Plant genetic resources newsletter No. 126, June 2001

Transcripción

ISSN 1020-3362
Plant Genetic Resources Newsletter
Bulletin de Ressources Phytogénétiques
Noticiario de Recursos Fitogenéticos
No. 126, 2001
Food and Agriculture Organization of the United Nations and the
International Plant Genetic Resources Institute
Organisation des Nations Unies pour l'alimentation et l'agriculture et
l'institut international des ressources phytogénétiques
Organización de las Naciones Unidas para la Agricultura y la Alimentación y
el Instituto Internacional de Recursos Fitogenéticos
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rédaction
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Cover: Mature fruits of Cucurbita
pepo—a selection of species
characterized by Paris from the
collection at the Newe Ya’ar
Research Center, Israel: discussed
on pp. 41–45.
Couverture: Fruits mûrs des
Curcubita pepo—une sélection
d’espèces caractérisées par Paris,
provenant de la collection du Centre
de recherche Newe Ya’ar, Israël :
discussion aux pp. 41–45.
Portada: Frutos maduros de
Cucurbita pepo—una selección de
especies caracterizada por París de
la colección del Centro de
Investigación Newe Ya’ar, Israel:
comentario en pp. 41–45.
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© IPGRI/FAO 2001
Plant
Plant
Genetic
Genetic
Resources
Resources
Newsletter,
Newsletter,
2001,
2001,
No. No.
126:126
1 - 11
1
ARTICLE
Classification of Italian maize (Zea mays L.) germplasm
Aureliano Brandolini1 and Andrea Brandolini2
1
2
Centro Ricerca Fitotecnica, Via Mazzini 30, 24128 Bergamo Italy
Istituto Sperimentale per la Cerealicoltura, 26866 S. Angelo Lodigiano (LO), Italy; e-mail: [email protected]
Summary
Résumé
Resumen
Classification of Italian maize
(Zea mays L.) germplasm
Classification du germoplasme
Italien de maïs (Zea mays L.)
Clasificación del germoplasma
Italiano de maíz (Zea mays L.)
The early introduction of maize from the
newly discovered Americas and the central location of the Mediterranean Basin
ensured the Italian peninsula an important role in maize secondary evolution
and regional diffusion. The introduction
of USA dent hybrids in the late 1950s
significantly changed Italian maize cultivation and led to the loss of traditional
varieties. In anticipation of such an event,
samples of many Italian maize landraces
were collected in 1954 and during subsequent years, to be maintained, studied
and used as sources of useful genes in
breeding programmes. Phenological
studies and germplasm relationships
were assessed through principal components analysis of 17 important phenological, morphological and geographical
characters for 562 Italian maize accessions. These were grouped into 65
agroecotypes; major clusters contained
34 landraces in 9 racial complexes of common ancestors and/or place of origin. A
brief description of the landraces and racial complexes is presented. The Italian
maize collection is permanently maintained ex situ at the Bergamo Section of
the Istituto Sperimentale per la
Cerealicoltura, a research Institute of the
Ministero delle Politiche Agricole e
Forestali.
La prompte introduction du maïs de
l’Amérique tout récemment découverte
et la localisation centrale de l‘Italie dans le
bassin Méditerranéen ont joué un rôle de
grande importance dans la différentiation
secondaire et dans la diffusion régionale
du maïs dans le pays. L’arrivée des hybrides dentés des Etats Unis a changé profondément dés la décade ’50 le panorama
des maïs italiens jusqu’à la presque désapparition des variétés traditionnelles. En
prévision de telle possibilité, à partir du
1954 on a récueilli la plupart des variétés
italiennes, dans l’objectif de leur conservation, étude et utilisation comme source de
gènes
utiles
au
programme
d’amélioration génétique. Dans cette perspective, 562 populations de maïs italien
ont été évaluées phénotipiquement: leur
classification a été déterminée à travers de
l’analyse des components principaux de
17 importantes caractères phènologiques,
morphologiques et géographiques. Les
populations étudiées ont été groupées en
65 agroecotypes; à niveau supérieur on a
identifié 34 landraces, dérivées de 9 complexes raciaux d’origine commune ou
d’hybridation par proximité. Une synthétique description des landraces et des complexes raciaux accompagne la présentation des tables descriptives des caractères
moyens des groups identifiées aux différents niveaux. La collection des maïs
italiens est conservée ex situ par la Section
de Bergamo de l’Istituto Sperimentale per
la Cerealicoltura du Ministère des Politiques Agricoles et Forestières.
La temprana introducción del maíz desde las recién descubiertas Américas y su
localización céntrica en el Mediterráneo
desempeñaron un importante rol en la
diferenciación secundaria y difusión regional del maiz en la península Italiana.
La llegada de los híbridos Dent de Estados Unidos en la década de los cincuenta
cambió profundamente el panorama de
los cultivos Italianos de maíz y condujo a
la desaparición de las variedades tradicionales. En previsión de este evento,
muestras de muchas landraces Italianas
fueron coleccionadas a partir de 1954,
para ser conservadas, estudiadas y utilizadas como fuentes de genes útiles en
los programas de mejoramiento. Con
esta finalidad, 562 muestras de maíces
Italianos fueron evaluadas fenológicamente y sus relaciones fueron determinadas a través del análisis de las componentes principales de 17 importantes
carácteres fenológicos, morfológicos y
geográficos. Las muestras fueron divididas en 65 agroecotipos; clusters mayores
evidenciaron 34 landraces, derivadas de 9
complejos raciales de antepasados u origen común. Se presenta una sintética descripción de las landraces y de los complejos raciales. La colección de maíces Italianos es conservada ex situ en la Sección
de Bergamo del Istituto Sperimentale per
la Cerealicoltura, un Instituto de investigación del Ministero delle Politiche Agricole e Forestali.
Key words: Agroecotype, germplasm,
Italy, landrace, population, racial
complex, Zea mays
Introduction
Maize (Zea mays L.) was introduced into Italy shortly after the
return of Columbus from his first voyage to the New World (1492–
93). However, due to the poor adaptation of Caribbean maize
cultivars to the 38–45° latitude photoperiod, the crop did not
spread until mid-1500, when better adapted varieties of the Everta
(popcorns) and Indurata (flint) groups were imported from subtropical and temperate regions of Central and South America.
The diffusion of maize in Italy and from Italy into southern
Europe was subsequently rapid: new environments, new uses and
crossing of cultivars of different provenance resulted in numerous
new populations particularly suited to the range of agroecological
environments. In the seventeenth and eighteenth centuries, white
Indentata maize cultivars (Caragua and Gourdseed beaked dents)
were introduced from the subtropics into the farms of eastern
Veneto. During the nineteenth century they were gradually substituted by higher yielding USA Corn Belt dent cultivars.
During four centuries the central position in the Mediterranean area conferred a special role on Italy as a hub of diffusion
towards central and eastern Europe (Brandolini 1969b, 1970,
1971) along the various commercial routes of those periods
(Messedaglia 1924). In the late 1950s maize landraces still predominated in Italian agriculture. Subsequently, however, the introduction of USA dent cultivars radically changed Italy’s
germplasm spectrum, substituting old varieties for new highyielding hybrids. Nevertheless, small populations of traditional
landraces continue to be grown in remote valleys by amateur
farmers, for use in making traditional foods such as polenta.
Maize classification
Maize variability, in terms of grain colour, texture and food use,
as well as general appearance, was appreciated and outlined by
both early explorers of America and by sixteenth and seventeenth
2
Plant Genetic Resources Newsletter, 2001, No. 126
century herbalists in Europe. Their descriptions of maize usually
included a simplified iconography of plant and ear type, as
observed in the ‘herbalist gardens’ of southern Europe.
The first systematic maize variety classification was attempted by Bonafous (1836, 1842), at a plurispecific level, based
on his experience of southern European types. This classification,
reduced to a subspecific level, was later (1885) amplified and
detailed by several European authors, including Harz, Koernicke,
Werner and Heuzé (cited by Succi’ 1931). A more broadly accepted classification was produced by Sturtevant (1899), who
based his grouping of USA varieties (at a subspecific level) on the
merceological type of the kernels. Kuleshov (1929) later used
such a classification as the basis for his studies on the variability
of maize in the Americas.
Sturtevant’s classification scheme and criteria were criticized by
Anderson and Cutler (1942) as not adequately representing the
existing abundant global diversity and for the monogenic basis of
some of the discriminating characters (Amylacea, Amilosaccharata,
Tunicata, Saccharata, Ceratina). Therefore, they proposed a more natural classification scheme based on a holistic consideration of genetic,
phenotypic, cytological, phenological and ethno-historical elements.
Recently, the introduction of molecular markers able to detect
genetic differences at the DNA level, has spurred new studies on
maize lines and cultivar relationships (Smith and Smith 1991, Livini
et al. 1992, Pejic et al. 1998). However, their application to landraces
has lagged behind because the intra-population variation found in
allogamous species hinders data analysis and interpretation.
Furthermore, the assessment of many morpho-physiological
characters in germplasm is important for an accurate measurement
of the differences between populations as well as for rapid assessment of their breeding potential. The description of many traits leads
to analysis and interpretation problems. For the study of such
morphologically complex samples, multivariate analysis combines
the capacity to provide a synthetic summary of the most relevant
traits and assessment of the relative importance of different characters to the total differences (Camussi 1979, Abadie et al. 1998).
Maize collection and classification in Italy
After fragmented attempts during the nineteenth (Bonafous 1836)
and early twentieth centuries (Venino 1916, Zapparoli 1939, Succi
1931), in 1954 the collection and classification of Italian maize
germplasm was begun in a systematic way. This was done through
a national acquisition programme for the Indentata and Indurata
types promoted by the senior author at the Stazione Sperimentale
per la Maiscoltura of Bergamo (Director: L. Fenaroli), under the aegis
of the Italian Ministry of Agriculture. Through the active co-operation of the Ispettorati Provinciali dell’Agricoltura, seed samples of
different populations from 65 major maize–producing provinces
were harvested and transferred to Bergamo, to undergo reproduction and classification studies (Brandolini 1958). The collection was
expanded during the following years, to fill possible gaps, and was
further studied for karyotype morphology (Bianchi et al. 1963,
Lorenzoni 1965) and for biochemical and texture composition of the
kernels (Brandolini 1967, 1969a, Camussi et al. 1980).
Brandolini and Mariani (1968) proposed classification of the
entire Italian collection by identifying and studying, on the basis
of morphological and historical information, a core-collection
comprising the ‘typical strains’ of each putative landrace.
Camussi (1979) and Camussi et al. (1980) confirmed this classification in principle through multivariate analysis of the quantitative characters of 102 core-collection samples.
A new step in the classification of the 562 accessions collected
is presented here, through multivariate analysis of 17 phenotypical and geographic characters important as hierarchic grouping
criteria for the Italian situation and accurately representing plant,
ear and adaptation characters.
Materials and methods
Accessions
Accessions (562) representing the Indurata and Indentata groups
were collected in 1954 from all over Italy (Fig. 1). These were grown in
1955 in nursery fields of the Stazione Sperimentale di Maiscoltura in
Bergamo, Italy, at 45°41’N, 9°37’E and 240 masl. The experimental
plots were 8 m long, 0.9 m wide and comprised 30 plants per plot.
Fig. 1. Collection sites of the 562 Italian maize accessions
discussed in this publication.
Traits scored
Thirty phenological, morphological and agronomic traits (five for
plant architecture, three for phenology, five for ear morphology,
seven for tassel morphology, six for grain morphology, three for
disease susceptibility and one set for internode pattern) were
measured at flowering and at harvest on ten representative plants
per plot. A preliminary correlation analysis of all the traits recorded suggested consideration of only the 14 most relevant and
independent plant, ear and geographic characters as well as three
morphological indices (Table 1). The semi-quantitative traits (kernel type and kernel colour) were coded by attributing them a
numeric value, taking into account the probable pairing of type
and colour of the kernels, as evident in each of the populations
studied, in order to provide meaningful averages.
Plant Genetic Resources Newsletter, 2001, No. 126 3
Table 1. Mean values and ranges of the variables measured in analysis of 562 Italian maize accessions
Ear length (cm)
Ear diameter (mm)
Row number
Kernel height (mm)
Kernel width (mm)
Kernel thickness (mm)
1000-kernel weight (g)
Female GDU
Plant height (cm)
Leaf number
Ear/plant height ratio
Leaf area (cm2)†
Conicity index‡
Kernel type§
Kernel colour¶
Latitude (°N)
Altitude (masl)
Mean
Standard
deviation
Minimum
Maximum
17.3
42.6
13.5
9.5
9.2
5.2
344.4
760.7
197.1
11.6
55.9
62.7
3.8
3.4
5.0
43.7
282.9
3.13
5.51
2.32
1.16
1.21
0.59
78.54
70.72
36.22
1.59
5.66
16.72
1.52
1.27
1.75
2.28
256.21
10.0
29.3
7.7
7.1
6.2
3.2
157.5
567.0
122.0
7.7
31.2
22.9
1.1
2.0
1.0
38.0
2.0
25.6
62.1
22.9
14.3
13.3
7.1
610.0
927.0
314.0
16.9
67.8
106.9
11.7
8.0
6.0
47.0
1275.0
†
¾ (leaf width x leaf length)
½ [(ear diameter at 2/3–ear diameter at 1/3) / 1/3 ear length]
§
1=floury, 2=semi-floury, 3=semi-flint, 4=flint, 5=semi-dent, 6=dent, 7=dent semi-rostrate, 8=flint rostrate
¶
1=white, 2=purple, 3=brown, 4=red, 5=yellow, 6=orange
‡
Statistical analysis
Mean values, standard errors and variation indices were computed
for each accession. The data matrix of 562 accessions and 17 traits
was standardized (mean=0 and variance=1) and analysed using
principal components analysis. Principal components (PC) were
calculated from the matrix of correlations between the characters,
and the original variables were transformed into a new set of
independent variables. To detect natural groupings in the collection, the Euclidean dissimilarity measures computed from this
new set of variables were clustered by the unweighted pair-group
method, arithmetic average (UPGMA). The resulting dendrogram
was pruned at a level that revealed most distinctive groups, but
still retained highly related cultivars within a single group. No
formal statistical criteria were used. The analysis was performed
using the statistical software Numerical Taxonomy and Multivariate Analysis System Version 1.70 (NTSYS-pc; Rholf 1993).
610 g, kernel height between 7.1 and 14.3 mm, width from 6.2 to
13.3 mm and thickness from 3.2 to 7.1 mm.
Growth environments ranged from the high alpine valleys and
slopes to the Padanian and coastal plains. A special case of
adaptation was represented by material from the Sicilian highlands, where several cultivars were cropped to 1275 masl at 39°N.
Results and discussion
Trait variation
The average, minimum and maximum values and standard
deviations for the 17 traits considered, for the entire Italian
collection, are presented in Table 1. Most characters showed ample
variation as a result of centuries of selection and adaptation to
different microenvironments and uses.
Plant height varied from 122 cm for those cultivars grown in
dry areas to 314 cm for those from the humid plains. Leaf number
ranged from 7.7 to 16.9, while the GDU at female anthesis was
between 567 and 927 and the leaf area between 22.9 and 106.9 cm2.
Ear type varied from true cylindrical to extra conical, with a
row number going from 7.7 to 22.9 and an ear diameter from 29.3
up to 62.1 mm. Average ear length was from 10 to 25.6 cm.
Figure 2 shows some ear types.
Kernel type and colour were very variable, with a strong
prevalence of flint (18%) and semi-flint (52%) types, generally
orange (64%), yellow (14%) or white (14%). Kernel shape and size
were also highly variable, with 1000-kernel weight from 157 to
Fig. 2. Ear morphology of some Italian maizes.
–0.11**
0.14**
–0.19**
0.37**
–0.18**
Kernel
colour
–0.30**
0.27**
–0.20**
0.08*
–0.52**
0.42**
–0.37**
0.45**
–0.28**
Kernel
type
Conicity
index
Leaf
area
0.56**
–0.23**
0.28**
–0.05
0.11**
–0.24**
Ear/plant
height
0.76**
0.61**
0.85**
–0.54**
0.46**
–0.33**
0.31**
–0.22**
0.55**
0.66**
–0.44**
0.35**
–0.24**
0.08*
–0.22**
Leaf
number
Plant
height
* and ** indicate statistical significance at P=0.05 and P=0.01, respectively.
–0.03
–0.01
–0.01
–0.14**
0.03
–0.07*
–0.36**
–0.16**
–0.20**
0.11**
0.12**
0.13**
0.11**
0.19**
0.15**
0.18**
0.21**
0.11**
0.18**
–0.05
0.68** 0.17**
0.71** 0.27**
0.54** 0.22**
0.35** 0.19**
0.69** 0.35**
–0.70** 0.14**
0.20** 0.07*
–0.28** –0.13**
0.16** 0.15**
–0.6
–0.07*
0.51**
0.64**
0.52**
0.35**
0.64**
–0.32**
0.42**
–0.42**
0.21**
–0.18**
0.41**
0.86**
–0.37**
0.48**
0.40**
0.56**
0.58**
Ear diameter
Row number
Kernel height
Kernel width
Kernel
thickness
1000-kernel
weight
Female GDU
Plant height
Leaf number
Ear/plant ratio
Leaf area
Conicity Index
Kernel type
Kernel colour
Latitude
Altitude
0.17**
–0.58**
–0.24**
0.55**
0.60**
0.24**
0.15**
0.17**
–0.04
0.41**
0.18**
–0.10*
0.25**
–0.25**
–0.32**
–0.36**
–0.31**
–0.32**
–0.34**
0.22**
–0.48**
0.08
–0.30**
0.22**
0.19**
0.30**
0.20**
0.05
0.36**
–0.22**
–0.10**
–0.30**
0.06
–0.02
0.78**
0.64**
0.56**
0.66**
–0.48*
0.31**
–0.35**
0.03
–0.09
Female
GDU
Kernel
1000thickness kernel
Kernel
width
Ear
Row
Kernel
diameter number height
Ear
length
Table 2. Correlation coefficients for morphological, agronomical and geographical traits measured in 562 Italian maize accessions
Trait association
Correlations among the 17 characters studied are presented in
Table 2. The large number of observations raised the test power,
giving significance to most of the correlations: hence, only values
above 0.6 are discussed. Several vegetative traits (plant height,
time to female anthesis, number of leaves, leaf area) appeared
highly correlated and were correlated with ear length. Tall, latematuring accessions had more and larger leaves and longer ears.
Plant height was also associated with ear/plant height ratio, as
expected in vigorous plants, and also with kernel height which
was also linked to leaf area. Ear length was negatively correlated
with conicity index: long ears were mainly cylindrical, while short
ears were generally conical. A high 1000-kernel weight was mostly
a consequence of thicker kernels.
–0.34**
Latitude
4
Clustering
Cluster analysis is often used to assess genetic diversity and to
classify species (Stanton et al. 1994, Rincon et al. 1996, Van
Beuningen and Busch 1997). In our study, the correlations existing between traits suggested the need to transform the variables
to have independent linear variables as input for cluster analysis.
Projection of the standardized original values on to the eigenvectors of the correlation matrix provides variable independence and
balanced weighting of traits. Generally, only PC with eigenvalues
larger than 1 are considered, but Jolliffe (1986) suggested retaining PC with eigenvalues as low as 0.75 if the input matrix is a
correlation type. Therefore, eigenvalues as low as 0.87 were considered in our study. Hence, six eigenvalues accounting for 82%
of the variation between populations, with the first two PC
explaining 52% of the variation, were considered and are presented in Table 3 along with correlation coefficients among eigenvectors and original variables.
The first eigenvector is mainly linked to the vegetative vigour
of the plant, as shown by its correlation with ear length, kernel
height, length of the vegetative cycle (GDU to female anthesis),
plant height, number of leaves, ratio between ear and plant
height, leaf area and, to a minor degree, kernel type and (negatively) conicity index. Populations with high scores for the first
eigenvector are, hence, late flowering, tall, leafy plants with long,
cylindrical ears and deep kernels. The second eigenvector is mostly
connected with kernel traits; kernel width, thickness and 1000kernel weight, as well as (negatively) with row number. Hence,
populations with large and heavy kernels are consequently characterized by a low number of rows. The third eigenvector summarizes the shape of the ear, since it relates to conicity index, ear
diameter and row number. The other traits are linked to the
remaining eigenvectors. In particular, the geographical origin of
the samples is mainly summarized by the fourth (correlated
mostly with latitude and altitude of the collection site) and by the
fifth (connected with altitude) eigenvectors, while kernel colour is
mainly associated with the sixth eigenvector.
Multivariate analysis of the Italian maize collection led to the
definition of several clusters. A preliminary UPGMA dendrogram, obtained from the Euclidean distances computed from the
data matrix obtained by the projection of the original scores on to
the eigenvectors (not presented) revealed the existence of major
similarities among accessions. Therefore, accessions clustering in
Table 3. Correlation coefficients among original values and eigenvectors for a principal components analysis
of important traits in 562 Italian maize accessions (eigenvalues and cumulative proportions of variance are
also reported)
Eigenvectors
Ear length
Ear diameter
Rows number
Kernel height
Kernel width
Kernel thickness
1000-kernel weight
Female GDU
Plant height
Leaf number
Ear/plant height ratio
Leaf area
Conicity index
Kernel type
Kernel colour
Latitude
Altitude
Eigenvalues
Cumulative proportion
of total variance
1
2
3
4
5
6
0.75
0.38
0.17
0.76
0.03
–0.40
0.38
0.80
0.93
0.78
0.63
0.91
–0.61
0.53
–0.44
0.36
–0.29
5.97
0.35
0.24
–0.25
–0.55
–0.22
0.96
0.69
0.86
–0.02
–0.03
–0.03
–0.20
0.01
–0.09
–0.43
–0.23
–0.27
0.20
2.95
0.52
0.28
–0.85
–0.67
–0.35
–0.01
–0.16
–0.18
0.18
–0.07
0.08
–0.02
–0.03
–0.54
0.00
–0.07
–0.17
0.10
1.81
0.63
0.15
0.11
0.27
–0.08
–0.08
0.11
–0.16
0.35
0.09
0.24
0.31
-0.08
0.07
–0.26
0.30
–0.67
0.52
1.39
0.71
0.05
0.05
0.15
0.18
–0.10
–0.09
–0.04
0.04
–0.05
–0.16
–0.45
–0.06
–0.16
0.29
–0.42
0.03
0.60
0.96
0.77
0.32
0.07
0.15
–0.19
–0.02
0.17
0.05
–0.12
0.05
–0.14
–0.10
0.12
–0.29
–0.05
0.52
0.44
0.25
0.87
0.82
the terminal branches of the tree were considered to be similar or
duplicate samples and were merged into 65 basic entities, after
Gregor (1931, 1933), agroecotypes: population units, differing in
adaptive characters, with a narrow genetic base but having certain genetic characters in common with the other varieties. Each
agroecotype included a minimum of 1 and a maximum of 26
accessions. The means and standard errors for each original
variable are presented in the Appendix. Identification of the
agroecotypes appears strongly based on their phenological performance as well as on regional diffusion of the samples.
A second-level principal components analysis (Fig. 3), performed on the mean values of the agroecotypes, provided evidence of hierarchical relationships between and among
agroecotypes, which resulted in a further grouping at a level of
races or, more accurately, landraces. These are related individuals with enough broad-based characteristics in common to permit their recognition as a group (Anderson and Cutler 1942),
maintained through panmictic reproduction in the populations
and occupying defined areas (Brieger 1950, Brieger et al. 1958).
At an even higher level evidence of racial complexes (RC) was
found. These are broader groups formed by a number of races
having some decisive characters in common: morphological,
locational and/or phylogenetical (Brieger, 1950, Brieger et al.
1958).
The relationships within and among landraces and their attribution to different racial complexes are presented in the plot of
first and second principal components (Fig. 4). The average values and standard errors associated with measured variables for
the 34 landraces, grouped according to the racial complexes, are
presented in the Appendix.
Landraces
A detailed description of each landrace and of its specific
agroecotypes will be reported in a more comprehensive
publication.We provide here a very synthetic description of the
OttofileN
Der10-12f
OttofilePen
Tajolone
OttofileTar
Meliun
Cannellino
Granturchella
Nostrale
Monachello
Barbina
Cavolone
Bufano
Maggese
Cinquantino
Poliranghi
Maggengo
Spadone
Culaccione
Ciociarino
Montano
Costarolo
Biancone
CinquantinoB
Montoro
BiancoSud
Rodindia
Pannaro
Primitivo
Trentinella
Primaticcio
Dindico
Lucano
Altosiculo
Poliota
Paesan
Trenodi
Nanoprecoce
Agostinello
Tirolese
Zeppetello
Pufano
Trecchinese
Marano
Cinquantino
Brigantino
Quarestivo
Cadore
Ostesa
Pignolino
Pignolo
Scagliolo
Rostrato
NostranoIn
NostranoVe
BaniScaiola
ZamengoB
BiancoPerla
RighettaB
DentatoB
RostratoB
DentatoBII
Cimalunga
DentatoG
DentatoSc
0.00
0.40
0.79
Coefficient of dissimilarity
1.19
1.58
Fig. 3. Dendrogram for 65 Italian maize agroecotypes. Pruning levels for landraces (solid line) and for racial complexes
(broken line) are indicated.
6
1.00
BaniScaiola
Pignolo
Cadore
0.55
Zeppetello
Marano
Rostrato
Scagliolo
Ostesa
Quarantino estivo
PC2 (17%)
Nostrano
dell’Isola
Pannaro
Barbina
0.10
Agostinello
Trenodi
Poliota
-0.35
Biancone
Dentati antichi
Poliranghi
Montano
Dindico
Trentinella
Altosiculo
Dentati moderni
Rodindia
Monachello
Cannellino
Montoro
Cimalunga
Bianco Perla
Derivato 10-12
Ottofile
Ottofile tardivo
Tirolese
Righetta bianca
Tajolone
-0.80
-1.20
-0.55
PC1 (35%)
0.10
0.75
1.40
Fig. 4. Plot of principal components 1 and 2 for a group of Italian maize landraces. The lines group landraces belonging to
the same racial complex.
races included in each racial complex, with an indication of their
regional localization and phylogenetic relationships.
Of the three major groups (subspecies, after Sturtevant 1899)
introduced into Italy during the last four centuries, the Everta did
not play a very important role, due to small-scale production and
to genetic isolation mechanisms. They are unable to accept pollen
from other subspecies, allowing survival as a pure crop in farms
and gardens, limiting passive intercrossing. However, some active pollination contributed to the building of a number of the
Indurata landraces, of the Microsperma racial complex, which
even today maintain three main characteristics inherited from the
Everta introgression: a very small kernel, an extra-hard texture
and a tendency to multiple ear-bearing. A multivariate test performed on major Everta strains and on representative
Microsperma landraces confirmed, however, the absence of other
significant similarities (data not presented).
Traditional maize varieties from the various regions of Italy
belong mainly to the Indurata group. Seventy percent of the
samples studied had a flint or semi-flint kernel texture. Only few
landraces, mainly grown in southern and eastern Veneto, can be
traced back to Meso-American beaked dents (Rostrato bianco)
and, in the late nineteenth century, to the dent varieties of the
USA Corn Belt. A special type of dent maize—the Scaiola—is
included as a dent ancestor in the Rostrato-Scagliolo group of the
Insubrian racial complex, but may be included in the Gourdseed
varieties of the southern USA. It was identified by Venino (1916)
as an import from Philadelphia.
In general we recognize within the Italian Indurata maizes
several phenological characters that permit tracing their origin to
different regions of the Americas.The landraces are the result of
their mixing and intercrossing. There is historical evidence that
maize cultivation in Italy began only in the second half of the
sixteenth century, notwithstanding the documented earlier introduction of maize seeds (Martire d’Anghiera 1514). Maize types
from the subtropics (West Indies, Central America) originally
met serious obstacles in their photoperiodic adaptation to 38–
45°N.Maize cropping was thus possible only when samples from
higher latitudes or the cold highlands of the Cordillera, and/or
genetic materials insensitive to daylength, were received. Again,
the long shipment routes favoured survival of hard, horny seeds
of the Indurata group (Morocho type), more resistant to moulds
and insects, and easily accepted in regions where the milling
activity was millenarian.
In addition to the detailed presentation of phenological and
adaptation data included in the Appendix, a synthesis of the
different complexes and identified landraces is provided, with a
short description and a few comments on evidence of relationships.
RACIAL COMPLEXES:
(A) OTTOFILE VITREI E DERIVATI
EIGHT-ROWED FLINTS AND DERIVED RACES
Agroecotypes of this racial complex are present in every Italian
region and are adapted to a range of agroclimatic conditions.Their
adaptation ranges from extra-early maturity in the dry regions of
central and southern Italy up to a full season cycle in the Padania
region and in high rainfall or irrigated locations of southern Italy
and Sicily. In situations of proximity to other RC (conical flints,
Microsperma flints) many derived forms (10–12–14 rowed flints)
exist as a result of the introgression of neighbouring racial groups
and appear to accompany the true eight-rowed forms, especially
in central Italy.
The following landraces, presented in two groups can be
included in this complex:
(D) CILINDRICI VITREI MERIDIONALI DI CICLO MEDIO
MID-SEASON SOUTHERN CYLINDRICAL FLINTS
These derive from southern Italy and the highlands of Sicily, this
RC is characterized by a mid-season growing cycle and mediumshort plants with rather reduced leaves.
Landrace
Landrace
Description
True-bred eight –rowed flints
(1) Ottofile
8-row
semi-flint
(3) Tajolone
large grain
8-row semifloury
(4) Ottofile
long-eared
tardivo
8-row flint
Derived landraces
(2) Derivati
8-row
10–12 file
derived
semi-flint
(5) Cannellino
long-eared
semi-flint
(6) Monachello long-eared
semi-flint
No.
entries
Region
Kernel
size
Maturity
14
large
medium
3
Northcentral
North
extralarge
medium–
late
6
North
large
medium–
late
7
central
large
medium
26
Northcentral
South
large
medium–
late
medium–
late
8
medium
(B) CONICI VITREI E DERIVATI
CONICAL-EARED FLINTS AND DERIVED RACES
A large number of accessions with traits suited to non-irrigated
plains and hill slopes of central and northern Italy, bearing conical
or subconical ears with medium sized, isodiametric, thick grains.
Bonciarelli (1961) noted that the large conical cob, rich in soft
parenchyma, is an adaptation to low-moisture growing conditions, where its structure functions as a water reservoir able to
maintain stigma turgidity during periods of high-transpiration.
An extreme form, the Ostesa landrace, characterized by extreme
conicity index (11.7) and high row number (18.8), can be included
in this RC.
Landrace
Description
No.
Region
entries
(7) Barbina
conical eared
79
Northyellow flint
central
(8) Poliranghi multi-rowed
65
Northyellow semi-flint
central
(9) Montano subalpine
17
North
flint
(10) Biancone Apennine
7
Central
yellow flint
(25) Ostesa
extra-conical
1
North
yellow flint
Maturity
Description
(11) Montoro
long-eared
white semi-flint
long-eared
8
orange semiflint
long-eared
7
Microsperma
(12) Rodindia
(13) Pannaro
No.
entries
13
Region
No.
entries
(14) Trentinella subcylindrical 33
yellow semiflint
(15) Dindico
yellow-red
22
semi-flint
(16) Altosiculo long
5
subcylindrical
yellow semiflint
Landrace
Description
(17) Poliota
extra-early
6
hills flint
extra early
23
Tyrrhenian flint
Alpine valleys
3
early flint
(18) Tre nodi
(19) Tirolean
No.
entries
Landrace
medium
medium
a) Apennine
(20) Agostinello early conical
orange flint
(21) Zeppetello early conical
yellow flint
b) Subalpine orange flint
(22) Marano
prolific
orange flint
(23) Quarantino extra-early
estivo
summer
orange flint
(24) Cadore
early multi-row
conical flint
medium–
early
medium medium
medium
medium
South
Kernel
size
large
South
large
South
small
Maturity
medium–
late
late
medium–
late
This last landrace is distinguished from the others by its
characteristic small seed (Microsperma).
South
Kernel
size
medium
South
medium
Maturity
medium–
late
medium–
late
Sicilian
medium medium–
highlands
late
Region
Kernel
size
Maturity
central
medium
central
medium
Alpine
valleys
medium
extraearly
extraearly
extraearly
(F) MICROSPERMA VITREI
MICROSPERMA FLINTS
These are cropped in northern and central Italy and include
landraces with small, hard, horny textured seeds. The seeds are
generally yellow, orange or reddish, and the plants are relatively
short with low ear insertion and reduced leaf area.
medium
medium
Region
(E) NANI PRECOCISSIMI VITREI
EXTRA-EARLY DWARF FLINTS
These are present in the mountain valleys of North and central
Italy, as well as in the plains of the Tyrrhenian coast. The group is
composed of three landraces characterized by extreme earliness
and reduced plant and ear size.
Kernel
size
medium
(C) CILINDRICI MERIDIONALI TARDIVI
LATE SOUTHERN CYLINDRICAL FLINTS
This is a RC characterized by lines with long cylindrical or
subcylindrical ears, a medium-late to late growing season, large
grains and large, leafy plants. The following landraces are ascribed to this group:
Landrace
Description
Description
No.
entries
Region
Kernel
size
Maturity
7
small
early
14
centralSouth
South
small
early
41
North
small
medium
7
North
small
extraearly
4
North
small
early
The Pignolo landrace, which is placed in the following RC but is
of typical Microsperma size, could be also related to this group.
(G) INSUBRICI VITREI E SEMI-VITREI
INSUBRIAN FLINTS AND SEMI-FLINTS
This complex is the result of convergent adaptation to a particular agrosystem centred in the peneplains of the InsubrianEuganean region, where maize found a preferred habitat. It
contains genetic material that can be traced back to various
American sources and races. Their proximity and easy intercrossing generated intermediate forms, especially in the ’elliptical
seed‘ group [Scaiola (Dent) x Pignoletto x Rostrato].
8
Two main subsections exist:
Landrace
Description
No.
Region
entries
a) multi-rowed, elliptical seeded
(26) Pignolo
elliptical seed 5
orange flint
(27) Rostrato deep grain
19
Scagliolo
orange semiflint
(29) Scaiola
gourd seed
3
yellow dent
b) long-eared, isodiametric (square) seeds
(28) Nostrano long
39
dell’Isola
cylindrical
flint
Kernel
size
Maturity
Euganean
hills
Insubria
small,
deep
small,
deep
mediumlate
medium–
late
Insubria
oblong, medium–
deep
late
Transpadania cubic
medium–
late
The Nostrano dell’Isola landrace, widely grown in different parts of
Italy, originally developed in the sub-Alpine region of Bergamo
province. It is characterized by a medium-late growing cycle and
by a typical long ear with enlarged butt and isodiametric orange
flint grains. It can be traced back to the Caribbean cylindrical
maizes. Similar types are endemic in other maize-growing countries, of southern Europe, including Portugal, Spain and Romania.
Bianchi veneti – Veneto white landraces
Due to historical and commercial routes, the diversification
of maize in the eastern and southern Venetian provinces followed
specific paths from the outset of its introduction, both within the
white dent group and the white flint group. It is thus possible to
distinguish two major sections (white and yellow dents, and
white pearls).Convergent evolution gave rise to landraces
(Cimalunga and Righetta Bianca) combining dent or pearl seed in
different plant types.
(H) BIANCO PERLA
PEARL WHITE FLINTS
This is a large group of accessions mainly found in the southern
Venetian plains, characterized by large cylindrical ears and white
grains of pearly appearance, very similarin many traits, to the Pearl
White group of varieties collected and studied in several Latin-American countries. The Italian collection includes three agroecotypes:
Cimalunga, Righetta Bianca and Bianco Perla, differing mainly in their
kernel type and maturity. The Bianco Perla landrace includes
introgressed Hickory King strains of recent importation.
Landrace
Description
(30) Bianco
Perla
(31) Righetta
Bianca
(33) Cimalunga
pearl white
semi-flint
eight-rowed
white dent
long-eared
white semiflint
No.
entries
25
Region
Veneto
Kernel
size
medium
4
Veneto
huge
6
Veneto
medium
Maturity
medium–
late
medium–
late
late
(I) DENTATI BIANCHI
WHITE DENTS
This RC includes agroecotypes of the Indentata group, either as
ancient introductions from Central America or from recent imports as high-yielding open-pollinated varieties from the USA.
The ancient dents include both the beaked white dents (extra
floury) from Mesoamerica and the equally late-maturing dent
varieties from Central America and southern USA. The modern
dents include open-pollinated varieties (white and yellow) imported from the USA during the first half of the twentieth century
and the new local varieties developed by Italian technicians from
Indurata x Indentata hybridization in the period between the two
world wars.
Landrace
Description
(32) Dentati
antichi
(34) Dentati
moderni
ancient
dentsdents
modern
dents O.P.
No.
entries
19
16
Region
eastern
Veneto
Po plain
Kernel
size
medium
large
Maturity
medium–
late
medium–
late
Some white dents were also grown in Novara province,
Piedmont.
Conclusions
The multivariate analysis of the major morphological, phenological and adaptive characteristics of the Italian Maize Collection
allowed a realistic clustering of the 562 entries into hierarchical
groups of different levels: 65 agroecotypes, 34 landraces and,
finally, 9 racial complexes. The classification confirms, at a phenotypic level, the relationships advanced by Brandolini and
Mariani (1968). A better and deeper knowledge of the different
types and of their similarities, supported by historical records,
Table 4. Presence of accessions, agroecotypes, landraces and racial complexes in the Italian administrative regions
Administrative region
Accessions
Agroecotypes
Landraces
Racial complexes
Piemonte
Liguria
Lombardia
Trentino-Alto Adige
Veneto
Friuli
Emilia Romagna
Toscana
Marche
Lazio
Abruzzi
Campania
Puglia
Basilicata
Calabria
Sicilia
Sardegna
34
17
42
24
91
35
26
72
45
51
32
27
9
23
4
25
5
14
12
18
12
28
16
16
19
12
20
13
12
7
8
4
9
4
11
11
12
9
17
9
11
10
8
13
8
8
6
7
2
6
4
5
4
6
5
5
5
6
5
4
8
4
5
3
5
2
3
2
allowed highlighting of several similarities as well as systematic
differences. Furthermore, it was possible to identify some
agroecotypes and landraces bearing ’primitive‘ or more evolved
traits, for reasons of similarity with American races of known
ancient origin. Finally, it was possible to trace a number of
evolutionary paths followed by maize in the course of five centuries under the multifarious conditions that characterize Italy.
The information contained in Table 4 highlights the distribution of the accessions, agroecotypes, landraces and racial complexes with regard to their presence in the various administrative
regions of Italy.
In conclusion, we would like to emphasize the importance of
the Italian maize germplasm during the four centuries after its
introduction in Europe. Italian naturalists and farmers received
germplasm from America, via Spain and Portugal and, later,
from the European regions of the Turkish Empire as well as
directly from Italians in America. This maize germplasm, after
adaptation to temperate climates, was consciously transferred,
via trade routes and technical exchange, to central Europe and
the Balkans (Pavlicic and Trifunovic 1967) and to North Africa.
During the Napoleonic era another exchange route included France
and Switzerland: Bonafous worked both in Paris and Turin.
Finally, in the twentieth century the Stazione Sperimentale per la
Maiscoltura of Bergamo, the first institution purposely created in
Europe for maize improvement, collaborated openly with many
maize researchers and freely exchanged germplasm, experiences
and human resources.
Acknowledgements
We would like to thank Drs C. Elitropi, G. Mariani, G. Orio and G.
Vandoni for their precious collaboration, valuable suggestions
and priceless friendship.
References
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de Andrade. 1998. A classification for Brazilian maize
landraces. Plant Genet. Resour. Newsletter. 114:43-44.
Anderson, E. and H. Cutler. 1942. Races of maize: their recognition and classification. Ann. Mo. Bot. Garden 29:69-88.
Bianchi, A., M.V. Ghatnekar and A. Ghidoni. 1963. Knobs in
Italian maize. Chromosoma 14:601-617.
Bonafous, M. 1836. Histoire naturelle agricole et économique du
maïs – Paris.
Bonafous, M. 1842. Annali della Società Agraria di Torino. Torino,
Volume 2.
Bonciarelli, F. 1961. Studio agronomico comparato della
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Brandolini, A. 1958. Il germoplasma del mais e la sua
conservazione. Maydica 3:4-14.
Brandolini, A. 1967. Examples of plant exploration: Zea mays L.
Technical conference on Gene Resources in: IBP Handbook
no.11, FAO, Rome, Italy. Pp. 273-309.
Brandolini, A. 1969a. Preliminary report on South Europe and
Mediterranean maize germplasm. Proc. V Meeting of the
maize and sorghum sections of Eucarpia. Martonvasar, Hungary. Pp. 108-118.
Brandolini, A. 1969b. European races of maize. XXIV corn and
sorghum research conference. American Seed Trade Association. Chicago, USA. 24:36-48.
Brandolini, A. 1970. Razze europee di mais. Maydica 15:5-27.
Brandolini, A. 1971. Report of the Southern Europe-Mediterranean committee, Proc. VI Meeting of the maize and sorghum
sections of Eucarpia, Freising-Weihenstephan, Germany. Pp.
57-93.
Brandolini, A. and G. Mariani. 1968. Il germoplasma italiano
nella fase attuale del miglioramento genetico del mais.
Genetica agraria 22:189-206.
Brieger, F.G. 1950. The genetic basis of heterosis in maize. Genetics 35:420-445.
Brieger, F.G., J.T.A. Gurgel, E. Paterniani, A. Blumenschein and
M.R. Alleoni. 1958. Races of maize in Brazil and other Eastern
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Washington. No. 593, Pp. 1-283.
Camussi, A.1979. Numerical taxonomy of Italian populations of
maize based on quantitative traits. Maydica 24:161-174.
Camussi, A., M.D. Jellum and E. Ottaviano. 1980. Numerical
taxonomy of Italian maize populations: fatty acid composition and morphological traits. Maydica 25:149-165.
Gregor, J.W. 1931. Experimental definition of species. New
Phytology 30:204-217.
Gregor, J.W. 1933. The ecotype concept in relation to registration
of crop plants. Ann. Appl. Biol. 20:205-219.
Jolliffe, I.T. 1986. Principle component analysis. Springer Verlag,
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Kuleshov, N.N. 1929. The geographical distribution of the varietal diversity of maize in the world. Bull. Appl. Bot. Genet.
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Livini, C., P. Ajmone-Marsan, A.E. Melchinger, M.M. Messmer
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10
Appendix
12
Plant Genetic Resources Newsletter, 2001, No. 126 Plant Genetic Resources Newsletter, 2001, No. 126: 12 - 16
ARTICLE
Collecting landscape trees and shrubs in Ukraine
for the evaluation of aesthetic quality and
adaptation in the north central United States†
Mark P. Widrlechner1, Robert E. Schutzki2, Vasily Y. Yukhnovsky3,
and Victor V. Sviatetsky3
USDA-Agricultural Research Service, North Central Regional Plant Introduction Station, Iowa State University,
Ames, Iowa 50011-1170, USA
2 Department of Horticulture, Michigan State University, East Lansing, Michigan 48824-1325, USA
3 Department of Forestry, National Agricultural University, Kyiv, 252041 Ukraine
1
Summary
Résumé
Resumen
Collecting landscape trees and
shrubs in Ukraine for the
evaluation of aesthetic quality
and adaptation in the north
central United States
Un assemblage des arbres et
arbustes ornementaux dans
l’Ukraine pour l’évaluation de
l’esthétique et de l’adaptation
dans la région nord-centrale
des États-Unis
Colección de árboles y
arbustos de paisaje en Ucrania
para la evaluación de la
calidad estética y adaptación
en la parte central norte de
Estados Unidos
Certaines expériences avec les évaluations à long terme des plantes ligneuses
de la Slovénie, de la Croatie, et de la
Bosnie-Herzégovine dans la région
nord-centrale des États-Unis ont indiqué
qu’une portion relativement petite de
ces introductions a été bien adapté aux
climats et terroirs régionaux. Fondé sur
ces résultats, quelques critères ont été
développées
afin
de
diriger
l’exploration future pour les arbres et
arbrustes ornementaux des environnements plus analogues dans l’Europe orientale et centrale. L’application de ces
critères a reconnu la zone de transition
entre le bois et la steppe dans l’Ukraine
du nord comme une région du potentiel
considérable, à cause des ressemblances
à la région nord-centrale des États-Unis
en les extrêmes climatiques, les types de
sols, et les associations naturelles de la
végétation. En 1999, le Système National du Matériel Génétique Végétal a financé une mission collaborative afin de
rassembler les semences, engageante
les chercheurs du Département
d’Agriculture
des
États-Unis,
l’Université de l’État du Michigan, et
l’Université Agricole de l’Ukraine.
L’excursion d’exploration a eu lieu entre
le 7 et le 26 septembre 1999 et a embrassé vers 3200 km des voyages aux
bois à travers la zone de transition entre
le bois et la steppe. Nous avons obtenu
89 échantillons de semences, contenant
26 genres et 45 espèces d’arbres, arbustes, et herbacées plantes vivaces.
Dans notre rapport, nous décrirons la
mission, ses collections, et les conditions
aux lieux de collection.
Experiencias en el pasado con evaluaciones de largo plazo en plantas leñosas
de paisaje provenientes de Eslovenia,
Croacia y Bosnia-Herzegovina en la
parte central norte de Estados Unidos,
han indicado que una proporción relativamente baja de estas introducciones
estuvieron bien adaptadas a las condiciones climáticas y de suelo. Con base en
estos resultados, se desarrollaron criterios para enfocar futuras exploraciones
de árboles y arbustos de paisaje de ambientes más análogos en el este y centro
de Europa. Con la aplicación de estos
criterios se identificó a la zona de transición bosque–estepa en la mitad norte
de Ucrania como una región con gran
potencial, debido a similitudes con la
parte central norte de Estados Unidos
con respecto a los extremos climáticos,
tipos de suelo y comunidades naturales
de plantas. En 1999, el Sistema Nacional
de Germoplasma Vegetal de los Estados
Unidos financió una misión colaborativa para colección de semilla involucrando investigadores del Departamento de
Agricultura de Estados Unidos, la Universidad Estatal de Michigan y la Universidad Agrícola Nacional de Ucrania.
El viaje de exploración se realizó entre el
7 y el 26 de septiembre de 1999 y abarcó
aproximadamente 3200 km de recorrido a sitios arbolados a través de la zona
de transición bosque-estepa. Se obtuvieron ochenta y nueve colecciones de
semilla, incluyendo 26 géneros y 45 especies de árboles, arbustos y herbáceas
perennes. En este reporte se describe la
misión, sus colecciones y las condiciones
de los sitios de colecta.
Past experiences with long-term evaluations of woody landscape plants from
Slovenia,
Croatia,
and
BosniaHerzegovina in the north central United
States indicated that a relatively low
proportion of these introductions were
well adapted to climatic and soil conditions. Based on these results, criteria
were developed to focus future exploration for landscape trees and shrubs
from more analogous environments in
eastern and central Europe. Application
of these criteria identified the forest–
steppe transition zone in the northern
half of Ukraine as a region with great
potential, because of similarities to the
north central United States in climatic
extremes, soil types and natural plant
communities. In 1999, the National Plant
Germplasm System of the United States
funded a collaborative seed-collection
mission involving researchers from the
United States Department of Agriculture, Michigan State University and the
National Agricultural University of
Ukraine. The exploration trip took place
between 7 and 26 September 1999 and
encompassed ca. 3200 km of travel to
wooded sites through the forest–steppe
transition zone. Eighty-nine seed collections, including 26 genera and 45 species
of trees, shrubs and herbaceous perennials were obtained. The mission, its collections and conditions at collection sites
are described in this report.
Key words: Climatic analogue, forestry,
genetic resources, ornamental, plant
community, plant exploration
†
Journal Paper No. J-19028 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa, Project No. 1018,
and supported by Hatch Act and State of Iowa.
Introduction
The north central United States is a region of climatic extremes,
and many parts of the region have alkaline soil that developed
under grasslands. Because of these conditions, the diversity found
in commercially available woody plants adapted to the region is
considerably less than that found in many other parts of the
United States.
Eastern and central Europe are potentially important sources
of well-adapted landscape plants for the nursery industry in the
United States. Many commonly produced shade trees and shrubs
cultivated in urban areas in the eastern United States, such as
Acer campestre and platanoides, Ligustrum vulgare, Quercus robur
and Tilia cordata, are native throughout much of Europe, but are
thought to be primarily of western European provenance because
of previously restricted access to regions further east. Western
European sources are often poorly adapted to the climatic and
edaphic stresses found in the north central region, leading to
plant loss and frequent replacement. But the native ranges of
many valuable European landscape plants extend east into the
more continental climates and grassland soils of central and
eastern Europe, creating opportunities to acquire and evaluate
plants that may be of direct utility in the north central region and
that should also serve as a reservoir of stress-tolerant genotypes
for plant improvement research.
The most comprehensive landscape-plant evaluation program in the north central region is the NC-7 Regional Ornamental
Plant Trials, which were begun in 1954 with the ultimate goal of
expanding the range of useful plants in the nursery trade in the
north central region (Widrlechner 1990). The emphasis in these
trials is placed on detailed, long-term evaluations across a diverse array of sites and the broad sharing of such performance
data rather than on direct promotion of new plants, with results
made available to horticultural professionals and the general
public via the Internet (Becker 2000). One of the first opportunities to evaluate landscape plants from central Europe occurred in
the early 1970s, when a U.S. government-sponsored project resulted in extensive collections of horticultural germplasm
throughout former Yugoslavia. Many of the tree and shrub collections made by that project were evaluated in the 1970s and 1980s
in the NC-7 Regional Ornamental Plant Trials. A detailed analysis of the performance of these plants in relation to climatic
variables at trial sites (Widrlechner et al. 1992) indicated that a
relatively low proportion of Yugoslavian plants were well
adapted in the north central region and that climates at the
collection sites were not analogous to those at the trial sites.
Based on these results, criteria were developed to focus a
search for better-adapted landscape plants from more analogous
environments in eastern and central Europe (Widrlechner 1994a,
1994b). It became obvious from this research that the northern
half of Ukraine met important climatic criteria and needed to be
examined more closely. Thus, in 1994, Widrlechner developed a
list of woody plants native to the area in question and circulated
that list widely among American botanical gardens, nursery
professionals and academic horticulturists, as part of a survey to
identify target species for collection. In 1995, more detailed climatic data (Slabkovich 1968) and soil maps (Anonymous 1960;
Ganssen and Hädrich 1965) were obtained for Ukraine. And in
1996, distribution maps for many of the target species were
located (Sokolov et al. 1977–1986). Taken together, the results of
these efforts indicated that an exploration focusing on native tree
and shrub populations adapted to the transition zone between
the central European deciduous forest and the Ukrainian steppes
should be most productive.
Most of the woody vegetation in the forest–steppe transition
has been cleared for agriculture. A figure in Sheljag-Sosonko et al.
(1982) indicates that remnant forests constitute 10% or less of the
historic forest–steppe transition zone. Ukraine is densely populated (>100 inhabitants/km2 in the target region) and, as landuse patterns change with changing economic systems, native
vegetation will likely face new threats from both urbanization
and the modernization of agricultural practices. The nation’s two
largest cities, Kyiv and Kharkiv, are also located in the target
region.
A status report of the Woody Landscape Plant Crop
Germplasm Committee (1996) identified Ukraine as a geographic
priority for exploration. A survey of the Germplasm Resources
Information Network (GRIN) database (http://www.arsgrin.gov/npgs) and of U.S. arboreta and botanical gardens also
indicated that there was almost no landscape plant germplasm
available from the target region. Academic exchange and collaboration agreements between Iowa State University (ISU) and the
National Agricultural University of Ukraine (NAUU) led to the
development of our team to conduct the exploration in 1998. We
then collectively developed an itinerary for exploration to sample
a broad range of sites on an east–west gradient ranging from
small outliers of woody vegetation within the steppe zone in the
east to rather diverse forests in the west. Sites focused on the
forest–steppe transition zone in U.S. Department of Agriculture
(USDA) hardiness zone 5 (mean minimum annual temperature
between –23.4 and –28.9°C) with grassland and/or brown forest
soils and moderate moisture deficits, avoiding podzolic soils in
the north, cool, moist habitats of the Carpathian Mountains, and
warm summer, mild winter habitats near the Black Sea.
Expedition and samples collected
The exploration trip was facilitated by an established collaboration
between ISU and NAUU and was also aided by a Memorandum of
Understanding between the National Plant Germplasm System
(NPGS) of the United States and the National Center for Plant
Genetic Resources of Ukraine. The trip took place between 7 and 26
September 1999 and encompassed approximately 3200 km of
travel to collection sites (Fig. 1). NAUU served as our base, with
excursions to the east and west. The first excursion went east
through Pryluky towards Okhtyrka, southeast to Kharkiv, southwest through Poltava and Cherkasy to Uman and finally returned
north to Kyiv. A second excursion involved traveling west through
Zhytomyr to Rivne, south through Kremenets to Ternopil, southeast through Khmelnytskyi to Vinnitsa and returning northeast to
Kyiv.
Military maps of each oblast (state) in 1:200 000 scale were
extremely helpful in locating natural forests, potentially interesting topographic features, and navigating through both the cities
and countryside. More exact locations for collection sites were
verified by the use of a Global Positioning System (GPS) receiver
14
Fig. 1. Route map.
and occasional comparisons of elevational contours with a handheld altimeter. Positional data were also verified upon return to
the U.S. by comparison with coordinates held by the GEOnet
Names Server (http://164.214.2.59/gns/html/).
Plant exploration focused on the forest–steppe transition
zone. Eighty-nine seed collections (most with herbarium vouchers) including 26 genera and 45 species of trees, shrubs and
herbaceous perennials were obtained (Table 1). Collections were
made between 49°14’ and 50°48’ north latitude and between
25°43’ and 35°48’ east longitude. Elevations ranged from 80 to
370 m above sea level. Based on climatological data, collection
sites correspond to USDA winter hardiness zones 5a and 5b
(mean minimum temperatures between –23.4 and –28.9°C). Mean
annual rainfall is highest in the west and decreases towards the
east/southeast. Rivne, Kremenets and Ternopil receive ca. 575
mm of annual precipitation; Khmelnytskyi is between 550 and
575 mm; Zhitomir, Kyiv, Pryluky, and Sumy between 525 and
550 mm; Vinnitsa and Okhtyrka between 500 and 525 mm; and
our driest collection areas were 475–500 mm through Kharkiv,
Poltava, Cherkasy and Uman. Annual rainfall for 1999 was at
76% of normal levels according to the Agricultural Attaché at the
U.S. Embassy in Kyiv, with slight regional variations. The crops
and natural vegetation in the eastern oblasts often showed visible
signs of drought stress, especially on sandy sites.
Soils throughout the region can be generally described as
transitional between podzolic forest soils, more specifically, grey
forest soils, and chernozem soils of the steppes. Soil texture
classifications were predominantly sandy loams and clay loams,
with isolated areas of sands near Poltava, Okhtyrka and
Cherkasy.
Natural vegetation occupies approximately 18 million ha in
Ukraine, with 8 million in forests, 1 million in steppes and the
balance in meadows and marshes. Sixty-five percent of the for-
ests are in the Carpathian Mountains. In our travels, we encountered five basic forest types. The first was Quercus–Carpinus
forest, with predominant species including Quercus robur, Carpinus
betulus, Corylus avellana, Cornus sanguinea and Euonymus
verrucosus. These species typically exist in conditions with 500–
700 mm rainfall and annual January mean temperatures not
lower than –6°C. A variant of this forest type was found in the
hills above Kremenets, where Q. petraea and Acer pseudoplatanus
were growing with many of the other species typically found in
the Quercus–Carpinus association. The second type was Quercus–
Tilia forest, with predominant species including Q. robur, Tilia
cordata, Corylus avellana, Cornus sanguinea, A. tataricum, Fraxinus
excelsior, A. campestre and Crataegus spp. Precipitation associated
with this forest type is approximately 400–500 mm with a mean
January temperature of about –4°C. A third type was riparian
forest, with Q. robur, F. excelsior, A. campestre, Ulmus glabra, Pyrus
communis, Populus spp. and Alnus glutinosa as dominant species.
The fourth and fifth forest types were dominated by Pinus
sylvestris, sometimes in nearly pure stands. On riverine sands, P.
sylvestris were found growing in scrubby to impressive stands,
depending upon water availability. Associated plants included
Betula pendula, Corylus avellana, Sambucus racemosa, Q. robur, Rosa
spp., Chamaecytisus spp. and Genista tinctoria. The other type of
pine forest was observed in Zhitomir and Rivne states, where
acid-soil indicator plants, such as Vaccinium spp., Rhamnus
frangula and Calluna vulgaris, were growing among the pines.
Tremendous diversity in Salix species was observed in or near
poorly drained areas, and extensive populations of naturalized
F. pennsylvanica, Acer negundo, Parthenocissus quinquefolia and
Robinia pseudoacacia were commonly located along roadsides and
in disturbed forests.
Two general types of steppe vegetation were seen on the
eastern excursion. One type was associated with very sandy
Table 1. Species list (taxonomy following the GRIN
database) and number of accessions collected
Species
No. accessions
collected
Acer campestre
Acer negundo
Acer platanoides
Acer pseudoplatanus
Acer tataricum
Acer tegmentosum
Betula pendula
Carpinus betulus
Chamaecytisus sp.
Cornus mas
Cornus sanguinea
Cotinus coggygria
Crataegus meyeri
Crataegus rhipidophylla
Crataegus sanguinea
Crataegus x kyrtostyla
Crataegus sp.
Daphne mezereum
Dianthus campestris
Dianthus carthusianorum
Euonymus europaeus
Euonymus verrucosus
Fragaria vesca
Fraxinus excelsior
Genista tinctoria
Juniperus communis
Juniperus sabina
Laburnum anagyroides
Ligustrum vulgare
Mentha longifolia
Pinus sylvestris
Quercus robur
Rosa canina
Rosa sp.
Rubus caesius
Sambucus ebulus
Sambucus nigra
Sambucus racemosa
Sorbus aucuparia
Sorbus torminalis
Staphylea pinnata
Tanacetum parthenium
Tanacetum vulgare
Tilia cordata
Tilia tomentosa
Viburnum opulus
3
1
4
4
5
1
3
4
1
1
2
1
1
1
1
1
1
1
1
1
4
3
1
5
2
1
1
1
3
1
2
4
2
1
1
2
3
1
2
2
1
1
1
4
1
1
soils, often adjoining P. sylvestris stands. The other was found on
more typical chernozem soils, usually in very small remnants or
on steep, eroded slopes. Many interesting wildflowers, including
Dianthus, Limonium, Salvia, Campanula, Lavatera and Thymus,
could be found on the steppe remnants, but these were not the
focus of the trip.
The vegetation around NAUU and Kyiv consisted of plantations of P. sylvestris edged with A. tataricum, Cotinus coggygria and
other shrubs along the highways at the outskirts of the city.
Boulevards and city streets were shaded by Aesculus
hippocastanum, T. cordata and Betula pendula. Genetic diversity in
Aesculus populations was obvious through their susceptibility to
leaf blotch; damage varied from tree to tree throughout most of
the country. Quercus–Carpinus forests surrounded the cultivated
landscapes of NAUU. Acorn collection was not very productive
on campus or in the countryside due to damage by weevil larvae.
The Botanical Garden of NAUU was developed from the
Golosiyeve Forest nursery in 1938. The grounds include a 9.5-ha
arboretum and a 15-ha dendropark. The arboreal collections
include 541 species, 60 forms and 22 hybrids. Samples were
collected from several tree and shrub species from within these
gardens. Directly south of the Botanical Garden lies small lakes
and natural riparian forests, which produced interesting collections of Sambucus, Cornus and Euonymus. Kyiv is also home to the
Central Botanical Garden of Ukraine, a 200-ha garden developed
in 1935 under the direction of the Ukrainian Academy of Sciences. There, research has been conducted on three native species
of Daphne, the rare and endangered D. cneorum and D. sophia and
the somewhat more common D. mezereum. The decline of D.
cneorum and D. sophia are related to the disruption and development of forested lands (Melnik 1996). Timing did not allow for
seed collection; however, these species were observed and photographed in the rare plant collection. An accession of D. mezereum
was collected earlier in the season from plants cultivated at
NAUU.
Traveling east towards Pryluky, we observed large masses of
Chamaecytisus growing along the roadsides; however, their seeds
were already dispersed. Pryluky has approximately 22 000 ha of
forest in the forest–steppe transition zone. Samples of A.
platanoides, T. cordata and Crataegus spp. were collected along the
edge of a dense-canopied Quercus–Carpinus forest. In the Romny
forests, populations of Quercus, Fraxinus, Acer and Corylus were
prevalent, but there was little or no seed production due to late
spring frosts. Seed production increased in the region southwest
of Kharkiv, with collections made of Acer spp. and Fraxinus
excelsior. An expansive meadow with a combination of grasses,
Genista, Dianthus, Tanacetum and other herbaceous plants was
found as we travelled toward Kremenchuk. We suspected this to
be a typical example of steppe vegetation.
Laburnum anagyroides was observed in three gardens in
Ukraine. In two gardens, the plants had obviously suffered significant winter damage and produced no seeds. But at Ustimovka
Dendrological Park, northwest of Kremenchuk, very large shrubs/
small trees with heavy seed production and no obvious winter
injury were observed. A large seed collection was made from
these plants in the hope that they have increased minimum
temperature tolerance.
Sofiyivka Park in Uman is a state preserve under the authority of the Ukrainian Academy of Sciences. It serves as a scientific
research park with four units of operations: science, administration, reserve, and exhibitions. Presently, the park covers 157.6 ha,
houses over 2000 species and is an important centre for plant
introduction in Ukraine. Seeds of Fraxinus excelsior were abundant, Carpinus betulus and other trees towered overhead, and we
were able to collect seeds of Sorbus torminalis, an extremely attractive, but uncommon tree native to Ukraine.
The western excursion passed through expansive forests.
F. excelsior was the dominant species on moist sites, with occasional populations of Quercus, Carpinus, Alnus and Ulmus. Near
Kremenets, collection occurred on rougher terrain that surrounds
the historic city. Carpinus, Fraxinus, A. pseudoplatanus, Betula and
16
Tilia covered the hillsides. The route to Vinnitsa passed through a
few natural forests with large Tilia specimens. Vinnitsa was
locally considered to have the best growing conditions in Ukraine.
Rich soils and abundant moisture contribute to excellent growth
and development of the forest. Time did not allow for extensive
collecting in Vinnitsa oblast, though attractive plants, such as
Sorbus torminalis, are reportedly native in its forests.
We also observed opportunities for future germplasm exploration for plants other than landscape ornamentals. Collections
from steppe remnants may yield useful forage and rangeland
germplasm, and fruit and nut germplasm in the genera Prunus,
Pyrus, Corylus, and Juglans was often abundant and may provide
useful sources of genes for adaptation to extreme environments.
Seed samples were shared with the NAUU and have now
been accessioned into the NPGS through the North Central Regional Plant Introduction Station, Ames, Iowa. These have also
been shared with appropriate NPGS sites for maintenance. Herbarium vouchers were divided between Ukrainian and American
institutions, with most deposited at the National Arboretum,
Washington DC, and the National Agricultural University of
Ukraine. Many of the seed collections will be propagated for longterm evaluation in the NC-7 Regional Ornamental Plant Trials. It
is expected that many of them will possess superior genetic
adaptation to climatic and edaphic stresses in comparison to
germplasm of these same taxa from western and other central
European provenances.
The sharing of landscape plant performance data from sites
in the north central United States that experience climatic patterns and soil types resembling those in Ukraine and of theoretical
and empirical models that relate climatic data to plant adaptation should benefit horticulturists and foresters in both Ukraine
and the United States. We are confident that our experiences
from this exploration serve as a first step in the development of
more extensive collaboration and germplasm exchange.
Acknowledgements
We greatly appreciate the financial support and cooperation of
the USDA-ARS National Plant Germplasm System, and especially of the Plant Exchange Office, in making this trip possible. In
addition, there were many individuals both in the United States
and Ukraine who gave generously of their time and expertise.
They include Dr David Topel, Dr David Acker, Dr Victor Udin,
Ms Lori Wilson-Voss, Dr Tatyana Shulkina, Dr Victor Ryabchoun,
Dr Victor Melnik, Dr Victor Kalensky, Mr Larry Panasuk, Mr
Dmitri Prikhodko, Ms Lois Simms, Mr V.M. Brezhniev, Mr Amalio
Santacruz-Varela, Ms Simone Kimber, and Mr Rex Heer. Critical
reviews of this report by Dr David Acker, Dr Edward Garvey and
Dr Harold Pellett are also much appreciated.
References
Anonymous. 1960. Pochvennaya Karta SSSR. Glavnoe Pravlenie
Geodezii i Kartografii MVD SSSR, Moscow.
Becker, H. 2000. Locate outstanding woody ornamentals—online.
Agricultural Research 48(9): 18-19.
Ganssen, R., and F. Hädrich. 1965. Atlas zur Bodenkunde.
Bibliographisches Institut AG, Mannheim.
Melnik, V. 1996. Distribution and plant communities of Daphne
cneorum and Daphne sophia in Ukraine. Thaiszia 6: 49-66.
Sheljag-Sosonko, Y.V., V.V. Osichnjur, and T.A. Andrienko. 1982.
Geography of vegetation cover in the Ukraine. (in Russian).
Science Publ., Kyiv.
Slabkovich, G.I. (ed.) 1968. Klimaticheskii Atlas Ukrainskoi SSR.
Gidrometeorogicheskoe Izdatel’stvo, Leningrad.
Sokolov, S.Ja., O.A. Svjazeva, and V.A. Kubli. 1977-1986. Arealy
derev’ev i kustarnikov SSSR. 3 vols. Izdatel’stvo Nauka,
Leningrad.
Widrlechner, M.P. 1990. NC-7 regional ornamental trials: Evaluation of new woody plants. Proc. Metropolitan Tree Improvement Alliance 7: 41-47. (Updated version available at http:/
/www.ars-grin.gov/ars/MidWest/Ames/trial.html)
Widrlechner, M.P. 1994a. Environmental analogs in the search for
stress-tolerant landscape plants. J. Arboric. 20: 114-119.
Widrlechner, M.P. 1994b. Is eastern Europe a useful source of
new landscape plants for the midwest? Comb. Proc. International Plant Propagators’ Soc. 42: 451-455.
Widrlechner, M.P., E.R. Hasselkus, D.E. Herman, J.K. Iles, J.C.
Pair, E.T. Paparozzi, R.E. Schutzki, and D.K. Wildung. 1992.
Performance of landscape plants from Yugoslavia in the
north central United States. J. Environ. Hortic. 10: 192-198.
Woody Landscape Plant Crop Germplasm Committee. 1996.
Report of the Woody Landscape Crop Germplasm Committee, 4 June 1996. Status report for the US National Plant
Germplasm System. 18 pp.
Plant
Plant
Genetic
Genetic
Resources
Resources
Newsletter,
Newsletter,
2001,
2001,
No. No.
126:126
17 - 17
20
ARTICLE
Colecta de germoplasma en la ecoregión de la
Península de Paria, Estado Sucre, Venezuela
Elena Mazzani y Víctor Segovia
Centro Nacional de Investigaciones Agropecuarias, Fondo Nacional de Investigaciones Agropecuarias,
Apdo. Postal 4653, Maracay 2101. Venezuela. E-mail: [email protected], [email protected]
Resumen
Résumé
Summary
Colecta de germoplasma en la
ecoregión de la Península de
Paria, Estado Sucre, Venezuela
Collecte de matériel génétique
dans l’éco-région de Paria
Peninsuala, Estado Sucre,
Venezuela
Germplasm collection in the
ecoregion of Paria Peninsula,
Estado Sucre, Venezuela
En comunidades de pequeños y medianos
agricultores de la Península de Paria, Estado Sucre, Venezuela, se recolectó material cultivado de diversas especies alimenticias y de interés actual y potencial para la
agricultura. La colecta fue realizada visitando pequeños y medianos agricultores
y patios caseros, y permitió reunir un
total de 141 muestras, correspondientes
a 19 especies y 21 localidades. El mayor
número de muestras recolectadas correspondió a maíz (Zea mays) (54); especies
de raíces como yuca (Manihot esculenta)
(19), ocumo o taro blanco (Xanthosoma
sagittifolium) (10) y ocumo chino (Colocasia esculenta) (4); ají dulce (Capsicum annuum) (20); auyama (Cucurbita moschata) (9)
y onoto o achiote (Bixa orellana) (8). Las
especias o condimentos recolectados fueron una muestra de jengibre morado
(Zingiber officinale), una de cúrcuma (Curcuma sp.) y una de pimienta (Piper nigrum
L.). Los frutales recolectados fueron tres
muestras de piña (Ananas comosus), dos
de lechosa, papaya o fruta bomba (Carica
papaya), una de jobo (Spondias cytherea),
una de cambur manzano (Musa AAB) y
dos de parchita maracuyá o fruto de la
pasión (Passiflora edulis var. flavicarpa). Las
unidades de producción visitadas fueron
en su mayoría “conucos” o propiedades
de pequeños agricultores que siembran
reducidas extensiones de cultivos asociados (maíz, yuca, auyama, ají y musáseas)
en terrenos distantes del lugar de vivienda.
Une mission envoyée dans des communautés de petits exploitants agricoles de
la Paria Peninsula, dans l’État de Sucre, au
Venezuela, a recueilli des espèces cultivées et non cultivées pouvant présenter une utilité agronomique. La mission,
qui s’est rendue dans des exploitations et
jardins privés de petite et moyenne taille,
dans 21 localités, a rapporté 141 échantillons appartenant à 19 espèces. Les échantillons les plus nombreux correspondaient au maïs (Zea mays) (54), suivi par
des espèces de racines comestibles telles
que le manioc (Manihot esculenta) (19), le
taro blanc (Xanthosoma sagittifolium) (10)
la colocase (Colocasia esculenta) (4), le
poivron doux (Capsicum annuum) (20), la
courge musquée (Cucurbita moschata) (9),
et le rocou (Bixa orellana) (8). Les épices
recueillies consistaient en un échantillon
de gingembre (Zingiber officinale), un de
curcuma (Curcuma sp.) et un de poivre
noir (Piper nigrum L.). Les fruits se composaient de trois échantillons d’ananas,
de deux de papaye (Carica papaya), d’un
de pomme de Cythère (Spondias
cytherea), d’un de banane (Musa AAB) et
de deux de fruit de la passion jaune (Passiflora edulis var. flavicarpa). La plupart des
exploitations visitées étaient des « conucos », dans lesquelles de petits agriculteurs plantent des cultures associées, y
compris le maïs, le manioc, la courge, le
poivron et la banane sur des surfaces peu
étendues.
A mission in communities of smallholder farmers in the Paria Peninsula, Sucre
State, Venezuela, collected cultivated
and other species of potential agronomic importance. The visit to small and
medium-sized local farms and home
gardens in 21 localities yielded a total of
141 samples belonging to 19 species.
The largest number of samples corresponded to maize (Zea mays) (54), species of edible roots as cassava (Manihot
esculenta) (19), white ocumo (Xanthosoma sagittifolium) (10) and Chinese taro
(Colocasia esculenta) (4), chilli peppers
(Capsicum annuum) (20), pumpkin (Cucurbita moschata) (9), and annatto (Bixa
orellana) (8). Spices collected were one
sample of purple ginger (Zingiber officinale), one of curcuma (Curcuma sp.) and
one of black pepper (Piper nigrum L.).
Fruits collected included three samples
of pineapple, two of carica or papaya
(Carica papaya), one of otaheite apple
(Spondias cytherea), one of banana (Musa
AAB) and two of ‘maracuya’ passion
fruit (Passiflora edulis var. flavicarpa).
Most of the visited farms were so-called
‘conucos’ where small-scale farmers
plant limited areas of associated crops
including maize, cassava, pumpkin,
sweet pepper and banana.
Key words: Collecting, home gardens,
Paria
Peninsula,
Sucre
State,
smallholder farmers, Venezuela
Introducción
“Tierra de Gracia” y “Paraíso Terrenal” fue como denominó el
Almirante Cristóbal Colón a las hermosas tierras que descubriera
el 1º de agosto de 1498 en su tercer viaje, a su llegada a la
Península de Paria, siendo ésta la primera vez que tocara tierras
continentales del nuevo mundo. Tal descubrimiento se produjo
por la parte sur-oriental de la Península y el primer punto donde
arribó fue el pueblo de Macuro (Vila et al. 1963).
Las etnias de la región practicaban la agricultura. Cultivaban
en sus conucos, maíz (Zea mays), yuca (Manihot esculenta) y otras
plantas alimenticias (Vila 1965). Durante la época de la Colonia,
se cultivaba en el Estado Sucre maíz, yuca, batatas (Ipomoea
batata), auyamas (Cucurbita spp.) y cocotero (Cocos nucifera)
(Humboldt 1799). Luego, durante el siglo XIX, se cultivaba
cacao (Theobroma cacao), cocotero, café (Coffea arabica), caña de
azúcar (Saccharum officinarum), maíz, plátano (Musa AAB), yuca,
arroz (Oryza sativa) y frijol (Vigna unguiculata) (Vila 1965). Los
productos de valor comercial en las exportaciones de la región,
según Codazzi (1841), eran el cacao, el café y el cocotero, para los
cuales “cada hacienda, cada valle tiene un punto para embarcar
sus frutos; y la inmediación de la Isla de Trinidad ofrece un
mercado ventajoso para ellos.”
Texera (1991) reseña las expediciones botánicas realizadas en
Venezuela desde 1754 hasta 1950. En ese período fueron ejecutadas
101 exploraciones botánicas en el territorio nacional. De éstas,
señala siete que tuvieron que ver con el Estado Sucre. Ellas fueron
las de Lofling en 1756, von Jacquin en 1760, Humboldt y Bonpland
de 1799 a 1800, Otto de 1840 a 1841, Grossourdy en 1867, Pittier
de 1913 a 1950, y Netting de 1929 a 1930.
18
Aun hoy en día, esta región recibe una gran influencia de la
Isla de Trinidad por su cercanía a las costas de Paria. Esta
influencia se manifiesta en alimentos consumidos y condimentos,
entre otros. Sin embargo, en su gran mayoría los habitantes de la
zona mantienen materiales autóctonos de Venezuela.
Esta región ha sido cacaotera por excelencia, siéndolo aún. A
diferencia de las demás regiones cacaoteras, en Oriente las haciendas estaban alejadas de los centros poblados y por ello se da el
caso especial del desarrollo urbano de Yaguaraparo e Irapa,
testimonio de la importancia cacaotera regional (Vila 1965), que
aunque en menor cuantía aún persiste, sembrándose, para 1997,
23.735 has (Venezuela 1997).
La región visitada durante la colecta comprende
principalmente los valles de la Península de Paria, los cuales
están sembrados con plantaciones de cacao, cocotero, frutales
como mango (Manguifera indica) y aguacate (Persea americana), así
como conucos donde se planta maíz, yuca, ocumo (Xanthosoma
sagittifolium) y algunas musáseas. Los ríos de estos valles nacen
en el sistema montañoso de la Península de Paria. Según Luque y
Mazzi (1977), sus suelos son de texturas medias y arcillosas,
moteados, a profundidades variables, con alta saturación de
bases. Según COPLANARH (1974), la mayoría de estos suelos
posee mal drenaje ya sea por inundaciones localizadas o por
elevadas mesas de agua. En el norte de la Península existe un
área ocupada por bosque húmedo tropical, con cultivo de café,
que es una zona ecológica diferente a la región de valles, ubicada
al sur de la Península.
El presente trabajo tiene como objetivo presentar los
resultados de la colecta efectuada en comunidades de pequeños
y medianos agricultores de la Península de Paria con el fin de
obtener material cultivado de diversas especies alimenticias y de
interés actual y potencial para la agricultura.
Descripción de las zonas de colecta
bajo pastizales naturales en algunas áreas, con limitaciones de
relieve, erosión de las tierras y suelos pobres. Esta región abarca
toda el área norte de la Península de Paria, y es parte del sistema
de la cordillera de la Costa. El macizo montañoso es relativamente
bajo, oscilando sus alturas entre 500 y 1000 metros. Pertenece a
las unidades fisiográficas “montaña baja premontano húmedo”,
con clima de bosque húmedo premontano, temperatura media
anual de 24,2 ºC y precipitación media de 1920 mm; y ”bosque
tropical semi-deciduo” con dosel bajo (Ewel y Madriz 1968).
El paisaje de piedemonte presenta una agricultura de
subsistencia y de plantaciones, con algunas limitantes de
topografía y suelos de baja fertilidad. Comprende la parte sur de
la Península de Paria, cuya costa da hacia el golfo del mismo
nombre. Esta región pertenece a la zona de vida “bosque seco
tropical” (Ewel y Madriz 1968).
El piedemonte ondulado tropical seco abarca las
inmediaciones de las localidades de Bohordal, Municipio Cajigal
y Parroquia Yaguaraparo. Presenta un relieve plano con
pendientes de 2 a 4%, de permeabilidad media a alta. El clima es
del tipo “bosque seco tropical” (Ewel y Madriz 1968), la
precipitación media anual es de alrededor de 1100 mm y la
temperatura media anual de 27,7ºC. Posee tierras agropecuarias,
existiendo “espinar tropical ralo” con moderada intervención en
la formación vegetal. Los suelos son de baja fertilidad y
frecuentemente erosionados por efecto del escurrimiento. Existen
plantaciones y agricultura de subsistencia.
Los valles se ubican en el piedemonte. Los ubicados en el
extremo sur de la Península de Paria pertenecen a la unidad
fisiográfica denominada “valle bajo tropical seco”, con
pendientes entre 1 y 2%. En los mismos existen plantaciones
asociadas a cultivos de subsistencia, ganadería semi-intensiva y
cultivos anuales mecanizados. Las limitaciones para la
agricultura comprenden un alto riesgo de inundaciones cortas
generalizadas y una alta pedregosidad del suelo. Corresponde a
una zona de vida de “bosque seco tropical” (Ewel y Madriz 1968)
cuya temperatura media anual es de 26,6ºC y precipitación
media anual de 1300 mm. Predomina la formación boscosa de
tipo tropical decidua y semi-decidua, con alturas y densidades
de medias a altas, fuerte intervención de plantaciones y moderada
intervención de la agricultura de subsistencia.
El Estado Sucre se encuentra ubicado en el extremo nororiental de
Venezuela, entre las coordenadas 10º02’34”, 10º45’25” de latitud
Norte y los 61º51’17” y 64º31’42” de longitud Oeste. Su economía
se basa en la agricultura, pesca, artesanía y turismo.
La Península de Paria, región visitada durante 1995, está
ubicada en la zona oriental del Estado Sucre, aproximadamente
entre las latitudes 10º35' hasta 10º42' Norte y 62º10'
hasta 63º00' de longitud Oeste. Limita al Norte con
el mar Caribe y al Sur con el Golfo de Paria. Las
muestras fueron recolectadas entre 10 y 215
m.s.n.m.
Durante la colecta fueron visitados los
municipios Arismendi, Mariño, Valdés y Cagigal.
El Estado Sucre está conformado
fisiogeográficamente por cuatro paisajes naturales:
montaña, piedemonte, planicie y valles (Marín
1993). Estos paisajes determinan la variabilidad
climática, diversidad de formaciones vegetales y la
heterogeneidad litológica. Las regiones visitadas en
la Península de Paria y parte del sur-este del Estado
comprenden estos cuatro paisajes naturales.
En la región montañosa se practica la agricultura Figura 1. Mapa del Estado Sucre, Venezuela mostrando las rutas de
de subsistencia, cultivos permanentes y ganadería colecta en la Península de Paria.
El paisaje de planicie es de “planicie de desbordamiento
tropical húmeda”. Ubicadas al noreste de Irapa, estas planicies
presentan una pendiente de 1% con bioclima predominante de
bosque húmedo tropical (Ewel y Madriz 1968), precipitaciones
de alrededor de 1500 mm anuales y temperatura media anual de
27,7ºC; los suelos son de baja fertilidad. Allí se observa
predominio de áreas sin uso, y se practican cultivos anuales
mecanizados y ganadería semi-intensiva y extensiva.
Al sur de la Península de Paria, la carretera hacia Irapa
atraviesa una sabana donde se observan, entre otras especies, la
curata (Curatella americana) y el chaparro manteco (Birssomina
crasssifoliaa). Estas sabanas se encuentran al pie de colinas de
aproximadamente 100 m de altitud, presentando un suelo pobre
y pedregoso (Vila 1965).
Todo el litoral norte de la Península de Paria está cubierto de
vegetación xerófita (Vila 1965).
caratos o papillas, atoles, cachapas, harinas. El maíz se utiliza también
para la alimentación animal. En todos los sitios donde se obtuvo
muestras (21), se recolectó por lo menos una muestra de esta especie.
En cuanto a las raíces recolectadas, se encontró ocumo blanco
y ocumo chino sembrados en laderas de montañas, en la vía que
conduce a San Juan de las Galdonas. Estas especies son
sembradas con fines comerciales y para consumo.
Por otra parte, todas las muestras de yuca correspondieron al
tipo dulce que en su mayor parte se siembra en pequeñas extensiones
o en patios caseros, para venta en los mercados locales y para
consumo propio. Determinados productores poseen siembras
comerciales de mayor extensión. Algunos de los materiales habían
sido traídos originalmente del sur del país (Estado Bolívar).
La segunda especie en importancia—en función del número
de muestras recolectadas (20) y sitios de recolección (13)—fue el
Resultados
En la Península de Paria se visitó un total de 21 localidades,
ubicadas en los paisajes descritos anteriormente. En la Figura 1 se
presenta el mapa del Estado Sucre mostrando la ruta de colecta.
La colecta fue realizada visitando pequeños y medianos
agricultores y patios caseros donde se recolectó un total de 141
muestras correspondientes a 19 especies. El número de muestras
recolectadas de cada especie, el número de localidades de colecta
y los usos de los materiales se presentan en el Cuadro 1.
El mayor número de muestras recolectadas correspondió a
maíz (54) y especies de raíces (19 muestras de yuca, 10 de ocumo
blanco y 4 de ocumo chino).
El maíz recolectado pertenece mayormente a los tipos amarillo
caribeño, blanco de Sucre, canilla y cariaco. En la región existen
agricultores que siembran híbridos y variedades comerciales, así como
también pequeños agricultores que mantienen sus variedades. A
partir de los maíces recolectados en este trabajo, elaboran arepas,
Figura 2. Dos muestras de ají dulce recolectadas en la
Península de Paria, Estado Sucre, Venezuela.
Cuadro 1. Información general de las muestras recolectadas en la Península de Paria, Estado Sucre, Venezuela
Especies
Nombre local
Usos
N°
muestras
N° sitios
recolección
Zea mays
Maíz
54
21
Capsicum annuum
Manihot esculenta
Xanthosoma sagittifolium
Cucurbita spp.
Bixa orellana
Colocasia esculenta
Theobroma cacao
Ananas comosus
Passiflora edulis
Carica papaya
Musa AAB
Spondias cytherea
Phaseolus vulgaris
Ricinus comunis
Curcuma sp.
Piper nigrum
Zingiber officinale
Hibiscus esculentum
Ají Dulce
Yuca
Ocumo Blanco
Auyama
Achiote
Ocumo Chino
Cacao
Piña
Parchita
Lechosa
Cambur manzano
Jobo
Caraota negra
Higuerilla
Cúrcuma
Pimienta
Jengibre
Chimbombó
Atoles, arepas, papillas,
harinas, cachapas,
alimentación animal
Condimento
Consumo fresco
Consumo fresco
Sopas
Colorante para alimento
Consumo fresco
–
Consumo fresco
Consumo fresco
Consumo fresco
Consumo fresco
Consumo fresco
Consumo fresco
Medicinal
Condimento
Condimento
Condimento
Sopas
20
19
10
9
8
4
4
3
2
2
1
1
1
1
1
1
1
1
13
9
7
8
5
4
2
2
1
2
1
1
1
1
1
1
1
1
20
ají dulce (Capsicum annuum). Esta especie es normalmente
sembrada en los patios caseros para consumo propio, existiendo
variabilidad entre muestras. El ají dulce es un condimento muy
utilizado en la región, sobre todo en la elaboración de platos
típicos a base de pescados. En la Figura 2 se muestran dos
ejemplares recolectados en la región.
Otra especie de importancia es la auyama (Cucurbita
moschata), recolectada en patios caseros. Es utilizada para
consumo propio, vendiéndose en algunos casos el excedente en
los mercados locales. Se encontró una alta variabilidad.
También fueron recolectadas ocho muestras de onoto o
achiote (Bixa orellana), las cuales fueron encontradas mayormente
en patio caseros. Es utilizado para consumo propio, como
colorante de alimentos, destinándose el excedente a la venta en
mercados locales.
Las otras especies recolectadas fueron especias foráneas y
frutales de la región.
Las especias o condimentos recolectados fueron una muestra
de jengibre morado (Zingiber officinale), una muestra de cúrcuma
(Curcuma sp.) y una muestra de pimienta (Piper nigrum L.). Estas
muestras corresponden a materiales originalmente procedentes
de Trinidad, por su cercanía a la región visitada. Las especies
recolectadas en patios caseros son utilizadas para consumo
familiar y como ornamentales.
Los frutales recolectados fueron tres muestras de piña; dos
de lechosa o papaya llamada comúnmente lechosa pajarito
(Carica papaya) encontradas a la orilla de camino; una muestra de
jobo (Spondias cytherea), ejemplar silvestre también encontrado a
orilla de camino; una muestra de cambur manzano (Musa AAB)
y dos muestras de parchita maracuyá o fruta de la pasión
(Passiflora edulis var. flavicarpa). En la Figura 3 se presenta un
ejemplar de piña recolectado en patio casero.
Las muestras recolectadas fueron incorporadas a los respectivos
bancos de germoplasma del Centro Nacional de Investigaciones
Agropecuarias (CENIAP) del Instituto Nacional de Investigaciones
Agrícolas (INIA), Maracay, para su conservación.
Las unidades de producción visitadas fueron en su mayoría
conucos, donde los pequeños agricultores siembran, en terrenos
Figura 3. Plantas de piña en patio casero en una localidad
de la Península de Paria, Estado Sucre, Venezuela.
distanciados del lugar de vivienda y en asociación, pequeñas
extensiones de cultivos como maíz, yuca, auyama, ají y
Musáseas.
Los materiales recolectados en patios caseros fueron frutales
como parchita y piña, onoto y en algunos casos ají. Las muestras
de pimienta, cúrcuma y jengibre fueron encontradas en un patio
casero en una sola localidad.
Por otra parte, las muestras encontradas a orilla de camino,
silvestres o escapadas, fueron las dos de lechosa, la de jobo y la
de tártago. Sin embargo, éstas se encontraban cerca de zonas
pobladas.
La información recabada constó de 34 datos de colecta
recomendados por el IPGRI, y que constan de los registros de lugar,
muestra, etc., normalmente tomados en colectas de este tipo.
Los habitantes de la región guardan y multiplican su
germoplasma, y realizan intercambio con habitantes de la misma
región. Denominan a los materiales de igual manera, que en el
resto del país, en la mayoría de los casos, a excepción de los
materiales exóticos poco conocidos en otras regiones y el achiote,
que en el resto del país es conocido como onoto.
Referencias bibliográficas
Codazzi, A. 1841. Obras Escogidas: Resumen de la Geografía de
Venezuela. Caracas, Ven. 1960. Ed. Ministerio de Educación.
Dirección de Cultura y Bellas Artes. V. I :761 p.
Ewel, J. J. y A. Madriz. 1968. Zonas de Vida de Venezuela.
Memorias Explicativas sobre el Mapa Ecológico. Fondo
Nacional de Investigaciones Agropecuarias, Ed. Caracas.
Humboldt, A. de. (1799-1804). Viajes a las Regiones Equinocciales
del Nuevo Continente. Caracas, Ven. 1956. Trad. Lisandro
Alvarado. Ed. Ministerio de Educación. Dirección de Cultura
y Bellas Artes. Tomo II: 364 p.
Luque, O. y L. Mazzi. 1977. Estudio Agrológico Detallado.
Estación Experimental de Irapa. FONAIAP, CENIAP.
Boletín Técnico Nº 4. 71 p.
Marín, A. 1993. Sectorización Fisiográfica de la Sub-región
Carúpano Paria. Ministerio del Ambiente y de los Recursos
Naturales Renovables. División de Planificación y
Ordenamiento del Ambiente. Región Sucre. s.n.p.
(Mecanografiado)
Ministerio del Ambiente y los Recursos Naturales Renovables. s/
f. Atlas Práctico de Venezuela. Publicación de El Nacional y
Cartografía Nacional. Fasc. Nº 20. Sucre.
Texera A., Y. 1991. Las exploraciones botánicas en Venezuela
(1754-1950). Caracas, Fondo Editorial Acta Científica
Venezolana. 186 p.
Venezuela, s/f. Municipio Arismendi. Información Turística.
Mimeografiado. snp.
Venezuela. Comisión del Plan Nacional de Aprovechamiento de
Recursos Hidráulicos (COPLANARH). 1974. Inventario
Nacional de Tierras. Regiones Centro Oriental y Oriental.
Regiones 7 y 8. Sub regiones 7B, 7C, 8A. Caracas, 1974. 415 p.
Publicación Nº 35.
Venezuela. Ministerio de Agricultura y Cría (MAC). 1997.
Anuario Estadístico Agropecuario. Caracas.
Vila, M. A. 1965. Aspectos Geográficos del Estado Sucre.
Corporación Venezolana de Fomento. Serie “Monografías
Estadales”. Caracas, Ven. 266 p.
Vila P., F. Brito, A. L. Cárdenas y R. Carpio. 1963. Geografía de
Venezuela. No. 2 El Paisaje Natural y el Paisaje Humanizado.
Ediciones del Ministerio de Educación. 558 p.
Strauss, R. 1992. El Tiempo Prehispánico de Venezuela.
Fundación Eugenio Mendoza. Caracas, Ven. 279 p.
Plant
Plant
Genetic
Genetic
Resources
Resources
Newsletter,
Newsletter,
2001,
2001,
No.No.
126:
126
21- 21
26
ARTICLE
Plant exploration in the Talysch Mountains
of Azerbaijan and Iran
L. Frese1, Z. Akbarov2, V. I. Burenin3, M. N. Arjmand4 and V. Hajiyev5
Federal Centre for Breeding Research on Cultivated Plants (BAZ) – Gene Bank, Bundesallee 50, 38116
Braunschweig, Germany. Email: [email protected]
2 Scientific Research Institute of Agriculture, Vegetable village no. 2, Baku, Azerbaijan
3 N. I. Vavilov Institute of Plant Industry (VIR), 42-44, Bolhaya Morskaya Street, 190000 St. Petersburg, Russia
4 Sugar Beet Seed Institute (SBSI), P.O. Box 31585 – 4114 Karaj, Islamic Republic of Iran
5 Laboratory of Botany, Baku, Azerbaijan
1
Summary
Résumé
Resumen
Plant exploration in the
Talysch Mountains of
Azerbaijan and Iran
Découverte de plantes dans
les montagnes Talysch
d’Azerbaidjan et d’Iran
Exploración fitogenética de
las montañas Talysch de
Azerbaiján e Irán
Fifty-four accessions of wild species and
crops have been collected in Azerbaijan
and Iran. Particular attention was given
to the collection of wild species of Beta
section Beta and section Corollinae. Section Beta has a wide distribution area in
the Mediterranean basin and the northwest Atlantic coast, while section
Corollinae has its main distribution in Turkey. The western and southern part of
the Caspian Sea probably forms the eastern margin of the distribution area of the
wild beet species Beta lomatogona and B.
vulgaris subsp. maritima. Populations of
both species appeared to suffer from genetic erosion caused by land management changes and overgrazing of growing sites in Azerbaijan and Iran. The authors suggest that an in situ conservation
and on-farm management project
should be established in Azerbaijan and
northwest Iran to rescue the wild beet
populations still known to exist in the
area visited, and to restore genetic diversity in the areas concerned.
54 échantillons d’espèce sauvage et des
plantes cultivées ont été rassemblées en
l’Azerbaïdjan et en l’Iran. Une attention
particulière a été donnée de à la collection
d’espèce sauvage Beta section Beta et section Corollinae. La section Beta a une zone
de distribution large dans le bassin
méditerranéen et la côte atlantique du
nord-ouest tandis que la section
Corollinae a sa distribution principale en
Turquie. La partie occidentale et
méridionale de la mer caspienne forme
probablement la marge orientale de la
zone de distribution des espèces de
betterave sauvage de B. lomatogona et de
sous-espèce de B. vulgaris subsp. maritima. Les populations des deux espèces
ont semblé souffrir de l’érosion
génétique causée par voie de terre des
changements de gestion et surpâturage
des sites croissants en Azerbaïdjan et en
Iran. Les auteurs suggèrent d’établir un
projet entretien in situ et de gestion de
ferme en Azerbaïdjan et en Iran du nordouest avec l’objectif pour sauver les
populations sauvages de betterave
toujours connues pour exister dans la
zone visitée et pour restaurer la diversité
génétique dans les zones intéressées.
54 accesiones de especies salvajes y de
cosechas se han recogido en Azerbaijan e
Irán. La atención determinada fue dada a
la colección de especie salvaje de la Beta
sección Beta y de la sección Corollinae. La
sección Beta tiene un área de distribución
amplia en el lavabo mediterráneo y la
costa atlántica del noroeste mientras que
la sección Corollinae tiene su distribución
principal en Turquía. La parte occidental
y meridional del mar caspio forma
probablemente el margen del este del
área de distribución del salvaje especie de
la remolocha de la B. lomatogona y del
subespecies B. vulgaris subsp. maritima.
Las poblaciones de ambas especies
aparecían sufrir de la erosión genética
causada por tierra cambios de la gerencia
y overgrazing de sitios crecientes en
Azerbaijan e Irán. Los autores sugieren
para
establecer
una
proyecto
conservación in situ y en de la gerencia
de granja en Azerbaijan e Irán del
noroeste con el objetivo para rescatar las
poblaciones salvajes de la remolocha
todavía sabidas para existir en el área
Key words: Azerbaijan, Beta
lomatogona, Beta vulgaris, genetic
erosion, germplasm collection, Iran,
sugar beet
Introduction
The Talysch Mountains are located in the south of Azerbaijan
between latitudes 38°30’N and 39°00’N and between longitudes
48°00’E and 48°50’E. The eastern slopes face the Caspian Sea
while the southern foothills are in northwest Iran. In the 300 km
from Baku to the town of Astara on the Iranian frontier, climate
and soil conditions change considerably. South of Baku, desertlike areas with highly saline soil can be found, while fertile black
soils occur in the South. Within a distance of about 200 km, the
annual rainfall increases from 400 mm, south of Baku, to
1200 mm in the humid-subtropical provinces of south Azerbaijan
and northwest Iran at the Caspian Sea (Zohary 1973). This highly
variable environment has created a diverse flora with a high
number of endemic plant species.
The wild beet species Beta lomatogona Fischer & Meyer (Beta
section Corollinae) is an element of this flora. The species was
detected by Hohenacker (1838) in the Talysch Mountains at
Tatuni. Buttler (1977) considered B. lomatogona as a model plant
for the irano-turanian flora because the distribution limits of this
wild beet species are almost congruent with the oriental-turanian
geobotanical area. The species has its main distribution area in
Turkey. Its abundance decreases from east Turkey to northwest
Iran and Azerbaijan. A second wild beet species is believed to
occur in Azerbaijan, as indications were found in the herbarium
of the Institute of Botany in Tblisi (Frese and Burenin 1991) that
B. vulgaris subsp. maritima, similar to the germplasm identified
by M. Nasser Arjmand in Iran (Srivastava et al. 1992), can be
found in inland areas of Azerbaijan.
Germany and The Netherlands are co-operating closely in the
field of plant genetic resources conservation, and are managing
two joint germplasm collections: the Dutch–German potato collection at PRI Centre for Genetic Resources (CGN) in Wageningen,
The Netherlands and the German–Dutch Beta collection at the
22
BAZ Gene Bank in Braunschweig, Germany. The task-sharing
between both countries includes the collection of target species
such as Allium, Beta, Brassica, Lactuca and potatoes. Since 1990,
Beta collecting missions have often been organized together with
the N. I. Vavilov Institute of Plant Industry (VIR).
Two previous expeditions had been conducted in Armenia,
Georgia and Daghestan by Frese et al. (1990) and Frese and
Burenin (1991). This third mission was part of a framework
programme that was approved by the German–Dutch Board for
Plant Genetic Resources in 1993 and by the World Beta Network
(WBN) in 1996. Detailed plans were made in 1998, and official
consent was granted by the host countries in mid-1999.
Objectives
Buttler (1977) considers the Armenian highlands as the evolutionary centre of Beta section Corollinae because the distribution of
all three basic species (B. corolliflora, B. macrorhiza and B.
lomatogona overlap there. Several collectors have stressed the risk
of genetic erosion within the section Corollinae (Anonymous 1990;
Buttler 1977; Frese et al. 1990; Frese and Burenin 1991). Therefore,
the first major objective of the team was to search for and rescue
the remaining Beta populations. Though the interest of the team
was concentrated on the genus Beta, there was a common understanding that the journey would be conducted as a multicrop
collecting mission with the aim of safeguarding the genetic resources of selected plant species for use in beet, vegetable and
pharmaceutical crop improvement programmes. In the case of
the medicinal plant Hypericum perforatum, it was intended to
increase the genetic diversity within a working collection that is
being specifically established for screening on wilt resistance. For
a forthcoming Beta genome research programme, B. lomatogona
accessions from geographically distant sites are required for wide
species crosses.
The second objective of the exploration was an assessment of
the need for, and feasibility of, in situ conservation projects for
Beta species in Azerbaijan and northwest Iran. In particular,
species of section Corollinae are difficult to manage in ex situ
holdings. In addition to requiring tiresome manual preparation
of the hard-coated fruits to facilitate seed germination, all
Corollinae species are adapted to specific growing conditions.
There are, therefore, good reasons to assume that, when the
original seed sample is first multiplied far away from its natural
mountainous habitat, plants will be selected within the population for ‘genebank management adaptation’. It seems impossible
therefore to maintain the genetic integrity of Corollinae accessions
through ex situ conservation. The best way to safeguard the
genetic diversity of Corollinae species would be in situ conservation as already implemented for crops, and their related wild
species, such as Triticum, Aegilops, Vicia, Lens and Pisum in Turkey (Firat 1999).
additional information on potential growing sites of the target
species. Further information on the distribution of Beta was found
in literature (Grossheim 1945). In addition, a detailed geobotanical map of Azerbaijan (1:600 000) and maps of the target areas in
the Azerian part of the Talysch Mountains (1:100 000) were used
to plan the travel route. Photographs of different growing stages
of B. lomatogona were prepared by the BAZ Gene Bank to facilitate
discussion with farmers and shepherds familiar with the vegetation of potential growing areas.
At each collecting site a passport data sheet of the BAZ Gene
Bank was filled in. The sheet consists of a mandatory part,
almost identical to the ECP/GR multicrop passport descriptors,
and an optional part where notes on topography, soil type etc.
can be recorded. Each collected sample was identified by a
collection number of the format, country code, collection year and
a sequence number, for example ‘AZE 99 01’. Longitude and
latitude co-ordinates were determined by a handheld GPS system
(Garmin 50). The elevation of a site was measured with an
altimeter.
A survey of the samples collected is presented in Table 1, and the
collecting areas are indicated in Fig. 1. Some of the germplasm
collected in Azerbaijan was partly shared with team members,
according to their specific interests. A Material Acquisition Agreement (MAA) was signed by the Research Institute of Agriculture
(Azerbaijan) and the BAZ Gene Bank. Copies of the complete set of
passport data and a travel report were made available to all partners
Material and methods
To get an overview of material already existing in collections, the
main database of the BAZ Gene Bank and the International
Database for Beta (IDBB) were searched for collecting sites of
Beta, Brassica, Daucus, Hypericum and Lactuca in Azerbaijan and
Iran. The herbarium collection at Baku was also searched to gain
Fig. 1. Collection areas in Azerbaijan and Iran.
Table 1. Germplasm collected in Azerbaijan and Iran
Genus
Species
Allium
Allium
Anethum
Avena
Beta
Beta
Beta
Carthamus
Coriandrum
Daucus
Daucus
Daucus
Hypericum
Lactuca
Lactuca
Lathyrus
Lens
Lepidium
Linum
Petroselinum
Phaseolus
Phaseolus
Phaseolus
Raphanus
Solanum
Total
cepa
porrum
graveolens
lomatogona
vulgaris
vulgaris
tinctorius
sativum
carota
carota
carota
perforatum ?
saligna
serriola
sativus
culinaris
sativum
usitatissimum
crispum
vulgaris
vulgaris
vulgaris
sativus
tuberosum
Subspecific
name
maritima
vulgaris
carota
maximus ?
sativa
Range of elevation
by genus
420
1600
2000
20
2000
420 to 800
800 to 1680
20 to 1600
1600
260 to 2000
20 to 1680
800 to 2300
800 to 1100
420 to 1100
800
1680
var. nanus
var. vulgaris
?
var. sativus
of the collecting team. All seed samples were first left in Iran and,
after signature of a separate MAA by the Plant Genetic Resources
National Bank, samples were sent to the BAZ Gene Bank.
In the case of B. lomatogona, the International Data Base for
Beta (IDBB) documented 11 collecting sites in Iran and seven sites
in the former Soviet Union, while 177 sites are recorded for the
Turkish distribution area. In the literature only scant information
on the species’ distribution could be found. In addition, due to
translations and transcriptions between Russian and German and
vice versa, some of the described places could not be clearly
identified by the local experts in Azerbaijan.
The herbarium of the Laboratory of Botany (Baku) provided
striking additional information. Before World War II several
botanists collected 18 specimens of Beta vulgaris, most of them
probably subsp. maritima, and two specimens of B. lomatogona in
Azerbaijan, Georgia and Iran (Table 2). The botanist and expert
in the flora of the Caucasus and Transcaucasus, A. Grossheim,
investigated the flora of Azerbaijan in the 1930s. He determined
10 vouchers, as B. perennis no longer a valid taxon, and collected
B. lomatogona in the district of Lenkoran (Azerbaijan) and in the
Iranian province of Tabris. This discovery proves that wild or
weedy types of B. vulgaris occur or have occurred in Azerbaijan.
The populations probably belonged to the material detected close
to the Iranian coast of the Caspian Sea (Srivastava et al. 1992).
The field exploration started on 20 September and ended on 1
October, 1999. Between Baku and Lenkoran cultivated B. vulgaris
was found in a private garden. The population is not deliberately
cultivated in gardens, but survives as a kind of tolerated weed,
which is allowed to bolt and produce seed. At Shorsulu, in the
district of Salyany, a few plants of B. vulgaris subsp. maritima
420
420
1100
1680
1800
Number
1
1
1
1
1
1
6
2
4
3
1
1
6
4
4
2
4
1
1
1
2
1
2
1
2
54
were found in an alfalfa field. This is proof that the subspecies
still occurs in Azerbaijan. The west and south coasts of the
Caspian Sea appear to be the northeastern limit of the distribution area of the ‘seabeet’, B. vulgaris subsp. maritima. The seabeet
population at Shorsulu seemed to be heavily damaged by grazing of the alfalfa crop. The field was, until recently, used to grow
cotton but was ploughed the previous autumn and cropped with
alfalfa. Farmers explained that about eight plants per square
metre had been growing amongst the alfalfa, and that many
neighbours had come in spring to harvest leaves of the wild beet
for salads. After the alfalfa was harvested and grazed, only a few
short plants with mature seed remained. In an adjacent cotton
field, two young plants and a small plant with mature seeds were
found. It is doubtful if this wild beet population can survive at
the site, even though, its high soil salinity and the ruderal character of the field margins are well suited to B. vulgaris subsp.
maritima. The danger is that the use of the field may change again
as it did in 1998/1999.
On 22 September, a quince cultivation area was passed on the
way from Lenkoran to Lerik. Many different forms of quince exist
in this area, amongst them a type with soft flesh, which can be
eaten fresh like an apple.
In the eastern foothills of the Talysch Mountains, along the
Lenkoran Chay River, rice is produced. The cultivation of rice was
halted by the planned economy in 1930, and until 1990 traditional varieties were only used and maintained privately. After
the independence of Azerbaijan, the cultivation of the favourite
local variety ‘Ambarbo’ was increased again.
The search for B. lomatogona started in the Lerik province the
same day. At Tatuni, a growing site where B. lomatogona had
Leg. et Det. A.
Grossheim
Leg. et Det. P
Gurijskij
Leg. A.
Kolakovsky
Leg. L. Prilipko
Det. A. Grossheim
Leg. M. Kotov
Det. M. Kotov
Leg. M. Kotov
Det. M. Kotov
Det. A. Grossheim
Leg. A. Kolakovsky
Det. A. Grossheim
Leg. L. Prilipko
Det. A. Grossheim
(2 specimens)
Leg. M. Sachokla
Det. A. Grossheim
Leg. et Det. C.
Gurvitsh
Leg. et Det. C.
Gurvitsh
Leg. et Det. A.
Kolakovsky
Leg. J. Doroshko
et T. Heideman
Det. A. Grossheim
Det. A. Grossheim
Leg. L. Prilipko
Det. A. Grossheim
Leg. A. Kolakovsky
Det. A. Grossheim
Leg. A. Kolakovsky
Det. A. Kolakovsky
Leg. et Det. L.
Prilipko/A.
Grossheim
Leg. et Det. A.
Grossheim
28.09.1925
30.01.1926
11.11.1934
29.05.1928
08.05.1928
20.06.1924
17.07.1930
27.06.1930
23.12.1929
09.03.1934
May 1930
27.06.1930
09.05.1938
02.05.1938
31.05.1929
11.05.1928
05.06.1934
04.11.1934
16.6.1929
18.11.1929
Collected and
determined by
Collection
date
(L.)
(L.)
(L.)
(L.)
(L.)
(L.)
(L.)
(L.)
(L.)
Beta lomatogona
F. et M.
Beta perennis (L.)
Halaczy
Beta perennis (L.)
Halaczy
Beta perennis (L.)
Halaczy
Beta perennis (L.)
Halaczy
Beta lomatogona
F. et M.
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis
Halaczy
Beta perennis (L.)
Halaczy
Beta perennis (L.)
Halaczy
Beta vulgaris (L.)
var. maritima Boiss.
Beta perennis (L.)
Halaczy
Beta vulgaris (L.)
var. maritima
Classified as
Beta lomatogona F. et M.
Beta lomatogona F. et M.
Beta vulgaris (L.) subsp. maritima Arcang.
Beta vulgaris (L.) subsp. maritima Arcang
Large bracts in the upper part indicate subsp.
adanensis. Would be rather unusual.
Beta vulgaris (L.) subsp. maritima Cylindrical
25cm long thickened root is very well conserved.
Beta vulgaris (L.) subsp. maritima
?, Specimen was collected very late in the
season. Only few seed balls existing.
Beta vulgaris (L.) subsp. ? Collected
very late in the season. Bleached
stems and a few seed balls left.
Beta vulgaris (L.) subsp. maritima
Suggested classification
and remarks
Table 2. The herbarium collection of Beta species of the Laboratory of Botany (Baku)
Persia borealis, province Tabris, Atropatania, in Jugo
Meshau-Dagh, propre, st .Viae, ferr, Jam, 1960–2500 masl
Azerbaijan, province Baku, District Lenkoran, vr. Ruzjmay v p. Geledera v Pirazona ?
Azerbaijan, district Soljary, ad meridium m. tis Kjuram
dagh sorchus Kara-tschola, ad fossos
Azerbaijan, province Gandzha, district Agdam, inter p.
Steppa Karabach, inter pp. Chinziristan et Bardy
Georgia, province Tiflis, steppa Karajazy, inter pp.
Michalijovca et Tatjanovika
Azerbaijan, Steppa Shirvan, s. Chaladzh Kureams
Azerbaijan, Steppa Shirvan, inter Sorchuz Karatschola et la Choladzh
Azerbaijan, Steppa Shirvan, inter Sorchuz Karatschola et la Choladzh
Azerbaijan, district Baku, insula Los, in maritimis
Azerbaijan, district Baku, insula Bulla, in argillosis
Azerbaijan, district Nucha, in Steppa Adzhinaur
Azerbaijan, Steppa Shirvan, district Saljany, prope opp.
Saljany, circa Kara-Tshala, inter sagetes Gossypii
Azerbaijan, district Saljany, pr. Sorchos, Kara-Tshala,
inter sagetes
Azerbaijan, province Gandzha, district Kazach, inter p.
Karasachkal et steppam Djeiran-tshely
Azerbaijan, province Gandzha, district Agdam, inter p.
Steppa Karabach, inter st. Viae ferr. Jevlach et Borsunly
Azerbaijan, Steppa Shirvan, district Saljany, prope pag
Chaladzh, inter sagetes Gossypii
Azerbaijan, Steppa Shirvan, inter Tumin, dominium.
Kara-tschola et st.v.f. Pirsagat
Azerbaijan, Baku, district Saljany
Azerbaijan, Gandza province, inter Ganja
Azerbaijan, province Baku, district Saljany, dominium
Kara-Tshala in steppa Shirvan
Collection site and
notes on the voucher
24
previously been found (Grossheim 1945), farmers and shepherds
recognized the species from photos, and showed three plants
growing close to a garden. In the graveyard of the small village of
Geledera at 2000 masl the B. lomatogona population, described by
Prilipko and Grossheim in 1930 (Table 2), was found to be still
very large. The area is not as intensively grazed as sites visited
earlier. A farmer reported that this species used to be found at
several sites in the area. Sheep, goats and cattle prefer B.
lomatogona as forage just at the time of flowering, with the result
that a decreasing number of these perennial plants can produce
mature seeds. There are more farm animals in the area than
before because younger, jobless family members have returned
from the towns to the villages to start families. They earn their
living mainly by the production of domesticated animals on the
pastureland, which in fact is common property. Hence, the growing human population is causing genetic erosion in B. lomatogona
in the Talysch Mountains of Azerbaijan.
The report of the farmer living at Geledera agrees with the
account of a shepherd claiming to be 100 years old, who said that
almost all of the birds which used to be abundant in the area have
now disappeared and that the landscape has become silent.
At the village of Pirsara, another site of B. lomatogona known
from literature, the farmers did not fully recognize the plant, but
showed us species with leaf shapes and roots similar to B.
lomatogona, i.e. Rumex and others. Later, we learned that the
following day about 50 inhabitants of the village had searched
the local slopes for B. lomatogona, unfortunately without success.
The slopes of Kyz-Jurdur were described as another site of B.
lomatogona. The dry, southern slopes of this mountain were found
to be extremely overgrazed, while the humid slopes exposed to
the Caspian Sea are not suitable for B. lomatogona. Inhabitants of
the village in the Kyz-Jurdur valley remembered the wild beet
species, B. lomatogona, which grew on the southern slopes, but
was lost years ago through overgrazing. Because the intensity of
grazing did not differ very much within the mountainous plateau, the chance of detecting other populations of B. lomatogona
was considered to be low, and exploration of the Talysch Mountains in Azerbaijan was stopped.
At Tatuni and Pirsara two old potato varieties were collected.
The farmer at Tatuni presented a red-skinned potato type which
had been grown by his grandfather. The farmer living in Pirsara
presented a blue-skinned potato which had long been cultivated
and consumed by the family because of its good taste. Farmers
sometimes mentioned preferring their own old potato varieties
because of their taste. Potatoes imported from Iran would be
consumed only in case of need.
On the way back from Lerik to Lenkoran, at a small farm in
the village of Hamarmecha, a woman farmer showed us beet
seeds and explained that she had mixed fodder and garden beet
seed in the sample. Her explanations were interesting because
freshly collected seed is sometimes composed of roughly equal
amounts of different varieties or even cultivated taxa. Curators
of collections consider that this results from a seed samplehandling mistake by the collector but, in fact, is caused by
farmer’s seed stock management procedures.
The farmer had bought the lentil sample, AZE 99 20, a long
time ago in Lenkoran and had maintained it since then. The
sample is distinguishable from AZE 99 21 by its smaller seeds,
and is used for soup preparations, while the large– seeded lentil
sample is used as an ingredient in the traditional dish ‘Ploff’. She
had also bought the climber bean accession AZE 99 22 from a
neighbour who had produced the seeds himself. AZE 99 24 has a
lower thousand grain mass than AZE 99 22. The tender green
pods of AZE 99 24 are eaten as well as the dry seeds. Similar
crops and crop uses were found in other villages, such as
Hamuscham and Matlajatag, which are located in remote areas
of the eastern foothills of the Talysch Mountains in the districts of
Astara and Lenkoran along the Penserchay River.
From Astara, on the Iranian frontier, the journey continued to
Ardabil, a large city in the southern foothills of the Talysch
Mountains. In 1990 and 1991 the region was explored by a
national team which had found B. lomatogona at three different
locations. Some of the populations sampled then were very small,
and one of the objectives of the 1999 mission was to trace larger
populations in the area. Trips from Ardabil to known collecting
sites were disappointing. Only at one site (Gerdeh, Ardabil district) three plants had survived in a field margin. The main cause
of plant and population losses proved to be ploughing for cultivation of crops like alfalfa (for example at Namin, Ardabil
district).
Only single populations of B. lomatogona and B. vulgaris
subsp. maritima were sampled during the mission. As northwest
Iran and Azerbaijan are probably the northeastern limit of the
distribution area of both species, it was known in advance that
tracing new material would be difficult. The narratives of local
people and the experiences of the local agricultural experts show
that overgrazing and intensified use of arable land has caused
genetic erosion in B. lomatogona.
The distribution area of this wild beet is separated into
uneven parts, the major part in central Anatolia and the minor
part in Armenia, northwest Iran and southern Azerbaijan. Buttler
(1977) noted that B. lomatogona is not a typical plant of natural
habitats, but should be considered as segetal flora preferring
cultivated land. Land cultivation probably enabled the species to
invade from the arid highland steppe. Although land disturbance
by the early farmers favoured the distribution of the species in
cereal fields and field margins, modern agriculture and overuse
of land is threatening the existence of B. lomatogona today. This
wild beet has a narrow ecological tolerance and cannot survive
under more humid conditions. Land cultivation with modern
ploughs and the shift from dryland farming of wheat to irrigated
cultivation of sugar beet will therefore threaten the species. E. de
Meijer, a team member of the collecting expedition in Turkey
(Anonymous 1990), noticed that the central threshing places in
Turkish villages contributed to the dissemination of the B.
lomatogona, which is harvested together with the wheat, brought
to the central threshing place and redistributed from there. Modern seed threshing equipment will probably prevent this seed
distribution mechanism. In the Eskisehir province in Turkey the
species is abundant and almost continuously distributed. As the
species frequently occurs in field margins, land cleaning
programmes can also contribute to the decline of the species
(Anonymous 1990). There are, therefore, good reasons to assume
that there is a need to monitor the population density and distri-
26
bution to prevent unexpected loss of genetic diversity. The observations made in Turkey may explain why almost no wild beet
plants were found in Iran, where large fields with small field
margins and few field bushes dominate the landscape in the drier
area around Ardabil.
Due to time constraints, it was impossible to visit all the
historical sites of B. vulgaris subsp. maritima, or to get a clear idea
of today’s distribution in Azerbaijan. Some places are on islands
in the Caspian Sea (Table 2), the names or locations of which were
unknown even to our Azerian partners. A search for B. vulgaris on
islands and inland sites would be very interesting for two reasons: firstly, the subspecies may be endangered in Azerbaijan
and on the Iranian coast of the Caspian Sea; secondly, because of
the saline soils and warm climate, ‘maritima’ populations from
these areas could contain highly salt- and drought-tolerant forms.
In addition, while looking for ‘maritima’ populations, collectors
could search more systematically for landraces of B. vulgaris
subsp. vulgaris. Examples from this area are still under-represented in the world Beta holding.
Conclusions
This was one of the first plant explorations in the Talysch Mountains for many years. During the 2-week mission only a fraction
of the area could be visited, making an objective assessment of
the situation difficult. Nevertheless, there are strong indications
that in B. lomatogona, a key species of the irano-turanian flora,
genetic erosion has taken place at the margin of its distribution
area, which may progress to the distribution centre with the
intensification of agricultural production practices and increasing human population pressure. To allow the development of an
in situ and on-farm management programme specifically focussing on the wild flora and traditional crops of the Caucasus and
Transcaucaus region, a systematic assessment of the threat of
genetic erosion and factors causing it should be considered. In
some cases urgent measures must be undertaken to safeguard
unique plant populations in Azerbaijan, such as the historical
remainder of B. lomatogona at Geledera (district of Lerik) and B.
vulgaris subsp. maritima at Shorsulu (district of Salyany). The
wild beet population at Geledera could best be protected by
establishing an in situ conservation project, whereas B. vulgaris
subsp. maritima is used by the local people and would qualify for
an on-farm management project. Currently, with the use of satellite remote sensing and geographic information systems (GIS),
maps with a scale of 1:50,000 or 1:25,000 are being produced for
the Ministry of Agriculture in Azerbaijan. The management and
monitoring of in situ and on-farm projects could be assisted by
the national remote sensing centre that the Government of
Azerbaijan intends to establish (www.fao.org, News & Highlights, FAO, 13 Dec. 1999).
Acknowledgements
We are indebted to everyone in the local administrations who
supported our mission through their advice and the provision of
guides and experts. We are also very grateful to all the people we
met in the villages who shared their knowledge with us and
always offered their hospitality. Their enthusiasm and their interest in the objectives of our mission make us believe that, with their
help, it should be possible to develop in situ and on-farm conservation projects.
The exploration was funded by the German Ministry of Food,
Agriculture and Forestry (BML) through the project 100 of the
German–Russian programme on co-operation in agricultural science. Considerable logistic support was also provided by the
Sugar Beet Seed Institute (S.B.S.I) and the S.B.S.I. staff at the
branch office in Ardabil. We greatly appreciate the excellent
cooperation between the different Ministries and the local authorities in Azerbaijan, Iran, Russia and Germany.
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Plant
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Resources
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Newsletter,
Newsletter,
2001,
2001,
No. 126:
No. 126
27 - 27
30
ARTICLE
Evaluation of variability in natural populations of
peperina (Minthostachys mollis (Kunth.) Griseb.),
an aromatic species from Argentina
M. Ojeda1, R. Coirini2,, J. Cosiansi3, R. Zapata2 y J. Zygadlo4
1 Genética, Facultad de Ciencias Agropecuarias, Universidad Nacional de Córdoba, C.C.: 509,
Ciudad Universitaria, (5000) Córdoba, Argentina
2 Manejo de Agrosistemas Marginales, Facultad de Ciencias Agropecuarias, Universidad Nacional de Córdoba,
C.C.: 509, Ciudad Universitaria, (5000) Córdoba, Argentina
3 Maquinaria Agrícola, Facultad de Ciencias Agropecuarias, Universidad Nacional de Córdoba, C.C.: 509,
Ciudad Universitaria, (5000) Córdoba, Argentina
4 Instituto Multidisciplinario de Biología Vegetal, Facultad de Ciencias Exactas, Físicas y Naturales,
Universidad Nacional de Córdoba, Ciudad Universitaria, (5000) Córdoba, Argentina
Summary
Résumé
Resumen
Evaluation of variability in
natural populations of
peperina (Minthostachys mollis
(Kunth.) Griseb.), an aromatic
species from Argentina
Évaluation de la variabilité dans
des populations naturelles de
Minthostachys mollis (Kunth.)
Griseb., une espèce aromatique
d’Argentine
Estudio de variabilidad en
poblaciones naturales de
peperina (Minthostachys mollis
(Kunth.) Griseb.), una especie
aromática de la Argentina
Commerce in aromatic plant species is
expanding. Minthostachys mollis (Kunth.)
Griseb, peperina, is among the most intensively cultivated native aromatic species in Córdoba, Argentina. It is valued
for its digestive properties and it is also
used in the beverage and candy industries. An expanding market for peperina
products requires rational exploitation of
this valuable resource to prevent irreversible loss of germplasm resulting
from over-collecting. Peperina germplasm was collected in situ for characterization and to assess diversity among
accessions. Eight collection sites were selected covering a wide range of habitats
within the Chaqueña Serrana phytogeographical province . Morphological characteristics, essential oils and topographical, meteorological and soil data were
recorded. Regular observations in those
areas revealed that plants growing in different locations varied considerably in
height and leaf characteristics. Populations that contained a high number of
erect plants tended to be smaller and had
more branches, probably resulting from
intense pruning and subsequent regrowth. The identification of preferred
habitats and the characterization of meteorological and soil conditions there will
allow reintroduction into depleted areas.
The data summarized in this preliminary
report indicate the substantial variability
of peperina growing in Argentina. For
example, when plant height was 30.12–
250.28 cm, leaf length ranged from 0.70
to 5.30 cm and leaf width from 0.30 to
2.60 cm. There was also significant variation in essential oil content. For example,
pulegone content ranged from 3.90 to
65.10%.
Le commerce des espèces de plantes aromatiques se développe. Minthostachys mollis (Kunth.) Griseb, « peperina » en espagnol, est l’une des espèces aromatiques indigènes les plus abondamment cultivées à
Córdoba, Argentine. Elle est appréciée
pour ses propriétés digestives et est également utilisée dans des boissons et en confiserie. L’expansion du marché de M. mollis
nécessite une exploitation rationnelle de
cette ressource précieuse afin d’éviter une
perte irrémédiable de matériel génétique
résultant de récoltes excessives. Le matériel génétique de M. mollis a été récolté in situ
afin de le caractériser et d’évaluer la perte
de diversité parmi les accessions. Huit sites
de collecte ont été choisis, couvrant une
grande variété d’habitats dans la province
phytogéographique Chaqueña Serrana.
Les caractéristiques morphologiques, les
huiles essentielles et les données topographiques,
météorologiques
et
édaphiques ont été enregistrées. Des observations régulières dans ces régions font
apparaître des variations importantes dans
la hauteur et les caractéristiques foliaires de
plantes situées en différents endroits. Dans
les populations comportant un grand
nombre de plantes dressées, celles-ci tendent à être plus petites et à avoir des
branches plus nombreuses, probablement
en raison de coupes importantes et de repousse ultérieure. L’identification des habitats préférés et la caractérisation des conditions météorologiques et édaphiques permettront une réintroduction dans les régions où l’espèce s’est raréfiée. Les données résumées dans ce rapport préliminaire indiquent la variabilité importante de
M. mollis en Argentine. Par exemple, lorsque la plante atteint une hauteur de 30,12
à 250,28 cm, la longueur de la feuille varie
de 0,70 à 5,30 cm et la largeur du limbe de
0,30 à 2,60 cm. On observe également des
variations significatives dans les teneurs en
huiles essentielles. Par exemple, la teneur
en pulégone varie de 3,90 à 65,10 %.
El comercio de especies aromáticas se
encuentra en expansión. La peperina,
(Minthostachys mollis (Kunth.) Griseb), es
una especie aromática, muy explotada
en Córdoba, Argentina. Se la valora por
sus propiedades digestivas y se la usa en
las industrias de bebidas y dulces. El mercado en expansión de productos de especies aromáticas requiere una explotación
racional para prevenir perdidas irreversibles de germoplasma por prácticas indiscriminadas de explotación. Se
recolectó germoplasma para su caracterización in situ y para determinar su diversidad. Fueron tomados ocho sitios de
recolección dentro de un amplio rango
de hábitats dentro de la provincia fitogeográfica del Chaco Serrano. Se tomaron datos de caracteres morfológicos,
aceites esenciales, topográficos, meteorológicos y de suelo. Se encontró una
considerable variabilidad en altura de
planta y caracteres de las hojas, sobre
todo entre diferentes localidades. Algunas poblaciones tienen un número alto
plantas erectas, éstas son generalmente
más bajas y con muchas ramificaciones,
esto probablemente debido a la reiterada poda y posterior rebrote. La identificación de los hábitats, con las características meteorológicas y de suelo marca un
camino para la reintroducción de
propágulos. Los datos aquí evaluados
brindan una idea del amplio rango de
variabilidad de la peperina que crece en
la Argentina. Por ejemplo: altura de planta va de 30.12 a 250.28 cm, largo de hoja
de 0.70 a 5.30 cm y ancho 0.30 a 2.60 cm.
Se determinó una muy importante variación en aceites esenciales, como en pulegona que varió entre 3.9 y 65.10%. Se
encontró una fuerte interacción genotipo—ambiente que afecta caracteres
cuantitativos en general y a los componentes de aceites esenciales en particular.
Key words: Essential oil, germplasm,
Minthostachys mollis, peperina, population variability
28
Introduction
Trade in aromatic species is expanding in Córdoba Province,
Argentina, and is based mainly on harvesting natural stands
(87%) and not on agricultural production. A growing market for
products from aromatic species requires rational exploitation of
these valuable resources to prevent irreversible loss of germplasm
resulting from indiscriminate exploitation.
Minthostachys mollis (Kunth.) Griseb (= Minthostachys
verticillata (Griseb.) Epling) (Boelcke 1992; Bonzani and Ariza
1993; Gupta 1995; Retamar et al. 1996), peperina, is among the
most intensively harvested aromatic species. Due to its menthol
content its aroma resembles that of mint. It is valued for its
digestive properties and it is also used in the beverage and candy
industries. It is the only native aromatic species for which there is
international demand (Lagroteria de Galán et al. 1987; Bocco et al.
1993).
Peperina is a perennial shrub 0.30-2 m high. It grows naturally in the northwest and central areas of Argentina and is also
native to Bolivia, Perú and Ecuador. In Argentina it grows in the
mountainous areas of Córdoba at altitudes between 700 and
1200 masl. It is also found in the provinces of Catamarca,
Tucumán and La Rioja, (Epling 1935–1937; Dimitri 1972). Previous studies detailed the essential oil content of this species
(Retamar et al. 1995; Zygadlo et al. 1996).
Peperina is already endangered, and current exploitation
exceeds the rate of natural regeneration. For this reason, there is a
need for immediate conservation measures. Studies of its population biology and genetic diversity are important for the successful development of conservation strategies. Therefore, germplasm
of Minthostachys mollis was collected to determine the extent of its
variability.
Collecting expeditions were planned for a wide range of habitats
within the Chaqueña Serrana phytogeographical province , which
is characterized by mountains with xerophytic forests and native
perennial grasses (Cabrera 1976). Local expertise was used in the
planning stages. Within this area, the Sierras de Córdoba is where
peperina is most abundant and where most plants for sale are
collected.
Collections were undertaken in April and May 1997 and
included areas from 26ºS to 32ºS, and from 650 to 1600 masl.
Collection sites were photographed and representative herbarium
specimens from all locations were collected, classified and stored
at our Faculty Herbarium. Latitude, longitude, altitude and
meteorological data were recorded (Table 1). Notes were made on
the distribution of plants and their densities, harvesting pressure,
habitat and slope, species composition of the habitat and soil
characteristics (Table 2).
The plant material collected in the field was processed in the
laboratory. A sample of seeds from each plant was catalogued.
Part of the sample comprised a common population sample. The
remaining seeds were labelled, dried and stored in trifoliate bags
at –20 °C in a freezer for long-term conservation. Part of this
sample was desiccated and stored at 0°-5 °C in the refrigerator
for short-term conservation.
In soil samples, organic carbon and pH (H2O 1:2.5) were determined using standard methods (Sparks 1996). Plant height and leaf
characteristics of each population were recorded in the field and also
in the growth cabinet. The volatile leaf constituents from each population sample were steam distilled and analyzed using gas chromatography. Statistical analyses were made in order to determine the
existence of within- and between- population variability.
Table 1. Location and climatic data for accessions of Minthostachys mollis collected in Argentina
Province
Sites
Latitude
Longitude
Altitude
(masl)
Mean temperature °C
January
July
Rainfall
(mm)
Córdoba
Córdoba
Córdoba
Córdoba
Tucumán
Tucumán
Catamarca
San Luis
Villa Allende
Capilla del Monte
Tala Cañada
Cuesta Blanca
Tafí del Valle
Escaba
Balcosna
Merlo
31°17’15”
30°48’45”
31°20’15”
31 27’45”
26 52’00”
27 38’40”
27 54’40”
31 23’19”
64 16’16”
64 30’51”
64 54’00”
64 31’42”
65 38’47”
65 39’31”
65 43’10”
64 59’25”
650
1060
1600
800
1600
1100
900
1250
23.4
22.6
21.5
22.5
21.9
25.2
23.4
22.6
700
608
640
700
1300
1100
650
660
9.5
10.4
9.3
9.3
11.8
14.5
9.2
9.3
Table 2. Site characterization in Argentina from where Mintostachys mollis accessions were collected
Sites
Number
of plants
sampled
Distribution
pattern
Extraction
pressure
Local
exposition
Slope
(%)
Soil characteristic
Organic
pH
Matter (%)
Villa Allende
Capilla del Monte
Tala Cañada
Cuesta Blanca
Tafí del Valle
Escaba
Balcosna
Merlo
40
40
40
39
40
33
23
40
Isolated
Continuous
Ecodemo
Ecodemo
Ecodemo
Ecodemo
Ecodemo
Isolated
Moderate–High
Moderate
Moderate
Moderate–High
Low
Moderate
Low
High
S
S
E–SE
S
S–SW
S–SE
S
S
10–20
30–40
30–40
20–30
20–30
30–50
30–50
10–20
12.7
7.5
10.2
9.7
5.9
1.7
0.5
7.2
6.4
5.7
6.1
5.6
5.2
7.2
7.6
5.5
Ecogeographical distribution
Peperina was collected in the region from Tafi del Valle (26º52’S),
province of Tucumán, to Merlo (32º23’S), province of San Luis. Eight
collection sites were selected. Peperina is present in the phytogeographical province of Selva Tucumano-Oranense (Tafí and Escaba)
according to the classification of De Fina (1974) and in the Sierras de
Córdoba, (Cabrera 1976). Collection sites are shown in Fig. 1.
Seeds and leaves were collected from 33 to 40 individual plants
per site, except in Balcosna, where only 23 samples were taken due to
the low population density. Accessions exhibited considerable morphological variability throughout the area of distribution. (Table 1).
Germplasm collection
The accessions were collected in different ecogeographical regions.
Peperina is usually found on the southern slopes of the mountains.
In lower forested areas it is always found as a lower canopy plant,
whereas at unforested higher altitudes, it is found in the open.
Populations were distributed in very restricted pockets
(ecodemos) comprising isolated plants, probably due to overcollection. In order to collect a large number of plants it was
necessary to find sites of limited accessibility, where the slopes
were 10% to 50%, and where exploitation was slight (Table 2).
Plants with shallow roots and soils containing abundant
humus characterized all sites. Soils were classified into two
groups. The first group comprised Escaba and Balcosna soils
that were slightly alkaline (pH 7.2 and 7.6) and low in organic
matter content (1.7 and 0.5). The second group, supporting the
Tafí, Merlo and Córdoba populations, were acid soils (pH <7)
with high organic matter content (>5.9) (Table 2).
In situ variability
Epling (1939) described M. mollis as being between 100 and 200
cm tall and with 1-4 cm leaves, but Dimitri (1972) reported
plants having 30–200 cm stems and 1–5 cm leaves. In our study
plants growing in different locations showed considerable morphological variability (Table 3). Our data indicate a much wider
Fig.1 Collection sites for Minthostachys mollis.
Table 3. Morphological characterization of Mintostachys mollis accessions collected in Argentina
Leaf sizes
Plant height (cm)
Growth
Long (cm)
Broad (cm)
Ratio L/B
Sites
means
s.e
erect (%)
means
s.e.
means
s.e.
means
Villa
Allende
Capilla
del Monte
Tala
Cañada
Cuesta
Blanca
Tafí del
Valle
Escaba
Balcosna
Merlo
108.07 a
33.15
47.5
1.60 e f
0.50
0.86 c d
0.28
79.55 b
31.77
57.5
1.73 d e
0.59
0.87 c d
90.13 b
31.61
45.0
2.22 b
0.78
74.49 b c
24.52
87.5
1.48 f
118.27 a
49.75
20.0
89.85 b
120.00 a
62.12 c
40.01
44.52
21.30
45.0
52.2
85.0
Branching
means
s.e.
1.51 c d 0.92
2.65 e
1.81
0.34
1.67 c d 1.16
9.21 d
6.39
1.18 b
0.39
2.86 b
1.18
15.87 a.b 6.81
0.42
0.78 d
0.30
1.26 d
0.81
10.92 c d
7.63
1.99 c
0.67
0.95 c
0.27
2.01 c
1.11
16.50 a
6.65
2.62 a
1.92 c d
1.66 e f
0.88
0.62
0.59
1.31 a
0.97 c
0.92 c
0.40
0.30
0.33
3.73 a
2.23
2.01 c
1.33
1.69 c d 1.16
Means with the same letter are not significantly different at P=0.05.
s.e.
14.84 a b 7.21
12.74 b c 8.64
16.13 a b 6.92
30
range; average plant height was 62.13 cm in Merlo, but there were
plants only 30.12 cm tall. In Tafí del Valle, where the mean height
was 120 cm, some plants were 250.28 cm. Overall mean plant
height was 91 cm.
Leaf length ranged from 0.7 cm in Merlo and Villa Allende to
5.30 cm in Escaba. Leaf width ranged from 0.30 cm in Villa
Allende to 2.60 cm in Tafí del Valle, but the means, 1.89 and 0.98
cm, respectively, were low in contrast with the extreme values.
This indicates that leaves were small in a large number of the
samples.
Generally, plant height and leaf size decrease with increasing
altitude, as reported for other species (Bhadula and Purohit
1994; Bhadula et al. 1996). This was not exactly so with peperina.
Our analysis showed a tendency for a decrease in plant size in
those populations where harvesting was intense, in Cuesta Blanca
and Merlo. This was accompanied by a greater number of
branches (16.5 and 16.13) and a large percentage of erect plants
(>85%). This habit probably results from intense pruning and
subsequent regrowth.
Oil composition of M. mollis has been reported by other
authors (Retamar et al. 1995, 1996; Zygadlo et al. 1996), and its
variability from one population to another was remarked upon.
Composition of the essential oils was analysed in all populations
(Table 4) and so were their main constituents, pulegone and
menthone. The variation in pulegone was from 3.9 to 65.1%. In
nearly all of the populations pulegone and menthone together
constituted the bulk of the essential oils. Only in Balcosna were
the levels of menthone and pulegone low (about 9%). Such variation is an important consideration for future use of peperina. The
variability between populations is more important than that
within. It is necessary to initiate studies for developing conservation and selection strategies according to oil content characteristics.
The prevalence of low plant population densities suggests the
need for germplasm conservation measures. In this context, the
identification of preferred habitats and the characterization of
meteorological and soil conditions will facilitate preservation of
peperina.
The data summarized in this preliminary paper indicate the
extent of variability in peperina growing in Argentina. As genotype by environment interaction is high for morphological characters and oil content, genetic variability could be masked by
Table 4. Composition of essential oils and their main
constituents in different populations of Mintostachys
mollis
Population
Menthone
Pulegone
Other
essential
oils
V. Allende
C.Monte
Tala Cañada
C. Blanca
Tafí del Valle
Escaba
Balcosna
Merlo
54.1
27.5
29.5
32.1
20.4
17.2
5.2
24.4
36.0
63.7
44.4
35.6
41.0
32.0
3.9
65.1
9.9
8.8
26.1
32.3
38.6
50.8
90.9
10.5
environmental factors. Therefore, as a first step, a systematic trial
must be designed to analyze the genetic variability and to develop suitable strategies for conservation, evaluation, characterization and eventual domestication of this species.
Acknowledgements
The first author thanks the International Foundation for Science
and Universidad Nacional de Córdoba, Facultad de Ciencias
Agropecuarias, for the financial assistance provided to carry out
this study.
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Newsletter,
Newsletter,
2001,
2001,
No. 126:
No. 126
31 - 31
40
ARTICLE
Morphological and isoenzyme variability of taro
(Colocasia esculenta L. Schott) germplasm in Cuba
Arlene Rodríguez Manzano1, Adolfo A. Rodríguez Nodals1, María I.
Román Gutiérrez2, Zoila Fundora Mayor1 and Leonor Castiñeiras Alfonso1
Instituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT), Calle 1 esq. 2, Santiago de las Vegas,
La Habana, Cuba. Tel.: +53 7 579010; Fax: +53 7 579014; Email:[email protected]
2 Instituto de Investigaciones en Viandas Tropicales (INIVIT), Cuba
1
Summary
Résumé
Resumen
Morphological and isoenzyme
variability of taro (Colocasia
esculenta L. Schott)
germplasm in Cuba
Variabilité morphologique et
isoenzymatique du germeplasm
de Colocasia esculenta L.
Schott á Cuba
Variabilidad morfológica e
isoenzimática del germoplasma
de malanga isleña Colocasia
esculenta L. Schott en Cuba
Forty-two accessions of taro (Colocasia
esculenta L. Schott), from the genebank
of the Research Institute on Tropical
Roots and Tubers (INIVIT) were studied.
Forty-two characters showing variability in the subterranean and leafy organs
were selected. Principal component
analysis (PCA) was carried out independently for 16 subterranean and 26 leaf
characteristicss, in order to establish a list
of minimum descriptors (28) enabling
the identification of clones. These were
subjected to another PCA, leading to a
list of descriptors for the creation of a
core collection, with 13 characteristics
representing three main groups and
eight subgroups. The isoenzyme analysis of esterases and peroxidases allowed
characterization of the clones and confirmed that there were no duplicates in
the collection studied, as each clone had
its characteristic band pattern in the esterase system. Applying Jaccard’s similarity index to the groups derived, it was
possible to conclude that there was a
strong African and Japanese, as well as
Southeast Asian and Philippine influence
on the origin of the Cuban accessions.
This result could guide future research
on the origin of this species in Cuba.
On a étudié 42 clones de Colocasia esculenta L. Schott, appartenant à la Banque de
Gènes de l’Institut de Recherches en Racines et Tubercules Tropicaux (INIVIT).
Quarante-deux caractères ayant une
variabilité chez les organes souterrains et
foliaires. On a fait des analyses des composants principaux sur 16 caractères souterrains et 26 caractères foliaires de façon
indépendante. Cela a permis de dresser
une liste de descripteurs minima ayant
28 caractères pour faciliter lídentification
des clones. Ces descripteurs-là ont été
lóbjet dúne nouvelle analyse des composants principaux afin dóbtenir une
liste rendant plus aisée la formation
dúne collection ‘noyau´, laquelle a été
constituée par 13 caractères représentatifs de trois grands groupes comprenant
huit sous-groupes. Les analyses isoenzymatiques désterases et peroxidases ont
permis de caractériser les clones et de
vérifier lábsence de duplicata dans la collection étudiée car chaque clone a eu son
étalon de bandes caractéristique chez le
système isoenzymatique esterase. Les
groupes formés en employant lÍndex de
Similitude de Jaccard ont permis de conclure quíl y a une grande incidence africaine et japonaise concernant la provenance de la collection cubaine, ainsi qu´à
partir du lieu d’origine au sud-est asiatique et aussi à partir des îles Philippines.
Les résultats de cette étude peuvent
servir à des travaux plus profonds sur la
phylogénie de cette espèce à Cuba.
Se estudiaron 42 clones de malanga isleña
Colocasia
esculenta
L.
Schott,
pertenecientes al banco de germoplasma
del Instituto de Investigaciones en Viandas Tropicales (INIVIT). Se seleccionaron
para este trabajo 42 caracteres que presentaron variabilidad en los órganos
subterráneos y foliares. Se realizaron
análisis de componentes principales
(ACP) con 16 caracteres subterráneos y
26 caracteres foliares de forma independiente. Esto permitió formar un listado
de descriptores mínimos que facilite la
identificación de los clones, el cual quedó
constituido por 28 caracteres. Estos se
sometieron a un nuevo ACP para crear
un listado que facilitara la formación de
una colección núcleo quedando la misma
constituida por 13 caracteres representativos de tres grandes grupos y 8 subgrupos. Los análisis isoenzimáticos de esterasas y peroxidasas permitieron caracterizar los clones y además corroborar
que no existen duplicados en la colección
estudiada, ya que cada clon tuvo su
patrón de bandas característico en el sistema isoenzimático esterasa. Las agrupaciones formadas empleando el Indice de
Similitud de Jaccard, permitieron inferir
una fuerte incidencia africana y japonesa
en la procedencia de la colección cubana,
así como desde el centro de origen en el
sudeste asiático y también desde las islas
Filipinas. Estos resultados pueden servir
de base a trabajos más profundos sobre
la filogenia de esta especie en Cuba.
Key words: Colocasia esculenta,
Cuba, isoenzyme variability, morphological variability, principal component analysis (PCA), taro.
Introduction
The Global Action Plan for the Conservation and Sustainable Use
of Plant Genetic Resources (GPA) has among its high priority
activities ex situ conservation. In addition, the GPA emphasizes
the need for studies concerning characterization, evaluation and
development of core collections, as these studies are important in
the effective classification of the collections and allow the users
to access their information needs (FAO 1996a).
There are 6000 Colocasia accessions around the world; the six
largest germplasm collections are in Malaysia (22% of the total),
Papua-New Guinea (13%), India (11%), USA (8%), Indonesia
(7%) and the Philippine Islands (6%) (FAO 1996b). At the Re-
search Institute on Tropical Roots and Tubers (INIVIT), Cuba, a
collection of introduced, collected and genetically improved clones
of taro [Colocasia esculenta (L.) Schott] has been held since 1967.
Particular attention is paid to the maintenance and introduction
of economically important species in Cuba, among them the root
and tropical tuber crops that play an important role in human
nutrition.
One of the first attempts to classify and identify taro
germplasm in Cuba was carried out by Roig (1913) in the former
Agronomic Experimental Station of Santiago de las Vegas (now
INIFAT). Roig characterized and identified clones belonging to
32
Table 1. Passport of the clones of Colocasia
esculenta L. Schott studied
Name
Origin
1. Isleña Blanca #2
2. Isleña Mulata #1
5. Isleña Rosada #1
6. Isleña Rosada
Escambray
7. Isleña Rosada
Jibacoa
8. Isleña Rosada
Mayajigua
9. Isleña Rosada
Sancti Spiritus
10. Isleña Violácea
11. Isleña Japonesa
Cuba
Cuba
Cuba
Cuba
Cuba
Cuba
12. Isleña China
13. Madere Graines
14. Madere Soufre
15. Selección
Herradura
16. Camerun 2
17. Camerun 8
18. Camerun 9
19. Camerun 14
20. Camerun 22
21. Camerun 23
22. Isleña Miranda
Source
(Villa Clara)
(Villa Clara)
(Villa Clara)
(Villa Clara)
(Villa Clara)
(Cienfuegos)
26. Isleña Blanca #1
28. Isleña Rosada
Sabanilla
29. CEMSA 75-11
Field collection
Cuba
(Sancti Spíritus)
Cuba
(Sancti Spíritus)
Cuba (Villa Clara)
Cuba (Isla de la
Juventud)
Cuba (Cienfuegos)
Guadeloupe
(Domaine Duclos)
Guadeloupe
(Domaine Duclos)
Cuba (Pinar del Río)
Field collection
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cuba (Villa Clara)
Guadeloupe
(Domaine Duclos)
Cuba (Villa Clara)
Cuba (Matanzas)
Cuba (Matanzas)
Cuba (Villa Clara)
30. México 1
31. México 2
32. México 3
33. Rosada CEMSA
México (Tabasco)
México (Tabasco)
México (Veracruz)
Cuba (Villa Clara)
34. MC-2
Cuba (Villa Clara)
35. Isleña
Cienfueguera
36. Isleña Yabú
Cuba (Cienfuegos)
37. Francesa
38. Sao Tomé
Vietnam
Sao Tomé
and Príncipe
Cuba (Granma)
39. Isleña Rosada
Bayamo
40. Isleña Bayamesa
41. Isleña Granma
42. Panameña
collection
collection
collection
collection
collection
collection
Cuba (Villa Clara)
23. Isleña Rosada #2 Cuba (Villa Clara)
24. Isleña Rosada
Cuba (C. Habana)
Habana
25. Madere Blanc
Field
Field
Field
Field
Field
Field
Cuba (Villa Clara)
Cuba (Granma)
Cuba (Granma)
Panama (Chiriquí)
Field collection
Field collection
Field collection
Field collection
Introduction
Introduction
Field collection
Introduction
Introduction
Introduction
Introduction
Introduction
Introduction
Selection of
somatic
mutations
Field collection
Selection of
somatic
mutations
Introduction
Field collection
Field collection
Field collection
Selection of
somatic
mutations
Field collection
Field collection
Field collection
Selection of
somatic
mutations
Selection of
somatic
mutations
Field collection
the genera Xanthosoma and Colocasia whose identification was
ambiguous, and emphasized the importance of the corm, cormel,
leaf and petiole characters in the evaluation and identification of
the genetic mixtures, as not all the clones have inflorescences.
Rodríguez Nodals (1971, 1979) made several taxonomic studies based on morphological characters. Rodríguez Manzano et al.
(1994, 1998) described the germplasm during 1989–1991, taking
into account passport descriptors and morphological traits, including subterranean, leaf and inflorescence characteristics, as
well as cytogenetic and biochemical aspects. This led to a better
understanding of the systematics of this genus, although the lack
of a statistical analysis reduced the effectiveness of the study.
More recently Rodríguez Manzano et al. (1999a, 1999b) used
multivariate statistical analysis to study the diversity of Colocasia
esculenta existing in Cuba. In crops such as beans, chickpeas, onions
and peanuts, multivariate analysis of agronomic and morphological characters have been used for the characterization, evaluation
and classification of the germplasm in Cuba (Castiñeiras 1992;
Fraga et al. 1996; Shagarodsky et al. 1996; Fundora et al. 1997).
In this work subterranean and leaf morphological characteristics were used, along with esterase and peroxidase isozyme
analysis, to establish a list of minimum descriptors for characterization, genotype identification and formation of a core collection, and to verify that there are no duplicates in the germplasm
collection of Colocasia esculenta L. Schott in Cuba.
Materials
Forty-two clones introduced from Asia, Africa and America
(Table 1), collected in different Cuban regions (Fig. 1), and genetically improved, were used. These clones belong to the national
collection of Colocasia esculenta L. Schott maintained by INIVIT,
Santo Domingo municipality, Villa Clara province, Cuba. Each
clone was kept ex situ in a four-row plot, totalling 80 rows. The
planting distance was 0.90 m between rows and 0.35 m between
plants in row. The two central rows of each plot were evaluated
(40 plants).
Morphological traits
Plants were harvested 10 months after planting and two consecutive years were evaluated.
Sixteen descriptors were used to evaluate corm, cormel and
root characteristics, as well as quality, and 26 descriptors to
evaluate the leaf characteristics (Table 2). The characteristics and
modalities used were those reported by IPGRI (1999 and IBPGR
1980) and Rodríguez Manzano et al. (1999a, 1999b).
Selection of
somatic
mutations
Introduction
Introduction
Field collection
Field collection
Field collection
Introduction
Fig. 1. Geographic distribution of the local and advanced
clones of the Cuban collection.
The characterization results permitted the selection of minimum
descriptors to study the Cuban clones of Colocasia esculenta. For this
purpose principal component analysis (PCA) was carried out, starting from a standardized correlation matrix analysing independently
for the underground and foliar plant organs (n=16 and n=26).
Characteristics that contributed most to variability were determined on the basis of those original variables with greater
influence on the components (C1 at C5), according to the following approach. The mean values from the highest and lowest
eigenvectors were used as the threshold for the selection of the
most contributing variables (Fundora et al. 1992). Associations
between the factors reported by Rodríguez Manzano et al. (1999a,
1999b) were also taken into account.
Another PCA was done using the characteristics selected in
the first analysis in order to select those contributing most to the
variability, for creating the core collection.
According to the C1–C2 interaction, groups of representative
clones were formed. In order to select the significant associations
Table 2. Descriptors used in the clone characterization
according to subterranean and leaf characteristics
Palatability (PAL)
Consistency (CON)
Corm dry matter percentage (DMC)
Cormel dry matter percentage (DMS)
Corm shape (CS)
Corm weight (CW)
Corm flesh colour of the central part (CCF)
Fibre degree (FD)
Number of cormels (NC)
Percentage of cormels under 50 g (PCV)
Percentage of cormels over 100 g (PCO)
Percentage of cormels between 50 and 100 g (PCE)
Shape of cormels (CLS)
Flesh colour of cormels (CFL)
Bud colour (BC)
Root colour (ROC)
Growth habit (GH)
Shoots after 5 months (S5M)
Shoots after 6 months (S6M)
Shoots at harvest time (SHT)
Plant height (PH)
Petiole to lamina length ratio (PSR)
Maturity at harvest time (MHT)
Leaf blade margin colour (CSE)
Leaf lamina length to width ratio (SLW)
Leaf lamina surface (SS)
Leaf blade colour—upper (SCU)
Leaf blade colour—lower (SCL)
Petiole junction pattern (upper surface of leaf) (LPU)
Petiole junction pattern (lower surface of leaf)
Colour of petiole junction pattern (upper surface of leaf) (CUP)
Colour of petiole junction pattern (lower surface of leaf) (CLP)
Colour of V vein pattern (upper part of leaf) (VCV) (Fig. 2)
Colour of I vein pattern (upper part of leaf) (VCI) (Fig. 2)
Colour of A–B vein pattern (lower part of leaf) (VAB) (Fig. 2)
Petiole colour (PC)
Leaf sheath colour in outer part (CAO)
Leaf sheath colour in inner part (CAI)
Colour of the petiole to corm insertion point (CI)
Wax in the petiole (WP)
Petiole transverse section (PTS)
Ratio of sheath length to total petiole length (PLR)
between the characteristics of the leaf and subterranean organs,
limits of chance with n=40 df and a significance of 0.001%
(Sigarroa 1985) were used.
Isozyme analysis
To study peroxidase and esterase isozyme variation, the techniques of Gonzáles and Román (1982) and Gonzáles (1989) were
used for electrophoresis and preparation of leaf extracts.
From the zymogram results of the peroxidase and esterase
isozyme systems (Rodríguez Manzano et al. 1998) similarities between clones were calculated using the MAT–GENE statistical
programme (Sigarroa and Cornide 1995) based on Jaccard’s similarity index. Data were recorded as the presence or absence of bands.
Data on the similarity matrix were introduced in the programme
database and processed by a cluster analysis in order to represent the
phenetic relationships between clones by means of a dendrogram.
The number of loci and alleles per locus, as well as the
percentage of polymorphic loci and the number of alleles per
polymorphic locus, were determined in both the isozyme systems, according to the following formulae:
Number of polymorphic loci
Percentage of polymorphic loci =
x 100
Total number of loci
Number of alleles per polymorphic locus
Mean number of =
x 100
alleles per locus
Number of polymorphic locus
Morphological and statistical traits
From the results of the matrix of eigenvectors and values for the
subterranean organ characteristics, the major descriptors influencing 70.9% of the total variability were accumulated until the fifth
component could be selected (Table 3). Root colour (ROC), bud
Table 3. Matrix of eigenvectors and values of the
principal components for the subterranean characters
Principal components
Variance
% total
contribution
%
accumulated
Eigenvectors
CS
CW
CCF
FD
NC
PCV
PCE
PCO
CLS
CFL
BC
ROC
PAL
CON
DMC
DMS
C1
C2
C3
C4
C5
4.8831
30.5
1.9492
12.2
1.6611
10.4
1.6132
10.1
1.2338
7.7
30.5
42.7
53.1
63.2
70.9
–0.0018
–0.1532
–0.3399
0.1565
0.2471
0.1181
–0.0978
0.0455
–0.1310
–0.3436
–0.3468
–0.3763
–0.3362
–0.3268
–0.2610
–0.2510
0.4596
0.1717
–0.1783
0.1763
–0.1265
–0.0966
–0.1434
0.0121
0.4572
–0.1800
–0.1573
–0.1284
–0.0790
–0.1401
0.4606
0.3602
–0.0217
0.0931
–0.2354
–0.1275
0.1372
–0.4888
0.3384
0.4776
0.0981
–0.3018
–0.1173
–0.0038
0.3432
0.2571
–0.1115
–0.1018
–0.3450
0.2906
–0.3403
–0.0500
-0.1449
0.2169
0.5082
–0.5064
0.0849
–0.2653
0.0212
–0.0016
0.0795
–0.0252
0.0616
0.0813
–0.3662
–0.1716
0.0895
0.1266
–0.4620
–0.4796
0.1645
0.1168
–0.2847
–0.0749
–0.1013
–0.1768
–0.2210
–0.2495
0.1479
0.2590
34
colour (BC) and cormel flesh colour (CFL) showed the greatest
variability in the first component. Although the colour of the corm
flesh (CCF) had a high value, it was not selected for integrating the
list of minimum descriptors, as Rodríguez Manzano et al. (1999a)
had demonstrated that it was significantly and positively correTable 4. Matrix of eigenvectors and values of the
principal components for the leaf characteristics
Variance
% total
contribution
%
accumulated
Eigenvectors
GH
S5M
S6M
SHT
PH
CSE
CLW
SS
SCU
SCL
LPU
LPL
CUP
CLP
VCV
VCI
VAB
PC
CAO
CAI
WP
CI
PTS
MHT
PSR
PLR
C1
C2
C3
C4
C5
8.8275
34.1
3.8515
14.8
2.1860
8.4
1.7738
6.8
1.3883
5.3
34.1
48.9
57.3
64.1
69.4
0.2432
–0.0087
–0.0060
0.0951
0.0050
0.1790
–0.1621
0.1780
0.2084
0.2275
0.2997
0.2549
0.2948
0.2685
0.2263
0.2404
0.2800
0.2127
0.2459
0.1540
0.2626
–0.0396
0.0877
–0.0731
0.1684
0.0629
–0.0934
–0.2619
–0.2968
–0.3339
–0.3181
–0.2023
–0.0778
–0.2199
–0.1783
0.0115
0.1265
0.1852
0.1531
0.0963
0.1642
0.1107
0.1391
–0.2778
–0.2158
–0.2474
0.0469
0.3383
0.0217
–0.1635
0.1490
0.0450
– 0.0125
0.3852
0.3813
0.1861
0.1938
–0.0447
0.1730
– 0.3070
–0.2212
–0.1118
0.0731
0.1477
0.1368
0.1809
0.1575
0.1010
0.1225
–0.1381
–0.1166
0.0140
–0.0918
0.2783
0.3293
0.2724
–0.0020
–0.1278
0.1875
0.3064
0.1517
0.2542
–0.0661
–0.1209
0.2089
–0.0868
0.1565
0.2719
0.0237
–0.1872
0.0466
0.0622
–0.0788
–0.1509
–0.0067
–0.2096
–0.2118
–0.2644
0.2088
–0.0756
–0.0771
–0.3227
0.2931
0.3743
0.0133
–0.1102
0.0920
0.1214
–0.1009
–0.3481
–0.2757
–0.1068
0.2634
0.1091
–0.1501
–0.1901
–0.0677
0.0301
–0.1257
–0.1068
–0.1376
0.0868
0.0465
0.2121
0.0924
0.2241
0.3653
–0.1906
0.2642
–0.4483
Fig. 2. Vein by the sheet lower part. 1: V part, 2: I part, AB part.
lated with CFL. Consequently, it was sufficient to record CFL for
the characterization because the cormels can be easily removed
from the plant and their colour remains the same, independent of
the plant age. It should be emphasized that the characteristics
selected in the first component are qualitative in nature rendering
the clone characterization. Most of these traits are determined by
one or a few genes and have a discrete distribution. They can be
easily identified and are little affected by the environment, although sometimes their expression may be altered by the action of
modifying genes (Gálvez 1997).
Corm dry matter percentage (DMC), corm shape (CS) and
cormel shape (CLS) were the characteristics that showed the
greatest variability in component 2. Cormel dry matter percentage (DMS) was not selected as it is significantly and positively
correlated with DMC (Rodríguez Manzano et al. 1999a).
The characteristics showing greater influence in C3 were the
percentage of cormels under 50 g (PCV) and percentage of
cormels over 100 g (PCO), as well as the palatability (PAL). The
percentage of cormels between 50 and 100 g (PCE), the corm
weight (CW) and the total number of cormels (NC) were selected
when variables in components 4 and 5 were analysed.
Table 4 shows the matrix of eigenvectors and values from the
principal component analysis for leaf characters in Colocasia
esculenta L. Schott, and the descriptors influencing more in 64.9%
of the variability accumulated up to the fifth component.
The most important descriptors for clone identification were
petiole junction pattern in the upper part of the leaf (LPU), colour
of the petiole junction pattern in the upper part of the leaf (CUP),
and colour of the vein pattern from A to B in the lower part of the
leaf (VAB) (Fig. 2). VAB showed a higher value than the colour of
the veins in the V-shape (VCV) and I-shape (VCI) parts described
by IPGRI (1999) (Fig. 2). Rodríguez Manzano et al. (1999b)
reported a significant association among VCV, VCI and VAB. In
the second component, the colour of the petiole to corm insertion
point (CI), shoots at harvest time (SHT), plant height (PH) and
petiole colour (PC) presented the greatest variability.
Rodríguez Manzano et al. (1999b) reported a significant and
positive association between the shoots at harvest time (SHT)
and shoots at 5 (S5M) and 6 months (S6M). Hence only SHT was
used which, together with plant height, is an important agronomic characteristic for predicting yield per plant.
Considering up to the fifth component, where about 69.4% of
the total variability for leaf organs was accumulated, the ratio of
sheath length to total petiole length (PLR), maturity at harvest
time (MHT), petiole to lamina length ratio (PSR), leaf blade
colour in the lower part (SCL), and leaf sheath colour in the inner
part (CAI) for C4, as well as leaf blade margin colour (CSE) and
leaf lamina length to width ratio (SLW) for C5, were included.
Thus, CAI showed variability important for the characterization
and identification of clones of this genus in the collection studied,
although this was not reported by IPGRI (1999).
A list of 28 minimum descriptors for the correct characterization and evaluation of the Cuban collection of this genus was
established (Table 5): 12 of which represent the subterranean
organs and 16 leaf characteristics. These included both quantitative and qualitative characters, and permitted coverage of an
important part of the existing diversity. Rodríguez Manzano and
1.000
0.028 1.000
0.091 –0.151 1.000
–0.027 0.025 0.263 1.000
–0.341 0.240 –0.123 –0.213 1.000
0.191 –0.070 0.245 –0.485 –0.122 1.000
0.388 0.202 –0.149 –0.132 0.032 –0.086 1.000
–0.045 –0.114 –0.378 –0.075 - 0.076 –0.089 –0.014 1.000
–0.096 0.121 –0.256 –0.052 0.235 –0.176 0.160 0.639 1.000
–0.018 0.221 –0.371 –0.138 0.192 –0.114 0.256 0.625 0.745 1.000
0.000 0.268 –0.208 –0.255 0.332 0.073 0.212 0.338 0.537 0.621 1.000
0.267 0.348 –0.422 –0.178 0.016 –0.130 0.407 0.321 0.276 0.305 0.261 1.000
1.000
–0.209
–0.279
–0.267
–0.119
–0.013
–0.066
–0.100
–0.008
–0.154
–0.148
–0.014
–0.200
1.000
0.265
0.099
–0.147
0.161
–0.032
0.030
0.170
0.018
–0.352
0.084
–0.051
0.261
–0.115
1.000
–0.337
–0.239
0.267
0.229
0.152
0.192
–0.240
0.026
0.041
0.040
–0.103
0.000
0.022
–0.182
1.000
–0.024
0.178
–0.141
0.210
–0.060
0.110
–0.028
0.063
0.078
0.292
0.085
0.264
0.136
0.211
–0.023
1.000
0.289
–0.122
0.163
–0.174
–0.039
0.104
–0.167
–0.196
0.190
0.135
0.133
0.606
0.949
0.707
0.556
0.235
1.000
–0.190
0.160
0.114
0.114
–0.100
0.080
–0.197
0.206
0.097
–0.049
–0.102
0.227
–0.460
–0.160
–0.289
–0.319
–0.274
1.000
0.663
–0.437
0.188
–0.003
0.093
–0.006
0.071
–0.198
0.228
–0.022
0.018
–0.044
0.068
–0.538
–0.426
–0.627
–0.425
–0.260
The underlined correlations are significant at 0.001% probability.
1.000
0.354
0.111
0.133
0.222
–0.189
0.432
0.140
0.202
–0.047
0.014
–0.264
0.052
0.340
0.129
–0.274
0.013
–0.287
0.078
–0.021
1.000
0.923 1.000
0.896 0.852
0.371 0.340
0.148 0.185
0.056 0.122
0.225 0.281
–0.191 –0.239
0.466 0.503
0.217 0.169
0.182 0.209
0.041 –0.025
–0.037 - 0.081
–0.305 –0.310
0.170 0.132
0.147 0.184
0.235 0.118
–0.358 –0.303
0.000 0.051
–0.236 –0.246
0.148 0.143
0.000 0.057
1.000
0.576
0.565
0.441
0.310
0.203
–0.105
0.020
–0.258
0.397
0.154
–0.019
–0.121
0.045
0.052
0.006
–0.142
0.089
–0.445
–0.067
–0.348
–0.062
–0.161
1.000
0.321
0.369
0.264
0.301
0.692
0.383
–0.474
–0.039
–0.199
–0.044
0.181
–0.129
–0.186
0.060
–0.076
0.131
–0.173
0.040
–0.453
–0.421
–0.526
–0.352
0.004
1.000
–0.405
–0.369
–0.419
–0.362
–0.364
–0.276
–0.200
0.000
–0.143
0.121
–0.245
0.101
–0.053
–0.142
–0.030
–0.053
–0.245
–0.011
–0.105
0.329
0.003
0.097
0.046
–0.053
1.000
–0.183
0.492
0.333
0.493
0.352
0.421
0.486
0.314
–0.361
0.000
0.155
0.000
0.110
0.123
0.033
0.015
–0.155
0.086
–0.085
0.328
–0.383
–0.333
–0.319
0.037
–0.186
1.000
0.070
0.009
–0.172
0.116
–0.172
–0.215
–0.305
–0.170
0.084
–0.110
–0.064
0.054
0.264
0.054
–0.181
0.057
0.153
–0.011
0.150
0.030
–0.131
–0.322
–0.116
0.156
0.239
0.402
PAL
ROC
BC
CFL
CLS
PCO
PCE
PCV
NC
CW
CS
PLR
PSR
MHT
PTS
CI
CAI
PC
VAB
CUP
LPU
SCL
SS
SLW
CSE
PH
1.000
0.155
0.148
0.032
0.279
0.181
0.122
0.114
0.117
0.382
0.359
–0.390
0.09
0.102
0.102
0.038
0.014
–0.365
0.694
0.282
–0.241
0.046
–0.145
–0.564
–0.411
–0.551
0.465
–0.479
Roots colour (ROC)
Bud colour (BC)
Flesh colour of cormels (CFL)
Corm dry matter percentage (DMS)
Corm shape (CS)
Shape of cormels (CLS)
Percentage of cormels under 50 g (PCU)
Percentage of cormels over 100 g (PCO)
Percentage of cormels between 50 and 100 g (PCE)
Palatability (PAL)
Corm weight (CW)
Number of cormels (NC)
Shoots at harvest time (SHT)
Plant height (PH)
Leaf blade margin colour (CSE)
Leaf lamina length to width ratio (SLW)
Leaf lamina surface (SS)
Leaf blade colour by the lower part (SCL)
Petiole junction pattern (upper surface of leaf) (LPU)
Colour of the petiole junction pattern (upper surface of leaf) (CUP)
Colour of the vein pattern (A-B vein in the lower part) (VAB) (Fig.2)
Petiole colour (PC)
Leaf sheath colour in the inner part (CAI)
Colour of the petiole to corm insertion paint (CI)
Petiole transversal section (PTS)
Maturity at harvest time (MHT)
Petiole to leaf lamina length ratio (PSR)
Ratio of sheath length to total petiole length (PLR)
SHT
Table 5. Minimum descriptors for characterizing
C. esculenta (L.) Schott clones in Cuba
Table 6. Correlations among the most variable subterranean and leafy organs
Rodríguez Nodals (unpublished data) selected five characteristics of the inflorescences to include in the minimum descriptors
for the morphological characterization.
Table 6 shows the correlations among 28 characteristics included in the minimum descriptor list. Forty-nine significant
correlations were obtained, although for this work only the 20
correlations among the subterranean and leaf characters were
taken into account, as the associations within the leaf and subterranean organs considered independently, were studied by
Rodríguez Manzano et al. (1999a, 1999b).
Four subterranean organ characteristics were involved in correlation among those studied: CFL, ROC, DMS, and NC. Four
leaf characteristics showed significant correlations: SHT, SS, PC
and SCL. Tanimoto and Matsumoto (1986) did not report significant correlations between characteristics of different organs, perhaps because fewer characteristics were studied. Thus, it seems
to be important to use a large number of characteristics in biometric studies of germplasm collections.
The highest correlation (0.949) was found between the colour
of petiole–corm insertion and bud colour, as in all clones both
organs are the same colour, either pink or white, with the exception of ‘Madere Soufre’ whose buds were white and the insertion
point was pink. Incidentally, this clone is the only one with yellow
flesh in the corms and cormels.
The colour of the petiole–corm insertion point correlated
positively not only with bud colour, but also with root colour,
flesh colour of the cormels, palatability, and consistency.
Rodríguez Manzano et al. (1999a) reported significant correlations between cormel flesh, bud and root pigmentation, and palatability, and that 100% of the clones with pink pigmentation in these
SHT
PH
CSE
SLW
SS
SCL
LPU
CUP
VAB
PC
CAI
CI
PTS
MHT
PSR
PLR
CS
CW
NC
PCV
PCE
PCO
CLS
CFL
BC
ROC
PAL
DMC
DMC
36
three organs possessed a delicious, or at least good, palatability.
Therefore, the association of these characteristics with the pigmentation in the petiole–corm insertion point will permit the use of this
descriptor for indirect selection in taro breeding programmes. This
will permit early selections for quality before harvest.
Number of shoots at harvest time showed a positive correlation with number of cormels. Thus, it can be used as a selection
index in yield prediction. It is negatively correlated with the
cormel flesh, and bud and root colour, as well as with the corm
dry matter percentage. That is, when the dry matter percentage in
corms decreases, and when the cormel flesh, buds and roots are
white, the number of shoots per plant increases.
Pandey et al. (1996) studied the correlations between eight
subterranean characteristics influencing yield, and pointed out
that the mother corm and cormel weight can be used as selection
criteria for yield. However, the correlations obtained in the current
study have a great practical importance since the crop cycle is
long and leaf correlations would allow prediction of the future
qualitative and quantitative characteristics of corms and cormels.
The matrix of eigenvectors and values for the minimum descriptors set (Table 7) shows the leaf and subterranean organ traits influencing 61.1% of the variability accumulated up to the fifth component.
For the first three components, which accounted for 46.4% of
the total variability, the most important characteristics in differ-
entiation of clones were bud colour, flesh colour in the cormels
and petiole colour in C1; petiole junction pattern in the upper
part of the leaf, vein pattern colour from A to B in the lower part
of the lamina, colour of the petiole to corm insertion point, and
palatability in C2; and dry matter percentage in corms and plant
height in C3. Other important components of variation were
number of cormels and corm shape in C4, and corm weight in C5.
Although the percentage of different cormels weights (<50 g,
50–100 g and >100 g) contributed more than corm weight in C5
(Table 7), they were not included in the list, since these traits are
greatly influenced by the environment. Inclusion of number of
cormels was found to be sufficient for the formation of core
collections.
Among the characteristics contributing most to the variability
in leaf organs analysis, the petiole colour (PC) was less important
than LPU, CUP, VAB, CLP, CI, SHT, PH and S6M, for the first
three components (Table 4). However, when combining the most
important leaf characteristics with those of subterranean organs
(Table 7), petiole colour had the third highest value among all leaf
characteristics, thus, it is very important for clone identification
due to its great variability (Fig. 3). This result is supported by
Rodríguez Nodals (1979) and Rodríguez Manzano et al. (1994,
1999b). Other important attributes in C1 were bud colour (BC)
and colour of the cormel flesh (CCF).
Table 7. Matrix of eigenvectors and values of the
principal components resulting from the interactions
of the most important subterranean and leaf organs
Variance
% total
contribution
%
accumulated
Eigenvector
SHT
PH
CSE
SLW
SS
SCL
LPU
CUP
VAB
PC
CAI
CI
PTS
MHT
PSR
PLR
CS
CW
NC
PCV
PCE
PCO
CLS
CFL
ROC
BC
PAL
DMC
C1
C2
C3
C4
C5
6.2582
22.4
4.3889
15.7
2.3252
8.3
2.2031
7.9
1.9147
6.8
22.4
38.1
46.4
54.3
61.1
Percentage
50
50
45
40
35
30
25
20
15
10
5
0
11.9
7.14
–0.1750
–0.0879
0.1051
–0.1854
0.0250
0.1743
0.3634
0.3634
0.3375
0.0079
–0.0215
0.2644
0.1775
–0.1539
0.2218
0.0387
0.0856
0.0831
–0.1567
–0.2153
0.1605
0.0727
0.1935
0.0427
0.2372
0.1407
0.2637
0.1728
0.1592
0.3496
–0.1936
0.0620
–0.1845
0.1185
0.0119
0.0463
0.0160
–0.1658
–0.1040
0.1482
–0.0032
–0.2107
0.3592
0.2192
–0.2813
–0.2292
0.1777
–0.0194
0.0894
0.1661
–0.3411
–0.0750
0.0882
0.0376
0.1285
–0.3631
0.2274
0.0371
–0.0601
0.0337
–0.2727
–0.0925
–0.0032
0.0269
0.0656
0.0045
0.1498
0.1695
0.3222
0.3089
0.1074
–0.3346
0.3831
–0.0037
0.3860
0.0970
–0.2411
0.2662
0.1474
–0.0267
0.0846
0.0545
0.0608
–0.0994
0.0129
0.3679
0.1533
–0.2411
0.0645
0.1255
–0.0587
–0.1196
–0.2005
0.0778
0.2594
0.0293
0.1004
0.0989
0.0251
–0.1999
–0.1816
0.2431
0.0728
0.2469
0.3528
–0.3721
0.1627
–0.1675
0.1472
0.1792
0.1689
–0.1106
2
3
4
11.9
4.76
4.76
2.38
1
–0.2524
–0.0038
–0.2249
0.1415
–0.2753
–0.2193
–0.2208
–0.1999
–0.2039
–0.3160
–0.2169
0.2477
–0.0137
0.0270
–0.1136
–0.0713
–0.0348
0.1149
–0.1521
–0.0163
0.0323
–0.0317
0.0087
0.3262
0.2601
0.3272
0.1958
0.1580
7.14
5
6
7
8
Modalities
Fig. 3. Petiole color. (1) Green, (2) green with light
violaceous tint, (3) pinkish-green with purple tint, (4) verde
violaceous with pink tint, (5) green violaceous with white
edges, (6) green violaceous, (7) violaceous-green with
strips, (8) violaceous-greenish with uniform colour.
Table 8. List of descriptors for creating core
collections
Bud colour
Flesh colour of the cormels
Corm dry matter percentage
Corm shape
Corm weight
Number of cormels
Palatability
Colour of the petiole to corm insertion point
Plant height
Petiole junction pattern (upper surface of leaf)
Colour of the vein pattern (A–B pattern)
Petiole colour
Petiole to lamina length ratio
Inflorescence formation
Chromosome number
In addition to the 28 minimum descriptors, including subterranean and leafy organs, necessary to
identify the genotypes of the whole collection, 13 characteristics of more variable attributes (Table 8) were
included in a list of useful descriptors to establish a
core collection. Also included were the chromosome
number and presence or absence of inflorescences.
Twenty clones were selected as representatives of this
variability. Such a list is helpful for quickly accessing
information on variability during collecting missions,
of uncharacterized accessions in the genebank, or in
different regions where in situ conservation in home
gardens is taking place (Esquivel and Hammer 1994;
Esquivel et al. 1994b; Castiñeiras et al. 2000).
These results can be used for identifying variability
present in each geographic niche, for example the clone
‘Isleña Rosada Sabanilla’ collected by Rodríguez
Nodals in February of 1975 in Matanzas Province was
again found near the place where it was originally Fig. 4. Groups formed from the principal component analysis combining
collected (Rodríguez Manzano et al. 2000).
attributes from the subterranean and leafy organs in components 1 and 2.
Characteristics contributing more to the variability
in C1 were bud colour and cormel flesh colour, as well
petioles and violet pigmentation in the limb–petiole insertion
as petiole colour; in C2 they were colour of the petiole junction
point in the upper part of some lower leaves.
pattern in the leaf upper part, and the colour of the vein from A to
Subgroup 4 includes one clone showing green petioles as well
B in the leaf lower part. Based on these traits, three large groups
as pink buds and cormel flesh, located here because of the
and eight subgroups were formed, taking into account the interstrong incidence of the white colour of the root.
action of these characteristics (Fig. 4). It is important to note that
Group II. This group contains 18 clones with pink buds and
clones with a pink root were located at the right side of the X-axis,
except clones 11, 15 and 22, which were placed at the left side, cormel flesh, except ‘Francesa’ (37), which was white, but is
located in this group because of the strong incidence of the pink
due to the strong influence of petiole colour.
Groups I and II included clones exhibiting a green pigmenta- root colour. As in the clones of Group I, Group II did not show
tion in the vein from A to B in the lower part of the leaves, while violet pigmentation in the petiole junction pattern in the upper
group III showed a violet pigmentation. The descriptions of the part of the leaf, although some clones exhibited such pigmentation in some lower leaves: ‘MC-2’ (34), ‘Isleña Bayamesa’ (40)
groups are as follows:
Group I. This group includes 13 clones whose buds and and ‘Isleña Granma’ (41). The petiole colour (PC) was green in all
cormel flesh are white, except ‘Isleña Rosada Bayamo’ (39 in Fig. the clones, with the exception of ‘Isleña Rosada Habana’ (24) and
4), in which they are pink. Nevertheless, this clone belongs to the ‘MC-2’ (34) which were green with a light violet tint. All the clones
group due to the high incidence of white root colour. Only clone had pink roots.
Group III. This group consists of 11 clones with both buds and
38 showed pigmentation in some lower leaves while the others
did not exhibit violet pigmentation in the petiole junction pattern roots white, or with both organs pink. Pigmentation was purple or
in the upper part of the leaf. Vein colour from A to B was always intense purple in the petiole junction pattern in the upper part of
the leaf and veins from A to B, with different variations (Rodríguez
green and root colour was white in all cases.
Subgroup 1 comprised seven clones with violet–green peti- Manzano et al. 1999b). All clones had white cormel flesh, except
oles; four clones (2, 3, 17 and 27) had stripes and the other ‘Isleña China’ (12) and ‘Madere Soufre’ (14) which had white flesh
with violet and yellow tints, respectively.
three (4, 10 and 20) almost uniform colour.
Subgroup 1 includes only ‘Isleña China’ (12), the only clone
Subgroup 2 is made up of four clones with green petioles (26,
having cormel flesh with violet pigmentation, pink buds and
1, 21 and 18).
green–violet petioles. Its root was white.
Subgroup 3 is formed by only one clone (38), exhibiting green
Table 9. Quantitative analysis of the 42 clones zymograms
Isozymatic
system
Number
of loci
Number
of alleles
Number of
rare alleles
Polymorphic loci
percentage (%)
Allele average
number per
polymorphic locus
Peroxidase
Esterase
Total
3
9
12
7
20
27
0
20
20
100
100
100
2,3
2,22
2,25
38
Subgroup 2 is formed by three clones (11, 15 and 22), with
pink buds and white cormel flesh. Petioles were green—violet
with a pink tint. Their roots were pink.
Subgroup 3 includes five clones with pink buds and white
roots. Four of them (13, 30, 31 and 32), however, had white
cormel flesh and green–violet petioles, while the fifth, ‘Madere
Soufré’, had green–rose petioles with a violet tint and yellow
cormel flesh.
Subgroup 4 includes two clones (36 and 42) with white buds,
cormel flesh and roots, and green–violet petioles with whitish
edges.
Isozyme analysis
Peroxidase analysis did not differentiate all the clones, and only
seven bands were visually detected. However, the esterase banding patterns of all cultivars were determined and all showed a
characteristic banding pattern. Within this system, 40 different
bands were found.
Table 9 shows the result of the quantitative analysis of zymograms of the 42 clones. Great variation among individuals was
observed, since each enzyme showed 100% of polymorphic loci,
with an average of 2.25 alleles per polymorphic locus.
The grouping of the clones on the basis of esterase and
peroxidase isozyme patterns is shown in Fig. 5, showing the
associations among individuals, and possible genetic proximity
among them.
Clones were grouped in ascending order in six clusters, and
subgroups within groups III, IV and VI were formed. ‘Isleña
Rosada Escambray’ (6) and ‘Isleña Rosada Sancti Spiritus’ (9)
clustered in isozyme Group I. These clones were collected in
zones that were very close together in the central region, and are
diploid clones (Rodríguez Manzano et al. 1998). In contrast,
‘Isleña China’ was placed in isozyme Group II; it did not associate with other accessions and was the only one with violet pigmentation in the flesh of corms and cormels.
Group III was composed of 12 clones collected in Cuba and
one introduction from Panama, and was divided into three subgroups. Subgroup a included three clones collected in the eastern
region and one in the central region. Subgroup b had only one
clone, ‘Isleña Miranda’ (22), which was obtained by through
selection from ‘Isleña Japonesa’ (11), which was located in subgroup c. In subgroup c there were seven clones—six collected in
Cuba and one introduced from Panama. Within this subgroup,
four clones had very marked morphological similarities: ‘Isleña
Japonesa’ (11), collected in Isla de la Juventud and probably
introduced from Japan (Rodríguez Nodals, pers. comm.);
‘Selección Herradura’ (15) collected in southern Pinar de Rio
province; ‘Isleña Yabú’ (36), a mutation from ‘Isleña Japonesa’
and ‘Panameña’ (42), very similar to clone 11 with respect to leaf
characteristics.
The clones ‘Isleña Blanca #1’ (26), ‘Isleña Blanca #2’ (1) and
‘Isleña Violácea’ (10) were collected in the Punta Felipe municipality, Villa Clara province, in the central region, and differed
from the other clones in the group in leaf and petiole colour.
Clones of this group could have a common origin in Asia, and
could have been introduced directly from Japan or from the
Canary Islands, developing a great variability since their intro-
60
50
40
30
20
10
0
6
9
12
41
39
40
35
22
10
15
26
1
36
42
11
37
27
4
3
2
31
32
30
38
13
16
19
21
20
18
17
5
28
25
14
29
23
34
8
7
24
33
I
II
a
b
III
c
a
IV
b
V
a
b
VI
c
Fig. 5. Dendogram that illustrates genetic similarities among
42 clones from Colocasia esculenta, generated by a cluster
analysis of the peroxidases and esterases isozymatic systems.
The Roman numbers indicate groups and the letters, the subgroups.
duction. These results would confirm the suggestion that C.
esculenta in Cuba is a plant of Asiatic origin (Hammer and
Esquivel 1994). This fact was further demonstrated by the fact
that there are several clones from this species in the Cuban
western and central regions, but in the eastern part there are more
clones from Xanthosoma spp. (Rodríguez Nodals 1984; Esquivel
et al. 1994b; Castiñeiras et al. 2000).
Group IV consisted of eight clones; three collected in Mexico
(30, 31 and 32), four with striped petioles from Cuba (2, 3, 4 and
27) and one introduction from Asia (37). There is much evidence of
the introduction of many Asian plants in Mexico from the Philippines Islands through the route from Manila to Acapulco, and the
exchange between Mexico and Cuba from Veracruz to Havana was
also very intense in the colonial period. Thus it is not difficult to
assume the probable common origin of these clones (Fig. 5).
Four clones with striped petioles (2, 3, 4 and 27), named
‘mulatos’ by Rodríguez Nodals (1979), and a clone from Asia
(37) belong to subgroup a, while subgroup b contains three clones
collected in Mexico with morphological similarities. These are the
only clones with white cormel flesh, pink buds and white roots,
except ‘Madere Graines’, which has a greater African influence.
Group V contained only one clone (38). This clone was recently introduced into Cuba from Sao Tomé and Principe, and
showed no association with any of the other clones studied. Thus,
during the colonial period no germplasm was introduced into
Cuba from the west African Islands.
Group VI consists of 18 clones, six of which were from
continental West Africa (16, 17, 18, 19, 20 and 21), three introduced from Guadeloupe (13, 14 and 25) and the other nine
collected or obtained by selection of somatic mutations in Cuba
from clones of African origin. The germplasm coming from Africa
has a strong influence in this group, since it gave rise to the other
clones from the French Antilles and Cuba by the selection of
somatic mutations.
It is likely that several Colocasia clones were introduced from
tropical Africa, a secondary centre of genetic variation of the
Asian taro, through the slave trade and also with the Spaniards
from the Canary Islands. These results confirm the hypothesis of
Gonzalo Oviedo (cited for Esquivel et al. 1994a). This crop is
related to African customs because this food was prepared in
home gardens in Cuba using ‘pilones’—traditional instruments
used for the slave trade (Esquivel et al. 1994a, Tirado and
Martínez 1994).
Within this group there are three subgroups. Subgroup a,
consists of ‘Madere Graines’ (13). Subgroup b, is formed by
‘Camerun 2’ (16). Subgroup c, includes 16 clones, five of which
came from West Africa (Cameroon), two from the French Antilles,
six collected in Cuba and the other six obtained through clonal
selection from spontaneous mutations of some of the clones
included in this subgroup.
No association was found between the ploidy level reported
by Rodríguez Manzano et al. (1998) and the isozyme groups.
These results confirm those published by Tanimoto and
Matsumoto (1986) and Lebot and Aradhya (1992).
These earlier authors found no correspondence between the
zymotypes and the morphological characteristics studied. However, in this study a correspondence was found between some
groups of clones based on morphological characters and subgroups from the cluster analysis formed on the basis of the
isozyme analysis. This is the case with clones with striped petioles with a probable Asian origin (Group IV, subgroup a) and
those from Mexico (Group IV, subgroup b), which are the only
ones with white cormel flesh and roots and pink buds.
Conclusions
Twenty-eight descriptors, 12 for subterranean organ traits and 16
for leaf characteristics, have been included in the list of minimum
descriptors for taro [Colocasia esculenta (L.) Schott]. These descriptors enable correct characterization and evaluation of collections
of this species in Cuba, covering most of the existing diversity.
PCA for the interaction of the 28 descriptors used in characterizing the clones showed that colour of the limb to petiole
insertion point and the distribution of pigmentation in the upper
part of the leaf, vein colour from A to B in lower part of the sheet,
bud colour, colour of the cormel flesh and petiole colour were the
descriptors contributing most to the total variability and were
determinant in the formation of the groups of clones.
Twenty significant correlations among the characteristics of the
leaf and subterranean organs were found. Some of them are important for the indirect selection in taro improvement programmes.
On the basis of the esterase and peroxidase isozyme analysis,
clones were classified in six groups and several subgroups.
Clones with striped petioles, as well as those with white
cormel flesh, pink buds and white roots were grouped together in
the dendrogram derived from the cluster analysis of the esterase
and peroxidase isozymes, and possibly originate from introductions from Asia.
Twelve polymorphic loci with 27 alleles, 20 of which were rare
alleles, were found for the esterase isozyme system. The average
allele number per polymorphic locus was 2.25.
A taro core collection can be formed using only the proposed
13 descriptors that contributed to the variability, and should be
composed of clones representing the three main groups of variability and the eight subgroups within them. Thus, information
on the variability of the accessions collected or maintained in situ,
as well as on the uncharacterized accessions in the gene bank can
be obtained quickly.
The Cuban taro collection has a strong African and Japanese
influence, as well as influence from the centre of origin in Southeast Asia and the Philippines.
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Inst.
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Plant
Plant
Genetic
Genetic
Resources
Resources
Newsletter,
Newsletter,
2001,
2001,
No. 126:
No. 126
41 - 41
45
ARTICLE
Characterization of the Cucurbita pepo collection
at the Newe Ya’ar Research Center, Israel†
Harry S. Paris
Department of Vegetable Crops, Agricultural Research Organization, Newe Ya’ar Research Center, PO Box 1021,
Ramat Yishay 30-095, Israel. Tel: +972-4-953-9511; Fax: +972-4-983-6936; email: [email protected]
Summary
Résumé
Resumen
Characterization of the
Cucurbita pepo collection at
the Newe Ya’ar Research
Center, Israel
Caractérisation de la
collection de Cucurbita pepo
du centre de recherche Newe
Ya’ar, Israël
Caracterización de la
colección de Cucurbita pepo
en el Centro de Investigación
Newe Ya’ar, Israel
The Cucurbita pepo (pumpkins, squash
and gourds) collection at the Newe Ya’ar
Research Center consists of seed
samples of 320 cultivars, landraces and
wild forms. The seed samples were obtained from both public and private
sources and include 133 hybrids and 187
open-pollinated forms. Most are named
cultivars, but those obtained from plant
introduction organizations usually bore
only numbers. The samples have been
grown out, observed and classified according to subspecies and cultivar-group.
Represented are 241 samples of subsp.
pepo, 69 of subsp. ovifera, two of subsp.
fraterna, two intersubspecific hybrids,
and six mixed, subsp. pepo and subsp.
ovifera gourds. The cultivar-groups of
subsp. pepo contain more than double
the number of cultivars of the cultivargroups of subsp. ovifera. The Pumpkin
Group and the Cocozelle Group (both
subsp. pepo) contain the most open-pollinated cultivars. The Zucchini Group
(subsp. pepo) and the Straightneck Group
(subsp. ovifera) contain the highest proportion of hybrid cultivars.
La collection de Cucurbita pepo (citrouilles, courges et coloquintes) du centre de
recherche Newe Ya’ar comprend des
échantillons de semences de 320 cultivars, variétés locales et formes sauvages.
Les échantillons de semences ont été obtenus auprès d’organismes publics et
privés et ils représentent 133 hybrides et
187 formes à pollinisation ouverte. La
plupart sont des cultivars désignés par
un nom, mais ceux qui ont été obtenus
auprès d’organismes chargés de
l’introduction de plantes sont habituellement uniquement identifiés par des
numéros. Les échantillons ont été cultivés, observés et classés selon les sousespèces et groupes de cultivars. Parmi les
échantillons, 241 appartiennent à la sousespèce pepo, 69 à la sous-espèce ovifera,
deux à la sous-espèce fraterna, deux sont
des hybrides entre sous-espèces différentes, et six constituent un mélange
des sous-espèces pepo et ovifera. Les
groupes de cultivars de la sous-espèce
pepo sont deux fois plus nombreux que
les groupes de la sous-espèce ovifera. La
majorité des cultivars à pollinisation ouverte appartiennent au Groupe citrouille
et au Groupe cocozelle (tous deux faisant
partie de la sous-espèce pepo). La proportion de cultivars hybrides la plus élevée
s’observe dans le Groupe courgette
(sous-espèce pepo) et le Groupe straightneck (sous-espèce ovifera).
La colección de Cucurbita pepo (calabazas,
calabacines) en el Centro de Investigación
Newe Ya’ar consta de muestras de semillas de 320 cultivares, variedades naturales y formas silvestres. Las semillas se
obtuvieron de fuentes públicas y privadas y comprenden 133 híbridos y 187
formas de polinización abierta. La mayoría son cultivares con nombre, pero los
obtenidos de organizaciones dedicadas a
la introducción de especies vegetales
suelen llevar simplemente números. Las
muestras han sido cultivadas, observadas y clasificadas en subespecies y grupos de cultivares. Están representadas
241 muestras de la subespecie pepo, 69 de
la ovifera, dos de la fraterna, dos híbridos
de subespecies y seis calabazas mixtas de
las subespecies pepo y ovifera. Los grupos
de cultivares de la subespecie pepo contienen más del doble de cultivares que
los de los grupos de la subespecie ovifera.
El Grupo Pumpkin y el Grupo Cocozelle
(ambos de la subespecie pepo) son los que
contienen más cultivares de polinización
abierta. El Grupo Zucchini (subesp. pepo)
y el Grupo Straightneck (subesp. ovifera)
contienen la mayor proporción de cultivares híbridos.
Key words: Collection, Cucurbita
pepo, gourd, pumpkin, squash
Introduction
Cucurbita pepo is perhaps the most variable species for fruit characteristics in the plant kingdom (Duchesne 1786; Naudin 1856). It is
native to North America, where it has been cultivated for at least
10 000 years (Smith 1997), but was introduced to Europe only
about 500 years ago (Whitaker 1947). This species includes ediblefruited forms, known as pumpkins and squash, and small-fruited,
often bitter, non-edible forms, known as gourds. Much of the
variability in fruit characteristics among cultivated C. pepo can be
attributed to the different quality characteristics needed for the
culinary use of the mature fruit flesh and seeds as opposed to the
use of young fruits. Today, C. pepo is among the economically
most important vegetable crops worldwide and is grown in almost
all temperate and subtropical regions (Paris 1996).
†
The Cucurbita pepo collection at the Newe Ya’ar Research
Center was begun in 1978 as part of a breeding programme.
Initially, samples were collected on the basis of horticulturally
desirable characteristics. Later, emphasis was placed on obtaining samples from a wide variety of locations and sources, especially of named open-pollinated cultivars, so as to understand
better the history, variability and potential of C. pepo germplasm.
Many of the original seed samples were small and were maintained in the collection through self- and sib-pollination. Although
such inbreeding results in a narrowing of the genetic base of the
original material, it was the only practicable method available for
reproducing and maintaining the genetic material. This method
was also employed for commercial hybrids, as this was the only
Contribution No. 104/00 from the Institute of Field & Garden Crops, Agricultural Research Organization, Bet Dagan, Israel.
42
way of maintaining their genetic constitution. At present seeds of
some 320 accessions are available for distribution.
This paper describes the breakdown of these 320 accessions
into subspecies, cultivar-groups and, for the open-pollinated
sorts among them, continents of origin, and in so doing aims at
contributing to better understanding of how this crop species has
developed.
Seed samples were obtained over the past 22 years from commercial sources, seed cooperatives, genebanks, geneticists, botanists,
breeders, plant introduction organizations, extension agents, family members, friends and travellers, almost entirely in North
America, Europe and Asia. The samples obtained from commercial sources included cultivars described and illustrated in catalogues of seed companies. Mostly, these were modern hybrids.
Information was scanty for most of the old cultivars and landraces.
Some samples had no name. For many, the location from which it
was collected, other than the country, was not known.
Nearly all the samples have been grown out in the field at least
twice. Although only a small part of the collection could be grown
out in a single season, the conditions at Newe Ya’ar allowed for two
seasons in the field per year, a spring–summer season (sowing in
March or April) and a summer–autumn season (sowing in late July
or early August). Cultural practices included direct sowing in the
field, drip irrigation, preplant and drip fertilization, and either bare
ground (spring–summer seasons) or silver plastic mulch (summer–autumn seasons). Based on information available before
growing out the samples, similar accessions were grouped together
in the same season and next to or near one another in the field for the
purpose of comparison. Plants were observed for stem colour, leaf
size and shape, presence or absence of silver mottling of the leaves,
vine or bush growth habit, presence or absence of branching, relatively closed or open growth habit, fruit shape, developmental fruit
colour and other characteristics.
There is much confusion regarding names of cultigens in many
crop species, Cucurbita pepo among them. Often, different names
have been used for the same cultivar and the same name for
different cultivars. For the purpose of this study, two samples were
considered to be from the same cultivar if their plants did not differ
phenotypically from one another in at least one trait, as expressed
by the majority of the plants grown out from each sample, even if
the two samples bore different names. If the plants of two samples
differed in one or more phenotypic traits (as expressed by the
majority of the plants of each sample), the samples were considered to be different cultivars, even if bearing the same name. For
example, the ‘Grey Zucchini’ samples I obtained from the U.S.A.
had plants that were identical phenotypically to the sample of
‘Faentina’ from Italy and to those of ‘Verte Petite d’Alger’ from
France: the three names are used for the same cultivar. On the other
hand, the plants of the sample of ‘Verte Non-Coureuse d’Italie’
from one French company were distinct from those grown from a
sample of identical name from another French company. These I
considered to be separate cultivars.
Another problem of nomenclature is presented by the misleading names of some cultivars. For example, the ‘Grey Zucchini’ of the U.S.A. has short, tapered, cylindrical fruits and thus
is a cultivar of vegetable marrow and not a zucchini (Fig. 1).
Likewise, ‘Golden Zucchini’ from South Korea is not the same as
‘Golden Zucchini’ from the USA. The latter has uniformly cylindrical fruits and is indeed a zucchini, but the Korean cultivar has
long bulbous fruits and is, therefore, a cocozelle cultivar.
Classification of samples
Beginning with Duchesne (1786), there have been a number of
attempts at subspecific classification of Cucurbita pepo. I prefer to
use a two-tiered, botanical-horticultural approach (Paris 1996).
Botanically, the species is divided into three subspecies, based on
allozyme variation, seed morphology and various phenotypic characteristics (Decker 1988): subsp. pepo, subsp. ovifera and subsp.
fraterna, the last of these representing wild Mexican C. pepo gourds.
The first two include gourds as well as edible forms. Horticulturally, the edible-fruited forms are divided into cultivar-groups
(Trehane et al. 1995), based on the highly polygenic characteristic of
fruit shape. There are eight such groups: Cocozelle, Pumpkin,
Vegetable Marrow, Zucchini, Acorn, Crookneck, Scallop and
Straightneck (Paris 1986; Figure 1). The first four groups belong in
subsp. pepo and the latter four in subsp. ovifera. Of the non-edible
forms, the ball, orange and warted gourds are more closely allied
with subsp. pepo whilst the egg and pear gourds, as well as wild
gourds of the United States, belong in subsp. ovifera (Decker 1988).
Recent results, obtained using cluster analysis of an inter-simple
sequence repeat multilocus marker system of the DNA (Katzir et
al. 2000), are consistent with this classification.
The 320 accessions were observed and classified according to
subspecies, cultivar-group and whether they are open-pollinated
or hybrid. No attempt was made to determine whether the purported hybrids were indeed such, or were merely said to be so by
the seed company. The open-pollinated forms were further classified according to continent of origin. The hybrid sorts were not
classified as to geographic origin because for many of them that
is uncertain; many have been developed by multinational companies and their subsidiaries and are commercially available in
many countries, but the parents and their countries of origin are
kept secret.
Fig. 1. Fruit shape profiles of the edible-fruited cultivargroups of Cucurbita pepo (after Paris 1986). Peduncular
end of the fruit at top, stylar end at bottom. Top row, C. pepo
subsp. pepo, left to right: pumpkin, vegetable marrow,
cocozelle, zucchini. Bottom row, C. pepo subsp. ovifera, left
to right: scallop, acorn, crookneck, straightneck.
Table 1. Summary description of the 320 samples of Cucurbita pepo at the Newe Ya’ar Research Center
Geographic source of open-pollinated sorts
C. pepo subsp. pepo
Pumpkin
Cocozelle
Vegetable marrow
Zucchini
Mixed/intermediate
Unique
Gourd
C. pepo subsp. ovifera
Acorn
Scallop
Crookneck
Straightneck
Unique
Gourd
C. pepo subsp. fraterna
C. pepo subsp. ovifera x
C. pepo subsp. pepo
C. pepo subsp. ovifera +
C. pepo subsp. pepo
(Gourd mixtures)
Total
Hybrid
Openpollinated
North
America
Europe
Asia
Africa
241
43
43
50
81
16
2
6
69
17
16
9
9
5
13
2
2
111
4
12
25
70
0
0
0
20
6
4
3
7
0
0
0
2
130
39
31
25
11
16
2
6
49
11
12
6
2
5
13
2
0
38
21
3
2
5
1
0
6
42
11
6
5
2
5
13
2
0
68
14
25
13
5
11
0
0
6
0
5
1
0
0
0
0
0
21
4
2
10
1
4
0
0
1
0
1
0
0
0
0
0
0
3
0
1
0
0
0
2
0
0
0
0
0
0
0
0
0
0
6
0
6
6
0
0
0
Results
Fig. 2. Mature fruits of Cucurbita pepo subsp. pepo. Left to right, top row,
Pumpkin group: ‘Connecticut Field’, ‘Khutorianka’, ‘Ukrainska
Nogoplodna’, ‘Cinderella’, ‘Uzbekistan Local Pumpkin’, ‘Porqueira’;
second row, Pumpkin group: ‘Jack O’Lantern’, ‘Winter Luxury’, ‘Tondo
Verde Scuro di Piacenza’, ‘Dagestan’, ‘Small Sugar’, ‘Gourmet Globe’,
‘Spookie’, ‘Ronde de Nice’, ‘Nonkadi’; third row, Vegetable Marrow group:
‘Beirut’, ‘Blanche non-coureuse’, ‘Bianco di Palermo’, ‘Bolognese’,
‘Caserta’, PI 181763 (from Lebanon), ‘Table Dainty’, ‘Gornurekhovskiye’,
‘Vegetable Marrow’, ‘Verte Petite d’Alger’, ‘Vegetable Spaghetti’, ‘Yakor’;
fourth row, left, Zucchini group: ‘Black Beauty’, ‘Fordhook Zucchini’, ‘Nero di
Milano’, ‘Nano Verde di Milano’, ‘True French’, ‘Verde di Milano’; fourth
row, right, Cocozelle group: ‘Alberello di Sarzane’, accession from Burkina
Faso, ‘Cocozelle Tripolis’, ‘Lungo Bianco di Sicilia’, ‘Long Cocozelle’,
‘Lungo di Toscana’, PI 165018 (from Turkey), ‘Striato d’Italia’, ‘Striato
Pugliese’, accession from Slovenia, ‘Verte Non-Coureuse d’Italie’;
foreground, gourds: ‘Flat’, ‘Miniature Ball’, ‘Orange Ball’, wild C. pepo
subsp. fraterna.
Of the 320 samples in the C. pepo collection, 241
(75%) are C. pepo subsp. pepo (Table 1; Fig. 2). Of
these 241, 217 (90%) are classified into one of the
four edible-fruited cultivar-groups of this subspecies. The remaining 24 are gourds (non-edible
fruited) or classified as ‘mixed/intermediate’ or
as ‘unique’. The Zucchini Group has by far the
largest number of cultivars. Each of the other
edible-fruited groups of C. pepo subsp. pepo has
over 40 representatives, more than twice the
number of any of the cultivar-groups of C. pepo
subsp. ovifera. Each of the C. pepo subsp. pepo
cultivar-groups has representatives from three
or more continents. The mixed/intermediate
C. pepo subsp. pepo accessions are not uniform or
they have intermediate fruit shape; most are from
Yugoslavia and Turkey. The unique forms are
the two edible-fruited but small, gourd-size
‘Little Gem’ and ‘Rolet’ from South Africa.
There are 69 samples of C. pepo subsp.
ovifera, 51 (74%) of which are classified into the
four edible-fruited cultivar groups of this subspecies (Table 1). The remainder of the C. pepo
subsp. ovifera are gourds or classified as
‘unique’. The unique forms are ‘Delicata’ and
similar forms from the northern U.S.A. and
‘Jack Be-Little’. C. pepo subsp. ovifera contains a
greater proportion of gourds (19%) than does
C. pepo subsp. pepo (2%). Of the C. pepo subsp.
ovifera gourds in the collection, six are culti-
44
vated whilst the other seven are wild forms. The collection also
contains two samples of wild gourds from Mexico (C. pepo
subsp. fraterna), two commercial hybrids of C. pepo subsp. pepo
x C. pepo subsp. ovifera and six ornamental mixtures of gourds
from the two cultivated subspecies.
The Zucchini Group contains far more cultivars than any
other group, but 70 of its 81 cultivars are hybrids (Table 1). Of the
C. pepo subsp. pepo cultivar-groups, the zucchini contains the
fewest number of open-pollinated cultivars, 11, but this number
is nevertheless greater than the average number of such sorts
possessed by the cultivar-groups of C. pepo subsp. ovifera. The
Zucchini and the Straightneck Groups have the highest proportion of hybrid cultivars: 86 and 78%, respectively. For the other
groups, the proportion of hybrid cultivars is distinctly lower.
Only four of the 43 pumpkins are hybrids and none of the mixed,
unique and gourd forms is.
The Pumpkin Group contains the most open-pollinated cultivars (39), followed by the Cocozelle Group (31). The majority of
the open-pollinated forms of C. pepo subsp. pepo are European.
The cocozelles and vegetable marrows have mainly an Old World
distribution whilst the pumpkin and zucchini cultivars have
approximately equal numbers of cultigens in the Old and New
Worlds. In contrast, the majority of the open-pollinated sorts of
C. pepo subsp. ovifera are North American.
Discussion
Before attempting to interpret what the content of the Cucurbita
pepo collection at Newe Ya’ar might mean historically and with
regard to crop development, the biases that may have affected
the content of the collection need to be considered. The collection
began with the goal of breeding improved forms of C. pepo,
especially zucchini, pumpkins, acorn squash, scallop squash
and, later, cocozelle and vegetable marrow squash. Initially,
emphasis was placed on gathering cultivars from American seed
companies, whose catalogues were easily available. Later, the
collection was supplemented by germplasm ordered from the
United States Plant Introduction System and other collections, as
well as Italian seed companies and cooperatives and colleagues
from North America, Europe and Asia, and by myself and
friends who had visited various localities and purchased seed
packets.
For many of the accessions, only a small quantity of seeds
was supplied and these had to be reproduced in order to assure
their being maintained. Some, most notably plant introductions
from Mexico, were very late to flower and set fruit and therefore
could not be reproduced and maintained. Thus, it could be
expected that this collection would be biased against crooknecks,
straightnecks and gourds, local cultivars from areas to which I
have not had access, and late Mexican forms. On the other hand,
this inherent bias is tempered by the fact that for the past few
years I have made a concerted effort to obtain seeds of all named
open-pollinated C. pepo cultivars. Another tempering of the bias
is that some common cultivars are grown in various, perhaps
unexpected places, under a local name or no name at all. For
example, I received an unnamed seed sample from Nepal in
1982. This was a cocozelle cultivar then unknown to me, and I
listed it as ‘Cocozelle Nepal’. However, in 1989, I received a
sample of ‘Romanesco’ from Italy and when I compared it with
the Nepalese sample, I discovered that they had identical plants
and fruits. Yet another tempering factor is the fact that many
hybrids are short-lived and cannot be reproduced because their
parents are not publicly available. Thus, the collection contains
several crookneck and straightneck hybrids that no longer exist.
However, newer ones have replaced them and the number of
extant cultivars in these groups appears to have changed little in
20 or 30 years.
C. pepo hybrids, first promulgated by Curtis (1940), have
been available commercially for half a century. The number of
hybrid cultivars and the proportion they form of the total number
of cultivars provides an indication of how heavily a given cultivar-group has been bred since the commercial introduction of
hybrid cultivars in the 1950s. From Table 1, it is clear that the
zucchini, with dozens of hybrids, is far and away the most
intensively bred C. pepo. Hybrid zucchini squash are offered in
catalogues of seed companies from North America, Europe and
Asia. The straightneck squash is a small group of cultivars but,
nonetheless, most straightnecks are hybrids, as are a fair proportion of crooknecks; these necked squash hybrids have been developed and commercialized by American seed companies.
Most of the open-pollinated sorts pre-date the hybrids and
therefore give a better indication of the possible geographic origins of the cultivar-groups of this species and of their state of
development before the 1950s. The Pumpkin, Cocozelle and
Vegetable Marrow groups have by far the greatest number of
open-pollinated sorts. I have observed great genetic variation,
expressed in both vegetative and reproductive characteristics,
among the open-pollinated cultivars in each of these groups
(Paris 1996), suggesting that each has been cultivated for a
considerable length of time. The majority of cocozelles and vegetable marrows are from Europe, suggesting that these groups
developed there between 50 and 500 years ago. There are nearly
equal numbers of pumpkins on the two sides of the Atlantic
Ocean. Those of Mexico are often grey–green or black–green and
grey–green striped with thick lignified rinds; those of the U.S.A.
are orange at maturity, grooved and not lignified. Those from
Europe are slightly ribbed, often black-green and orange striped,
with thin lignified rinds; others have bush plants and are small,
black-green or orange, with thin lignified rinds. Thus, distinct
kinds of pumpkins have been developed in both North America
and Europe. The zucchini group contains fewer open-pollinated
cultivars than any of the other C. pepo subsp. pepo groups.
Perhaps other open-pollinated zucchinis had existed formerly
but became obsolete with the advent of many improved hybrids.
I have not observed as much variation among the extant openpollinated zucchini cultivars as I have observed for the other C.
pepo subsp. pepo groups. Thus, it appears that the zucchini group
is a relatively recent development.
The cultivar-groups of C. pepo subsp. ovifera contain fewer
open-pollinated cultivars than their counterparts of C. pepo
subsp. pepo. Again, it is possible that more cultivars had existed
but were replaced by modern hybrids. On the other hand, except
for the scallop group, these are almost entirely North American in
their distribution; only the scallops have any commercial importance elsewhere. The small number of open-pollinated cultivars
of these groups can be attributed to their limited geographical
range of popularity.
The collection is maintained at the Newe Ya’ar Research
Center in a cool (10°C), relatively dry (50% R.H.) seed storage
chamber. Periodically, the collection has been and will continue to
be renewed by self- and sib-pollination. The collection, in addition
to being a germplasm source for breeders, is expected to be
invaluable for obtaining a better understanding of relationships
within C. pepo, especially using the latest techniques of DNA
analysis.
The list of the 320 accessions in the collection is maintained in
a database using Excel® for Windows™ format. The list includes
the name of each cultivar and its synonyms, seed source, whether
the existing seed stock is original or from an increase, whether it is
open-pollinated or hybrid and cultivar-group affiliation. This list
can be obtained in printed or electronic form from the author.
Small seed samples are available to all interested in the genetics,
breeding and history and development of C. pepo, on formal
written request.
References
Curtis, L.C. 1940. Heterosis in summer squash (Cucurbita pepo)
and the possibilities of producing F1 hybrid seed for commercial planting. Proc. Amer. Soc. Hort. Sci. 37(1939):827-828.
Decker, D.S. 1988. Origin(s), evolution and systematics of
Cucurbita pepo (Cucurbitaceae). Econ. Bot. 42:4-15.
Duchesne, A.N. 1786. Essai sur l’histoire naturelle des courges.
Panckoucke, Paris, France.
Katzir, N., Y. Tadmor, G. Tzuri, E. Leshzeshen, N. Mozes-Daube,
Y. Danin-Poleg and H.S. Paris. 2000. Further ISSR and preliminary SSR analysis of relationships among accessions of
Cucurbita pepo. in: Proceedings of Cucurbitaceae 2000: the 7th
Eucarpia Meeting on Cucurbit Genetics and Breeding (N.
Katzir and H.S. Paris, eds.). Acta Hort. 510:433-439.
Naudin, C. 1856. Nouvelles recherches sur les caractères
spécifiques et les variétés des plantes du genre Cucurbita.
Ann. Sci. Nat. Bot. IV(6):5-73.
Paris, H.S. 1986. A proposed subspecific classification for
Cucurbita pepo. Phytologia 61:133-138.
Paris, H.S. 1996. Summer squash: history, diversity and distribution. HortTechnol. 6:6-13.
Smith, B.D. 1997. The initial domestication of Cucurbita pepo in
the Americas 10,000 years ago. Science 276:932-934.
Trehane, P., C.D. Brickell, B.R. Baum, W.L.A. Hetterscheid, A.C.
Leslie, J. McNeill, S.A. Spongberg and F. Vrugtman. 1995.
International code of nomenclature for cultivated plants.
Quarterjack, Wimborne, UK.
Whitaker, T.W. 1947. American origin of the cultivated cucurbits.
Ann. Missouri Bot. Gard. 34:101-111.
46
Plant Genetic Resources Newsletter, 2001, No. Plant
126 Genetic Resources Newsletter, 2000, No. 123: 68 -77
Book Reviews
Biotechnology in Agriculture Series, No. 24.
The Biotechnology Revolution in Global Agriculture:
Invention, Innovation and Investment in the Canola Sector
P.W.B. Phillips and G.G. Khachatourians editors. 2001. CABI Publishing. ISBN 0-851-99513-6
At one level, the book is the story of the development of the Canola
industry in Canada, and I think that a brief synopsis is in order.
Brassica rapa was introduced into Canada in 1927 by an immigrant farmer from Poland. Brassica napus was introduced from the
Argentine via the USA in 1942 as a government response to the
Second World War blockade of European and Asian sources of
rapeseed oil, which was used as a lubricant in marine engines. The
spring forms of these species proved suited to western Canada and
provided farmers with a new crop, with the faster growing and earlier
maturing B. rapa becoming established in northern regions and B.
napus in the longer growing seasons further south.
Public sector plant breeding began in 1944 and was followed in
the 1950s by research into oil quality, which culminated in 1963 in
the discovery of low erucic acid seed and opened the way to
rapeseed oil for human consumption. The development and use of
gas-liquid chromatography for analysing small quantities of seed
had proved vital, and so did the half seed technique for breeding,
whereby half a seed could be analysed for oil quality and the other
half germinated and grown into a fertile plant. The first Canadianbred low erucic acid (single low) cultivars were Oro (B. napus) and
Span (B. rapa), released in 1968 and 1971, respectively. Meal
quality for animal feed was then vastly improved by selecting
seeds with low glucosinolate content, with the first double low B.
napus cultivar, Tower, released in 1974. By 1978, double low B.
rapa was also readily available and the ‘meaningless’ name Canola
(Can=Canada) was accepted as the Trademark for such cultivars
and their seed.
The establishment of Plant Breeders’ Rights in Canada in 1990
encouraged private sector breeding; by 1997 the private sector
share of new varieties had increased to 74%. By 1999 F1 hybrid
cultivars (B. napus) accounted for 30% of the acreage. However, a
much larger private sector investment had already taken place in the
1980s as new technologies and patents allowed chemical companies such as Monsanto and AgrEvo to develop new transgenic (GM)
cultivars tolerant to their patented herbicides. Rapid adoption followed the release of the first such cultivar in 1994 so that by 1999
over 70% of the Canadian acreage was herbicide tolerant. Recent
transgenic field trials indicate that the first generation of transgenic
cultivars with changed agronomic attributes (herbicide, stress, insect, virus and fungal resistance) will be followed by even more
profitable second generation ones with altered end-use attributes
(modified oil composition, changed nutritional balance, ability to produce nutra or pharmaceutical products).
The purpose of the book is to analyze this story of Canola
development in Canada as an example of how the agri-food sector is
being transformed worldwide into an innovation-driven, predominantly privately managed and vertically coordinated life science
business that exports differentiated high-value products and in which
knowledge has replaced capital, land and labour as the new basic
economic resource (the means of
production). The eight contributors
to the book are from Agricultural
Economics, Commerce, and Applied Microbiology and Food Sciences in the University of
Saskatchewan, Canada. The 17
chapters are grouped into six parts:
The Setting, Innovation and Canola, The Actors, Regulating Biotechnology-based Growth, Winners and Losers, and Policy Implications.
The book is a rich mixture of ideas, data and statistics, new
economic theories, and analyses and interpretation, and makes a
thought-provoking read. Each reader will no doubt find different
parts of greatest interest and value and I am being selective in my
highlights. I was particularly interested in the analysis of the changing roles of public-sector institutions, private firms and government, and their increasingly complex interactions. For example, on
the one hand, Government wants to promote wealth creation so
that its citizens can gain from the economic benefits flowing from
the new technologies; in pursuing this objective it has introduced
new intellectual property rights and supportive domestic and international trade rules and has engaged as a partner and promoter of
research activity. On the other hand, Government also wants to
act as regulator and custodian for consumer concerns over health
and safety and environmental and biodiversity issues arising from
the new technology, and also wants to ensure an equitable distribution of the benefits. Indeed, I was also fascinated by the analysis
of who gains from research and the distribution of its benefits
between biotechnologist, breeder, farmer, processor and consumer. Furthermore, the distribution is likely to shift as a new
generation of transgenic cultivars changes the value of products to
end consumers in contrast to lowering the cost of production. In
this context, the authors conclude that North American regulations
are more conducive than European ones to the development and
commercialisation of agricultural biotechnology because the former
are based on the end product, whereas the latter tend to apply the
precautionary principle to the technology.
Finally, and perhaps not surprisingly, on the structure and
geographic location of the industry, the authors conclude that
Saskatchewan in Canada has found a niche in the research and
production chain, assembling the basic scientific knowledge and
proprietary technologies, and using them to breed novel traits into
Canola cultivars which are then planted, grown and crushed to a
first stage of processing, before being exported to global markets.
In conclusion, I found the book a fascinating and stimulating
read, which I can thoroughly recommend.
John Bradshaw
SCRI, Dundee, Scotland
Monitoring and Surveillance of Genetically Modified Higher Plants: Guidelines for
Procedures and Analysis of Environmental Effects
G. Kjellson and M. Strandberg. 20001. Birkhaurser. ISBN: 3-764-36227-8
The book is a compendium of procedures that may address the
environmental risks and public concerns over large-scale cultivation of genetically modified higher plants (GMHPs). It is readable and practical and offers a concise yet wide presentation of
the technical issues surrounding the commercial application of
GMHPs. It presents a helpful specification of the range of agronomic and ecological dimensions of understanding the environmental impacts of GMHPs.
We agree with the book’s underlying assumption that the widescale introduction of GMHPs has risks: achieving superior crop
qualities (e.g. stress tolerance, better growth characteristics, enhanced resistance to pests) through genetic modification, has the
potential of altering natural ecosystems, or even of being toxic to
non-target organisms, and hence would pose risks to humans and
biota. The possibility is not remote, nor has it been sufficiently
discounted, that multiple gene inserts that could create ‘super
variants’ might out-compete less endowed endemic species and
gene transfers may create unwanted effects on a chain of organisms in the intricate web of life across ecosystems. Yet, as the
book also assumes, while the risks are real, so is the need to
improve the world’s ability to produce foods, under, we note, a
variety of ecological conditions that have been themselves much
altered, more extensively in the past 100 years, that they pose
unnaturally high risks to crops (e.g. higher virulence of diseases,
altered cycles of drought and rainfall, lower soil fertility, disturbed
balance of soil biota, and altered behaviour of pests and pest
populations). We agree that, as the book suggests, developing
robust and rigorous assessment, monitoring, and surveillance
systems will allow for more effective control of the risks associated
with GMHPs.
However, the monitoring and surveillance (MS) procedure the
book describes is restricted to the biophysical impacts of commercial-scale applications of GMHPs in environments that are mainly
temperate to cold (i.e. NW Europe). If it were to be useful in more
tropical countries like the Philippines, or where (a) environmental
concerns heavily encompass social, political and cultural dimensions of ecological changes, (b) biodiversity is high enough that
even field trials are heavily debated activities, and (c) the environment is much more complex and fragile, we anticipate that a more
comprehensive MS procedure would have to include assessing
(1) social impacts (effects of wide-scale GMHP applications on
existing patterns of tenure and of income opportunities of farmers
and other farming-related income earners—who wins, who loses,
particularly where farming opportunities are thin and tenuous?);
and (2) the risks of small-scale applications.
The book does include discussion of environmental effects
when GM plants are cultivated but it could perhaps also address
the variation of biological features across ecological systems like,
for example, variations of evolutionary rates in temperate, arctic
and tropical environments, and varying hybridization potentials of
GMHPs across landscapes with increasingly higher diversity and
closer genetic affinity of endemic weed species.
Regarding who will carry the risk assessment and monitoring
ecosystem effects, the book suggests formal institutions or standing regulatory agencies, but in our view these would not serve well
in countries such as the Philippines where core political and economic institutions (particularly those established with colonial roots)
are still to acquire full legitimacy and widespread social acceptance
and credibility. Ad hoc committees of scientists, members of civil
society and industry, and credible personalities from among the
general public would be more workable alternatives.
In summary, we believe that the book should have been expanded to cover assessment, monitoring and surveillance of GMHP
applications in different scales, and across different ecological,
agronomic, agrarian and ‘eco-political’ conditions of the world.
Ben S. Malayang III and Pacifico C. Payawal
University of the Philippines Los Baños, Philippines
48
First Announcement
PROTA — Plant Resources of Tropical Africa
Ressources Végétales de l’Afrique Tropicale
FIRST INTERNATIONAL WORKSHOP
PREMIER ATELIER INTERNATIONAL
23–25 September 2002
Nairobi, Kenya
PROTA will be the subject of an International Workshop to review the progress made, and to reach international consensus on
the structure, organization, activities, and finances of the First Implementation Phase 2003–2007.
The workshop is a forum for scientists, policy-makers and donors, in order:
●
to highlight the importance of the Plant Resources of Tropical Africa through Commodity Group Reports, Country
Reports and Plant Resources Reports;
●
to review the progress made in the Phase 2000–2002 towards international cooperation, the documentation and
information system, and the publication of the monographs;
●
to make the recommendations for the Implementation Phase 2003–2012 on all aspects of the programme including
organization, manpower, finances, publication policy and the databank.
PROGRAMME OUTLINE
The Workshop will be bilingual (English and French).
Section 1: General
Invited papers on the many facets of the PROTA programme; its relation with new trends in agricultural, silvicultural and
environmental policies and new developments in information technology.
Section 2: Commodity Group Reports
Invited papers on general aspects of a number of Commodity Groups, like ‘Cereals and Pulses’, ‘Vegetables’, ‘Timbers’, ‘Auxiliary
plants’ and ‘Medicinal plants’.
Section 3: Country Reports
Invited papers on the plant resources of the various parts of Tropical Africa, based on the documentation work of the PROTA
Regional and Country offices.
Section 4: Plant Resources Reports
Contributed papers (Posters) on subgroups of plant resources, in particular treatment of the ecology, agronomy/management,
uses and improvement of neglected and potentially important species.
Section 5: Phase 2003–2012
Working Groups and Plenary discussions on organization, financial aspects, publication policy and databank. Formulation of
recommendations.
Excursion
Half-day field-trip to interesting projects on plant resources in the surroundings of Nairobi.
PREREGISTRATION
Please contact:
Or for more information:
Secretariat
PROTA FIRST INTERNATIONAL WORKSHOP
c/o ICRAF
PO Box 30677
Nairobi
Kenya
PROTA PROGRAMME
Wageningen University
PO Box 341
6700 AH Wageningen
The Netherlands
email: [email protected]
Web: www.prota.org
Plant Genetic Resources
Newsletter
Bulletin des ressources
phytogénétiques
Boletín de Recursos
Fitogenéticos
Aims and scope
Domaine d’intérêt
Objetivos y temas
The Plant Genetic Resources Newsletter publishes papers in English, French or Spanish, dealing
with the genetic resources of useful plants, resulting from new work, historical study, review and
criticism in genetic diversity, ethnobotanical and
ecogeographical surveying, herbarium studies, collecting, characterization and evaluation, documentation, conservation, and genebank practice.
Le Bulletin des ressources phytogénétiques publie des articles en anglais, en espagnol et en
français, sur les ressources génétiques de plantes utiles, fruit de nouvelles recherches, d’études
historiques, d’examens et de critiques concernant la diversité génétique, d’études ethnobotaniques et écogéographiques, d’études d’herbiers,
d’activités de collecte, de caractérisation et
d’évaluation, de documentation, de conservation
et les pratiques des banques de gènes.
El Noticiario de Recursos Fitogenéticos publica
documentos en inglés, francés y español que
tratan de los recursos genéticos de plantas útiles,
fruto de nuevos trabajos, estudios históricos,
revisiones y análisis críticos relacionados con la
diversidad genética, investigaciones etnobotánicas y ecogeográficas, estudios de herbarios,
actividades de colección, caracterización y evaluación, documentación, conservación, y prácticas en bancos de germoplasma.
Parrainage
Dirección
Le Bulletin des ressources phytogénétiques est
publié sous les auspices de l’Institut international
des ressources phytogénétiques (IPGRI) et de la
Division de la production végétale et de la protection des plantes de l’Organisation des Nations
Unies pour l’alimentation et l’agriculture (FAO)
El Noticiario de Recursos Fitogenéticos se publica bajo los auspicios conjuntos del Instituto Internacional de Recursos Fitogenéticos y la Dirección de Producción y Protección Vegetal de la
Organización de las Naciones Unidas para la
Agricultura y la Alimentación.
Distribution
Distribución
Le Bulletin des ressources phytogénétiques paraît
une fois par an en un volume regroupant quatre
numéros publiés en mars, juin, septembre et
décembre. Il est distribué gratuitement aux bibliothèques des banques de gènes, universités,
services gouvernementaux, instituts de recherche, etc. s’intéressant aux ressources phytogénétiques. Il est aussi envoyé sur demande à tous
ceux pouvant démontrer qu’ils ont besoin d’un
exemplaire personnel de cette publication.
El Noticiario de Recursos Fitogenéticos aparece
como un volumen anual compuesto por cuatro
números, que se publican en marzo, junio, septiembre y diciembre. Se distribuye gratuitamente a
las bibliotecas de bancos de germoplasma, facultades universitarias y servicios gubernamentales,
centros de investigación, etc. que se interesan
en los recursos fitogenéticos. También pueden
obtener este noticiario las personas que demuestren necesitar una copia personal.
Types de documents publiés
Tipos de documentos
Articles
Artículos
Un article contient les résultats de travaux nouveaux et originaux qui apportent une contribution
importante à la connaissance du sujet dont traite
l’article. Les articles, qui doivent être d’une
longueur raisonnable, sont d’abord examinés par
le Comité de rédaction qui en évalue la portée et
la validité, puis par un expert qui en examine le
contenu et l’intérêt scientifiques.
Los artículos divulgarán los resultados de trabajos nuevos y originales que contribuyan de modo
importante al conocimiento del tema tratado.
Dichos artículos, que deberán tener una longitud
razonable, serán examinados por el Comité de
Redacción en cuanto a su pertinencia e idoneidad
y posteriormente un experto juzgará su contenido
y validez científicos.
Brèves communications
Comunicaciones breves
On entend par brève communication un texte
contenant, sous une forme abrégée, les résultats
de travaux présentant un intêrêt pour tous ceux
qui s’occupent de ressources phytogénétiques.
Elle contient en particulier des comptes rendus
des missions d’acquisition de matériel génétique.
Las comunicaciones breves informarán de modo
conciso sobre los resultados de trabajos de interés para las personas que se ocupan de los
recursos fitogenéticos. Las comunicaciones
breves incluirán, en particular, resúmenes sobre
las misiones de adquisición de germoplasma.
Autres documents
Otros documentos
Le Bulletin des ressources phytogénétiques publie d’autres types de rapport tels que des documents de synthèse, des études critiques et des
articles commentant des problèmes actuels concernant les ressources phytogénétiques. Le Bulletin publie une revue de livres ainsi qu’une section intitulée Nouvelles et Notes. Les auteurs
sont invités à envoyer leurs suggestions pour les
livres à passer en revue ainsi que des contributions aux Nouvelles et Notes.
El Noticiario de Recursos Fitogenéticos publicará otros tipos de informes, como documentos
de trabajo, análisis críticos, y documentos que
examinen cuestiones de actualidad relacionadas
con los recursos fitogenéticos. El Noticiario publicará una reseña de libros así como una sección
de Noticias y Notas. Las propuestas de libros
para reseñar y las contribuciones a la sección de
Noticias y Notas serán bien acogidas.
Management
The Plant Genetic Resources Newsletter is published under the joint auspices of the International Plant Genetic Resources Institute (IPGRI) and
the Plant Production and Protection Division of
the Food and Agriculture Organization of the
United Nations (FAO).
Availability
The Plant Genetic Resources Newsletter appears as one volume per year, made up of four
issues, published in March, June, September and
December. Plant Genetic Resources Newsletter
is available free of charge to interested libraries
of genebanks, university and government departments, research institutions, etc. The periodical
may also be made available to individuals who
can show that they have a need for a personal
copy of the publication.
Types of paper
Articles
An article will publish the results of new and
original work that makes a significant contribution to the knowledge of the subject area that the
article deals with. Articles, which should be of a
reasonable length, will be considered by the Editorial Committee for scope and suitability, then
assessed by an expert referee for scientific content and validity.
Short communications
A short communication will report results, in an
abbreviated form, of work of interest to the plant
genetic resources community. Short communications in particular will contain accounts of germplasm acquisition missions. The papers will be
assessed by an expert referee for scientific content and validity.
Other papers
The Plant Genetic Resources Newsletter will
publish other forms of reports such as discussion
papers, critical reviews, and papers discussing
current issues within plant genetic resources.
Book reviews will be printed, as well as a News
and Notes section. Suggestions for books to
review are invited, as are contributions to News
and Notes.
Submission
In the first instance papers may be submitted in
typescript form or as an Email message. The
final version may be submitted as an Email file or
as a Windows-readable file on diskette. Manuscripts submitted for publication and other communications on editorial matters should be addressed to IPGRI's Editorial and Publications
Unit.
Presentación
Présentation
En premier lieu, les documents doivent être soumis dactylographiés ou par courrier électronique.
La version définitive doit être présentée en fichier
de courrier électronique ou sur disquettes compatibles Windows. Prière d’adresser les manuscrits
présentés pour être publiés et d’autres communications sur des questions de rédaction au Bureau
de rédaction de l'IPGRI.
Los documentos deben entregarse, incialmente,
en forma de texto mecanografiado o a través del
correo electrónico. La versión final debe presentarse como un archivo de correo electrónico o en
disquete compatible con el sistema operativo
Windows. Los manuscritos para publicar y otras
comunicaciones sobre asuntos relativos a la redacción deberán dirigirse a la Oficina de Redacción del IPGRI.
No. 126, June 2001
Contents
Articles
Classification of Italian maize (Zea mays L.) germplasm
A. Brandolini and A. Brandolini (Italy) .................................................................................................................. 1
Collecting landscape trees and shrubs in Ukraine for the evaluation of aesthetic quality and
adaptation in the north central United States
M.P. Widrlechner, R.E. Schutzki, V.Y. Yukhnovsky (USA) and V.V. Sviatetsky (Ukraine) .............................. 12
Colecta de germoplasma en la ecoregión de la Península de Paria, Estado Sucre, Venezuela
E. Mazzani y V. Segovia (Venezuela) ................................................................................................................ 17
Plant exploration in the Talysch Mountains of Azerbaijan and Iran
L. Frese (Germany), Z. Akbarov (Azerbaijan), V.I. Burenin (Russia), M.N. Arjmand (Iran)
and V. Hajiyev (Azerbaijan) ................................................................................................................................ 21
Evaluation of variability in natural populations of peperina (Minthostachys mollis (Kunth.) Griseb.),
an aromatic species from Argentina
O. Marta, R. Coirini, J. Cosiansi, R. Zapata y J. Zygadlo (Argentina) ............................................................... 27
Morphological and isoenzyme variability of taro (Colocasia esculenta L. Schott) germplasm in Cuba
A. Rodríguez Manzano, A.A.Rodríguez Nodals, M.I. Román Gutiérrez, Z. Fundora Mayor and
L. Castiñeiras Alfonso (Cuba) ............................................................................................................................. 31
Characterization of the Cucurbita pepo collection at the Newe Ya'ar Research Center, Israel
H.S. Paris (Israel) ............................................................................................................................................... 41
Book Reviews .................................................................................................................................................... 46
First Announcement...................................................................................................................................48

Plant genetic resources newsletter No. 126, June 2001

Transcripción

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