Genet Resour Crop Evol (2012) 59:655–681 DOI 10.1007/s10722-011-9709-2 RESEARCH ARTICLE Inventory of related wild species of priority crops in Venezuela Chiara Berlingeri • Manuel B. Crespo Received: 17 December 2010 / Accepted: 16 May 2011 / Published online: 15 June 2011 Ó Springer Science+Business Media B.V. 2011 Abstract A prerequisite in any conservation programme of Plant Genetic Resources is estimation of diversity. The inventory of wild and naturalized relatives of priority crops in Venezuela (CWR) is based on the main Catalogues of Flora in the country, selecting taxa closely related to crops, according to the concepts of ‘‘gene pool’’ and ‘‘taxonomic group’’. We included 47 genera, 217 species and 228 taxa belonging to 28 plant families. Among them, those with higher richness are: Fabaceae, Solanaceae, Araceae, Lauraceae, Dioscoreaceae, Poaceae, Rosaceae and Myrtaceae. Genera with a higher number of species are Xanthosoma, Persea, Dioscorea, Prunus, Psidium, Phaseolus, Solanum, Vigna, Capsicum, Manihot, Theobroma, Ipomoea and Oryza. A total of 26 endemic species are found, which belong to genera Xanthosoma, Persea, Dioscorea, Prunus and Manihot. The primary gene pool of crops include native species from genera such as Manihot, Solanum (Section Petota), Lycopersicon, Ananas, Capsicum, Dioscorea, Xanthosoma, Phaseolus, Theobroma, Ipomoea, Gossypium, Arracacia and Psidium. Genera C. Berlingeri (&)  M. B. Crespo CIBIO (Instituto de la Biodiversidad), Universidad de Alicante, P.O. Box 99, 03080 Alicante, Spain e-mail: cabg1@alu.ua.es; cberlingeri@gmail.com C. Berlingeri Instituto Nacional de Investigaciones Agrı́colas (INIA), Estación Experimental Trujillo, Avenida Principal de Pampanito, 3152 Estado Trujillo, Venezuela with native species weakly related to crops are Saccharum, Persea, Ipomoea, Prunus, Vigna, Solanum (Section Melongena) and Daucus. Crop genera without native species in Venezuela are Allium, Musa, Brassica, Spinacia, Helianthus, Pisum, Lactuca, Citrus, Elaeis, Beta, Glycine and Triticum. Only a few taxa have already been evaluated according to the IUCN criteria, and Venezuelan accessions of crop wild relatives in national and international genebanks are very scarce. Keywords Crop wild relatives  Inventory  Plant genetic resources  Venezuela Introduction Venezuela shows a higher biological diversity among the Caribbean countries of South America due to the convergence of several biogeographic regions together with a tropical climate, which places it among the top most biodiverse countries worldwide (Huber et al. 1998). Plant diversity with real or potential value for agriculture is known as Plant Genetic Resources (hereafter PGR) and in a broad sense it includes traditional varieties, modern cultivars and Crop Wild Relatives (hereafter CWR; FAO 1996a, 2001). Those resources are basic for the improvement of cultivated plants and agricultural ecosystems sustainability. 123 656 In the same way, as the rest of biodiversity, PGR are threatened by mismanagement of landscape. Habitat fragmentation, climate change and agricultural intensification put at risk the crop wild relatives and traditional varieties. Accordingly it is urgent to take actions to reduce genetic erosion. A prerequisite to define strategies for conservation and sustainable use of crop genetic resources is to generate an inventory of CWR to assess the diversity and to keep track of it (FAO 1996b). The study of PGR can be considered at different scales. In a global perspective, it seems that the number of crops to meet food requirements is relatively small (staple crops), whereas in a local analysis the number of useful plants may increase, becoming then important the underutilized crops, local varieties and wild useful plants. In this frame, the present study was aimed at creating a national inventory of wild and naturalized relatives of priority crops in Venezuela, as a useful tool for the establishment of PGR conservation strategies. Methods The inventory of CWR was mainly based on the New Catalogue of the Vascular Flora of Venezuela (Hokche et al. 2008). Taxa recorded there were selected according to its closeness to priority crops of Venezuela, following the ‘‘gene pool’’ concept of Harlan and de Wet (1971) and the ‘‘taxonomic group’’ concept of Maxted et al. (2006). Both native and introduced taxa were included in order to bring a true vision about the importance that these PGR represent in the national economy. Authorships of the names of taxa correspond to those accepted in the International Plant Names Index (IPNI 2011). The ‘‘gene pool’’ concept takes into account the genetic distance and hybridization ability between the crop and the relatives to assign the latter to a primary (GP1), secondary (GP2) or tertiary (GP3) gene pool of the crop (Harlan and de Wet 1971). In those cases where information on genetic diversity and hybridization was insufficient, the taxonomic hierarchy was used to estimate the relationship degree. ‘‘Taxon groups’’ were defined according to taxa pertainance to either the same species, series (or section), subgenus, genus or tribe as the crop (TG1, TG2, TG3, TG4 and TG5, respectively) (see Maxted et al. 2006). 123 Genet Resour Crop Evol (2012) 59:655–681 Finally, in cases of crop genera in which no formal taxonomic treatments are available (i.e. those in which series, sections or subgenera have not been defined), all species of the concerned genus occurring in Venezuela are mentioned as a potential crop gene pool. Records in Venezuela for each genus included in the CWR inventory were taken from databases of major herbaria and genebanks worldwide, which were accessed on line through the Global Biodiversity Information Facility, GBIF (www.gbif.net). In those cases in which doubtful records were found, original providers of databases and taxonomists were contacted. Since the Venezuelan flora is not yet accessible in a database that can be matched digitally with the existing crop databases, the process to produce the national CWR inventory may be carried out manually (Maxted pers. comm. 2008). Therefore, a list of priority crops in Venezuela was first established by consulting national and international statistics. Secondly, it was compared with the taxonomic treatments and genetic diversity studies of genera to select those taxa belonging to the corresponding ‘‘gene pool’’ or ‘‘taxonomic group’’ of the priority crops. Once the list of species was completed, the database was supplemented with additional information to facilitate the selection of taxa requiring further research or conservation actions. Table 1 shows criteria followed here to generate the CWR datable in Venezuela, which is mostly based on those used by previous authors to priorize species for conservation (Maxted et al. 1997; Magos 2008). Results and discussion Taxonomic diversity of crop wild relatives The inventory of priority CWR for Venezuela (‘‘Appendix’’) is arranged according to the economic importance of the crop. It includes 47 genera, 217 species and 228 taxa belonging to 28 plant families. Of these, those with higher taxa richness are Fabaceae, Solanaceae, Araceae, Lauraceae, Dioscoreaceae, Poaceae, Rosaceae and Myrtaceae, which comprise 67% of the whole taxa included in the inventory. The families with most genera with native species are Solanaceae and Fabaceae (Table 2). Genet Resour Crop Evol (2012) 59:655–681 657 Table 1 Criteria considered for the database elaboration of CWR in Venezuela Criteria Categories Scientific name – Sources of information Hokche et al. (2008) IPNI, The International Plant Names Index. Published on the Internet http://www.ipni.org (accessed May 2011) Synonyms – Hokche et al. (2008). TROPICOS (2009), Missouri Botanical Garden, http://www.tropicos.org (accessed January 2009) Status Endemic, native, naturalized, introduced (archaeophyte, neophyte), Uncertain. Hokche et al. (2008) Economic importance Production value of the related crop in Venezuela FAO (2005) FAOSTAT, http://faostat.fao.org (accessed September 2008) Taxonomic relationship with the crop TG1a, TG1b, TG2, TG3, TG4 and TG5. Accepted taxonomic treatments for the genus (see ‘‘Appendix’’ for references) Harlan (1992) Mabberley (2008) USDA, ARS, Germplasm Resourses Information Network (GRIN), http://www.ars-grin.gov/cgi-bin/npgs/html/index.pl (accessed February to July 2009) Genetic relationship with the crop GP1a, GP1b, GP2, GP3 See ‘‘Appendix’’ for references Threat degree Least Concern (LC), Near Threatened (NT), Vulnerable (VU), Endangered (EN), Critically Endangered (CR), The IUCN Red List of Threatened Species (2009), http://www.iucnredlist.org (accessed April 2009) Llamozas et al. (2003) Distribution Pantropical, paleotropical, neotropical, Endemic. Mabberley (2008) National and International ex situ conservation Number of collections and accessions in national and international genebanks Hokche et al. (2008) TROPICOS (2009), Missouri Botanical Garden, http://www.tropicos.org (accessed April 2009) GBIF, Global Biodiversity Information Facility (2009), http://www.gbif.net (accessed June 2009) Bioversity International (2009), http://www.bioversityinternational.org (accessed June 2009) SINGER, The System—wide Information Network for Genetic Resources (2009), http://www.singer.cgiar.org/index.jsp?page=collections Knudsen (2000) Only 24 (51%) out of 47 genera related with priority crops have native species in Venezuela, while 23 (49%) are introduced. As regards to the taxa listed, 140 are native, 55 are introduced and 33 have a doubtful origin (Table 2). Among the introduced species, both archaeophytes (12) and neophytes (44) can be found. The former are species whose introduction was long before the discovery of America, so it is likely that evolutionary adaptations have occurred during all that time. The neophytes are plants introduced after the American discovery, so they probably have a narrower diversity. However, no generalization is possible, because this issue has much to do with the reproduction way and biology of each species. The taxonomic representation of the genera was estimated by the ratio between the number of Venezuelan species and the total number of species known for each genus, following Mabberley (2008). According to that, the genera of cultivated plants best represented in Venezuela are Capsicum, Ananas, Xanthosoma, Theobroma, Cucurbita, Oryza, Zea, Psidium, and Phaseolus. 123 658 Genet Resour Crop Evol (2012) 59:655–681 Table 2 Summary of taxa diversity of major crop relatives in Venezuela Crop Family Genus Corn Poaceae Zea Species number Taxa number 1 2 Endemic Native Archaeophyte Neophyte 1 Doubtful 1 Rice Poaceae Oryza 6 6 2 4 Sugarcane Poaceae Saccharum 4 4 1 3 Cassava Euphorbiaceae Manihot 7 8 Plantain, Banana Musaceae Musa 1 2 Potato Solanaceae Solanum 9 10 8 1 Pineapple Bromeliaceae Ananas 4 4 Onion, Garlic Alliaceae Allium 2 2 1 7 2 2 2 1 2 Coffee Rubiaceae Coffea 3 3 3 Sorghum Poaceae Sorghum 4 4 4 Tomato Solanaceae Lycopersicon 1 2 Melon, Cucumber Cucurbitaceae Cucumis 5 5 1 1 4 Peppers Solanaceae Capsicum 8 10 5 Carrot Apiaceae Daucus 2 2 1 5 1 Coconut Arecaceae Cocos 1 1 Watermelon Cucurbitaceae Citrullus 1 1 1 Orange, other citruses Rutaceae Citrus 4 4 4 Papaya Caricaceae Carica 1 1 Tobacco Solanaceae Nicotiana 2 2 1 1 1 Yam Dioscoreaceae Dioscorea 18 18 5 11 Tannia Araceae Xanthosoma 22 22 13 9 Avocado Beans Lauraceae Fabaceae Persea Phaseolus 20 10 21 10 5 15 7 Guava Myrtaceae Psidium 13 13 Mango Anacardiaceae Mangifera 1 1 Cacao Sterculiaceae Theobroma 7 8 Sesame Pedaliaceae Sesamum 1 1 1 2 1 2 1 12 1 1 8 1 Grape Vitaceae Vitis 2 2 1 Cabbage, Cauliflower Brassicaceae Brassica 1 2 2 Lettuce Asteraceae Lactuca 1 1 1 Cotton Malvaceae Gossypium 2 2 Strawberry Rosaceae Fragaria 1 1 Peach Beet Rosaceae Chenopodiaceae Prunus Beta 13 1 13 1 Sweet potato Convolvulaceae Ipomoea 6 6 1 1 2 8 1 1 5 Solanaceae Solanum 1 1 1 Fabaceae Cajanus 1 1 1 Agavaceae Agave 3 4 Fabaceae Glycine 1 1 Peanut Fabaceae Arachis 3 3 Sunflower Asteraceae Helianthus 1 1 123 2 1 Eggplant Sisal 1 1 Pigeon pea Soya 1 3 1 1 1 2 1 Genet Resour Crop Evol (2012) 59:655–681 659 Table 2 continued Crop Family Genus Species number Taxa number Peas Fabaceae Pisum 1 1 Wheat Poaceae Triticum Cowpea Fabaceae Vigna Pumpkin, Squash Cucurbitaceae Cucurbita 1 1 10 10 4 4 Arracacha Apiaceae Arracacia 4 4 Palm oil Arecaceae Elaeis 1 1 Spinach Chenopodiaceae Spinacia Total 1 1 217 228 The inventory of CWR contains 26 endemic species to Venezuela (12%), belonging to genera Xanthosoma (13), Persea (5), Dioscorea (5), Prunus (2) and Manihot (1) (Table 2). The highest percentage of endemic species is found in Amazonas and Bolı́var states, followed in smaller numbers by Lara, Merida and Táchira (Hokche et al. 2008). Likewise, genera with distributions restricted to the Americas are also important (e.g. such as Xanthosoma, Psidium, Phaseolus, Capsicum, Manihot, Theobroma, Ananas, Arracacia, Cucurbita, Arachis, Agave, Lycopersicon, Carica, Helianthus and Zea), which account for 90 species (41%). Other infrageneric taxa are also endemic to America, such as Solanum sect. Petota and the CCDD genome species of Oryza (Hawkes 1990; Hijmans and Spooner 2001; Subudhi et al. 2006). Genera including the highest number of species are Xanthosoma, Persea, Dioscorea, Prunus, Psidium, Phaseolus, Solanum, Vigna, Capsicum, Manihot, Theobroma, Ipomoea and Oryza (Table 2). This fact responds to the inclusion of species according to the genetic and taxonomic proximity of wild taxa to the crop, since more accurate genetic data were not available. Diversity assessment of crop genera in Venezuela According to FAO statistics (FAO 2005) the most commercially valuable crop in Venezuela is maize (Zea mays L.). Although Venezuela has no related wild species of this crop, there exists great infraspecific diversity with a high potential for genetic Endemic Native Archaeophyte Neophyte Doubtful 1 1 7 1 2 4 3 1 1 1 26 114 11 44 33 improvement and direct use (Grant et al. 1965; Alfaro and Segovia 2000). Rice (Oryza sativa L.) and sugarcane (Saccharum officinarum L.) occupy respectively the second and third position in commercial value. Venezuela is neither the origin centre nor the diversity centre for both species, this showing the dependence of the country for those food genetic resources. In the case of the genus Saccharum, although the diversity of native species has not been studied, they are geographically and taxonomically distant from the crop species complex (D’Hont et al. 2008). The genus Oryza is usually classified into 21 wild and two cultivated species, which have been assigned to ten distinct genomes following hybridization studies and molecular analyses (Khush 1997; Subudhi et al. 2006). In Venezuela only three species are found belonging to the O. sativa complex with AA genome (O. sativa L., O. rufipogon Griff. and O. glumaepatula Steud.) and three species from the O. officinalis complex with CCDD genome (O. alta Swallen, O. grandiglumis (Döll) Prodoehl and O. latifolia Desv.). The species O. glumaepatula, O. alta, O. grandiglumis and O. latifolia are endemic to Neotropics (Khush 1997; Subudhi et al. 2006). Although O. glumaepatula is not mentioned in floristic catalogues of Venezuela, the website ‘‘Cereal Knowledge Bank’’ managed by the IRRI and CIMMYT (http://www.knowledgebank.irri.org/default. htm) includes Venezuela in the range of distribution for this species. Populations of O. glumaepatula show a great variation and are difficult to distinguish morphologically from other species with AA genome 123 660 (Juliano et al. 1998), being often referred as a synonym of O. rufipogon (www.tropicos.org). However, there are reproductive barriers (Naredo et al. 1998) and molecular differences (Ge et al. 1999) that warrant segregation of O. glumaepatula from the Asian AA genome of O. rufipogon. Similarly the genetic diversity of CCDD genome species from Latin America is unclear. Molecular and hybridization data show that CCDD species are very closely related and appear to have varied according to ecological conditions (Jena and Kochert 1991). The presence of wild (or feral) rice with AA genome in Venezuela, which continuously are exchanging genes with the crops (Chen et al. 2004), and the scarce knowledge about the genetic identity of AA and CCDD populations makes a priority the phylogeographic study of the genus in Venezuela (Pérez-Almeida pers. comm. 2009). Manihot, Solanum (section Petota), Ananas, Lycopersicon, Capsicum, Dioscorea, Xanthosoma, Phaseolus, Psidium, Theobroma, Gossypium, Ipomoea and Arracacia are economically the most important genera with native species in Venezuela, which belong to the primary gene pool of the crops. Among them, Manihot, Solanum, Capsicum, Dioscorea, Xanthosoma, Phaseolus, Psidium, Theobroma and Ipomoea, include over 10 species each (Table 2). The most recent monograph of Manihot classifies M. esculenta as the only species within the section Manihot (Rogers and Appan 1973). However, phylogenetic studies have shown that sections accepted by Rogers and Appan (1973) are not monophyletic, therefore the genus is now in need of revision (Chacón et al. 2008). Furthermore, although M. esculenta belongs to a particular section, it can produce hybrids with some wild species of other sections (Nassar et al. 2008), suggesting that some of those sections may be closer. Since in Venezuela there are wild populations of M. esculenta and related wild species, including the endemic species M. filamentosa Pittier, it would be interesting to deal with phylogeographic studies of the genus for conservation purposes. The classification system most widely used for Solanum (D’Arcy 1972, 1991) arranges the genus into seven subgenera, including subg. Potatoe (G. Don) Walp., to which S. tuberosum L. (potato) belongs, and the subgenus Leptostemonum (Dunal) Bitter, which includes S. melongena (egg-plant). No wild relatives of the latter species are found in Venezuela. The wild 123 Genet Resour Crop Evol (2012) 59:655–681 relatives of potato are grouped into Solanum section Petota Dumort subsection Potatoe, according to the classification of Hawkes (1990), being very frequent the interspecific hybridization within subsection (Spooner and Hijmans 2001). In the Catalogue of the Flora of Venezuela (Hokche et al. 2008) are reported seven wild species of Solanum, section Petota in the Andean states of Venezuela (Solanum colombianum Dunal, Solanum juglandifolium Humb. et Bonpl. ex Dunal, Solanum otites Dunal, Solanum paramoense Bitter ex Pittier, Solanum subpanduratum Ochoa, Solanum tuberosum L., Solanum woodsonii Correll). However, the database of the International Potato Center (CIP) shows a record identified as S. curtilobum Juz. et Bukasov collected in Trujillo state, and S. flahaultii Bitter is recorded by several authors (Nee 1999; Hawkes 1990; Luteyn 1999) as present in Venezuela. The existence of S. curtilobum in Venezuela is not surprising, as it is a cultivated species (Spooner, pers. comm.). The results of morphological, enzymatic, karyological, hybridization and molecular analyses support the hypothesis that three independent evolutionary lineages are involved in the origin of the domesticated Capsicum. C. annuum, C. chinense and C. frutescens integrate the ‘‘white flowered C. annuum group’’, in which few reproductive barriers and high morphological and molecular similarities exist. A second lineage with yellow-spotted white flowers includes C. baccatum, and finally, a third lineage includes the shrubby purple-flowered Capsicum to which C. pubescens belongs (McLeod et al. 1982; Walsh and Hoot 2001; Moscone et al. 2006). In terms of hybridization ability, the species of C. pubescens complex are fully compatible to one another, but show unilateral incompatibility with species from other groups. Crosses between species of the remaining two groups show no incompatibility (Naci and Pickersgill 2004). Relationships of C. cumanense Fingerh. and C. rhomboideum Kuntze with the rest of Venezuelan Capsicum species and their taxonomic status, are rather uncertain due to the absence of studies. Although the Catalogue of the Flora of Venezuela (Hokche et al. 2008) does not include C. chinense, the database of the National Plant Germplasm System of the United States Department of Agriculture, (USDA—ARS) (http://sun.ars-grin.gov/npgs/search grin.html) and the Centre for Genetic Resources at Wageningen University, CGN (http://www.cgn.wur. Genet Resour Crop Evol (2012) 59:655–681 nl/applications/cgngenis/) registered accessions collected in Venezuela. Cultivated taxa of Dioscorea appear to have been originated in tropical regions of three continents. Among these, D. trifida L.f is the only cultivated species originated in South America (Mignouna et al. 2007), and also it is the only Venezuelan species included by Knuth (1924) in the section Macrogynodium (subgenus Eudioscorea). Knuth’s classification divides the genus into four subgenera and 58 sections, but recent phylogenetic studies have provided evidence for the reclassification of Dioscoreaceae and its genera (Caddick et al. 2002), as well as for simplifying the classification proposed by Knuth (Wilkin et al. 2005). Engler (1920) arranged the genus Xanthosoma in two sections: Euxanthosoma and Acontias. After that, many species have been reported for Venezuela (Bunting 1975, 1979, 1986, 1988; Madison 1981; Croat and Lambert 1986), though only describing the morphology and distribution of the species. There is not an updated taxonomic treatment of the genus or references about phylogenetic relationships among species. In the present inventory, therefore, all species of Xanthosoma reported for Venezuela are included. With regard to Phaseolus, the Catalogue of the Vascular Flora of Venezuela (Hokche et al. 2008) includes eight species; however three of these were transferred to the related genera Macroptilium and Vigna according to a narrower circumscription of the genus. Also, P. aborigineus Burkart is regarded as Phaseolus vulgaris L. var. aborigineus (Burkart) Baudet. The presence of P. tuerckheimii in Venezuela is cited for the states of Monagas and Táchira, although Delgado-Salinas et al. (2006) restricted its range only to Mexico and Central America. Likewise, P. unilobatus was published as a new species by Pittier (1944) from a single sample collected in the Monagas state, this species being only accepted in the Venezuelan catalogue (Hokche et al. 2008). Moreover, the database of the USDA-ARS (http://sun.arsgrin.gov/npgs/searchgrin.html) includes an accession labelled P. dumosus Macfad, collected in Mérida, Venezuela. This species is referred as native to Central America but it is cultivated and widely naturalized in northwestern South America. In addition, in the Nationaal Herbarium Nederland (NHN) and the Missouri Botanical Garden (MO) there are samples 661 identified as P. coccineus subsp. darwinianus collected in Venezuela. Since the taxonomic definition and nomenclature of the Venezuelan species of Phaseolus is uncertain and confusing, there is a need for a revision of the genus for the country. We are currently undertaking a revision to define key characters for taxa identification, to establish their geographical distribution and to infer their phylogenetic relationships. The sweet potato (Ipomoea batatas (L.) Lam.) and its closest wild relatives belong to subgenus Eriospermum Verdc., section Eriospermum Hallier f., series Batatas (Choisy) D.F. Austin (Austin and Huáman 1996). Crossing of species within this group is complex because of genetic, cytogenetic and physiological interactions. Studies with molecular markers and cytogenetics have been used to infer phylogenetic relationships in the group, confirming a close relationship between I. batatas and I. trifida G.Don and a greater genetic distance with the rest of species in the series (Huang et al. 2002; Rajapakse et al. 2004; Srisuwan et al. 2006). Threat degree of taxa Estimating the threat degree of native taxa related to cultivated species is very difficult in Venezuela, due to the lack of detailed studies on that subject. In the present work the list of endangered species of IUCN (www.iucnredlist.org) and the Red Book of Venezuela (Llamozas et al. 2003) were checked. The latter assigned threat categories to 1,598 out of 15,820 native species reported in the Catalogue of Venezuela (Hokche et al. 2008). Table 3 shows the crop relatives included in any of the IUCN categories. Only seven species in the present inventory have been evaluated, which are in the lower risk categories. The scarce number of taxa so far evaluated under the IUCN criteria as well as the poor knowledge about conservation status of populations of crop relatives, highlight the urgent need to conduct such studies. Another approach useful to estimate the threat of taxa is to consider their geographic distribution and endemicity. This information was taken from the New Catalogue of the Vascular Flora of Venezuela (Hokche et al. 2008), in which the distributions were based on data from the herbarium specimens and literature. Since some areas of the country have not been adequately sampled yet and many taxa are 123 662 Genet Resour Crop Evol (2012) 59:655–681 Table 3 Crop relatives included in lists of threatened species Family Taxon Euphorbiaceae Manihot brachyloba Müll.Arg. Native MR/pm Euphorbiaceae Manihot filamentosa Pittier Endemic IC Euphorbiaceae Euphorbiaceae Manihot surinamensis D.J. Rogers et Appan Manihot tristis Müll.Arg. subsp. tristis Native Native MR/pm IC Lauraceae Persea caerulea (Ruiz et Pav.) Mez Native MR/pm Myrtaceae Psidium cinereum DC. Native MR/ca Solanaceae Solanum flahaultii Bitter Doubtful MR scarcely collected in the herbaria, the present analysis must be considered preliminary in order to select priority taxa for future studies. As mentioned before, endemic species related to crops belong to Xanthosoma, Persea, Dioscorea, Prunus and Manihot. Llamozas et al. (2003) cited Solanum paramoense and S. subpanduratum as endemic to the Venezuelan Andes. However, several authors consider S. subpanduratum as a synonym of S. colombianum, and others doubt about the status of S. paramoense which is a matter of current work (Spooner, pers. comm.). Most endemic species are distributed in the south of Orinoco River (Bolivar and Amazonas states), excepting Xanthosoma, Manihot and Solanum that have endemic species towards the north of Venezuela. In this regard, it is noteworthy that the states located in the south of Venezuela have a population density and habitat fragmentation considerably smaller than those in the north, and therefore the degree of threat to those populations is very likely weaker. When the genera with native species are considered, the highest species richness is concentrated in Guayana and Andes regions. Dioscorea, Theobroma and Ananas have the highest species representation in the southern parts of the country (Amazonas and Bolivar states). The genus Arracacia, Solanum (section Petota) and Phaseolus vulgaris var. aborigineus (the closest wild relative of beans) are concentrated in the Andean states. Some genera have a wide distribution in the country, such as Xanthosoma, Manihot and Capsicum. Conservation status of taxa Only 44 taxa (19%) out of those included in the present inventory have at least one entry in 123 Status IUCN category international genebanks. Of these, 25 are cultivated species (57%) and 19 are wild ones (43%). Regarding to ex situ conservation in Venezuela, 37 taxa (16%) are present in national genebanks, most of which correspond to cultivated species (32 taxa, 86%). Only five taxa (14%) belong to non cultivated species: Lycopersicon esculentum var. cerasiforme (Dunal) A.Gray, Solanum tuberosum subsp. andigenum, Theobroma grandiflorum (Willd. ex Spreng.) K.Schum., Ananas parguazensis Camargo et L.B.Sm. and Ananas nanus (L.B.Sm.) L.B.Sm. (‘‘Appendix’’). This suggests a low representation of Venezuelan plant genetic resources in national and international genebanks, especially of wild relatives of crops. However, the above must be analyzed carefully, since many genebanks, mainly those of Venezuela, lack updated online database and there are still many materials not fully identified (Pérez pers. comm. 2009). The genera with greater representation of taxa collected in Venezuela conserved in international genebanks are Lycopersicon (2), Gossypium (2), Nicotiana (2), Vigna (7), Solanum (6), Zea (1), Phaseolus (5), Capsicum (5) and Ipomoea (3). The species with the highest number of Venezuelan accessions in international genebanks are Zea mays, Phaseolus vulgaris, Manihot esculenta, Sorghum bicolor, Gossypium hirsutum, Sesamum orientale, Cajanus cajan and Ipomoea batatas. Genera with a higher number of taxa included in national collections are probably Ananas, Zea and Gossypium. The species with the highest number of accessions in national genebanks are Zea mays, Phaseolus vulgaris, Oryza sativa, Theobroma cacao, Manihot esculenta, Sorghum bicolor, Sesamum orientale, Vigna unguiculata, Cocos nucifera, Mangifera indica, Ipomoea batatas, Cajanus cajan, and those of Coffea, Lycopersicon, Gossypium and Capsicum. Genet Resour Crop Evol (2012) 59:655–681 No or very few Venezuelan germplasm of cultivated and related wild species of Dioscorea, Xanthosoma, Arracacia, Agave, Cucurbita and Carica have been collected, despite the regional or local importance of those crops. No ex situ germplasm accessions exist of endemic Venezuelan crop relatives of Xanthosoma, Persea, Dioscorea, Prunus and Manihot (‘‘Appendix’’). Estimating the risk of gene escape from genetically modified organisms (GMOs) In Venezuela, any activity related to GMOs that involves its release, production, trade and public use is not legally permitted. However, large quantities of seeds and agricultural products are imported from countries where production of transgenic crops is a common practice. In addition, several Venezuelan institutions are conducting genetic engineering projects (Delgado 2005). Consequently, the risk of escape of transgenic genes to nature and the commercial release of GMOs exists in a medium-term. In September 2003 the Cartagena Protocol, which considers the evaluation of the risks associated with transit, handling and use of GMOs for biodiversity conservation, became effective in Venezuela. Also the law of Biological Diversity of Venezuela (Ley de Diversidad Biológica 2000) establishes biosafety measures. The transgenic gene transfer to native or naturalized species may occur through intraspecific and interspecific hybridization, which would allow the continuation of these genes in nature and likely could affect agricultural activity and biodiversity. Although in the present inventory 58 priority crops were considered, the risk of gene escape varies depending on the crop. In Venezuela wild forms of cultivated species exist, such as Solanum tuberosum subsp. andigena, Lycopersicon esculentum var. cerasiforme, Ananas comosus var. parguazensis, Capsicum annuum var. glabriusculum and Phaseolus vulgaris var. aborigineus. All these have high reproductive compatibility with crops, which means that production of fertile hybrids is possible and probable. Other cultivated species natives to Venezuela are cotton (Gossypium barbadense), sweet potato (Ipomoea batatas), cacao (Theobroma cacao), yam (Dioscorea trifida), tannia (Xanthosoma sagittifolium) and cassava (Manihot esculenta). 663 Special care must also be taken with transgenic crops of introduced species that are regularly found in the natural vegetation and are able to naturalize in the wild, such as corn (Zea mays), squashes (Cucurbita sp. pl.), tobacco (Nicotiana tabacum), avocado (Persea americana), sesame (Sesamum orientale), cotton (Gossypium hirsutum), rice (Oryza sativa), coffee (Coffea sp. pl.), coconut (Cocos nucifera), strawberry (Fragaria vesca), guava (Psidium guajava), mango (Manguifera indica), melon (Cucumis melo), cucumber (Cucumis sativus), papaya (Carica papaya), pigeon pea (Cajanus cajan) and watermelon (Citrullus lanatus). On the other hand, it is important to consider those cultivated species capable of exchanging genes with other related species. It is the case fo Solanum tuberosum and the rest of species of section Petota; Ipomoea batatas and the wild species I. trifida; Oryza sativa and genome AA species (O. rufipogon and O. glumaepatula); Capsicum annuum complex species; annual and perennial taxa of Sorghum; cultivated species of Cucurbita; and Vitis vinifera and the American species V. tiliifolia. The experimental hybridization between Cucumis sativus and Cucumis melo is also possible (Deakin et al. 1971). At the present, commercial genetic modified crops exist of potato, tomato, cotton, corn, rice, melon, tobacco, squash and papaya, among others (Delgado 2005). Besides, many others transgenic are in progress to be produced, among which bean and pineapple are found. In this regard, especial attention may be payed to erosion risk assessment in these crops. Another important issue in biodiversity conservation is the estimation of genetic contamination risks for endemic species. In Venezuela, endemic taxa are found in Xathosoma, Dioscorea, Manihot, Persea and Prunus, but their real relationships with the related crops are unknown. However, tannia (Xanthosoma sagittifolium), yam (Dioscorea trifida) and cassava (Manihot esculenta) are regional important crops, in which no transgenic varieties are known to date. Avocado (Persea americana) and peach (Prunus persica) are taxonomically distant from the Venezuelan species of those genera, and they are likely to be also genetically distant. Genera of priority crops without native or naturalized species in Venezuela are Allium, Musa, Brassica, Spinacia, Helianthus, Pisum, Lactuca, Citrus, Elaeis, 123 664 Beta, Glycine and Triticum. The cultivated species Solanum melongena, Saccharum officinarum, Prunus persica, Vigna unguiculata and Daucus carota seem to be remotely related to the Venezuelan native species, although no specific studies on their crossing ability have been made. Conclusions The present work was aimed at creating a national inventory of crop relatives for Venezuela, establishing a list of priority crops, and selecting manually taxa closest to the crops according to their genetic and/or taxonomic relationships. In this regard, a large amount of related literature was reviewed and synthesized in order to generate a list of priority species that can facilitate taxa selection for future studies and activities related to conservation and use of plant diversity. The inventory allows estimating the importance and research needs of taxa according to the following criteria: (1) economic importance of their related crop; (2) their taxonomic richness and representativeness; (3) the presence of endemic and native taxa; (4) the genetic relationships with the crop (their potential as gene donors and genetic pollution risks); (5) extinction risk, and (6) their inclusion in genebanks. Although a more general perspective could have been developed if a digitalized flora were available, the number of taxa possible to be managed for conservation purposes is limited. Results from CWR inventories in Europe have shown that a high proportion of the wild flora has potential as crop relatives, and in these cases a posterior prioritization process has been recommended (Maxted et al. 2007; 123 Genet Resour Crop Evol (2012) 59:655–681 Magos et al. 2008). In the present contribution the prioritization criteria were initially established and the practical utility of the methodology was confirmed to select priority taxa for conservation. The genera with native species that are part of the gene pool of the crop are: Manihot, Solanum (Section Petota), Lycopersicon, Ananas, Capsicum, Dioscorea, Xanthosoma, Phaseolus, Theobroma, Ipomoea, Gossypium, Psidium and Arracacia. Further studies are still needed to determine the diversity and relationships between most of the genera and the related crop, though the Venezuelan species of Saccharum, Persea, Ipomoea, Prunus, Vigna, Solanum (Section Melongena) and Daucus seem to be weakly related to crops. The genera of priority crops that probably have no native or naturalized species in Venezuela are Allium, Musa, Brassica, Spinacia, Helianthus, Pisum, Lactuca, Citrus, Elaeis, Beta, Glycine and Triticum. The number of taxa that have been evaluated to date according to the IUCN criteria is virtually zero; and a low representation of Venezuelan CWR accessions exist in national and international genebanks. This is particularly true for the major genera Phaseolus, Capsicum, Ananas, Lycopersicon, Oryza, Manihot and Psidium, and for genera regionally important such as Dioscorea, Xanthosoma, Arracacia, Agave, Cucurbita and Carica. There are no germplasm in genebanks from Venezuelan endemic species of Xanthosoma, Persea, Dioscorea, Prunus and Manihot. Appendix See Table 4. Family Poaceae Poaceae Poaceae Euphorbiaceae Crop Corn Rice Sugarcane Cassava Manihot Saccharum Oryza Zea Genus Native Manihot esculenta Crantz Endemic Native Manihot carthagenensis (Jacq.) Müll. Arg. Manihot filamentosa Pittier Native Manihot brachyloba Müll. Arg. Doubtful Saccharum villosum Steud. Native Neophyte Saccharum officinarum L. Manihot anomala Pohl subsp. pubescens D.J.Rogers et Appan Doubtful Saccharum asperum Steud. Naturalized Doubtful Saccharum angustifolium Trin. Saccharum trinii (Hack.) Renvoize Oryza perennis Moench Oryza rufipogon Griff. Oryza sativa L. Doubtful Oryza alta Swallen Oryza latifolia Desv. Naturalized Doubtful Oryza grandiglumis (Döll) Prodoehl Doubtful Doubtful Oryza latifolia Desv. Oryza glumaepatula Steud. Oryza alta Swallen Doubtful Zea mays L. subsp. mexicana (Schrad.) H.H.Iltis Status Archaeophyte Synonymous Zea mays L. subsp. mays Taxon Table 4 Inventory of major crop relatives in Venezuela TG4 TG1 TG4 TG4 TG4 TG4 TG1 TG4 TG4 TG1 TG2 TG3 TG3 TG2 TG3 TG1 TG1 Taxon group Section Carthaginenses Section Manihot Section Carthaginenses Section Peruvianae Section Sinuatae Taxonomic classification ? GP1 ? ? GP1 GP1 GP2 GP3 GP3 GP2 GP3 GP1 GP1 Gene pool X X X X International X X X X National ex situ Conservation Rogers and Appan (1973), Nassar et al. (2008). D’Hont et al. (2008). Khush (1997), Vaughan and Tomooka (1999), Vaughan et al. (2003) Grant et al. (1965), Alfaro and Segovia (2000). References Genet Resour Crop Evol (2012) 59:655–681 665 123 Family Musaceae Musaceae Solanaceae Crop Plantain Banana Potato Table 4 continued 123 Solanum Musa Musa Genus Solanum tuberosum L. subsp. andigena (Juz. et Bukasov) Hawkes Solanum colombianum Dunal Solanum paramoense Bitter ex Pittier Solanum subpanduratum Ochoa Solanum tuberosum L. Native Solanum colombianum Dunal Solanum otites Dunal Native Native Native Native Doubtful Solanum flahaultii Bitter Solanum juglandifolium Humb. et Bonpl. ex Dunal Doubtful Solanum curtilobum Juz. et Bukasov Solanum paucijugum Bitter Native Neophyte Solanum colombianum Dunal Musa 9 paradisiaca L. Neophyte Native Manihot tristis Müll. Arg. subsp. tristis Musa 9 paradisiaca L. Native Manihot tristis Müll. Arg. subsp. saxicola D.J. Rogers et Appan Status Native Synonymous Manihot surinamensis D.J. Rogers et Appan Taxon TG1 TG2 TG2 TG2 TG2 TG2 TG2 TG2 TG1 TG1 TG4 TG4 TG4 Taxon group SubSection Potatoe, series Tuberosa. SubSection Potatoe SubSection Potatoe SubSection Potatoe SubSection Estolonifera SubSection Potatoe SubSection Potatoe SubSection Potatoe Section Heterophyllae Section Heterophyllae Section Heterophyllae Taxonomic classification Clade Potato Clade Potato Clade Potato Clade Potato Clade Potato Clade Potato Clade Potato Clade Potato Gene pool X X X X X International X X X National ex situ Conservation Hawkes (1990), Spooner et al. (1995), Spooner and Hijmans (2001), Huamán and Spooner (2002), Bohs (2005), Weese and Bohs (2007) References 666 Genet Resour Crop Evol (2012) 59:655–681 Family Solanaceae Bromeliaceae Alliaceae Rubiaceae Poaceae Solanaceae Crop Eggplant Pineapple Onion Coffee Sorghum Tomato Table 4 continued Lycopersicon Sorghum Coffea Allium Ananas Solanum Genus Native Ananas parguazensis Camargo et L.B.Sm. Neophyte Sorghum 9 drummondii (Nees ex Steud.) Millsp. et Chase Doubtful Neophyte Sorghum halepense (L.) Pers. Lycopersicon esculentum Mill. Neophyte Sorghum bicolor (L.) Moench Naturalized Coffea liberica W.Bull ex Hiern Neophyte Doubtful Coffea canephora Pierre ex A.Froehner Sorghum arundinaceum (Desv.) Stapf Naturalized Coffea arabica L. Neophyte Doubtful Ananas nanus (L.B.Sm.) L.B.Sm. Allium cepa L. Archaeophyte Ananas lucidus Mill. Neophyte Archaeophyte Ananas comosus (L.) Merr. var. comosus Solanum melongena L. Native Solanum colombianum Dunal Solanum woodsonii Correll Status Native Synonymous Solanum tuberosum L. subsp. andigena (Juz. et Bukasov) Hawkes Taxon TG1 TG1 TG2 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG2 TG1 Taxon group S. bicolor subsp. drummondii Section Sorghum S. bicolor subsp. bicolor S. bicolor subsp. arundinaceum SubSection Potatoe SubSection Potatoe, series Tuberosa Taxonomic classification GP1 GP2 GP1 GP1 Clade Potato Clade Potato Gene pool X X X X International X X X X X X X X X X National ex situ Conservation De Wet (1978). Leal and Coppens d’Eeckenbrugge (1996), Duval et al. (2001, 2003), Ruas et al. (2001). References Genet Resour Crop Evol (2012) 59:655–681 667 123 Family Cucurbitaceae Cucurbitaceae Solanaceae Crop Melon Cucumber Peppers Table 4 continued 123 Capsicum Cucumis Cucumis Genus Doubtful Doubtful Capsicum annuum L. var. glabriusculum (Dunal) Heiser et Pickersgill Capsicum chinense Jacq. Capsicum pendulum Willd. Capsicum pubescens Ruiz et Pav. Capsicum baccatum L var. pendulum (Willd.) Eshbaugh Capsicum microcarpum Cav. Doubtful Native Native Native Capsicum baccatum L. var. baccatum Capsicum frutescens L. Native Doubtful Capsicum annuum L. var. aviculare (Dierb.) D’Arcy et Eshbaugh Capsicum cumanense Fingerh. Doubtful Capsicum annuum L. Naturalized Naturalized Cucumis sativus L. TG1 Naturalized Cucumis melo L. Cucumis metuliferus Naudin ? ? ? ? ? ? TG1 TG1 TG1 TG1 TG3 TG3 Doubtful TG3 TG1 Taxon group Cucumis dipsaceus Ehrenb. ex Spach Native Status Naturalized Capsicum annuum L. var. annuum Synonymous Cucumis anguria L. Lycopersicon esculentum Mill. var. cerasiforme (Dunal) A. Gray Taxon C. pubescens complex GP2 GP2 GP2 C. baccatum complex GP2 C. annuum complex ? GP2 GP1 GP1 GP2 GP3 GP1 GP3 GP3 Gene pool C. baccatum complex ? C. annuum complex C. annuum complex C. annuum complex Section Cucumis Section Metuliferi Section Cucumis Section Aculeatosi Section Aculeatosi Taxonomic classification X X X X X X International X X National ex situ Conservation Moscone et al. (2006), Walsh and Hoot (2001), Naci and Pickersgill (2004). Deakin et al. (1971), Kho et al. (1980) References 668 Genet Resour Crop Evol (2012) 59:655–681 Family Apiaceae Arecaceae Cucurbitaceae Rutaceae Rutaceae Rutaceae Rutaceae Caricaceae Solanaceae Dioscoreaceae Crop Carrot Coconut Watermelon Lime Lemon Mandarin Orange Papaya Tobacco Yam Table 4 continued Dioscorea Nicotiana Carica Citrus Citrus Citrus Citrus Citrullus Cocos Daucus Genus Endemic Endemic Neophyte Dioscorea atrescens R.Knuth Dioscorea bolivarensis Steyerm. Dioscorea cayenensis Lam. Native Native Dioscorea amazonum Mart. ex Griseb. Dioscorea crateriflora R.Knuth Neophyte Dioscorea alata L. Archaeophyte Endemic Nicotiana tabacum L. Dioscorea abysmophila Maguire et Steyerm. Native Naturalized Neophyte Neophyte Neophyte Neophyte Naturalized Naturalized Nicotiana glauca Graham Carica papaya L. Citrus sinensis Osbeck Citrus reticulata Blanco Citrus limon (L.) Osbeck Citrus aurantiifolia (Christm.) Swingle Citrullus lanatus (Thunb.) Matsum. et Nakai Cocos nucifera L. Neophyte Native Native Status Daucus carota L. Synonymous Daucus montanus Humb. et Bonpl. ex Schult. Capsicum rhomboideum (Humb., Bonpl. et Kunth) Kuntze Taxon TG3 TG2 TG3 TG3 TG3 TG1 ? TG1 TG4 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG4 TG1 ? Taxon group Section Macrothyrsa Section Enantiophyllum Section Sarcantha Section Strutantha Section Sarcantha Section Enantiophyllum ? Section Nicotiana Section Noctiflorae Section Anisactis Section Daucus ? Taxonomic classification GP1 GP3 GP1 GP3 Gene pool X X International X X National ex situ Conservation Knuth (1924). Knapp et al. (2004), Trojak-Goluch and Berbeć (2003). Sáenz-Laı́n (1981). References Genet Resour Crop Evol (2012) 59:655–681 669 123 Family Araceae Crop Tannia Table 4 continued 123 Xanthosoma Genus Endemic Xanthosoma caulotuberculatum G.S.Bunting Native Dioscorea trifida L.f. Endemic Native Dioscorea trichanthera Gleason Xanthosoma bolivaranum G.S.Bunting Endemic Dioscorea sororopana Steyerm. Endemic Native Dioscorea polygonoides Humb. et Bonpl. ex Willd. Xanthosoma bayo G.S.Bunting Native Dioscorea meridensis Kunth Native Native Dioscorea laxiflora Mart. ex Griseb. Xanthosoma aristeguietae (G.S.Bunting) Madison Endemic Dioscorea holmioidea Maury Endemic TG3 Native Dioscorea discolor Hort.Berol. ex Kunth Xanthosoma akkermansii (G.S.Bunting) Croat TG3 Native Dioscorea decorticans C.Presl ? ? ? ? ? TG1 TG3 TG3 TG3 TG3 TG3 ? TG3 Native TG3 Taxon group Dioscorea cuspidata Humb. et Bonpl. ex Willd. Status Native Synonymous Dioscorea crotalariifolia Uline Taxon Section Macrogynodium Section Epistemon Section Hemidematostemon Section Lychnostemon Section Lychnostemon Section Cryptantha ? Section Lasiogyne Section Strutantha Section Apodostemon Section Apodostemon Taxonomic classification Gene pool International National ex situ Conservation Engler (1920). References 670 Genet Resour Crop Evol (2012) 59:655–681 Crop Family Table 4 continued Genus Endemic Endemic Native Endemic Endemic Endemic Endemic Native Native Native Xanthosoma mafaffoides G.S.Bunting Xanthosoma maroae G.S.Bunting Xanthosoma mexicanum Liebm. Xanthosoma nitidum G.S.Bunting Xanthosoma orinocense G.S.Bunting Xanthosoma pariense G.S.Bunting Xanthosoma peltatum G.S.Bunting Xanthosoma sagittifolium (L.) Schott Xanthosoma saguasense G.S.Bunting Xanthosoma striatipes (Kunth) Madison Native Native Xanthosoma longilobum G.S.Bunting Xanthosoma undipes K.Koch Native Xanthosoma helleborifolium Schott Endemic Endemic Xanthosoma exiguum G.S.Bunting Xanthosoma trilobum G.S.Bunting Endemic Xanthosoma contractum G.S.Bunting Status Native Synonymous Xanthosoma conspurcatum Schott Taxon TG2 ? ? ? TG1 ? ? ? ? TG2 ? ? ? TG4 ? ? TG4 Taxon group Section Euxanthosoma Section Euxanthosoma Section Euxanthosoma Section Acontias Section Acontias Taxonomic classification Gene pool International National ex situ Conservation References Genet Resour Crop Evol (2012) 59:655–681 671 123 Family Lauraceae Crop Avocado Table 4 continued 123 Persea Genus Native Native Persea grandiflora L.E.Kopp Persea pseudofasciculata L.E.Kopp Endemic Persea fluviatilis van der Werff Native Native Persea ferruginea Kunth Native Native Persea fendleri van der Werff Persea perseiphylla (C.K.Allen) van der Werff Native Persea fastigiata var.sericea L.E.Kopp Persea mutisii Kunth Endemic Persea fastigiata L.E.Kopp var. fastigiata Native Native Persea cuneata Meisn. Persea meridensis L.E.Kopp Native Persea croizatii van der Werff Native Endemic Persea croatii van der Werff Endemic Native Persea caerulea (Ruiz et Pav.) Mez Persea maguirei L.E.Kopp Native Persea bernardii L.E.Kopp Persea jenmanii Mez Endemic Persea areolatocostae (C.K.Allen) van der Werff Status Naturalized Synonymous Persea americana Mill. Taxon TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG4 TG1 Taxon group Subgenus Eriodaphne Subgenus Persea Taxonomic classification GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP3 GP1 Gene pool International X National ex situ Conservation Kopp (1966), Bergh and Ellstrand (1986). References 672 Genet Resour Crop Evol (2012) 59:655–681 Family Alliaceae Fabaceae Anacardiaceae Myrtaceae Crop Garlic Beans Mango Guava Table 4 continued Psidium Mangifera Phaseolus Allium Genus Naturalized Phaseolus vulgaris L. Native Native Psidium appendiculatum Kiaersk. Psidium brownianum Mart. ex DC. Native Psidium acutangulum DC. Naturalized Native Phaseolus unilobatus Pittier Mangifera indica L. Native Phaseolus tuerckheimii Donn.Sm. Phaseolus vulgaris L. var. vulgaris Vigna speciosa (Kunth) Verdc. Phaseolus speciosus Kunth Native Native Native Vigna lasiocarpa (Mart. ex Benth.) Verdc. Phaseolus pilosus Kunth Native Phaseolus erythroloma Mart. ex Benth. Phaseolus lunatus L. Doubtful Doubtful Macroptilium erythroloma Urb. Neophyte Native Phaseolus dumosus Macfad. Phaseolus vulgaris L. var. aborigineus (Burkart) Baudet Phaseolus coccineus L. Phaseolus aborigineus Burkart Allium sativum L. Native Persea subcordata Ruiz et Pav. Status Native Synonymous Persea rigens C.K.Allen Taxon ? ? ? TG1 TG1 ? TG4 TG5 TG5 TG4 TG5 TG2 TG4 TG1 TG1 TG4 TG4 Taxon group Section Phaseoli ? Section Brevilegumeni Section Paniculati Section Phaseoli Section Coccinei Section Phaseoli Taxonomic classification ? ? ? GP1 ? ? GP3 GP3 GP3 GP3 GP1 GP3 GP3 Gene pool X X X X X International X X National ex situ Conservation Freytag and Debouck (2002), Debouck (1999), Delgado-Salinas et al. (1999, 2006). References Genet Resour Crop Evol (2012) 59:655–681 673 123 Family Sterculiaceae Crop Cacao Table 4 continued 123 Theobroma Genus Native Native Native Native Native Theobroma cacao L. subsp. sphaerocarpum (A. Chev.) Cuatrec. Theobroma grandiflorum (Willd. ex Spreng.) Schum. Theobroma microcarpum Mart. Theobroma obovatum Klotzsch ex Bernoulli Theobroma speciosum Willd. Native Psidium striatulum DC. Native Native Psidium sartorianum (O.Berg) Nied. Theobroma cacao L. subsp. cacao Native Psidium salutare (Kunth) O.Berg Native Native Psidium maribense Mart. ex DC. Theobroma bicolor Humb. et Bonpl. Native Psidium laruotteanum Cambess. TG4 TG4 TG4 TG4 TG1 TG1 TG4 ? ? ? ? ? TG1 ? Naturalized Native ? Psidium guineense Sw. Native Psidium friedrichsthalianum Nied. ? ? Taxon group Psidium guajava L. Native Psidium densicomum Mart. ex DC. Status Native Synonymous Psidium cinereum Mart. ex DC. Taxon Section Oreanthes Section Glossopetalum Section Telmatocarpus Section Glossopetalum Section Theobroma Section Theobroma Section Rhytidocarpus Taxonomic classification ? ? GP2 GP2 GP1 GP1 ? ? ? ? ? ? ? ? ? ? Gene pool X International X X X National ex situ Conservation Cuatrecasas (1964), Rondón and Cumana (2005), Sousa et al. (2004). References 674 Genet Resour Crop Evol (2012) 59:655–681 Family Pedaliaceae Vitaceae Brassicaceae Brassicaceae Asteraceae Malvaceae Rosaceae Rosaceae Crop Sesame Grape Cabbage Cauliflower Lettuce Cotton Strawberry Peach Table 4 continued Prunus Fragaria Gossypium Lactuca Brassica Brassica Vitis Sesamum Genus Native Native Neophyte Native Prunus myrtifolia (L.) Urb. Prunus persica (L.) Batsch Prunus pleuradenia Griseb. Doubtful Prunus integrifolia Walp. Prunus moritziana Koehne Endemic Prunus espinozana C.L.Li Native Native Prunus amplifolia Pilg. Prunus lichoana G.Aymard Native Prunus accumulans (Koehne) C.L.Li et G.Aymard Prunus recurviflora Koehne Naturalized Naturalized Gossypium hirsutum L. Fragaria vesca L. Native Neophyte Neophyte Gossypium barbadense L. Lactuca sativa L. Brassica oleracea L. var. botrytis L. Neophyte Neophyte Brassica oleracea L. var. capitata L. Vitis vinifera L. Naturalized Native Status Doubtful Potentilla vesca Scop. Synonymous Vitis tiliifolia Humb. et Bonpl. ex Roem. et Schult. Sesamum orientale L. Theobroma subincanum Mart. Taxon ? TG1 TG4 ? TG4 ? TG4 TG4 TG4 TG1 TG1 TG1 TG1 TG1 TG1 TG1 TG3 TG1 TG4 Taxon group ? Subgenus Laurocerasus ? Subgenus Laurocerasus Subgenus Laurocerasus Subgenus Laurocerasus Subgenus Laurocerasus Subgenus Vitis Subgenus Vitis. Section Glossopetalum Taxonomic classification ? ? ? ? ? GP2 ? Gene pool X X X International X X National ex situ Conservation Li and Aymard (1997), Lersten and Horner (2000), Wen et al. (2008). References Genet Resour Crop Evol (2012) 59:655–681 675 123 123 Chenopodiaceae Convolvulaceae Fabaceae Agavaceae Fabaceae Beet Sweet Potato Pigeonpea Sisal Soya Family Crop Table 4 continued Glycine Agave Cajanus Ipomoea Beta Genus Doubtful Native Native Ipomoea ramosissima Choisy Ipomoea tiliacea (Willd.) Choisy Ipomoea trifida G.Don Agave rigida Mill. Native Ipomoea longeramosa Choisy Glycine max (L.) Merr. Neophyte Native Introduced Agave sisalana Perrine ex Engelm. Agave vivipara L. TG3 Native Agave cocui Trel. var. laguayrensis Hummelinck TG1 TG3 TG1 TG3 Native TG1 TG2 TG2 TG2 TG2 TG2 TG1 TG1 TG4 ? ? TG4 Taxon group Agave cocui Trel. var. cocui Neophyte Native Ipomoea cordatotriloba Dennst. Cajanus cajan (L.) Millsp. Native Ipomoea batatas (L.) Lam. Neophyte Endemic Prunus wurdackii C.L.Li Beta vulgaris L. Native Prunus urotaenia Koehne Doubtful Native Prunus serotina subsp. capuli (Cav.) McVaugh; Prunus capulli Cav. Prunus salicifolia Kunth Status Prunus skutchii I.M.Johnst. Synonymous Taxon Subgenus Agave Subgenus Agave Subgenus Agave Subgenus Agave Section Eriospermum Section Eriospermum Section Eriospermum Section Eriospermum Section Eriospermum Section Eriospermum Subgenus Laurocerasus ? ? Subgenus Padus Taxonomic classification ? ? ? ? GP2 ? GP3 ? GP3 GP1 ? Gene pool X X X X International X X X National ex situ Conservation Gentry (1982). Austin and Huáman (1996), Dı́az et al. (1996), Huang et al. (2002), Rajapakse et al. (2004), Srisuwan et al. (2006). References 676 Genet Resour Crop Evol (2012) 59:655–681 Poaceae Cucurbitaceae Arecaceae Chenopodiaceae Fabaceae Pumpkins, Squash Palm Oil Spinach Cowpea Fabaceae Peas Apiaceae Asteraceae Sunflower Arracacha Fabaceae Peanut Wheat Family Crop Table 4 continued Vigna Spinacia Elaeis Cucurbita Arracacia Triticum Pisum Helianthus Arachis Genus Archaeophyte Archaeophyte Cucurbita moschata Duchesne Cucurbita pepo L. Vigna juruana (Harms) Verdc. Native TG4 TG4 Native Vigna candida (Vell.) Maréchal, Mascherpa et Stainier TG1 TG4 Native Neophyte TG1 TG1 TG1 TG1 TG1 TG1 ? ? ? TG1 TG1 TG1 ? TG4 TG1 Taxon group Vigna adenantha (G.Mey.) Maréchal, Mascherpa et Stainier Spinacia oleracea L. Neophyte Archaeophyte Cucurbita maxima Duch. ex Lam. Elaeis guineensis Jacq. Archaeophyte Doubtful Arracacia xanthorrhiza Bancr. Cucurbita ficifolia Bouché Native Arracacia vaginata J.M.Coult. et Rose Native Neophyte Native Arracacia elata H. Wolff Neophyte Arracacia tillettii Constance et Affolter Arracacia pennellii Constance Triticum aestivum L. Pisum sativum L. Neophyte Doubtful Arachis prostrata Benth. Helianthus annuus L. Native Arachis pintoi Krapov. et W.C.Greg. Status Doubtful Synonymous Arachis hypogaea L. Taxon Subgenus Lasiospron Subgenus Sigmoidotropis, Section Sigmoidotropis Subgenus Sigmoidotropis, Section Leptospron Section Caulorrhizae Section Arachis Taxonomic classification ? ? ? GP1 ? ? ? GP3 GP1 Gene pool X X X X X International X X X X National ex situ Conservation Marechal et al. (1978), Sonnante et al. (1996). Krapovickas and Gregory (1994), Gimenes et al. (2002). References Genet Resour Crop Evol (2012) 59:655–681 677 123 Crop Family Table 4 continued Genus 123 Native Native Doubtful Native Neophyte Doubtful Vigna linearis (Kunth) Maréchal, Mascherpa et Stainier Vigna longifolia (Benth.) Verdc. Vigna luteola (Jacq.) Benth. Vigna peduncularis Fawc. et Rendle Vigna unguiculata (L.) Walp. Vigna vexillata (L.) A.Rich. Status Native Synonymous Vigna lasiocarpa (Benth.) Verdc. Taxon TG4 TG1 TG4 TG3 TG4 TG4 TG4 Taxon group Subgenus Plectotropis, Section Plectrotropis Subgenus Vigna, Section Catiang Subgenus Sigmoidotropis, Section Pedunculares Subgenus Vigna, Section Vigna Subgenus Lasiospron Subgenus Sigmoidotropis, Section Caracallae Subgenus Lasiospron Taxonomic classification GP3 GP1 ? ? ? ? ? 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