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Home , Archaeobiology, Domestication, Fertile Crescent, Founder crops

PERSPECTIVE

Domestication and early in the : Origins, diffusion, and impact

Melinda A. Zeder* Archaeobiology Program, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013

Edited by Jeremy A. Sabloff, University of Pennsylvania Museum of and , Philadelphia, PA, and approved May 27, 2008 (received for review March 20, 2008)

The past decade has witnessed a quantum leap in our understanding of the origins, diffusion, and impact of early agriculture in the Mediterranean Basin. In large measure these advances are attributable to new methods for documenting in and animals. The initial steps toward and animal domestication in the can now be pushed back to the 12th millennium cal B.P. Evidence for management and crop cultivation appears at least 1,000 years earlier than the morphological changes traditionally used to document domestication. Different seem to have been domesticated in different parts of the Fertile , with genetic analyses detecting multiple domestic lineages for each species. Recent evidence suggests that the ex- pansion of domesticates and agricultural economies across the Mediterranean was accomplished by several waves of seafaring colonists who established coastal farming enclaves around the Mediterranean Basin. This process also involved the adoption of do- mesticates and domestic by indigenous populations and the local domestication of some endemic species. envi- ronmental impacts are seen in the complete replacement of endemic island by imported mainland and in today’s anthropogenic, but threatened, Mediterranean landscapes where sustainable agricultural practices have helped maintain high bio- diversity since the .

archaeology ͉

he transition from foraging and fusion, and impacts of domesticates and therein). Rather than domestic status, to farming and herding agriculture in the Mediterranean Basin, sex is the primary factor affecting body is a significant threshold in hu- outlining our current understanding of size in these ungulates, manifested by a man history. Domesticates and these developments and highlighting marked and consistent difference be- T tween larger males and smaller females the agricultural economies based on promising areas for future study. them are associated with radical restruc- in essentially all skeletal elements. Envi- turing of human societies, worldwide Initial Animal Domestication in the ronment also strongly influences body alterations in , and signifi- size, with increasing heat and aridity cant changes in the ’s landforms Until the late 1990s archaeozoologists positively correlated with smaller size. and its atmosphere. Given the momen- relied on morphological changes in tar- What archaeozoologists had originally tous outcomes of this transition it comes get species to identify where and when interpreted as body size reduction asso- as little surprise that the origin and wild prey animals were transformed into ciated with initial domestication can now be attributed to differences in the spread of domesticates and the emer- herded livestock (2). A proposed sharp culling strategies of herders as opposed gence of agriculture remain topics of and rapid reduction in overall body size to hunters. In most prey species, hunters enduring interest to both the scholarly among archaeological prey populations focus on large adult animals (particu- community and the general public. was the most widely accepted morpho- larly males) to maximize return, and the The past decade has seen remarkable logical marker of this threshold (3, 4). bones of these larger animals generally analytical advances in documenting do- Based on this size reduction criterion, dominate in prey assemblages generated mestication (1), particularly in tracking the established consensus was that ani- by hunters. Archaeological assemblages the domestication of four major Near mal domestication (beginning with generated by herders, on the other Eastern livestock species (, goats, and then sheep) occurred at ca. 10,000– hand, are usually dominated by the , and ) and their subsequent 9,500 B.P.†, Ϸ1,000 years after the bones of smaller females slaughtered dispersal throughout the Mediterranean domestication of crop plants in the after their prime reproductive years. Ex- Basin. New morphometric methods are southern (3, 5). Domestication of cess males not needed for herd propaga- tracking changes in human prey strate- these two animal species was thought to tion were harvested at young ages and gies that mark the transition from hunt- have occurred somewhere to the north their more friable bones are usually less ing to herding. Genetic analyses bring and east of the heartland of plant do- represented in these assemblages. fresh insights into initial livestock mestication (5), although a second, inde- Although the linkage between domes- domestication and their dispersal. Small- pendent domestication of goats was tication and body size was called into sample atomic mass spectrometry proposed for the (6). radiocarbon dating provides refined The utility of this size reduction chronological frameworks for these de- marker, and indeed of all morphological This article grew out of a presentation given by M.Z. at the Calpe 2007 Symposium: People in the Mediterranean–A velopments. These recent analytical markers, has come under increasing History of Interaction, September 27–30, 2007, the Gibraltar advances, in turn, have produced an ex- scrutiny (7). My own work on both mod- Museum, Gibraltar. plosion of new information that is call- ern skeletal collections and archaeologi- Author contributions: M.A.Z. designed research, per- ing into question prevailing hypotheses cal caprine (sheep and ) remains formed research, synthesized research in referenced publi- about the origin and early spread of ani- from the finds little support cations, and wrote the paper. mal domesticates and the Neolithic life- for the almost axiomatic acceptance that The author declares no conflict of interest. ways of which they were a part. Here, I domestication results in an automatic This article is a PNAS Direct Submission. bring together these different sources of overall reduction in body size in man- *E-mail: [email protected]. information to consider the origins, dif- aged animals (ref. 8 and references †All dates are reported in calibrated years before present.

www.pnas.org͞cgi͞doi͞10.1073͞pnas.0801317105 PNAS ͉ August 19, 2008 ͉ vol. 105 ͉ no. 33 ͉ 11597–11604 Downloaded by guest on September 27, 2021 question by this research, the marked degree of sexual dimorphism in caprines it documented offered another approach to tracking the transition from hunting PIGS SHEEP to herding. Pronounced differences in 10,000 10,500 the size of male and female skeletal 10,000 11,000 8,500 CATTLE GOATS 10,000 elements make it possible to separate 8,500 11,000 archaeological assemblages into 9,600 9,000 sex-specific subpopulations, which, based 10,500 10,500 on a refined understanding of the se- 10,500 10,500 10,000 quence and timing of long bone fusion 9,000 (9), can be used to generate high- 9,600 9,500 resolution harvest profiles for male and 9,000 8,500 female animals capable of distinguishing 8,500 8,500 the prey strategies of hunters from the 8,500 harvest strategies of herders. Application of this approach to ar- chaeological assemblages from and Fig. 1. The origin and dispersal of domestic livestock species in the Fertile Crescent. Shaded areas show has identified the clear signature of the general and the approximate dates in calibrated years B.P. in which initial domestication is a managed herd of goats (harvesting of thought to take place. Dates outside of the shaded areas show the approximate date when the domes- young males and prolonged survivorship ticate first appears in a region. Orange, goats ( hircus); blue, sheep (Ovis aries); green, cattle ( of females) at the site of in taurus); fuscia, pigs (Sus scrofa). highland Iran (8). Directly dated to 9,900 B.P., the goats from this site show no evidence of size reduction or any elsewhere in the Fertile Crescent are slower, to that of sheep (Fig. 1) (10, 14). other domestication-induced morpholog- detecting parallel patterns to those doc- Morphologically altered domestic pigs ical change. Smaller body size and umented in the Zagros. Changes in the are not found in the southern Levant or changes in the size and shape of horns age of harvested caprines, and possibly lowland Iran until ca. 8,500–8,000 B.P. [a morphological change clearly linked demographically driven changes in size Recent demographic evidence suggests to domestication (2)] appear 500–1,000 consistent with early herd management, that taurine cattle were initially domesti- years later than this demographic shift, are found in southeastern at cated somewhere in the upper Eu- when managed animals were moved ca. 10,500 B.P. (12). Sheep seem to be phrates Valley between ca. 11,000 and from the natural habitat of wild goats the initial early focus of the transition 10,000 B.P. (15), but, like sheep and and introduced into hotter and more from hunting to herding in this region, pigs, they arrived relatively late in more arid lowland Iran. These follow-on mor- with managed goats arriving from out- distant parts of the Fertile Crescent phological changes likely reflect re- side the area at ca. 10,200 B.P. (12). (Fig. 1). Morphologically altered domes- sponses to new selective pressures, plus Similarly, demographic evidence for the tic pigs and cattle are not found in Cen- the now more limited opportunities for management of morphologically unal- tral Anatolia until after 8,500 B.P. (16). introgression between managed and wild tered caprines (mostly sheep) is found in Genetic data from modern and ar- animals or the restocking of with Central Anatolia between 10,400 and chaeological specimens both support wild animals. 9,400 B.P. (13). These results suggest and enhance this picture of initial ani- Looking back before Ganj Dareh, un- that both sheep and goats were brought mal domestication. Recent work has suc- usual demographic profiles detected in under domestication (probably indepen- ceeded in definitively identifying the sheep bone assemblages from northeast- dently of one another and possibly mul- progenitors of both domestic sheep and ern Iraq (10) and southeastern Anatolia tiple times) in the region that stretches goat as belonging to species found in (11) dating to 12,000 B.P. may reflect from the northern Zagros to southeast- the Fertile Crescent (Ovis orientalis and early attempts at manipulating herd de- ern Anatolia between ca. 11,000 and Capra aegagrus, respectively) (17, 18). mographics to maximize returns. These 10,500 B.P., and perhaps even earlier Moreover, in both of these livestock spe- assemblages show an almost exclusive (Fig. 1). Morphologically wild, but man- cies there are at least four and, in the focus on 2- to 3-year-old males, which is aged, goats appear to have been moved case of goats, as many as six (19), genet- older than expected with herd manage- relatively rapidly through the region, ically distinguishable domestic lineages, ment but younger than expected with reaching the southernmost tips of both or haplotypes. It is not entirely clear, hunting. This pattern is argued to result the eastern and western arms of the however, whether these different lin- from a prime male hunting strategy de- Fertile Crescent by ca. 9,500 B.P. Do- eages represent spatially and temporally veloped under conditions of intensive mestic sheep were spread more slowly discrete ‘‘domestication events’’ in which pressure on local wild herds during the and first appear in these Ϸ500– different populations of animals were climatic downturn (11). 1,000 years later than managed goats brought under domestication indepen- The proposed scenario suggests that, (10, 12). dently of one another (20). Genetic data rather than broadening the prey strategy Recent research has also clarified the for taurine cattle have identified five to include both males and females, hunt- spatial and temporal context of the do- different domestic haplotypes, at least ers conserved female breeding stock mestication of two other major livestock three and possibly four of which origi- while at the same time relying on a species in the Near East: pigs and cattle. nated in the Fertile Crescent (21). Simi- steady immigration of younger males Archaeological evidence now suggests larly, as many as four of the many drawn from surrounding territories to that pigs were first domesticated some- different lineages of domestic pigs fill the vacuum left by the kill-off of where in southeastern Anatolia by originated in the Near East (22, 23). local prime-age males. 10,500–10,000 B.P. and that the timing Animal domestication in the Near Lower-resolution demographic meth- of their geographical expansion as do- East can then be seen as arising from a ods used by archaeozoologists working mesticates was similar, although perhaps period of prolonged human interaction

11598 ͉ www.pnas.org͞cgi͞doi͞10.1073͞pnas.0801317105 Zeder Downloaded by guest on September 27, 2021 with the ancestors of core livestock spe- cies that unfolded across much of the Fertile Crescent. Over time hunting strategies aimed at maximizing local 7,8007,800 availability of wild ungulates developed 7,7007,700 into active management, with all four 8,100 7,300 major livestock species coming under 7,300 9,000-8,500 9,000-8,500 7,700 management over a period from ca. 7,7,60

7,400 600 11,000 to 10,000 B.P. Even species like 7,400 7,7007,700 0 gazelle, which are behaviorally unsuited 7,0007,000 to domestication, may have been audi- 10,50010,500 tioned for management in the southern and northern Levant, where they were 7,0007,000 the most abundant wild ungulate (11). Clear-cut morphological responses to domestication (i.e., changes in horns in Fig. 2. An integrated model of the Neolithic expansion in the Mediterranean Basin. The location of bovids and tooth size in pigs) are not colonist farming enclaves is shown in the red ellipses. Approximate dates of these enclaves are given inside evident in these four livestock species the ellipses in calibrated years B.P. Red dots represent areas that are proposed to have been settled by until ca. 9,500–9,000 B.P. colonist ; green dots indicate areas where indigenous foragers adopted elements of the Neolithic As is the case with animal domestica- package; and blue dots indicate areas of proposed integration of colonist farmers with indigenous tion in the Near East, the leading edge foraging groups. Data were complied from refs. 52, 54, 56, 57, and 65 and figure 7.1 of ref. 74. of plant domestication in the region is now recognized as an extended process (24, 25). Evidence from multiple loca- in Upper and Meso- not demic diffusion was the primary en- tions point to a prolonged period of hu- lithic levels at Francthi on the gine driving this transition. Specifically, man manipulation of morphologically eastern coast of Greece, followed by the proponents maintained that the selective wild, but possibly cultivated, plants appearance of fully domesticated adoption of various elements of the which, in certain species, resulted in the and lentils in later Neolithic levels, was Neolithic package by indigenous popula- development of morphologically altered interpreted as evidence for the local tions around the Mediterranean could domesticated crops (26–28). This period crop domestication (34). recov- have happened through trade and tech- of intensified plant management dates at ered from cave deposits in nology transfer alone without any direct least as far back as ca. 12,000 B.P., with southern France were seen as evidence contact between indigenous hunter– morphological markers of crop domesti- of incipient cultivation, if not domestica- gatherers and colonizing farming popu- cation (i.e., nonshattering seed heads in tion, of local wild plants (35). Evidence lations from the east (41–46). ) not well established until ca. for indigenous animal domestication was The shortcomings of these different 10,500 B.P. (24, 25, 29). Agricultural based on the identification of wild sheep single-agent models have become economies reliant on a mix of domesti- in age deposits in southern increasingly apparent as new archaeo- cated crops and livestock apparently do France and the presence of domestic logical data come to light and older not fully crystallize in the region until sheep and goat remains in Mesolithic collections are reanalyzed by using new ca. 9,500–9,000 B.P. (5, 10). contexts in France and Spain (36, 37). methods and new perspectives. Recent Reports of domesticated and cattle genetic analyses of livestock species and The Diffusion of Animal Domesticates in remains in Mesolithic (pre-8,000 B.P.) their progenitors have also contributed the Mediterranean Basin levels from sites in southern Spain (38) important new insights into this process. The last two decades has witnessed the were also cited as evidence for the local As a result, a much more complex, and rise and fall of a number of models of domestication of these species. more interesting, scenario is emerging Neolithic expansion across the Mediterra- Genetic studies have subsequently for the Neolithic transition across the nean Basin. In the early 1980s Ammer- ruled out European ancestry for domes- Mediterranean Basin. man and Cavalli-Sforza (30) combined tic , barley, and pulses, confirming Beginning in the early 1990s a num- archaeological and human genetic data to the Near East as the source of these ber of sites have been discovered and frame their ‘‘wave and advance’’ model. crops (26, 39). Morphological, cytologi- excavated on that have radically This model attributed the westward cal, hemoglobin, and, most recently, ge- transformed our understanding of Neo- spread of the Neolithic to Near Eastern netic studies have shown that the ‘‘wild’’ lithic emergence in the Mediterranean colonists who, driven by agriculture-fueled sheep and goats found on Mediterra- Basin (47). Until the early 1990s Cyprus population growth, slowly pushed aside nean islands, once argued to be the was thought to have been colonized ca. indigenous hunter–gatherers at a pre- descendents of the progenitors of indig- 8,500 B.P. by a derived offshoot of fully dicted average pace of Ϸ1 km per year. enous domestic caprines, are instead the established Neolithic mainland cultures Objecting to the passive role this descendents of Near Eastern ca- (48). The new sites, however, date 2,000 model assigned to indigenous Mesolithic prines (for a review in chronological years earlier (10,500–9,000 B.P.) and people, a number of researchers subse- order, see refs. 40, 41, 17, and 18). document the arrival of early pioneers quently countered with alternative mod- Ruling out the indigenous domestica- hypothesized to have originated some- els that awarded local populations a tion of caprines and major crop plants where in the Northern Levant (Figs. 1 starring role in Mediterranean Neolithic did not, however, to an embrace of and 2) (47, 49). Traveling the 60 k to emergence. Early models within this in- colonist expansion diffusion models for Cyprus by boat, these colonists trans- digenist perspective argued for autocho- the emergence of Neolithic lifeways in ported the full complement of economi- nous domestication of crops and live- the Mediterranean. Instead, researchers cally important mainland fauna (50). stock in a process parallel to, but subscribing to the indigenist perspective, including all four major livestock species independent of, the Near East (31–33). especially those working in the western (sheep, goat, cattle, and pig). Early The presence of wild , barley, and Mediterranean, argued that cultural and colonists also imported mainland game

Zeder PNAS ͉ August 19, 2008 ͉ vol. 105 ͉ no. 33 ͉ 11599 Downloaded by guest on September 27, 2021 animals like fallow deer and fox that, sula ca. 8,000 B.P. by maritime colonists favorable coastal locals around the pe- although perhaps kept in (48), who first established farming villages on riphery of the at a were never truly domesticated. None of the Apulian ‘‘boot heel’’ region of steady and quite rapid pace, appearing these animals are endemic to Cyprus. southeastern Italy (Fig. 2). These tradi- first on the eastern and southern coasts Although imported livestock species did tions appear in northwest coastal Italy of Spain at ca. 7,700–7,600 B.P. and on not show any of the morphological fea- Ϸ200–300 years later (ca. 7,800–7,600 the Atlantic coast of Portugal ca. 7,400– tures traditionally used to mark domes- B.P.). In southern France, a compelling 7,300 B.P. (Fig. 2). tic status when they arrived on the case can be made for a marked geo- However, Zilha˜o’s (61, 62) work in island, demographic profiles of these graphic, ecological, and cultural break Portugal has also shown that Mesolithic animals are consistent with human man- between interior Mesolithic settlements cultures focusing on the intensive exploi- agement. In contrast, demographic pro- and coastal Neolithic colonies (58) Re- tation of estuary resources persist for files of the fallow deer are indicative of cent excavation of a coastal settlement several hundred years after the estab- hunting, suggesting that early colonists in southern France, dating to 7,700– lishment of these farming enclaves and were engaged in game stocking and 7,600 B.P. and characterized as a beach- that the subsequent spread of agricul- herd management (13, 48). Deep head colony of seafaring migrant - tural economies into the interior likely constructed at one of these early sites ers from mainland Italy, has yielded proceeded through a combined process yielded abundant evidence of domesti- , domestic sheep, einkorn, and of colonist expansion, selective local cated einkorn and wheat and emmer wheat (59). adoption of Neolithic technologies, and lentils, none of which are native to Questions have also been raised about the integration of colonist and indige- Cyprus, and domestic barley, which in the evidence for the early occurrence nous populations. Similar patterns of the wild is endemic to the island (26, of domestic animals and pottery in Me- development are hinted at in interior 51). Other introduced plants include solithic contexts in the western Mediter- and northern Italy, which seem to lag pistachios and , as well as figs possi- ranean, which had formed a primary several hundred years behind coastal bly domesticated in the Levant by this foundation of earlier culture diffusion areas in the appearance of plant and time (28). Thus the initial diffusion of models. Based on a reappraisal of the animal domesticates and other markers the nascent Neolithic package out of the complicated cave stratigraphy of Iberian of Neolithic adaptations (56, 57). In Fertile Crescent to Cyprus involved the sites and a reanalysis of their associated southern France, the initial, essentially transplant of all aspects of daily life radiocarbon dates, Zilha˜o (60–62) ar- exclusive, focus on domestic livestock (i.e., subsistence resources, technologies, gues that the pottery and domesticated evidenced at the early coastal pioneer- and, most likely, social networks and caprines recovered from Mesolithic lev- ing sites stands in stark contrast to sub- belief systems) by seafaring colonists els actually derive from higher Neolithic sistence strategies of later interior sites who, for unclear reasons, were seeking a levels. Domestic sheep reported as re- that show persistence of hunting along fresh start in a new land (52). Far from covered from Mesolithic and earlier de- with the utilization of domesticates, a being an isolated event, the colonization posits in southern France can also be pattern that points to the blending of of Cyprus provides a clear and valuable argued to have been derived from over- Neolithic and Mesolithic traditions after template for the subsequent diffusion of lying Neolithic contexts, or, in the case initial colonization (65). Farther east, the Neolithic across the rest of the Med- of higher elevation sites, represent misi- the disjunction between later Neolithic iterranean Basin. dentified native chamois (Rupicapra sites and their Mesolithic and early Neo- Recent archaeological evidence from rupicapra) and European ibex (Capra lithic predecessors in the Aegean signals the Aegean, for example, no longer sup- ibex) (41, 58, 63). Similarly, Rowley- a similar process of dispersal, adoption, ports a model of gradual in-place transi- Conwy’s (64) reexamination of the argu- and integration (54) (Fig. 2). tion of ancestral Mesolithic cultures into ment for the domestic status of pigs and Genetic studies of modern and an- Neolithic cultures (53–55). Instead, cattle in Mesolithic contexts in southern cient DNA from Mediterranean Basin there appears to have been a sharp Spain suggests that the smaller size of livestock species and their progenitors decline in Late Mesolithic population these animals is not a sign of their do- adds further support, and nuance, to levels, combined with the sudden ap- mestication as originally argued, but is this emerging picture. A study of ancient pearance of radically different Neolithic instead a reflection of body size re- mtDNA has shown that two haplotypes settlements in previously unoccupied sponses to different climatic regimes of domestic goats (the A and C lin- locations. As on Cyprus, recent work in among native wild animals. eages) had arrived in southern France the Aegean argues for the arrival of Having discounted evidence for piece- by 7,300 B.P., suggesting their dispersal maritime colonists who, at ca. 9,000 to meal cultural diffusion of various ele- out of the Near East as a single package 8,000 B.P., carried many components of ments of Neolithic economy and their (66). Among modern goat breeds in the full Neolithic package (plant and selective adoption by indigenous Meso- Portugal researchers have found both animal domesticates, new lithic tradi- lithic populations in the western Medi- the ubiquitous A haplotype and the tions, and, perhaps a bit later, pottery) terranean, Zilha˜o (61, 62) has gone on much more restricted haplotype C (67). (Fig. 2). Following a leapfrog pattern, to demonstrate that, as in other parts of Three domestic lineages were found these seafaring pioneers established the Mediterranean Basin, the Late Me- among modern breeds of sheep in Por- farming communities that selectively solithic of the Iberian Peninsula was a tugal, including those previously found focused on favorable environments in period of population decline and reloca- only in the and (68). coastal Greece and on various Aegean tion. Also as elsewhere, Neolithic settle- Both Portuguese sheep and goat show a Islands. ments with apparently fully formed much higher degree of within-breed ge- Based on a careful reevaluation of agro-pastoral economic systems sud- netic diversity than expected at the west- archaeological evidence, especially avail- denly appear in the Iberian Peninsula as ernmost periphery of sheep and goat able radiocarbon dates, researchers now coastal enclaves occupying limestone- expansion. This diversity is attributed to see major discontinuities between Meso- based soils abandoned by earlier Meso- multiple introductions of caprines into lithic and Neolithic cultures in Italy (56, lithic peoples. The initial establishment the Iberian Peninsula, not only through 57). They argue that Neolithic lifeways of these colonies follows a familiar pat- maritime colonization from Italy and were introduced into the Italian penin- tern, with farming enclaves appearing in France, but through subsequent intro-

11600 ͉ www.pnas.org͞cgi͞doi͞10.1073͞pnas.0801317105 Zeder Downloaded by guest on September 27, 2021 Table 1. The of large endemic and human colonization of Mediterranean islands Island/

Tyrrhenian Islands/ Gymnesic Islands/ Megaloceros Crete/ Cyprus/ Time period Myotragus balearicus cazioti Candiacervus sp. Phanourios minutus

Most recent endemic 5,000* 10,000† Ͼ 9,000 11,500*† Earliest human presence 7,000 10,000 None 11,500 Neolithic colonization 4,000 7,600 9,000 10,500

All dates in calibrated years B.P. References are as follows: Gymnesic Islands (69), Tyrrhenian Islands (57), Crete (62), and Cyprus (39, 61). *Directly dated skeletal element. †Radiocarbon-dated stratigraphic context.

ductions out of or overland dispersal of Near Eastern pigs into and populations. Genetic studies of and through . across Europe between 7,500 and 5,000 oats also indicate that the modern vari- Genetic data also support a pattern of B.P. (23). Surprisingly, subsequent to eties of these major crop plants have a multiple introductions of cattle into the this initial diffusion, Near Eastern swine European and not Near Eastern ances- region. The T3 haplotype of domestic are later replaced by domestic pigs of try (24). Future interpretive frameworks cattle, which dominates among modern European maternal ancestry, even will have to take a more integrated ap- and ancient European cattle, seems to within the Near East. Indigenous do- proach, which recognizes colonization, have followed a relatively rapid path of mestication of European boar also ap- diffusion, and independent domestica- expansion around the Mediterranean parently happened several times, with tion as all playing a role in Neolithic Basin without any significant introgres- two major European clades indicative of expansion across the Mediterranean sion with female European aurochsen two separate domestication events, and (65, 74) (Fig. 2). (refs. 21, 69, and 70 and contra ancient an additional clade, currently restricted DNA evidence reported in ref. 71). to Italy and Sardinia, representing an- Environmental Impacts Modern DNA, however, indicates that T other (22, 23). The impact of Neolithic economies on and T2 haplotype cattle were included Thus it appears that none of the ear- the biotic communities of the Mediter- in migratory movements into the Bal- lier models for Neolithic emergence in ranean Basin is most clearly seen on the kans and (71). T1 cattle, the Mediterranean accurately or ade- large islands scattered across the region, which dominate among modern North quately frame the transition. Clearly where highly endemic and disharmonic African taurine cattle, were initially ar- there was a movement of people west- faunas were replaced by a mixture of gued to represent a separate North Afri- ward out of the Near East all of the way domestic and wild mainland fauna (75– can domestication event (21). This lin- to the Atlantic shores of the Iberian 77). Although are clearly the eage now seems more likely to have Peninsula. But this demic expansion did agent of island introductions of main- been brought under domestication with not follow the slow and steady, all- land faunas, their role in the extirpation other T haplogroup cattle in the Near encompassing pace of expansion pre- of endemic island faunas is unclear. East (72) and subsequently radiated dicted by the wave and advance model. Once again, Cyprus reflects a general across through trade and Instead the rate of dispersal varied, with pattern for the Mediterranean Basin. human migration. The patchy occur- Neolithic colonists taking 2,000 years to The endemic mammalian fauna of rence of the TI haplotype among mod- move from Cyprus to the Aegean, an- Cyprus was impoverished and unbal- ern cattle in the Iberian Peninsula, Sicily other 500 to reach Italy, and then only anced, limited to pygmy hippopotamus and central Italy, and the sug- 500–600 years to travel the much (Phanourios minutus), a pygmy elephant gests that T1 cattle entered southern greater distance from Italy to the Atlan- (Elephas cypriotes), a genet (Genetta Europe out of North Africa through tic (52). Moreover, rather than entirely plesictoides), the only carnivore on the multiple points of entry (71). It is also replacing or engulfing indigenous forag- island), and a mouse (Mus cyprinacus, possible that T1 cattle traveled overland ing populations, these colonists seem to the only endemic to survive to the across the Dardanelles into Eastern Eu- have been restricted to scattered coastal present day) (76, 78). None of the larger . The high-diversity T haplogroup farming enclaves established around the endemics are represented among the taurine cattle found among modern Tus- Mediterranean Basin. Although cultural imported mainland fauna associated can cattle has been linked to a post- diffusion can no longer be argued to with the sites of colonists of the 11th Neolithic migration of Etruscans who, provide a universal explanation for Neo- millennium B.P. (50). It is now clear based on both historical evidence and lithic expansion into the Mediterranean, that these pioneer settlers were not the modern human genetic data, are be- it is clear that the movement of Neo- first humans on Cyprus and that main- lieved to have been of Eastern Mediter- lithic lifeways out of these beachhead land hunters made periodic visits to the ranean origin (73). settlements involved selective adoption island 1,000 years earlier during the Pigs a different story. Research and adaptation of elements of the Neo- Younger Dryas climatic downturn (79) by Larson et al. (22) has shown that lithic package by indigenous peoples. (Table 1). Simmons (79) argues that a current-day domestic pigs in Europe Moreover, although caprines, cattle, and large accumulation of pygmy hippopota- bear no trace of Middle Eastern ances- primary crop plants were most certainly mus remains found in a collapsed - try, but instead are most closely related not independently domesticated in Eu- side is directly associated to European wild boar. Subsequent rope, recent genetic data for pigs points with an overlying, but apparently con- analysis by the same team of mtDNA to indigenous domestication of local temporaneous, containing stone extracted from archaeological remains wild boar, possibly occurring multiple and dated to ca. 11,775 has found convincing evidence for the times in geographically separate sub- B.P. Other researchers, although ac-

Zeder PNAS ͉ August 19, 2008 ͉ vol. 105 ͉ no. 33 ͉ 11601 Downloaded by guest on September 27, 2021 cepting the evidence for early visits of land fauna just before, or slightly after, philous small vertebrate species (76) hunter–gatherers to the island, question substantial human colonization was, clearly implicates humans in these island the evidence for human predation on until recently, thought to be a small . the endemic mammals discovered at the derived caprine species (Mytotragus The impact of the Neolithic transition site (refs. 80 and 81, but see ref. 82). balearicus), found only on the Gymnesic on mainland environments throughout Large endemic mammals on Crete islands off the eastern coast of Spain the Mediterranean, although perhaps included pygmy hippopotamus (Hippo- (89). Here initial dating argued for a less dramatic, is no less pronounced. potamus creutzburgi), elephants (Elephas Ͼ3,000-year overlap between initial hu- Blondel and Aronson (92), in fact, argue creutzburgi), and several species of man presence on Mallorca at 8,000 B.P. that the entire Mediterranean Basin is megalocerine deer (Candiacervus sp.) and the last documented Myotragus characterized by highly anthropogenic (76). Dating the last appearance of specimens on the island dated at ca. environments shaped by thousands of these species has proven problematic 5,000 B.P.. A case was even mounted years of human landscape management, (83), although it appears that the Cre- for domestication of this species before species introductions, and associated tian hippopotamuses and elephants be- its extinction (90), which, if true, made responses by indigenous faunas and flo- came extinct before the endemic deer. Myotragus the only domestic animal to ras, all dating to the Neolithic emer- There is as yet no evidence for a tempo- undergo extinction. Recent reevaluation gence. They also note that the various ral overlap between humans and larger of the primary arguments for Myotragus forms of traditional human landscape endemic mammals on Crete. As on domestication has, however, overturned engineering in the Mediterranean have Cyprus, Cretian endemics do not occur this hypothesis (89, 91). A reexamina- created viable, sustainable ecosystems, among the faunal remains recovered tion of the dates for earliest human oc- which have, in fact, been highly benefi- from the earliest known Neolithic settle- cupation of these islands and the most cial to Mediterranean biodiversity. This ment on the island, which dates to ca. recent evidence for Myotragus, more- balanced system is undergoing increas- 9,000 B.P. (84) (Table 1). It is possible over, has all but eliminated any overlap ing threat from urban growth and agri- that the larger endemic mammals of between humans and Myotragus (89). cultural intensification, however, threats Crete became extinct long before Neo- The earliest evidence of substantial hu- that can only be met with a clear under- lithic farmers and herders colonized the man settlement of Mallorca dates to ca. standing of the long-term role of human island, thus limiting the island’s appeal 4,000 B.P., whereas the most recent management in shaping current-day to earlier mainland hunting parties. It is solid evidence for Myotragus on the biodiversity in the Mediterranean Basin. also possible, however, that the ephem- island dates to ca. 5,000 B.P., with Myo- eral camps of these hunters have yet to tragus extinction and initial human colo- Future Research be discovered. nization happening sometime between Recent research on the initial develop- A closer connection between humans these two dates (Table 1). ment and subsequent expansion of domes- and now extinct endemic island faunas has Even lacking definitive evidence for tication and agricultural economies in the been proposed for the islands of Sardinia humans involvement in the extirpation of Mediterranean Basin provides a clear and Corsica. The case for an overlap be- larger endemic mammals on Mediterra- roadmap for future research. This is espe- tween humans and indigenous megalocer- nean islands, the progressive east-to-west cially true for the Fertile Crescent where ine deer (Megaloceros cazioti) on Sardinia disappearance of larger endemics coinci- recent advances are transforming our un- in the (ca. 20,000 B.P.) dent with human settlement of Mediterra- derstanding of the origins of plant and (85) is contested (86). But there is firmer nean islands (Table 1) clearly suggests animal domestication in this key heartland evidence that Mesolithic hunter–gatherers that humans played some role in their region. Traditional approaches to docu- at least visited (if not colonized) Sardinia extinction. Having evolved in the absence menting domestication relied on the by ca. 10,000 B.P. and that they encoun- of major predators, these larger herbivores appearance of genetically driven morpho- tered endemic deer and, possibly, the is- probably lacked protective behaviors, logical change (i.e., the development of land’s sole carnivore, a small fox-size making them especially vulnerable to sus- nonshattering seed heads in cereals and canid (Cynotherium sardous) (refs. 76 and tained human predation. Moreover, all of body size reduction in animals). The de- 86 and Table 1). Neither of these species the species that became extinct around the velopment of new analytical approaches survived much beyond this initial contact, time of initial human settlement likely had has, however, provided a window into the however, and there is no evidence of over- low reproductive rates, a trait commonly preceding processes of human interaction lap between Mesolithic hunter–gatherers found among island endemics, which fur- with target plant and animal species and and these two endemics on Corsica (76). ther limited their ability to withstand any the genetic responses to this interaction A number of sites on both islands attest to degree of hunting pressure (76). Extermi- that eventually resulted in morphological heavy utilization of smaller endemic mam- nation of these island endemics by hu- change. Researchers in the Fertile Cres- mals, especially a -size endemic lago- mans could have happened within a cent are detecting early signs of human morph (Prolagus sardus), which seems to generation. Smaller endemic mammals ecosystem engineering aimed at encourag- be the primary source of animal protein seem to have survived initial human colo- ing plant production (24, 25); they are until Neolithic colonizers brought domes- nization somewhat better, perhaps as a able to document the manipulation of tic livestock to these islands at ca. 7,600 result of their higher reproductive rates herd structure to promote a secure and B.P. (87). All of these smaller endemics, and because they were less attractive prey predictable yield of animal products (7, 8, however, were extripated sometime be- species. However, they, too, eventually 10, 11). In both plants and animals these tween the Late Roman period and the could not withstand the combination of new indicators precede the manifestation early Middle Ages, probably through com- overhunting, loss of habitat, and competi- of traditional morphological markers of bined pressures associated with the intro- tion with invasive mainland imports. The domestication by hundreds, if not thou- duction of Rattus rattus during the Early almost complete turnover of island faunas sands, of years. Estimating exactly when Roman colonization of the island and throughout the region [involving essen- during this extended coevolutionary pro- subsequent episodes of deforestation and tially 100% of mammal species, plus many cess a plant or animal species crossed the agricultural intensification (75, 88). avian and herpetological species (77)] and domestic threshold is now more a seman- The apparent exception to this pat- their replacement with domestic livestock, tic issue than a substantive research ques- tern of rapid extinction of endemic is- game species, and an array of anthropolo- tion (20, 93). Although some researchers

11602 ͉ www.pnas.org͞cgi͞doi͞10.1073͞pnas.0801317105 Zeder Downloaded by guest on September 27, 2021 may require the appearance of specific come into focus. Maritime colonization of transfer rather than truly independent morphological traits before conferring the Mediterranean clearly involved not domestication. Local, culturally indepen- domestic status, others may be more will- one, but multiple unrelated seaborne mi- dent domestication of indigenous wild ing to consider a managed animal or a grations (52). The cultural context of pigs, however, still cannot be ruled out. cultivated plant as having achieved this these migratory movements, their causes, Application of enhanced archaeological status. Regardless of where one chooses their routes, their timing, and their tempo and genetic techniques for detecting and to draw the line between wild and domes- all call out for additional investigation. dating domestication to both extant and tic, recent advances have provided re- The southern margin of the Mediterra- yet-to-be-recovered assemblages is key to searchers with powerful new tools capable nean Basis along coastal North Africa is understanding patterns of indigenous do- of examining the entire process of domes- essentially terra incognita for understand- mestication around the Mediterranean tication, not just its morphological impacts ing the course of Neolithic emergence and Basin. which, if they occur at all, only appear seems an especially promising region for Finally, although we may never be able after the process is well underway. future research (60). to detect the final coup de graˆce for en- Just how far back this process of active The subsequent inland transfer of do- demic island faunas, the future holds con- human resource management goes or how mesticates, agriculture, and associated siderable promise for a much fuller widespread it was in the Fertile Crescent Neolithic lifeways from newly arrived understanding of the human impact on Mediterranean biodiversity. As it has in is, at this point, an open question. The colonists to indigenous populations the Gymnesic islands and on Cyprus, early dispersal of an integrated economy around the Mediterranean Basin is an- ‘‘carpet-dating’’ large numbers of archaeo- based on crop plants and managed ani- other intriguing research area that will logical materials by the small-sample mals to Cyprus at least 2,000 years before benefit from recent advances in our atomic mass spectrometry radiocarbon the apparent crystallization of agricultural ability to detect and date domesticates method will certainly help refine the chro- economies on the mainland, however, sug- in the archaeological record. Careful nology of the disappearance of endemic gests that our understanding of plant and analysis of increasingly more precise ra- island faunas and the arrival of human animal domestication and agricultural diocarbon dates will continue to be criti- colonists. Application of demographic emergence on the mainland is, at best, cal in discriminating between demic profiling techniques to the remains of incomplete. Researchers working in this diffusion and selective adoption of Neo- these animals may make it possible to dis- area are only just beginning to realize the lithic components in different parts of tinguish between natural-death accumula- potential of these newly available analytic the Mediterranean (e.g., ref. 94). New tions and prey assemblages resulting from tools. Additional even more effective ana- demographic techniques for profiling human predation. Similarly, recognition of lytical approaches will almost certainly be prey strategies, morphometric tech- the broader role of humans in shaping developed in the future as researchers niques capable of tracking genetic and post-Neolithic environments is central to embrace this broader concept of domesti- responses to human management, understanding how Mediterranean biodi- cation and begin to exploit the many op- isotopic analysis, and the increasing suc- versity evolved and how we might best portunities available in the Fertile Cres- cess of ancient DNA studies of domesti- work to conserve it. The archaeobiological cent region for documenting it. cates will enhance our understanding of sciences have a valuable role to play in Recent research has also shown that the the ways in which both colonists and providing greater time depth to biodiver- dispersal of domesticates and the Neo- local populations adapted management sity studies by monitoring the creation of lithic way of life west across the Mediter- strategies to these new environments. anthropogenic ecosystems and tracing the ranean Basin was much more complex There are also obvious opportunities development and impacts of both environ- and multifaceted than previous prime for those interested in understanding the mentally sustainable and destructive agri- mover models could accommodate. To independent domestication of European cultural economies over thousands of varying degrees, in different areas, this wild species. Larson et al. (23), for exam- years of human occupation. process involved elements of demic diffu- ple, suggest that European wild boar were sion, local adoption, and independent domesticated only after the introduction ACKNOWLEDGMENTS. This work has benefited from the comments of Greger Larson, Bruce domestication. But the outlines of this of Near Eastern domestic swine, repre- Smith, and Jean-Denis Vigne and from the art- complex process are just beginning to senting a case of apparent istry of Marcia Bakry, who drew the maps.

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