Your Federal Quarterly Tax Payments are due April 15th Get Help Now >>

Domestication and early agriculture in the Mediterranean Basin by yaosaigeng


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

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 domestication in plants and
animals. The initial steps toward plant and animal domestication in the Eastern Mediterranean can now be pushed back to the 12th
millennium cal B.P. Evidence for herd management and crop cultivation appears at least 1,000 years earlier than the morphological
changes traditionally used to document domestication. Different species seem to have been domesticated in different parts of the
Fertile 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 seafaring
colonists 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’s
anthropogenic, but threatened, Mediterranean landscapes where sustainable agricultural practices have helped maintain high bio-
diversity since the Neolithic.

archaeology   livestock

           he transition from foraging and     these developments and highlighting           ronment also strongly influences body
           hunting to farming and herding      promising areas for future study.             size, with increasing heat and aridity
           is a significant threshold in hu-                                                 positively correlated with smaller size.
           man history. Domesticates and       Initial Animal Domestication in the           What archaeozoologists had originally
the agricultural economies based on            Fertile Crescent                              interpreted as body size reduction asso-
them are associated with radical restruc-      Until the late 1990s archaeozoologists        ciated with initial domestication can
turing of human societies, worldwide           relied on morphological changes in tar-       now be attributed to differences in the
alterations in biodiversity, and signifi-      get species to identify where and when        culling strategies of herders as opposed
cant changes in the Earth’s landforms          wild prey animals were transformed into       to hunters. In most prey species, hunters
and its atmosphere. Given the momen-           herded livestock (2). A proposed sharp        focus on large adult animals (particu-
tous outcomes of this transition it comes      and rapid reduction in overall body size      larly males) to maximize return, and the
as little surprise that the origin and         among archaeological prey populations         bones of these larger animals generally
spread of domesticates and the emer-           was the most widely accepted morpho-          dominate in prey assemblages generated
gence of agriculture remain topics of          logical marker of this threshold (3, 4).      by hunters. Archaeological assemblages
enduring interest to both the scholarly        Based on this size reduction criterion,       generated by herders, on the other
community and the general public.              the established consensus was that ani-       hand, are usually dominated by the
   The past decade has seen remarkable         mal domestication (beginning with goats       bones of smaller females slaughtered
analytical advances in documenting do-         and then sheep) occurred at ca. 10,000–       after their prime reproductive years. Ex-
mestication (1), particularly in tracking      9,500 B.P.†, 1,000 years after the            cess males not needed for herd propaga-
the domestication of four major Near           domestication of crop plants in the           tion were harvested at young ages and
Eastern livestock species (sheep, goats,       southern Levant (3, 5). Domestication of      their more friable bones are usually less
cattle, and pigs) and their subsequent         these two animal species was thought to       well represented in these assemblages.
dispersal throughout the Mediterranean         have occurred somewhere to the north             Although the linkage between domes-
Basin. New morphometric methods are            and east of the heartland of plant do-        tication and body size was called into
tracking changes in human prey strate-         mestication (5), although a second, inde-     question by this research, the marked
gies that mark the transition from hunt-       pendent domestication of goats was            degree of sexual dimorphism in caprines
ing to herding. Genetic analyses bring         proposed for the southern Levant (6).         it documented offered another approach
fresh insights into initial livestock             The utility of this size reduction         to tracking the transition from hunting
domestication and their dispersal. Small-      marker, and indeed of all morphological       to herding. Pronounced differences in
sample atomic mass spectrometry                markers, has come under increasing            the size of male and female skeletal
radiocarbon dating provides refined            scrutiny (7). My own work on both mod-        elements make it possible to separate
chronological frameworks for these de-         ern skeletal collections and archaeologi-     archaeological assemblages into
velopments. These recent analytical            cal caprine (sheep and goat) remains          sex-specific subpopulations, which, based
advances, in turn, have produced an ex-        from the Near East finds little support
plosion of new information that is call-       for the almost axiomatic acceptance that
                                                                                             This article grew out of a presentation given by M.Z. at the
ing into question prevailing hypotheses        domestication results in an automatic
                                                                                             Calpe 2007 Symposium: People in the Mediterranean–A
about the origin and early spread of ani-      overall reduction in body size in man-        History of Interaction, September 27–30, 2007, the Gibraltar
mal domesticates and the Neolithic life-       aged animals (ref. 8 and references           Museum, Gibraltar.
ways of which they were a part. Here, I        therein). Rather than domestic status,        Author contributions: M.A.Z. designed research, per-
bring together these different sources of      sex is the primary factor affecting body      formed research, synthesized research in referenced publi-
information to consider the origins, dif-      size in these ungulates, manifested by a      cations and wrote the paper.

fusion, and impacts of domesticates and        marked and consistent difference be-          The author declares no conflict of interest.
agriculture in the Mediterranean Basin,        tween larger males and smaller females        *E-mail:
outlining our current understanding of         in essentially all skeletal elements. Envi-   †All   dates are reported in calibrated years before present. cgi doi 10.1073 pnas.0801317105                                     PNAS    August 19, 2008          vol. 105      no. 33      11597–11604
on 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 and
female animals capable of distinguishing                                             10,000
                                                                                                 SHEEP         10,500
the prey strategies of hunters from the                                            10,000                    CATTLE          GOATS
harvest strategies of herders.                                                       8,500
                                                                                                           10,000              11,000
   Application of this approach to ar-                                                                    9,600           9,000
chaeological assemblages from Iraq and
                                                                               10,500 10,500
Iran has identified the clear signature of                                              10,500
a managed herd of goats (harvesting of                                            10,500                                                  10,000
young males and prolonged survivorship                                                            9,600
of females) at the site of Ganj Dareh in                                                         9,000                                8,500
highland Iran (8). Directly dated to                                                             8,500                                 8,500

9,900 B.P., the goats from this site show                                                            8,500
no evidence of size reduction or any
other domestication-induced morpholog-
ical change. Smaller body size and               Fig. 1. The origin and dispersal of domestic livestock species in the Fertile Crescent. Shaded areas show
changes in the size and shape of horns           the general region and the approximate dates in calibrated years B.P. in which initial domestication is
[a morphological change clearly linked           thought to take place. Dates outside of the shaded areas show the approximate date when the domes-
to domestication (2)] appear 500–1,000           ticate first appears in a region. Orange, goats (Capra hircus); blue, sheep (Ovis aries); green, cattle (Bos
years later than this demographic shift,         taurus); fuscia, pigs (Sus scrofa).
when managed animals were moved
from the natural habitat of wild goats
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 herds 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 regions 500–                      different populations of animals were
Younger Dryas 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
elsewhere in the Fertile Crescent are            slower, to that of sheep (Fig. 1) (10, 14).                 with the ancestors of core livestock spe-
detecting parallel patterns to those doc-        Morphologically altered domestic pigs                       cies that unfolded across much of the
umented in the Zagros. Changes in the            are not found in the southern Levant or                     Fertile Crescent. Over time hunting
age of harvested caprines, and possibly          lowland Iran until ca. 8,500–8,000 B.P.                     strategies aimed at maximizing local
demographically driven changes in size           Recent demographic evidence suggests                        availability of wild ungulates developed
consistent with early herd management,           that taurine cattle were initially domesti-                 into active management, with all four
are found in southeastern Anatolia at            cated somewhere in the upper Eu-                            major livestock species coming under
ca. 10,500 B.P. (12). Sheep seem to be           phrates Valley between ca. 11,000 and                       management over a period from ca.
the initial early focus of the transition        10,000 B.P. (15), but, like sheep and                       11,000 to 10,000 B.P. Even species like

11598 cgi doi 10.1073 pnas.0801317105                                                                                                  Zeder
gazelle, which are behaviorally unsuited
to domestication, may have been audi-
tioned for management in the southern
and northern Levant, where they were                                                          000
the most abundant wild ungulate (11).

                                                  7,400 7,300

Clear-cut morphological responses to                                                                               10


domestication (i.e., changes in horns in

bovids and tooth size in pigs) are not



evident in these four livestock species





until ca. 9,500–9,000 B.P.                                                       7,000


                                                                                                                                                               , 50
   As is the case with animal domestica-                                                                                                                     0
tion in the Near East, the leading edge
of plant domestication in the region is                                                                                      7,000

now recognized as an extended process
(24, 25). Evidence from multiple loca-
tions point to a prolonged period of hu-      Fig. 2. An integrated model of the Neolithic expansion in the Mediterranean Basin. The location of
man manipulation of morphologically           colonist farming enclaves is shown in the red ellipses. Approximate dates of these enclaves are given inside
wild, but possibly cultivated, plants         the ellipses in calibrated years B.P. Red dots represent areas that are proposed to have been settled by
which, in certain species, resulted in the    colonist farmers; green dots indicate areas where indigenous foragers adopted elements of the Neolithic
development of morphologically altered        package; and blue dots indicate areas of proposed integration of colonist farmers with indigenous
domesticated crops (26–28). This period       foraging groups. Data were complied from refs. 52, 54, 56, 57, and 65 and figure 7.1 of ref. 74.
of intensified plant management dates at
least as far back as ca. 12,000 B.P., with
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).
cereals) 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 Pleistocene 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 pig 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 wheat, 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 Cyprus 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 de-                      was thought to have been colonized ca.
indigenous hunter–gatherers at a pre-         scendents of the progenitors of indige-                      8,500 B.P. by a derived offshoot of fully
dicted average pace of 1 km per year.         nous domestic caprines, are instead the                      established Neolithic mainland cultures
   Objecting to the passive role this         feral 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, lead 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 oats, barley, and        Mediterranean, argued that cultural and                      colonists also imported mainland game
lentils in Upper Paleolithic and Meso-        not demic diffusion was the primary en-                      animals like fallow deer and fox that,
lithic levels at Francthi Cave on the         gine driving this transition. Specifically,                  although perhaps kept in captivity (48),
eastern coast of Greece, followed by the      proponents maintained that the selective                     were never truly domesticated. None of
appearance of fully domesticated barley       adoption of various elements of the                          these animals are endemic to Cyprus.
and lentils in later Neolithic levels, was    Neolithic package by indigenous popula-                      Although imported livestock species did
interpreted as evidence for the local         tions around the Mediterranean could                         not show any of the morphological fea-
crop domestication (34). Legumes recov-       have happened through trade and tech-                        tures traditionally used to mark domes-
ered from Mesolithic cave deposits in         nology transfer alone without any direct                     tic status when they arrived on the
southern France were seen as evidence         contact between indigenous hunter–                           island, demographic profiles of these

Zeder                                                                                               PNAS            August 19, 2008            vol. 105   no. 33        11599
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 wells             head colony of seafaring migrant farm-       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-            pottery, domestic sheep, einkorn, and        of colonist expansion, selective local
cated einkorn and emmer 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 flax, 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, Zilhao (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, Zilhao (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 Middle East and Asia (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-
sula ca. 8,000 B.P. by maritime colonists        favorable coastal locals around the pe-      ductions out of Africa or overland
who first established farming villages on        riphery of the Iberian Peninsula at a        through Europe.
the Apulian ‘‘boot heel’’ region of              steady and quite rapid pace, appearing          Genetic data also support a pattern of
southeastern Italy (Fig. 2). These tradi-        first on the eastern and southern coasts     multiple introductions of cattle into the
tions appear in northwest coastal Italy          of Spain at ca. 7,700–7,600 B.P. and on      region. The T3 haplotype of domestic
   200–300 years later (ca. 7,800–7,600          the Atlantic coast of Portugal ca. 7,400–    cattle, which dominates among modern
B.P.). In southern France, a compelling          7,300 B.P. (Fig. 2).                         and ancient European cattle, seems to
case can be made for a marked geo-                                  ˜
                                                    However, Zilhao’s (61, 62) work in        have followed a relatively rapid path of
graphic, ecological, and cultural break          Portugal has also shown that Mesolithic      expansion around the Mediterranean

11600 cgi doi 10.1073 pnas.0801317105                                                                                Zeder
          Table 1. The extinction of Late Pleistocene large endemic mammals and human colonization of
          Mediterranean islands

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

          Most recent endemic                        5,000*                        10,000†                        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.

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 Central Europe (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 humans 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 North Africa 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 Balkans 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
rope. 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 tell 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 sea-
try, but instead are most closely related              not independently domesticated in Eu-                      side rock shelter 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, stratum containing stone
extracted from archaeological remains                  wild boar, possibly occurring multiple                     tools and hearths dated to ca. 11,775
has found convincing evidence for the                  times in geographically separate sub-                      B.P. Other researchers, although ac-
dispersal of Near Eastern pigs into and                populations. Genetic studies of rye and                    cepting the evidence for early visits of
across Europe between 7,500 and 5,000                  oats also indicate that the modern vari-                   hunter–gatherers to the island, question
B.P. (23). Surprisingly, subsequent to                 eties of these major crop plants have a                    the evidence for human predation on
this initial diffusion, Near Eastern swine             European and not Near Eastern ances-                       the endemic mammals discovered at the
are later replaced by domestic pigs of                 try (24). Future interpretive frameworks                   site (refs. 80 and 81, but see ref. 82).
European maternal ancestry, even                       will have to take a more integrated ap-                       Large endemic mammals on Crete
within the Near East. Indigenous do-                   proach, which recognizes colonization,                     included pygmy hippopotamus (Hippo-
mestication of European boar also ap-                  diffusion, and independent domestica-                      potamus creutzburgi), elephants (Elephas
parently happened several times, with                  tion as all playing a role in Neolithic                    creutzburgi), and several species of

Zeder                                                                                                     PNAS      August 19, 2008      vol. 105     no. 33   11601
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 Upper Paleolithic (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 rat-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
land fauna just before, or slightly after,       philous small vertebrate species (76)         may require the appearance of specific
substantial human colonization was,              clearly implicates humans in these island     morphological traits before conferring
until recently, thought to be a small            extinctions.                                  domestic status, others may be more will-
derived caprine species (Mytotragus                 The impact of the Neolithic transition     ing to consider a managed animal or a
balearicus), found only on the Gymnesic          on mainland environments throughout           cultivated plant as having achieved this
islands off the eastern coast of Spain           the Mediterranean, although perhaps           status. Regardless of where one chooses
(89). Here initial dating argued for a           less dramatic, is no less pronounced.         to draw the line between wild and domes-
   3,000-year overlap between initial hu-        Blondel and Aronson (92), in fact, argue      tic, recent advances have provided re-
man presence on Mallorca at 8,000 B.P.           that the entire Mediterranean Basin is        searchers with powerful new tools capable

11602 cgi doi 10.1073 pnas.0801317105                                                                                 Zeder
of examining the entire process of domes-                        all call out for additional investigation.                      pigs, however, still cannot be ruled out.
tication, not just its morphological impacts                     The southern margin of the Mediterra-                           Application of enhanced archaeological
which, if they occur at all, only appear                         nean Basis along coastal North Africa is                        and genetic techniques for detecting and
after the process is well underway.                              essentially terra incognita for understand-                     dating domestication to both extant and
   Just how far back this process of active                      ing the course of Neolithic emergence and                       yet-to-be-recovered assemblages is key to
human resource management goes or how                            seems an especially promising region for                        understanding patterns of indigenous do-
widespread it was in the Fertile Crescent                        future research (60).                                           mestication around the Mediterranean
is, at this point, an open question. The                            The subsequent inland transfer of do-                        Basin.
early dispersal of an integrated economy                         mesticates, agriculture, and associated                            Finally, although we may never be able
based on crop plants and managed ani-                            Neolithic lifeways from newly arrived                                                          ˆ
                                                                                                                                 to detect the final coup de grace for en-
mals to Cyprus at least 2,000 years before                       colonists to indigenous populations                             demic island faunas, the future holds con-
the apparent crystallization of agricultural                     around the Mediterranean Basin is an-                           siderable promise for a much fuller
economies on the mainland, however, sug-                         other intriguing research area that will                        understanding of the human impact on
gests that our understanding of plant and                        benefit from recent advances in our                             Mediterranean biodiversity. As it has in
animal domestication and agricultural                            ability to detect and date domesticates                         the Gymnesic islands and on Cyprus,
                                                                                                                                 ‘‘carpet-dating’’ large numbers of archaeo-
emergence on the mainland is, at best,                           in the archaeological record. Careful
                                                                                                                                 logical materials by the small-sample
incomplete. Researchers working in this                          analysis of increasingly more precise ra-
                                                                                                                                 atomic mass spectrometry radiocarbon
area are only just beginning to realize the                      diocarbon dates will continue to be criti-
                                                                                                                                 method will certainly help refine the chro-
potential of these newly available analytic                      cal in discriminating between demic                             nology of the disappearance of endemic
tools. Additional even more effective ana-                       diffusion and selective adoption of Neo-                        island faunas and the arrival of human
lytical approaches will almost certainly be                      lithic components in different parts of                         colonists. Application of demographic
developed in the future as researchers                           the Mediterranean (e.g., ref. 94). New                          profiling techniques to the remains of
embrace this broader concept of domesti-                         demographic techniques for profiling                            these animals may make it possible to dis-
cation and begin to exploit the many op-                         prey strategies, morphometric tech-                             tinguish between natural-death accumula-
portunities available in the Fertile Cres-                       niques capable of tracking genetic and                          tions and prey assemblages resulting from
cent region for documenting it.                                  plastic responses to human management,                          human predation. Similarly, recognition of
   Recent research has also shown that the                       isotopic analysis, and the increasing suc-                      the broader role of humans in shaping
dispersal of domesticates and the Neo-                           cess of ancient DNA studies of domesti-                         post-Neolithic environments is central to
lithic way of life west across the Mediter-                      cates will enhance our understanding of                         understanding how Mediterranean biodi-
ranean Basin was much more complex                               the ways in which both colonists and                            versity evolved and how we might best
and multifaceted than previous prime                             local populations adapted management                            work to conserve it. The archaeobiological
mover models could accommodate. To                               strategies to these new environments.                           sciences have a valuable role to play in
varying degrees, in different areas, this                           There are also obvious opportunities                         providing greater time depth to biodiver-
process involved elements of demic diffu-                        for those interested in understanding the                       sity studies by monitoring the creation of
sion, local adoption, and independent                            independent domestication of European                           anthropogenic ecosystems and tracing the
domestication. But the outlines of this                          wild species. Larson et al. (23), for exam-                     development and impacts of both environ-
complex process are just beginning to                            ple, suggest that European wild boar were                       mentally sustainable and destructive agri-
come into focus. Maritime colonization of                        domesticated only after the introduction                        cultural economies over thousands of
the Mediterranean clearly involved not                           of Near Eastern domestic swine, repre-                          years of human occupation.
one, but multiple unrelated seaborne mi-                         senting a case of apparent technology
grations (52). The cultural context of                           transfer rather than truly independent                          ACKNOWLEDGMENTS. This work has benefited
                                                                                                                                 from the comments of Greger Larson, Bruce
these migratory movements, their causes,                         domestication. Local, culturally indepen-                       Smith, and Jean-Denis Vigne and from the art-
their routes, their timing, and their tempo                      dent domestication of indigenous wild                           istry of Marcia Bakry, who drew the maps.

 1. Zeder MA, Emshwiller E, Smith BD, Bradley DG, eds             8. Zeder MA (2006) A critical examination of markers of                                                             ˜
                                                                                                                                 15. Helmer D, Gourichon L, Monchot H, Peters J, Sana Segui
    (2006) Documenting Domestication (Univ of California             initial domestication in goats (Capra hircus). Docu-            M (2005) Identifying early domestic cattle from prepot-
    Press, Berkeley).                                                menting Domestication, eds Zeder MA, Emshwiller E,              tery Neolithic sites on the middle Euphrates using sex-
 2. Zeder MA (2006) Archaeological approaches to docu-               Smith BD, Bradley DG (Univ of California Press, Berke-          ual dimorphism. The First Steps of Animal Domestica-
    menting animal domestication. Documenting Domes-                 ley), pp 181–208.                                               tion, eds Vigne J-D, Peters J, Helmer D (Oxbow Books,
    tication, eds Zeder MA, Emshwiller E, Smith BD, Bradley       9. Zeder MA (2006) Reconciling rates of long bone fusion           Oxford), pp 86 –95.
    DG (Univ of California Press, Berkeley), pp 171–180.             and tooth eruption and wear in sheep (Ovis) and goat        16. Martin L, Russell N, Carruthers D (2002) Animal remains
 3. Uerpmann H-P (1979) Problems of the Neolithization of            (Capra). Ageing and Sexing Animals from Archaeolog-             from the central Anatolian Neolithic. The Neolithic of
    the Mediterranean Region (original in German). Supple-           ical Sites, ed Ruscillo D (Oxbow Press, Oxford), pp 87–                                  ´
                                                                                                                                     Central Anatolia, eds Gerard F, Thissen L (Ege Yayınları,
    ments to the Tuebingen Atlas of the Near Orient, series B.       118.                                                            ˙
                                                                                                                                     Istanbul, Turkey), pp 193–206.
    (Ludwig Reichert, Weisbaden).                                10. Zeder MA (2009) The Neolithic (macro)-revolution. J         17. Bruford M, Townsend SJ (2006) Mitochondrial DNA
 4. Meadow RH (1989) Osteological evidence for the pro-
                                                                     Archaeol Res, in press.                                         diversity in modern sheep: Implications for domestica-
    cess of animal domestication. The Walking Larder, ed
                                                                 11. Redding RW (2005) Breaking the mold, a consideration            tion. Documenting Domestication, eds Zeder MA, Em-
    Clutton-Brock J (Unwin Hyman, London), pp 80 –90.
                                                                     of variation in the evolution of animal domestication.          shwiller E, Smith BD, Bradley DG (Univ California Press,
 5. Bar-Yosef O, Meadow RH (1995) The origins of agricul-
                                                                     The First Steps of Animal Domestication, eds Vigne J-D,         Berkeley), pp 306 –316.
    ture in the Near East. Last Hunters, First Farmers, eds
                                                                     Peters J, Helmer D (Oxbow Books, Oxford), pp 41– 48.                              ´
                                                                                                                                 18. Luikart G, Fernandez H, Mashkour M, England PR,
    Price TD, Gebauer, A-B (School of American Research
                                                                 12. Peters J, von den Driesch A, Helmer D (2005) The                Taberlet P (2006) Origins and diffusion of domestic
    Press, Santa Fe, NM), pp 39 –94.
                                                                     upper Euphrates-Tigris Basin, cradle of agro-pasto-             goats inferred from DNA markers. Documenting Do-
 6. Horwitz LK (1993) The development of ovicaprine do-
    mestication during the PPNB of the southern Levant.              ralism? The First Steps of Animal Domestication, eds            mestication, eds Zeder MA, Emshwiller E, Smith BD,
    Archaeozoology of the Near East I, eds Buitenhuis H,             Vigne J-D, Peters J, Helmer D (Oxbow Books, Oxford),            Bradley DG (Univ of California Press, Berkeley), pp 294 –
    Clason AT (Universal Book Service, Leiden, The Nether-           pp 96 –124.                                                     305.
    lands), pp 27–36.                                            13. Vigne J-D, Buitenhuis H, Davis S (1999) The first steps in   19. Naderi S, et al. (2007) Large-scale mitochondrial DNA
 7. Vigne J-D, Peters J, Helmer D (2005) New archaeozoologi-         animal domestication into the east of the Euphrates:            analysis of the domestic goat reveals six haplogroups
    cal approaches to trace the first steps of animal domesti-        Cypres and Central Anatolia (original in French). Pale-´        with high diversity. PLos One 10:1–23.
    cation. The First Steps of Animal Domestication, eds             orient 25/2:49 – 62.                                        20. Dobney K, Larson G (2006) Genetics and animal domes-
    Vigne J-D, Peters J, Helmer D (Oxbow Books, Oxford), pp      14. Ervynck A, Dobney K, Hongo H, Meadow RH (2001)                  tication. J Zool 286:261–271.
    1–16.                                                                            ´
                                                                     Born free! Paleorient 27/2:47–73.

Zeder                                                                                                                     PNAS      August 19, 2008         vol. 105      no. 33       11603
21. Bradley DG (2006) Genetics and the origins of domestic       45. Dennell R (1992) The origin of crop agriculture in Eu-       68. Pereira F, et al. (2006) Genetic signatures of a Mediter-
    cattle. Documenting Domestication, eds Zeder MA,                 rope. The Origins of Agriculture, eds Cowan CW,                  ranean influence in Iberian Peninsula sheep hus-
    Emshwiller E, Smith BD, Bradley DG (Univ of California           Watson PJ (Smithsonian Institution Press, Washington,            bandry. Mol Biol Evol 23:1420 –1426.
    Press, Berkeley), pp 317–328.                                    DC), pp 71–100.                                              69. Edwards C, et al. (2007) Mitochondrial DNA analysis
22. Larson G, et al. (2005) Worldwide pylogeography of           46. Lewthwaite J (1986) The transition to food production.           shows a Near Eastern origin for domestic cattle and no
    wild boar reveals multiple centers of pig domestica-             Hunters in Transition: Mesolithic Societies in Temper-           indication of domestication of European aurochs. Proc
    tion. Science 307:1618 –1621.                                    ate Eurasia and Their Transition to Farming, ed Zvelebil         R Soc London Ser B 274:1377–1385.
23. Larson G, et al. (2007) Ancient DNA, pig domestication,          M (Cambridge Univ Press, Cambridge, UK), pp 53– 66.          70. Cymbron T, et al. (2005) Microsatellite diversity sug-
    and the spread of the Neolithic into Europe. Proc Natl       47. Guilaine J, LeBrun eds (2003) The Neolithic of Cyprus            gests different histories for Mediterranean and
    Acad Sci USA 104:15276 –15281.                                   (original in French) École Français d’Athèns, Athens,            Norther Eurpean cattle populations. Proc R Soc London
24. Weiss E, Kislev ME, Hartmann A (2006) Autonomous                 Bull Corr Hellenique Suppl 43.
                                                                                 ´                                                    Ser B 272:1837–1843.
    cultivation before domestication. Science 312:1608 –         48. LeBrun A, Cluzan S, Davis SJM, Hansen J, Renault-            71. Beja-Pereira A, et al. (2006) The origin of European
    1610.                                                            Mislkovsky (1987) The pre-ceramic Neolithic of Cyprus            cattle. Proc Natl Acad Sci USA 103:8113– 8118.
25. Wilcox G, Fornite S, Herveux L (2007) Early Holocene             (original in French). L’Anthropologie 91:283–316.            72. Achilli A, et al. (2008) Mitochrondrial genomes of ex-
    cultivation before domestication in northern Syria.          49. Peltenburg E (2004) Introduction. Neolithic Revolution,          tinct aurochs survive in domestic cattle. Curr Biol
    Vegetative History Archaeobot, 10.1007/s00334 – 007-             eds Peltenburg E, Wasse A (Anthony Row, Chippen-                 18:157–158.
    0121-y.                                                          ham, UK), pp. xi–xx.                                         73. Pellecchia M, et al. (2007) The mystery of Etruscan
26. Wilcox G (2002) Geographical variation in major cereal       50. Vigne J-D, Carrere I, Guilaine J (2003) Unstable status of
                                                                                     ´                                                origins. Proc R Soc London Ser B 274:1175–1179.
    components and evidence for independent domestica-               early domestic ungulates in the Near East. The Neo-          74. Zvelebil M (2000) The social context of the agricultural tran-
    tion events in Western Asia. The Dawn of Farming in              lithic of Cyprus (original in French), eds Guilaine J,           sition to Europe. Archaeogenetics: DNA and the Population
    the Near East, eds Cappers RTJ, Bottema S (Ex Oriente,           LeBrun A École Français d’Athèns, Athens, Bull Corr              Prehistory of Europe, eds Renfrew C, Boyle K (McDonald
    Berlin), pp 133–140, Studies in Near Eastern Production,         Hellenique, Suppl 43, pp. 239 –251.
                                                                        ´                                                             Institute Monographs, Oxbow, Oxford), pp 57–80.
    Subsistence, and Environment no. 6.                          51. Colledge S (2004) Reappraisal of the archaeobotanical        75. Blondel J, Vigne J-D (1993) Space, time, and man as
27. Wilcox G (2005) The distribution, natural habitats, and          evidence for the emergence and dispersal of the                  determinants of diversity of birds and mammals in the
    availability of wild cereals in relation to their domesti-       ‘‘founder crops.’’ Neolithic Revolution, eds Peltenburg          Mediterranean Region. Species Diversity in Ecological
    cation in the Near East. Vegetative History Archaeobot           E, Wasse A (Anthony Row, Chippenham, UK), pp 49 –                Communities, eds Ricklefs, RE, Schluter D (Univ Chicago
    14:534 –541.                                                     60.                                                              Press, Chicago), pp 135–146.
28. Kislev ME, Hartmann A, Bar-Yosef O (2006) Early do-          52. Guilaine J (2003) The Neolithization in the Mediterra-       76. Vigne J-D (1999) The large ‘‘true’’ Mediterranean is-
    mesticated fig in the Jordan Valley. Science 312:1375–            nean and France. The Widening Harvest, eds Ammer-                lands as a model for the Holocene human impact on the
    1374.                                                            man A, Biagi P (Archaeological Institute of America,             European vertebrate fauna? The Holocene History of
                                                                                                                                      the European Vertebrate Fauna, ed Benneke N (Deut-
29. Nesbitt M (2002) When and where did domesticated                 Boston), pp 189 –206.
                                                                                                                                      sches Archaologisches Institut, Eurasien-Abteilung,
    cereals first occur in Southwest Asia? The Dawn of            53. Broodbank C (1999) Colonization and configuration in
                                                                                                                                      Berlin), pp 295–322.
    Farming in the Near East, eds Cappers RTJ, Bottema S             the insular Neolithic of the Aegean. Neolithic Society in
                                                                                                                                  77. Alcover JA, Seguı B, Bover P (1999) Extinctions and local
    (Ex Oriente, Berlin), pp 113–132, Studies in Near Eastern        Greece, ed Halstead P (Sheffield Academic, Sheffield,
                                                                                                                                      disappearances of vertebrates in the western Mediter-
    Production, Subsistence, and Environment no. 6.                  UK), pp 15– 41.
                                                                                                                                      ranean islands. Extinctions in Near Time, ed McPhee
30. Ammerman AJ, Cavalli-Sforza LL (1984) The Neolithic                ´
                                                                 54. Perles C (2001) The Early Neolithic in Greece (Cam-
                                                                                                                                      RDE (Kluwer, New York), pp 165–188.
    Transition and the Genetics of Populations in Europe             bridge Univ Press, Cambridge, UK).
                                                                                                                                  78. Cucchi T, et al. A new endemic species of the subgeneus
    (Princeton Univ Press, Princeton).                           55. Runnels C (2003) The origins of the Greek Neolithic. The
                                                                                                                                      Mus (Rodentia Mammalia) on the island of Cyprus.
31. Higgs E, Jarman MR (1969) The origins of agriculture.            Widening Harvest, eds Ammerman A, Biagi P (Archae-
                                                                                                                                      Zootaxa 1241:1–36.
    Antiquity 43:31– 41.                                             ological Institute of America, Boston), pp 121–132.
                                                                                                                                  79. Simmons A (1999) Faunal Extinction in an Island Society
32. Dennell R (1983) European Economic Prehistory (Aca-          56. Biagi P (2003) A review of the late Mesolithic in Italy
                                                                                                                                      (Kluwer, New York).
    demic, London).                                                  and its implication for the Neolithic transition. The
                                                                                                                                  80. Bunimovitz S, Barkai R (1996) Ancient bones and mod-
33. Barker G (1985) Prehistoric Farming in Europe (Cam-              Widening Harvest, eds Ammerman A, Biagi P (Archae-
                                                                                                                                      ern myths. J Med Arch 9:85–96.
    bridge Univ Press, Cambridge, UK).                               ological Institute of America, Boston), pp 133–156.
                                                                                                                                  81. Ammerman A, Noller J (2005) New light on Aeotokrem-
34. Hansen JM, Renfrew JM (1978) Paleolithic-Neolithic           57. Skeates R (2003) Radiocarbon dating and interpreta-
                                                                                                                                      nos. World Archaeol 37:533–543.
    seed remains at Francthi Cave, Greece. Nature                    tions of the Mesolithic-Neolithic transition in Italy. The
                                                                                                                                  82. Simmons A, Mandel R (2007) Not such a new light.
    271:349 –352.                                                    Widening Harvest, eds Ammerman A, Biagi P (Archae-
                                                                                                                                      World Archaeol 39:475– 482.
35. Vaquer J, Geddes D, Barbaza M, Erroux J (1986) Meso-             ological Institute of America, Boston), pp 157–188.
                                                                                                                                  83. Reese DS, Belluomini G, Ikeya M (1996) Absolute dates
    lithic plant exploitation at the Balma Abeurador             58. Binder D (2000) Mesolithic and Neolithic interaction in
                                                                                                                                      for the Pleistocene fauna of Crete. Pleistocene and
    (France). Oxford J Arch 5:1–18.                                  southern France and northern Italy. Europe’s First
                                                                                                                                      Holocene Fauna of Crete and its First Settlers, ed Reese
36. Ducos P (1976) Some evidence for the origin of domes-            Farmers, ed Price TD (Cambridge Univ Press, Cam-                 DS (Prehistory Press, Madison, WI), pp 47–51.
    tication in France (original in French). The Prehistory of       bridge, UK), pp 117–145.                                     84. Broodbank C, Strasser TF (1991) Migrant farmers and the
    France (original in French), ed Guilaine J (Centre Na-       59. Guilaine J, Manen C, Vigne J-D (2008) New Insights into          Neolithic colonization of Crete. Antiquity 65:233–245.
    tional de la Recherche Scientifique, Paris), pp 165–177.          the Neolithization of Mediterranean France (original         85. Sondaar PY, Martini F, Ulzega A, Klein G, Hofmeijer
37. Estevez J (1988) Study of the faunal remains (original in        in French) (Centre d’Anthropologie, Toulouse, France),           (1991) Pleistocene Man in Sardinia (original in French).
    Spanish). Cova Fosca, ed Olaria C (Servico de Publica-           in press.                                                        L’Anthropologie 95:181–200.
                      ´              ´          ´
    ciones Diputacion de Castellon, Castellon, Spain), pp                 ˜
                                                                 60. Zilhao J (1993) The spread of agro-pastoral economies        86. Vigne J-D (2005) The first evidence of modern humans in
    281–337.                                                         across Mediterranean Europe. J Med Archaeol 6:5– 63.             Corsica and Sardinia (original in French). Human Migra-
38. Boessneck J, von den Driesch A (1980) Finds of animal                 ˜
                                                                 61. Zilhao J (2000) From the Mesolithic to the Neolithic in          tion and Environmental Variation in the Quaternary
    bones from four caves in southern Spain (original in             the Iberian Peninsula. Europe’s First Farmers, ed Price          (original in French), eds Hedges J, Hedges E (Archaopress,
    German). In Studies on finds of early animal bones                TD (Cambridge Univ Press, Cambridge, UK), pp 144 –               Oxford), pp 139 –145, BAR International Series 1352.
    from the Iberian Peninsula 7, eds Boessneck J, von den           183.                                                         87. Vigne J-D, Desse-Berset N (1995) The exploitation of
    Driesch A (Institut fur Palaeoanatomir, Domestika-                    ˜
                                                                 62. Zilhao J (2001) Radiocarbon evidence for maritime pi-            animal resources in the Mediterranean Islands during
    tionsforschung und Geschichte de Tiermedizin der Uni-            oneer colonization at the origins of farming in west             the Preneolithic, the example of Corsica. Man and Sea
           ¨    ¨
    versitat Munchen, Munich), pp 1– 83.                             Mediterranean Europe. Proc Natl Acad Sci USA                     in the Mesolithic, ed Fisher A (Oxbow Books, Oxford),
39. Zohary D (1996) The mode of domestication of the                 98:14180 –14185.                                                 pp 309 –318.
    founder crops of Southwest Asian agriculture. The Or-        63. Guilaine J, et al. (1993) Dourgne (Centre                    88. Vigne J-D (1996) Small mammal fossil assemblages as
    igins and Spread of Agriculture and Pastoralism in               d’Anthropologie de Societes Rurales, Archeologie en              indicators of environment change in northern Corsica
    Eurasia, ed Harris D (Smithsonian Institution Press,             Terre d’Aude, Toulouse/Carcassonne, France).                     during the last 2,500 years. J Archaeol Sci 23:199 –215.
    Washington DC), pp 142–158.                                  64. Rowley-Conwy P (2003) Early domestic animals in Eu-          89. Bover P, Alcover JA (2003) Understanding Late Quater-
40. Geddes DS (1985) Mesolithic domestic sheep in west               rope. The Widening Harvest, eds Ammerman A, Biagi P              nary extinctions, J Biogeog 30:771–781.
    Mediterranean Europe. J Archaeol Sci 12:25– 48.                  (Archaeological Institute of America, Boston), pp 99 –       90. Waldren WH (1982) Balearic Prehistoric Ecology and
41. Poplin F, et al. (1986) The beginning of herding in              120.                                                             Culture (Archaeopress, Oxford), British Archaeological
    France (original in French). The French Neolithic (orig-     65. Tresset A, Vigne J-D (2007) Substitution of species,             Reports IS 149.
    inal in French), edd Demoule J-P, Guilaine J (Picard,            techniques and symbols at the Mesolithic-Neolithic           91. Ramis D, Bover P (2001) A review of the evidence for
    Paris), pp 37–51.                                                transition in western Europe. Proc Br Acad 144:189 –             domestication of Myotragus balearicus Bate 1909 (Ar-
42. Guilaine J (1976) First Herders and Farmers in the               210.                                                             tiodactyla, Caprinae) in the Balearic Islands. J Archaeol
    Occidental Mediterranean (original in French) (Mou-                     ´
                                                                 66. Fernandez H, et al. (2006) Divergent mtDNA of goats in           Sci 28:265–282.
    ton, Paris).                                                     an Early Neolithic site, far from the initial domestica-     92. Blondel J, Aronson J (1999) Biology and Wildlife of the
43. Price TD (1987) The Mesolithic of Western Europe. J              tion areas. Proc Natl Acad Sci USA 103:15375–15379.              Mediterranean Region (Oxford Univ Press, Oxford).
    World Prehist 1:225–305.                                     67. Pereira F, Pereira L, Van Asch B, Bradley DG, Amorim A       93. Zeder MA (2006) Central questions in the domestica-
44. Zvelebil M (1989) On the transition to farming in Eu-            (2005) The mtDNA catalogue of all Portuguese auto-               tion of plants and animals. Evol Anthropol 15:105–117.
    rope, or what was spreading with the Neolithic. Antiq-           chothonous goat (Capra hircus) breeds. Mol Ecol              94. Gkiastra M, Russell T, Shennan, Steele J (2003) Neolithic
    uity 63:379 –383.                                                14:2313–2318.                                                    transition in Europe. Antiquity 77:45– 62.

11604 cgi doi 10.1073 pnas.0801317105                                                                                                                                    Zeder

To top