Domestication and early agriculture in the MediterraneanBasin: Origins, diffusion, and impactMelinda A. Zeder*Archaeobiology Program, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013

Edited by Jeremy A. Sabloff, University of Pennsylvania Museum of Archaeology and Anthropology, Philadelphia, PA, and approved May 27, 2008 (receivedfor 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 theMediterranean Basin. In large measure these advances are attributable to new methods for documenting domestication in plants andanimals. The initial steps toward plant and animal domestication in the Eastern Mediterranean can now be pushed back to the 12thmillennium cal B.P. Evidence for herd management and crop cultivation appears at least 1,000 years earlier than the morphologicalchanges traditionally used to document domestication. Different species seem to have been domesticated in different parts of theFertile Crescent, 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 seafaringcolonists who established coastal farming enclaves around the Mediterranean Basin. This process also involved the adoption of do-mesticates and domestic technologies by indigenous populations and the local domestication of some endemic species. Human envi-ronmental impacts are seen in the complete replacement of endemic island faunas by imported mainland fauna and in today’santhropogenic, but threatened, Mediterranean landscapes where sustainable agricultural practices have helped maintain high bio-diversity since the Neolithic.

archaeology � livestock

The transition from foraging andhunting to farming and herdingis a significant threshold in hu-man history. Domesticates and

the agricultural economies based onthem are associated with radical restruc-turing of human societies, worldwidealterations in biodiversity, and signifi-cant changes in the Earth’s landformsand its atmosphere. Given the momen-tous outcomes of this transition it comesas little surprise that the origin andspread of domesticates and the emer-gence of agriculture remain topics ofenduring interest to both the scholarlycommunity and the general public.

The past decade has seen remarkableanalytical advances in documenting do-mestication (1), particularly in trackingthe domestication of four major NearEastern livestock species (sheep, goats,cattle, and pigs) and their subsequentdispersal throughout the MediterraneanBasin. New morphometric methods aretracking changes in human prey strate-gies that mark the transition from hunt-ing to herding. Genetic analyses bringfresh insights into initial livestockdomestication and their dispersal. Small-sample atomic mass spectrometryradiocarbon dating provides refinedchronological frameworks for these de-velopments. These recent analyticaladvances, in turn, have produced an ex-plosion of new information that is call-ing into question prevailing hypothesesabout the origin and early spread of ani-mal domesticates and the Neolithic life-ways of which they were a part. Here, Ibring together these different sources ofinformation to consider the origins, dif-

fusion, and impacts of domesticates andagriculture in the Mediterranean Basin,outlining our current understanding ofthese developments and highlightingpromising areas for future study.

Initial Animal Domestication in theFertile CrescentUntil the late 1990s archaeozoologistsrelied on morphological changes in tar-get species to identify where and whenwild prey animals were transformed intoherded livestock (2). A proposed sharpand rapid reduction in overall body sizeamong archaeological prey populationswas the most widely accepted morpho-logical marker of this threshold (3, 4).Based on this size reduction criterion,the established consensus was that ani-mal domestication (beginning with goatsand then sheep) occurred at ca. 10,000–9,500 B.P.†, �1,000 years after thedomestication of crop plants in thesouthern Levant (3, 5). Domestication ofthese two animal species was thought tohave occurred somewhere to the northand east of the heartland of plant do-mestication (5), although a second, inde-pendent domestication of goats wasproposed for the southern Levant (6).

The utility of this size reductionmarker, and indeed of all morphologicalmarkers, has come under increasingscrutiny (7). My own work on both mod-ern skeletal collections and archaeologi-cal caprine (sheep and goat) remainsfrom the Near East finds little supportfor the almost axiomatic acceptance thatdomestication results in an automaticoverall reduction in body size in man-aged animals (ref. 8 and references

therein). Rather than domestic status,sex is the primary factor affecting bodysize in these ungulates, manifested by amarked and consistent difference be-tween larger males and smaller femalesin essentially all skeletal elements. Envi-ronment also strongly influences bodysize, with increasing heat and ariditypositively correlated with smaller size.What archaeozoologists had originallyinterpreted as body size reduction asso-ciated with initial domestication cannow be attributed to differences in theculling strategies of herders as opposedto hunters. In most prey species, huntersfocus on large adult animals (particu-larly males) to maximize return, and thebones of these larger animals generallydominate in prey assemblages generatedby hunters. Archaeological assemblagesgenerated by herders, on the otherhand, are usually dominated by thebones of smaller females slaughteredafter their prime reproductive years. Ex-cess males not needed for herd propaga-tion were harvested at young ages andtheir more friable bones are usually lesswell represented in these assemblages.

Although the linkage between domes-tication and body size was called into

This article grew out of a presentation given by M.Z. at theCalpe 2007 Symposium: People in the Mediterranean–AHistory of Interaction, September 27–30, 2007, the GibraltarMuseum, Gibraltar.

Author contributions: M.A.Z. designed research, per-formed research, synthesized research in referenced publi-cations, and wrote the paper.

The author declares no conflict of interest.

This article is a PNAS Direct Submission.

*E-mail: zederm@si.edu.

†All dates are reported in calibrated years before present.

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question by this research, the markeddegree of sexual dimorphism in caprinesit documented offered another approachto tracking the transition from huntingto herding. Pronounced differences inthe size of male and female skeletalelements make it possible to separatearchaeological assemblages intosex-specific subpopulations, which, basedon a refined understanding of the se-quence and timing of long bone fusion(9), can be used to generate high-resolution harvest profiles for male andfemale animals capable of distinguishingthe prey strategies of hunters from theharvest strategies of herders.

Application of this approach to ar-chaeological assemblages from Iraq andIran has identified the clear signature ofa managed herd of goats (harvesting ofyoung males and prolonged survivorshipof females) at the site of Ganj Dareh inhighland Iran (8). Directly dated to9,900 B.P., the goats from this site showno evidence of size reduction or anyother domestication-induced morpholog-ical change. Smaller body size andchanges in the size and shape of horns[a morphological change clearly linkedto domestication (2)] appear 500–1,000years later than this demographic shift,when managed animals were movedfrom the natural habitat of wild goatsand introduced into hotter and morearid lowland Iran. These follow-on mor-phological changes likely reflect re-sponses to new selective pressures, plusthe now more limited opportunities forintrogression between managed and wildanimals or the restocking of herds withwild animals.

Looking back before Ganj Dareh, un-usual demographic profiles detected insheep bone assemblages from northeast-ern Iraq (10) and southeastern Anatolia(11) dating to 12,000 B.P. may reflectearly attempts at manipulating herd de-mographics to maximize returns. Theseassemblages show an almost exclusivefocus on 2- to 3-year-old males, which isolder than expected with herd manage-ment but younger than expected withhunting. This pattern is argued to resultfrom a prime male hunting strategy de-veloped under conditions of intensivepressure on local wild herds during theYounger Dryas climatic downturn (11).The proposed scenario suggests that,rather than broadening the prey strategyto include both males and females, hunt-ers conserved female breeding stockwhile at the same time relying on asteady immigration of younger malesdrawn from surrounding territories tofill the vacuum left by the kill-off oflocal prime-age males.

Lower-resolution demographic meth-ods used by archaeozoologists working

elsewhere in the Fertile Crescent aredetecting parallel patterns to those doc-umented in the Zagros. Changes in theage of harvested caprines, and possiblydemographically driven changes in sizeconsistent with early herd management,are found in southeastern Anatolia atca. 10,500 B.P. (12). Sheep seem to bethe initial early focus of the transitionfrom hunting to herding in this region,with managed goats arriving from out-side the area at ca. 10,200 B.P. (12).Similarly, demographic evidence for themanagement of morphologically unal-tered caprines (mostly sheep) is found inCentral Anatolia between 10,400 and9,400 B.P. (13). These results suggestthat both sheep and goats were broughtunder domestication (probably indepen-dently of one another and possibly mul-tiple times) in the region that stretchesfrom the northern Zagros to southeast-ern Anatolia between ca. 11,000 and10,500 B.P., and perhaps even earlier(Fig. 1). Morphologically wild, but man-aged, goats appear to have been movedrelatively rapidly through the region,reaching the southernmost tips of boththe eastern and western arms of theFertile Crescent by ca. 9,500 B.P. Do-mestic sheep were spread more slowlyand first appear in these regions �500–1,000 years later than managed goats(10, 12).

Recent research has also clarified thespatial and temporal context of the do-mestication of two other major livestockspecies in the Near East: pigs and cattle.Archaeological evidence now suggeststhat pigs were first domesticated some-where in southeastern Anatolia by10,500–10,000 B.P. and that the timingof their geographical expansion as do-mesticates was similar, although perhaps

slower, to that of sheep (Fig. 1) (10, 14).Morphologically altered domestic pigsare not found in the southern Levant orlowland Iran until ca. 8,500–8,000 B.P.Recent demographic evidence suggeststhat taurine cattle were initially domesti-cated somewhere in the upper Eu-phrates Valley between ca. 11,000 and10,000 B.P. (15), but, like sheep andpigs, they arrived relatively late in moredistant parts of the Fertile Crescent(Fig. 1). Morphologically altered domes-tic pigs and cattle are not found in Cen-tral Anatolia until after 8,500 B.P. (16).

Genetic data from modern and ar-chaeological specimens both supportand enhance this picture of initial ani-mal domestication. Recent work has suc-ceeded in definitively identifying theprogenitors of both domestic sheep andgoat as belonging to species found inthe Fertile Crescent (Ovis orientalis andCapra aegagrus, respectively) (17, 18).Moreover, in both of these livestock spe-cies there are at least four and, in thecase of goats, as many as six (19), genet-ically distinguishable domestic lineages,or haplotypes. It is not entirely clear,however, whether these different lin-eages represent spatially and temporallydiscrete ‘‘domestication events’’ in whichdifferent populations of animals werebrought under domestication indepen-dently of one another (20). Genetic datafor taurine cattle have identified fivedifferent domestic haplotypes, at leastthree and possibly four of which origi-nated in the Fertile Crescent (21). Simi-larly, as many as four of the manydifferent lineages of domestic pigsoriginated in the Near East (22, 23).

Animal domestication in the NearEast can then be seen as arising from aperiod of prolonged human interaction

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Fig. 1. The origin and dispersal of domestic livestock species in the Fertile Crescent. Shaded areas showthe general region and the approximate dates in calibrated years B.P. in which initial domestication isthought to take place. Dates outside of the shaded areas show the approximate date when the domes-ticate first appears in a region. Orange, goats (Capra hircus); blue, sheep (Ovis aries); green, cattle (Bostaurus); fuscia, pigs (Sus scrofa).

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with the ancestors of core livestock spe-cies that unfolded across much of theFertile Crescent. Over time huntingstrategies aimed at maximizing localavailability of wild ungulates developedinto active management, with all fourmajor livestock species coming undermanagement over a period from ca.11,000 to 10,000 B.P. Even species likegazelle, which are behaviorally unsuitedto domestication, may have been audi-tioned for management in the southernand northern Levant, where they werethe most abundant wild ungulate (11).Clear-cut morphological responses todomestication (i.e., changes in horns inbovids and tooth size in pigs) are notevident in these four livestock speciesuntil ca. 9,500–9,000 B.P.

As is the case with animal domestica-tion in the Near East, the leading edgeof plant domestication in the region isnow recognized as an extended process(24, 25). Evidence from multiple loca-tions point to a prolonged period of hu-man manipulation of morphologicallywild, but possibly cultivated, plantswhich, in certain species, resulted in thedevelopment of morphologically altereddomesticated crops (26–28). This periodof intensified plant management dates atleast as far back as ca. 12,000 B.P., withmorphological markers of crop domesti-cation (i.e., nonshattering seed heads incereals) not well established until ca.10,500 B.P. (24, 25, 29). Agriculturaleconomies reliant on a mix of domesti-cated crops and livestock apparently donot fully crystallize in the region untilca. 9,500–9,000 B.P. (5, 10).

The Diffusion of Animal Domesticates inthe Mediterranean BasinThe last two decades has witnessed therise and fall of a number of models ofNeolithic expansion across the Mediterra-nean Basin. In the early 1980s Ammer-man and Cavalli-Sforza (30) combinedarchaeological and human genetic data toframe their ‘‘wave and advance’’ model.This model attributed the westwardspread of the Neolithic to Near Easterncolonists who, driven by agriculture-fueledpopulation growth, slowly pushed asideindigenous hunter–gatherers at a pre-dicted average pace of �1 km per year.

Objecting to the passive role thismodel assigned to indigenous Mesolithicpeople, a number of researchers subse-quently countered with alternative mod-els that awarded local populations astarring role in Mediterranean Neolithicemergence. Early models within this in-digenist perspective argued for autocho-nous domestication of crops and live-stock in a process parallel to, butindependent of, the Near East (31–33).The presence of wild oats, barley, and

lentils in Upper Paleolithic and Meso-lithic levels at Francthi Cave on theeastern coast of Greece, followed by theappearance of fully domesticated barleyand lentils in later Neolithic levels, wasinterpreted as evidence for the localcrop domestication (34). Legumes recov-ered from Mesolithic cave deposits insouthern France were seen as evidenceof incipient cultivation, if not domestica-tion, of local wild plants (35). Evidencefor indigenous animal domestication wasbased on the identification of wild sheepin Pleistocene age deposits in southernFrance and the presence of domesticsheep and goat remains in Mesolithiccontexts in France and Spain (36, 37).Reports of domesticated pig and cattleremains in Mesolithic (pre-8,000 B.P.)levels from sites in southern Spain (38)were also cited as evidence for the localdomestication of these species.

Genetic studies have subsequentlyruled out European ancestry for domes-tic wheat, barley, and pulses, confirmingthe Near East as the source of thesecrops (26, 39). Morphological, cytologi-cal, hemoglobin, and, most recently, ge-netic studies have shown that the ‘‘wild’’sheep and goats found on Mediterra-nean islands, once argued to be thedescendents of the progenitors of indig-enous domestic caprines, are instead theferal descendents of Near Eastern ca-prines (for a review in chronologicalorder, see refs. 40, 41, 17, and 18).

Ruling out the indigenous domestica-tion of caprines and major crop plantsdid not, however, lead to an embrace ofcolonist expansion diffusion models forthe emergence of Neolithic lifeways inthe Mediterranean. Instead, researcherssubscribing to the indigenist perspective,especially those working in the westernMediterranean, argued that cultural and

not demic diffusion was the primary en-gine driving this transition. Specifically,proponents maintained that the selectiveadoption of various elements of theNeolithic package by indigenous popula-tions around the Mediterranean couldhave happened through trade and tech-nology transfer alone without any directcontact between indigenous hunter–gatherers and colonizing farming popu-lations from the east (41–46).

The shortcomings of these differentsingle-agent models have becomeincreasingly apparent as new archaeo-logical data come to light and oldercollections are reanalyzed by using newmethods and new perspectives. Recentgenetic analyses of livestock species andtheir progenitors have also contributedimportant new insights into this process.As a result, a much more complex, andmore interesting, scenario is emergingfor the Neolithic transition across theMediterranean Basin.

Beginning in the early 1990s a num-ber of sites have been discovered andexcavated on Cyprus that have radicallytransformed our understanding of Neo-lithic emergence in the MediterraneanBasin (47). Until the early 1990s Cypruswas thought to have been colonized ca.8,500 B.P. by a derived offshoot of fullyestablished Neolithic mainland cultures(48). The new sites, however, date 2,000years earlier (10,500–9,000 B.P.) anddocument the arrival of early pioneershypothesized to have originated some-where in the Northern Levant (Figs. 1and 2) (47, 49). Traveling the 60 k toCyprus by boat, these colonists trans-ported the full complement of economi-cally important mainland fauna (50).including all four major livestock species(sheep, goat, cattle, and pig). Earlycolonists also imported mainland game

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Fig. 2. An integrated model of the Neolithic expansion in the Mediterranean Basin. The location ofcolonist farming enclaves is shown in the red ellipses. Approximate dates of these enclaves are given insidethe ellipses in calibrated years B.P. Red dots represent areas that are proposed to have been settled bycolonist farmers; green dots indicate areas where indigenous foragers adopted elements of the Neolithicpackage; and blue dots indicate areas of proposed integration of colonist farmers with indigenousforaging groups. Data were complied from refs. 52, 54, 56, 57, and 65 and figure 7.1 of ref. 74.

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animals like fallow deer and fox that,although perhaps kept in captivity (48),were never truly domesticated. None ofthese animals are endemic to Cyprus.Although imported livestock species didnot show any of the morphological fea-tures traditionally used to mark domes-tic status when they arrived on theisland, demographic profiles of theseanimals are consistent with human man-agement. In contrast, demographic pro-files of the fallow deer are indicative ofhunting, suggesting that early colonistswere engaged in game stocking andherd management (13, 48). Deep wellsconstructed at one of these early sitesyielded abundant evidence of domesti-cated einkorn and emmer wheat andlentils, none of which are native toCyprus, and domestic barley, which inthe wild is endemic to the island (26,51). Other introduced plants includepistachios and flax, as well as figs possi-bly domesticated in the Levant by thistime (28). Thus the initial diffusion ofthe nascent Neolithic package out of theFertile Crescent to Cyprus involved thetransplant of all aspects of daily life(i.e., subsistence resources, technologies,and, most likely, social networks andbelief systems) by seafaring colonistswho, for unclear reasons, were seeking afresh start in a new land (52). Far frombeing an isolated event, the colonizationof Cyprus provides a clear and valuabletemplate for the subsequent diffusion ofthe Neolithic across the rest of the Med-iterranean Basin.

Recent archaeological evidence fromthe Aegean, for example, no longer sup-ports a model of gradual in-place transi-tion of ancestral Mesolithic cultures intoNeolithic cultures (53–55). Instead,there appears to have been a sharpdecline in Late Mesolithic populationlevels, combined with the sudden ap-pearance of radically different Neolithicsettlements in previously unoccupiedlocations. As on Cyprus, recent work inthe Aegean argues for the arrival ofmaritime colonists who, at ca. 9,000 to8,000 B.P., carried many components ofthe full Neolithic package (plant andanimal domesticates, new lithic tradi-tions, and, perhaps a bit later, pottery)(Fig. 2). Following a leapfrog pattern,these seafaring pioneers establishedfarming communities that selectivelyfocused on favorable environments incoastal Greece and on various AegeanIslands.

Based on a careful reevaluation ofarchaeological evidence, especially avail-able radiocarbon dates, researchers nowsee major discontinuities between Meso-lithic and Neolithic cultures in Italy (56,57). They argue that Neolithic lifewayswere introduced into the Italian penin-

sula ca. 8,000 B.P. by maritime colonistswho first established farming villages onthe Apulian ‘‘boot heel’’ region ofsoutheastern Italy (Fig. 2). These tradi-tions appear in northwest coastal Italy�200–300 years later (ca. 7,800–7,600B.P.). In southern France, a compellingcase can be made for a marked geo-graphic, ecological, and cultural breakbetween interior Mesolithic settlementsand coastal Neolithic colonies (58) Re-cent excavation of a coastal settlementin southern France, dating to 7,700–7,600 B.P. and characterized as a beach-head colony of seafaring migrant farm-ers from mainland Italy, has yieldedpottery, domestic sheep, einkorn, andemmer wheat (59).

Questions have also been raised aboutthe evidence for the early occurrenceof domestic animals and pottery in Me-solithic contexts in the western Mediter-ranean, which had formed a primaryfoundation of earlier culture diffusionmodels. Based on a reappraisal of thecomplicated cave stratigraphy of Iberiansites and a reanalysis of their associatedradiocarbon dates, Zilhao (60–62) ar-gues that the pottery and domesticatedcaprines recovered from Mesolithic lev-els actually derive from higher Neolithiclevels. Domestic sheep reported as re-covered from Mesolithic and earlier de-posits in southern France can also beargued to have been derived from over-lying Neolithic contexts, or, in the caseof higher elevation sites, represent misi-dentified native chamois (Rupicaprarupicapra) and European ibex (Capraibex) (41, 58, 63). Similarly, Rowley-Conwy’s (64) reexamination of the argu-ment for the domestic status of pigs andcattle in Mesolithic contexts in southernSpain suggests that the smaller size ofthese animals is not a sign of their do-mestication as originally argued, but isinstead a reflection of body size re-sponses to different climatic regimesamong native wild animals.

Having discounted evidence for piece-meal cultural diffusion of various ele-ments of Neolithic economy and theirselective adoption by indigenous Meso-lithic populations in the western Medi-terranean, Zilhao (61, 62) has gone onto demonstrate that, as in other parts ofthe Mediterranean Basin, the Late Me-solithic of the Iberian Peninsula was aperiod of population decline and reloca-tion. Also as elsewhere, Neolithic settle-ments with apparently fully formedagro-pastoral economic systems sud-denly appear in the Iberian Peninsula ascoastal enclaves occupying limestone-based soils abandoned by earlier Meso-lithic peoples. The initial establishmentof these colonies follows a familiar pat-tern, with farming enclaves appearing in

favorable coastal locals around the pe-riphery of the Iberian Peninsula at asteady and quite rapid pace, appearingfirst on the eastern and southern coastsof Spain at ca. 7,700–7,600 B.P. and onthe Atlantic coast of Portugal ca. 7,400–7,300 B.P. (Fig. 2).

However, Zilhao’s (61, 62) work inPortugal has also shown that Mesolithiccultures focusing on the intensive exploi-tation of estuary resources persist forseveral hundred years after the estab-lishment of these farming enclaves andthat the subsequent spread of agricul-tural economies into the interior likelyproceeded through a combined processof colonist expansion, selective localadoption of Neolithic technologies, andthe integration of colonist and indige-nous populations. Similar patterns ofdevelopment are hinted at in interiorand northern Italy, which seem to lagseveral hundred years behind coastalareas in the appearance of plant andanimal domesticates and other markersof Neolithic adaptations (56, 57). Insouthern France, the initial, essentiallyexclusive, focus on domestic livestockevidenced at the early coastal pioneer-ing sites stands in stark contrast to sub-sistence strategies of later interior sitesthat show persistence of hunting alongwith the utilization of domesticates, apattern that points to the blending ofNeolithic and Mesolithic traditions afterinitial colonization (65). Farther east,the disjunction between later Neolithicsites and their Mesolithic and early Neo-lithic predecessors in the Aegean signalsa similar process of dispersal, adoption,and integration (54) (Fig. 2).

Genetic studies of modern and an-cient DNA from Mediterranean Basinlivestock species and their progenitorsadds further support, and nuance, tothis emerging picture. A study of ancientmtDNA has shown that two haplotypesof domestic goats (the A and C lin-eages) had arrived in southern Franceby 7,300 B.P., suggesting their dispersalout of the Near East as a single package(66). Among modern goat breeds inPortugal researchers have found boththe ubiquitous A haplotype and themuch more restricted haplotype C (67).Three domestic lineages were foundamong modern breeds of sheep in Por-tugal, including those previously foundonly in the Middle East and Asia (68).Both Portuguese sheep and goat show amuch higher degree of within-breed ge-netic diversity than expected at the west-ernmost periphery of sheep and goatexpansion. This diversity is attributed tomultiple introductions of caprines intothe Iberian Peninsula, not only throughmaritime colonization from Italy andFrance, but through subsequent intro-

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ductions out of Africa or overlandthrough Europe.

Genetic data also support a pattern ofmultiple introductions of cattle into theregion. The T3 haplotype of domesticcattle, which dominates among modernand ancient European cattle, seems tohave followed a relatively rapid path ofexpansion around the MediterraneanBasin without any significant introgres-sion with female European aurochsen(refs. 21, 69, and 70 and contra ancientDNA evidence reported in ref. 71).Modern DNA, however, indicates that Tand T2 haplotype cattle were includedin migratory movements into the Bal-kans and Central Europe (71). T1 cattle,which dominate among modern NorthAfrican taurine cattle, were initially ar-gued to represent a separate North Afri-can domestication event (21). This lin-eage now seems more likely to havebeen brought under domestication withother T haplogroup cattle in the NearEast (72) and subsequently radiatedacross North Africa through trade andhuman migration. The patchy occur-rence of the TI haplotype among mod-ern cattle in the Iberian Peninsula, Sicilyand central Italy, and the Balkans sug-gests that T1 cattle entered southernEurope out of North Africa throughmultiple points of entry (71). It is alsopossible that T1 cattle traveled overlandacross the Dardanelles into Eastern Eu-rope. The high-diversity T haplogrouptaurine cattle found among modern Tus-can cattle has been linked to a post-Neolithic migration of Etruscans who,based on both historical evidence andmodern human genetic data, are be-lieved to have been of Eastern Mediter-ranean origin (73).

Pigs tell a different story. Researchby Larson et al. (22) has shown thatcurrent-day domestic pigs in Europebear no trace of Middle Eastern ances-try, but instead are most closely relatedto European wild boar. Subsequentanalysis by the same team of mtDNAextracted from archaeological remainshas found convincing evidence for the

dispersal of Near Eastern pigs into andacross Europe between 7,500 and 5,000B.P. (23). Surprisingly, subsequent tothis initial diffusion, Near Eastern swineare later replaced by domestic pigs ofEuropean maternal ancestry, evenwithin the Near East. Indigenous do-mestication of European boar also ap-parently happened several times, withtwo major European clades indicative oftwo separate domestication events, andan additional clade, currently restrictedto Italy and Sardinia, representing an-other (22, 23).

Thus it appears that none of the ear-lier models for Neolithic emergence inthe Mediterranean accurately or ade-quately frame the transition. Clearlythere was a movement of people west-ward out of the Near East all of the wayto the Atlantic shores of the IberianPeninsula. But this demic expansion didnot follow the slow and steady, all-encompassing pace of expansion pre-dicted by the wave and advance model.Instead the rate of dispersal varied, withNeolithic colonists taking 2,000 years tomove from Cyprus to the Aegean, an-other 500 to reach Italy, and then only500–600 years to travel the muchgreater distance from Italy to the Atlan-tic (52). Moreover, rather than entirelyreplacing or engulfing indigenous forag-ing populations, these colonists seem tohave been restricted to scattered coastalfarming enclaves established around theMediterranean Basin. Although culturaldiffusion can no longer be argued toprovide a universal explanation for Neo-lithic expansion into the Mediterranean,it is clear that the movement of Neo-lithic lifeways out of these beachheadsettlements involved selective adoptionand adaptation of elements of the Neo-lithic package by indigenous peoples.Moreover, although caprines, cattle, andprimary crop plants were most certainlynot independently domesticated in Eu-rope, recent genetic data for pigs pointsto indigenous domestication of localwild boar, possibly occurring multipletimes in geographically separate sub-

populations. Genetic studies of rye andoats also indicate that the modern vari-eties of these major crop plants have aEuropean and not Near Eastern ances-try (24). Future interpretive frameworkswill have to take a more integrated ap-proach, which recognizes colonization,diffusion, and independent domestica-tion as all playing a role in Neolithicexpansion across the Mediterranean(65, 74) (Fig. 2).

Environmental ImpactsThe impact of Neolithic economies onthe biotic communities of the Mediter-ranean Basin is most clearly seen on thelarge islands scattered across the region,where highly endemic and disharmonicfaunas were replaced by a mixture ofdomestic and wild mainland fauna (75–77). Although humans are clearly theagent of island introductions of main-land faunas, their role in the extirpationof endemic island faunas is unclear.

Once again, Cyprus reflects a generalpattern for the Mediterranean Basin.The endemic mammalian fauna ofCyprus was impoverished and unbal-anced, limited to pygmy hippopotamus(Phanourios minutus), a pygmy elephant(Elephas cypriotes), a genet (Genettaplesictoides), the only carnivore on theisland), and a mouse (Mus cyprinacus,the only endemic to survive to thepresent day) (76, 78). None of the largerendemics are represented among theimported mainland fauna associatedwith the sites of colonists of the 11thmillennium B.P. (50). It is now clearthat these pioneer settlers were not thefirst humans on Cyprus and that main-land hunters made periodic visits to theisland 1,000 years earlier during theYounger Dryas climatic downturn (79)(Table 1). Simmons (79) argues that alarge accumulation of pygmy hippopota-mus remains found in a collapsed sea-side rock shelter is directly associatedwith an overlying, but apparently con-temporaneous, stratum containing stonetools and hearths dated to ca. 11,775B.P. Other researchers, although ac-

Table 1. The extinction of Late Pleistocene large endemic mammals and human colonization ofMediterranean islands

Time period

Island/mammal

Gymnesic Islands/Myotragus balearicus

Tyrrhenian Islands/Megaloceros

caziotiCrete/

Candiacervus sp.Cyprus/

Phanourios minutus

Most recent endemic 5,000* 10,000† � 9,000 11,500*†

Earliest human presence 7,000 10,000 None 11,500Neolithic 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.

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cepting the evidence for early visits ofhunter–gatherers to the island, questionthe evidence for human predation onthe endemic mammals discovered at thesite (refs. 80 and 81, but see ref. 82).

Large endemic mammals on Creteincluded pygmy hippopotamus (Hippo-potamus creutzburgi), elephants (Elephascreutzburgi), and several species ofmegalocerine deer (Candiacervus sp.)(76). Dating the last appearance ofthese species has proven problematic(83), although it appears that the Cre-tian hippopotamuses and elephants be-came extinct before the endemic deer.There is as yet no evidence for a tempo-ral overlap between humans and largerendemic mammals on Crete. As onCyprus, Cretian endemics do not occuramong the faunal remains recoveredfrom the earliest known Neolithic settle-ment on the island, which dates to ca.9,000 B.P. (84) (Table 1). It is possiblethat the larger endemic mammals ofCrete became extinct long before Neo-lithic farmers and herders colonized theisland, thus limiting the island’s appealto earlier mainland hunting parties. It isalso possible, however, that the ephem-eral camps of these hunters have yet tobe discovered.

A closer connection between humansand now extinct endemic island faunas hasbeen proposed for the islands of Sardiniaand Corsica. The case for an overlap be-tween humans and indigenous megalocer-ine deer (Megaloceros cazioti) on Sardiniain the Upper Paleolithic (ca. 20,000 B.P.)(85) is contested (86). But there is firmerevidence that Mesolithic hunter–gatherersat least visited (if not colonized) Sardiniaby ca. 10,000 B.P. and that they encoun-tered endemic deer and, possibly, the is-land’s sole carnivore, a small fox-sizecanid (Cynotherium sardous) (refs. 76 and86 and Table 1). Neither of these speciessurvived much beyond this initial contact,however, and there is no evidence of over-lap between Mesolithic hunter–gatherersand these two endemics on Corsica (76).A number of sites on both islands attest toheavy utilization of smaller endemic mam-mals, especially a rat-size endemic lago-morph (Prolagus sardus), which seems tobe the primary source of animal proteinuntil Neolithic colonizers brought domes-tic livestock to these islands at ca. 7,600B.P. (87). All of these smaller endemics,however, were extripated sometime be-tween the Late Roman period and theearly Middle Ages, probably through com-bined pressures associated with the intro-duction of Rattus rattus during the EarlyRoman colonization of the island andsubsequent episodes of deforestation andagricultural intensification (75, 88).

The apparent exception to this pat-tern of rapid extinction of endemic is-

land fauna just before, or slightly after,substantial human colonization was,until recently, thought to be a smallderived caprine species (Mytotragusbalearicus), found only on the Gymnesicislands off the eastern coast of Spain(89). Here initial dating argued for a�3,000-year overlap between initial hu-man presence on Mallorca at 8,000 B.P.and the last documented Myotragusspecimens on the island dated at ca.5,000 B.P.. A case was even mountedfor domestication of this species beforeits extinction (90), which, if true, madeMyotragus the only domestic animal toundergo extinction. Recent reevaluationof the primary arguments for Myotragusdomestication has, however, overturnedthis hypothesis (89, 91). A reexamina-tion of the dates for earliest human oc-cupation of these islands and the mostrecent evidence for Myotragus, more-over, has all but eliminated any overlapbetween humans and Myotragus (89).The earliest evidence of substantial hu-man settlement of Mallorca dates to ca.4,000 B.P., whereas the most recentsolid evidence for Myotragus on theisland dates to ca. 5,000 B.P., with Myo-tragus extinction and initial human colo-nization happening sometime betweenthese two dates (Table 1).

Even lacking definitive evidence forhumans involvement in the extirpation oflarger endemic mammals on Mediterra-nean islands, the progressive east-to-westdisappearance of larger endemics coinci-dent with human settlement of Mediterra-nean islands (Table 1) clearly suggeststhat humans played some role in theirextinction. Having evolved in the absenceof major predators, these larger herbivoresprobably lacked protective behaviors,making them especially vulnerable to sus-tained human predation. Moreover, all ofthe species that became extinct around thetime of initial human settlement likely hadlow reproductive rates, a trait commonlyfound among island endemics, which fur-ther limited their ability to withstand anydegree of hunting pressure (76). Extermi-nation of these island endemics by hu-mans could have happened within ageneration. Smaller endemic mammalsseem to have survived initial human colo-nization somewhat better, perhaps as aresult of their higher reproductive ratesand because they were less attractive preyspecies. However, they, too, eventuallycould not withstand the combination ofoverhunting, loss of habitat, and competi-tion with invasive mainland imports. Thealmost complete turnover of island faunasthroughout the region [involving essen-tially 100% of mammal species, plus manyavian and herpetological species (77)] andtheir replacement with domestic livestock,game species, and an array of anthropolo-

philous small vertebrate species (76)clearly implicates humans in these islandextinctions.

The impact of the Neolithic transitionon mainland environments throughoutthe Mediterranean, although perhapsless dramatic, is no less pronounced.Blondel and Aronson (92), in fact, arguethat the entire Mediterranean Basin ischaracterized by highly anthropogenicenvironments shaped by thousands ofyears of human landscape management,species introductions, and associatedresponses by indigenous faunas and flo-ras, all dating to the Neolithic emer-gence. They also note that the variousforms of traditional human landscapeengineering in the Mediterranean havecreated viable, sustainable ecosystems,which have, in fact, been highly benefi-cial to Mediterranean biodiversity. Thisbalanced system is undergoing increas-ing threat from urban growth and agri-cultural intensification, however, threatsthat can only be met with a clear under-standing of the long-term role of humanmanagement in shaping current-daybiodiversity in the Mediterranean Basin.

Future ResearchRecent research on the initial develop-ment and subsequent expansion of domes-tication and agricultural economies in theMediterranean Basin provides a clearroadmap for future research. This is espe-cially true for the Fertile Crescent whererecent advances are transforming our un-derstanding of the origins of plant andanimal domestication in this key heartlandregion. Traditional approaches to docu-menting domestication relied on theappearance of genetically driven morpho-logical change (i.e., the development ofnonshattering seed heads in cereals andbody size reduction in animals). The de-velopment of new analytical approacheshas, however, provided a window into thepreceding processes of human interactionwith target plant and animal species andthe genetic responses to this interactionthat eventually resulted in morphologicalchange. Researchers in the Fertile Cres-cent are detecting early signs of humanecosystem engineering aimed at encourag-ing plant production (24, 25); they areable to document the manipulation ofherd structure to promote a secure andpredictable yield of animal products (7, 8,10, 11). In both plants and animals thesenew indicators precede the manifestationof traditional morphological markers ofdomestication by hundreds, if not thou-sands, of years. Estimating exactly whenduring this extended coevolutionary pro-cess a plant or animal species crossed thedomestic threshold is now more a seman-tic issue than a substantive research ques-tion (20, 93). Although some researchers

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may require the appearance of specificmorphological traits before conferringdomestic status, others may be more will-ing to consider a managed animal or acultivated plant as having achieved thisstatus. Regardless of where one choosesto draw the line between wild and domes-tic, recent advances have provided re-searchers with powerful new tools capableof examining the entire process of domes-tication, not just its morphological impactswhich, if they occur at all, only appearafter the process is well underway.

Just how far back this process of activehuman resource management goes or howwidespread it was in the Fertile Crescentis, at this point, an open question. Theearly dispersal of an integrated economybased on crop plants and managed ani-mals to Cyprus at least 2,000 years beforethe apparent crystallization of agriculturaleconomies on the mainland, however, sug-gests that our understanding of plant andanimal domestication and agriculturalemergence on the mainland is, at best,incomplete. Researchers working in thisarea are only just beginning to realize thepotential of these newly available analytictools. Additional even more effective ana-lytical approaches will almost certainly bedeveloped in the future as researchersembrace this broader concept of domesti-cation and begin to exploit the many op-portunities available in the Fertile Cres-cent region for documenting it.

Recent research has also shown that thedispersal of domesticates and the Neo-lithic way of life west across the Mediter-ranean Basin was much more complexand multifaceted than previous primemover models could accommodate. Tovarying degrees, in different areas, thisprocess involved elements of demic diffu-sion, local adoption, and independentdomestication. But the outlines of thiscomplex process are just beginning to

come into focus. Maritime colonization ofthe Mediterranean clearly involved notone, but multiple unrelated seaborne mi-grations (52). The cultural context ofthese migratory movements, their causes,their routes, their timing, and their tempoall call out for additional investigation.The southern margin of the Mediterra-nean Basis along coastal North Africa isessentially terra incognita for understand-ing the course of Neolithic emergence andseems an especially promising region forfuture research (60).

The subsequent inland transfer of do-mesticates, agriculture, and associatedNeolithic lifeways from newly arrivedcolonists to indigenous populationsaround the Mediterranean Basin is an-other intriguing research area that willbenefit from recent advances in ourability to detect and date domesticatesin the archaeological record. Carefulanalysis of increasingly more precise ra-diocarbon dates will continue to be criti-cal in discriminating between demicdiffusion and selective adoption of Neo-lithic components in different parts ofthe Mediterranean (e.g., ref. 94). Newdemographic techniques for profilingprey strategies, morphometric tech-niques capable of tracking genetic andplastic responses to human management,isotopic analysis, and the increasing suc-cess of ancient DNA studies of domesti-cates will enhance our understanding ofthe ways in which both colonists andlocal populations adapted managementstrategies to these new environments.

There are also obvious opportunitiesfor those interested in understanding theindependent domestication of Europeanwild species. Larson et al. (23), for exam-ple, suggest that European wild boar weredomesticated only after the introductionof Near Eastern domestic swine, repre-senting a case of apparent technology

transfer rather than truly independentdomestication. Local, culturally indepen-dent domestication of indigenous wildpigs, however, still cannot be ruled out.Application of enhanced archaeologicaland genetic techniques for detecting anddating domestication to both extant andyet-to-be-recovered assemblages is key tounderstanding patterns of indigenous do-mestication around the MediterraneanBasin.

Finally, although we may never be ableto detect the final coup de grace for en-demic island faunas, the future holds con-siderable promise for a much fullerunderstanding of the human impact onMediterranean biodiversity. As it has inthe Gymnesic islands and on Cyprus,‘‘carpet-dating’’ large numbers of archaeo-logical materials by the small-sampleatomic mass spectrometry radiocarbonmethod will certainly help refine the chro-nology of the disappearance of endemicisland faunas and the arrival of humancolonists. Application of demographicprofiling techniques to the remains ofthese animals may make it possible to dis-tinguish between natural-death accumula-tions and prey assemblages resulting fromhuman predation. Similarly, recognition ofthe broader role of humans in shapingpost-Neolithic environments is central tounderstanding how Mediterranean biodi-versity evolved and how we might bestwork to conserve it. The archaeobiologicalsciences have a valuable role to play inproviding greater time depth to biodiver-sity studies by monitoring the creation ofanthropogenic ecosystems and tracing thedevelopment and impacts of both environ-mentally sustainable and destructive agri-cultural economies over thousands ofyears of human occupation.

ACKNOWLEDGMENTS. This work has benefitedfrom the comments of Greger Larson, BruceSmith, and Jean-Denis Vigne and from the art-istry of Marcia Bakry, who drew the maps.

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