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Allozyme Variation of Oleaster Populations (Wild Olive Tree) (Olea Europaea L.) in the Mediterranean Basin

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Allozyme Variation of Oleaster Populations (Wild Olive Tree) (Olea Europaea L.) in the Mediterranean Basin Heredity (2004) 92, 343–351 & 2004 Nature Publishing Group All rights reserved 0018-067X/04 $25.00 www.nature.com/hdy Allozyme variation of oleaster populations (wild olive tree) (Olea europaea L.) in the Mediterranean Basin R Lumaret1, N Ouazzani2, H Michaud1, G Vivier1, M-F Deguilloux1 and F Di Giusto1 1Centre d’Ecologie Fonctionnelle et Evolutive CEFE/CNRS (U.P.R. 9056), 1919 Route de Mende, F-34293 Montpellier Cedex 05, France; 2Ecole Nationale d’Agriculture, De´partement d’Arboriculture, B.P. S40, Mekne`s, Morocco As a result of the early domestication and extensive collected in forests potentially containing genuinely wild cultivation of the olive tree throughout the Mediterranean forms according to environmental, historical and demo- Basin, the wild-looking forms of olive (oleasters) presently graphic criteria. As reported previously from cytoplasmic observed constitute a complex, potentially ranging from wild and RAPDs analysis, substantial genetic differentiation was to feral forms. Allozyme variation was analysed at 10 loci in observed between the eastern oleaster populations geneti- 31 large and 44 small oleaster populations distributed in cally close to most olive clones cultivated in the Mediterra- various habitats of the Mediterranean Basin and in two nean Basin, and the western populations that are related to populations of the wild subspecies Olea europaea subsp the wild Canarian populations. In addition, the occurrence of (ssp) guanchica, endemic to the Canary islands and closely significantly lower heterozygosity in cultivated olive than in related to oleasters. At eight polymorphic loci, 25 alleles were oleasters, whatever their origin, suggests that intensive identified. Genetic evidence that nondomesticated oleasters selection involving inbreeding has taken place under cultiva- still survive locally was provided by the occurrence of four tion to obtain particular characteristics in the olive cultivars. and one alleles shared exclusively by the eight western and Heredity (2004) 92, 343–351, advance online publication, 25 two eastern oleaster populations, respectively, which were February 2004; doi:10.1038/sj.hdy.6800430 Keywords: allozyme variation; Olea europaea L. ssp Sylvestris; oleaster populations; olive domestication Introduction planted directly or, more recently, grafted onto indigen- ous oleasters. From these propagated individuals, olive The olive tree (Olea europaea L.) is one of the most ancient cultivars were developed and distributed by various domestic, cultivated plants characteristic of the Mediter- successive human migrations throughout the Mediterra- ranean Basin (Zohary and Spiegel-Roy, 1975). The crop nean Basin, especially from East to West although the (O. europaea subsp (ssp) europaea ( ¼ O. europaea var. selection of olive cultivars was multilocal and included sativa Lehr.)) has been of immense importance as the originally western plant material, as shown by genetic principal source of edible oil for the peoples of the studies based on allozyme polymorphism (Lumaret et al, Mediterranean region for many millennia. Palynological 1997), on RAPD profiles and on mitochondrial RFLPs and anthracological (fossil charcoal) evidence supports (Besnard and Berville´, 2000; Claros et al, 2000; Besnard the occurrence of wild olive forms (O. europaea sssp et al, 2001a, b; Bronzini de Caraffa et al, 2002a) as well as sylvestris (Miller) Hegi ( ¼ var. oleaster (Hoffmanns and inter-simple sequence repeat (ISSR) markers (Vargas and Link) DC), in forests of the Mediterranean Basin during Kadereit, 2001). Deforestation probably reduced native the Paleolithic (Liphschitz et al, 1991). According to oleaster populations and the geographic dissemination anthracological studies, during the Neolithic (10 000– of cultivars favoured multiple contact areas and hybri- 7000 years ago), use of wild olive (oleasters) persisted in disation between cultivated olive and indigenous olea- several parts of the Basin (Liphschitz et al, 1991; Zohary sters that are fully interfertile. This may have led to the and Hopf, 1993). Clear signs of olive domestication (olive progressive disappearance of the original genetic char- oil presses) come from the Near East during the Early acteristics of the wild oleasters (Zohary and Spiegel-Roy, Bronze Age (second half of the fifth millennium BP) 1975). (Zohary and Spiegel-Roy, 1975; Liphschitz et al, 1991). It Throughout the Mediterranean Basin, oleaster olives is currently assumed that individual oleaster trees differ from the cultivated clones by the presence of showing superior performance for size and/or oil spinescent juvenile shoots, smaller fruits characterised content of fruit were selected empirically and propa- by less fleshy mesocarp and lower oil content, as well as gated vegetatively as clones, using cuttings that were by a long juvenile stage that may last for several decades in some individuals. As reported previously (Zohary and Spiegel-Roy, 1975), whatever their parental origin (wild Correspondence: R Lumaret, Centre d’Ecologie Fonctionnelle et Evolutive, CEFE 1919, Route de Mende, F-34293 Montpellier Cedex 05, France. or cultivated) any olive progeny grown from seed E mail: [email protected] usually shows the morphological characters specific to Received 18 February 2003; accepted 12 December 2003 oleasters. These therefore constitute a complex of Allozyme variation in wild olive populations R Lumaret et al 344 wild-looking olives potentially ranging from wild to feral of the same species, which reproduce sexually and forms (ie secondary sexual derivatives of the cultivated vegetatively, respectively. clones or products of hybridisation between cultivated trees and nearby oleasters) (Zohary and Spiegel-Roy 1975; Zohary and Hopf, 1993). Materials and methods During the last five decades, controversy has devel- oped as to whether the oleasters observed at present Tree sampling have originated exclusively by naturalisation from At least 20 trees (38.25 individuals on average) were cultivated plants (Chevalier, 1948) or whether genuinely sampled in 31 large oleaster populations distributed over wild native forms still survive in some forests of the the Mediterranean Basin (Figure 1 and Table 1). Two Mediterranean Basin. In this latter case, only the oleasters additional large populations (nos. 32 and 33) of the wild observed in secondary habitats (the edges of cultivated subspecies, O. europaea ssp guanchica P. Vargas, J. Hess, or abandoned orchards) would be feral (Zohary and Mun˜os Garm and Kadereit (O. cerasiformis (Webb and Spiegel-Roy, 1975; Zohary and Hopf, 1993). In Australia, Berth) Kundel and Sunding), endemic to the Canary oleasters derived from olive cultivars introduced 200 Islands, were sampled in two sites on Grand Canary years ago, and which are bird-dispersed plants, have Island (Figure 1). There is no evidence that this proved to be successful invaders in numerous biotopes, subspecies has been involved with olive domestication including forests (Spennemann and Allen, 2000). More- (Zohary, 1994), but it was found to be very closely related over, knowledge about the dissemination distance of to Mediterranean oleasters (O. europaea ssp sylvestris)on olive pollen by wind (Griggs et al, 1975), from recent the basis of chloroplast DNA variation and RAPD and studies of seed dispersal of wild olive by birds in Spain ISSR markers (Hess et al, 2000; Lumaret et al, 2000; (eg Alcantara et al, 2000), and from local reports about Me´dail et al, 2001). the extensive use of olive stones from olive groves for Of the large populations, 10 were sampled in forest forestation in several Mediterranean countries suggest areas considered to be good candidates to contain that genuinely wild olive populations may be restricted genuinely wild oleasters according to three criteria: (i) to very few isolated forest areas. past and present climatic conditions suitable for wild In several previous studies, the nuclear genetic olive growth (humid and subhumid variants of the variation of oleasters was analysed over the Mediterra- thermo-Mediterranean climate with an average mini- nean Basin using RAPD markers (Besnard et al, 2001a, b) mum temperature of the coldest month and annual and, in more restricted areas, using RAPDs (Bronzini de rainfall exceeding 51C and 450 mm, respectively (Daget, Caraffa et al, 2002a, b), AFLPs (Angiolillo et al, 1999) and 1977; Rivas-Martinez, 1981; Terral and Arnold-Simard, allozyme polymorphism (Lumaret et al, 1997; Lumaret 1996); (ii) past and present isolation (usually more than and Ouazzani, 2001). When wide areas were sampled, a 10 km) from areas of cultivated olive trees; and (iii) large clear nuclear genetic differentiation was observed be- oleaster populations (several thousands of trees) to tween oleasters from the eastern and western parts of the reduce the influence of occasional pollen and seed flow Mediterranean Basin. In the present work, we analysed from cultivated olive areas. In the eastern part of the in detail the genetic variation of numerous oleaster Mediterranean Basin, where olive trees have been populations distributed over the Mediterranean Basin. extensively cultivated for long periods, we used less These populations were found to grow in various habitat restrictive criteria to identify candidate areas, more conditions, such as the edge of olive groves, abandoned particularly for population size and isolation distance groves and noncultivated areas including forests. More- from cultivated olive areas. Candidate areas were not
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