Diversity of common bean (Phaseolus vulgaris L.) germplasm from Portugal

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1 Genetic Resources and Crop Evolution 48: , Kluwer Academic Publishers. Printed in the Netherlands. Diversity of common bean (Phaseolus vulgaris L.) germplasm from Portugal 1, A.P. Rodino *, M. Santalla, I. Montero, P.A. Casquero and A.M. De Ron 1 Legumes Breeding Group, Mision Biologica de Galicia (MBG)-Spanish Council for Scientific Research 2 (CSIC), University of Santiago de Compostela (USC), P.O. Box 28, Pontevedra, Spain; Current address: ESTIA, University of Leon, Ponferrada, Spain; * Author for correspondence Received 27 March 2000; accepted in revised form 23 February 2001 Key words: Bean, Breeding, Germplasm, Landraces, Phaseolus vulgaris, Variation Abstract Common bean (Phaseolus vulgaris L.) is a traditional crop in Portugal, where farmers grow varieties selected and maintained by themselves. A collection of 88 landraces of common bean was evaluated for 17 quantitative and qualitative traits and the biochemical marker phaseolin to display the degree of variation of this germplasm. Agronomic data were subjected to cluster analysis and several groups were identified, with three groups clustering most of the landraces. Regarding phaseolin variation the C and T banding patterns are the most frequent ones, so the origin of the Portuguese beans is thus probably the Andean region of South America. These results give information about the origin, diversity and breeding value of the Portuguese germplasm, that could be useful to widen the genetic base of currently cultivated bean varieties in Europe. South American Andes since the Mesoamerican varieties are not currently very popular in Europe. Gepts and Bliss (1988) suggested that the bean grown on the Iberian Peninsula was originated in a different point from those of the rest of Europe. Nevertheless, the introduction of beans in Europe is unclear and cur- rently under discussion. There is evidence that com- mon bean reached France in 1508, probably as an ornamental plant without value for human consump- tion in that time (Zeven 1997). Sailors and traders, 500 years ago, could have brought beans as a curiosi- ty, only for fun, as children use to do currently at home and school in the Andean region. The sensitivi- ty to photoperiod and low temperature during the growing season could have been a limiting factor for cultivated bean in many European latitudes in the early times. In fact it is possible to grow primitive Andean landraces and wild populations in the North of Spain (Pontevedra, 428N) but only under greenhouse conditions during the fall-winter-spring period (De Ron et al. 1999). Nevertheless, Zeven (1997) has found some descriptions of common bean as early as Introduction Common bean is a species of American origin whose two principal areas of domestication are South America and Middle America (Gepts et al. 1986). The cultivated gene pools of common bean can be distinguished by their morphology (Singh et al. 1991a), phaseolin seed protein electrophoretic type (Koenig et al. 1990) and isozymes (Singh et al. 1991b). Varieties from Middle America usually have small seeds and through electrophoresis produce phaseolin (S and B types) patterns different from those of their South American counterparts (T, C, H and A phaseolin types) (Gepts et al. 1986; Singh et al. 1991b). Three races and six different gene pools (Singh et al. 1991c) have been proposed both for common bean from Middle America (Mesoamerica, Durango and Jalisco races) and from the Andean region in South America (Chile, Peru and Nueva Granada races). Concerning the origin of the European beans, Mc Clean et al. (1993) suggested that the germplasm dispersed to Europe was probably domesticated in the

2 which indicate, in fact, the wide distribution of and Beira Litoral, yielding together up to 80% of the this species in Europe, starting obviously in 1492 by Portuguese national bean production (Anonymous means of its introduction in Spain by C. Columbus. 1993). The production has been decreasing from Feijao is the Portuguese name for common bean t in 1980 to t in 1992, but the average that reminds the Latin name of faseolum or yield has increased from 368 kg/ ha to 554 kg/ ha phaseolum as well as the Greek one phaselos, and (Anonymous 1993). This increase could have been it is similar to the popular name frijol in many due to the introduction of improved cultivars, which countries of Latin America. However, the most popu- would result in a reduction of the genetic variability lar Spanish name is judıa, which reminds the jews within the species (Carnide et al. 1997a; 1997b), and (or judıos in Spanish) as usual bean consumers. the increase of the genetic vulnerability (Esquinas- Hence, the knowledge of the local names does not Alcazar 1983). contribute to clarify the problem of the bean intro- This genetic diversity maintained in the Portuguese duction and evolution in Southern Europe. The origin unimproved bean landraces and garden forms could of Portuguese bean germplasm is unclear because include appropiate characteristics for the improvethere are no evidences of a clear exchange of seeds ment of European current cultivars. Since Europe is with the Spanish surroundings in spite of the strong quite far from being self-sufficient in bean production links between both countries according to their his- (as in grain legumes in general), contributions to the tory, culture and language. In the 15th and 16th genetic improvement of cultivars deserve special centuries there were some harbours maintaining ac- attention. However, in spite of its relevance, this tive commerce with the New World in the Northwest genetic source of common bean variation has been of Spain (Galicia). The introduction of some crops scarcely studied and only a reduced morphoagrosuch as bean and maize and the distribution to other nomic and isoenzymatic characterization exists for a areas could have ocurred in this area. The traditional small number of landraces (Carnide et al. 1997a; cropping systems for the bean crop similar to those 1997b; Casquero et al. 1997; Matos et al. 1998). used in many areas of America (Santalla et al. 1994) These are the reasons why the Legumes Breeding are strong arguments to support this hypothesis. Addi- Group has performed some field trials for Portuguese tionally, there are evidences of seed exchange among germplasm. These Portuguese landraces were cofarmers and gardeners in many countries of Europe llected in the North of Portugal in 1988 and 1993 (Zeven 1999) for testing some new material or for jointly with the University of Tras-os-Montes e Alto avoiding the degeneration of varieties cultivated year Douro (UTAD, Vila Real, Portugal) and they are after year. currently maintained at the germplasm collection of Agriculture in Portugal is still traditional and far- UTAD and BPGV (Portuguese Bank of Vegetable mers cultivate their own varieties, commonly associ- Germplasm, Braga, Portugal) and the MBG-CSIC ated with other crops such as the maize, obtained and (De Ron et al. 1997) (Table 1). Additionally, some maintained by themselves after many years of mass Portuguese bean landraces maintained at the Spanish selection. Hence, this genetic material must be consid- Centre for Plant Genetic Resources (CRF) were transered as landraces or garden forms (Zeven 1997). For ferred to the MBG-CSIC in order to be also evaluated. this reason, a high degree of diversity is still main- A morphological and agronomical characterization tained within this species, and it is therefore possible and the biochemical marker phaseolin were used in to collect valuable germplasm in this region. this work to display the variation of common bean 76% of the bean production is located in the North germplasm from Portugal. The objectives of the preof Portugal, in the regions Entre Douro and Minho sent research are: 1) to display the diversity of 88 Table 1. Portuguese germplasm collections. Germplasm Banks Country Accessions number University of Tras-os-Montes e Alto Douro (UTAD) Vila Real, Portugal 22 Portuguese Bank of Vegetable Germplasm (BPGV) Braga, Portugal 1125 Spanish Centre for Plant Genetic Resources (CRF) Madrid, Spain 293 Germplasm Bank of the Agronomic and Genetic National Station Oeiras, Portugal 559 Mision Biologica de Galicia (MBG-CSIC) Pontevedra, Spain 121 International Centre of Tropical Agriculture (CIAT) Cali, Colombia 1001

3 bean accessions from Portugal, which are a sample of used for the identification of the phaseolin patterns germplasm from several areas of this region, 2) to (Figure 2). clarify the origin of the Portuguese bean germplasm The characters previously mentioned, except for and 3) to assess about the role of this germplasm in the growth habit and phaseolin type, were used to the improvement of European current bean cultivars. perform the numerical classification by means of a cluster analysis (Sneath and Sokal 1973; Rohlf 1993). 411 Material and methods Results and discussion A collection of 88 common bean accessions from The study of the dendrogram and the previous know- Portugal which came from the collections of MBG- ledge of this genetic material permitted to classify the CSIC, CRF and UTAD was evaluated (Table 2). The 88 populations studied into 13 groups in broad sense map of Figure 1 shows the geographical origin of but in fact some of them could be considered as these populations. The material consists of landraces subgroups inside the major groups IV, VI and VII or native populations maintained by farmers for ge- (Figure 3) which clustered most the landraces. The nerations, and representative of the different bean mean values and the standard error of the variables types growing in the North of Portugal. used for each of the groups and subgroups are dis- The common bean landraces were grown in field played in Table 4. trials carried out during the Summer season and from The group IV was divided into four subgroups, the 1987 to 1998 in different environments (Table 3). The group VI into two subgroups and the group VII into map of Figure 1 shows the location of the different three subgroups. The groups I, II, III and V, include 13 trials. landraces and do not present differential characteris- The experimental design used from 1987 to 1995 tics. These groups are a misinterpretation and contriwas a randomized complete block design with two bute little information from the viewpoint of studied replications. Each cultivar was planted in a single 15 germplasm. plants-row plot with row to row distance of 80 cm and Five groups and subgroups could be considered as plant to plant distance of 25 cm equivalent to a germplasm for dry seed consumption, the groups III, planting density of plants/ ha. In 1998 the V and VI and the subgroups IV.1 and VII.1 which experimental design used was a hill- plot with 8 include the typical dry bean cultivars usually grown plants-plot spaced m between plots and a and consumed in the Northwest of Spain such as planting density of plants/ ha. Canela, Pinto, Alubia and Garbanzo. These The following 17 quantitative characters were de- groups have phaseolin types T, C and H of Andean terminated (IBPGR 1982; Santalla et al. 1994): begin- origin but group III includes landraces which have ning of flowering, end of flowering, period of flower- small seeds with the phaseolin types S and B of ing, pod length, number of pods per plant, seed Mesoamerican origin and group V presents a mixture dimensions (length, width and thickness), 100 seed of seeds of Mesoamerican and Andean origin. The weight (g), water absorption (%), coat proportion group II and the subgroup IV.2 include germplasm for (%), protein content (%), humidity content (%), fat fresh pod consumption, with T or C phaseolin types content (%), starch content (%) and total sugar con- and Pinto seeds. Finally, the group I and the subtent (%) (by using Near Infrared Transmission spec- groups IV.3, IV.4, VII.2 and VII.3 represent germtroscopy method). The growth habit was also evalu- plasm for dual purposes which include the cultivars as ated according to CIAT (Hidalgo 1991). Canela, Pinto and White kidney. The groups I The phaseolin pattern was analysed from a sample and IV.4 present T or C phaseolin type which indicate of the cotyledons of five seeds of each bean landrace an Andean origin, the groups IV.3 have Pinto and by using one-dimensional sodium dodecyl sulphate small seeds of Mesoamerican origin and the groups polyacrylamide gel electrophoresis (SDS-PAGE), ac- VII.2 and VII.3 have a mixture of seeds of cording to the method described by Ma and Bliss Mesoamerican and Andean origin. This information (1978), Brown et al. (1981), Gepts et al. (1986). The would permit to choose some populations to be infollowing cultivars: Boyaca 22 (B), Contender (C), cluded in different breeding programmes. Pampa (H), Sanilac (S) and Tendergreen (T) were From the view point of the breeder, it must be

4 412 Table 2. Phaseolin seed type and growth habit of the studied landraces. Landrace n Phaseolin type Colour Seed type and local name Growth habit PHA-0035 C white large, kidney, I PHA-0036 T white medium, round, Garbanzo II PHA-0037 S/ T white large, kidney, Great Northern I PHA-0038 C dark brown large, oval, III PHA-0039 T/ C pinto (light and dark brown) large, oval, Pinto I PHA-0040 T pinto (light brown and red) medium, oval, Cranberry III PHA-0041 H yellow medium, round II PHA-0042 T cream large, oval I PHA-0043 C white medium, oval, Garbanzo I PHA-0044 T light brown large, kidney I PHA-0047 S white medium, round, Garbanzo II PHA-0049 T light brown large, oval II PHA-0050 C cream medium, oval II PHA-0053 T white large, kidney, I PHA-0054 C pinto (light and dark brown) medium, oval, Pinto IV PHA-0055 S white medium, oval, Garbanzo III PHA-0056 T cream large, kidney I PHA-0057 S pinto (light and dark brown) medium, oval, Pinto II PHA-0058 T dark red medium, kidney, Dark red kidney II PHA-0059 S/ C white medium, oval IV PHA-0061 T/ C cream large, oval IV PHA-0062 T/ H white large, oval, III PHA-0063 C pinto (light brown and red) large, kidney, Cranberry IV PHA-0064 C pinto (light and dark brown) large, round, Pinto I PHA-0065 B white medium, round, Garbanzo II PHA-0068 C dark brown large, round III PHA-0070 C pinto (light and dark brown) large, oval, Pinto IV PHA-0071 B white medium, kidney IV PHA-0072 C pinto (light brown and red) large, kidney, Cranberry IV PHA-0073 C dark red large, oval I PHA-0075 S pinto (light and dark brown) medium, oval, Pinto II PHA-0076 T cream large, oval I PHA-0077 S pinto (light and dark brown) medium, oval, Pinto II PHA-0080 C white large, kidney, I PHA-0081 C pinto (light brown and red) medium, oval, Cranberry III PHA-0082 C brown large, kidney III PHA-0084 C pinto (light and dark brown) medium, round, Pinto I PHA-0085 B pinto (light and dark brown) medium, oval, Pinto II PHA-0086 C white medium, oval I PHA-0087 T/ H white medium, oval, Garbanzo I PHA-0088 T cream large, kidney I PHA-0089 T pinto (light and dark brown) large, round, Pinto I PHA-0091 T dark brown large, kidney IV PHA-0092 T dark brown medium, kidney IV PHA-0095 T cream medium, oval I PHA-0097 C pinto (light and dark brown) large, oval, Pinto IV PHA-0098 B white medium, oval III PHA-0100 C pinto (light brown and red) large, oval, Cranberry IV PHA-0101 C/ T pinto (light brown and red) large, oval, Cranberry II PHA-0102 C/ T cream large, kidney II PHA-0103 S white medium, oval IV PHA-0104 C pinto (light brown and red) large, oval, Cranberry II PHA-0105 T brown large, oval II PHA-0106 S white medium, oval III

5 413 Table 2. (continued) Landrace n Phaseolin type Colour Seed type and local name Growth habit PHA-0107 B white small, oval IV PHA-0108 T cream large, kidney I PHA-0298 C pinto (light and dark brown) large, round, Pinto I PHA-0459 T/ C white medium, kidney I PHA-0460 C brown medium, round IV PHA-0462 T white small, oval I PHA-0463 T brown large, kidney II PHA-0464 C cream medium, oval II PHA-0465 C brown medium, oval IV PHA-0648 C pinto (light brown and red) large, oval, Cranberry IV PHA-0649 T dark red medium, kidney, type dark red kidney I PHA-0650 C brown medium, kidney I PHA-0651 T/ C cream medium, oval I PHA-0652 T/ C pinto (light brown and red) large, oval, Cranberry I PHA-0653 C white medium, kidney I PHA-0654 T pinto (red and white) large, kidney I PHA-0655 T/ C brown medium, kidney I PHA-0656 T/ C cream large, oval I PHA-0657 T pinto (light brown and red) large, oval, Cranberry II PHA-0658 T/ C pinto and brown large, kidney, Cranberry IV PHA-0660 T white small, oval I PHA-0662 C red large, oval I PHA-0663 B brown small, kidney III PHA-0666 H white medium, kidney IV PHA-0668 C pinto (red and cream) and white large, oval IV PHA-0672 S white large, kidney II PHA-0673 C/ T pinto and red medium, oval IV PHA-0674 C/ T black, brown, red medium, oval IV PHA-0676 S white and brown small, kidney II PHA-0677 T/ S white, red, brown and pinto small, kidney II PHA-0678 B white, red, black and brown small, kidney II PHA-0680 T brown medium, oval I PHA-0681 S black small, oval I PHA-0682 C/ B white medium, kidney II PHA: accession code. Type Garbanzo : medium or small, round or oval and white seeds. Growth habit: I (determinate upright), II (indeterminate upright bush), III (indeterminate prostrate), IV (indeterminate strong climbers) The C banding type was present in 40.6% of the material, T banding pattern was present in 35.5% and S and B banding patterns were present in 12.3% and 7.5% of the material, respectively, while the H band- ing type was present in only 3.8% of the landraces. These results confirm that most of the bean germ- plasm of Portugal came from South America. Gepts and Bliss (1988) also analysed 112 accessions from the Iberian Peninsula and showed that beans grown in this area were characterized by a high frequency of C phaseolin pattern, contrary to other regions or countries investigated, which suggests that the bean land- races from Iberian Peninsula were originated in a different point from those of the rest of Europe. Lioi (1989) evaluated the variation of the phaseolin pat- terns of 372 accessions from the Mediterranean area pointed out that these old Portuguese landraces after many years of cultivation are highly adapted to specific environmental conditions and they could be a diverse genetic source to widen the genetic base of the common bean crop in Europe. The process of adaptation of the old bean varieties to different European areas has probably produced changes in plant architecture, including the adaptation to different cropping systems as the change from intercropping with maize in America and Northwestern Iberian Peninsula to sole cropping in central Europe (Vanderberg and Nleya 1999). This traditional farming system makes mechanization more difficult but provides a more efficient use of environmental resources. The distribution of the electrophoretic banding types of phaseolin is reported in the Table 2 and 4.

6 414 Figure 1. Geographical origin of the studied landraces and locations where the field trials have been performed. Table 3. Field trials carried out in Spain from Trials Location Year Latitude Longitude Altitude Average temperature Average rainfall Accessions 1 Pontevedra N 88399W 40 masl 14 8C 1600 mm 56 2 Pontevedra N 88399W 40 masl 14 8C 1600 mm 36 3 Pontevedra N 88399W 40 masl 14 8C 1600 mm 27 4 Pontevedra N 88399W 40 masl 14 8C 1600 mm 26 5 Puentecaldelas N 88329W 300 masl 14 8C 1600 mm 26 6 Pontevedra N 88399W 40 masl 14 8C 1600 mm 8 7 Mabegondo N 88169W 200 masl 13 8C 970 mm 8 8 Pontevedra N 88399W 40 masl 14 8C 1600 mm 8 9 Mabegondo N 88169W 200 masl 13 8C 970 mm 8 10 Pontevedra N 88399W 40 masl 14 8C 1600 mm Lalın N 88089W 600 masl 11 8C 1100 mm Vitoria N 28389W 500 masl 11 8C 840 mm Pontevedra N 88399W 40 masl 14 8C 1600 mm 4 14 Mabegondo N 88169W 200 masl 13 8C 970 mm 4 15 Pontevedra N 88399W 40 masl 14 8C 1600 mm 4 16 Mabegondo N 88169W 200 masl 13 8C 970 mm 4 17 Mabegondo N 88169W 200 masl 13 8C 970 mm 16

7 415 Figure 2. Photograph of phaseolin patterns used in this work. B (Boyaca), C (Contender), H (Pampa), S (Sanilac) and T (Tendergreen). (Italy and Cyprus) and the results agree with the study of Gepts and Bliss (1988). Escribano et al. (1998) studied 66 accessions from the Nortwestern Iberian Peninsula and the results did not agree with those of other authors, since the T and H phaseolin patterns were more frequent than the type C. According to these results there is not an easy conclusion about the origin of the germplasm from the Iberian Peninsula although several bean introductions in Europe could be considered. Nevertheless, the abundance of the phaseolin types C and T (that represent 76.4%) indicate the Andean origin of the Portuguese germplasm. Andean types include medium and large, white and pinto seeded cultivars as well as indeterminate habits, usually to associate with maize. These are the types preferred by the Iberian growers and consumers and the reasons why Andean landraces have been more successful in the Iberian Peninsula than in other places throughout Europe. Hence, most of the European bean germplasm from Mesoamerica and the Andean region could have arrived in Europe by means of Spanish and Portuguese sailors and traders. Large seeded faba bean, which was usually cultivated by farmers in the Figure 3. Dendrogram compiled by the UPGMA method showing the grouping of the studied landraces.

8 416 Table 4. Mean of the traits and phaseolin types in the different groups. Groups Subgroups in Subgroups in Subgroups in the group IV the group VI the group VII Character I II III V SE Beginning of flowering (days) End of flowering (days) Period of flowering (days) Pod length (mm) Pod per plant Seed length (mm) Seed width (mm) Seed thickness (mm) Growth habit I I I/ II I III/ IV II II/ IV II/ III/ IV I/ II I/ II/ IV I/ II I/ III/ IV I/ III/ IV 100 seed weigth (g) Water absorption (%) Coat percentage (%) Protein content (%) Humidity content (%) Fat content (%) Starch content (%) Total sugars (%) Phaseolin types T C S, B T, C T, C S, B T, C C, B T, C T, C T, C T, C T, S T T, S S, B B H, B C SE: Standard error. Some groups can present more than one growth habit and phaseolin types. Northwest of the Iberian Peninsula was probably Portuguese germplasm and to the Centro de Recursos displaced by white large seeded bean cultivars with a Fitogeneticos (CRF, Ministry of Agriculture, Madrid, better culinary quality and probably popular name Spain) for supplying some of the studied populations. faba given in this area to common bean could support this statement. Later an asymmetric distribution of the varieties grown by farmers was produced, on based upon preferences of the different regions. References Hence, the Iberian Peninsula displays a wide diversity Anonymous Portugal Agrıcola. Instituto Nacional de Estaon bean germplasm but with an asymmetry displaced dıstica, Lisboa, Portugal, 181 pp. to the white large-seeded Andean types. Brown J.W.S., Ma Y. and Bliss F.A Genetic variation in the In conclusion, it would point out the valuable subunits of globulin-1 storage protein of French bean. Theor. information that is now available about the origin, Appl. Genet. 59: Carnide V., Guedes-Pinto H. and Quintao J. 1997a. Caracterizaçao diversity and breeding value from the underutilized morfologica de germoplasma de feijao (Phaseolus vulgaris L.) Portuguese varieties. This information would be use- de Tras-os-Montes. Actas de Horticultura 17: ful for breeders and geneticists in order to use sources Carnide V., Jaques J. and Guedes-Pinto H. 1997b. Diversidade para of Portuguese germplasm to widen the genetic base of caracterısticas morfologicas e de produçao em cultivares re- currently cultivated bean varieties in Europe. gionais de feijao do norte interior de Portugal. In: De Ron A.M (ed.), Situacion actual y perspectivas del cultivo de la judıa. Universidad de Santiago de Compostela, Spain, pp Casquero P.A., Ruız de Galarreta J.I., Santalla M. and De Ron A.M. Acknowledgements Evaluacion preliminar de variedades locales de judıa comun (Phaseolus vulgaris L.) del Norte de Portugal. Actas de The authors thank V. P. Carnide for the information on Horticultura 17: Escribano M.R., Santalla M., Casquero P.A. and De Ron A.M. bean production in Portugal. Gratitude is also extend Patterns of genetic diversity in landraces of common bean ed to the Centro Internacional de Agricultura Tropical (Phaseolus vulgaris L.) from Galicia. Plant Breeding 117: 49 (CIAT, Cali, Colombia) for its information on the 56.

9 417 Esquinas-Alcazar J.T Los recursos fitogeneticos: una inver- De Ron A.M., Santalla M., Barcala N., Rodino A.P., Casquero P.A. sion segura para el futuro. IBPGR-FAO, Roma e InIA, Madrid. and Menendez M.C Phaseolus spp. at the Mision Bio- Gepts P. and Bliss F.A Dissemination paths of common bean logica de Galicia, Spain. Plant Genetic Resources Newsletter (Phaseolus vulgaris Fabaceae) deduced from phaseolin electro- 112: 100. phoretic variability. II. Europe and Africa. Econ. Bot. 42: 86 De Ron A.M., Rodino A.P., Menendez-Sevillano M.C., Santalla 104. M., Barcala N. and Montero I Variation in wild and Gepts P., Osborn T.C., Rashka K. and Bliss F.A Phaseolin primitive Andean bean varieties under European conditions. protein variability in wild forms and landraces of the common Annual Report of the Bean Improvement Cooperative 42: bean (Phaseolus vulgaris): evidence for multiple centers of Santalla M., De Ron A.M. and Escribano M.R Effect of domestication. Econ. Bot. 40: intercropping bush bean populations with maize on agronomic Hidalgo R CIAT S world Phaseolus collection. In: traits and their implications for selection. Field Crop Research Schoonhoven A. and Voysest O. (eds), Common Bean: Research 36: for Improvement. C.A.B. International and CIAT, pp Singh S.P., Gepts P. and Debouck D.G. 1991a. Races of common IBPGR Descriptors for Phaseolus vulgaris. International bean (Phaseolus vulgaris Fabaceae). Economic Botany 45: 379 Board for Plant Genetic Resources Secretariat, Rome, Italy Koenig R., Singh S.P. and Gepts P Novel phaseolin types in Singh S.P., Nodari R. and Gepts P. 1991b. Genetic diversity in wild and cultivated common bean (Phaseolus vulgaris cultivated common bean. I. Allozymes. Crop Science 31: Fabaceae). Economic Botany 44: Singh S.P., Gutierrez A.J., Molina A., Urrea C. and Gepts P. 1991c. Lioi L Geographical variation of phaseolin patterns in an Old Genetic diversity in cultivated common bean. II. Marker-based World collection of Phaseolus vulgaris. Seed Sci. and Technol. analysis of morphological and agronomic traits. Crop Science 17: : Ma Y. and Bliss F.A Seed proteins of common bean. Crop Sneath P.H.A. and Sokal R.R Numerical Taxonomy. W. H. Sci. 18: Freeman Co., San Francisco. Matos M., Carnide V., Martins S. and Guedes-Pinto H Vanderberg A. and Nleya T Breeding to improve plant type. Allozyme diversity in landraces of the common bean from North In: Singh S.P (ed.), Common bean improvement in the twenty- Interior Portugal. In: De Ron A.M (ed.), International Sym- first century. Kluwer Academic Publishers, The Netherlands, pp. posium on breeding of Proteins and Oil Crops. Fundacion Pedro Barrie de la Maza, Pontevedra, Spain, pp Zeven A.C The introduction of the common bean (Phaseolus Mc Clean P.E., Myers J.R. and Hammond J.J Coefficient of vulgaris L.) into western Europe and the phenotypic variation of parentage and cluster analysis of North American dry bean dry bean collected in the Netherlands in Euphytica 94: cultivars. Crop Science 33: Rohlf F.J NTSYS-pc. Numerical Taxonomy and Multi- Zeven A.C The traditional inexplicable replacement of seed variate Analysis System. Exeter Software, Dep. of Ecology and and seed ware of landraces and cultivars: A review. Euphytica Evolution, State University of New York. 110:

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