The classification of olive germplasm – A review

Review Article

The classification of olive germplasm – A review

By T. GANINO1*, G. BARTOLINI2 and A. FABBRI1

1Dipartimento di Biologia Evolutiva e Funzionale, Università degli Studi di Parma, Parco Area delleScienze, 11/A - 43100 Parma, Italy 2Istituto per la Valorizzazione del Legno e delle Specie Arboree, CNR-IVALSA, Via Madonna delPiano - 50019 Sesto Fiorentino (FI), Italy(e-mail: [email protected]) (Accepted 6 February 2006)

SUMMARYOlive cultivar classification has been a primary concern for olive growers since the earliest times of olive cultivation.As one of the earliest fruit crops to be domesticated, the number of olive genotypes cultivated in the Mediterraneanregion and elsewhere is high, and the assessment of an easy and undisputed system of classification has becomeincreasingly urgent. This is also because of the high number of synonyms and homonyms in existence. The presentreview examines the history of attempts, in all olive-producing countries and by many researchers, to achieve anultimate and exhaustive descriptor list that is valid for all past, present and future genotypes, to support technicians,growers and scholars. After a thorough review of the numerous elaiographical lists proposed, the modern researchtools that have been applied to olive (i.e., ultrastructural, biochemical, DNA and molecular markers) are examinedcritically. A balance sheet has been prepared, taking into account initiatives by major international and nationalinstitutions.

In Roman times, the need to identify and classifycultivated olive plants present in each area was

already clear. Agronomists of the time summarilydescribed all those varieties they could distinguish whichhad interesting agronomic features. Descriptions exist byCato, Varro, Virgil and other scholars. However it isdifficult to find any correspondence between thosevarieties and varieties cultivated today. Many of theformer are likely to have disappeared due to abiotic andbiotic environmental stresses (e.g., frost, drought,parasites), or may have altered due to outcrossing,mutation and/or changes in selection pressure, includinggrower requirements.

Over time, a large number of olive varieties have beenidentified by numerous scholars who described them bytaking into consideration their most varied characters,with remarkable descriptive meticulousness.This led to alarge amount of data as the description often concernedan individual tree, irrespective of whether or not itbelonged to a known cultivar. The result was “to confusethe mind of even the most intelligent agronomists and ofthe sharpest growers” (Caruso, 1883).

One of the main difficulties was understanding thenames of all these varieties, often described in localdialect or vernacular. Another problem was to relateresults from different regions. The first result was anelevated homonymy (i.e., when different olive cultivarshave the same name in different zones) and synonymy(i.e., a given variety is named in one area, and recordedwith another name in a different area) within a givenolive district and between different districts.

The same problem exists for scientists and growerstoday. Determination of characters that can be observedand expressed in all environments (i.e., varietalidentification) is the first objective in order to: a) classifyand describe the existing varietal heritage; b) describeand isolate all genotypes with valuable characteristics;and c) overcome existing confusion concerning varietalnames.

Hence the importance of distinguishing and identifyingolive genotypes. Knowledge of the morphological,ecological and agronomical characteristics of all availablegermplasm is indispensable for selection of the mostpromising individuals, best-suited to modern cultivationtechniques. At the same time, commercial requirementscan be satisfied, particularly those of nurserymen whoneed to know the main characters of a genotype to beintroduced onto the market, to be able to certify it and, attimes, patent it.

TAXONOMIC ENTITIES THAT ARE THE OBJECTOF CLASSIFICATION

The smallest taxonomic entity that can usually bedefined with distinction is universally termed a “variety”.In botany, “variety” implies some morphologicalcriterion of differentiation, while among agriculturalvarieties we may have identical morphologies alongsidemarked distinctions in other characters such as bearingprecocity, productivity, or resistance to biotic and/orabiotic factors (Trehane et al., 1995).

The term “cultivar” (cultivated variety) is alsocommonly used in agricultural terminology. “Cultivar” isdefined by the International Code of Nomenclature for*Author for correspondence.

Journal of Horticultural Science & Biotechnology (2006) 81 (3) 319–334

History of cultivated olive classification and description320

Cultivated Plants as “an ensemble of cultivated plants,clearly distinguished by any character (morphological,physiological, cytological, chemical, etc) which, whenreproduced (sexually or asexually), preserves itsdistinctive features” (Trehane et al., 1995).A cultivar mayconsist of a single clone, or a number of very similarclones (polyclonal variety), where clone is meant as “anensemble of genetically uniform individuals, derivedoriginally from a single individual by asexualpropagation, i.e., by cutting, division, grafting or obligateapomixis” (Roselli and Scaramuzzi, 1974).

Morphological and biological characters may notalways appear constant in all areas and for all clones, dueto the influence of the particular pedoclimatic conditionsin which they may be cultivated. It is therefore necessaryto introduce the concept of “ecotype”; that is, thepeculiar biological, morphological and/or agronomicappearance that cultivars and clones may assume indifferent environments, usually over a limitedgeographical range (Lorenzetti et al., 1996).

In perennial plants, the concept of a variety is oftenundetermined, as varieties are often grouped withreference to a number of common characters, allowing adegree of variation for other phenological features.However, it is precisely in perennial plants, includinggrapevines, olives and the most important fruit crops,that we have devised an empirical selective method,vegetative propagation. This method permits thepreservation of genetic inheritance in the progeny,unchanged and easily usable. The most ancientdomesticatied fruit crops in the Mediterranean Basinhave therefore been those that could be easilypropagated from plant parts (e.g., fig, date palm, olive,pomegranate, grape).

The number of olive cultivars described has beenincreasing since ancient times due to progress intaxonomy and plant breeding techniques.

To comply with the need for a practical classificationof cultivated plants, Marinucci (1908) suggestedsubstitution of the term “variety” with “agrarian race”.For olive, this term has not been further explained oradopted. For Marinucci (1908) it applied to groups ofindividuals, agamically propagated over severalgenerations, and therefore each group should be termed“a clone” (Baldini and Scaramuzzi, 1952), or a set ofclones.

The term “race” was eventually replaced by “cultivar”,although perhaps too quickly. In fact, all known cultivarsshould still be considered as wide and heterogeneouspopulations of clones, separated by a number ofcharacters.

CHRONOLOGY OF THE PROPOSEDCLASSIFICATION SYSTEMS

We owe the first olive classification system to thebotanist Pitton de Tournefort (1719). He was soonfollowed by numerous other botanists, who suggested oradapted several systems to describe large number ofolive varieties, but they never achieved an accurate orpractical classification (Prevost and Mostardini, 1999;Table I).

In Spain, the first to attempt an olive classification in1815 was Simòn de Rojas Clemente (Barranco et al.,1984), based on features of leaves and fruits in a numberof varieties.

The proposal to take the olive stone into considerationas a discriminating character in the classification ofItalian germplasm was made by Tavanti (1819). He wasconvinced that endocarp characters had fundamentalimportance in the study he was to undertake on olive“races”. The stone was considered both in its wholeshape, and in its separate and minute aspects: the base,the apex, the valves and the course of the suture lines ofthe valves that form the stone. Tavanti (1819) based thedistinction he made of Tuscan cultivated olive varietieson these five characters, and accordingly divided theminto 21 groups.

Analogous classification systems were eventuallyproposed by Marinucci (1908), who reduced the numberof groups listed by Tavanti to five, by Zito (1932), byFrezzotti (1937) and by Savastano (1939). But endocarpcharacters were no longer considered sufficient, andother characters (morphological, biological andecological) were added, in order to satisfy technicalrequirements and to take into account the influence ofenvironment. With this in mind Ciferri and Breviglieri(1942) made the first attempt at a morpho-ecologicalclassification by enforcing the criteria adopted by theRussian school of Vavilov. In all papers of the time, theuse of morphological characters was aimed at achievinga sub-division of cultivars according to morphological

TABLE IComparison of different classification systems in the history of olive identification from 1719 to the present-day in various olive-producing countries

Year Authors Country Discriminant characters

1719 Pitton de Tournefort France Botanic characters1815 Simón de Rojas Clemente Spain Leaf and fruit1819 Tavanti Italy Endocarp1908 Marinucci Italy Endocarp1917 Ruby France Leaf, fruit and endocarp1932 Miliani Italy Ecological characters1932 Zito Italy Endocarp1937 Bracci Italy Leaf, inflorescence, fruit, endocarp and ecological characters1937 Frezzotti Italy Endocarp1939 Savastano Italy Endocarp1942 Ciferri et al. Italy Morphological, biological and agronomic characters1954 Patac Spain Morphological, botanical and agronomic characters1984 Barranco and Rallo Spain Morphological and agronomic characters1985 UPOV World Morphological and agronomic characters1986 Leitão et al. Portugal Morphological and agronomic characters1998 Bartolini et al. World Passport, agronomic characters, biochemical and molecular markers, collection, etc.2000 Barranco et al. World Passport and agronomic characters2000 Pannelli et al. Italy Morphological, agronomic and commercial characters2005 Rallo et al. Spain Passport, morphological, agronomic and commercial characters

T. GANINO, G. BARTOLINI and A. FABBRI 321

variations in fruits, leaves, inflorescences, endocarp andother organs. Progress lay in increased numbers ofcharacters taken into account.

In 1917, in France, Ruby proposed a classificationsystem in which three characters were considered: fruits,stones and leaves. With these, and with the ratiosbetween maximum transverse and longitudinal lengths,data were obtained which were typical of groups ofplants, and which enabled differences between varietiesexisting in the territory to be distinguished andevaluated.

Another tentative classification, this time based onecological grounds, was that of Miliani (1932), whodistinguished some Italian olive races with reference totheir resistance to low temperatures and arid conditions,to hardiness, to productivity and to the quality of theproduct.

According to the considerations of Ciferri andBreviglieri (1942), and because of the inadequacy andlimited usefulness of schemes suggested in previous years,Ciferri and other researchers (Ciferri et al., 1942)elaborated more acceptable solutions. In the classificationthen put forward, numerous morphological andbiological characters were considered, as well as criteriaadopted to evaluate the importance of each character.

Based on these elements, a very detailed elaiographic(from ancient Greek “elaion”, pertaining to olive oil)descriptor list was prepared, and used for data collectionon approx. 70 cultivars grown in Central Italy (Tuscany,Latium, Marches, Umbria). Where data collection wascarried out with care, in a circumscribed environmentand in such conditions as to exclude environmentalvariables, this list allowed varietal identification with anacceptable degree of reliability. The introduction ofagronomic and morphological-ecological characters hadtherefore become necessary, as had been proposed byCaruso in his olive monograph (1883). Caruso hadstressed the fact that “an ordering of races must be clearand able to put anyone in the condition of being able toappreciate and choose the olive trees according to theiragrarian prerequisites” (Morettini, 1972). Zonallimitations, however, do not allow the identification ofsynonyms (i.e., the various names by which the samecultivar is identified in different environments) which isprecisely one of the main tasks of any classification.

However, a study in 1942 by Ciferri et al. is stillconsidered today when studying new classificationsystems. Ciferri et al. (1942) acknowledged that theirsystem had some limitations, largely because olive is avery homogeneous species, and consequently sub-division of Italian proles into sub-proles is difficult. Inaddition, the values considered were so variable, due todifferent cultivation techniques and environmentalconditions, that their exact definition is impossible,including the classification of the same cultivars intoproles and sub-proles (Zito, 1942).

Further discussion was on the terminology adopted.The word “average”, as used to describe characters, has adifferent meaning for statisticians, defines nothing forthe practical technician (grower), and is never sufficientto define a given character. Despite criticisms raised atthe time of its publication, the 1942 Descriptor Listeventually became the starting point and a fundamentalreference for all researchers in the field.

Towards the end of the 1940’s, Patac propounded asystem for the identification of cultivars existing in Spain(Patac et al., 1954). The assumption that morphologicalcharacters were highly influenced by environmentalconditions and by human activities led Patac to consideras important, all characters that may give informationconcerning olive transformation, without omitting thedescription of characters deemed to be more strictlybotanical. This innovation consisted of introducing ahierarchy of importance among the characters used. Indecreasing order of importance, therefore, were theshape and size of the endocarp, the shape and size offruits, features of the tree, shoots, inflorescences andseed, and organoleptic characters of the oil. For eachcharacter, coefficients and biometric indices weredeveloped and included. Thus, the descriptor aimed tosatisfy classification, agronomic and industrial needs.Thesystem, and the problems related to it, were discussed atthe International Olive Congress in Seville, in 1950(Patac et al., 1954).

Over the next 30 years, no major innovations wereintroduced into olive classification. The innovativeproposal of Patac’s descriptor was not well-received, andthe reference document remained the pomological listsuggested by Ciferri et al. (1942).

The need to understand the complexity of olivegermplasm was particularly strong in countries in whichthe olive industry was most developed. In fact, in the1980’s interesting proposals were made in Spain,Portugal and by UPOV (Union Internationale pour laProtection des Obtentions Végétales) in Geneva. In1984, Barranco et al. put forward pomological lists inwhich the description of the tree and of its fruitingbranches, leaves, inflorescences, fruits and endocarpswere considered fundamental characters, to catalogueand identify all Andalusian olive cultivars. Thisdescriptive scheme, though, was confined by taking intoaccount that of Ciferri et al. (1942), with a few changesintroduced by Bottari and Spina (1953) and by theauthors. The changes introduced included using thelargest possible amount of information, eliminatingcharacters that were most difficult to obtain and thosemost influenced by the environment, as well as othersconsidered irrelevant and/or subjective. In addition, foreach character, Barranco et al. (1984) tried to use as fewcategories as possible, to separate cultivars withoutcreating too many classes. To make interpretation andreading easier, each file was accompanied byphotographs of those organs important for thedescription.

In 1985, UPOV proposed a specific descriptor list forolive. In that list, whole plant characters, fruiting shoots,leaves, inflorescences, flowers, fruits and endocarp weretaken into consideration. The aim was to standardise themethodology of data collection, and to indicate a “leaderlist” usable by all researchers who seek to describe olivecultivars. This initiative represented the second mostsignificant attempt, after Ciferri et al. (1942), atclassifying all olive germplasm existing in variouscountries with an homogeneous method.

In 1986, Leitâo described 22 Portuguese olivecultivars. For each cultivar, tree, shoot, leaves, flowersand inflorescences, endocarp and fruit characters wereconsidered, together with agronomic characters. For the


author, this technique had the merit of allowing an easyand fast plant description, without resorting tosophisticated and difficult techniques. The aim was toprovide a summary description of the plant, whilefocussing interest on some characters (leaf, inflorescence,fruit and endocarp) that were considered interesting forthe identification and characterisation of cultivars.Leitâo also took general and agronomic aspects, such asrooting ability, disease resistances and susceptibilitiesinto account. The whole description of each cultivar wasaccompanied by photographs of plant organs.

In 1993, in Tuscany, the problem of describing andidentifying olive germplasm was again on the agenda,this time involving plants kept in collections in theProvinces of Pistoia, Grosseto and Firenze. Thedescriptor list proposed by Cimato et al. (1993) includeddescriptions of the tree, shoots, leaves, inflorescences,fruits and endocarp. For a description of the differentcharacters, reference was made to the UPOV descriptor.By applying a few changes, Cimato et al. (1993) deemedit indispensable to reduce environmental effects on thephenotypic expression of characters. Such changes werenot clear, and no adequate explanation was given. In thesame year, an elaiographic descriptor was produced forolive germplasm in Catalonia (Tous Martì and RomeroAroca, 1993). No major methodological innovationswere adopted.

Cristoferi et al. (1997) propounded a list for theidentification and classification of Romagna olivecultivars. For each cultivar, the list contained informationon its diffusion, synonyms, elaiographic characters (e.g.,tree, leaf, inflorescence, fruit, endocarp, etc.), biologicaland agronomic characters.

In the following year, Bartolini and collaborators(1998) advocated the use of a simplified pomologicaldescriptor for the World’s olive germplasm. Such a listhad plant passport data, tree data and agronomiccharacters (e.g., productivity, oil yield, tolerance tobiotic and abiotic stresses, etc.). For the first time,important information was added such as the existenceof patents, any presence in germplasm collections, andwhere, and data concerning biochemical and molecularcharacterisations.

Soon a scheme was elaborated for the identificationof Marches olive cultivars (Pannelli et al., 1998), whichwas also eventually used for Umbrian cultivars(Pannelli et al., 2000). The first scheme was generatedby describing the tree, shoots, leaves, inflorescences,fruits and agronomic behaviour. For the description ofmorphological characters, reference was made to theterminology adopted by UPOV, with a few minorchanges concerning the elimination of some charactersthat were hard to detect, and the addition ormodification of others. As regards agronomiccharacters, for each cultivar judgments were expressedaccording to results obtained over several years and indifferent situations. The work was thus simplified, asonly essential characters were used. Practicalconsiderations were also expressed on harvest time,taking into account oil accumulation in the drupes, andjudgments on oil quality characteristics and plantproductivity; information was also given on thesuitability of fruits for milling and extraction with apressure system. For the first time, technical

information was introduced for the producer, relatingto the milling and extraction process.

A “World Catalogue of Olive Varieties” (Barrancoet al., 2000a) has recently been published by TheInternational Olive Oil Council (IOOC), with aclassification system based on descriptor lists which arevalid for all olive producing areas. Once more, the need touse a unified and valid classification system was stressed,to guarantee the conservation of olive germplasm.The fileconsists of a list of descriptive characters, accompanied bya synthetic bio-agronomic evaluation. The information isdivided into three groups: plant passport data,morphological characters, and agronomic and commercialconsiderations. The first group contains the most commonname of the cultivar, its synonyms, its country of origin[i.e., the country of most probable cultivar provenance, orin which it has reached the largest distribution, usingInternational Standards Organisation (ISO) codes], themain growing areas, the importance of the cultivar in theseareas, and the purpose(s) of production.The morphologicalcharacters are structured as primary descriptors used forthe identification and characterisation of each cultivar.Thecharacters have been structured according to qualitative(shapes, expression of morphological features) andquantitative (biometric indices) descriptors. Theagronomic and commercial considerations contain datathat are useful for growers, scientists and techniciansinterested in the olive industry.

The IOOC proposal, which was the result of a jointstudy by experts and scientists throughout the oliveworld, is the latest attempt to unify the methodologies ofolive classification. In recent years, others have suggestedother descriptor lists, starting from existing ones. AgainCimato et al. (2001) published elaiographic lists of Tuscanolive germplasm, relying on former work elaborated bythe same authors in 1993. Several other elaiographic listshave been elaborated recently (Cicoria et al., 2000;Pugliano et al., 2000; Rotundo and Marone, 2002;Trigui etal., 2002; Parlati and Pandolfi, 2003; Lombardo et al.,2003; 2004; Bassi, 2003). That of Rotondi et al. (2004)appears to be quite interesting. Besides the charactersformerly listed in previous work (Cristoferi et al., 1997),detailed information is added on the organoleptic andchemical properties of the oil and, for the first time, onthe phytosanitary situation of each cultivar. Amonograph by Rallo et al. (2005) has very recently beenpublished in which the elaiographic lists of all Spanishcultivars are collected together (including plant passport,morphological, agronomic and commercial characters).The starting point for this descriptor was that ofBarranco et al. (2000a) which also contained informationon plant passport, tree, leaf, fruit, endocarp andagronomic features.

The distinctive features of the most recentelaiographic lists undoubtedly lie in the introduction ofmore refined methods of study such as molecularmarkers and the chemical and organoleptic analyses ofoil. Varietal identification by such methods is madedifficult by the nature of the species, which is oftencharacterised by the presence of cultivar-populations orpopulations of clones, rather than by clear cultivars(Bartolini et al., 1992; 1994a).

Thanks to statistical analyses of the variability of theleast variable individual characters, most suitable

https://www.researchgate.net/publication/244189341_Polymorphism_and_Discrimination_Capacity_of_Randomly_Amplified_Polymorphic_Markers_in_an_Olive_Germplasm_Bank?el=1_x_8&enrichId=rgreq-ccca519b-d3ee-4ff0-871b-6d56824652f1&enrichSource=Y292ZXJQYWdlOzIzNDExNTgyMjtBUzoxMDIxNDAzMzI3NDA2MTFAMTQwMTM2MzU2MTkwNg==


characters were identified: leaves, drupes, stones (Baldiniand Scaramuzzi, 1952; 1955; Damigella, 1960). Morerecently, several methods of multivariate statisticalanalysis have shown the existence of numerousbiometric variables related to the fruit and to the plant,or to the acid composition of the oil, which have asatisfactory discriminating power (Barone et al., 1994a,b;Perri et al., 1995; Cantini et al., 1999; Barranco et al.,2000a). Notwithstanding, morphological markers remainpoorly discriminating for varietal identification, as theyare characterised by high environmental variability.Their study must therefore be integrated by recentultrastructural, artificial neural networks (ANN),biochemical and molecular genetic techniques. Thesenew research tools represent a turning point to resolveproblems in olive taxonomy.

Ultrastructural analyses: Some authors (Quiros, 1975;Maas, 1977) proposed pollen morphology (section,shape, etc.) as a useful tool for varietal identification.Pollen features, although part of the morphologicalcharacters, are considered intermediate between fullymorphological characters and biochemical ones(Petruccelli, 1992; Bartolini et al., 1994b). In particular,exine structure (i.e., the outer coating of the cell wallsurrounding the pollen grain at maturity) is considered,as its characteristics (grain diameter, net mesh size andshape, etc.) are under the control of the sporophyte(pollen mother cells; Pandey and Troughton, 1974). Wallstructure is therefore a more stable character comparedto other morphological characters, as it is specificallygenetic, with little or no influence from the environment.Roselli (1979) was the first to use pollen ultrastructuralanalysis for olive cultivar identification. He studied 13table olive cultivars; however, the research did notproduce sufficient information for the identification ofcultivated varieties. In 1995, Lanza et al., used scanningelectron microscopy to study the morphology of exinepatterns on pollen grains of four olive cultivars. The dataobtained allowed complete differentiation of the fourgenotypes (Lanza et al., 1995). These analyses, however,give little information for the identification of theindividual cultivar (Bartolini and Petruccelli, 1994).Further studies on several other anatomical andultrastructural aspects have also not yielded encouragingresults.

ANN: Interesting results were obtained by applying thetechnique of backpropagation neural networks (BPNN)to ten olive cultivars. Sets of phyllometric parameterswere used for cultivar identification and the results,albeit preliminary, could be considered encouraging(Mancuso and Nicese, 1999).

Biochemical markers: Total and storage proteins,isozymes and molecular markers (RFLPs, RAPDs,AFLPs, micro- and mini-satellites sequences) permit theelimination of two negative aspects of phenotypicanalyses: the use of polygenic (quantitative) charactersand genotype � environment interactions.

The basic concept of biochemical and molecularmethodologies has been presented by Larsen (1969). Hemaintained that “All morphological displays of varietiesmust, in the end, have a biochemical differentiation, but

not all biochemical characterisations necessarily have amorphological display; hence, biochemical characterswill be more numerous than morphological characters”.For example, the genotypes of several individuals mightbe distinguished by their phenolic content, as shown fora number of species by several authors (Singh andThompson, 1961; Brown et al., 1971). In olive, thesecompounds have been used to separate ten cultivars, butthe results did not prove to be sufficient to discriminateamong plants (Heimler et al., 1994).

Since the 1970s, protein analyses (total proteins and/orstorage proteins) have been as a method to identify anddistinguish species and cultivars. Storage proteins arewell-suited for use as genome markers. Protein analysis,compared to other biochemical markers, has theadvantage of simplicity of extraction and separationtechnique, although the information it provides ongenetic variability is less exhaustive. In olive, seedstorage proteins (Durante et al., 1992) and total leafproteins (Petruccelli, 1992) have been studied.

Enzymes are specialised proteins that catalyse thenumerous chemical reactions that occurr in livingbeings. The term “isoenzyme” (or isozyme) was firstproposed by Markert and Moller (1959) to indicatemultiple molecular forms of an enzyme, catalysing thesame biochemical reaction. Isoenzymaticelectrophoresis permits the detection of geneticvariability in several enzymatic proteins. Varietaldifferences are shown by means of enzymicpolymorphism on “activity gels”.Therefore “the analysisof a protein structure, by means of electrophoresis, isapproximately an analysis of the gene” (Gottlieb, 1977).Isozyme polymorphism became a simple marker toanalyse genetic relationships within a population(Gottlieb, 1981). However, isoenzymes suffer severallimitations: the genetic (isoenzymatic) loci testedrepresent a very small portion of the structural genes inthe whole genome; the variation present in a populationcan not be expressed in full; only some nucleotidesubstitutions that occur at the DNA level can beidentified because variations in amino acid sequence donot always take place. Nevertheless, numerous authors(Pontikis et al., 1980; Trujillo et al., 1989; Roselli et al.,1990; Petruccelli et al., 1992; Seker et al., 2005) employedisozymes to identify olive cultivars.

In recent years, in parallel with progress inbiochemical and bio-molecular marker techniques(Pontikis et al., 1980; Ouazzani et al., 1993; Trujillo et al.,1995; Hatzopoulos et al., 2002), molecular fingerprintingmethodologies have been established. In particular, theintroduction of DNA markers and of DNAfingerprinting techniques provide a high resolutiondiscrimination system that is independent ofenvironmental conditions and able to contribute to theresolution of problems such as mis-identification ingermplasm collections, nursery certification, varietalprotection, and/or guaranteed labelling for certificationof geographical origin (Powell et al., 1996;Testolin, 2000).The application of these technologies has alreadypermitted several advances: i) taxonomical revisionwithin the genus Olea and forms connected to Oleaeuropaea L.; ii) clarification of the relationships betweenwild and cultivated forms; and iii) contributions to thestudy of variability within the olive genome, by


identifying markers associated with specific characters ofagronomic interest (Rugini and Baldoni, 2003).

The first molecular marker technique employed onolive was Restriction Fragment Length Polymorphisms(RFLPs; Gallitelli et al., 1991). RFLP markers confirmedthat the Mediterranean Basin was the site ofdomestication of this fruit crop (Besnard et al., 2001).This technique has also been used by De la Rosa et al.(2003), when 95 plants obtained by crossing two cultivars(‘Leccino’ and ‘Dolce Agogia’) were analysed.

Randomly Amplified Polymorphic DNA (RAPD)has been applied successfully in many investigationsaimed at the study of polymorphism in olive cultivars,and at ways to distinguish them (Bogani et al. 1994;Fabbri et al., 1995; Cresti et al., 1996; Mekuria et al., 1999;Nikoloudakis et al., 2003; Perri et al., 1999; GonzaloClaros et al., 2000; Russo et al., 2001; Bronzini de Caraffaet al., 2002; Guerin et al., 2002; Mailer and May, 2002;Fabbri and Ganino, 2003; Belaj et al., 2001; 2002; 2003a;2004a; Mir Ali and Nabulsi, 2004; Ganino and Fabbri,2005; Hossein-Mazinani, 2005). The technique was alsoemployed by Gemas et al. (2000) and by Roselli et al.(2002) to evaluate intra- and inter-cultivar variability.Both RAPD and RFLP have contributed greatly to thetaxonomic classification within the genus Olea (Besnardet al., 2002).

Mini-satellites, or Variable Number of TandemRepeats (VNTR), are based on the study of smallgenome fragments which can be isolated due to theirspecific length. They consist of tandem repeats of a shortDNA sequence, and polymorphism is due to the numberof times the element is repeated in the genome (Sollerand Beekman, 1983). To use mini-satellites, it is notnecessary to create genomic libraries, but the techniqueis expensive. Contento et al. (2002) carried out a studybased on tandem repeats in three Olea europaea L.nuclear DNA sequences.

Amplified Fragment Length Polymorphisms(AFLPs) are very sensitive when seekingpolymorphism in the whole genome(Ranamukraarachchi et al., 2000). The use of AFLPmarkers made it possible to evaluate the geneticrelations between different forms of Olea and thecultivated olive, thus contributing to our understandingof the relative genetic distances separating forms thatbelong to different geographic areas (Angiolillo et al.,1999; Ambrosino and Rao, 2001; Ambrosino et al., 2002;Labombarda and Fontanazza, 2002; Ricciolini et al.,2003; Sanz-Cortes et al., 2003; Grati Kamoun et al., 2005;Kalaitzis et al., 2005; Owen et al., 2005; Strikic et al.,2005). By employing AFLPs, genetic affinity was alsoanalysed, in an attempt to clarify situations ofsynonymy and homonymy in a number of cultivarsfrom several Italian regions: Sicily (Angiolillo et al.,1999; Baldoni et al., 2002; 2003a; Labombarda andFontanazza, 2004); Sardinia (Angiolillo et al., 1998);Umbria (Marchionni et al., 1999; Labombarda et al.,2002; Baldoni et al., 2003b); Apulia (Resta et al., 2002);Marche (Baldoni et al., 2001); Molise (Pilla et al., 1999);Campania (Ambrosino et al., 2003); and Colli Euganei(Ziliotto et al., 2002). The simultaneous use of AFLPs,RAPDs and morphological characters madediscrimination possible between olive cultivars(Hagidimitriou et al., 2005)

Simple Sequence Polymorphisms, Simple SequenceRepeats (SSR) or microsatellites locate the DNAregions that are characterised by a tandem repeat of only1–6 bp in a sequence. Microsatellites are widespread anddispersed in the genomes of all plants, and display anelevated level of hypervariability within each species.These features makes them excellent both for genemapping and fingerprinting. Presently, the SSRtechnique has provided reliable markers to solve theproblems of varietal identification and for geneticcharacterisation of olive because of their hightransferability, elevated polymorphism and co-dominance, which is particularly interesting for olive(Sefc et al., 2000; Bandelj et al., 2002; Carriero et al., 2002;Marrazzo et al., 2002; Belaj et al., 2004b; 2005a; Bracciet al., 2005; Diaz et al., 2005a, b; Lopes et al., 2005).

Further research using microsatellites to identify olivecultivars was carried out by Rallo et al. (2000) whoanalysed 46 genotypes from several countries. In 2001,the SSR technique was also used to characterise ancientolive cultivars from Central Italy and Lake Garda(Cipriani et al., 2001).

Morphological and phenological aspects, as well asmolecular investigations (SSR markers) were used inSicily to characterise germplasm existing in the region(Caruso et al., 2005).

With the combined use of RAPDs and SSRs, thesolution to some cases of synonymy was made possible(Barranco et al., 2000b). The same combination ofmarkers led to the identification and characterisation ofIranian olive cultivars (Shahriari et al., 2005).

Simultaneous use of AFLPs and SSRs clarifiedrelationships between some Slovenian and Italian olivecultivars (Bandelj et al., 2004), within Italian, Spanish,French and Greek cultivars (Montemurro et al., 2005),and within Emilia-Romagna germplasm (Rotondi et al.,2003). RAPDs,AFLPs and SSRs were used by Belaj et al.(2003b) to evaluate the genetic relationships betweenItalian and Spanish cultivars. Later, Belaj et al. (2005b)used isozymes, RAPDs and SSRs to investigate therelationships between olive cultivars conserved in theWorld Olive Collection at Córdoba.

Single nucleotide polymorphisms (SNPs) represent anew and alternative technique for molecularidentification. By this type of analysis it is possible toidentify, in a very short time, differences betweengenomes due to a single base-pair (Cooper et al., 1985).These markers have been shown to be highlydiscriminatory both in the human genome (Collins et al.,1998) and with plants, particularly maize (Rafalski et al.,2001) and soybean (Rafalski, 2002). In olive, SNPs wereused for the first time to classify 51 olive cultivars, withsatisfactory results (Diaz Bermudez, 2005).

The integration between statistical methods applied tobiometry and to bio-molecular analyses is less well-developed. Some recent applications of bothmethodologies to the same genetic pool (Bassi et al.,2002; Rotondi et al., 2003) have demonstrated that thetwo procedures do not always give matchingrepresentations of similarity relations between cultivars.Although, in some cases, only molecular analyses wereable to characterise specific olive cultivars and toidentify erroneous classifications and synonyms, therehave also been cases in which molecular analysis alone





did not allow the separation and distinction of presumedclones which, however, presented visible differences inagronomic and morphological characteristics, and in theproperties of their oils (Perri et al., 1995).

To conclude, improvements in biochemical andmolecular techniques are proving to be valid instrumentsfor research in plant taxonomy, and for the identificationand characterisation of varieties. The improvement ofsuch techniques is still in progress and, although anincreasing number of papers are being published inwhich these are employed, the old variety descriptor list,with its indications of morphological and agronomictraits, retains its importance and usefulness, and is farfrom having become obsolete.

THE IMPORTANCE OF A CLASSIFICATION A journey through successive classifications since 1900

(Table II) shows a profound change in the choice ofcharacters adopted for cultivar identification. In thebeginning, botanical characters were of greatest interest,with some agronomic indications. A typical example isthe famous descriptor of Ciferri et al. (1942) which, in itsdiscrimination into “families”, merely returns to thesystematic concept of likeness between individuals.

Next, a change occurred as to choice of characters todescribe morphological and agronomic features. Thiscaused the loss of the “systematic” aspect in allsubsequently proposed descriptors. By the end of the1990s, it was felt necessary to introduce moretechnological and scientific characters, such as accuratechemical and organoleptic oil analyses and molecularcharacterisation.

It is also true that adoption of a new descriptionprocedure was not universal. On the contrary, eachauthor tended to represent and satisfy the needs of thecontext in which s/he operated. This led to thepublication of descriptor lists which were similar butdifferent, useful and interesting for one sector of theindustry, but less so for another. For example, adescriptor list that focusses on morphological charactersis seldom used by geneticists. Geneticists aim to separategenotypes that differ in a number of characters byeliminating, or at least reducing, the environmentalcomponent, and this is possible only within a commonenvironment, or by using methodologies that are notinfluenced by the environment.

An elaiographic methodology for olive is importantfor, and must satisfy the needs of: a) the botanist, toobtain valid information to begin any classification;b) the agronomist, to optimise cultivation techniques;c) the oil mill operator, to know the characteristics of thefruit and its oil content; d) the geneticist, as a startingpoint for genetic analyses and various breedingtechniques; and e) the nurseryman, who must be certainhe is propagating the most suitable material to marketwith due guarantees.

The need to construct an elaiographic descriptor thatincludes all characters useful for the completedescription of genotypes, therefore has the purpose ofsupporting both the production sector and scientificresearch.

The industry-oriented role is clear, there is a perpetualneed to improve production, both quantitatively and

qualitatively. This can only be achieved if thecharacteristics of cultivars from various regions areassessed, and there is easy access to certified propagationmaterial. These requirements can only be satisfied usingidentification tools of undisputed efficacy.

We have seen that a consideration of morphologicalcharacters alone is not sufficient to completely identifycultivars, although it represents an important advance inknowledge. For a better, more definitive description it isnecessary to grow and study plants in the sameenvironment, in which the features of each genotype canbe evaluated and compared according to their intrinsiccharacters. Such evaluation is more accurate if more thanone environment is used. Hence the usefulness ofreuniting those genotypes to be studied in the samecollection fields, possibly covering national or regionalgermplasm. A collection field, in order to deserve such adefinition, must contain plants of indisputed origin, withsufficient numbers of each accession to allow correctevaluation and to guarantee survival. In addition, thecollection must be accessible to all scholars, to facilitatecomparisons between germplasm, and to start breedingprogrammes.

From a scientific point of view, individual cultivars canbe regarded as potential sources of genetic material, tobe used to transmit characters concerning productivity,disease resistance or tolerance, etc., to new genotypesproduced through breeding programmes. Thus, theprotection of olive germplasm is particularly important,as it includes the gene pools of both domestic varietiesand related wild species, amenable to cross-breeding.Theneed to safeguard our existing genetic heritage thereforeappears increasingly urgent. But this is only feasibleinsofar as it is possible to identify, catalogue, andeliminate synonyms and homonyms still present ingermplasm in all olive-producing countries.

From the above, the need to establish a “varietalstandard” (as has been done with other fruit species)becomes apparent. This requires the creation of areference point, obtained by using averages of all knowncharacters, so that all fluctuations in a population canserve to define the cultivar more precisely. The more wecan eliminate environment-related variables by creatingcollections and by working in a restricted environmentwith a given number of trees per variety, the morereliable will be this reference. At the same time, it isextremely important to describe each plant in its originalenvironment, where it grew and developed, and atdifferent sites.

The IOOC acknowledged the need to have the largestnumber of olive genotypes in the same environment, andadvocated the organisation of three World germplasmcollections located in Spain (Cordoba), Italy (Cosenza)and Morocco (Marrakech). The Spanish collection isalready established. The other two are now beingrealised.

Today, although all published monographs on oliveclassification refer to clearly defined areas, theirreliability is fairly low. This is because all authors tend todescribe only those cultivars present in their ownterritory, and disregard those present in adjacent areasand/or in other olive-producing countries, with theirfrequent homonyms, synonyms and discrepancies as tothe classification of the same variety. In addition, this


TABLE IIComparison of descriptor lists established since 1942 according to the characters they describe and olive growing areas

LIST A – PLANT PASSPORT DATAAUTHOR*

CHARACTER† A B C D E F G H I J K L M N O P Q R S T U V W Y

Cultivar name X X X X X X X X X X X X X X X X X X X X X X XSynonyms X X X X X X X X X X X X X X XOrigin X X X X X X XDiffusion X X X X X X X X X X X X X XArea of major diffusion X X XFrequency of cultivation X X X% Presence of the cultivar X XValue XFruit purpose X X X X X X X X X X XBiochemical characterisation XMolecular characterisation X XPatents XExistence of collections X X

LIST B – CHARACTERS OF THE TREEAUTHOR*


Average height X XVigour X X X X X X X X X X X X X X X X X X X X X X X XGrowth habit X X X X X X X X X X X X X X X X X X X X XCanopy colour XCanopy width XCanopy shape X XCanopy section X X XCanopy volume XCanopy vigour XCanopy density X X X X X X X X X X X X X XInflorescence emission XFruit set XTrunk size XTrunk shape XTrunk surface XBark appearance and colour X XTrunk section X

LIST C – CHARACTERS OF MAIN SCAFFOLDSAUTHOR*


Average size XShape XSurface XColour XGrowth habit X

LIST D – CHARACTER OF FRUITING SHOOTSAUTHOR*


Abundance X XSize XShape XSurface XGrowth habit X X X X X X XLength X X X X XSection XInternode length X X X X X X X X X XColour X X X X XShoot XFeathers X X X X

*Ciferri et al., 1942, Italy (A); Patac et al., 1954, Spain (B); D Amore et al., 1977, Calabria (Italy) (C); Loussert et al., 1978, World (D); Barranco et al.,1984, Andalusia (Spain) (E); UPOV, 1985, World (F); Leitão et al., 1986, Portugal (G); Cimato et al., 1993, Tuscany (Italy) (H); Tous Martí and RomeroAroca, 1993, Catalonia (Spain) (I); Cristoferi et al., 1997, Emilia Romagna, (Italy) (J); Bartolini et al., 1998, World, (K); Pannelli et al., 1998, Abruzzo(Italy) (L); Cicoria et al., 2000, Molise (Italy) (M); Pannelli et al., 2000, Umbria (Italy) (N); Barranco et al., 2000a, World (O); Pugliano et al., 2000,Campania (Italy) (P); Cimato et al., 2001, Tuscany, (Italy) (Q); Rotundo and Marone, 2002, Lucania (Italy)(R); Trigui et al., 2002, Tunisia (S); Parlatiand Pandolfi, 2003, Lazio (Italy)(T); Lombardo et al., 2003, Calabria (Italy)(U); Bassi, 2003, Lombardy (Italy) (V); Lombardo et al., 2004, Apulia(Italy)(W); Rallo et al. 2005, Spain (Y).†The characters column carries as accurate as possible a translation of the original terms. When terminology was similar and obviously indicated thesame character, the terms were unified in English and listed in the same row; otherwise the character was given a separate row, especially if, in theoriginal publication, no clear indications were given as to the meaning of the character.


LIST E – CHARACTERS OF THE LEAFAUTHOR*


Abundance XShape X X X X X X X X X X X X X X X X X X X XLength/ Width ratio X X X X X X X XSize X X X X X X X X X X X X XLength X X X X X X X X XWidth X X X X X X X X XSymmetry XTile curvature XVeins XMidrib XSecondary veins XMucro XPetiole XTransverse section X XLeaf bearing X% Stomata open in the hottest hours XMargins XBlade XLongitudinal curvature of the blade X X X X X X X X X X X X X X XSurface X X X X X X X X XProfile of blade X XTorsion XApex angle X X X X X X X X XBase angle X X X X X X X X XPosition of maximum width X X X X X X XConsistency X X XColour X XDorsal colour X X X X X X X X X X XVentral colour X X X X X X X XBrightness X X XAbnormal leaves X XShape of abnormal leaves X X

LIST F – CHARACTERS OF THE INFLORESCENCEAUTHOR*


Time of development XPosition XShape X X X X X X X X XRachis length X X X X X X X X X X X X X X X XPeduncle length X XMaximum length XBranching type XColour at anthesis XPresence of axillary flowers XDensity XArrangement XNumber of inflorescence per branch XTotal number and position of flowers XFlower number/inflorescence X X X X X X X X X% Terminal inflorescences X% Simple and compound inflorescences XSize of swollen buds X X X XFrequency of supernumerary flowers X X XRachis structure X X X X X X X X X X X XRachis branching X X X XTime of full bloom X X X X X X X XFlower diameter XCorolla colour XPetal length XStylus length XShape and size of stigma X% Abortive flowers X X X X XNumber of fertile flowers X% Fertilisation XSelf-pollination or heterogamy X X XLevel of incompatibility XIncidence and localisation XAxillary flowers XPollen X


LIST G – CHARACTERS OF THE FRUITAUTHOR*


Fruit number per infructescence XSize X X X X X X X XWeight X X X X X X X X X X X X X XVolume and weight: averages and upper

values XLength X X X XSymmetry X X XShape X X X X X X X X X X X X X X X X X X X X X XDiameter medium X XAverage length/average diameter ratio X XMaximum diameter X XShape of transverse section X XSymmetry in position A X X X X X X X X X X X X X XSymmetry in position B X XSection XShape maximum transverse section XPosition maximum transverse diameter X X X X X X X X X X X X X X XShape of apex in position A X X X X X X X X X X X X X X X X XShape of apex in position B X X XShape of base in position A X X X X X X X X X X X X X X X XShape of base in position B X X XPedicel insertion XPedicel length XPedicel diameter XShape of pedicel cavity X X X X X X X X X XSize of pedicel cavity X X X X X X X X X XDepth of pedicel cavity X X XUmbo X X X X X XMucro X X X X X XPulp/endocarp ratio X X X X X X XSarcocarp type (at maturity) XSarcocarp consistency (at maturity) XSarcocarp (%) XEpicarp surface X XEpicuticular wax coating X X X X X X XLenticels, coloration, frequency X X X X X X X X X X X XLenticels size X X X X X XPerceptibility of lenticels at maturity X XEpicarp X XFruit distribution in 2–3 year-old plants XColour uniformity at maturity XColour before maturity XColour at maturity X X X X X XColour at harvest X X X X X X XColour evolution XPulp consistency X X XAdhesion pulp to epidermis XMesocarp colour XCuticle thickness XPosition of pistil scar X XResistance to abscission X XTime of maximum oil content X X XMesocarp clinginess X XTurning to dark colour (Veraison) X X X X X X X X X X X X XBiometrics indices XAverage weight/single fruit weight ratio XAverage volume/volume 100 olive fruits XSarcocarp/endocarp ratio XSarcocarp weight/weight of 100 olive fruits X

Average fruit volume/volume of singlefruit ratio X

Fruit drop (after fruit set) X X X XAmount of olive drop (% total olive

fruits at end season) X XTime of onset of olive drop X

*Ciferri et al., 1942, Italy (A); Patac et al., 1954, Spain (B); D Amore et al., 1977, Calabria (Italy) (C); Loussert et al., 1978, World (D); Barranco et al.,1984, Andalusia (Spain) (E); UPOV, 1985, World (F); Leitão et al., 1986, Portugal (G); Cimato et al., 1993, Tuscany (Italy) (H); Tous Martí and RomeroAroca, 1993, Catalonia (Spain) (I); Cristoferi et al., 1997, Emilia Romagna, (Italy) (J); Bartolini et al., 1998, World, (K); Pannelli et al., 1998, Abruzzo(Italy) (L); Cicoria et al., 2000, Molise (Italy) (M); Pannelli et al., 2000, Umbria (Italy) (N); Barranco et al., 2000a, World (O); Pugliano et al., 2000,Campania (Italy) (P); Cimato et al., 2001, Tuscany, (Italy) (Q); Rotundo and Marone, 2002, Lucania (Italy)(R); Trigui et al., 2002, Tunisia (S); Parlatiand Pandolfi, 2003, Lazio (Italy)(T); Lombardo et al., 2003, Calabria (Italy)(U); Bassi, 2003, Lombardy (Italy) (V); Lombardo et al., 2004, Apulia(Italy)(W); Rallo et al. 2005, Spain (Y).†The characters column carries as accurate as possible a translation of the original terms. When terminology was similar and obviously indicated thesame character, the terms were unified in English and listed in the same row; otherwise the character was given a separate row, especially if, in theoriginal publication, no clear indications were given as to the meaning of the character.


LIST H – CHARACTERS OF THE ENDOCARPAUTHOR*


Weight X X X X X X X X X X X XShape (positions A and B) X X X X X X X X X X X X X X X X X XThickness XThickness/weight ratio XLength X X X X XAverage diameter X X XLength/diameter ratio X XSize X X X X X XSymmetry (positions A and/or B) X X X X X X X X X X X X X X XVolume and weight: averages and upper

values XDiameters ratio (longitudinal and

transverse) XSection XBiometrics indices XMaximum diameter X XShape of largest transverse section XPosition of longest transverse diameter X X X X X X X X X X X X X X XApex (positions A and B) X X X X X X X X X X X X X X X X XBase (positions A and B) X X X X X X X X X X X X X X X XColour XSurface: colour, roughness and groove

number XSurface X X X X X X X X X X X X X X XGroove type XGroove number X X X X X X X X X X X X X XGroove gait X X X X X X XGroove distribution X X X XGroove depth X X X X X X XSuture line X XSuture curvature XApex termination X X X X X X X X X X X X X X

LIST I – CHARACTERS OF THE SEEDAUTHOR*


Shape XColour XSize XFlavour XAverage weight X XNumber of seeds per fruit XSeedless fruits X

LIST J – AGRONOMIC AND COMMERCIAL CHARACTERSAUTHOR*


Precocity X XOnset of bearing X X X X X X X XProductivity X X X X X X X X X X X X X X X X XProduction regularity X X X X X X X X X X X XDrupe oil content X X X X X X X X X X X XOil yield X X X X X X X X X X X X XOil quality X X X X X X XCourse of oil accumulation X XMesocarp clinginess X XRooting ability X X X X X X X XBloom time XTime of recovery of vegetative activity

and inflorescence emission XBloom duration XDays from full bloom to fruit ripening XPollinators X XCompatibility X X X X X X X X XFlowers/ripe fruits ratio XFruit-set after free pollination XFruit-set after self pollination XAverage no of seeds per drupe (%) XFruit ripening time X X X X X X X X XSusceptibility to abiotic stress X X X X X X X X X X X X X X X XSusceptibility to biotic stress X X X X X X X X X X XHardiness X X X XSuitability to mechanical harvest XParticular conditions for variety

identification XInfluence on the plant of environment

and cultivation techniques X XAdditional information X X


makes it particularly difficult to transfer data obtained ina given environment to other environments, due to a lackof common references.

As a consequence of these considerations, theproposed classification methods, even the most recentones, suffer grave limitations due to: a) polygenic controlof morphological characters and the strong influence ofenvironment and cultivation techniques on theirexpression; b) the absence of identical collections indifferent environments, which would be extremely usefulin the study of genotype � environment interactions;c) the need to consider a large number of characters todraw up a suitable descriptor list; and d) the need togather data concerning the required characters for aminimum number of years.

In conclusion, we must acknowledge that we do notyet possess an accurate methodology to identify olivevarieties. This causes a delay in olive breeding, andconsequently a delay in improvements in oliveproduction. The unification of all methodologies, bycreating a common pomological descriptor list for allscholars and scientists in the field, consisting of a seriesof characters useful both for genotype description andscientific research, would be a significant advance. Inaddition, any new list should include informationobtained by new research methods (physiological dataand ultrastructural analysis, serological, biochemical andmolecular marker analyses). We must admit that the useof morphological, biological and agronomic charactersalone does not permit an accurate varietal identificationof olive cultivars. The sole use of molecular markers,which are receiving increasing interest and credit, doesnot necessarily provide the accuracy and reliability theyseem to inspire. Although DNA marker techniques are

theoretically valid, in practice the methodology needs tobe better tested and improved. Actually, there are limitsin the application of the new techniques. It must beremembered that: a) even reproducible differences inDNA profiles only offer the certainty of the negativeresult, and never genetic identity; consequently, b) whenthe molecular profiles between two plants are identical,we still do not have absolute certainty that we aredealing with the same genotype. We can only reasonablydeduce that two individuals can be referred to the samegenotype, with very low or no significant (≤ 1%) marginfor error, provided the numerical acceptabilitythresholds of the molecular markers being tested, whichvary according to species and type of marker, are passed.

The problem of olive classification is therefore ofprime importance. Equally apparent is the need for theconstruction of an univocal descriptor list, theestablishment of collection fields, and the creation of acomplete data-bank.

Nevertheless, this review can close on an optimisticnote. The Italian Accademia Nazionale dell’Olivo edell’Olio of Spoleto has sponsored a Committee of themost eminent Italian olive scientists to prepare adescriptor list (plant passport data, morphological,biological, agronomic and commercial characters,biochemical and molecular markers, collections, etc.), atdifferent levels, and valid for all olive-producingcountries, which will soon be submitted to theinternational research community.

The “new” descriptor list, once accepted by all thoseinvolved in the olive industry, will represent a model thatcan eliminate, or at least contain the confusionaccumulated so far in attempts to describe the cultivatedolive.

AMBROSINO, O. and RAO, R. (2001). Marcatori AFLP per la valu-tazione della variabilità genetica intra-intervarietale in cultivardi olivo. Atti VI Convegno Nazionale Biodiversità, Valenzano,Italy. 2, 397–405.

AMBROSINO, O., MANZO, M. and RAO, R. (2002). Identificazione erelazioni genetiche tra cultivar di olivo mediante marcatoriAFLP. Atti “Convegno Internazionale di Olivicoltura”, Spoleto,Italy. 332–337.

AMBROSINO, O., MANZO, M., PUGLIANO, G., MONTI, L. M. and RAO,R. (2003). Identificazione di sinonimi ed omonimi di varietà diolivo incluse nei disciplinari DOP Campani mediante marcatoriAFLP. Atti Convegno “Germoplasma olivicolo e tipicità dell’olio”, Perugia, Italy. 253–257.

ANGIOLILLO, A., BALDONI, L., BANDINO, G. and MULAS, M. (1998).Analisi molecolare con marcatori AFLP delle risorse genetichedi olivo della Sardegna. Atti IV Convegno NazionaleBiodiversità, Alghero, Italy. 413–416.

ANGIOLILLO, A., MENCUCCINI, M. and BALDONI, L. (1999). Olivegenetic diversity assessed using amplified length polymor-phisms. Theoretical and Applied Genetics, 98, 411–421.

BALDINI, E. and SCARAMUZZI, F. (1952). Sul valore dei dati bio-metrici nella descrizione e classificazione delle razze di olivo incoltura. Annali Istituto Sperimentale Agronomico, 6, 1597–1636.

BALDINI, E. and SCARAMUZZI, F. (1955). Ulteriori indagini sullavalidità del metodo bio-statistico nella descrizione e classifi-cazi-one delle cultivar di olivo. Annali Istituto SperimentaleAgronomico, 9, 171–186.

BALDONI, L., ALFEI, B., SANTINELLI, A., ANGIOLILLO, A. and PAN-

NELLI, G. (2001). Esame dei principali casi di sinonimia edomonimia in varietà di olivo marchigiane mediante analisiAFLP. Atti VI Convegno Nazionale Biodiversità, Valenzano,Italy. 2, 431–440.

BALDONI, L., PANNELLI, G., ALFEI, B. and BANDINO, G. (2002).Relazioni genetiche tra le varietà di olivo Bosana, Coroncina ePeranzana. Atti Convegno Internazionale di Olivicoltura,Spoleto, Italy. 387–390.

BALDONI, L., RICCIOLINI, C., DI MARCO, L. and GERMANÀ, M.A.(2003a). Variabilità e relazioni genetiche tra le varietà di olivodella Sicilia. Atti Convegno Germoplasma olivicolo e tipicitàdell’olio, Perugia, Italy. 211–215.

BALDONI, L., RICCIOLINI, C., MUNARI, C., PANNELLI, G. andARCIONI, S. (2003b). Caratterizzazione del DNA di varietà edecotipi di olivo dell Umbria. Atti Convegno Germoplasma olivi-colo e tipicità dell’olio, Perugia, Italy. 312–315.

BANDELJ, D., JAKS̆E, J. and JAVORNIK, B. (2002). DNA fingerprintingof olive varieties by microsatellite markers. Food Technologyand Biotechnology, 40, 185–190.

BANDELJ, D., JAKS̆E, J. and JAVORNIK, B. (2004). Assessment ofgenetic variability of olive varieties by microsatellite and AFLPmarkers. Euphytica, 136, 93–102.

BARONE, E., CARUSO, T., MARRA, F. P. and MOTISI, A. (1994a).Caratteristiche biometriche di 25 cultivar di olivo del germo-plasma siciliano. Atti Convegno “Tecniche, norme e qualità inolivicoltura”, Potenza, Italy. 623–630.

REFERENCES


BARONE, E., DI MARCO, L., MOTISI, A. and CARUSO, T. (1994b). TheSicilian olive germplasm and its characterization by using sta-tistical methods. Acta Horticulturae, 356, 66–69.

BARRANCO, D. and RALLO, L. (1984). Las Variedades de OlivoCultivadas en Andalucia. Ed. Junta de Andalucia, Conserjeriade Agricoltura, Pesca e Alimentacion, Cordoba, Spain. 33–63.

BARRANCO, D., CIMATO, A., FIORINO, P., RALLO, L., TOUZANI, A.,CASTANEDA, C., SERAFIN, F. and TRUJILLO, I. (2000a). WorldCatalogue of Olive Varieties. Consejo Oleìcola Internacional,Madrid. 360 pp.

BARRANCO, D., TRUJILLO, I. and RALLO, P. (2000b). Are “Oblonga”and “Frantoio” olives the same cultivar? Journal of theAmerican Society for Horticultural Science, 35, 1323–1325.

BARTOLINI, G. and PETRUCELLI, R. (1994). Metodi biochimici e mol-ecolari per l’identificazione delle specie coltivate. RivistaSementi Elette, 5, 45–56.

BARTOLINI, G., BARONCELLI, S., PETRUCCELLI, R. and PUGLIESI, C.(1992). La caratterizzazione varietale dell’olivo e relative prob-lematiche. Atti Congresso “Salvaguardia e valorizzazione dellerisorse genetiche”, Alghero, Italy. 637–642.

BARTOLINI, G., PREVOST, G. and MESSERI, C. (1994a). Olive treegermplasm: descriptor list of cultivated varieties in the world.Acta Horticulturae, 356, 116–118.

BARTOLINI, G., PETRUCCELLI, R., TOPONI, M. A., DI MONTE, G. andPANICUCCI, M. (1994b). Morphological and biochemical evalu-tion of the Olea europaea L. cv ‘Leccino’. Acta Horticulturae,356, 78–81.

BARTOLINI, G., PREVOST, G., MESSERI, C. and CARIGNANI, G. (1998).Olive Germplasm: Cultivars and World-wide Collections. Seedand Plant Genetic Resources Service, F.A.O., Rome, Italy. 459pp.

BASSI, D. (2003). Il Germoplasma dell’Olivo in Lombardia. RegioneLombardia-Università degli Studi, Milano, Italy. 88 pp.

BASSI, D., TURA, D., GEUNA, F., FAILLA, O. and PEDÒ, S. (2002).Characterisation of local olive (Olea europaea L.) accessions byoil composition, morphological and molecular markersmethods. Acta Horticulturae, 586, 57–60.

BELAJ, A., TRUJILLO, I., DE LA ROSA, L. and RALLO, L. (2001).Polymorphism and discrimination capacity of randomly ampli-fied polymorphic markers in an olive germplasm bank. Journalof the American Society for Horticultural Science, 126, 64–71.

BELAJ, A., SATOVIC, Z., RALLO, J. and TRUJILLO, I. (2002). Geneticdiversity and relationships in olive (Olea europaea L.)germplasm collections as determined by randomly amplifiedpolymorphic DNA. Theoretical and Applied Genetics, 105,638–644.

BELAJ, A., CABALLERO, J. M., BARRANCO, D., RALLO, L. and TRU-

JILLO, I. (2003a). Genetic characterization and identification ofnew accessions from Syria in an olive germplasm bank bymeans of RAPD markers. Euphytica, 134, 261–268.

BELAJ, A., SATOVIC, Z., CIPRIANI, G., BALDONI, L., TESTOLIN, R.,RALLO, L. and TRUJILLO, I. (2003b). Comparative study of thediscriminating capacity of RAPD, AFLP and SSR markers andof their effectiveness in establishing genetic relationships inolive. Theoretical and Applied Genetics, 107, 736–744.

BELAJ, A., SATOVIC, Z., RALLO, J. and TRUJILLO, I. (2004a). Optimaluse of RAPD markers for identifying varieties in olive (Oleaeuropaea L.) germplasm collection. Journal of the AmericanSociety for Horticultural Science, 129, 266–270.

BELAJ, A., CIPRIANI, G., TESTOLIN, R., RALLO, L. and TRUJILLO, I.(2004b). Characterization and identification of the mainSpanish and Italian olive cultivars by simple-sequence-repeatmarkers. Journal of the American Society for HorticulturalScience, 39, 1557–1561.

BELAJ, A., DIEZ, C., SATOVIC, Z., BALDONI, L. and BARRANCO, D.(2005a). Collection and study of wild olive populations in Spainby means of SSR markers. Vth International Symposium onOlive Growing, Izmir, Turkey. 123.

BELAJ, A., DE LA ROSA, R., RALLO, P., ANGELES OJEDA, M., DIAZ,A., MUÑOZ, C., TRUJILLO, I., BALDONI, L., MARTIN, A., BAR-

RANCO, D. and RALLO, L. (2005b). Molecular markers in olive:an integrated approach. Vth International Symposium on OliveGrowing, Izmir, Turkey. 98.

BESNARD, G., BARADAT, P., CHEVALIER, D., TAGMOUNT, A. andBERVILLÉ, A. (2001). Genetic differentiation in the olivecomplex (Olea europaea) revealed by RAPDs and RFLPs inthe rRNA genes. Genetic Resources and Crop Evolution, 48,165–182.

BESNARD, G., KHADARI, B., BARADAT, P. and BERVILLÉ, A. (2002).Olea europaea (Oleaceae) phylogeography based on chloro-plast DNA polymorphism. Theoretical and Applied Genetics,104, 1353–1361.

BOGANI, P., PETRUCCELLI, R., POLSINELLI, L. and ROSELLI, G. (1994).Identification olive tree cultivars by using random amplifiedpolymorphic DNA. Acta Horticulturae, 356, 98–101.

BOTTARI, V. and SPINA, P. (1953). Le varietà di olivo coltivate inSicilia. Annali Istituto Sperimentale Agronomico, 7, 937–1004.

BRACCI, F. (1937). Le varietà di olivo coltivate in Toscana. In: LeVarietà di Olivo Coltivate in Italia. REDA, Rome, Italy. 3–15.

BRACCI, T., BUSCONI, M., FOGHER, C. and SEBASTIANI, L. (2005).Evaluation of genetic diversity in Liguria region olive (Oleaeuropaea L.) germplasm by SSR markers. Vth InternationalSymposium on Olive Growing, Izmir, Turkey. 102.

BRONZINI DE CARAFFA, V., GIANNETTINI, J., GAMBETTI, C. andMAURY, J. (2002). Genetic relationships between cultivated andwild olives of Corsica and Sardinia using RAPD markers.Euphytica, 123, 263–271.

BROWN, G. B., DEAKIN, J. R. and HOFFMAN, J. C. (1971).Identification of snap bean cultivars by paper chromatographyof flavonoids. Journal of the American Society for HorticulturalScience, 96, 477–481.

CANTINI, C., CIMATO, A. and SANI, A. (1999). Morphological evalu-ation of olive germplasm present in Tuscany region. Euphytica,109, 173–181.

CARRIERO, F., FONTANAZZA, G., CELLINI, F. and GIORIO, G. (2002).Identification of simple sequence repeats (SSRs) in olive (Oleaeuropaea L.). Theoretical and Applied Genetics, 104, 301–307.

CARUSO, G. (1883). Monografia dell’Olivo. Enciclopedia AgricolaItaliana. U.T.E.T., Torino, Italy. 533 pp.

CARUSO, T., CAMPISI, G., MOTISI, A., LA MANTIA, M., OCCORSO, G.and TESTOLIN, R. (2005). Morphological, phenological andmolecular characterization of the Sicilian indigenous olive(Olea europaea sativa L.) germplasm. Vth InternationalSymposium on Olive Growing, Izmir, Turkey. 99.

CICORIA, M., CORBO, M., D’UVA, T., D’UVA, T. and RUGGIERO, A.(2000). Il germoplasma dell’Olivo nel Molise. ERSA-Molise,Campobasso, Italy. 64 pp.

CIFERRI, R. and BREVIGLIERI, N. (1942). Introduzione ad una classi-ficazione morfo – ecologica dell olivo coltivato in Italia.L’Olivicoltore, 1, 1–2.

CIFERRI, R., MARINUCCI, M. and MORETTINI, A. (1942). Dati pre-liminari per una sistematica delle razze di olivo in coltura.L’Olivicoltore, 1, 3–7.

CIMATO, A., CANTINI, C., SANI, G. and MARRANCI, M. (1993). IlGermoplasma dell’Olivo in Toscana. Regione Toscana,Florence, Italy. 81 pp.

CIMATO, A., CANTINI, C. and SANI, G. (2001). L’olivo in Toscana: ilGermoplasma Autoctono. Ed. ARSIA Toscana-IPSL, CNR-Regione Toscana, Florence, Italy. 215 pp.

CIMATO, A., CANTINI, C., SANI, G., ROMANI, A., AUTINO, A. andCRESTI, M. (2004). Il germoplasma autoctono dell’olivo inToscana. ARSIA Toscana-Regione Toscana, Florence, Italy.56 pp.

CIPRIANI, G., MARRAZZO, M. T., MARCONI, A., CIMATO, A. andTESTOLIN, R. (2001). Microsatellite markers isolated in olive(Olea europeaea L.) are suitable for individual fingerprintingand reveal polymorphism within ancient cultivars. Theoreticaland Applied Genetics, 104, 223–228.

COLLINS, F. S., BROOKS, L. D. and CHARKRAVARTI, A. (1998). ADNA polymorphism discovery resource for research on humangenetic variation. Genome Research, 8, 1229–1231.

CONTENTO, A., CECCARELLI, M., GELATI, M. T., MAGGINI, F.,BALDONI, L. and CIONINI, P. C. (2002). Diversity of Olea geno-types and the origin of cultivated olives. Theoretical andApplied Genetics, 104, 1229–1238.

COOPER, D. N., SMITH, B. A., COOKE, H., NIEMANN, S. andSCHMIDTKE, J. (1985). An estimate of unique sequence het-erozygosity in the human genome. Human Genetics, 69,201–205.

CRESTI, M., LINSKENS, H. F., MULCAHY, D. L., BUSH, S., DI STILIO,V.,XU, M. Y., VIGNANI, R. and CIMATO, A. (1996). Preliminarycommunication about the identification of DNA in leaves andin olive oil of Olea europaea. Advances in Horticultural Science,10, 105–107.


CRISTOFERI, G., ROTONDI, A. and MAGLI, M. (1997). IlGermoplasma dell’Olivo in Emilia Romagna. ISTEA, Bologna,Italy, 80 pp.

D’AMORE, R., IANNOTTA, N. and PERRI, L. (1977). Contributo alloStudio delle Principali Cultivar d’Olivo Presenti in Calabria.ISOl, Cosenza, Italy. 3–9.

DAMIGELLA, P. (1960). Variabilità dei caratteri biometrici dell’olivoe impiego delle funzioni discriminanti. La Ricerca Scientifica, 4,522–530.

DE LA ROSA, R., ANGIOLILLO, A., GUERRIERO, C., PELLEGRINI, M.,RALLO, L., BESNARD, G., BERVILLÈ, A., MARTIN, A. andBALDONI, L. (2003).A first linkage map of olive (Olea europaeaL.) cultivars using RAPD, AFLP, RFLP and SSR markers.Theoretical and Applied Genetics, 106, 1273–82.

DIAZ BERMUDEZ, A. (2005). Desarrollo y caracterización de nuevosmicrosatélites y SNPs y aplicacion en la mejora genètica delolivo (Olea europaea L.). Ph.D. Thesis, Departamento deGenètica, Universidad de Córdoba, E.T.S.I.A.M. Córdoba.Spain. 183 pp.

DIAZ, A., DE LA ROSA, R., MARTIN, A. and RALLO, P. (2005a).Development and characterization of 12 new microsatellites inolive (Olea europaea L.). Vth International Symposium on OliveGrowing, Izmir, Turkey. 118.

DIAZ, A., DE LA ROSA, R., MARTIN, A. and RALLO, P. (2005b).Cultivar identification and elucidation of genetic relationshipswithin the species Olea europaea L. using microsatellites. Vth

International Symposium on Olive Growing, Izmir, Turkey. 94.DURANTE, M., PETRUCELLI, R., BARTOLINI, G. and BERNARDI, R.

(1992). Impiego delle proteine di riserva per l’identificazionedelle cultivar di olivo (Olea europaea L.). Atti Convegno “OliveOil Quality”, Florence, Italy. 57–60.

FABBRI, A. and GANINO, T. (2003). Il germoplasma olivicolo inEmilia. Atti Convegno “Germoplasma olivicolo e tipicitàdell’Olio”, Perugia, Italy. 48–52.

FABBRI, A., HORMAZA, J. I. and POLITO, V .S. (1995). RandomAmplified Polymorphic DNA analysis of olive (Olea europaeaL.) cultivars. Journal of the American Society for HorticulturalScience, 120, 538–542.

FREZZOTTI, G. (1937). Studio biometrico sulle olive di diversevarietà e provenienze. In: Le Varietà di Olivo Coltivate in Italia.REDA, Rome, Italy.

GALLITELLI, M., SEMERARO, L. and ANTONELLI, N. M. (1991). RFLPanalysis in the olive (Olea europaea L.). EMBO Course,Cologne. Germany.

GANINO, T. and FABBRI, A. (2005). Genetic characterization of Oleaeuropaea L. germplasm in northern Italy. Vth InternationalSymposium on Olive Growing, Izmir, Turkey. 127.

GEMAS,V. J., RIJO-JOHANSEN, M. J.,TENREIRO, R. and FEVEREIRO, P.(2000). Inter- and intra-varietal analysis of three Olea europaeaL. cultivars using the RAPD technique. Journal of the AmericanSociety for Horticultural Science and Biotechnology, 75,312–319.

GONZALO CLAROS, M., CRESPILLO, R., AGUILAR, M. L. andCÁNOVAS, F. M. (2000). DNA fingerprinting and classification ofgeographically related genotypes of olive-tree (Olea europaeaL.). Euphytica, 116, 131–142.

GOTTLIEB, L. D. (1977). Electrophoretic evidence and plant system-atic. Annals of the Missouri Botanical Garden, 64, 161–180.

GOTTLIEB, L. D. (1981). Electrophoretic evidence and plant popula-tions. Progress in Phytochemistry, 7, 1–46.

GRATI KAMOUN, N., LAMY MAHMOUD, F., REBAÏ,A., GARGOURI,A.,PANAUD, O. and SAAR, A. (2005). Genetic diversity (inter- andintra-varietal) of some Tunisian olive tree cultivars detected byAFLP markers. Vth International Symposium on Olive Growing,Izmir, Turkey. 91.

GUERIN, J., SWEENEY, S., COLLINS, G. and SEDGLEY, M. (2002). Thedevelopment of a genetic database to identify olive cultivars.Journal of the American Society for Horticultural Science, 127,977–983.

HAGIDIMITRIOU, M., KATSIOTIS, A., MENEXES, G., PONTIKIS, C. andLOUKAS, M. (2005). Genetic diversity of major Greek olive cul-tivars using molecular (AFLPs and RAPDs) markers and mor-phological traits. Journal of the American Society forHorticultural Science, 130, 211–217.

HATZOPOULOS, P., BANILAS, G. GIANNOULIA, K., GAZIS, F.,NIKOLOUDAKIS, N., MILIONI, D. and HARALAMPIDIS, K. (2002).Breeding, molecular markers and molecular biology of theolive tree. European Journal of Lipid Science and Technology,104, 574–586.

HEIMLER, D., CIMATO, A., PIERONI, A., SANI, G. and TATTINI, M.(1994). Seasonal trend of flavonoids, flavonoid glycosides, andbiflanoids in ten olive cultivars. Acta Horticulturae, 356,372–374.

HOSSEIN-MAZINANI, M. (2005). Olive growing and olive research inIran: a status report. Vth International Symposium on OliveGrowing, Izmir, Turkey. 113.

KALAITZIS, P., OWEN, C.A., CRAITA-BITA, E., HAJJAR, S.E., SELL-

IANAKIS, V., AKSOY, U., HEPAKSOY, S., CHAMOUN, R., TALHOOK,S., BANILAS, G., HATZOPOULOS, P. and METZIDAKIS, I. (2005).AFLP reveals structural details of genetic diversity within cul-tivated olive germplasm from the eastern Mediterranean. Vth

International Symposium on Olive Growing, Izmir, Turkey. 130.LABOMBARDA, P. and FONTANAZZA, G. (2002). Caratterizzazione

genetica del gruppo varietale del “Moraiolo”: utilizzo dei mar-catori molecolari AFLP per chiarire casi di omonimie e sinon-imie. Atti “Convegno Internazionale di Olivicoltura”, Spoleto,Italy. 406–410.

LABOMBARDA, P. and FONTANAZZA, G. (2004). I marcatori AFLPnello studio della biodiversità di olivo in Sicilia, con particolareriferimento alla provincia ennese. Atti convegno “Germoplasmaolivicolo e tipicità dell’olio”, Perugia, Italy. 196–201.

LABOMBARDA, P., CIPRIANI, M. and FONTANAZZA, G. (2002). AnalisiAFLP della biodiversità di germoplasma olivicolo umbro. Atti“Convegno Internazionale di Olivicoltura”, Spoleto, Italy.400–405.

LANZA, B., MARSILIO,V. and MARTINELLI, N. (1995). Identificazionevarietale di cultivars di olivo (Olea europaea L.). Approccianalitici quantitativi del pattern esinico del granello pollinico.Atti del convegno: “L’olivicoltura mediterranea: stato e prospet-tive della coltura e della ricerca”, Rende, Italy. 219–224.

LARSEN, A. L. (1969). Isoenzymes and varietal identification. SeedWorld, 104, 5–6.

LEITÃO, F., DE FATIMA POTES, M., LEONILDE CALADO, M. and JOSÉ

DE ALMEIDA, F. (1986). Descrição de 22 variedades de oliveiracultivadas em Portugal. Ministério da Agricultura, Pescas eAlimentação, Direccão Geral de Planeamento e Agricultura,Lisboa. 5–13.

LOMBARDO, N., PERRI, E., MUZZALUPO, A., MADEO, A., GODINO, G.and PELLEGRINO, M. (2003). Il Germoplasma OlivicoloCalabrese. ISOl-Regione Calabria-Co.R.Ass.Ol, Cosenza, Italy.26 pp.

LOMBARDO, N., MADEO, A., MAZZALUPO, I., ALESSANDRINO, M.,BELFIORE, T., CILIBERTI, A., GODINO, G., PELLEGRINO, M.,RIZZATI, B., PERRI, E., MAZZOTTI, F., RUSSO, A., SALERNO, R.,PARISE, A. and NOCE, M.E. (2004). Contributo allaCaratterizzazione del Germoplasma Olivicolo Pugliese. RegionePuglia-I.S.Ol.-Comunità Europea, Cosenza, Italy. 114 pp.

LOPES, M. S., MENDONÇA, D., SEFC, K. M., GIL, F. S. and MACHADO,A. C. (2005). Evidence of intra-cultivar genetic variability inolive cultivars. Vth International Symposium on Olive Growing,Izmir, Turkey. 133.

LORENZETTI, F., CECCARELLI, S. and VERONESI, F. (1996). GeneticaAgraria. Patron Editore, Bologna, Italy. 454 pp.

LOUSSERT, R. and BROUSSE, G. (1978). L’Olivier. G. P. Maisonneuve& Larose, Paris, France. 92–110.

MAAS, J. L. (1977). Pollen ultrastructure of strawberry and othersmall-fruit crops. Journal of the American Society forHorticultural Science, 102, 560–571.

MAILER, R. J., and MAY, C. E. (2002). Variability and inter-relation-ships of olive trees and cultivars using RAPD analysis.Advances in Horticultural Science, 16, 192–197.

MANCUSO, S. and NICESE, F. P. (1999). Identifying olive (Oleaeuropaea) cultivars using artificial neural networks. Journal ofthe American Society for Horticultural Science, 124, 527–531.

MARCHIONNI, C., BALDONI, L., ANGIOLILLO, A., PANNELLI, G. andPANARA, F. (1999). Caratterizzazione genetica delle più impor-tanti cultivar di olivo dell’Umbria. Atti 5° Convegno nazionalesulla biodiversità, Caserta, Italy. 13, 243–250.

MARINUCCI, M. (1908). Schema di sistemazione delle razze di olivocoltivate nell Italia meridionale. Atti Istitituto Incoragg, Naples,Italy. 4, 1–18.

MARKERT, C. L. and MOLLER, F. (1959). Multiple forms of enzymes:tissue, antogene and species specific patterns. Proceedings of theNational Academy Sciences of the USA, 45, 753–763.


MARRAZZO, T., CIPRIANI, G., MARCONI, R., CIMATO, A. andTESTOLIN, R. (2002). Isolation and characterisation ofmicrosatellite DNA in Olive (Olea europaea L.). ActaHorticulturae, 586, 61–64.

MEKURIA, G., COLLINS, G. and SEDGLEY, M. (1999). Genetic vari-ability between different accessions of some common commer-cial olive cultivars. Journal of Horticultural Science &Biotechnology, 74, 309–314.

MILIANI, R. (1932). Classificazione ecologica dell’olivo in Italia.Annali di Tecnica Agraria, V(III), 247–256.

MIRALI, N. and NABULSI, I. (2004). Molecular characterization andgenetic relatedness among olive cultivars using RAPDmarkers. Advances in Horticultural Science, 18, 173–180.

MONTEMURRO, C., SIMEONE, R., PASQUALONE, A., FERRAR, E. andBLANCO, A. (2005). Genetic relationships and cultivar identifi-cation among 112 olive accessions using AFLP and SSRmarkers. Journal of Horticultural Science & Biotechnology, 80,105–110.

MORETTINI, A. (1972). Olivicoltura. REDA, Rome, Italy. 522 pp.NIKOLOUDAKIS, N., BANILAS, G., GAZIS, F., HATZOPOULOS, P. and

METZIDAKIS, J. (2003). Discrimination and genetic diversityamong cultivated olives of Greece using RAPD markers.Journal of the American Society for Horticultural Science, 128,741–746.

OUAZZANI, N., LUMARET, R.,VILLEMUR, P. and GIUSTO, F. D. (1993).Leaf allozyme variation in cultivated and wild olive trees (Oleaeuropea L.). Journal of Heredity, 84, 34–42.

OWEN, C. A., BITA, E. C., BANILAS, G., HAJJAR, S. E., SELLIANAKIS,V., AKSOY, U., HEPAKSOY, S., CHAMOUN, R., TALHOOK, S. N.,METZIDAKIS, I., HATZOPOULOS, P. and KALAITZIS, P. (2005).AFLP reveals structural details of genetic diversity within cul-tivated olive germplasm from the Eastern Mediterranean.Theoretical and Applied Genetics, 110, 1169–1176.

PANDEY, K. M. and TROUGHTON, J. N. (1974). Scanning electronmicroscope observations of pollen grains and stigma and theself incompatible species. Euphytica, 23, 337 – 344.

PANNELLI, G., ALFEI, B. and SANTINELLI, A. (1998). Varietà di’Olivonelle Marche. ASSAM. Ancona, Italy. 108 pp.

PANNELLI, G., ALFEI, B., D AMBROSIO, A., ROSATI, S. and FAMIANI,F. (2000). Varietà di Olivo in Umbria. Pliniana, Perugia, Italy. 98pp.

PARLATI, M. V. and PANDOLFI, S. (2003). Catalogo delle principalivarietà di olivo selezionate nel Lazio. Unione Europea-RegioneLazio-ARSIAL, Rome, Italy. 46 pp.

PATAC DE LA TRAVIESAS, L., CADAHIA CIRCUENDEZ, P. and DEL

CAMPO SANCHEZ, E. (1954). Tratado de Olivicultura. SindicatoNacional del Olivo, Madrid, Spain.

PERRI, E., PARLATI, M. V., PALOPOLI, A., SIRIANNI, R. and GION-

FRIDDO, F. (1995). Caratterizzazione varietale in olivo (Oleaeuropaea L.) mediante analisi multivariata applicata alla com-posizione acidica di oli monovarietali. Atti Convegno “Tecniche,norme e qualità in olivicoltura”, Potenza, Italy. 687–698.

PERRI, E., LOMBARDO, N., PALOPOLI, A. and MIELE, D. (1999).Indagine sul germoplasma di olivo della Sicilia medianteRAPD. Vth Convegno nazionale sulla biodiversità, Caserta,Italy. 13, 299–304.

PETRUCCELLI, R. (1992). Analisi di caratteri biochimici e morfo-agronomici ai fini della identificazione varietale dell olivo (Oleaeuropaea L.). Ph.D. Thesis, Dipartimento di Ortoflorofrutti-coltura, Università di Firenze. Italy. pp. 1–40.

PILLA, F., VELTRI, C., LAGONIGRO, R., ANGIOLILLO, A., MAR-

CHIONNI, C. and BALDONI, L. (1999). Esame di alcune cultivar diolivo del Molise mediante marcatori AFLP. Vth Convegnonazionale sulla biodiversità, Caserta, Italy, 13, 305–309.

PONTIKIS, C. A., LOUKAS, N. and KOUSSONIG, G. (1980). The use ofbiochemical markers to distinguish olive cultivars. Journal ofthe American Society for Horticultural Science, 55, 333–343.

POWELL, W., MORGANTE, M., ANDRE, C., HANAFEY, M., VOGEL, J.,TINGEY, S. and RAFALSKI, A. (1996). A comparison of RFLP,RAPD, AFLP and SSR (microsatellite) markers for germplasmanalysis. Molecular Breeding, 2, 225–238.

PREVOST, G. and MOSTARDINI, S. (1999). Gli studiosi dell’olivo e lasua classificazione botanica. Olivæ, 78, 60–77.

PUGLIANO, G., FLAMINIO, G., PUGLIANO, G., PUGLIANO, M. L.,SANNINO, G. and SCHIAMONE, S. (2000). La Risorsa Geneticadell’Olivo in Campania. Regione Campania – GiuntaRegionale, Naple, Italy. 160 pp.

QUIROS, C. F. (1975). Exine pattern of a hybrid betwenLycopersicum esculentum and Solanum pennelli. Journal ofHeredity, 66, 45–77.

RAFALSKI, A. (2002). Applications of single nucleotide polymor-phism in crop genetics. Current Opinion in Plant Biology, 5,94–100.

RAFALSKI, A., CHING, A., BHATTRAMAKKI, D., MORGANTE, M.,DOLAN, M., REGISTER, J. C., SMITH, O. S. and TINGEY, S. (2001).SNP markers in maize: discovery and applications. IXth

International Plant and Animal Genome Conference, SanDiego, CA, USA. W149.

RALLO, P., DORADO, G. and MARTIN, A. (2000). Development ofsimple sequence repeats (SSRs) in olive tree (Olea europaeaL.). Theoretical and Applied Genetics, 101, 984–989.

RALLO, L., BARRANCO, D., CABALLERO, J. M., DEL RIO, C., MARTIN,A., TOUS, J. and TRUJILLO, I. (2005). Variedades de Olivo enEspaña. Junta de Andalucia, Miniterio de Agricultura, Pesca yAlimentacion, Ediciones Mundi-Prensa, Madrid, Barcelona,México, 496 pp.

RANAMUKHAARACHCHI, D. G., KANE, M. E., GUY, C. L. and LI, Q. B.(2000). Modified AFLP technique for rapid genetic characteri-zation in plants. BioTechniques, 29, 858–866.

RESTA, P., LOTTI, C., FANIZZA, G., GODINI, A., MARIANI, R. andPALASCIANO, M. (2002). Use of AFLP to characterize Apulianolive varieties. Acta Horticulturae, 586, 73–77.

RICCIOLINI, C., BALDONI, L., ARCIONI, S. and ROSELLI, G. (2003).Primi studi di genomica dell olivo: analisi molecolare di popo-lazioni da incrocio intervarietale. Atti Convegno “Germoplasmaolivicolo e tipicità dell’olio”, Perugia, Italy. 207–210.

ROSELLI, G. (1979). Identificazione di cultivar di olivo da alcunicaratteri del polline. Rivista di Ortoflorofrutticoltura, 63,435–445.

ROSELLI, G. and SCARAMUZZI, F. (1974). Definizione dei termini dicultivar e clone. Accademia Nazionale di Agricoltura, Estrattodagli annali, Bologna, Italy. 2, 119–126.

ROSELLI, G., VENDRAMIN, G. and ROSSI, P. (1990). Patterns isoenzi-matici in cultivar di olivo. Atti XXXIV Convegno Società digenetica Agraria, Lecce, Italy.

ROSELLI, G., PETRUCCELLI, R., POLSINELLI, L. and CAVALIERI, D.(2002). Variability in five olive cultivars (Olea europea L.).Journal of Genetics and Breeding, 56, 51–60.

ROTUNDO, A. and MARONE, E. (2002). Il Germoplasma OlivicoloLucano. Unione Europea-Regione Basilicata-MiPAF, Potenza,Italy, 156 pp.

ROTONDI, A., MAGLI, M., RICCIOLINI, C. and BALDONI, L. (2003).Morphological and molecular analyses for the characterizationof a group of Italian olive cultivars. Euphytica, 132, 129–137.

ROTONDI, A., MARI, M., BABINI, A. R., COVONI, M. and CRISTOFERI,G. (2004). L’attitudine alla Propagazione e la CertificazioneGenetica e Sanitaria dell’Olivo in Emilia-Romagna. LaMandragora, Bologna, Italy. 117 pp.

RUBY, M. J. (1917). Recherches morphologiques et biologiques surl olivier et sur ses variétés cultivées en France. Annales desSciences Naturelles-Paris, 20, 1–287.

RUGINI, E. and BALDONI, L. (2003). Le biotecnologie. In: Olea -Trattato di Olivicoltura. (Fiorino, P., Ed.). Edagricole, Bologna,Italy. 145–167.

RUSSO, G., RESTA, P., CORONA, M. G. and CANIZZA, G. (2001).Analisi RAPD della diversità in varietà pugliesi di olivo. Atti VIConvegno Nazionale Biodiversità, Valenzano, Italy.

SANSAVINI, S. and PANCALDI, M. (2000). Brevettabilità e tutela dellidentità genetica delle nuove varietà. Frutticoltura, 2, 38–44.

SANZ-CORTES, F., PARFITT, D. E., ROMERO, C., STRUSS, D., LIÁCER,G. and BADENES, M. (2003). Intraspecific olive diversityassessed with AFLP. Plant Breeding, 122, 173–177.

SAVASTANO, L. (1939). Identificazione delle varietà di olivo. Annalidella Stazione Sperimentale per l’Olivicoltura, Pescara, Italy, I,99–102.

SEFC, K. M., LOPES, M. S., MENDONÇA, D., RODRIGUES DOS SANTOS,M., LAIMER DA CÂMARA MACHADO and DA CÂMARA

MACHADO, A. (2000). Identification of microsatellite loci inolive (Olea europaea) and their characterization in Italian andIberian olive trees. Journal of Molecular Ecology, 9, 1171–1173.

SEKER, M., DULGER, S. and KAYNAS, N. (2005). Investigation ofgenetic diversity in olive (Olea europaea L.) cultivars and typesusing isozyme analysis. Vth International Symposium on OliveGrowing, Izmir, Turkey. 105.


SHAHRIARI, M., NANKALI, A. and GHAREYAZI, B. (2005).Identification of Iranian olive cultivars by using RAPD andSSR markers. Vth International Symposium on Olive Growing,Izmir, Turkey. 117.

SINGH, L. and THOMPSON, B. D. (1961). The possibility of identifica-tion of vegetable varieties by paper chromatography offlavonoid compounds. Proceedings of the National Academy ofSciences of the USA, 77, 520–527.

SOLLER, M. and BECKMANN, J. S. (1983). Genetic polymorphism invarietal identification and genetic improvenement. Theoreticaland Applied Genetics, 67, 25–33.

STRIKIC, F., BANDELJ, D., PERICA, S., CMELIK, Z., SATOVIC, Z. andJAVORNIK, B. (2005). Genetic variation within the olive (Oleaeuropaea L.) cultivar ‘Oblica’ detected using AFLP markers. Vth

International Symposium on Olive Growing, Izmir, Turkey. 120.TAVANTI, G. (1819). Trattato Teorico – Pratico Completo sull’Olivo.

Volume I. Ed. Piatti. Florence. Italy.TESTOLIN, R., CIPRIANI, G., MARRAZZO, T., MARCONI, R. and

CIMATO, A. (2000). Identificazione genetica delle varietà diolivo del Garda. L’Informatore Agrario, 35, 71–76.

TOURNEFORT, J. P. (1719). Institutiones rei Herbariae. ExTypographia Regia. Parisiis. France. 695 pp.

TOUS MARTÍ, J. and ROMERO AROCA, A. (1993). Variedades delOlivo. Fundación “la Caixa”, Barcelona, Spain. 172 pp.

TREHANE, P., BRICKELL, C. D., BAUM, B. R., HETTERSCHEID, W. L. A.,LESLIE, A. C., MCNEILL, J., SPONGBERG, S. A. and VRUGTMAN, F.(1995). International Code of Nomenclature for Cultivated Plants.Quarterjack Publishing, Wimborne, UK. 175 pp.

TRIGUI,A., MSALLEM, M.,YENGUI,A., BELGUITH, H., KHECHEREM, J.,MELIÈNE, A., MALEK, S., BOUSSELMI, A., SAMET, A. and TRA-

BELSI, E.B. (2002). Oliviers de Tunisie: Catalogue des VariétésAutochtones et Types Locaux. Reliure d’Art, Sfax,Tunisia. 159 pp.

TRUJILLO, I., RALLO, L., CARBONELL, E. A. and ASINS, M. J. (1989).Isoenzymatic variability of olive cultivars according to theirorigin. Acta Horticulturae, 286, 137–139.

TRUJILLO, I., RALLO, L. and ARUS, P. (1995). Identifying olive culti-vars by isozyme analysis. Journal of the American Society forHorticultural Science, 120, 318–324.

UPOV (1985). Guidelines for the Conduct of Tests for Distinctness,Homogeneity and Stability: Olive. International Union for theProtection of New Varieties of Plants. Genéve, Switzerland.

ZILIOTTO, F., BARCACCIA, G., BALDONI, L. and TONUTTI, P. (2002).Identificazione e caratterizzazione di alcune cultivar di olivo.L’Informatore Agrario, 15, 115–118.

ZITO, F. (1932). L’esame biometrico del nocciolo delle olive comebase complementare di classificazione delle varietà.L’Olivicoltore, 24, 2–5.

ZITO, F. (1942). La classificazione delle razze di olivo coltivate.L’Olivicoltore, 4, 79–80.

Date post:	23-Nov-2023
Category:	Documents
Upload:	unipr
View:	1 times
Download:	0 times

The classification of olive germplasm – A review

Documents