This is an author version of the contribution published on:
Questa è la versione dell’autore dell’opera:
Schneider A., Raimondi S., Pirolo C.S., Torello Marinoni D., Ruffa P., Venerito P., La
Notte P. (2014) Genetic characterization of grape cultivars from Apulia (southern
Italy) and synonymies in other mediterranean regions, 65(2): 244-249.
doi:10.5344/ajev.2013.13082
The definitive version is available at:
La versione definitiva è disponibile alla URL:
http://www.ajevonline.org/content/65/2/244
Genetic Characterization of Grape Cultivars from Apulia (Southern Italy) and
Synonymies with other Mediterranean Regions
Anna Schneider1*, Stefano Raimondi1, Costantino S. Pirolo2, Daniela Torello Marinoni3, Paola Ruffa1, Pasquale Venerito4, Pierfederico Lanotte4,5
1 National Research Council of Italy (CNR), Institute of Plant Virology – Support Unit Grugliasco, Via L. da Vinci 44, 10095 Grugliasco (Italy)
2 University of Bari, Department of Soil, Plant and Food Sciences (DISSPA), Via Amendola 165, 70126 Bari (Italy)
3 Department of Agricultural, Forest and Food Sciences (DISAFA), Largo Paolo Braccini 2, 10095 Grugliasco (Italy)
4 Research, Experimentation and Education Centre in Agriculture "Basile Caramia" (CRSFA), Via Cisternino 281, 70010 Locorotondo (Italy)
5 National Research Council of Italy (CNR), Institute of Plant Virology – Support Unit Bari, Via Amendola 165, 70126 Bari (Italy)
*Corresponding author: email: a.schneider@ivv.cnr.it, phone: 0116708745, fax: +39-0116708658
Acknowledgments
This work has been partially funded by the Apulia Region with the project “Clonal and sanitary selection of table, wine grape and rootstock varieties" (SELMIVI). Authors are grateful to Frances Cooper for language revision.
Short version of title: Apulian Cultivars: Genetic Characterization and Synonyms
Abstract: Forty-five grape accessions, traditional and historically mentioned in Apulia (south
eastern Italy), were genotyped at 13 microsatellite (SSR) markers and observed for their morphological features with the aim of characterizing and identifying the local grape diversity relevant for economic or historical significance, or for endangered germplasm conservation. Twelve of the 45 accessions examined were found to be synonyms or somatic mutants, leaving 33 distinct genotypes. Attempts were then made to verify the true identities of the accessions investigated and to determine their appropriate denominations. This entailed comparing them with published allelic profiles and morphological features of cultivars from Apulia and from surrounding areas linked historically to the region.
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While confirming the identity of the major Apulian cultivars, further matches with varieties from other Mediterranean regions were revealed. Approximately half of the Apulian cultivars investigated were found to have a foreign counterpart mainly along the Adriatic Sea (Croatia), in Greece, or in other southern Italian regions. The new synonymies found with cultivars traditional to other areas shed light on the migration of cultivars following the settlement of colonies and the historical establishment of Mediterranean trade routes.
Key words: microsatellites; SSR markers; Vitis vinifera; variety identification; ampelography.
Introduction
Apulia is a region in southeastern Italy, jutting between the Adriatic and Ionian Seas towards Albania and the western coast of Greece. Grapevines were already cultivated in Apulia in the Bronze Age. In the eighth century B.C. Greek colonists settled on the Ionian coast of the Apulia and developed the emporion, trading stations for unloading goods, including wine. In Roman times, Apulia continued to produce and trade wheat, wine and olive oil so that strategic ports were built to connect the area with the eastern Mediterranean. In the Middle Ages, the sea played a key role: the Italian cities of Venice and Genoa, both trading in Apulian ports, increased their influence. A further link with eastern Meditterranean was established during the Crusades. In the early and middle 1800s, a large number of grape varieties were inventoried in the region. During the great expansion of viticulture to 300,000 300,000 hectares in the second half of the 19th century, because of the phylloxera crisis in France, the main varieties planted were Uva di Troia, Primitivo, and Negro Amaro.
Viticulture now occupies about 119,000 ha in Apulia (http://www.agri.istat.it), with 59 wine-grape cultivars currently authorized to be grown (http://www.regione.puglia.it), including 22 local, historical varieties. Nevertheless, there are still numerous minor varieties grown in the region, in moderate or very limited quantities, that several programs aim to recover and possibly exploit for 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66
commercial use. In the last few years, six local ancient varieties were officially registered in the Italian Grape Variety Catalogue (La Notte et al. 2011-2012). Other threatened grapes maintained in field collections are under study.
Microsatellite or Simple Sequence Repeat (SSR) genotyping, together with vine morphological descriptions, are the most widely used approaches to characterize and identify grape varieties (Sefc et al. 2009; This et al. 2011) and comprise a valuable tool for the management of collected genetic resources (Ortiz et al. 2004; Laucou et al. 2011). In grapevines, nuclear SSRs reveal highly diverse allelic patterns, codominant inheritance, and high repeatability and interlaboratory reproducibility, due to the simplicity of allele score adjustment (This et al. 2004). SSR profiles provide direct comparison of genotype similarity at common loci, while plant morphology observations support synonymy hypotheses and can be related to historical references.
Twenty grape varieties from Apulia had been characterized by the six microsatellites of international use (Zulini et al. 2002); the genetic origin of the local Malvasia nera was also demonstrated (Crespan et al. 2008a); among a large set of Italian grape varieties, several cultivars from Apulia were investigated by SSRs with three to five nucleotide core repeats (Cipriani et al. 2010). In the present study, 45 traditional, historical cultivars from Apulia were examined for their morphological and genetic profiles. Despite their economic significance in the region, modern table grape varieties issued form breeding programs were excluded from this study, which focuses on the ancient germplasm.
The study aimed to help catalog, characterize and identify local grape varieties of relevance for their economic or historical significance, and for endangered germplasm conservation.
Since intense cultivar migration has been demonstrated for the prominent grape Primitivo (Maletić et al. 2004; Calò et al. 2008), published allelic profiles of cultivars from neighboring and surrounding areas were compared with their Apulian counterparts, in order to discover any new synonyms and provide historical evidence on grape circulation.
67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92
Materials and methods
Most of the investigated accessions are maintained in the grapevine collection of the Research, Experimentation and Education Centre in Agriculture "Basile Caramia" (CRSFA) at Locorotondo (Bari, Apulia). Several accessions (including presumed bud sports) were located in commercial vineyards. The list of the 45 accessions examined is in Table 1.
DNA was extracted from fresh shoot tissues sampled in the field, following the procedure described by Thomas et al. (1993) with some modifications. Samples were genotyped using a set of 13 SSR loci, nine of them developed under the projects GenRes 081 and GrapeGen06 (http://www1.montpellier.inra.fr/grapegen06/page_summary/summary.php) as common markers for international use. The 13 markers used were: VVMD5, VVMD7, VVMD25, VVMD27, VVMD28, VVMD32, VVMD36, VVS2, VrZAG21, VrZAG62, VrZAG64, VrZAG67, VrZAG79 ( for detailed
information on markers see www.eu-vitis.de/index.php and
www.vitisdb.it/descriptors/loci_descriptors). PCR products were then analyzed on a 3130 Genetic Analyzer (Applied Biosystems, Foster City, Calif., USA). Data were processed using GeneMapper Software (ver. 4.0; Applied Biosystems) and alleles were defined by their size in base pairs, by comparison with the standard size (GeneScan-500 LIZ, Applied Biosystems).
To evaluate the markers used, the following statistical parameters were calculated from data on the 33 genotypes that were found to be unique, in the set of 45 accessions examined, using IDENTITY software (ver. 4.0; Centre for Applied Genetics, University of Agricultural Sciences, Vienna): numbers of alleles, observed and expected heterozygosity, estimated frequency of null alleles and probability of identity (PI) for the analyzed loci.
The SSR profiles obtained were then compared with those provided by Vitis genetic databases, standardizing data to common true-to-type cultivars: the European Vitis database (http://www.eu-vitis.de/index.php), the nuclear microsatellite Greek Vitis database (http://gvd.biology.uoc.gr/gvd/contents/index.htmse), the National Clonal Germplasm Repository from UC-Davis (NCGR-Davis, http://www.ars.usda.gov/Main/docs.htm?docid=13743), Pl@nt 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118
Grape from French Catalogue (http://plantgrape.plantnet-project.org/) as well as the 735 unique
Vitis vinifera microsatellite allelic patterns developed by the CNR - Institute of Plant Virology
(unpublished data). Bibliographic sources presenting nuclear microsatellite profiles of traditional grapes from other southern Italian regions, or other European regions surrounding or related to Apulia by presumed historical relationships were also used for genetic data comparison. SSR loci such as VVMD36, VrZAG21, VrZAG64 and VrZAG67, not included in the international lists promoted by OIV and GrapeGen06, were analyzed to cover published allelic patterns more completely. In the comparison process at least the six international markers, but generally from 7 to 10 and often more were in common. Probability of identity for all loci (PItot) was calculated within the set of 735 Vitis vinifera genotypes analyzed by the CNR - Institute of Plant Virology, for the 6, 7, 10 nSSR loci found to be in common with published profiles and for all the 13 analyzed loci. The varieties investigated were described in terms of vine morphological features, according to the major OIV descriptors selected from the European Vitis database (http://www.eu-vitis.de/index.php) and compared with published references.
Results and discussion
Most of the investigated varieties have been mentioned in historical references from the early nineteenth century. Approximately one-third have a national or regional relevance; most are local or neglected, but often scattered in throughout the region (Table 1).
Among the 45 accessions analyzed, 33 unique genotypes were found. Statistical parameters on these unique profiles indicated a number of alleles/locus ranging from 6 to 10 (8 on average), a level of observed and expected heterozygosity similar to that found in previous studies, an estimated frequency of null allele positive only for 4 loci out of 13, and a very low overall loci probability of identity (4.93 x 10-16). It can therefore be assumed that the identical profiles found are mutants or synonyms (Table 2). The two Baresana accessions differing for grape color were confirmed to be somatic mutations, as were the four Regina clones distinguishable for berry shape 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144
and size: broad elliptic in the Regina bianca, narrow elliptic in the mennavacca, elongated elliptic in the pizzutella, significantly larger in the mennavaccone. Similarly, sport mutations never described before are the two Somarello, showing black and red grapes. The other equal genotypes were identified as “internal” synonyms: that is, synonyms within the set of the varieties analyzed. Most of these synonyms refer to minor, neglected varieties. Montonico-Pagabebiti-Uva della scala is a prolific grape variety that is widespread under different names (and therefore not recognized) in commercial vineyards from central to southern Italy. Pampanuto and Verdeca, although evidence of their synonymy has already been reported, are still erroneously registered as distinct cultivars in the Italian catalogue, thus resulting as duplicates.
The 33 unique genetic profiles at 13 SSR markers and profiles from two international varieties used as references are in Supplemental Table 1. For the accessions provided of genetic and/or morphological references, hence identified, the varietal names according to the Italian Grape Variety Catalogue are indicated. A morphological profile of these grapes can be found (http://catalogoviti.politicheagricole.it/ricerca.php), except for the recently registered Antinello, Minutolo, Marchione, Maresco and Somarello rosso (descriptions reported in Lanotte et al. 2011-2012). For Palumbo, Passera rossa, Santa Teresa, Uva Attina, and Uva di Angela, no genetic or morphological matches were found among the Italian grape cultivars. The examined accession Fiano di Avellino (belonging to variety Fiano B.) showed three alleles at VrZAG67, 139, 150 and 153 bp in size, a condition not uncommon in grapes and explained by the occurrence of a chimeric structure.
Comparison of the allelic profiles of the Apulian cultivars with those of varieties from other regions, while confirming known identities and synonyms, also revealed new matches, significant for reconstructing the migration of varieties over time (Table 3). The general probability of identity (PItot) calculated on the 735 Vitis vinifera genotypes (accessions from throughout Italy) included in the nSSR database of the CNR - Institute of Plant Virology (unpublished data) ranged from 5,57 x 10-9 for accessions with 6 loci in common with other publications, to 4.96 x 10-14 for those with 10 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170
loci in common. It is thus likely that identical microsatellite profiles correspond to synonyms or to mutants. Comparison of plant morphological features (photos and descriptions) of presumed synonyms corroborated the genetic data, except for Pampanuto (Verdeca)/Lagorthi, where the genetic profile of Lagorthi from the Greek Vitis Database indeed matched with that of Pampanuto (alias Verdeca) from Apulia, whose phenotype differed from Lagorthi shown elsewhere (Kotinis 1985). The NCGR-Davis database reports three different SSR profiles relating to the name Lagorthi.
The identity of the grapes mainly grown in Apulia, such as Aglianico, Aleatico, Malvasia nera, Negro amaro, Primitivo, Uva di Troia, was confirmed by comparison with published genetic profiles. The association of the unspecified Malvasia bianca (historically mentioned in all southern Italian regions) with Malvasia del Chianti from Tuscany, Maraština from Croatia, and Pavlos from Greece reinforces the ancient origins of this cultivar, widespread along the Adriatic and Ionian coasts and one of the source of Italian grape genetic diversity (Crespan et al. 2007; 2008a).
In addition to the synonyms Malvasia/Maraština/Pavlos and Primitivo/Crljenak/Kratošija, our findings demonstrated further correspondences between grape cultivars traditional to Apulia and varieties from other Mediterranean coastal regions, especially from Croatia and Greece. Half of the genotypes examined were found to be synonyms of grapes from Slovenia or Croatia (Bianco d’Alessano/Beretinjok and Topol; Francavidda/Zlatarica vrgorska; Maruggio/Posipica, Bratkovina, Popetre and Stradunska; Santa Teresa/Frmentum), from Greece (Baresana/Kolokythas, Korithi and Roditis lefkos; Pampanuto/Bampa-Hasan and Lagorthi; Sgarraparete/Karystino), or from other southern Italian regions (Baresana/Cessalà; Bianco d’Alessano/Iuvarello; Montonico bianco/Greco bianco; Pampanuto/Bracaù; Negro amaro/Zagarese; Susumaniello/Nerello; Uva di Troia/Summariello) (Table 3). Although no correspondences were found among grapes from Albania (Ladoukakis et al. 2005), Macedonia (Štajner et al. 2009), or the Maltese islands (Giannetto et al. 2010), the synonymies revealed suggest that intense movement of grape material occurred along the trade routes of the Ionian and Adriatic Seas, with a remarkable influence on the 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196
assortment and evolution of Apulian varieties. This reflects what can be assumed for the entire Italian peninsula, whose grape cultivars appeared by far the most admixed among geographic groups (Bacilieri et al. 2013). As for Apulia, it is not yet clear whether the exchange/introduction of materials occurred during the intense trade of the Venetians in the modern age, or whether it should be traced back to Greek colonization. Grape movements also occurred across southern Italian regions, from Apulia to Sicily or vice versa. Moreover, the synonymies revealed enable materials masked by local names to be identified, such as Cessalà and Bracaù analyzed in Sicily (Carimi et al. 2010), which have now been identified with Baresana and Pampanuto/Verdeca, respectively.
The table grape Baresana (Baresana meaning “from Bari”, the largest Apulian city), attested in the region since the mid-19th century, is considered a historical variety in Apulia, where it is currently the object of local valorization. The white and the pink-skinned sports have both long been present in southern Italian regions. Its white synonyms Kolokythas, Korithi and Roditis Lefkos are widespread in many Greek inland regions and islands, demonstrating the historical importance of the variety. The pink-skinned variant does not seem to occur in Greece: the cultivar Roditis Rs (with pink-skinned grape) included in the Greek catalogue depicted in Kotinis (1985) and described by Branas and Truel (1965) is a variety different from Baresana.
Bombino bianco (Trevolina in Slovenia) is regarded as one of the recurrent genitors in the origins of traditional grapes from Croatia and Apulia (Lacombe et al. 2013), giving rise in the latter region to Uva di Troia (Bombino bianco X Bouteillan, alias Quagliano), Moscatello selvatico (Bombino bianco X Muscat of Alexandria), and Impigno (Bombino bianco X Bouteillan) (Cipriani et al., 2010). In addition to confirm these findings, our data further suggest Bombino nero and Palumbo as other varieties likely belonging to the Bombino kin-group, sharing half of their alleles at each SSR locus analyzed.
Sangiovese has been shown to possibly originate in Apulia (Bergamini et al. 2013). Among the cultivars examined in this study, Susumaniello (as also reported in Crespan et al. 2008b) Notardomenico, and Uva di Angela are likely related to Sangiovese by half-kinship.
197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222
Cinsaut, the ancient variety traditional to Provence (southern France), also occurs in southern Italian regions: Ottavianello is the official name in Apulia, where it is also present as Impigno rosso, while our findings show that, in Sicily, it is concealed beneath the name of Grecaù. The question thus arises as to whether Cinsaut reached southern Italian regions from the French coast (or from the related north African colonies), or whether it spread across southern Italy on its way from the eastern Mediterranean.
Four cultivars, Palumbo, Passera rossa, Uva Attina and Uva di Angela, did not show any matches neither in SSR allelic nor in vine morphological profiles. Since they are likely unique genotypes, they are worth maintaining as endangered genetic resources, currently neglected in Apulia. Fiano minutolo (officially named Minutolo) must be clearly distinguished from the more popular Fiano di Avellino from Campania. Finally, few cases of variety misnaming (Magliocco instead of Notardomenico and Susumaniello instead of Somarello) are reported.
Conclusions
Data from genotyping several historical grape varieties from Apulia, including minor and neglected cultivars, are presented here for the first time. Particular efforts were made to determine the true identity of the accessions investigated and their appropriate denominations. Cultivars from surrounding areas or from regions related to Apulia by historical relations and analyzed in other studies were considered for this purpose.
Comparison of nuclear SSR profiles from bibliographic sources and from available molecular databases was confirmed as a modern and reliable tool for determining identical genotypes and synonyms. However, at least basic morphological information is crucial in order to pair the genetic profiles obtained with the true-to-type variety and appropriate denominations. Published references for variety morphological profiles were therefore checked, and are reported together with genetic data. 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247
New synonymies, sometimes unexpected, found among grapes traditional to other areas, shed some light on the migration of grapes following the development of colonies and the establishment of trade routes in ancient times. Links with Greece and the Adriatic coastal areas, and to a lesser extent with other overseas parts of the central Mediterranean, were evident in the traditional grape varietal assortment of Apulia, a Mediterranean land jutting out into the sea.
Table captions
Table 1. The 45 grape cultivars/accessions examined in this study.
Table 2. Matching genetic profiles within the set of accessions examined.
Table 3. Synonyms from other regions revealed by matching with genetic and morphological profiles.
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Table 1. The 45 grape cultivars/accessions examined in this study: color, use (winegrape, W; table grape, T), cultural area (Italian province), cultural importance, and current registration in the Italian catalogue (catalogoviti.politiche agricole.it/home.php)
N. Cultivar/accession Berry
colora Use Cultural area (ItalianProvince)b importanceCultural c registrationCurrent
1 Aglianico N W FG N yes
2 Aleatico N W FG – BA – BR – LE - TA N yes
3 Antinello B W BA L yes
4 Baresana bianca B T BA – BR - LE L yes
5 Baresana rosa Rs T BA – BR - LE L yes
6 Bianco d’Alessano B W BA – BR -TA R yes
7 Bombino bianco B W FG - BA N yes
8 Bombino nero N W FG - BA L yes
9 Cuccimaniello N W BR - TA L no
10 Fiano d’Avellino B W FG - BA N yes
11 Fiano minutolo B W BA – BR -TA L yes
12 Francavidda B W BA – BR -TA L yes
13 Impigno B W BA – BR -TA L yes
14 Impigno rosso N W,T BA – BR -TA NE no
15 Malvasia bianca B W FG – BA – BR – LE - TA N yes
16 Malvasia nera N W BR - LE R yes
17 Marchione B W BA – BR – TA L yes
18 Maruggio B W BA – BR – TA L yes
19 Montonico bianco B W FG N yes
20 Moscatello selvatico B W FG - BA L yes
21 Moscato reale B W FG - BA L yes
22 Negro amaro N W BR – LE - BA R yes
23 Notardomenico N W-T BA – BR -TA L yes
24 Pagadebiti B W FG - BA NE no
25 Palumbo B W BA – BR -TA NE no
26 Pampanuto B W BA - FG L yes
27 Passera rossa Rs W BR NE no
28 Primitivo N W BA – TA - LE N yes
29 Regina bianca B T BA – TA - FG N yes
30 Regina bianca mennavacca B T BA L no
31 Regina bianca pizzutella B T BA L no
32 Regina bianca mennavaccone B T BA L no
33 San Nicola N W TA NE no
34 Santa Teresa B W TA NE no
35 Sgarraparete Rs W BR - TA NE no
36 Somarello nero N W BA – BR – TA L no
37 Somarello rosso R W BA – BR – TA NE yes
38 Susumaniello N W BA – BR – TA L yes
39 Uva Attina B W BA – BR – TA NE no
40 Uva Carrieri B W BA – BR – TA NE no
41 Uva della scala B W BA – BR – TA NE no
42 Uva del monaco B W FG - BA NE no
43 Uva di Angela B W BR - TA NE no
44 Uva di Troia N W FG R yes
45 Verdeca B W BA – BR -TA R yes
351 352 353
aAccording to OIV descriptor 225: B = white, R = red, Rs = Rose, V = dark red-violet, N = blue-black bApulian provinces: Foggia (FG), Bari (BA), Brindisi (BR), Lecce (LE), Taranto (TA)
cN = national (also present in Italian regions other than Apulia), R = regional; L = local; NE = neglected/risk of
extinction
Table 2. Matching genetic profiles within the set of accessions examined.
Accession name Synonyms Mutants
Baresana bianca Baresana rosa
Maruggio (Maresco) Uva del monaco Montonico bianco PagadebitiUva della scala
Notardomenico San Nicola
Palumbo Uva Carrieri
Pampanuto Verdeca
Regina bianca
Regina bianca mennavacca Regina bianca mennavaccone Regina bianca pizzutella
Somarello rosso Somarello nero
Table 3. Synonyms from other regions revealed by matching with genetic (published genotypes, geographical area, and related references) and morphological (description and related references) profiles
aDatabases: GrVDB: Greek Vitis Database (http://gvd.biology.uoc.gr/gvd/index.htm); IVV-DB: the CNR-Institute of Plant Virology database (unpublished);
EuVDB: European Vitis Database (http://www.eu-vitis.de/index.php); NCGR: National Clonal Germplasm Repository-Davis (http://www.ars.usda.gov/Main/docs.htm?docid=13743); Pl@ntgrape (http://plantgrape.plantnet-project.
Accession name Genetic matching Morphological matching
Baresana bianca
Cessalà (Italy, Sicily): Carimi et al. 2010 Kolokythas lefkos (Greece): GrVDB Korithi aspro (Greece): Lacombe et al. 2013 Korithi lefko (Greece): GrVDB
Roditis lefkos (Greece, Peloponnese): GrVDB
Baresana: Branas and Truel 1965 Kolokythas lefkos: Kotinis 1985 Korithi lefko: Kotinis 1985 Bianco d’Alessano
Beretinjok (Croatia): Zdunić et al. 2013 Iuvarello (Italy, Calabria): IVV-DB Topol (Croatia): EuVDB (HRV041)
Tikvar bijela (synonym of Topol and Bratinijak): Bulič 1949
Francavidda Kadarun IV, Surac VI (Bosnia and Herzegovina): Tomić et al. 2012 Kadarun (Bosnia and Herzegovina): Žulj Mihaljevć et al. 2013 Zlatarica vrgorska (Croatia): EuVDB (HRV041), Zdunić et al. 2013
Kadarun crni (Bulič 1949) a distinct, black grape Impigno rosso Cinsaut (France): IVV-DB, Pl@ntgrapeGrecaù (Italy, Sicily): Carimi et al. 2010 Cinsaut: Pl@ntgrape
Maruggio (Maresco)/Uva del monaco
Bratkovina bijela (Croatia): EuVDB (HRV041) Bratkovina Blatska (Croatia): Zdunić et al. 2013 Popetre (Slovenia): Štajner et al. 2011
Posipica (Croatia): EuVDB (HRV041)
Stradunska (Croatia): Žulj Mihaljević et al. 2013
Bratkovina Bijela (syn. Pošipica): Bulič 1949
Montonico bianco/
Pagadebiti/Uva della scala Greco bianco del Pollino (Italy, Calabria): Costacurta et al. 2004 Moscatello selvatico Moscato di Barletta (Italy, Apulia): Zulini et al. 2002
Negro amaro Zagarese (Italy, Campania): Costantini et al. 2005 Notardomenico/San Nicola Magliocco (Italy): Lacombe et al. 2013 (misnomer) Pampanuto/Verdeca Bampa-Hasan (Greece, Ipiros): GrVDBBracaù (Italy, Sicily): Carimi et al. 2010
Lagorthi (Greece, Ionian Islands-Akaia-Arkadia): GrVDB; Lacombe et al. 2013, NCGR
Bampa-Hasan (Baba-hassan): Kotinis 1985 Santa Teresa Frmentum (Croatia): Zdunić et al. 2013Frmentun (Croatia): EuVDB (HRV041)
Sgarraparete Barbarossa (Italy, Apulia): Zulini et al. 2002Karystino (Greece): GrVDB, Lacombe et al. 2013
Japudzak (Montenegro): EuVDB (YUG027) Karystino: Kotinis 1985 Somarello nero/rosso Susumaniello (Italy): Di Vecchi Staraz et al. 2007; Lacombe et al. 2013 (misnomer)
Susumaniello/Cuccimaniello Greco nero di Cosenza (Italy, Calabria): Di Vecchi Staraz et al. 2007Nerello (Italy, Calabria): IVV-DB Uva di Troia Summariello (Italy, Campania): Costantini et al. 2005
354 355
357 358 359
