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Pedigrees of Fifty Table-Grape Cultivars

Article in American Journal of Enology and Viticulture January 2009

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Pedigrees of Fifty Table-Grape Cultivars

Alba M. Vargas,1 M. Teresa de Andrs,1 Joaqun Borrego,1
and Javier Ibez1,2*
Abstract: Information on the origin of elite table-grape cultivars is essential for breeding programs. A parentage study on table-grape accessions based on analysis of 25 nuclear and five chloroplast microsatellite loci
was carried out. The pedigree of 50 cultivars was analyzed, including morphology studies when necessary.
The chlorotypes clarify the direction of the crosses in the majority of cases, while likelihood ratios indicate
high reliability of the proposed crosses. Nineteen new pedigrees are proposed for varieties from highly diverse
geographical origins, such as Circ, Imperial Roja, Misket Vratchanskii, Mistress Hall, and Pizzutello Nero.
Mistakes were also detected in previously described pedigrees for 13 cultivars, including Delizia di Vaprio and
Madeleine Angevine. Alternative parents were identified for these 13 varieties. For another 18 varieties the
previously proposed pedigree was confirmed. The cultivars Muscat of Alexandria, Afus Ali, Muscat Hamburg,
and Chasselas were the most frequent parents of these 50 varieties.
Key words: chlorotype, genotyping, microsatellite, parentage analysis, Vitis vinifera L.

Grapevine (Vitis vinifera L.) is an ancient crop with a

long history of human selection. Although winegrapes are
economically more important, world production of grapes
intended for fresh consumption grew by 26% between the
end of the 1990s and the beginning of the 2000s (OIV 2005).
Table-grape breeding is also becoming more significant, as
indicated by the rising number of applications for plant variety rights at the European Community Plant Variety Office (http://www.cpvo.eu.int/main/). Historically, table-grape
breeding made significant advances during the 19th century
and the beginning of the 20th century because of the work
of breeders such as Bruni and Prosperi in Italy, Vibert and
Moreau-Robert in France, Foster and Thomson in the United
Kingdom, and Mathiasz in Hungary. In Italy, Luigi and Alberto Pirovano produced more than 500 new table-grape
cultivars (Galet 2000), some of which, like Italia, are still in
use. These breeders often used a limited number of common
parents such as Chasselas, Afus Ali, and Muscat of Alexandria. The latter, for instance, is involved in 191 crosses, according to the Vitis International Variety Catalogue (VIVC;
www.vivc.bafz.de). Table-grape breeding today is of global
interest, and cultivars used in current breeding programs

are often progeny from past breeding programs, thus taking

advantage of previously selected favorable traits.
Information on the origin of table-grape cultivars is of
great value for breeding programs, but often that information is unknown or incorrect. Among widely distributed
elite grapevine cultivars there are many synonyms (syn.,
different names for the same cultivar) and homonyms (different cultivars with the same name) (This et al. 2006).
This complication occurs not only in wine cultivars but
also in table-grape cultivars (OIV 1987), especially older
ones, which makes it difficult to determine the parentage
relationships from historical archives. For example, when
different homonyms exist for a published parent, it is difficult to know which cultivar is the one cited in the literature. Moreover, some synonyms or homonyms have existed
at certain times and places but then disappeared from use,
leaving their modern equivalent, if any, unknown.
In grape-breeding programs, there is certainty about the
female parent, from which the seeds are collected, but the
putative pollen donor cultivar is frequently mistaken, due
to, for example, pollen contamination during the pollination process. That is why in some crosses the male parent is
unknown or does not correspond to the described progenitor (Ibez et al. 2009). Today, molecular markers can help
detect mistakes by identifying the putative male parent.
Sequence-tagged microsatellite sites (STMS or microsatellites) are excellent markers for grapevine characterization
(Bowers et al. 1996, Thomas and Scott 1993). In general,
all plants belonging to the same cultivar (i.e., from a monoclonal origin) show identical genotypes at all microsatellite
loci. Co-dominant Mendelian inheritance of microsatellites
has been previously used to discover the genetic origin of
some important wine cultivars (Bowers and Meredith 1997,
Bowers et al. 1999, Sefc et al. 1998). Moreover, chloroplast
microsatellites can uncover the direction of the cross, as
chloroplasts are maternally inherited in grapevine (ArroyoGarca et al. 2002).

Instituto Madrileo de Investigacin y Desarrollo Rural, Agrario y Alimentario (IMIDRA), Finca El Encn, Carretera A-2, PK 38,200, 28800 Alcal
de Henares, Spain; 2present address, Instituto de Ciencias de la Vid y del
Vino (ICVV), Complejo Cientfico Tecnolgico, C/Madre de Dios 5, 26006
Logroo, Spain.
*Corresponding author (email: javier.ibanez@icvv.es; fax: 34 94 1299721)
Acknowledgments: This study was made possible with the funding from the
RF99-009 project (INIA, Agriculture Ministry of Spain) and of the GrapeGen
project (joint venture between Genome Canada and Genoma Espaa). A.M.
Vargas was funded by a predoctoral fellowship from IMIDRA.
Manuscript submitted Mar 2009, revised May 2009, accepted Jun 2009
Supplementary data is freely available with the online version of this article
at www.ajevonline.org.
Copyright 2009 by the American Society for Enology and Viticulture. All
rights reserved.
1

525
Am. J. Enol. Vitic. 60:4 (2009)

526 Vargas et al.

Given the great economic importance of this species

and the numerous breeding programs in progress worldwide, it is essential to determine the actual genetic origin
of historical cultivars. Previously, we confirmed 35 historical pedigrees and suggested eight new pedigrees, paying
special attention to seedless grape cultivars (Ibez et al.
2009). This expanded work reveals the genetic origins of
50 table-grape cultivars through genotyping of nuclear and
chloroplast microsatellite loci. The existence of synonyms
and homonyms makes it necessary to accompany molecular
analysis with thorough literature revisions and morphological evaluation to ascertain that the described cultivars are
those mentioned in the historical records.

Materials and Methods

Plant material was obtained from the Vitis Germplasm
Bank (BGV) at the Finca El Encn (IMIDRA, Alcal de
Henares, Spain). A total of 94 grape accessions were involved in the studied pedigrees (Table 1). Most are tablegrape accessions, but some wine cultivars were included as
possible parents of some table-grape cultivars. Among the
94 accessions, 24 in our collection have a name different
from the prime name established in the Vitis International
Variety Catalogue (VIVC; www.vivc.bafz.de).
DNA extractions were carried out from young leaves
using the DNeasy 96 Plant Kit (Qiagen, Hilden, Germany).
Twenty-five nuclear STMS were analyzed through three
multiplex PCR assays as described previously (Ibez et
al. 2009). Five additional chloroplast microsatellites were
analyzed in 91 accessions through a PCR multiplex (Ibez
et al. 2009). Chloroplast haplotypes were named using published nomenclature (Arroyo-Garca et al. 2006).
Pedigree analysis was accomplished using Identity 1.0
software (Wagner and Sefc 1999). Statistical analysis was
performed using likelihood ratios (Hagelberg et al. 1991)
where parent/offspring combinations were detected. The
likelihood ratio (LR) is the quotient between the probability
of the progeny genotype if it had the presumptive parents
and the probability of the progeny genotype for alternative
possibilities.

Results and Discussion

In a previous study of 25 nuclear microsatellites in 381
accessions, compatible crosses were found for 43 tablegrape cultivars (Ibez et al. 2009). In the present work, the
pedigrees of 50 other table-grape cultivars were determined
by using the same 25 microsatellite loci (Supplementary
Table 1). Because of the existence of synonyms, homonyms,
and possible mistakes within the collection, the morphology
of many cultivars involved in these pedigrees was studied. These data were compared with published descriptions
(Branas and Truel 1965) to confirm, as far as possible, that
the accessions used were the correct ones. Chloroplast haplotypes (Supplementary Table 2) were used to uncover the
direction of the crosses.
The pedigrees reported were divided into three groups:
(1) 19 cultivars for which no pedigree information was

available in literature (Table 2), (2) 13 cultivars whose published pedigree was incorrect and for which a corrected
pedigree is proposed (Table 3), and (3) confirmation of 18
pedigrees previously described (Table 4). We analyzed the
chlorotypes of all parents and progeny and the likelihood
ratios for each pedigree using a reference database of 301
mostly table-grape cultivars (data not shown).
New proposed pedigrees. The Italian breeder Bruni
named his new cultivars with his name followed by a number. Bruni 12 and Bruni 36 are described as progeny of
open pollinations of Madeleine Angevine (Branas and Truel
1965). We identified Baresana, an Italian cultivar from the
region of Bari, as the male parent of Bruni 12, which is also
cultivated in this region (Galet 2000). We also identified
another Italian cultivar, Luglienga Bianca (Lignan blanc),
as the pollen donor of Bruni 36 based on microsatellite genotypes. Furthermore, Bruni 36 is similar to Lignan blanc in
traits such as dense bunches and early ripening.
Bruni 707 is compatible with a self-pollination of Cardinal (LR 7.07 x 10 21). They differ at only four microsatellite loci and are morphologically very similar. Alternatively, Bruni 707 could be a cross between Cardinal and
Koenigin der Weingaerten if a null allele is accepted at
locus VMC1B11 (LR 1.25 x 1017 ). Cardinal is a progeny of
Koenigin der Weingaerten (Ibez et al. 2009), and thus
they share at least one allele per locus, making it difficult
to eliminate one of them as a parent, even after analysis
of five additional loci (data not shown). Given the higher
likelihood ratios and full compatibility, the hypothesis of a
self-pollination of Cardinal is more probable.
Vibert and his successor Moreau-Robert were active
table-grape breeders in Angers, France, during the 19th
century. According to microsatellite data, three of their selected cultivarsSulivan blanc, Circ, and Duc de Magentaarose from a cross between Schiras and Chasselas. Sulivan blanc is similar to Chasselas (Galet 2000), while Circ
has female f lowers, like Schiras. Duc de Magenta could
also come from the same cross, assuming the presence of a
null allele at locus VVIP60. Schiras and Chasselas are also
the parents of another cultivar, Black Prince, but we found
no available information about its breeder. According to the
VIVC (www.vivc.bafz.de), there are three homonymous cultivars named Black Prince: one corresponds with Cinsaut,
another with Schiava Grossa, and the third is cultivated in
South Africa and corresponds with the actual Black Prince.
The latter is presumably the accession present in El Encn,
which comes from South Africa and is different from Cinsaut and Schiava Grossa.
Gradisca is another cultivar bred in Angers by MoreauRobert, using Chasselas as the pollen donor, while Bicane
(female flower) was the mother. As Chasselas Dor, Gradisca
is a white cultivar with golden tone berries at maturity and
has a synonym called Moranet Dor (www.vivc.bafz.de).
Though no pedigree information was found in the literature for the cultivar Malingre Precoce, bred by Malingre
in France in 1840, its genotype suggests that it could be
derived from a cross between Bicane and Pinot. Malingre

Am. J. Enol. Vitic. 60:4 (2009)

Pedigrees of Fifty Table-Grape Cultivars 527

Precoce is a white cultivar with loose bunches like Bicane

and ripens early like Pinot. Another early ripening French
cultivar, Roi des Precoces, could also be a progeny of Pinot,
with Cinsaut as the female parent.
Muscat f lavor has been a desired trait in many crosses,
which sometimes supports a pedigree. Muscat Petits
Grains Blancs may be the male parent of cultivars Duch-

ess of Buccleugh and Misket Vratchanskii, whose female

progenitors would be Bicane and Coarna Alba (a female
Romanian cultivar), respectively. Duchess of Buccleugh
was bred by Thomson in 1863 in Scotland. It has Muscat
f lavor and dense bunches, as does Muscat Petits Grains
Blancs, while it could have inherited late ripening and the
liability of producing millerandage from Bicane. Misket

Table 1 Accessions studied with prime names for those accessions that were collected under a name different than the one
established in the Vitis International Variety Catalogue.
Accession
Ahmeur bou Ahmeur
Almera Nera
Alphonse Lavalle
Angelo
Barbableu
Baresana
Beba
Bicane
Black Prince
Black Seedless
Bruni 12
Bruni 36
Bruni 41
Bruni 90
Bruni 415
Bruni 707
Cardinal
Chasselas
Cinsaut
Circa
Ciruela Roja
Coarna Bianca
Colgar Roja
Dabouki
Dalmasso 3-33
Dalmasso 6-6
Dalmasso 7-3
Dalmasso 11-8
Dalmasso 11-20
Danam
Danlas
Dawn Seedless
Delizia di Vaprio
Don Mariano
Dona Maria
Duc de Magenta
Duchess of Buccleugh
Fosters White Seedling
Frankenthal
Gold
Gradiska
Gros Colman
Gros Maroc
Imperial Roja
Jantar
Lady Downes Seedling
Lival

Prime name

Angelo Pirovano

Circ
Coarna Alba

Emilia

Schiava Grossa
Gradisca
Kharistvala Kolkhuri
Marocain noir

Accession
Luglienga Bianca
Madeleine Angevine
Madeleine Royal
Mistress Hall
Moscatel de Alejandra
Moscatel de Grano Menudo
Moscatel de Hamburgo
Moscato d Adda
Moscato Gustav Szauter
Moscato di Terracina
Muscat Madame Mathiasz
Muscat Ottonel
Ohanes
Olivetta Nera
Olivette Barthelet
Olivette Blanche
Panse Precoce
Perlette
Pinot noir
Pirovano 93
Pirovano 159
Pirovano 188
Pirovano 235
Pirovano 315
Pirovano 531
Pizzutello Bianco
Pizzutello Nero
Precoce de Malingre
Princeps
Prosperi 285
Prune de Cazouls
Prunesta Nera
Queen
Ragol
Regina dei Vigneti
Roi des Precoces
Roseti
Schiraz
Sovrana
Sulivan
Sultana Moscata
Sultanina
Thalloczy Lajos
Therme
Verdea
Vivona 378
Wratchanski Misket

Am. J. Enol. Vitic. 60:4 (2009)

Prime name
Lignan blanc
Madeleine Royale
Muscat of Alexandria
Muscat Petits Grains Blancs
Muscat Hamburg
Szauter Gusztav
Mathiasz Janosne

Olivette Noire

Sicilien
Pinot

Cornichon blanc
Malingre Precoce

Prunesta

Koenigin der Weingaerten

Afus Ali
Schiras
Sulivan blanc

Misket Vratchanskii

528 Vargas et al.

Vratchanskii is a white Muscat-f lavored Bulgarian cultivar bearing bunches and berries similar to Muscat Petits
Grains Blancs.
The female-f lowered cultivar Marocain noir and Muscat of Alexandria could be the parents of Mistress Hall,
an English cultivar. Mistress Hall has a slight Muscat f lavor, probably inherited from Muscat of Alexandria, and
the dense clusters, dark black berries, and late ripening of
Marocain noir. The unknown breeder of this cultivar may
be Foster, as he obtained other cultivars in England using
Marocain noir as the female progenitor.
The cultivar Vivona 378 (probably bred by Antonino
Vivona in Italy) could be the offspring of Muscat of Alexandria and Bruni 415. It would have inherited its loose
bunches, flat berries, and early ripening from Bruni 415.
There are two Italian accessions both named Pizzutello
Nero with unknown origins and different genotypes. One
is synonymous with Olivette Noire and the other is an accession present in different collections (www.vivc.bafz.de).
The parents of the second Pizzutello Nero could be Cornichon blanc and Prune de Cazouls. The latter cultivar is a
black cultivar of possible Eastern origin cultivated in Sicily
(Longo, cited by Galet 2000).
Finally, there exist four Spanish cultivars of unknown
origin that could be highly related: Imperial Roja, Ciruela
Roja, Colgar Roja, and Almera Nera. Their genotypes re-

veal that they could all share Ohanes as their mother. This
female-f lowered cultivar originated in Rgol, a village in
the province of Almera (Andalusia, Spain). The cultivar
was carried to Ohanes, a village 13 km from Rgol (Alonso
et al. 2006). The cultivar Ragol, named after Rgol village, is not a direct relative of Ohanes (Ibez et al. 2009).
The four accessions proposed here as offspring of Ohanes
were collected in Almera, and three of them have Ragol
as the male parent. Imperial Roja has been described as
a synonym for Ahmeur bou Ahmeur (Galet 2000), likely
because of morphological similarity, but the genotype of
the accession present in El Encn is distinct from Ahmeur
bou Ahmeur and from the rest of the cultivars studied. We
found that a cross between Ohanes and Ragol could have
produced this cultivar. Ragol has been frequently used as
a commercial pollinator for Ohanes in Almera (Hidalgo
1999). On the other hand, Ragol itself is also considered a
synonym for Ahmeur bou Ahmeur according to the Vitis
International Variety Catalogue. Ragol is called Encarnada de Ragol and Uva de Colgar in Almera (Alonso et al.
2006), and Galet (2000) considers those names synonyms
of Ahmeur bou Ahmeur. However, the genotype of Ragol
present at El Encn does not match Ahmeur bou Ahmeur.
We cannot exclude the existence of two homonyms called
Ragol, one a synonym for Ahmeur bou Ahmeur and the
other from Almera, studied here.

Table 2 Nineteen new pedigrees proposed. Compatible pedigrees, haplotypes (H) for parents and progeny, and likelihood ratios
for the crosses calculated on the basis of 25 microsatellite loci.
Progeny
Almera Nera
Black Prince
Bruni 12
Bruni 36
Bruni 707
Circ
Ciruela Roja
Colgar Roja
Duc de Magentab
Duchess of
Buccleugh
Gradisca
Imperial Roja
Malingre Precoce
Misket Vratchanskii

H
C
C
C
C
B
C
C
C
C
C

Female parent
Ohanes
Schiras
Madeleine Angevine
Madeleine Angevine
Cardinal
Schiras
Ohanes
Ohanes
Schiras
Bicane

H
C
C
C
C
B
C
C
C
C
C

C
C
C
C

Bicane
Ohanes
Bicane
Coarna Alba

C
C
C
C

Mistress Hall

Marocain noir

Pizzutello Nero
Roi des Precoces
Sulivan blanc
Vivona 378

A
C
C
B

Cornichon blanc
Cinsaut
Schiras
Muscat of Alexandria

A
C
C
B

Male parent
Prunesta
Chasselas
Baresana
Lignan blanc
Cardinal
Chasselas
Ragol
Ragol
Chasselas
Muscat Petits
Grains Blancs
Chasselas
Ragol
Pinot
Muscat Petits
Grains Blancs
Muscat of
Alexandria
Prune de Cazouls
Pinot
Chasselas
Bruni 415

H
A
D
C
D
B
D
A
A
D
D

XxY
9.22E+21
3.25E+18
7.27E+21
3.63E+20
7.07E+21
2.84E+22
5.38E+20
5.07E+17
4.51E+18
2.39E+18

Likelihood ratioa
(1)xX
rel(2)x(1)
(2)xX
5.29E+11 5.17E+03 4.51E+10
5.50E+10 2.32E+03 2.62E+13
2.02E+13 1.15E+04 9.32E+11
1.62E+12 6.55E+03 4.52E+11
2.93E+10 4.41E+03 2.93E+10
1.47E+15 2.88E+04 2.71E+12
6.19E+13 9.00E+03 9.33E+07
5.39E+11 5.20E+03 1.22E+08
5.22E+12 1.14E+04 7.79E+11
1.19E+14 2.52E+04 4.62E+09

rel(1)x(2)
3.26E+03
6.66E+03
4.99E+03
6.90E+03
4.41E+03
3.76E+03
6.12E+02
8.64E+02
4.78E+03
2.31E+03

D
A
A
D

1.50E+17
1.63E+21
1.37E+20
1.06E+19

1.43E+10
3.09E+12
7.50E+14
6.73E+10

1.94E+03
9.57E+03
1.81E+04
5.26E+03

1.44E+10
4.77E+10
6.58E+08
6.22E+11

3.33E+03
3.94E+03
1.33E+03
4.61E+03

1.72E+17

1.19E+10

2.73E+03

5.00E+10

2.55E+03

D
A
D
C

7.35E+19
7.88E+16
1.12E+20
2.92E+21

5.07E+10
1.85E+10
4.49E+11
5.77E+16

3.50E+03
1.89E+03
7.39E+03
3.95E+04

3.31E+14
2.55E+10
4.57E+14
1.75E+10

1.54E+04
3.05E+03
1.11E+04
3.23E+03

Likelihood ratio: quotient between the probability of the progeny genotype if it had the presumptive parents and the probability of the progeny
genotype for alternative possibilities: XxY, where X and Y are random unrelated cultivars; (1)xX and (2)xX, the identity of one parent is assumed but that of the other is unknown (a random unrelated cultivar); rel(2)x(1) and rel(1)x(2), the identity of one parent is assumed but the
possibility is considered that the other parent is a close relative of the cultivar proposed as the second parent. A close relative is a parent,
progeny, or full sibling.
bLikelihood ratio is based on 24 microsatellites because of the presence of null alleles or mutations.
a

Am. J. Enol. Vitic. 60:4 (2009)

Additionally, crosses between Ohanes and

Ragol gave rise to Cir uela Roja and Colgar
Roja. Both crosses have the late ripening characteristic of Ohanes. Colgar Roja is listed as
an American cultivar in the VIVC, but given
its parents, it should be considered of Spanish
origin. Furthermore, one synonym for Colgar
Roja is Colgar Rioja, named after Rioja village,
which is close to the villages of Ohanes and
Rgol.
These data point to a natural origin for these
three cultivars: spontaneous pollinations in the
Almera growing region. Among the millions
of natural crosses between Ohanes and Ragol
needed to produce grapes in the female Ohanes
plants, seedlings growing from unharvested
fruits were probably produced. Three of them
were selected as new cultivars which have survived until the present time.
Almera Nera is the fourth Ohanes progeny
found here. It has been considered a synonym of
Alphonse Lavalle (Galet 2000). Nevertheless,
Alphonse Lavalle is dark black, while Almera
Nera is red, and the berry size and shape are
different as are their microsatellite genotypes.
Almera Nera was probably conf used with
Almer a Negra (which is the true synonym for
Alphonse Lavalle) because of the similarity in
their names. According to the Vitis International Variety Catalogue, Almera Nera is Imperial
Napolen, a synonym for Don Mariano. However, Don Mariano is genetically different from
Almera Nera. Microsatellite analysis shows
that Almera Nera could derive from a cross
between Ohanes and Prunesta (syn. Prunesta
Nera). Moreover, Almera Nera has late ripening like Ohanes and is red, like Prunesta, from
which it could have inherited its second name.
Possible mistakes in literature. Parentage
analyses using microsatellite genotypes also
allowed detection of several possible errors in
the literature. We used the most complete and
extended information sources (Branas and Truel
1965, Galet 2000) unless otherwise stated. Compatible progenitors for the mistaken pedigrees
are suggested and their chlorotypes and likelihood ratios are provided (Table 3).
According to the literature, the parents of
Fosters White Seedling are Black Morocco (syn.
Marocain noir) and Sweet Water (syn. Chasselas). These two cultivars are also specified as the
parents of Lady Downes Seedling, also bred by
Foster in England in 1835. However, Chasselas
is noncompatible as it does not share at least
one allele at 10 loci with Fosters White Seedling and at 12 loci with Lady Downes Seedling.
Our results show that Marocain noir and Schiava

Am. J. Enol. Vitic. 60:4 (2009)

Marocain noir x Chasselas

Madeleine Royale x Malingre

Precoce
Chasselas rouge x Muscat Ottonel
Olivette Blanche x Cornichon blanc
Moscato dAdda x Olivette Noire
Delizia di Vaprio x Madeleine
Royale
Kharistvala Kolkhuri x Olivette Noire
Angelo Pirovano x Sultana Moscata
Moscato di Terracina x Bicane
Koenigin der Weingaerten x
Muscat of Alexandria

B
C
C
C

B
D
D
C

Fosters White
Seedling
Lady Downes
Seedling
Madeleine Angevine

Mathiasz Janosne
Olivette Barthelet
Pirovano 159
Pirovano 315

Pirovano 531
Princeps
Prosperi 285
Thalloczy Lajos

1
1, 2
1, 2
2, 3

1, 2
2
1
1

1, 2

1, 2

1, 2

Ref
1, 2
1, 2

Kharistvala Kolkhuri
Muscat Hamburg
Moscato di Terracina
Sicilien

Muscat of Alexandria
Olivette Blanche
Delizia di Vaprio
Madeleine Angevine

Circ

Marocain noir

Marocain noir

Female parent
Sicilien
Sicilien

B
D
D
C

B
C
C
C

H
C
C

Barbableu
Kharistvala Kolkhuri
Muscat of Alexandria
Muscat of Alexandria

Chasselas Rouge
Beba
Angelo Pirovano
Delizia di Vaprio

Madeleine Royale

Muscat of Alexandria

Schiava Grossa

Male parent
Szauter Gusztav
Muscat of Alexandria

C
B
B
B

D
A
D
C

H
C
B

1.05E+16
1.41E+17
4.09E+19
1.04E+16

1.95E+18
1.06E+14
2.30E+19
1.13E+17

2.01E+08
1.00E+11
9.25E+10
4.72E+10

1.75E+11
4.62E+08
2.60E+12
2.42E+10

9.45E+02
4.22E+03
5.42E+03
4.67E+03

3.96E+03
9.17E+02
8.71E+03
1.67E+03

9.91E+11
1.38E+10
7.19E+11
5.38E+08

7.78E+10
1.08E+08
2.18E+12
2.44E+12

3.77E+03
3.24E+03
8.77E+03
9.24E+02

6.06E+03
8.21E+02
1.06E+04
1.03E+04

1.06E+19 9.98E+09 2.58E+03 1.92E+12 8.61E+03

3.41E+20 2.58E+10 3.01E+03 8.66E+13 7.63E+03

3.16E+19 1.98E+11 5.92E+03 6.19E+12 4.11E+03

Likelihood ratioa
XxY
(1)xX
rel(2)x(1)
(2)xX
rel(1)x(2)
2.61E+16 6.99E+08 9.62E+02 3.08E+09 1.54E+03
4.51E+16 1.80E+10 2.71E+03 9.79E+09 1.61E+03

Likelihood ratio: quotient between the probability of the progeny genotype if it had the presumptive parents and the probability of the progeny genotype for alternative possibilities:
XxY, where X and Y are random unrelated cultivars; (1)xX and (2)xX, the identity of one parent is assumed but that of the other is unknown (a random unrelated cultivar); rel(2)x(1)
and rel(1)x(2), the identity of one parent is assumed but the possibility is considered that the other parent is a close relative of the cultivar proposed as the second parent. A close relative is a parent, progeny, or full sibling.
bReferences: 1, Branas and Truel 1965; 2, Galet 2000; 3, OIV 1987.
cThe female parent could not be identified because both parents are hermaphrodite and presented identical chlorotype.

Described pedigree
Mathiasz Janosne x Sicilien
Muscat of Alexandria x
Fosters White Seedling
Marocain noir x Chasselas

H
C
C

Progeny
Bruni 41c
Delizia di Vaprio
b

Table 3 Thirteen suggested parents for pedigrees with putative mistakes in the literature. Compatible pedigrees, haplotypes (H) for parents and progeny,
and likelihood ratios for the crosses calculated on the basis of 25 microsatellite loci.

Pedigrees of Fifty Table-Grape Cultivars 529

530 Vargas et al.

Grossa (syn. Frankenthal) could be the actual parents of

Fosters White Seedling. This confusion between Chasselas
and Frankenthal could be due to the existence of a sport of
Frankenthal known as Frankenthal blanc, synonym Buckland Sweet Water. This homonym, Sweet Water, could be
the source of the mistake. Moreover, Frankenthal (Schiava
Grossa) has an herbaceous flavor, as does Fosters White
Seedling. On the other hand, Marocain noir and Muscat
of Alexandria could be the real parents of Lady Downes
Seedling. The accession named Lady Downes Seedling in
our collection has a slight Muscat flavor in ripe berries, as
Perold indicated (Branas and Truel 1965), supporting that
one of its parents could be Muscat of Alexandria.
Madeleine Angevine was bred in 1857 and is a presumed
progeny of a cross between Madeleine Royale and Malingre Precoce. Although Madeleine Royale is compatible as
progenitor, Malingre Precoce is noncompatible at 11 loci,
with Circ the most likely other parent. Both Madeleine
Angevine and Circ were released from the same breeding program in Angers, France. Madeleine Angevine could
have inherited female flowers from Circ.
Several mistakes have also been detected in the described pedigrees of cultivars bred by Pirovano, an Italian
who consistently used some cultivars as progenitors, including Delizia di Vaprio, Angelo, and Muscat of Alexandria.
Delizia di Vaprio is an Italian cultivar obtained in 1908
from a supposed cross between Muscat of Alexandria and
Fosters White Seedling. Parentage analysis indicates that
Sicilien (syn. Panse Precoce) could be its female parent
instead of Fosters White Seedling. Fosters White Seedling was called Panse Precoce in Italy (Galet 2000), so the
existence in Italy of two homonymous Panse Precoce could
be responsible for the confusion.
Pirovano 159 is described as the progeny of a cross between Moscato dAdda and Olivette Noire. However, the
genotype of Pirovano 159 is noncompatible at 5 loci with
the former and at 12 with the latter. Our analysis showed
that Delizia di Vaprio and Angelo Pirovano (both bred by
Pirovano) could be the true parents of this cross. Pirovano
315 is described as a progeny of Delizia di Vaprio and
Madeleine Royale, but the genotype of the latter is not consistent with this pedigree. Madeleine Angevine (a descent
of Madeleine Royale, see above) is the most likely female
progenitor instead. Pirovano 531 is a similar case, where
the true parent is likely a progeny of the described parent.
The presumed ancestors of Pirovano 531 are Kharistvala
Kolkhuri (syn. Gros Colman) and Olivette Noire. However,
Olivette Noire is noncompatible at four loci, while Barbableu, a progeny of Olivette Noire and Muscat Hamburg
(Ibez et al. 2009), could be the actual male progenitor.
Princeps is also a Pirovano release. Its described parents are Angelo Pirovano and Sultana Moscata, but they
are noncompatible at seven different loci. Muscat Hamburg
and Kharistvala Kolkhuri could be the real progenitors of
Princeps.
Antoine Besson bred Olivette Bar thelet in 1864 in
France. It is described as a cross between Olivette Blanche

and Cornichon blanc. However, the accession analyzed

in this work arises from crossing Olivette Blanche with
Beba. Cornichon blanc is noncompatible at 11 microsatellites, while Beba and Olivette Barthelet share a special
trait: their berry pedicels separate easily from the cluster.
The mistake could be due to a confusion between Beba and
Cornichon blanc, since Beba is an ancient cultivar mainly
cultivated in Andalusia and Cornichon blanc was described
by a 13th-century Arabian author in Mlaga (Andalusia)
(Rojas Clemente y Rubio 1807).
Mathiasz bred the cultivar Mathiasz Janosne in 1902 in
Hungary, reportedly from a cross between Chasselas rouge
and Muscat Ottonel. Muscat Ottonel is noncompatible with
Mathiasz Janosne at three loci and at 13 loci when Chasselas rouge is considered as the second parent. Muscat of
Alexandria could be the female progenitor. In this case,
the male parent was correct, while the mistake occurred
in the mother, probably because of confusion between two
Muscat cultivars at the breeding center: Muscat Ottonel and
Muscat of Alexandria. Mathiasz also bred Thalloczy Lajos
from a reported cross between Royal Vineyard and Muscat
of Alexandria (Galet 2000, OIV 1987). The prime name
for Royal Vineyard is Koenigin der Weingaerten and it is
noncompatible with Thalloczy Lajos. Nevertheless, there is
a less common homonym for Royal Vineyard whose prime
name is Sicilien (syn. Panse Precoce), which is compatible
as a parent of this progeny. In this instance, the problem
stems from the use of a little-extended synonym for the
parent Sicilien. To avoid confusion with Koenigin der Weingaerten, this pedigree has been included as a possible
mistake.
Bruni 41 was described as a progeny of Mathiasz Janosne
and Panse Precoce. However, Mathiasz Janosne is noncompatible with this pedigree at eight microsatellites. Szauter
Gusztav, also bred by Mathiasz, could be the actual male
progenitor. Lastly, Prosperi 285 is described as the offspring of Moscato di Terracina and Bicane, but results are
noncompatible with Bicane at 12 loci. Muscat of Alexandria
could be the male parent.
Confirmation of previously described pedigrees. We
confirmed 18 previously described pedigrees (Branas and
Truel 1965, Cosmo 1975, Galet 2000, OIV 1987, Wagner
and Truel 1988). The direction of the crosses, frequently
unknown, was established through chlorotype analysis (Table 4). The presence in these pedigrees of certain cultivars
with revised pedigrees discussed in the previous section,
such as Delizia di Vaprio and Mathiasz Janosne, supports
findings described above.
The crosses that produced Danam and Bruni 90 are compatible only if a null allele is present at the locus VVIV37.
The presence of this possible mutation in two pedigrees
described in the literature supports the existence of null
alleles at this locus.
Reliability of described crosses. The reliability of the
50 crosses studied was quantified through the calculation
of likelihood ratios. The values ranged from 1016 to 1022 for
the new and amended pedigrees (Table 2, Table 3), while

Am. J. Enol. Vitic. 60:4 (2009)

Pedigrees of Fifty Table-Grape Cultivars 531

ratios for the confirmed pedigrees were lower (1013 to 1021,

Table 4), supporting the new proposed pedigrees. When
considering the possibility of close relatives involved in
the crosses (rel(2)x(1) and rel(1)x(2) in Tables 2, 3, 4), likelihood ratios are much lower, but still above 103 for most
crosses. Again, there are no substantial differences between
the values obtained for the new and for the previously described pedigrees. Low likelihood ratios are expected as a
consequence of the high endogamy existing within tablegrape cultivars, but our values are in the same order as
published pedigrees for wine cultivars such as Chardonnay,
with 32 microsatellite loci (Bowers et al. 1999). In general,
the values are similar or slightly higher than those obtained
by other authors (Bowers et al. 1999, di Vecchi Staraz et al.
2007, Piljac et al. 2002, Sefc et al. 1998, Vouillamoz et al.
2003), although lower than that obtained for the pedigree of
Lafnetscha, an alpine grape cultivar, using 31 microsatellites (Vouillamoz et al. 2004).
Only four chloroplast haplotypes (A, B, C, and D) were
found among the studied cultivars. Despite this (expected)
low diversity, the chlorotypes allowed discrimination of the
female parent in 42 of the 49 outcrosses. In two instances
unsolved by chlorotypes, maternity could be resolved because one parent was a female (Madeleine Angevine). We
found no prospective pedigree with full compatibility for
the nuclear microsatellites and no compatibility for the

chlorotype of both parents, which also supports the reliability of the suggested crosses.
Historical table-grape breeding. The 50 cultivars for
which compatible parents were found in this work as well
as those previously published (Ibez et al. 2009) were not
chosen or selected for any particular reason and could be
then considered as representative results of breeding efforts
during the 19th and 20th centuries. Hence, some general remarks can be made regarding historical table-grape breeding by pooling data for the 90 pedigrees examined in these
two studies. While 54 cultivars were used as parents, only
10 of them were used six or more times: Sultanina (15),
Muscat of Alexandria (14), Muscat Hamburg (12), Afus
Ali (10), Bicane (9), Delizia di Vaprio (8), Chasselas (8),
Koenigin der Weingaerten (7), Alphonse Lavalle (6), and
Angelo Pirovano (6). Together, these 10 cultivars appear 95
times in the 90 crosses studied. There is a balance between
the use of cultivars of natural origin (such as Sultanina,
Afus Ali, and Chasselas) and bred cultivars (such as Muscat Hamburg, Alphonse Lavalle, and Angelo Pirovano).
Sultanina is involved in most seedless table-grape crosses,
while the two main Muscat table-grape cultivars (Muscat
of Alexandria and Muscat Hamburg) appeared in 26 of the
90 different crosses, indicating the importance of Muscat
f lavor in table-grape breeding. Female cultivars have also
been relevant: Bicane, Madeleine Angevine, and Ohanes

Table 4 Eighteen pedigrees confirmed as previously described in the literature. Compatible pedigrees, haplotypes (H) for parents and
progeny, and likelihood ratios for the crosses calculated on the basis of 25 microsatellite loci.
H
B
D
C
D
D
D
D
D
B
C

Male parent
Black Seedless
Afus Ali
Afus Ali
Afus Ali
Afus Ali
Muscat Hamburg
Dabouki
Perlette
Afus Ali
Afus Ali

H
C
A
A
A
A
D
D
A
A
A

Ref
1, 2
1, 2
1, 2
1, 2
1, 2
2, 3, 4
2, 3, 4
2, 4
2, 3, 4
1, 2, 4

XxY
1.05E+20
6.09E+14
1.22E+16
1.09E+16
3.43E+16
9.51E+19
1.28E+21
2.15E+17
2.26E+19
1.68E+16

A
B
D
C
D
D
D

Female parent
Mathiasz Janosne
Muscat Hamburg
Bicane
Muscat Hamburg
Muscat Hamburg
Dabouki
Chasselas
Gold
Muscat of Alexandria
Bicane
Koenigin der
Weingaerten
Alphonse Lavalle
Angelo Pirovano
Delizia Di Vaprio
Schiava Grossa
Muscat Hamburg
Schiava Grossa

Likelihood ratioa
(1)xX
rel(2)x(1)
(2)xX
1.35E+15 1.25E+04 1.93E+10
1.45E+08 8.23E+02 1.93E+10
6.74E+10 3.30E+03 3.10E+09
2.87E+09 1.38E+03 2.22E+09
3.37E+10 2.88E+03 6.36E+09
1.22E+09 1.87E+03 2.73E+15
2.63E+14 9.77E+03 3.58E+13
3.58E+11 6.75E+03 5.19E+08
3.77E+11 4.80E+03 2.03E+12
2.13E+11 4.16E+03 6.87E+08

A
B
D
C
D
D
D

A
D
D
D
C
C
C

2
2, 3, 4
1
1
1
1, 2, 3
1, 2

5.75E+18
1.10E+20
1.30E+18
1.06E+17
4.30E+15
8.24E+13
4.24E+17

1.19E+10
3.72E+13
3.18E+12
1.46E+11
2.24E+07
3.01E+09
2.31E+11

2.79E+03
1.17E+04
5.92E+03
5.15E+03
6.49E+02
6.77E+02
5.03E+03

3.53E+09
4.97E+11
9.05E+09
2.33E+11
1.49E+12
6.53E+08
1.36E+11

2.54E+03
5.95E+03
2.11E+03
6.72E+03
9.13E+03
1.69E+03
6.93E+03

Cinsaut

Afus Ali
Lignan blanc
Verdea
Angelo Pirovano
Delizia di Vaprio
Sultanina
Delizia di Vaprio
Koenigin der
Weingaerten

1.36E+16

5.87E+09

2.61E+03

1.01E+10

2.29E+03

Progeny
Bruni 90c
Dalmasso 3-33
Dalmasso 6-6
Dalmasso 11-8
Dalmasso 11-20
Danamcd
Danlasd
Dawn Seedless
Dona Maria
Emilia

H
B
D
C
D
D
D
D
D
B
C

Jantard
Lival
Pirovano 93d
Pirovano 188
Pirovano 235
Queen
Sovrana
Therme

rel(1)x(2)
2.85E+03
4.53E+03
2.24E+03
2.66E+03
2.82E+03
1.17E+04
1.63E+04
1.23E+03
1.11E+04
1.50E+03

Likelihood ratio: quotient between the probability of the progeny genotype if it had the presumptive parents and the probability of the progeny
genotype for alternative possibilities: XxY, where X and Y are random unrelated cultivars; (1)xX and (2)xX, the identity of one parent is assumed but that of the other is unknown (a random unrelated cultivar); rel(2)x(1) and rel(1)x(2), the identity of one parent is assumed but the
possibility is considered that the other parent is a close relative of the cultivar proposed as the second parent. A close relative is a parent,
progeny, or full sibling.
bReferences: 1, Branas and Truel 1965; 2, Galet 2000; 3, OIV 1987; 4, Wagner and Truel 1988.
cLikelihood ratio is based on 24 microsatellites because of the presence of null alleles or mutations.
dThe female parent could not be identified because both parents are hermaphrodite and presented identical chlorotype.
a

Am. J. Enol. Vitic. 60:4 (2009)

532 Vargas et al.

appeared in 18 crosses, most of which produced seeded

cultivars.

Conclusions
This work contributes to the definition and clarification
of the parentage of 50 table-grape cultivars and provides
additional information on the existence of possible synonyms and homonyms for cultivars involved in the crosses.
Nineteen new pedigrees with high likelihood were suggested. Thirteen mistakes in the literature were detected and
putative parents identified. Finally, 18 pedigrees previously
described were confirmed by our results. In most instances,
the direction of the cross was determined using chloroplast
microsatellites that allow definition of chlorotypes. Results
confirm that several cultivars like Muscat of Alexandria,
Muscat Hamburg, and Afus Ali have contributed very significantly to the present table-grape gene pool.

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