Introduction

Montenegro is a small country located in the Balkan Peninsula and overlooking the Adriatic Sea. Grapevine growing in Montenegro dates back before the Roman period (Vuksanović, 1977). The climatic, soil and orographic conditions of southern Montenegro define two wine regions: the Montenegro Skadar Lake basin, encompassing the Podgorica, Crmnica, Rijecki, Bjelopavlicki and Katunski sub regions, and the Montenegrin seacoast region, including the Boko-Kotorski, Budvansko-Barski, Ulcinjski and Grahovsko-Nudolski sub regions. Ancient and probably autochthonous cultivars dominate the present grapevine assortment of these two regions, and very heterogeneous populations appear as a consequence of perennial growing. Furthermore, there are numerous local grapevine cultivars lacking heterogeneity in their population, which have been registered as autochthonous even though they were not definitively identified. Most of these cultivars can be recognized using OIV codes, but it is not enough for their reliable identification (Ulićević, 1966; Pejović, 1988; Maraš, 2000).

Current viticulture and wine production in Montenegro is based mainly on presumed autochthonous grapevine cultivars, such as Vranac, Kratošija, Krstač, and Žižak. The dominant cultivars for red wine production are Vranac and Kratošija. Significantly less prevalent is Krstač, used for white wine production, while no commercial wine is produced from Žižak. The presence of Muškaćela and Trojka is significantly lower.

The results of Calò et al. (2008) showed that Kratošija is a synonym of the Californian Zinfandel, the Italian Primitivo and the Croatian Crljenak Kaštelanski, and suggested a first-degree relationship with the Montenegrin Vranac. Contrary to Vranac, which has stable morphological traits and has undergone expansion in the recent years, Kratošija is very ancient and a lot of biotypes have been found within its population, not fully investigated yet. Because of its heterogeneity, Kratošija is less present alone in the vineyards, and is found mostly in combination with Vranac, which dominates in the assortment of red grape cultivars. The heterogeneity of Kratošija has been described by many authors (Stojanović, 1929; Bulić, 1949; Jelaska, 1954; Ulićević, 1966; Avramov, 1988; Pejović, 1988; Burić, 1995; Bozinovik et al., 1998; Maraš, 2000; Maraš et al., 2004). It is well known that the features of an old grapevine cultivar may change with the accumulation and fixation of mutations in the shoot apical meristem, giving rise to many biotypes within the population, even with noticeable morphological or phenological differences. In this context, our work had different purposes. Using simple sequence repeat (SSR) markers, we wanted i) to investigate the identity of almost all grapevine cultivars grown in Montenegro; ii) to determine which cultivars are likely native to Montenegro and which are common to neighbouring countries; iii) to ascertain the identity of presumed Kratošija biotypes collected in the experimental field of the Biotechnical Institute in Ljeskopolje and coming from different Montenegro private vineyards, where they are cultivated under many different names, depending on the area; and iv) to verify the correct identity of Vranac mother vines chosen for clonal selection process.

Materials and methods

1. Plant material

Seventy samples were selected for genotyping during the period from 2006 to 2010. The vines were collected from different viticultural areas of Montenegro (Table 1).

2. Microsatellite DNA analysis

DNA was extracted from young leaves obtained from cuttings, and genotyping was performed with 11 SSR loci routinely employed at CRA-VIT (Centro di Ricerca per la Viticoltura, Italy) for cultivar identification: VVS2 (Thomas and Scott, 1993); VVMD5, VVMD7, VVMD27 and VVMD28 (Bowers et al., 1996 and 1999); VrZAG62 and VrZAG79 (Sefc et al., 1999); ISV2, ISV3 and ISV4 (Crespan, 2003); and VMCNG4b9 (Welter et al., 2007), as described by Crespan et al. (2006). The varietal identification was achieved by comparing the obtained SSR profiles with available molecular databases and literature data.

3. Statistics on SSR data

To evaluate the discrimination power of the 11 SSRs used in this work, the following parameters were calculated: the polymorphic information content (PIC) and the combined non-exclusion probability for identity of unrelated and full-sib genotypes. SSR data were computed using 1418 unique profiles of Vitis vinifera L. cultivars present in the CRA-VIT molecular database, including the new genotypes found in the present work. CERVUS 3.0.3 software was used for calculations (Marshall et al., 1998; Kalinowski et al., 2007; http://www.fieldgenetics.com). Genotypes showing a single allele were considered as homozygous.

Results and discussion

Fourteen genotypes have been found within the 70 samples (Table 2). Using our set of 11 SSR markers, the mean PIC computed on 1418 unique profiles of Vitis vinifera L. cultivars was 0.8036; the estimated probability of identity to deal with unique genotypes was 3.38E-15 for unrelated genotypes and 9.52E-6 for full sibs. These parameters showed the significant reliability of this set of markers for varietal identification.

The identification work showed that five genotypes match well-known cultivars, namely Kratošija, Muscat blanc à petits grains, Muscat rouge de Madere (alias Moscato violetto), Plavina and Vranac. In particular, the accessions named Kratošija (plus an adjective), Bikača, Ljutica, Rehuljaca, Srednji Vranac, Velji Vranac, Vran, Vrancic, Vrancina and a wrongly named Vranac (id 44 in Table 1) all belong to the Kratošija cultivar, as previously supposed based on ampelographic descriptions (Maraš, 2000). Crni Krstač matches the Kratošija profile and is another clear misnomer. The first remarkable descriptions of Vranac and Kratošija cultivars have been given by Petar Plamenac in the Ampelography of Viala and Vermorel (1910). Later on, many scientists (Stojanović, 1929; Bulić, 1949; Ulićević, 1959 and 1966) state that Vranac is present only in Montenegro. Ulićević (1959) writes the following: "characteristic cultivar of Crmnica from the Skadar Lake viticultural area is Vranac, covering about 40% of the assortment of grape cultivars. As a matter of fact, this is the only viticultural area where it prevails and the only one where it is represented to a greater extent. It is in a proper sense a Crmnica cultivar". Nastev (1967) states that Vranac is a Montenegrin autochthonous grapevine cultivar mostly cultivated in the Skadar lake region (Crmnica), but also on the Montenegrin seacoast. This author declares that Vranac has been spread in the 1950s in Macedonia (experimental field of Butel). Nowadays, it represents the leading cultivar for red wine production in this country (Stajner et al., 2009). Ulićević (1959) states that Kratošija is almost as remarkable as Vranac. In the same paper he writes that Kratošija dominated in Montenegrin vineyards for over 70 years and that it often represented 90% of the assortment. Therefore, Kratošija is the main and probably the oldest Montenegrin grapevine cultivar. Ulićević (1959) argues that the Californian Zinfandel is identical to Kratošija and that Zinfandel likely originates from Montenegro and has been exported by Montenegrin emigrants. This was supported by the work of Calò et al. (2008), confirming that Kratošija has the same genetic profile as Zinfandel. Avramov (1988), Pejović (1988), Cindrić (1994), Cindrić et al. (2000), Burić (1995), Božinovik (1996), Maraš (2000) and Maraš et al. (2004) write that Vranac and Kratošija are autochthonous cultivars from Montenegro and that Kratošija has a very heterogeneous population. The wines of these cultivars are complementary: Kratošija usually provides a higher amount of total acids and sugars, while Vranac is richer in coloured substances. Bulić (1949) gives a description of Kratošija with synonyms (Gartosija, Grakosija, Kratkosija) from the nine municipalities of the Montenegrin seacoast. This cultivar is rarely found in the Dalmatia region, therefore, the author states that it has been possibly introduced to Dalmatia from Montenegro. By 2001, only about twenty living vines of Crljenak Kaštelanski had been found in Dalmatia (Maletić et al., 2009), confirming that this cultivar was not widespread in Dalmatia in the 19th and 20th century (Savić, 2003). Stojanović (1929) writes that the best vineyards in Montenegro are in the Skadar lake region (Crmnica), which is famous for its red wines that can reach a high quality, while the wines produced along the Montenegrin seacoast are similar to those from Dalmatia. This author states that Plavac mali, Dobričić and some others are the main cultivars grown in Dalmatia, whereas he does not mention Kratošija.

Ampelographic descriptions are available for all Kratošija accessions listed in Table 1 and grown at the Experimental Estate of the Biotechnical Institute in Podgorica - Ljeskopolje (Maraš, 2000). Some examples of ampelographic variability in Kratošija accessions are reported in Table 3 and Figure 1. The name of each Kratošija biotype is in connection with some of its particular characteristics. In particular, these biotypes showed a wide range of variability in grape yield, mainly dependent on bud fertility rather than cluster weight. Cluster weight was highly variable among accessions and correlated with bunch morphology; these traits showed to be stable within each biotype (Maraš, 2000). One mother vine was selected per biotype for the sanitary check-up and all showed the presence of viruses. Furthermore, additional investigation of the sanitary status of newly selected vines of Kratošija from all viticultural regions of Montenegro showed a high level of viral infection (Maraš et al., 2014).

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Figure 1. Examples of cluster variability among Kratošija biotypes (id number in parenthesis refers to table 1).

By putting together different pieces of information from the literature cited, and given the results of our study, we can conclude that Kratošija with all its biotypes has been grown in Montenegro for centuries. Given its presence in all the viticultural regions of this little country, the wide variability of the population, the impressive high number of different names, the close genetic relationship with Vranac, and its current presence (survival) in viticultural areas in Montenegro, we suggest that this area be considered a likely region for the origin and spreading centre of this cultivar. Therefore, the place of origin of this cultivar should be moved further south than the Dalmatian coast, unlike what has been previously suggested by Maletic et al. (2004).

The SSR profile of Muškaćela accessions matches that of Muscat blanc à petits grains. This outcome adds a new synonym to the long list of this very ancient cultivar, largely spread all over the viticultural areas in Europe and recently showed to be the main ancestor of the Muscat family (Cipriani et al., 2010). Trojka accessions have the same profile as Muscat rouge de Madere (alias Moscato violetto), another important member of the Muscat family (Di Vecchi-Staraz et al., 2007; Lacombe et al., 2013), but the berry colour is different (it is white in our analyzed Trojka cultivar). Muscat rouge de Madere is a progeny of Muscat blanc à petits grains and Sciaccarello (Lacombe et al., 2013) and has little importance. Supposed to be native to Portugal, it does not have any agronomic relevance. Actually, it is not included in any official list of grapevine cultivars authorized for cultivation in 24 national catalogues (Lacombe et al., 2011). It is considered a grape for enthusiasts (Galet, 2000) and is grown mostly in germplasm repositories. That is why it was a surprise to find it among Montenegrin cultivated grapevines. Plavina is a progeny of Kratošija and Verdeca (Cipriani et al., 2010) and is an important cultivar in the sub region of North Dalmatia. All mother plants chosen for clonal selection of Vranac (NCV 62/13; NCV 14/39; NCV 15/13; NCV 62/7; NCV 62/8) showed to be correctly identified.

Although in literature Bioka is considered as a synonym of Krstač in Crmnica (Ulićević, 1959), SSR data showed that it has a different molecular profile. Therefore, Bioka is another cultivar. We have discovered that Bioka shares the same genotype as the Italian Francavidda, a cultivar registered in the Italian Catalogue and grown in Apulia, the Italian region facing Montenegro on the other side of the Adriatic Sea (http://catalogoviti.politicheagricole.it). Moreover, Bioka has the same SSR profile as the Croatian Zlatarica Vrgorska (Zdunić et al., 2013). Therefore, these three cultivars represent a new case of previously unknown synonyms grown in different countries facing the shores of the Adriatic Sea (Dalmatia and Apulia).

Comparing our data with the European Vitis database (http://www.eu-vitis.de/index.php), we found one match at all seven common SSR loci between the Montenegrin Kadarun and a Kadarun accession from Croatia (HRV041-SA#11-95) lacking the true-to-type validation. In the Vitis International Variety Catalogue (VIVC, http://www.vivc.de), Kadarun is reported as a Turkish cultivar.

The red berry Razaklija accession from the Skadar Lake region (Doljani), a known synonym of the Serbian Drenak crven, matches the SSR profile of Crven Drenok (Vitis-WBC, Western-Balkans Vitis Database, http://vitis.atcglabs.com). Table grapes in Montenegro only accounts for 5%, of which 95% is the Razaklija cultivar (Drenak crven) from both the Skadar Lake and the seacoast region (Ulićević, 1959). Many authors from Yugoslavia consider that Razaklija originates from Asia Minor (Ulićević, 1966). It is not known how and when it arrived in Montenegro, Macedonia and other countries and how it spread.

Crna Loza, Čubrica, Krstač, Lisičina, Razaklija crna and Žižak show unique SSR profiles. Crna Loza was considered as a Kratošija synonym, but our analysis shows a different SSR profile for this cultivar. Based on SSR allele sharing at all analyzed loci, Crna Loza could be another offspring of Kratošija, but further research should be done to confirm this hypothesis.

Ulićević (1959) states that Čubrica is a cultivar for red wine production and is represented in very small percentage in the vineyards of the Podgorica sub region (Doljani and Kuci). The sample of Čubrica from Kuci has a DNA profile confirming that it is an independent cultivar.

Ulićević (1959) writes that in the Skadar Lake region, in the sub region of Podgorica (Beri and Doljani), Krstač was dominant among the white grapevine cultivars used for white wine production, produced in small amount in that period. In the sub region of Podgorica, it represents only 10% of the vineyard area, mostly in Beri and Doljani, and very little in the Ljesanska district. Krstač is believed to be autochthonous of Montenegro and probably originated in Beri (near Podgorica); its name comes from the look and shape of the bunch that resembles a cross (Viala and Vermorel, 1910; Vujović, 1956). Žižak (or Žižak bijeli after Bulić, 1949) is considered another autochthonous cultivar of Montenegro and its origin is unknown. Individual specimens can be found nearby Podgorica, but it is mostly grown on the Montenegrin seacoast (Boko-Kotorski sub region), where it is used for dessert wine (Ulićević, 1959).

About Lisičina, Stojanović (1929) reports the use of this cultivar for white wine production in Montenegro. Bulić (1949) and Ulićević (1959) found Lisičina (synonyms are Lisica, Ružica, Sjemerava, Sjeruša) in Montenegro around Bar, Ulcinj, Podgorica and Virpazar (Crmnica). The VIVC catalogue reports that Lisičina is a synonym of the Croatian Lisica. According to Nastev (1967), Lisičina is a wrong synonym of the Plovdina (Pamid) cultivar. Accordingly, the SSR profile of our Lisičina sample is different from that of the Bulgarian Pamid (Hvarleva et al., 2004; Dzhambazova et al., 2009).

Razaklija crna (black), having a dark blue coloured skin, was found only in Kuci (sub region of Podgorica) in old vineyards and under this name. It has to be noted that this name indicates a table grape cultivar originated by crossing Razaklija Rossa and Vranac (performed by Ulićević M. at the Biotechnical Institute in Podgorica). Based on SSR allele sharing at all analyzed loci, Razaklija crna could really be a progeny of Drenak Crven, but our SSR data rule out Vranac. Further data are required to validate this partial pedigree.

Conclusion

Seventy accessions belonging to the Montenegrin grapevine germplasm were analyzed using SSR markers. Fourteen genotypes have been found, three of them being native to Montenegro (i.e., Vranac, Krstač and Žižak). Kratošija has been recently recognized as an international cultivar, being a synonym of Italian Primitivo, Californian Zinfandel and Croatian Crljenak Kaštelanski, whose origin has been widely investigated. Given the large number of different names used in Montenegro for Kratošija and the wide morphological variability of Kratošija biotypes, we propose here, for the first time, Montenegro as the best candidate country for the origin and spreading centre of this group of synonyms. Crna Loza, Čubrica, Lisičina and Razaklija crna are cultivars likely native to Montenegro. Muškaćela and Trojka, considered autochthonous of Montenegro, are clearly of different origin. We discovered that Montenegrin Bioka, Italian Francavidda and Croatian Zlatarica Vrgorska represent a new group of synonyms.

Overall, our genotyping study gave us the opportunity to check the available information based on ampelographic descriptions and comparisons, providing a more stringent and sometimes new perspective on the origins and relatedness of grapevine cultivars in Montenegro.

Further analyses should be done in order to clarify the genetic relationships among Montenegrin grapevine cultivars as well as the degree of relatedness and origin of commercially important Montenegrin cultivars.

The sharing of common cultivars among neighbouring countries was expected as a natural consequence of easily vegetative propagation of grapevine, but only recently we have started to get accurate information on their identity and the present research contributes to shed some light on this topic.

Acknowledgments: This research was supported by the following projects: "Variability of Kratošija cultivar in Montenegrin viticultural regions" and "Clonal selection of Kratošija cultivar" funded by the Ministry of Science of Montenegro; "Identification and characterization of Montenegrin autochthonous grapevine cultivars" and "Clonal selection of autochthonous grapevine cultivars" funded by Company 13 jul Plantaze and the Ministry of Agriculture of Montenegro; and "IDENTIVIT/ASER" funded by the Italian Ministry of Agricultural, Alimentary and Forestry Policies. Special thanks to Dordjije Rajkovic for useful suggestions and support and to Angelo Costacurta for initiating us to cultivar characterization and identification.

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