates intermediate behaviour is thus unlikely to be correct; the status of the Molino Casarotto pigs remains uncertain. Can Dwyer’s ‘female breeding’ (option 2) be recognised in the archaeological record? At Hallan C¸emi in eastern Turkey, no plants or other animals were domesticated, but ‘female breeding’ of pigs is suggested at the start of the Pre-Pottery Neolithic as a possible precursor to full domestication (Redding and Rosenberg 1998; Rosenberg et al. 1998). The authors stress that the size range of the Hallan C ¸ emi pigs corresponds entirely with wild boar; and the five lower M3s fall in the wild range (Fig. 8). ‘Female breeding’ would not cause a size change (Redding and Rosenberg 1998, p. 68), but the high proportion of juveniles is a supporting argument: bone fusion indicates that some 40% of animals died under 12 months—and Jarman advanced a similar argument for Molino Casarotto, where 50% died under 2 years of age (1976, p. 528). The issue of age at death will be discussed below. Separation of ‘wild’ and ‘domestic’ pigs may thus be substantially more complex than earlier researchers assumed, and these may indeed not be the only possible categories that should be considered. Nothing discussed in this section has cleared up the situation with regard to Gotland (Fig. 5). Jarman did not discuss these animals; had he done so he might well have suggested an intermediate status for them, but as noted above, metrical intermediacy is not evidence of behavioural intermediacy. The Significance of Variability Discussion so far has been based on the variations visible in archaeological measurements. But how much variation is there in a living population? Most comparative collections do not contain enough individuals to answer this. A major step was the publication of 18 wild boar skeletons from a single population at Kızılcahamam in Turkey (Payne and Bull 1988). Payne and Bull stress that the sample is relatively small, and that its age distribution limits some conclusions: only five animals were old enough to have erupted M3s, for example. Studies of larger samples of modern skeletons are definitely needed. A very large sample of archaeological material from the Neolithic site of Durrington Walls in Britain has partly filled the gap: there are no complete skeletons, but only one population is present, assumed

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to be domestic pig (Albarella and Payne 2005). These studies have resulted in two major methodological improvements. The first improvement is a better understanding of age-related size change. Zooarchaeologists have often assumed that once bones are fused, full adult size has been attained; but some elements actually continue to grow after fusion. In the Kızılcahamam skeletons, most forelimb measurements increase after fusion (Payne and Bull 1988, pp. 30–31 and table 2). At Durrington Walls the same was true, although individual bones could not be aged by associated jaws: proximal radius breadth (measurement BpP of Albarella and Payne 2005) was substantially greater when fusion was complete than when the fusion line was visible (Albarella and Payne 2005, p. 595). This reduces the value of most forelimb measurements. Rear limb measurements increase rather less, but often do not survive very well. Distal tibia is common, and shows little post-fusion growth. Astragalus rapidly reaches adult size and thereafter shows little change. Teeth do not grow after eruption, but a tooth may by impacted by other teeth growing next to it; this is particularly the case for M1 and M2. The lengths of these teeth may therefore decrease slightly, though their widths do not (Payne and Bull 1988, pp. 30–31). At Durrington Walls, heavily worn lower M1s were on average shorter than lightly worn ones (Albarella and Payne 2005, fig. 4). Lower M3 length and width is rather more variable than the widths of M1 and M2; M3 remains a useful tooth, but its measurements should not be used in isolation (Albarella and Payne 2005, pp. 596–597). The result is a better understanding of which measurements are most appropriate for particular tasks. Simply measuring more and more skeletal elements does not increase information return. Albarella and Payne (2005, p. 598) argue that assemblages are most usefully compared using: • • • •

widths of lower M1 and M2 width and length of lower M3 tibia BdP astragalus GLl.

Distal humerus is often common. Bd shows considerable post-fusion growth but the trochlea shows less, so it is worth adding: • humerus BT and HTC. Other measurements may also be taken but may be more useful for other purposes. Scapula SLC and longbone shaft widths are subject to so much growth that they are more useful for determining site seasonality (Rowley-Conwy 1998, 2001a). The second improvement concerns the variability to be expected in any particular measurement. This is expressed mathematically by calculating: 1. the mean of a measurement 2. the standard deviation 3. The coefficient of variation (V), which expresses the standard deviation as a percentage of the mean (Payne and Bull 1988, p. 29). For example, the Kızılcahamam astragalus GLl mean is 47.4. The standard deviation is 2.70. The coefficient of variation is thus 2.70/47.4 9 100, or 5.7 (Payne and Bull 1988, table 1). Coefficients of variation for all measurements from Kızılcahamam and Durrington Walls with a sample size larger than 8 are presented in Table 1. Astragalus GLl from Zambujal in Portugal (Fig. 10 top) provides a useful application of this. The distribution forms a major peak, with a ‘tail’ of larger specimens projecting to the

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Author's personal copy J World Prehist Table 1 Metrical attributes of pig bones from the Kızılcahamam population, for all measurements where a sample of 8 or more is available, and for selected elements from Durrington Walls Element and measurement

Kızılcahamam N

Mean

Durrington Walls SD

V

N

Mean

V

Upper dp4 L

23

14.0

4.6

WA

22

11.0

3.7

WP

23

10.7

4.8 5.8

Upper M1 L

18

18.9

1.06

5.6

82

17.1

WA

18

15.3

0.44

2.9

84

13.7

4.0

WP

18

15.6

0.41

2.6

86

13.5

3.9

L

15

24.8

1.10

4.4

82

22.1

4.6

WA

15

19.7

0.64

3.2

77

17.0

4.6

WP

15

19.2

0.66

3.4

71

16.5

5.7

L

39

32.9

6.3

WA

44

18.5

4.4

L

74

18.9

4.8

WA

69

6.3

4.3

WP

73

8.6

4.5

Upper M2

Upper M3

Lower dp4

Lower M1 L

18

18.9

1.20

6.3

128

17.3

5.2

WA

18

11.5

0.39

3.4

127

10.3

4.5

WP

18

12.5

0.46

3.7

125

10.9

5.0

L

15

24.9

0.92

3.7

81

21.8

4.6

WA

15

15.4

0.53

3.4

74

13.7

4.3

WP

15

16.3

0.61

3.7

68

14.2

4.5

L

39

34.5

5.5

WA

42

15.7

6.0

Lower M2

Lower M3

Scapula GLP

14

39.4a

2.59

6.6

96

36.7

7.1

SLC

14

25.5a

2.34

9.2

115

23.8

10.6

Bd

15

46.3a

2.73

5.9

102

41.1

5.0

BT

15

32.6a

1.92

5.9

67

31.3

5.7

HTC

15

21.5a

1.12

5.2

110

19.7

5.4

15

32.6

1.84

5.6

172

29.6b

5.8b

13

34.8a

1.83

5.2

97

33.3

4.9

Humerus

Radius BpP Pelvis LAR

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Author's personal copy J World Prehist Table 1 continued Element and measurement

Kızılcahamam

Durrington Walls

N

Mean

SD

V

N

Mean

V

8

33.2a

1.62

4.9

116

30.8b

4.5b

17

47.4

2.70

5.7

160

40.8

6.0

Tibia BdP Astragalus GLl a

Fusing as well as fused bones are included

b

Single large (possibly wild) individuals included. Kızılcahamam from Payne and Bull (1988: table 1b), where V is calculated only to the nearest whole number; here Payne and Bull’s figures are used to recalculate values to the nearest 0.1. Durrington Walls from Albarella and Payne (2005, tables 2, 3 and 4)

right. In the original report the peak was interpreted as domestic animals, the tail as a smaller number of large wild boar, dividing at 45 mm (von den Driesch and Boessneck 1976). The V for the entire Zambujal sample is 8.2, compared to 5.7 at Kızılcahamam and 6.0 at Durrington Walls. This higher value suggests that there were indeed two populations of differing sizes at Zambujal, as von den Driesch and Boessneck (1976) concluded. When calculated only for the domestic group, V falls to 5.7, very close to Kızılcahamam and Durrington Walls, suggesting that this is a single breeding population. But when calculated for the wild group above 45 mm, V falls to 3.4, considerably lower than at Kızılcahamam and Durrington Walls. The full wild range at Zambujal is therefore probably greater than appears in Fig. 10; the lower end must extend below 45 mm, down into the ‘domestic’ zone. The minor peak at 43–44 mm supports this suggestion. Thus if the V of any sample is much greater than at Kızılcahamam and Durrington Walls, two populations of different sizes must be present. V is not the perfect statistic: in small samples it may be unduly influenced by individual outliers, and it is not clear how large it must be before we can conclude that two populations must be present. But where V does indicate two populations, this has huge implications, because if two populations are present, one must be wild and the other domestic, with little or no interbreeding between them: domestic males and females must therefore both have been under close human control— only humans can be responsible for the breeding barrier between wild and domestic. Therefore when two populations are present, no ‘intermediate’ situations, such as Jarman’s ‘semi-domestic’ pigs or Dwyer’s ‘reproductive alienation’ or ‘female breeding’ strategies (see above), have occurred: the pigs are conventionally ‘wild’ and ‘domestic’. Variability in the Archaeological Record The method discussed in the previous section is widely applicable. The V for all the elements plotted in the figures in this paper are listed in Table 2. Too great accuracy should not be assumed for this table. Current practice is to measure to the nearest 0.1 mm, but in many early publications (Ertebølle, Aalborg, Dyrholm II, Bundsø, Seeberg Burga¨schiseeSu¨d) most measurements are given to the nearest millimetre. For C¸ayo¨nu¨ the individual measurements are not listed, but are grouped in millimetre intervals in the published graph (Ervynck et al. 2002, fig. 13), so all specimens between (for example) 41.0 and 41.9 mm are here treated as if they measured 41.0 mm. Similarly, the Egolzwil 2 specimens in Fig. 10 were read from Higham (1968, fig. 6) to the nearest millimetre. Variations between

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Author's personal copy J World Prehist 25

Zambujal v:

20

whole sample 8.2 domestic 5.7 wild 3.4

15 10

domestic wild 5 0 30

32

34

36

38

40

42

44

46

48

50

astragalus GLl (mm)

Star evo v: 16.7 2 1 0 26

28

30

32

34

36

38

40

42

44

46

48

50

52

54

lower M3 L (mm) 7 6 5

Selevac v: 26.0

4 3 2 1 0 26

28

30

32

34

36

38

40

42

44

46

48

50

52

54

lower M3 L (mm) 10

Egolzwil 2 v: 14.9

9 8 7 6 5 4 3 2 1 0 26

28

30

32

34

36

38

40

42

44

46

48

50

52

54

lower M3 L (mm) Fig. 10 Lengths of astragalus and lower M3 from various sites, with coefficients of variation (V). Zambujal 1–4 from von den Driesch and Boessneck (1976); fig. 12 of that work plots 94 domestic specimens and all are included here, while table 22 gives measurements of only 84 domestic specimens, which are used to calculate V. Starcˇevo from Clason (1982, table 38); Selevac from Legge (1990, pp. 239–240); Egolzwil from Higham (1968, fig. 6)

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Author's personal copy J World Prehist Table 2 Metrical attributes of the various samples of tooth and bone measurements presented in the graphs in this paper Figures

Series

N

Mean

SD

V

Lower M3: length (L) 3

Ertebølle and others

16

44.8

3.6

3

Aalborg and others

18

36.1

2.8

7.6

3

Dyrholm II

17

46.7

3.9

8.3

3

Bundsø (all)

15

36.7

5.1

13.9

12

34.8

3.2

9.2

4

Same, omitting three wild Seeberg Burga¨schisee-Su¨d

17

43.7

2.6

5.9

5

Ringkloster

60

44.7

2.9

6.5

5

Troldebjerg

68

35.9

2.6

7.2

5

15

41.9

2.0

4.8

9

42.8

2.4

5.6

8

Gotland burials C¸ayo¨nu¨ PPN 2 C¸ayo¨nu¨ PPN 3

8

C¸ayo¨nu¨ Pottery Neolithic

9

Molino Casarotto Starcˇevo

40

39.9

3.2

8.0

10

15

40.1

6.7

16.7

10

Selevac

40

35.4

9.2

26.0

10

Egolzwil 2

94

38.8

5.8

14.9

8

8.0

15

40.4

3.4

8.4

9

37.0

3.7

10.0

Lower M3: breadth (B) 3

Dyrholm II

17

19.2

1.0

5.2

3

Bundsø (all)

15

16.3

1.6

9.9

12

15.8

1.0

6.3

4

Same, omitting three wild Seeberg Burga¨schisee-Su¨d

17

19.0

1.1

5.8

5

Ringkoster

60

19.9

0.9

4.5

5

Troldebjerg

68

16.0

1.0

6.2

5

Gotland burials

15

18.1

0.8

4.4

Upper M3: length (L) 4

Seeberg Burga¨schisee-Su¨d (all)

9

Molino Casarotto

9

37.0

4.7

12.7

38

37.4

2.3

6.1

9

20.4

2.1

10.3

Upper M3: breadth (WA) 4

Seeberg Burga¨schisee-Su¨d (all)

Lower M2: length (L) 11

Gomolava wild

12

22.7

1.9

8.4

11

Gomolava domestic

15

18.9

0.8

4.4

Lower M2: posterior width (WP) 11

Gomolava wild

12

18.1

0.8

4.5

11

Gomolava domestic

15

12.5

1.0

8.1

Lower M1: length (L) 11

Gomolava wild

11

Gomolava domestic

9

17.8

1.3

7.3

22

15.4

1.0

6.8

9

13.1

0.7

5.6

22

9.8

0.7

6.8

Lower M1: posterior width (WP) 11

Gomolava wild

11

Gomolava domestic

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Author's personal copy J World Prehist Table 2 continued Figures

Series

N

Mean

SD

V

Upper M1: anterior width (WA) 12

Peschany 1 (all)

52

15.2

1.0

Same, omitting three wild

49

15.0

0.7

4.6

9

17.9

1.2

6.7

Peschany 1 (all)

52

15.6

1.0

6.4

Same, omitting three wild

49

15.4

0.7

4.6

9

18.6

1.0

5.2

21

44.5

5.6

12.6

19

46.0

3.4

7.4

21

31.2

4.0

12.8

19

32.3

2.4

7.4

Seeberg Burga¨schisee-Su¨d (all)

45

48.1

6.4

13.3

Same, omitting seven domestic

38

50.5

3.4

6.7

Seeberg Burga¨schisee-Su¨d (all)

31

49.7

3.7

7.4

Same, omitting two domestic

29

50.4

2.4

4.8

6

Ringkloster

39

52.2

2.2

4.2

10

Zambujal, all

106

40.3

3.3

8.2

6, 10

Same, wild only

11

47.1

1.6

3.4

10

Same, domestic only

94

39.4

2.3

5.7

6

Muge middens

29

43.5

1.4

3.2

6

Sado middens

13

42.5

2.8

6.6

6

Fiais

10

40.2

3.5

8.7

12

Modern wild ussuricus

6.6

Upper M1: posterior width (WP) 12 12

Modern wild ussuricus

Scapula: length of glenoid process (GLP) 4 Seeberg Burga¨schisee-Su¨d (all) Same, omitting two domestic Scapula: breadth of glenoid (BG) 4 Seeberg Burga¨schisee-Su¨d (all) Same, omitting two domestic Humerus: distal breadth (Bd) 4

Astragalus: length (GLl) 4

The figure number in which each appears is listed in the left column—see the figures for the sources of the measurements. See the text for potential sources of variation or error

observers will also play a part, and we cannot assume that measurements taken a long time ago correspond exactly with the modern definitions of von den Driesch (1976) and Albarella and Payne (2005). The values in Table 2 are therefore indicative rather than absolute, but do present interesting information. Lower M3 lengths in many samples where a single population is likely (Seeberg Burga¨schisee-Su¨d, Ringkloster, Troldebjerg) produce a V around 5 to 7, close to the 5.5 at Durrington Walls. However, some sites where single populations would be expected (Ertebølle, Aalborg, Dyrholm II, Bundsø domestic) have a rather larger V, and would benefit from being re-measured. At Seeberg Burga¨schisee-Su¨d (Fig. 4), the V of upper M3 length and breadth, scapula GLP, humerus Bd and astragalus GLl all decrease substantially when the presumed domestic specimens are omitted, bringing them closer to what is expected from a single population. Not all problems can be resolved, however. At Molino Casarotto (Fig. 9), lower M3 has a V of 8.0, slightly larger than expected for a

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single population; upper M3, however, has a V of 6.1, closely comparable to the single population at Durrington Walls. Measurements from other teeth and bones might resolve this, but at the moment the situation at Molino Casarotto remains unclear. Three other Neolithic populations with large V are shown in Fig. 10. At Starcˇevo in Serbia, Clason (1982) argued that both wild and domestic animals were present. The V is 16.7, which clearly supports this interpretation; it is not certain whether the domestic specimens are really Neolithic, or are intrusive from overlying Iron Age layers (Clason 1982, p. 158). At Selevac, also in Serbia, the distribution of M3 lengths is somewhat bimodal, and Legge (1990) identified both wild and domestic animals; the large V of 26.0 confirms this. Egolzwil 2 in Switzerland is less bimodal, but the V of 14.9 again confirms that both wild and domestic animals are present. All the samples presented in Fig. 10 thus demonstrate that some close-herded domestic animals were present (assuming the small Starcˇevo specimens are Neolithic). C¸ayo¨nu¨ produces interesting results regarding initial domestication. There is a clear decrease in size, attributed to domestication, through the PPN 2, PPN 3 and Pottery Neolithic layers (Fig. 8). This is, however, accompanied by an increase in V through these layers: PPN 2 PPN 3 Pottery Neolithic

5.6 8.4 10.0

This suggests the increasing divergence of two separate populations through time: the newly domesticated population, now isolated from the wild gene pool; and wild boar, evidently still being hunted. Measurements of M1 and M2 can be very informative. This is shown by the Neolithic site of Gomolava in Serbia, where Clason (1979, table 6) published lengths and widths of these teeth. Of the lower M3s, Clason concluded that 17 came from wild boar and 6 from domestic pigs, but many of the M1s and M2s remained undiagnosed. For M2 (Fig. 11 bottom), length produces no separation—but width clearly separates wild from domestic, showing that most of Clason’s undiagnosed M2s were in fact from domestic pigs. Lengths of both domestic and wild M2 have rather high V, 8.1 and 8.4 respectively (Table 2); perhaps there was age-related shortening of the tooth crowns, as discussed above. For M1, width again shows the best separation (Fig. 2 top); 3 teeth originally identified as domestic fall in the wild scatter. If the identifications suggested in Fig. 11 are correct, the wild/ domestic ratios are as follows: Tooth

Wild

Domestic

% Domestic

M3

17

6

M2

12

15

56

M1

9

22

71

26

The percentage of domestic animals is highest in M1, the earliest erupting tooth, and lowest in M3, which erupts last. This suggests that wild boar and domestic pigs were killed at different ages: if most domestic pigs were killed young, few would have erupted M3s; so if more wild boar were killed as adults, their M3s would be relatively more common. At

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Fig. 11 Lower M1 and M2 from Gomolava (Clason 1979, table 6), compared with the modern sample from Kızılcahamam (Payne and Bull 1988, table 1a). The key gives Clason’s identifications as wild, domestic or unknown; the divisions marked on the graphs are as explained in the text

Gomolava, therefore, the pig economy was based on the slaughter of juvenile domestic pigs and the hunting of more adult wild boar. Is the two-population pattern also found in eastern Eurasia? It certainly is at Peschany 1, in the Russian Far East (see Fig. 7). The site dates from the first millennium BC (RowleyConwy and Vostretsov 2009). Figure 12 presents one pair of measurements, upper M1 WA and WP. Peschany 1 forms a tight distribution, with three large outliers falling in the size range of modern wild boar from the region. The V for the total sample is rather larger than for Kızılcahamam and Durrington Walls; but when the three presumed wild specimens are excluded, it comes much closer (Table 2). Figure 12 also indicates the size ranges from Kızılcahamam, which makes clear the outstandingly large size of Sus scrofa ussuricus wild boar; the Peschany 1 domestic pigs are also considerably larger than those from Durrington Walls.

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Fig. 12 Upper M1 from Peschany 1 (Russian Far East) compared to Sus scrofa ussuricus wild boar from the same region (Rowley-Conwy and Vostretsov 2009). Size ranges from Durrington Walls and Kızılcahamam are included for comparison (Albarella and Payne 2005, table 2; and Payne and Bull 1988, table 1b; see Table 1 of this paper for details)

Metrical Methods: Conclusions The outcome of this section is rather paradoxical. Wide and diffuse scatters (those with a high V), that were formerly poorly understood, now yield the clearest information: two populations were present, and one of them must be fully domestic pigs under close human control—even though the precise proportion of wild and domestic cannot be established. For the European Neolithic sites considered above, this goes for Ru¨timeyer’s Swiss sites, Bundsø, Zambujal, Seeberg Burga¨schisee-Su¨d, Egolzwil, Gomolava, Starcˇevo and Selevac; Argissa in Greece can be added (Payne and Bull 1988, p. 35); C ¸ ayo¨nu¨ shows the emergence of this pattern; and Peschany 1 demonstrates it at the other end of Eurasia. In spite of the theoretical possibility that a simple division into ‘wild’ and ‘domestic’ may be an oversimplification, in most cases the animals do divide clearly into ‘wild’ and ‘domestic’. Undefined further categories like ‘semi-domestic’ are not usually needed. Sites with a single population (a small V) are less easy to interpret. Neolithic sites with a single population of small animals (Durrington Walls, Troldebjerg) are probably safely interpreted as having only domestic animals. Mesolithic sites with a single population of large animals (Ertebølle, Ringkloster, the Muge and Sado middens, and Fiais) probably have only wild boar—although as we have seen, we cannot technically exclude the possibility of ‘female domestication’. Most difficult to interpret are single-population assemblages from Neolithic sites, where the pigs are large (of ‘wild’ size), but are found with other domestic animals and cultivated plants. This is the situation in Italy. Jarman (1976) showed that the Neolithic pigs at Molino Casarotto were larger than domestic pigs in central Europe (see above). At Grotta dell’Uzzo, the teeth of Neolithic pigs are the same size as those in the underlying Mesolithic layers, while the postcranial elements show a very slight size drop (Albarella, Tagliacozzo et al. 2006). Only much later in the Neolithic do samples with a large V

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appear, for example at Arene Candide (Rowley-Conwy 1997), various sites in the Po Valley (Rowley-Conwy et al. in press), and at Connelle, dating to the Eneolithic—and here, the wild boar increase in size at the same time as the domestic pigs decrease (Albarella, Tagliacozzo et al. 2006). The Italian Neolithic animals are so large that they are unlikely to be imported domestic specimens. They may represent local domestication from Italian wild boar, although it is possible that they remained wild until the end of the Neolithic (Albarella, Tagliacozzo et al. 2006, pp. 216–220). The Middle Neolithic pigs from Gotland (Fig. 5) are also large, but are associated only with wild animals and plants (domestic species had been present in the Early Neolithic, but hunting and gathering had re-advanced during the Middle Neolithic). Many teeth other than M3 have been measured; their metrical attributes are as follows: Tooth and measurement

N

Mean

SD

V

Lower dp4 L

99

21.0

0.7

3.2

WA

100

6.8

0.3

4.0

WP

99

9.7

0.3

3.5

L

88

19.1

0.8

4.0

WA

89

11.6

0.5

4.1

WP

87

13.2

0.6

4.2

L

25

25.1

0.8

3.1

WA

25

15.9

0.7

4.3

WP

25

16.8

0.6

3.8

Lower M1

Lower M2

This table pools the data from the three sites. The measurements group remarkably tightly: V is mostly lower than at Durrington Walls (cf Table 1), so only a single population is present—but this does not reveal whether the population was wild, domestic, or subject to ‘female breeding’. Non-metrical Methods A variety of non-metrical methods have been employed. These range from some that are widely and generally applicable, to others that are relevant in only a restricted set of circumstances, or are too difficult and expensive to apply widely. Researchers must decide which method(s) can most profitably be applied in which circumstances. Age at Death Age at death is often discussed as a possible means of distinguishing wild from domestic animals. The assumption is that domestic animals are killed younger, because herders can control and manage their slaughter to maximise productivity. Such arguments are often applied to pigs. Three instances were mentioned above, but as this table shows, the contexts differed greatly:

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Site

Were other domestic species present?

Did the pigs show a size reduction?

Hagoshrim

Yes

Yes

Grotta dell’Uzzo

Yes

No/minimal

Hallan C ¸ emi

No

No

The arguments are therefore complex and need to be considered carefully. Demographic analysis has recently been applied to Pre-Pottery Neolithic goat domestication in the Zagros Mountains. This work is based on (a) a large sample of modern comparative skeletons (Zeder 2001), and (b) the fact that goats are sexually dimorphic: differences in male and female slaughter patterns can be revealed by different epiphyseal fusion frequencies (Hesse 1982, 1984; Zeder and Hesse 2000; Zeder 2005, 2006, 2008). The modern skeletons show that goats vary in size for climatic reasons, just as pigs do (see above). Demographic analysis reveals that around 10,000 years ago more juveniles were killed, and these juveniles were predominantly males: most females survived into adulthood. Zeder argues that this selection for young males is a domestic pattern that could not result from hunting. The pattern appears up to 1,000 years before the first size reduction in goats, suggesting that domestication preceded morphological change (Zeder 2006, 2008). Since goats (a) are amenable to close herding, and (b) were undoubtedly domestic shortly afterwards, this argument is persuasive. Most telling is the fact that the aged females are also present in the cull: an intensive hunting strategy concentrating on young males would not include many of these. But distinguishing between intensive hunting, close herding, and a hypothetical intermediate state of ‘proto-domestication,’ is problematic. There is no unvarying ‘natural’ age structure in wild herds, no single ‘hunting pattern’, and young males may be particularly vulnerable to predation (e.g. Jarman and Wilkinson 1972; Collier and White 1976). Gazelle in the Levantine Epipalaeolithic and early PPN provide a good example. At Nahal Oren, gazelle dominated the fauna in PPNA, and over 50% of metapodials were unfused (Legge 1972). Since juvenility was used as an argument that caprines were domestic, this ‘raises the question of the extent to which the gazelle can be regarded as a ‘‘domestic animal’’’ (Legge 1972, p. 122). More recent work has suggested that the majority of the Epipalaeolithic gazelle were probably male, although sexual separation is not as clear as in goats (Horwitz et al. 1990; Bar-Oz et al. 2004). The predominance of young males thus parallels that in the domestic goats. However, various aspects of gazelle behaviour make this species unsuitable for close herding, and Legge no longer argues for something approaching domestication: at Abu Hureyra, seasonal hunting was probably the factor causing a high juvenile kill (Legge and Rowley-Conwy 2000). Territorial behaviour by male gazelle during the mating season militates against human herders being able to control gazelle flocks. Sheep and goat do not display such behaviour, so they are a more amenable domesticate (Garrard 1984; RowleyConwy and Layton 2011). Cope argues that the gazelle pattern represents ‘proto-domestication,’ defined as ‘an intermediate area between management of wild animals and true domestication’ (Cope 1991, p. 357). Some aspects of Cope’s analysis have, however, been contested (Dayan and Simberloff 1995; Bar-Oz and Dayan 1999). Davis (2005) distinguishes three main phases of subsistence in the Levant: (1) during the Epipalaeolithic, gazelle became predominant, and the proportions of young increased;

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(2) during PPNA the prey spectrum broadened to include smaller species such as hare and fox; and (3) during PPNB domestic species appeared. Davis argues convincingly that this represents a progressive intensification, the first two phases involving intensification of hunting, as increased sedentism led to increased hunting pressure (Davis 2005). Increased hunting intensity may thus produce a kill pattern approaching that of a domestic herd. Modern studies have confirmed this: kills dominated by juvenile males can occur in species such as bison, both European (Krasinski 1978) and American (Van Vuren and Bray 1986), grizzly bear (McLellan and Shackleton 1988), and Bennett’s wallaby (Driessen 1992). Needless to say, in none of these species is intensive hunting a precursor to ‘protodomestication’. Problems increase for archaeologists when assemblages contain an increased frequency of juveniles, but where a male dominance cannot be demonstrated. High juvenile kills often indicate increased hunting intensity, not domestication. In a classic paper, Elder demonstrated that deer jaws from prehistoric sites in Missouri were predominantly adult; fewer adults were killed in the historic period, as hunting was intensified; and in the twentieth century 30% of the kill were fawns (Elder 1965). Deer behaviour may account for this: in Cervus canadensis young males are driven away from their mothers by the dominant stags as the rut approaches. These inexperienced young males range widely and behave erratically (Altmann 1960), thus becoming more vulnerable to hunting. The increase in young noted by Elder (1965) thus probably did include a majority of males, even though the mandibles were not sexed. This effect is exacerbated in animals that normally produce twin offspring, because the live population contains a greater proportion of juveniles. Roe deer are an example—one population when totally exterminated comprised 43% fawns and 19% between 1 and 2 years (Andersen 1953, fig. 3). During the summer, when the mature males are territorial for several months, juvenile males are driven out and form mobile groups, joined by the juvenile females when the does give birth. The juvenile groups are more vulnerable and exposed to greater hazards (McDiarmid 1978). This is probably why yearlings dominated the summer roe deer kill at Star Carr (Legge and Rowley-Conwy 1988, pp. 40–42)—like gazelle, roe deer cannot be close herded (Rowley-Conwy 1986, pp. 26–27). These problems culminate in wild boar. Firstly, sexual determination of post-cranial elements is difficult, and the many juveniles in most assemblages would make this still more problematic. Adult jaws can be sexed by the canine, but jaws are often broken, and the juveniles cannot be determined. Secondly, pigs have multiple offspring—Oliver et al. (1993, p.115) give a range of 4–7 for European wild boar. Wild boar populations therefore contain more young, and have a higher natural juvenile mortality rate, than other large mammals (Jezierski 1977). One five-year study produced the following age/sex breakdown (calculated from Andrzejewski and Jezierski 1978, Table 5): Up to 1 year

Yearlings

Over 1 year

Males

158

65

Females

165

79

36 98

Total

323

144

134

Age class as % of total

54

24

22

% Males in age class

49

45

27

These figures do not include an estimated 15% mortality rate among piglets under 3 months; if these are added, the first age class rises to 60% of the total. Males suffer

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higher mortality than females at 2–4 years of age (Jezierski 1977, pp. 341–342, table 3). After weaning in summer, groups comprise two or three females and their offspring, the females collectively defending the young (Haber 1961, p.75). When the adult males join the females for the rut in late autumn, the young males are driven away; subsequently they rejoin the females but again leave when the birth season approaches in spring. Female offspring stay with their mothers for longer (Fra¨drich 1971, p.135). Much juvenile mortality results from the separation of the inexperienced young from their mothers (Oliver et al. 1993, p.115). High juvenile kill rates are thus a likely outcome of various hunting strategies. Human hunters may achieve a high juvenile kill rate in two different ways: tactical or strategic. The tactical way need not involve intensification. At a summer campsite, the wild boar population would include many spring-born piglets, while in winter the inexperienced young males would be easy targets. Even low-intensity hunting would thus result in many young. This probably occurred at Early Mesolithic sites in Denmark, occupied in summer (Rowley-Conwy 1993). Kill ages are plotted in the lower part of Fig. 13; some sites have high proportions of first-summer animals. The strategic way would result from intensification: if the human population increased and became sedentary, wild boar would face increased hunting, and all year round, not just seasonally. Hunting intensification would be part of the strategy for maintaining the large sedentary settlement. This would probably increase the frequency of juveniles in both the live population and the kill. The Late Mesolithic sites plotted in Fig. 13 were occupied for much or all of the year (RowleyConwy 1998, 1999), and the high frequency of juveniles may result from this. The strategic option might result in a further increase in juvenile kills, in a most interesting way: the deliberate hunting of younger animals. This can increase the size and productivity of the hunted population, even though it runs counter to Western notions of ‘sportsmanship’ and preserving the young. Beaver produce more offspring when their populations are low and food plentiful. Various North American groups exploited this by targeting young beaver, thus ‘tricking’ the adults into producing larger numbers of offspring (Dods 2002, pp. 481–482). Modern moose hunters in Quebec, encountering a female and two calves, are advised to shoot one of the calves, not the female, because this increases the moose population in future years (Grenier 1979; see Rowley-Conwy 2001b, fig. 3.8). The way this might work for wild boar is shown in Fig. 14. This compares the outcomes of two hypothetical hunting strategies, assuming that a sow will produce five piglets and successfully wean four of them. The strategy on the left shows the outcome if a hunter encounters a family group, and kills the sow. The model assumes that, without her protection, two more of the piglets will die. By year 4, the two surviving piglets and their year 2 offspring will be adults, a total of ten; and their year 3 litters will add eight piglets in year 4. The strategy on the right shows the outcome if a piglet is killed, assuming that the three remaining weaned piglets are successfully reared. The sow and her year 1 and 2 piglets amount to nine adults in year 4, while her three surviving piglets and their year 2 litters make a further fifteen; and the year 1 litter will have produced further piglets in year 3. Killing the piglet is clearly the optimal choice for wild boar population management. The outcomes of these models will vary for many reasons (e.g. hunting methods), and also if the inputs (number of births, survivorship etc.) are changed, but Fig. 14 shows clearly how a strategy of hunting juveniles can be beneficial in the longer term. Such a strategy might be deliberately designed and fully understood, as the North American beaver hunting strategy apparently was (Dods 2002). Alternatively it might come about by accident, as cultural selection spread the most successful long-term hunting strategy

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Fig. 13 Age at death of pigs in Denmark and Gotland. Sludegaard, Nivaa, Kongemose and Bloksbjerg from Dobney et al. (in prep.), the rest from Rowley-Conwy unpublished records (modified and extended from Rowley-Conwy 2004, fig. 3)

throughout a human population. Sedentary hunter-gatherers are sometimes territorial and claim ownership of particular resources (e.g. Rowley-Conwy 2001b), and populations of wild boar might possibly be ‘owned’ by particular groups. The juvenile hunting strategy is, however, ‘maintenance by hunting,’ not ‘proto-domestication’: it does not imply that the wild boar are at the start of a process leading towards domestication. The conclusion of this discussion is that consideration of age at death is likely to be less helpful in pigs than in other species. While domestication will always lead to a high

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Fig. 14 Outcomes of two hypothetical hunting strategies, killing either an adult sow (left), or one of her piglets (right). See text for explanation. Note that descendants of the original sow are coloured white, those of her piglets pale grey, dark grey, and black

juvenile kill, hunting will often do so as well, in a variety of possible ways—and other categories like ‘semi-domestic’ will be even harder to diagnose. Age at death can therefore at best provide a supporting argument, considered alongside other lines of evidence. For the three sites mentioned at the start of this section, the increase in juveniles at Hagoshrim supports the other lines of evidence indicating domestication; at Grotta dell’Uzzo it is inconclusive; and at Hallan C¸emi it does not suggest domestication—though a ‘female breeding’ strategy cannot be ruled out. The suggestion by Zvelebil (1995, pp. 94–97; 2008, p.31) that an increase in juvenile pigs in Late Mesolithic Scandinavia indicates domestication is therefore unfounded; and in any case, no such increase occurred (see Fig. 13). For Neolithic Italy, age structure does not resolve the problem at Molino Casarotto. Jarman (1976, p. 528) states that 25% of the pigs there died in their first year, a further 25% in their second. Many of the Mesolithic sites in Fig. 13 have more juveniles than this, so the Molino Casarotto figures may equally well indicate hunting. Finally, the proportions for jaws from the Neolithic graves on Gotland are shown in Fig. 13 (top). These graves contain

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a higher proportion of juveniles than any Mesolithic site plotted—although not much greater than at Mullerup or Kongemose. The Gotland settlements were occupied all year (Rowley-Conwy and Stora˚ 1997), Gotland is not a large island, no other domesticates were present, and pigs were the only large terrestrial mammal. If the pigs were wild, intensive hunting would definitely be expected; the pattern in Fig. 13 may therefore not prove that they were domestic. Finally, the mandibles were in human graves, and we do not know on what grounds these specific ones were selected for this ritual purpose. Biogeography: Pigs on Islands A common method of identifying domestic animals in the archaeological record is to spot those that appear outside their wild range. Sheep are not native to Europe, for example, so when they appear in the Neolithic they must be domestic, brought in by humans. This approach is much less useful for pigs. They are dispersed across much of Eurasia (Oliver et al. 1993, fig. 10), so there are rather few areas where human agency is the only way that pigs could have arrived. Mostly these are restricted to islands which wild boar could not reach by swimming. Pigs transported to islands must have been under human control at least for the duration of the voyage. They would often be part of a transplanted agricultural system, remaining under close domestic control. But there are other possibilities. Domestic animals may escape and become feral, or they may be deliberately released to found a feral population for hunting—this has been repeatedly done by European explorers, for example Columbus in Cuba in 1493, and Cook in Tasmania in 1776 (Holden 1994, pp. 23, 27). Finally, wild animals may be captured, taken to an island, and released to form a new wild population. Such ‘seeding’ of islands by animal translocation has a very long history: the earliest instance so far known is the movement of cuscus (a marsupial, Phalanger orientalis) to New Britain 23,500 years ago (Flannery and White 1991; Swadling and Hide 2005). Much of our understanding of pigs on islands depends on the context in which they occur. The Vikings took pigs to the Faroe Islands and Iceland in the North Atlantic as part of a full agricultural package, attested both archaeologically and historically (Church et al. 2005). Domestic pigs were part of Neolithic economies taken to the Philippines (Piper et al. 2009), and beyond the Wallace Line into Island Southeast Asia and on into the Pacific (Dobney et al. 2008); and to the East Mediterranean islands of Cyprus (Davis 2003; Vigne et al. 2011), and Crete (Isaakidou 2008). There is no reason to doubt that these introduced animals were fully domestic—even though feral populations became established in Crete and New Guinea, and even though fallow deer were introduced to Cyprus at an early time to form a wild population for hunting. Introductions of wild boar to offshore islands are also known. Wild boar were introduced to Ireland during the Mesolithic. There is no suggestion that either the Irish animals or their parent stock (wherever they lived) were anything but fully wild (McCormick 1999; McCormick and Murray 2007). They were introduced to Cyprus during the Epipalaeolithic; although it has been suggested that the mainland wild boar were ‘managed’ (Vigne et al. 2009), they need not have been so. Wild boar were introduced during the Early Holocene to Okinawa (Matsui et al. 2005) and the Izu Islands (Yamazaki et al. 2005). In contrast to the Irish boar, the Cypriot, Okinawan and Izu boar subsequently reduced in size (modern Okinawan wild boar are the world’s smallest), but a review of wild boar sizes across the Old World shows that this follows the expected pattern: wild boar become smaller on small islands like Okinawa, but Ireland is so large that this did not happen (Albarella et al. 2009). A recent study has found no evidence that mainland Japanese boar

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were managed (Hongo et al. 2007), so the situation there appears similar to Ireland except for the insular dwarfing. The problems are greatest when the status of the parent population is unclear. Pigs were introduced into Corsica and Sardinia at the start of the Neolithic; Mesolithic sites on these islands contain none. They most probably came from Italy, and were the same size as the Neolithic pigs at Grotta dell’Uzzo (Albarella, Manconi et al. 2006, p. 292)—but the problems with this site were noted above. Wild, feral and domestic pigs are now present on Corsica and Sardinia. Insularity has had a major effect: size has reduced since the Neolithic, and the modern wild boar are the second smallest in the world, after Okinawa (Albarella, Dobney and Rowley-Conwy 2009). Red deer were introduced in the Neolithic (Vigne 1988)—but, as with the fallow deer on Crete (see above), this has no bearing on whether the pigs were wild or domestic. Perhaps the Neolithic pigs were domestic when introduced (as suggested by Vigne 1988), but some soon became feral and established the populations now considered wild; this remains uncertain. Also uncertain is the status of the Gotland Middle Neolithic pigs. No wild boar were present in the Mesolithic; domestic animals were imported in the Early Neolithic (Lindqvist and Possnert 1997). The Middle Neolithic saw a withdrawal of farming and the re-establishment of hunting and gathering (Rowley-Conwy 1999, fig. 12). The Middle Neolithic pigs might be the feral descendants of Early Neolithic domesticates. However, pigs in Australia that have been feral for 150 years retain the metrical attributes of their domestic ancestors, while the Gotland specimens align with wild boar in this respect (Rowley-Conwy and Dobney 2007, figs. 7.11 and 7.12). The Gotland pigs are a little smaller than mainland wild animals (Fig. 5), but this could be accounted for by insularity. ¨ sterholm (1989) has suggested that the pigs were domestic because they were imported O by humans. This may be so, but as we have seen above, it need not be so. Had the pigs been killed for a funerary feast, this would have implied that they were readily available and hence under close control. However, the pigs were in fact killed throughout the year and the jaws cached for funerary use (Rowley-Conwy and Dobney 2007, fig. 7.16), so this line of enquiry is also inconclusive. The status of pigs on islands is thus not always easy to resolve. Where the context is understood—that is, where we already ‘know the answer’—the situation is easier, as in Ireland, Crete, Iceland and the Faeroes, the Izu Islands, the Philippines, and Island Southeast Asia. But presence on an island by itself is no necessary indication of domesticity. Linear Enamel Hypoplasia The analysis of Linear Enamel Hypoplasia (LEH) is an area which is yielding interesting results. LEHs are abnormal incremental lines in tooth enamel, resulting from a period of physiological stress affecting the animal during the growth of the tooth. After the period of stress ends, normal growth is resumed; the stress episode is visible as a line or series of pits in the normal enamel; see Dobney and Ervynck (1998) and Dobney et al. (2002) for details and recording method. LEH yields information in two ways. First, the overall frequency of LEHs indicates the general level of stress to which a population was exposed. Second, the relative chronology of stress events can be examined. This is because the enamel crowns of M1, M2 and M3 form in that order, and from the tip of each cusp towards the root. Mandibular teeth are those most usually examined. The crown of lower M1 is in formation when piglets are born, and birth often leaves an LEH line some 5 mm above the bottom edge of the enamel. Another concentration often appears near the bottom of the crown of lower M1, probably

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associated with weaning. A concentration towards the bottom of lower M2 is likely to indicate dietary stress in the animal’s first winter, while another in lower M3 corresponds to the second winter (Dobney and Ervynck 2000; Dobney et al. 2004). Modern wild boar populations generally have low levels of LEHs, suggesting a low incidence of the physiological stresses implicated in LEH formation. Archaeological wild boar have similarly low frequencies, while pigs identified as domestic by conventional metrical means (see above) often have higher levels, suggesting that closely managed populations were exposed to more episodes of physiological stress (Dobney et al. 2004). Figure 15 (top) shows this for some Scandinavian assemblages. Mesolithic wild boar have consistently low frequencies. The Neolithic sites of Troldebjerg and Bundsø have higher levels. Wild and domestic populations are not always clearly separate, however: the Bronze Age domestic pigs in Fig. 15 (top) have frequencies as low as the wild populations. Some other domestic populations also have low frequencies: the PPN early domestic pigs at C¸ayo¨nu¨ had low levels, though these increased somewhat in the PN (Ervynck et al. 2002). Neolithic domestic pigs in China have higher frequencies than archaeological wild boar, thus conforming to the general European pattern (Dobney et al. 2007). Also plotted in Fig. 15 (top) are the Middle Neolithic pigs from Gotland, which have the highest values of any of the plotted populations. These pigs were clearly suffering high levels of physiological stress. But is this an unequivocal sign that they were domestic? The situation is complex. Figure 15 (bottom) shows the distribution of LEHs across the molar teeth. The domestic pigs at Troldebjerg have quite high frequencies on M1 (second cusp), corresponding to weaning (see above), and high levels on M2, forming during the first winter. This pattern is commonly found in domestic animals (Dobney et al. 2002, fig. 4; Dobney et al. 2004, figs. 4, 5 and 7). The Gotland pattern differs in having the highest levels on M3, forming during the animals’ second winter. One possibility to be considered is that this could result from intensive hunting, expected on Gotland in the Middle Neolithic (see above). It was noted above that mortality is high among male wild boar of this age, and the repeated disturbance of wild populations by frequent hunting might be a factor causing high levels of physiological stress. Jomon Japan also has high frequencies of LEH (Dobney et al. 2007). Jomon pigs were, however, almost certainly wild (Hongo et al. 2007). Jomon hunter-gatherers were often sedentary and achieved high population densities (e.g. Habu 2004; Imamura 1996; Matsui 1996). Wild boar populations within reach of large, sedentary Jomon settlements would probably be under considerable hunting pressure, which could be a cause of stress. However, if so, it operated differently to Gotland, because the Jomon pigs have high levels of LEH on M1 and M2, but not on M3 (Dobney et al. 2007, fig. 4.3). The analysis of LEH is at a relatively early stage, but is creating interesting patterns. While it is not an absolute indicator of domestication, high levels of LEH are more characteristic of domestic than wild populations in the samples studied to date. The high levels of LEH in the Gotland pigs could suggest that they were domestic, but more work is needed to reveal the effects of intensive hunting before this can be safely concluded. Palaeodiet: Stable Isotopes and Tooth Microwear The diets of domestic pigs (at least if under close human control) are likely to differ from those of wild boar. This section discusses two methods by which pig diets may be examined. Isotopic analysis is beginning to produce very useful information, which, particularly in East Asia, is casting light on the status of early pigs. Stable isotope ratios of nitrogen and carbon in bones and teeth reveal different aspects of the diet (see Fig. 16 top).

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index

1

0

Bundsø

Bronze Age

Late Mesolithic

Troldebjerg

Gotland

Jutland

Zealand

Middle Mesolithic

Early Mesolithic

-1

average number of LEH lines per cusp

1.0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0

M1 first

M1 second

M2 first

M2 second

M3 first

M3 second

Late Mesolithic Zealand

Troldebjerg

Late Mesolithic Jutland

Gotland

Fig. 15 Top frequency of Linear Enamel Hypoplasia in various assemblages of pigs from southern Scandinavia; the index is the mean of all the assemblages plotted (after Rowley-Conwy and Dobney 2007, fig. 7.13). Bottom frequency of LEH in different cusps of M1, M2 and M3 from southern Scandinavian assemblages (combined from Rowley-Conwy and Dobney 2007, figs. 7.14 and 7.15)

A higher d15 N ratio indicates a higher position in the food chain, involving a greater intake of animal products. An increase in d13C indicates an increase in either marine foods, or C4 plants, which include the millet species. C3 plants comprise the majority of terrestrial plants including wheat and barley. Being omnivores, both wild boar and domestic pigs can consume a wide variety of foods (Masseti 2007), so the patterns are potentially complex. Figure 16 shows various applications. The Chinese Neolithic sites of Xipo and Kangjia (7000–4000 BP) have pigs with very high d13C (Fig. 16 top). Since the sites are inland (Fig. 7), this must indicate that the pigs were consuming C4 plants. In China this must mean millet: people were feeding this non-native cultigen to their pigs. This in turn

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Fig. 16 d15 N and d13C ratios from various sites. Top Awazu and Gushibaru from Minagawa et al. (2005, table 1); Kangmung Dong from Matsui et al. (2005, fig. 5); Xipo and Kangjia from Pechenkina et al. (2005, table 7). Middle Nevalı C¸ori from Lo¨sch et al. (2006, table 3). Bottom Va¨sterbjers from Eriksson (2004, table 3); Korsna¨s from Fornander et al. (2008, table 3)

indicates that the pigs were domestic (Pechenkina et al. 2005). In contrast the wild boar from Awazu in Japan (4000–4500 BP) were consuming a ‘conventional’ terrestrial diet (Minagawa et al. 2005). Later sites have more pigs apparently fed on millet, for example Kangmung Dong in Korea (first century AD), and two animals from Gushibaru on Okinawa (500 BC–AD 300). Kangmung Dong is coastal, but the pigs did not consume much marine food: had they done so, their d15 N ratio would also have increased, because most marine foods are of animal origin (Matsui et al. 2005; Minagawa et al. 2005).

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The Gushibaru pattern is complex, forming three groups (Fig. 16 top). The two milletfed individuals have been mentioned. Another group overlaps with the wild boar from Awazu, while the most numerous group lies directly above these, having an elevated d15 N ratio. This is consistent with the pigs foraging around human settlements, eating agricultural food waste and human faeces. But the inhabitants of Okinawa apparently were not agricultural at this time—and the two-fold d15 N pattern is found at much earlier sites, such as Noguni B (4500–7000 BP), long before agriculture became established. One possibility is that the pigs were raised on agricultural settlements away from Okinawa (some fed on human waste, some on millet) and taken there by sea (Matsui et al. 2005; Minagawa et al. 2005); pigs were certainly domesticated in China much earlier (Yuan and Flad 2002). Early domestication is less easy to see when C4 plants are not present. The PPNB site of Nevalı C¸ori in Turkey provides an example (Fig. 16 middle). Lo¨sch et al. (2006) suggest that the pigs were domestic, but despite consuming human waste their d15 N was lowered because much of the waste came from pulses; these plants contain low levels of d15N. However, the Nevalı C¸ori pig isotopes correspond closely to wild boar (cf Awazu in Fig. 16 top), and to other species from Nevalı C¸ori, including sheep (which were domestic) and gazelle (which were not). The osteological evidence regarding the Nevalı C¸ori pigs is not clear (Peters et al. 2005). The isotopic evidence from Nevalı C¸ori is thus not easy to interpret. Isotopic studies have produced important results on Gotland (Fig. 16 bottom). Various species have been analysed at Va¨sterbjers (Eriksson 2004), and contemporary Korsna¨s on the Swedish mainland (Fornander et al. 2008). One argument that the Gotland pigs were wild was that there was no economic niche for them on a hunter-gatherer settlement generating no agricultural waste (Rowley-Conwy and Stora˚ 1997). However, it has been persuasively argued that if the settlement had a maritime economy, it would generate much fish and marine mammal waste, which the omnivorous pigs could consume (Masseti 2007). Isotopic analyses however reveal that the pigs at both Va¨sterbjers and Korsna¨s had a terrestrial diet (Fig. 16 bottom, cf Awazu and Nevalı C¸ori), as did those from Ko¨pingsvik (Eriksson et al. 2008). This contrasts markedly with the domestic dogs on all three sites, who ate so much marine food that they fall close to the seals. Thus the dogs were scavenging on the settlements, while the pigs were not. This makes it much more difficult to argue that the pigs were domestic. Microwear on pig teeth can distinguish between free-ranging animals that are able to root, and stall-fed animals that are not. The diet of the free-ranging pigs contains more abrasive material, so the tooth surfaces exhibit much more pitting and scratching (Ward and Mainland 1999). If early domestic pigs were stall-fed, this should therefore be detectable; but if they were allowed to range freely around the settlement their diet and behaviour would closely mimic wild boar. It is not known when pigs in any region began to be stall-fed. The East Asian pigs whose isotopes reveal that they were fed on millet (see above) would presumably show a stall-fed pattern, and microwear has identified stall-fed pigs at the Iron Age site of Elms Farm in Britain (Wilkie et al. 2007). It would be interesting to apply this method to Neolithic domestic pigs at European sites where osteometric methods reveal a close-herded population (see above). Population History: Genetics and Geometric Morphometrics Population history yields insights into pig domestication. DNA studies have concentrated on the non-mutating mitochondrial DNA (mtDNA), inherited only through the female line. There is substantial geographical variability in modern wild boar. Fourteen major Old

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World clusters have been identified (Larson, Dobney et al. 2005; Larson, Albarella, Dobney and Rowley-Conwy 2007; Larson, Cucchi and Dobney 2011). If domestic pigs carry the mtDNA of a particular wild boar cluster, pigs were domesticated in the area occupied by that cluster. Two of the major clusters are found in Europe: one in Italy only, the other across the rest of the subcontinent. A third cluster appears in the Near East. Many modern domestic pigs carry European mtDNA, so wild boar were domesticated in Europe. Remarkably, Near Eastern mtDNA is (with a single exception) not known in modern domestic pigs (Larson, Dobney et al. 2005). Ancient DNA from Neolithic pig bones from various sites across Europe has, however, shown that Near Eastern pigs were domesticated, and were carried into Europe in the Neolithic (Larson, Albarella et al. 2007). Sites in southeastern Europe have large wild boar and small domestic pigs (see the morphometric section). The wild boar belong to the European cluster, the domestic pigs to the Near Eastern one. But further west, small domestic pigs appear that belong to the European cluster, indicating local domestication. The oldest are at Bercy in France (Fig. 1), also the westernmost site with individuals from the Near Eastern cluster. By the Bronze Age, the Near Eastern cluster has disappeared from Europe. Thus in most of Europe the mtDNA pattern is consistent with morphometric information and indicates reasonably clear conclusions. In southern Europe the situation is less clear. The Near Eastern cluster contains two variants, and the central European Neolithic pigs carry just one of these. The other Neolithic expansion route into Europe, along the Mediterranean coasts, might possibly have involved the other Near Eastern variant. The sole modern domestic pig with Near Eastern mtDNA carries this variant, and comes from Corsica (Larson, Albarella et al. 2007). One Bronze Age specimen from Sardinia belongs to the Italian cluster, so at least some pigs were introduced to Sardinia from Italy. They were probably domestic—but not definitely so: wild boar may be carried to islands to found populations for hunting (see above). Several East Asian wild boar lineages appear in domestic pigs, indicating multiple domestication events (Larson, Dobney et al. 2005). Watanobe et al. (1999) demonstrated that modern Japanese wild boar are fairly closely related to mainland East Asian wild and domestic pigs, but wild boar on Okinawa are not closely related to those in Japan. The origin of the Okinawan animals was therefore unclear, until Hongo et al. (2002) showed that wild boar in Vietnam carried the same mtDNA lineage. This suggests that Okinawan wild boar are not feral, but an original wild boar population isolated by the rising postglacial sea (Hongo et al. 2002). Ancient DNA has shown that domestic pigs of the last 2,000 years in both Okinawa and the main Japanese islands are genetically similar to domestic specimens on the East Asian mainland, suggesting an introduction from there (Watanobe et al. 2002; Matsui et al. 2005). Poor preservation unfortunately means that little mtDNA has been recovered from earlier sites on Okinawa, and few metrical data can be taken (Matsui et al. 2005), so the possibility of an earlier import of pigs suggested on isotopic grounds (see above) cannot be further elucidated by these means. The pattern in Southeast Asia is complex. Two major wild boar clusters are found: one in Island Southeast Asia and the Malay Peninsula, incorporating Sus scrofa and related species, the other on the continental mainland (Larson, Cucchi et al. 2007). Three movements involving these clusters can be distinguished. The first is a movement of pigs from Sulawesi to Flores around 7000 BP, inferred from ancient DNA. Other domestic species were not taken to Flores so early, so the status of these pigs remains unknown; they could have been wild. The two later movements both involve the transporting of the mainland cluster. Remarkably, ancient and modern DNA reveals that one variant of the mainland cluster extends from Vietnam, through Sumatra and Java to New Guinea, and on to Near and

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Remote Oceania. In Vietnam, the pigs of this variant are wild; but the others are associated with other domestic species and form part of the Neolithic economy. The first pigs in Hawaii were thus the domestic descendants of Vietnamese wild boar. The other later movement was of pigs from eastern China to Taiwan, the Philippines, and the Mariana Islands (Larson, Cucchi et al. 2007; Dobney, Cucchi and Larson 2008; Cucchi et al. 2009). The date of the last one is at least as early as 4,000 years ago, since domestic pigs occur in the Philippines at this time (Piper et al. 2009). Geometric morphometrics is an exciting new line of study in the context of pig domestication. When applied to teeth, the technique involves constructing a detailed mathematical description of the shape of a tooth, independent of its size. Since shape of mammalian teeth does not vary much due to environmental factors, but is genetically controlled, mathematical comparisons between teeth are analogous to DNA comparisons, with the added advantage that every surviving tooth can provide information—while ancient DNA often does not survive (Dobney et al. 2008). Few applications of the technique have so far been published. Warman (2005) distinguished variations between some modern breeds. In Southeast Asia, the technique produces the same groupings as the genetic evidence (Larson, Cucchi et al. 2007; Dobney, Cucchi and Larson 2008). There is clearly potential for this method to be applied much more widely. Population history can thus reveal the origins of domestic pigs, as in Europe and Insular Southeast Asia, but it does not solve all problems, for example the status of the pigs transported to Flores and Sardinia. The Gotland pigs have not yet been examined. No doubt they belong to the European genetic cluster, but in itself this would tell us nothing—further work might be revealing. Within Japanese wild boar, there is considerable local geographical variability (Okumura et al. 1996). If such variability were also present in Europe, populations east and west of the Baltic Sea might show differences. The Early Neolithic domestic economy came to Gotland from southern Scandinavia. Ancient DNA of the Middle Neolithic hunter-gatherers, however, reveals that they came from the Northeast Baltic (Linderholm et al. 2008; Malmstro¨m et al. 2009). If the Gotland pigs also came from there, they would have been brought to the island by hunter-gatherers—suggesting that they were wild.

Conclusions It was stated in the introduction that pigs are the most difficult of the major domestic animals to categorise as ‘wild’ or ‘domestic’. It will by now be clear why: no method of separation provides a perfect answer by itself. That said, we have shown that pigs can be diagnosed as wild or domestic more often than is generally supposed, and much more often than theoretical problems such as ‘semi-domestication’ might lead us to believe. The closest to certainty so far achieved is when metrical analysis produces a two-population pattern. In such cases we can conclude that a close-herded domestic population was present—even though some individuals of intermediate size cannot be diagnosed, and a precise ratio cannot be established. The two-population pattern is found remarkably widely in time and space, demonstrating the existence of close-herded pigs more often than we might have expected on theoretical grounds. The non-metrical methods have not so far raised serious questions in relation to this conclusion. But many things remain unknown. We know that pigs were domesticated several times—but we know little of the circumstances of initial domestication. The size change associated with domestication is most clearly visible at C¸ayo¨nu¨ (Fig. 8), but interpretation

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of this is not straightforward. Perhaps domestication was a long drawn-out process paralleling the size change; or perhaps it was an abrupt event at the start of the size change that exerted a continuous selective pressure on size from then on. We also cannot distinguish between hunting and ‘female domestication’, where domestic females breed with wild males. This must be a high priority for future research: if we can start to detect ‘female breeding’ in the zooarchaeological record, we may be able to see whether this forms a stage between hunting and full domestication—or whether it is a practice unique to New Guinea. Many currently accepted conclusions have emerged because multiple lines of analysis have been applied. The best strategy for future work is simply to continue this, improving the methods we have and developing new ones, re-evaluating previous conclusions as we go. The pigs from Gotland have been considered in five different places above, and illustrate this process, as the following table shows: Approach

Result

Conclusion

Metrical

Large size, single population

Wild?

Age at death

High proportion of juveniles

Domestic?

Biogeography

Introduced on to island

Domestic?

Hypoplasia

High frequency

Domestic?

Isotopes

Terrestrial diet

Wild?

No certain conclusion can be reached at this stage, but two points are worth noting. First, it can be argued that the isotopes are the most revealing of these approaches: if the pigs had been kept on the settlements of maritime hunter-gatherers, their diet should have included marine waste—but it did not. Second, there are alternative arguments in each of the three approaches for which the conclusion is ‘domestic’: the high frequency of juveniles and hypoplasias might have been the result of intensive hunting, and the pigs might have been taken to Gotland to form a population for hunting. For these reasons our tentative conclusion is that the Gotland pigs were wild—but the work that proves this wrong may be being carried out as these words are being written. Acknowledgments This paper is an outcome of the Durham Pig Project, funded by the Wellcome Trust (GR060888, and GR071037) and the Arts and Humanities Research Board (B/RG/AN1759/APN10977), and we thank both organisations. Simon Davis, Annat Haber and Akira Matsui provided useful references and much help. Terry O’Connor and an anonymous referee provided comments that have materially improved this paper.

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