Identification by SNP Analysis of a Major Role for Cayetana Blanca in the Genetic Network of Iberian Peninsula Grapevine Varieties

Several Portuguese cultivars are siblings.

A total of six trios including Cayetana Blanca as a parent were found (Table 2, Figure 1). Five of them have the same variety as the other parent, a well-known Portuguese variety, Alfrocheiro Preto, which gave place to five varieties presently cultivated in Portugal: Periquita, Cornifesto, Malvasia Preta, Mouraton, and Camarate (Figure 2). These cultivars have been described by Galet (2000) and have different names (synonyms) in different locations of Portugal and/or Spain (Supplemental Table 2).

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Figure 1

Genetic relationships found for Cayetana Blanca. Solid lines indicate full pedigree (trio parents and offspring), while dashed lines link cultivars sharing at least one allele per locus (possible parent-offspring relationship).

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Figure 2

Map of the Iberian Peninsula showing the different locations of the offspring and other cultivars related to Cayetana Blanca: (1) Alfrocheiro Preto, (2) Periquita, (3) Cornifesto, (4) Malvasia Preta, (5) Mouraton, (6) Camarate, (7) Antão Vaz, (8) Rabo de Ovelha, (9) Cayetana Blanca, (10) Jaén Tinto, (11) Cigüente, (12) Castellana Blanca, (13) Plateado, (14) Puerto Alto, (15) Rocia, (16) Castillo de Arcos, (17) Água Santa.

Alfrocheiro Preto is a red grape, one of the most widely planted (220 ha) in the Dão region (Portugal), and also found in the vineyards of Alentejo, Ribatejo, and Bairrada. Periquita (its current Portuguese name is Castelão) is a red wine cultivar known in Terras do Sado and throughout southern Portugal. Wine produced from this grape is usually quite tannic and fruity. Cornifesto is a black grapevine cultivar planted in the Douro and Alto Trás-os-Montes regions in northern Portugal. Malvasia Preta is an autochthonous red winegrape from the Douro and Dão regions (Portugal) and is also known as Moreto do Dão. The cross originating this variety, under the name of Mureto, has been described (Lacombe et al. 2007). This variety has a rose mutation. Mouraton is a black Spanish grapevine mainly cultivated in the border between Spain and northeastern Portugal (Arribes del Duero). Tinta Gorda is a Portuguese variety cultivated in Dão and Douro. In this study, a full match has been obtained between the SNP genotypes of this variety and Mouraton, which has allowed for discovering that these two varieties are synonyms. Camarate, a black cultivar widely cultivated in Portugal, was largely planted during the 18th century and presently covers 947 ha (Galet 2000). This variety is characterized by small to medium bunches and is very susceptible to oidium disease. Água Santa is a black winegrape cultivar authorized in the Beira Litoral, Beira Interior, Ribatejo, Madeira, and Açores regions in Portugal. This variety was obtained by Leão Ferreira de Almeida in 1948, after crossing Mortágua (syn. Camarate) and João Santarém (syn. Castelão) (Ghira et al. 1982). SNP data confirmed that this variety originated from a cross between Periquita (syn. Castelão) and Camarate, and thus is a “double grandchild” of Cayetana Blanca and Alfrocheiro Preto (Figure 1).

Jaén Tinto is a descent of Jaén Blanco and Listán Prieto.

A cross between the cultivars Cayetana Blanca and Listán Prieto gave rise to Jaén Tinto (Table 2, Figure 1). This cross had been previously described (This et al. 2006) based on the genotypes for 20 microsatellite loci, with a very high LOD score of 61. Using SNP, this trio is supported with an even higher LOD score of 80. Jaén Tinto is an old Spanish cultivar (Valcarcel 1765, Rojas Clemente 1807, Abela 1885, García de los Salmones 1914) that is now cultivated in southern Spain (Galet 2000), with 1.145 ha in the Granada, Jaén, Almeria, and Badajoz provinces (Figure 2).

There is another red variety called Jaen that originates from the Bierzo region, Castilla-León (Spain) (Martins et al. 1997). This Jaen is a homonym for Jaén Tinto, which corresponds to a synonym of Mencía, the typical red variety of that region.

The relationship between Cayetana Blanca and Jaén Tinto was also expected, based on several morphological traits. The shape of the mature leaves is highly similar: pentagonal, with rectilinear teeth, and often a tooth in the margin of the petiole sinus. The clusters of the two cultivars are similar only in shape and density: Jaén Tinto clusters are smaller than those of Cayetana Blanca. Morphological similarities between Jaén Tinto and Cayetana Blanca are exemplified by the fact that a well-known Spanish synonym for Cayetana Blanca is Jaén Blanco. Conversely, Listán Prieto and Jaén Tinto are morphologically quite dissimilar.

There are doubts about the origin of Listán Prieto, and the fact that it is the parent of Jaén Tinto could provide information. Listán Prieto is presently cultivated in several of the Canary Islands, with ~1700 ha in Tenerife, Lanzarote, and Gran Canaria, but its cultivation in the Iberian Peninsula almost disappeared after phylloxera in the 19th century. This variety is also cultivated in the Americas from California to Chile, where it is known under many names, including País, Criolla, and Mission (Milla-Tapia et al. 2007, This et al. 2006). Some bibliographic sources consider Listán Prieto as originally from the Americas (VIVC), with the suggestion that it could have been developed in South America from grapevine seeds introduced by Spanish missionaries in the 16th century (Galet 1957). Nevertheless, given that Listán Prieto is a parent of Jaén Tinto, a variety mentioned by Valcarcel (1765), indicates its existence as a cultivar earlier in the Iberian Peninsula. Thus, it would be very difficult to consider that Listán Prieto was developed in America from a seed, was multiplied and distributed there, and later brought to the Iberian Peninsula and crossed with Cayetana Blanca to produce Jaén Tinto. In addition, Herrera (1513) cited a variety called Palomino Negro that, according to his description, could possibly correspond to Listán Prieto.

Based on the maternal transmissions of chloroplasts in the Vitis genus (Arroyo-García et al. 2002, Strefeler et al. 1992), the analysis of the chloroplast microsatellite loci leads us to propose Listán Prieto as the mother and Cayetana Blanca as the father of Jaén Tinto. For the other crosses, the coincidence of the parental haplotypes precludes the identification of maternal and paternal parents (Table 2).

Cayetana Blanca is the putative parent of several other Iberian varieties.

Cayetana Blanca is related to several Spanish varieties—Puerto Alto, Castellana Blanca, Cigüente, Plateado, Rocia, Garrido Macho, and Castillo de Arcos—sharing at least one allele per locus at the 243 SNPs studied (Figure 1). These results point to possible parent-offspring relationships between Cayetana Blanca and each of the mentioned varieties. In some cases this relationship is in agreement with some degree of morphological similarity between the related cultivars, while in others there are no conspicuous similarities.

Puerto Alto is a minor white winegrape cultivar grown in the Andalusian region (southern Spain) (García de los Salmones 1914). Castellana Blanca originated from the Rueda region (Spain) and is also known as Tolociriana and Temprana Agosteña (Yuste et al. 2006). Cigüente is considered a Spanish autochthonous cultivar and was first cited by Herrera (1513). This variety spread and is still cultivated along the Spanish-Portuguese border, following the Roman pathway known as Vía de la Plata (“silver way”) (Muñoz-Organero et al. 2006), where it is known by various names (Supplemental Table 2). In Portugal, its official name is Síria. Plateado is a white wine grapevine cultivated in Andalusia and first described by Candela (1971). Morphologically it is very similar to Cayetana Blanca. Rocia is a black wine cultivar from Spain also found in southern Spain (Martín et al. 2003). Garrido Macho is a white grapevine, long cultivated in Huelva (a southwestern Spanish region that borders Portugal), from where it spread to Cadiz and neighboring regions (Galet 2000). It is also known as Garrio Macho, Parron Garrido, and Parron Garrio (www.vivc.de). Castillo de Arcos is a white grape cultivated in Jerez, southwest Spain (www.vivc.de).

Origin and synonyms of Cayetana Blanca.

Alentejo-Badajoz, a region in the southwest Iberian Peninsula, has been suggested as the probable place of origin of Cayetana Blanca (Lopes et al. 2006). In Morocco, the name for Cayetana Blanca is Djinani and is also considered a local variety. Unfortunately, there were no written records on viticulture in Morocco until the 1950s (Vidal 1951). Galet (2000) mentioned Djiniani as a Moroccan variety considered by Truel as a synonym of Jaén Blanco. The accession used here was from the SODEA germplasm bank, a collection created in 1952 in Meknes by Vidal, who collected local vines from different regions, particularly from the Rif (northern Morocco). A comparison of the genetic profile of Djinani and Jaén using 20 SSR recently confirmed that they are synonyms (El Oualkadi et al. 2009). This study confirmed the identity of these synonyms and two color sports (rose) (Table 1) for the SNP markers used. Their local names and distribution are shown in Supplemental Table 2.

In Portugal, Cayetana Blanca is currently known as Sarigo and is cultivated in Douro. It is also known as Mourisco Branco (Alentejo), Boal Carrasquenho, and Dona Branca (Estremadura, Portugal).

The numerous Iberian cultivars identified here as progenies of Cayetana Blanca and the number of synonyms in Spain—Calagraño, Baladí, Blanca Cayetana, Cirial, Baladí Verdejo, Jaén Empinadillo, Maizancho, Robal, Balay, and Cayetana (Asensio 2000, Manso de Zúñiga 1905)—strongly suggest that Cayetana Blanca originated in the Iberian Peninsula, probably close to the present border between Portugal and Spain, where many relatives of Cayetana Blanca have originated and have been long cultivated (Figure 2). The probable exchange of grapevine cultivars between northern Africa and the Iberian Peninsula during centuries of close interaction would be responsible for its spread in Morocco.

The parents of Cayetana Blanca remain unknown.

Twenty-eight SSR markers were used to provide evidence that Cayetana Blanca was the progeny of a cross between Antão Vaz and Rabo de Ovelha (Lopes et al. 2006). Although Cayetana Blanca displayed only one of the parental alleles at three loci, the authors posited the presence of null alleles which would still make the cross feasible. Geographical and historical reasons also supported the parentage hypothesis: Antão Vaz is the most important wine white cultivar cultivated in the Alentejo region, around Virgueira, Portugal (Almadanim et al. 2007, Lopes et al. 2006). Rabo de Ovelha is a white grapevine which at present is found throughout Portugal (Lopes et al. 2006), but it probably originated from the Alentejo region (Almadanim et al. 2007). Cayetana Blanca is mainly cultivated in the Extremadura region, close to the Alentejo region, but on the Spanish side of the Iberian Peninsula.

The analysis of 243 SNP loci in Cayetana Blanca, Antão Vaz, and Rabo de Ovelha allowed for discarding the suggested trio because there were incompatibilities at 10 SNPs (Supplemental Table 3). Mutation rates are in general considerably lower for SNPs than for microsatellites (Helyar et al. 2011), and thus it is expected to find fewer incompatibilities due to mutations in parentage analyses using SNP markers than microsatellite markers. In order to definitively corroborate or discard this putative parentage, we genotyped the three cultivars involved at 20 SSR loci. The trio was noncompatible at three SSR loci (Supplemental Table 4): VMC1B11, VVIQ52, and VVMD32. Furthermore, since Cayetana Blanca is heterozygous at these three mismatching loci, noncompatibilities cannot be caused by null alleles.

We also tested the possibility of pair-wise relationships of parent-offspring type in the three varieties. Antão Vaz and Rabo de Ovelha are not compatible at four SNPs. A similar result was previously found using 28 SSR markers where the two varieties share only 36% of alleles and then cannot have a parent-offspring relationship (Lopes et al. 2006). Nevertheless, Antão Vaz with Cayetana Blanca and Rabo de Ovelha with Cayetana Blanca share one allele per locus in all markers analyzed in this study. Thus, we conclude that Antão Vaz x Rabo de Ovelha is not the cross that gave rise to Cayetana Blanca; however, one of them could be a parent and the other an offspring, or perhaps both are offspring of Cayetana Blanca. This result stresses the difficulties of rejecting false trios when very closely related cultivars are involved, even using a high number of markers.

As described above, we found that Cayetana Blanca is related to other old and minor cultivars from Spain and Portugal, sharing one allele per locus at 243 SNPs (Figure 1, Figure 2). Any of them could be one of the parents of Cayetana Blanca, but, given the known history of the varieties included, we suggest that Cigüente or Castellana Blanca would be more probable.