On the origins and spread of Olea europaea L.

(olive) domestication: evidence for shape

variation of olive stones at Ugarit, Late Bronze Age, Syria—a window on the
Mediterranean Basin and on the westward diffusion of olive varieties
Author(s): Claire Newton, Christine Lorre, Caroline Sauvage, Sarah Ivorra and Jean-
Frédéric Terral
Source: Vegetation History and Archaeobotany , September 2014, Vol. 23, No. 5
(September 2014), pp. 567-575
Published by: Springer

Stable URL: https://www.jstor.org/stable/43554270

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Veget Hist Archaeobot (2014) 23:567-575
DOI 1 0. 1 007/s00334-0 1 3-04 1 2-4

ORIGINAL ARTICLE

On the origins and spread of Olea europaea L. (olive)

domestication: evidence for shape variation of olive stone
Ugarit, Late Bronze Age, Syria - a window on the Mediter
Basin and on the westward diffusion of olive varieties

Claire Newton • Christine Lorre • Caroline Sauvage •

Sarah Ivorra • Jean-Frédéric Terral

Received: 15 April 2013 /Accepted: 11 August 2013 /Published online: 27 August 2013
© Springer- Verlag Berlin Heidelberg 2013

Abstract Charred archaeological stones of Olea euro- by comparison to archaeological material from that area.
paea L. (olive) from Late Bronze Age Ugarit, Syria, were
Combining morphometric and genetic data, modern refer-
analyzed with geometric morphometry and compared withence and archaeological material also guides us in under-
a morphological differentiation model established on thestanding the mechanisms that prevailed in the long-term
basis of analyses of modern spontaneous (uncultivated) agrobiodiversity of the olive.
olive populations and cultivated varieties of various origins
within the Mediterranean Basin. The results allow a rein- Keywords Agricultural biological diversity •
terpretation of the east-west morphological diversity Archaeological
pre- olive stones • Diffusion • Geometric
morphometry • Olea europaea
viously observed in wild olives. The archaeobotanical data
were compared in detail to the partly geographically
structured modern morphological diversity of the culti-
Introduction
vated olive. Ancient morphotypes could be distinguished,
among which one is dominant in the assemblage. Their
diffusion from east to west is shown, and their time Olive
of trees became a symbol of the Mediterranean world as
arrival in the northwestern Mediterranean can be evaluated a result of a long-term process which began several million
years ago with the establishment of the Mediterranean
Basin and climate (Sue 1984; Mai 1989; Krijgsman et al.
Communicated by G. Willcox. 1999; Quézel and Médail 2003) and finalized by the use
and manipulation of the olive by human societies
Electronic supplementary material The online version of this
article (doi: 10. 1007/s00334-0 13-04 12-4) contains supplementary (Kaniewski et al. 2012). Indeed in the Mediterranean Basin,
material, which is available to authorized users.

C. Newton (El) • S. Ivorra • J.-F. Terral C. Sauvage

Centre de Bio- Archéologie et d'Ecologie, Institut de Botanique,
UMR 5059 CNRS/Université Montpellier 2/EPHE/INRAP,
163, rue Auguste Broussonet, 34090 Montpellier, France
e-mail: clairennewton@gmail.com
J.-F. Terrai
e-mail: terral@univ-montp2.fr
Pitzer College, 1050 North Mills Avenue, Claremont,
CA 91711-6101, USA
C. Sauvage
Archeorient, UMR 5133 - CNRS/Université Lumière Lyon 2,
Maison de l'Orient et de la Méditerranée - Jean Pouilloux,
7 rue Raulin, 69365 Lyon Cedex 7, France

C. Newton
J.-F. Terral
Laboratoire d'Archéologie et de Patrimoine, Université du Université Montpellier 2, Place Eugène Bataillon,
Québec à Rimouski, 300, allée des Ursulines, Rimouski,
34095 Montpellier Cedex, France
QC G5L 3 Al, Canada

C. Lorre
Musée d'Archéologie Nationale, Château - Place Charles
de Gaulle, 78105 Saint-Germain-en-Laye, France

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 568 Veget Hist Archaeobot (2014) 23:567-575

olive tree distribution is used to delimit the Mediterranean domestication took place independently in several regions,
climate zone. This correlation seems relevant only for Oleaand not exclusively in the Near Eastern centre, from which
europaea ssp. europaea var. sativa Lehr (cultivated olive),derived the most prevalent agrolineages (Besnard et al.
but not for Olea europaea ssp. europaea var. sylvestris (Mill.)2002; Khadari 2005; Breton et al. 2006, 2009; Kaniewski
Lehr (wild olive, also referred to as oleaster or Olea oleaster et al. 2012). Nevertheless, the contribution of the occi-
Hoflmans. & Link) (Ozenda 1975; Rivas-Martinez 1987). dental genetic pool in the constitution of olive varietal
Therefore, as Daget (1984) suggested, olive trees, Medi-heritage seems to have been minor, compared to the
terranean climate and ecosystems may only be strictly related oriental one (Besnard et al. 2013).
inasmuch as societal, cultural and economic factors that The area of Ugarit, the ancient city of the present
emerged eight to nine millennia ago in the Levant are taken archaeological site of Ras Shamra, Syria, former capital of
into account (Zohary et al. 2012). the kingdom of Ugarit and economic centre at the crossroads
Olive history, particularly the origins of its cultivationof Hittite, Mesopotamian, Egyptian and Mediterranean
and domestication in the western Mediterranean, has spheres of influence (Fig. 1), yielded archaeological olive
always been a sensitive and controversial matter. Thestones which will be studied to characterize locally culti-
classic view is that a change from wild to cultivated olive vated varieties. Olive oil and fruit contributed to the com-
populations took place in the southern Levant during the mercial wealth of the city during the Late Bronze Age, and
Chalcolithic in the 4th millennium b.c.e. (Zohary andthe olive groves were located in the surrounding countryside,
Spiegel-Roy 1975; Neef 1990; Liphschitz et al. 1991;some of which at least were owned by the royal palace and
Kislev 1995). In areas where olive trees were not native,leased to farmers (Callot 1987). Local oil consumption and
such as Egypt, finds of remains such as wood charcoal or production were certainly important, as shown by the
numerous oil presses excavated in the city of Ugarit (Callot
olive stones indirectly provide a terminus ante quem for the
earliest cultivation practices. Locally, the development and1993, 1994). Texts found in the city also mention numerous
expansion of olive cultivation, technological developmentsexports of oil, probably olive oil. For instance, text RS 18.42
in agriculture, oil production and trade in conjunction withmentions 160 jars of oil released to a Cypriot and 144 jars of
environmental changes have been regarded as factorsoil to an Egyptian (Virolleaud 1965). Oil export lies at the
explaining the transition from the Chalcolithic to the Early
heart of this work, since the city was in a privileged position
Bronze Age, leading to the rise of city-states (Lovell 2002).
to engage in Late Bronze Age maritime trade of finely crafted
From this first Near Eastern centre there ensued a slow products such as ivory, metals or textiles, ores and agricul-
diffusion of domesticated olive forms, of technical tural produce, among which wine and oil were certainly
knowledge and cultivation practices, first to the Aegean in prominent (Sauvage 2012).
the 3rd millennium, then to the central and western By comparison to cultivated forms from northwestern
Mediterranean where domesticated olives have been Mediterranean sites of different periods (Terrai et al. 2004),
found in Italy and Spain in the Late Bronze Age,the around
analysis of olive stones from Ugarit may help to better
understand
1200-1000 B.C.E., and finally to southern France during the the timing of the diffusion of varieties which
1st millennium b.c.e. (van Zeist 1980). had been domesticated in the eastern Mediterranean and
Eco-anatomical and morphometrical approachessubsequently
devel- introduced westwards. This diffusion may
oped on wood/charcoal (Terral and Arnold-Simard have played an important role in the development and
1996;
Terral 1997) and olive stones respectively (Terraldiversification
et al. of olive agrobiodiversity in the western
Mediterranean.
2004), reconsidered that strict diffusionist model by dem-
onstrating that olive trees had probably been used and
cultivated in the northwestern Mediterranean since the end
of the Neolithic-Chalcolithic transition (Terral and Arnold- Materials and methods
Simard 1996; Terral 2000). Nevertheless, these results do
not in any way challenge the significant influence of The first step of this study consists in the analysis of the
Phoenicians, Etruscans, Greeks and Romans on the spread morphological structure of stones from extant populations.
of the olive across the whole Mediterranean Basin. It
Our reference collection comprises 498 stones collected
from
appears then, that the origins of olive cultivation and trees of 15 spontaneous (uncultivated) populations
domestication in the northwestern Mediterranean are more with 30 stones per population, except Slunfeh (Syria) for
ancient than previously thought. They seem to emerge two which we hold only 28 stones, and 1,517 stones belonging
millennia before the introduction of new varieties, know- to 51 cultivated varieties of various origins with 30 stones
how and techniques from various other Mediterranean per variety, except the one from the Menara garden, Mar-
lands (Terral et al. 2004). These results agree with models rakech, Morocco, for which only 17 were studied (Fig. la).
provided by molecular biology, which show that olive The geographical origin of the varieties is based on current

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 Veget Hist Archaeobot (2014) 23:567-575 569

Fig. 1 a, Geographical
France: origin
of the spontaneous populations
- Aglandeau
and cultivars of olive studied
- Belgentier
(the anonymous cultivar from - Cailletier
the Menara garden of - Colombaie
Marrakech, Morocco is named
-Comiale Italy: - Amygdalolia
"Menara" in Fig. 3); b, - Ghjermana (Corsica) - Ascolana Tenera . Carolia
Location of Ugarit and - Grossane . Bella di Spagna . Gaidouriola
Mahadou, Syria, with -Lucques -Cypressino - Kalamata
neighbouring cultural influences - Olivière . Grappola . Koroneiki
between 1500 and 1300 b.c.e - Picholine . Razzola . Kothreiki
-Tanche

-Verd
Spain: ' ' / / / -Ayvalik
- Arbequina r~~~ ' s - ^ ' / ? 7 / Domai
- - Arbequina Picual 'a r~~~ ' X* s - ^ ' : / / ? ¿ 7 /1 / -
- Ecijano ' Vv ' ; / / BUCAKKIÇLA2
- Manzanilla cr / / / / BUCAKXISLA3
j' _t''l '' ' SLUNFEH
MORAIRA - . / ^ J S r~
JOROX O* f) w'/ /Syria: r~Tir u w'

MEKNEŠ

/ 'C Mediterranean Sea - Khodeiry

/ 'C ' ^
rjf Spontaneous populations ' ' ' ^ Teffahi
/ it Cuhivuted varieties

M#rocco: . .
- Moroccan Picholine Algeria. . .

- Unnamed cultivar from - Ch

the Menara gardens of Kabylia . ßarouni ļ
(Marrakech)*

Meski Israel - Gaza Strip - Jordania:

ļj Hittite Empire ^ %*■ " Shami

v S Vj J (N Mitanni
V ^ ' A fand1/ '^e ^t>ł c' ^
[5 Ugarit Assyria
ť Cyprus i ' f
' sť ' /
/ A' ' Minet ei- Beidll
J ^-Mahadou
Mediterranean Sea J Egyptian Shamra

knowledge aboutIn the

where second
they sta
are g
consideration structure
recent of 199 ch
introductions.
tion, the Late
east-west Bronze Age
dividing a
line
Basin, which have
has beenshown
used thatru
here ch
Sea and the Libyan Desert;
stones this i
significantly
division tuting
proposed by the stone
Blondel ana
collection, at This stages
several batch,of n°76
its d
been used in d'Archéologie
previous studies Na
(T
Newton France,
et
2006; entered
Terral th
al.
2007). F
wild found
populations and in Minet
cultivated el-B
var
in Turkey and of the
Syria city
have of
been U
inc

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 570 Veget Hist Archaeobot (2014) 23:567-575

According to the excavation

olive stones belongs torecords,
locally produced olives it came
destined for fr
northern part of trench 8.IV,
oil production, in
with the anoccurring
pressing area in a interprete
workshop
olive press and located near
or house tomb
within the city of Mahadou. V and deposi
(Sauvage unpublished a). The
The olive precise
stones were date
studied by geometric morpho- of th
Bronze Age press is difficult
metry, which is to
a set ofknow.
methods allowingIt was or
the individual install
a pit burialgrouped
1) analysis of size
dating (sepulture
from and shape the
of the specimens
beginning
Late Bronze Age, in the(Adams et15th century
al. 2004). Previous b.c.e. (Sch
studies on similar biological
1949, Fig. 51; Sauvage and
unpublished
archaeological material showed thatb), and
the shape of the theref
olive stones is
press post-dates the burial. not significantly
The press dependent on growing
probably belo
conditions, V,
the building hosting tomb whether local or could
and regional in scale,
be nor contemp
is it
with, at least, the latest use
significantly of
altered this
by the tomb
process of during
charring (Terrai et al. t
century b.c.e. (Sauvage 2004). In the present paper, we use shape (standardizing
unpublished c). It thus seem
sonable to attribute asize)13th
parameters tocentury date and
compare olive stone morphologies to this
without however being able
reveal to confirm it due
distinct morphotypes. to a
associated datable The baseline
material. Thissuperimposition
press, methodlike
(Rohlf 1990)
other ex
from Ugarit, comprised developed
a millfor the characterization
where of the
olive stones (Terral
fruit were
and then crushed, and a circular stone slab used for oil et al. 2004; Newton et al. 2006) and later adapted to cherry
extraction (Fig. 2; Callot 1993). Schaeffer also mentions in stones (Burger et al. 201 1) was applied. The outline of each
his 1931 excavation notebook two or three large vessels valve (fertile valve: FV and sterile valve: SV) of the stone
(jars or pithoi) located near the olive stones. These vessels observed in lateral view is described by two configurations
may have been used for décantation, to store oil, or to store of point coordinates, using image analysis software. These
water used for a second warm extraction. They also may points are superimposed on a new reference grid, which is
have been used to store olives. However, the stones were defined so that the distance between the base (centre of the
found in the soil, and not inside these vases, suggesting that peduncle insertion zone) and the apex of the stone equals 1.
the olives were stored either directly on the floor or in The size of all stones is therefore standardized and the new
perishable containers such as bags or baskets. In any case, point configurations (two configurations of shape coordi-
it is likely from the archaeological context that this batch of nates per stone) are adjusted by a third-order fitted poly-
nomial regression. Finally, each stone is defined by a
system of two equations (fertile valve: Ypv = bo + bix+
b2X2 + bßX3; sterile valve: YSv = bó+b'1x + b^x2 + b'3x3).
Their quantitative parameters (b0, bb b2, b3, bó, b', b'2
and b3) are used as quantitative variables in a linear dis-
criminant analysis (LDA). The aim of this multivariate
statistical method is to evaluate the morphological variabil-
ity of olives by maximising differences between predefined
groups (spontaneous populations and varieties) compared to
the intra-group variation, and to quantify geometric differ-
ences between each of these spontaneous populations and
varieties using a dissimilarity matrix. That distance matrix is
eventually used in a statistical typological analysis (UPGMA
cluster analysis). Its role is to describe and partition the
morphological diversity into distinct morphotypes, whose
relevance is appreciated by a probability index that evaluates
their discrimination level as a percentage.
Finally, when different morphotypes are defined, the
quantitative data relative to the morphometric analysis of
the archaeological stones are compared with the initial
LDA. The archaeological stones are then put into a specific
reference morphotype with a probability of allocation. The
Fig. 2 Reconstruction of the oil workshop from the centre of the
ancient city of Ugarit. The installation comprised a press (< rectangular
archaeological stones are assigned if their allocation
stone slab and counterweight ), a paved mill, and a courtyard with probability is equal or superior to the discrimination level
three pithoi and a hearth (modified from Callot 1987, Fig. 7, p 202) of the group into which they have been put.

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Veget Hist Archaeobot (2014) 23:567-575 571

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 572 Veget Hist Archaeobot (2014) 23:567-575

Table 1 Allocation of archaeological olive stones from Ug

recognized at Ugarit in northwestern Mediterranean arch

Modern morphotypes Number of stones from Ugarit Earliest occurrence of modern morphotypes
allocated to a modern morphotype in Spain, France and Italya

p > 0.80 p > 0.65

Oli vière 0 0 Middle Ages (13- 15th century)
ORI 0 0-

OC-C 0 0 Chalcolithic/Bronze Age (2500-2000 b.c.e.)

OC-W 1 (0.5 %) 1 (0.5 %) Early Neolithic
OC-C+OC-W 2 (1 %) 5 (2.5 %)
Ecijano 0 0-
M1 70 (35.2 %) 124 (62.3 %)
MIOR 29 (14.6 %) 101 (50.7 %) Iron Age (end 3rd century b.c.e.)
OR2 0 1 (0.5 %) Roman Period (3rd-2nd century b.c.e.)
M2 3 (1.5 %) 7 (3.5 %)
M2a 2 (1 %) 5 (2.5 %) Roman Period (end 2nd century b.c.e.)
M2+OR2+M1 169 (85 %) 175 (88 %)
unclassified 28 (14.1 %) 19 (9.5 %)

Percentages presented in parentheses are calculated based on the total numbe

a After Terrai et al. (2004) and Terral (2007)

Results Depending on the level of assignment chosen (p = 0.65

or p = 0.80), the comparison of the morphological struc-
In modern material, the structure of the morphological
tures of the 199 archaeological stones to the reference model
shows that 19-28 samples of stones could not be identified
diversity of olive stones is observed in relation to their taxo-
(Table 1). The absence of modern analogues in the reference
nomie status and geographical origins (ESM 1). Olives from
collection or damage to the archaeological stones too small
spontaneous populations are distinguished from cultivars with
to be detected during preliminary scanning could have pre-
a discrimination rate of 62.5 %. Within spontaneous popula-
tions, western and eastern forms are discriminated with a
vented their association with an extant morphotype.
59 % rate. "Wild" types, originally of small size (length under Within the assigned archaeological stones, one is
assigned to OC-W, a group mostly constituted of wild
1 cm, although this criterion is not exclusive) and more sig-
western forms. At a lower aggregation level, two to five
nificantly, rather globular in shape, contrast with domesticated
stones are assigned to a group including OC-W and OC-C.
types which show a diversification of varieties which have
Eleven have a probability of allocation to OC-W between
distinct morphotypes, as shown by this study (Fig. 3): morpho-
0.50 and 0.65, and negligible (<0.15) to OC-C. We can
type ORI comprises 3 Egyptian varieties; morphotypes
Olivière and Bella di Spagna correspond to the names of the
therefore reasonably consider them as wild forms.
varieties, French and Italian, respectively; OC-C comprises aRegarding stones identified as cultivated forms, morpho-
group of varieties from the western Mediterranean; OC-Wtype
is MIOR, comprising mainly eastern varieties from
composed of western spontaneous populations and one
Greece, Turkey, Syria and Israel, is very well represented in
Syrian form. The Spanish variety Ecijano (syn. Lechin de
the archaeological material from Ugarit. One stone is assigned
to morphotype OR2 (p = 0.69) and two to five stones to
Sevilla ) constitutes a morphotype on its own. Morphotype
morphotype M2a. Finally, many specimens (169-175) are
Ml comprises varieties of diverse origins, including a sub-
group (MIOR), discriminated with a 74.2 % rate, composed
allocated to a super-group including morphotypes M2, OR2
of varieties mostly from the eastern Mediterranean and and
of Ml.
spontaneous forms (five eastern and two western, from
Tunisia and Italy). Morphotype OR2 is composed of varie-
Discussion
ties from Greece, Turkey and Syria. Morphotype M2 com-
prises, like Ml, varieties of diverse origins; it can be
partitioned in two sub-groups, one of which (M2a), dis- From a geographical standpoint, the present-day morpho-
logical differences between morphotypes, composed
criminated with a 74.7 % rate, is composed of eastern vari-
mostly of western (Olivière, OC-W, OC-C) or eastern
eties from Syria, Israel and Gaza, and three French varieties.

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 Veget Hist Archaeobot (2014) 23:567-575 573

(ORI, OR2, Ml OR) probability of allocation

types, seems (Table 1), to
appear to belong to
point to a
logical east-west western forms (OC-W and OC-C+OC-W).
differentiation, although Despite this at a h
of UPGMA aggregation two
affiliation discriminated
to western forms, the few Ugarit stones would grou
M2) comprise both eastern and
therefore represent local wildwestern
forms. Although they forms.
grow
graphical structure seems
on the marginsvalid
of cultivated both for
zones, wild forms are still cultivat
used
and spontaneous olive populations.
as rootstock today in traditional olive The
cultivation. status
Fruit of of
is as yet wild forms, as today, may have been pressed for oil
unclear, as feral forms, defined as unc
offspring of cultivated
production.individuals through sex
duction, may show phenotypic traits
Because it is morphologically close similar
to the wild morpho- to
wild olives (Zohary type
et OC-Wal. 2012).
and divergent from eastern Uncultivate
Mediterranean
(oleasters) therefore comprise both
varieties, morphotype OC-C may wild
represent forms which forms
evolved naturally, and feral
were independently individuals
domesticated in the western Medi- direc
from crosses between varieties or between wild forms and terranean. This morphotype was maintained over time,
cultivated varieties. In this way, the east-west structure ofeven if three of the constitutive varieties of OC-C are
wild olives as shown by morphometry (OC-W composed ofcharacterized by an oriental haplotype and thus possess at
western forms and Ml OR comprising eastern forms and least a maternal ancestry of Eastern origin (Besnard et al.
two central populations, from Tunisia and Italy) may be2013). Despite hybridization with an introduced Eastern
influenced by human-mediated movements of cultivated variety, repeated introgressions with local forms have
olives and hybridizations between trees of different origins.probably allowed this morphotype to remain unchanged. The
Genetic data based on maternally transmitted chloro- assignation to morphotype OC-C of archaeological olive
plastic DNA polymorphism (Besnard et al. 2007) validated stones from Spanish sites dating from the Chalcolithic to the
by studies on nuclear DNA (Besnard et al. 2001a, b; BretonBronze Age confirms this hypothesis, suggesting the exis-
et al. 2006) confirm the structure of diversity shown by tence of a primary olive domestication event in the west.
geometric morphometry. A unique maternal lineage was This morphotype has not been found in the studied Ugarit
detected in eastern Mediterranean populations, but three sample.
maternal lineages were found in western populations, and Morphotype Ml OR, composed mostly of varieties of
phylogeographic data indicate that one of them, namely El,eastern origin, represents 14.6-50.7 % of the assigned
was probably recently introduced by humans from East to olive stones from Ugarit, depending on the probability of
West (Besnard et al. 2001b, 2002, 2007, 2011, 2013; allocation. This morphotype also defines the most ancient
Khadari et al. 2008). This geographical structure suggests evidence of an eastern cultivated form in the western
that wild olive populations evolved before domestication, Mediterranean, dated to the end of the 3rd century b.c.e.,
thereby confirming palaeoecological (Terral and Arnold-from La Seña, Spain (Pérez Jorda 2000; Terral et al. 2004).
Simard 1996) and archaeobotanical conclusions (Terral Given the geographical origin of varieties that constitute
2000; Terral et al. 2004). this morphotype (essentially the Levant, Turkey and
The east-west structure of the morphological diversity Greece, but also Tunisia and Spain), this archaeological
observed in the wild specimens (59 % discrimination rate data can be interpreted as showing the introduction of
between eastern and western spontaneous forms) could in cultivated forms from a Near Eastern and Aegean gene
fact be linked to ferality, which would explain that Ml OR pool, probably under Phoenician and Greek influence. One
comprises spontaneous forms and eastern Mediterranean century later, the western Mediterranean olive agrobiodi-
varieties. The structure could therefore be not only theversity was enriched of two new morphotypes, OR2 and
result of an east-west differentiation which happenedM2a found at L' Almadrava, Spain (Terrai et al. 2004). The
before domestication, but also an expression of the close geographical distribution of their constitutive varieties is
relationship between wild populations and local agrobi-Mediterranean-wide, as is the area of the diffusion of olive
odiversity, thereby explaining the discrimination rate of cultivation in antiquity, caused in part by the extent of the
only 62.5 % between spontaneous and cultivated forms. AsRoman Empire. In this case, the morphological affinity
in the cases of the grapevine (Terrai et al. 2010; Picq 2012)between domesticated forms of diverse origins can be
and the date palm (Terrai et al. 2012), spontaneous indi-explained as the consequence of human migrations that
viduals, regardless of origin, show a morphological con-have encouraged crosses between western and eastern
vergence trend (such as OC-W and Ml OR) as a result of forms through time. The Moroccan Picholine (morphotype
similar selection pressures essentially related to environ- Ml) is a variety that fits this history, as described by
mental constraints. Khadari et al. (2008) on the basis of genetic markers.
Within the archaeological material from the Ugarit Indeed, its chloroplastic DNA haplotype indicates an
kingdom however, one to five stones, depending on eastern Mediterranean origin, whereas nuclear markers

â Springer

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 574 Veget Hist Archaeobot (2014) 23:567-575

suggest an admixed Besnard G, Baradatbetween

origin P, Breton C, Khadari B, eastern
Bervillé A (2001b) Olive
and
domestication from structure of oleasters and cultivars using
gene pools (Besnard et al. 2001b; Khadari et al. 200
nuclear RAPDs and mitochondrial RFLPs. Genet Sel Evol
Moroccan Picholine , which
33:S251-S268 is important in olive
duction in Morocco today,
Besnard G, is therefore
Khadari B, Baradat P, Berville a descendan
A (2002) Olea europaea
eastern varietal phylogeography based
pool, but its maternal ancestor on chloroplast DNA polymorphism. w
Theor Appi Genet 104:1,353-1,361
bly introduced into Morocco and repeatedly intro
Besnard G, Henry P, Wille L, Cooke D, Chapuis E (2007) On the
by local forms (Khadariorigin
etofal. 2008).
the invasive olives ( Olea europaea L., Oleaceae).
From a practical point of
Heredity view, these results
99:608-619
single batch of olive Besnard
stones G, Hernandez P, Khadari
also B, Dorado G, Savolainen
show thatV (201 1)
oliv
Genomic profiling of plastid DNA variation in the Mediterranean
produced using a mixture of olives collected from
olive tree. BMC Plant Biol 11:80
trees belonging to different
Besnard G, Khadaritypes, including
B, Navascués M, Fernández-Mazuecos M, El both
and cultivated
This may
Bakkali A, Arrigo N,mean ones.
Baali-Cherif D, that the
Brunini-Bronzini de oliv
Caraffa V, Santoni S, Vargas
collected both from tended olive groves and f P, Savolainen V (2013) The
complex history of the olive tree: from Late Quaternary
surrounding spontaneous vegetation where unc
diversification of Mediterranean lineages to primary domestica-
olive trees grew, whether
tion in theferal or
northern Levant. wild.
Proc Royal It under
Soc B 280:20122833.
role of wild olive populations in the
doi:10.1098/rspb.2012.2833 plant econom
Late Bronze Age as Blondel as
well J, Aronson J (1995) Biodiversity
today, evenand ecosystem
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a incontex
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agriculture is classically considered prominent.
Mediterranean-type ecosystems. In: Davis GW, Richardson DM
(eds) The function of biodiversity. Springer, Berlin, pp 43-119
Breton C, Tersac M, Bervillé A (2006) Genetic diversity and gene
Conclusion
flow between the wild olive (oleaster, Olea europaea L.) and the
olive: several Plio-Pleistocene refuge zones in the Mediterranean
The comparison of archaeological olive stones from Ugarit Basin suggested by simple sequence repeats analysis. J Biogeogr
33:1,916-1,928
with a reference model of morphological differentiation
Breton C, Terral JF, Pinatel C, Médail F, Bonhomme F, Bervillé A
based on modern material allows us to reinterpret the east-
(2009) The origins of the domestication of the olive tree. C R
west structure shown in wild olive populations by Terral Biol 332(12): 1,059-1,064
Burger P, Terral JF, Ruas MP, Ivorra S, Picq S (201 1) Assessing past
et al. (2004). In agreement with genetic data, it appears that
the division of oleasters into two geographical types in the agrobiodiversity of Prunus avium L. (Rosaceae) and related
cherry species: focus on the stones from the Hôtel-Dieu
Mediterranean could reflect, not only an ancient differen-
archaeological site (16th c., Tours, France) using morphometries.
tiation, but also a subsequent development resulting from Veget Hist Archaeobot 20:447-458
hybridizations between wild populations on the one hand,
Callot O (1987) Les huileries du Bronze Récent à Ougarit , premiers
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in the western archaeological record 1,000-1,500 years domestique. RSO X, ERC, Paris
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