Schriften zu Genetischen Ressourcen - Genres

Schriften zu Genetischen Ressourcen - Genres Schriften zu Genetischen Ressourcen - Genres

30.11.2012 Views

B. PICKERSGILL, M.I. CHACÓN SÁNCHEZ and D.G. DEBOUCK These data support BEEBE et al. (2000) in suggesting that at least three of the four Mesoamerican races were domesticated independently: Race Jalisco in Jalisco; Race Guatemala in Guatemala or Chiapas; Races Mesoamerica and Durango in northern and/or southern Mexico. If there was only a single domestication, as GEPTS (1990) argued, then it probably occurred in Oaxaca rather than Jalisco, since in Oaxaca the relevant chloroplast haplotypes all occur and S phaseolin is also present. Infraspecific taxonomy of Phaseolus vulgaris Three particularly significant events have occurred in the evolutionary history of P. vulgaris. Firstly, early intercontinental dispersal resulted in genetic isolation of wild populations in the two continents, so they differentiated into what have been called the Mesoamerican and Andean gene pools. Secondly, beans in each of these gene pools were independently domesticated, probably in more than one location in the case of Mesoamerica. Thirdly, domesticated beans diverged and diversified under human selection into different ecogeographic groups (races), different agronomic groups (e.g., climbing or bush beans) and different use groups (popping beans, black beans, navy beans, pinto beans, etc.). The question then arises, should any or all of these events be recognised in the infraspecific classification of P. vulgaris and if so, how? Intercontinental dispersal and divergence of gene pools The many differences between Mexican and Andean wild beans have led to their being recognised as distinct varieties (see DEBOUCK 1991 for further discussion). This provides a formal taxonomic rank for the informal category of gene pool. Wild beans from Ecuador and northern Peru are morphologically intermediate between those of the Mesoamerican and Andean gene pools (DEBOUCK et al. 1993), and combine isozyme alleles characteristic of both gene pools (GEPTS 1993). They are genetically unique in their RAPD and AFLP patterns (FREYRE et al. 1996, TOHME et al. 1996) and in their mitochondrial DNA (KHAIRALLAH et al. 1992). They also carry an apparently ancestral type of phaseolin (KAMI et al. 1995). It has been suggested that these populations are relicts of the ancestral stock from which the Mesoamerican and Andean gene pools are derived (KAMI et al. 1995, TOHME et al. 1996). Wild beans from Costa Rica south to Colombia are also morphologically intermediate (SINGH et al. 1991), while a study of AFLPs showed a genetic continuum between wild beans from the northern and southern ends of the range (TOHME et al. 1996). Under the Rules of the International Code of Botanical Nomenclature, populations that are intermediate between two varieties must be included in one or the other 77

Multiple domestications and their taxonomic consequences: Phaseolus vulgaris (which causes problems in distinguishing between the varieties) or else a third variety must be created for the intermediates. The advantage of an informal category such as gene pool is that the situation can be described without every accession having to be placed in a gene pool. In other words, the Mesoamerican and Andean gene pools can be recognised, together with the Ecuadorian / north Peruvian relict of the ancestral gene pool if desired. Wild beans from intervening areas can be treated in whatever way the available information seems to warrant. For example, they may be left as intermediates or placed in additional gene pools. Furthermore, gene pools, unlike taxonomic varieties, do not have to be morphologically distinguishable. Gene pool is a category used to express genetic differentiation, not necessarily a category that can be recognised in the field. The chloroplast haplotypes define three lineages of wild P. vulgaris (Fig. 1). These agree only partially with the picture based on gene pools. Haplotypes E and F are found in the putatively ancestral relict populations of Ecuador and northern Peru and seem to represent early derivatives of a lineage that then spread into Central America (haplotypes G and H). No elements of this lineage seem to have been domesticated. A second lineage, bearing haplotypes I to O, contains wild beans of the Mesoamerican gene pool and some of the intermediates. However, the third lineage (haplotypes A to D and P) combines morphologically distinct wild beans from both Mesoamerican and Andean gene pools. These chloroplast data are potentially significant with regard to both conservation and utilisation of genetic resources of wild beans. For example, Mesoamerican beans with haplotypes A, B and P appear to have evolved independently of Mesoamerican beans with haplotypes I to O for a long time so may carry significantly different diversity. They should therefore be prime candidates for both conservation and evaluation. However, we consider it premature to try to represent the chloroplast lineages in either formal or informal classifications, at least until some nuclear phylogenies are available and the chloroplast lineages checked for congruence with these. Domestication As in many species that include a crop and its conspecific wild progenitor, wild and domesticated common beans are usually placed in different subspecies, ssp. aborigineus and ssp. vulgaris respectively (see POLHILL and VAN DER MAESEN 1985 for discussion of this treatment for grain legumes in general, including Phaseolus). However, subspecies ranks above variety in the taxonomic hierarchy, so if the Mesoamerican and Andean gene pools are treated as varieties, they cannot then be subdivided into subspecies. Moreover, at whatever rank the gene pools are treated, four (not two) names at a lower rank would be needed to classify the wild and domesticated beans within each gene pool. The current Code of Nomenclature for Cultivated Plants (TREHANE et al. 1995) recommends that cultivated plants below the 78

Multiple domestications and their taxonomic consequences: Phaseolus vulgaris<br />

(which causes problems in distinguishing between the varieties) or else a third variety<br />

must be created for the intermediates. The advantage of an informal category such<br />

as gene pool is that the situation can be described without every accession having to<br />

be placed in a gene pool. In other words, the Mesoamerican and Andean gene pools<br />

can be recognised, together with the Ecuadorian / north Peruvian relict of the ancestral<br />

gene pool if desired. Wild beans from intervening areas can be treated in whatever<br />

way the available information seems to warrant. For example, they may be left<br />

as intermediates or placed in additional gene pools. Furthermore, gene pools, unlike<br />

taxonomic varieties, do not have to be morphologically distinguishable. Gene pool is<br />

a category used to express genetic differentiation, not necessarily a category that can<br />

be recognised in the field.<br />

The chloroplast haplotypes define three lineages of wild P. vulgaris (Fig. 1). These<br />

agree only partially with the picture based on gene pools. Haplotypes E and F are<br />

found in the putatively ancestral relict populations of Ecuador and northern Peru and<br />

seem to represent early derivatives of a lineage that then spread into Central America<br />

(haplotypes G and H). No elements of this lineage seem to have been domesticated.<br />

A second lineage, bearing haplotypes I to O, contains wild beans of the<br />

Mesoamerican gene pool and some of the intermediates. However, the third lineage<br />

(haplotypes A to D and P) combines morphologically distinct wild beans from both<br />

Mesoamerican and Andean gene pools. These chloroplast data are potentially significant<br />

with regard to both conservation and utilisation of genetic resources of wild<br />

beans. For example, Mesoamerican beans with haplotypes A, B and P appear to<br />

have evolved independently of Mesoamerican beans with haplotypes I to O for a long<br />

time so may carry significantly different diversity. They should therefore be prime<br />

candidates for both conservation and evaluation. However, we consider it premature<br />

to try to represent the chloroplast lineages in either formal or informal classifications,<br />

at least until some nuclear phylogenies are available and the chloroplast lineages<br />

checked for congruence with these.<br />

Domestication<br />

As in many species that include a crop and its conspecific wild progenitor, wild and<br />

domesticated common beans are usually placed in different subspecies, ssp. aborigineus<br />

and ssp. vulgaris respectively (see POLHILL and VAN DER MAESEN 1985 for<br />

discussion of this treatment for grain legumes in general, including Phaseolus). However,<br />

subspecies ranks above variety in the taxonomic hierarchy, so if the<br />

Mesoamerican and Andean gene pools are treated as varieties, they cannot then be<br />

subdivided into subspecies. Moreover, at whatever rank the gene pools are treated,<br />

four (not two) names at a lower rank would be needed to classify the wild and domesticated<br />

beans within each gene pool. The current Code of Nomenclature for Cultivated<br />

Plants (TREHANE et al. 1995) recommends that cultivated plants below the<br />

78

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