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Genet Resour Crop Evol DOI 10.1007/s10722-012-9923-6 RESEARCH ARTICLE National inventory and prioritization of crop wild relatives: case study for Benin Rodrigue Idohou • Achille Ephrem Assogbadjo • Belarmain Fandohan • Gerard Nounagnon Gouwakinnou Romain Lucas Glele Kakai • Brice Sinsin • Nigel Maxted • Received: 11 June 2012 / Accepted: 10 October 2012 Ó Springer Science+Business Media Dordrecht 2012 Abstract Species prioritization is a crucial step in any development of conservation strategy, especially for crop wild relatives (CWR), since financial resources are generally limited. This study aimed at: assessing the biodiversity of crop wild relatives in Benin and identifying priority species for active conservation. Data were collected through literature review to establish an exhaustive list of CWR in Benin. Eight prioritization criteria and different prioritization systems were used. The top 50 species obtained by each of these methods were identified and twenty final top CWR were shortlisted as those occurring as priority across methods. A total of 266 plant species belonging to 65 genera and 36 families were identified. The most represented are: Cyperaceae (12.50 %), Leguminosae-Papilionoideae (11.87 %), R. Idohou  A. E. Assogbadjo  B. Fandohan (&)  G. N. Gouwakinnou  R. L. Glele Kakai  B. Sinsin Laboratory of Applied Ecology, University of Abomey-Calavi, 01 BP 526 Cotonou, Benin e-mail: bfandohan@gmail.com; belarmain.fandohan@unep-iemp.org B. Fandohan International Ecosystem Management Partnership, United Nations Environment Programme, c/o Institute of Geography and Natural Resources Research, Chinese Academy of Sciences, No. 11A Datun Rd., Beijing 100101, China N. Maxted School of Biosciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK Convolvulaceae (11.25 %), Poaceae (10.31 %), Asteraceae (7.81 %), Solanaceae (6.87 %) and Dioscoreaceae (5.31 %). Among the 20 species of highest priority for conservation, Manihot glaziovii Müll. Arg. and Piper guineense Schumach. et Thonn., appeared as the most represented species on top of the list. Keywords Biodiversity  Conservation  Crop wild relatives  Threat  West Africa Introduction Millions of the world’s poor rely for a large part of their income/food intake on a wide variety of indigenous edible plants to sustain their livelihood. This is particularly the case in Sub-Saharan Africa, where over 70 % of the people reside in rural areas and use plant resources to meet their routine needs (Cavendish 2000; Mahapatra et al. 2005). Crop wild relatives (CWR) are wild plant taxa more or less closely related to species of direct socio-economic importance including food, fodder and forage crops, medicinal plants, condiments, ornamental and forestry species, as well as those related to crops used for industrial purposes such as oils and fibres (Maxted et al. 2007). CWR include the progenitors of crops as well as other species more or less closely related to them, and have been undeniably beneficial to modern agriculture, providing plant breeders with a broad pool of 123 Genet Resour Crop Evol potentially useful genetic resources (Hajjar and Hodgkin 2007). Hence, CWR represent a tangible resource of actual or potential economic benefit for humankind as they have contributed significantly to improvement of food production. Prescott-Allen and Prescott-Allen (1986) calculated that the yield and quality contribution by CWR to the US-grown or imported crops was over US$350 million a year. Phillips and Meilleur (1998) noted that losses of rare wild plants represent a substantial economic loss to agriculture, estimating that the endangered food crop relatives have a worth of about US$10 billion annually in wholesale farm values. Pimentel et al. (1997), for their part, estimate a global value of CWR at US$115 billion per annum. Like for the rest of biodiversity, CWR are threatened by mismanagement of landscape. Furthermore, habitat fragmentation, climate change and agricultural intensification put at risk the CWR and traditional cultivars. Accordingly, it is urgent to take actions to reduce genetic erosion or species extinction. FAO (2009) reported a significant increase in the number of CWR inventories. However, in Sub-Saharan Africa, there is a recurrent lack of knowledge regarding the breadth and/or potential use of CWR diversity. Presently, inventories are lacking for most countries and CWR diversity is largely uncharacterized or un-evaluated and not systematically conserved. With so much CWR diversity, it’s necessary to inventory the diversity of these species and establish priorities for conservation (Kell et al. 2008). Prioritization for conservation can be undertaken at different levels: species, ecosystem, etc. (Brehm et al. 2010). A method of prioritizing at species level is preferable because it allows conservationists to know which taxa should be primarily targeted for conservation, which are not priorities, and which have insufficient information to know whether they are priorities for conservation or not (Brehm et al. 2010). Numerous methods for setting species’ priorities have been developed over time (Rabinowitz et al. 1986; Coates and Atkins 2001). There has been considerable debate over which criteria should be considered when prioritizing species for conservation (see Maxted et al. 1997). Recently, Brehm et al. (2010) proposed various criteria and different prioritization schemes. This study aimed at developing an innovative prioritization scheme making use of the readily available data and to identify priority CWR and Wild Harvested Plants (WHP) for conservation 123 in Portugal (Brehm et al. 2010). This new scheme was applied in the current study. The objective of the present study was to create a national inventory of wild relatives of priority crops in Benin and highlight priority species as a useful case study for the establishment of Phytogenetic Genetic Resource (PGR) conservation strategies. Materials and methods Study area The study was carried out in the Republic of Benin, located between 6° and 12°500 N and 1° and 3°400 E in West Africa (Adomou 2005). Three large chorological climatic zones (Sudanian, Sudano-Guinean and Guineo-Congolian zones; Fig. 4) embody ten phytogeographic zones, which are Atacora chain, Bassila, coastal, Mekrou-Pendjari, North-Borgou, Plateau, Pobè, South-Borgou, Valley of Oueme and Zou (Adomou 2005). Vegetation in Benin comprises semi-deciduous rain forest, swamp forest, gallery forest, dense dry forest, open forest, woodland savanna and tree and shrub savanna (Adomou et al. 2010). The flora is estimated at 2,807 species of plants divided into 1,129 genera and 185 families (Akoègninou et al. 2006). The most diversified families in terms of number of species are: Leguminosae (14.8 %), Poaceae (9.3 %), Rubiaceae and Cyperaceae (5 % each), Asteraceae (4.6 %) and Euphorbiaceae (4.3 %). In the south of the country, mean monthly temperatures oscillate between 26 and 28 °C while in the north they are generally above 35 °C, and in some places they average out at 40 °C (Adomou 2005). Rainfall varies from 900 to 1,400 mm per year according to West-East and South-North gradient. Rainfall distribution shows two types of climates. In the south, the climate is tropical humid (Subequatorial or Guinean) with two rainfall maxima in April–July and September–October. In the North, the climate is tropical sub-humid dry to arid from 8°N northwards, with one maximum in June (Adomou 2005). In 2009, agriculture contributed 33.2 % to the Gross National Product (GNP) of Benin. The main crops are cereals (maize, rice, sorghum, etc.), legumes (cowpea, peanuts, etc.), tubers (manioc, yam, etc.), market garden products (lettuce, cabbage, etc.) and industrial crops (cotton, cashew nuts, pineapple, etc.) (MAEP 2010). Genet Resour Crop Evol Inventory of crop wild relatives in Benin The starting point for preparing a national CWR conservation strategy is the generation of a national CWR inventory. Here, we recorded Taxa selected on the basis of their closeness to priority crops of Benin, using the ‘‘taxonomic group’’ concept of Maxted et al. (2006). Since the flora of Benin 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 was carried out manually. This approach was recently successfully implemented for Bhutan (Tamang 2004), the Seychelles (Antoine 2004), and Venezuela (Chiara and Crespo 2012). It consisted, first, of making an exhaustive census of all the cultivated crops in Benin. Data were collected from libraries (public and private), agricultural extension services, research institutes, laboratories, and botanical garden of the University of Abomey Calavi. Then a list of the cultivated crops was matched with the existing floras (Flora of West Tropical Africa, Analytic flora of Benin and Biodiversity atlas for West Africa) to select the species in the same genus as the crops. The CWR inventory was then compiled from the species found in the same genus as the cultivated plant and that occur in the national flora. Records for each genus included in the CWR inventory were also taken from databases of major herbaria and gene banks worldwide, which were accessed online through the Global Biodiversity Information Facility, GBIF (www.gbif.net). Setting priorities for CWR conservation The process of establishing priorities is a first step in any conservation strategy (Maxted and Kell 2009a, b). The criteria used for the prioritization are those proposed by Brehm et al. (2010) and described as follows: 1. 2. Native status. Since the inventory has both native and introduced species, priorities were given to native (indigenous) taxa; Economic value. The CWR have their main potential application in genetic improvement of existing cultivars. Therefore, the economic importance of the related crop species is one good indicator of their value as a wild relative. 3. 4. 5. 6. 7. 8. Ethnobotanical value. This was assessed through local knowledge on the species uses. Priority was given to the species having a high importance for local communities. Global distribution. Priority increases with the more a restricted distribution, therefore, nationally- or regionally-restricted species (or endemics) were given higher priority than species occurring world-wide. National distribution. National distribution was considered here as an indicator of rarity. A species occurring in a few provinces was considered rarer than a species occurring throughout the country. In-situ and ex-situ conservation status. Before a taxon can be given high priority for conservation, current conservation activities relating to it should be reviewed. If sufficient genetic diversity is already being conserved in situ and/or ex-situ, additional conservation efforts may not be justified, and resources should focus on those species that are not being conserved. Legislation. A species under any kind of legislation requires conservation attention because national governments are responsible for protecting them. Threat assessment. The IUCN Red List threat status is probably the most used criterion for determining conservation priority. Endangered species received greater attention than those that are not. Four different methods of combining the above mentioned eight criteria were used as described in Brehm et al. (2010): point scoring procedure (PSP), point scoring procedure with weighting (PSPW), compound ranking system (CRS), and binomial ranking system (BRS). In the PSP, a series of scores for multiple criteria was assigned to each species, with the highest number always indicating the highest priority. For example, the overall score for each CWR was obtained by the sum of all individual criteria: (native status ? economic value ? ethnobotanical value ? global distribution ? national distribution ? conservation ? legislation ? threatened status). Then, higher scores indicate greater conservation concern. The PSPW is very similar to the PSP with the difference that to each criterion a particular weight is given. The CRS uses individual criteria ranking positions (not scores as in 123 Genet Resour Crop Evol Fig. 1 Methodology used for establishing conservation priorities for CWR in Benin, Adapted from Brehm et al. (2010) PSP), which are then combined in order to obtain a compound rank for each of the species and for each of the major criteria. The BRS is based on a series of Yes/No questions. A ‘‘Yes’’ answer is always a higher priority than a ‘‘No’’ answer. For both CRS and BRS, three types of ranking were used as described in Brehm et al. (2010). Then, the top 50 species were obtained for each method: PSP, PSPW, CRS (CRS1, CRS2 and CRS3) and BRS (BRS1, BRS2 and BRS3). The number of times each top 50 species occurred in the different sub lists was recorded. The priority species were those that occurred most commonly in individual lists (For further information see Brehm et al. 2010) (Fig. 1). Results Taxonomic diversity of the crop wild relatives Matching the list of the cultivated crops with the species present in the flora of Benin enabled generation of a CWR list for Benin. This original inventory contained 266 species belonging to 65 genera (Figs. 2, 3) and 36 families, of which the most represented were: Cyperaceae 123 (12.50 %), Leguminosae-Papilionoideae (11.87 %), Convolvulaceae (11.25 %), Poaceae (10.31 %), Asteraceae (7.81 %), Solanaceae (6.87 %) and Dioscoreaceae (5.31 %). Among the families, 67.57 % were represented by one genus; 18.82 % by 2 genera, while 13.51 % were represented by more than 2 genera. Note also that some crops (42.59 %) have no wild relatives in Benin, these are largely exotic crops introduced from outside of Africa. Priority CWR for active conservation Use of different methods of combining the data sets generated different lists of priority taxa for CWR. The PSP method yielded a list of priority species for conservation, with the Dioscoreaceae family widely represented (26 %), followed by the LeguminosaePapilionoideae (22 %). The two most prioritized species were: Dioscorea praehensilis Benth., (Dioscoreaceae) and Manihot glaziovii Müll. Arg. (Euphorbiaceae). For the PSPW method, Dioscorea burkilliana J. Miege (Dioscoreaceae) appeared as the highest priority. This family was widely represented (10 species) and four of these species were among the top 5 priority species for conservation. It was followed Genet Resour Crop Evol Fig. 2 Diversity of genera among the CWR’s family Fig. 3 Diversity of species among CWR’s family by the Convolvulaceae and Leguminosae-Papilionoideae, which have, respectively, 9 and 8 priority species for conservation. Also, the CRS (with variants) generated a list of 50 species for active conservation among which D. burkilliana (Dioscoreaceae), D. praehensilis (Dioscoreaceae) and M. glaziovii (Euphorbiaceae) were the most prioritized species for conservation. Among the families, Dioscoreaceae is the most diverse family in the group (containing 26 % of prioritized species). Each variant of the BRS generated a list of 50 priority species for conservation among which the most represented are Ipomoea beninensis Akoègninou, Lisowski et Sinsin (Convolvulaceae), M. glaziovii (Euphorbiaceae), Abelmoschus moschatus Medik. (Malvaceae) and Piper guineense Schumach. et Thonn. (Piperaceae). The Poaceae family is the most represented group (26 %), followed by the families Convolvulaceae (20 %), and Leguminosae-Papilionoideae (12 %). These three families constitute 58 % of priority species for conservation. Finally, appearance of the top 50 species on the various lists (obtained by the methods of prioritization) was noted. These lists were used to identify the first 20 species of highest priority for conservation (Table 1). The appearance of the species on each of the list is described below (Table 1). This table shows that 2 species, M. glaziovii and P. guineense appeared as the most important for conservation in Benin. These are followed by Corchorus trilocularis L., D. burkilliana, D. praehensilis, Dioscorea togoensis Knuth, Blighia welwitschii (Hiern) Radlk., Pennisetum glaucum (L.) R. Br. subsp. violaceum (Lam.) Rich, Pennisetum macrourum Trin, I. beninensis, Sesamum alatum Thonn., Cajanus kerstingii Harms, Celosia bonnivairii Schinz, Cucumis prophetarum L., Cyperus papyrus L., 123 Genet Resour Crop Evol Table 1 List of the 20 priority CWR for active conservation in Benin obtained using the methodology combining four different priority setting methods (with variants) Species PSP PSPW CRS1 CRS2 Manihot glaziovii Müll. Arg. X X X X Piper guineense Schumach. et Thonn. X X X X X X Corchorus trilocularis L. Dioscorea burkilliana J. Miège X X X Dioscorea praehensilis Benth. X X X Dioscorea togoensis Knuth X X X CRS3 X X X BRS1 BRS2 BRS3 Total X X X 7 X X X 7 X X X X X X X X X Blighia welwitschii (Hiern) Radlk. X X X X X Pennisetum glaucum (L.) R.Br. subsp. violaceum (Lam.) Rich. X X X X X Pennisetum macrourum Trin. X X X X X X X X X X X X X X Ipomoea beninensis Akoègninou, Lisowski et Sinsin X Sesamum alatum Thonn. Cajanus kerstingii Harms X X Celosia bonnivairii Schinz X X 6 6 X X X 6 6 6 6 X 6 X X X 6 X X X X 6 6 X X X X X X 6 Cucumis prophetarum L. X X X Cyperus papyrus L. X X X X X X Dioscorea preussii Pax X X X X X X 6 Dioscorea hirtiflora Benth. X X X X X 6 X Dioscorea mangenotiana J.Miège X X Jatropha neriifolia Müll. Arg. X X Vigna juruana (Harms) Verdc. X X X X X X 6 6 X X X 6 X X X 6 X X 6 PSP, point scoring procedure; PSPW, point scoring procedure with weighing; CRS, compound ranking system; BRS, binomial ranking system; 1, 2, 3 = variants of the methods. For further information see Brehm et al. (2010) Dioscorea preussii Pax, Dioscorea hirtiflora Benth., Dioscorea mangenotiana J. Miège, Jatropha neriifolia Müll. Arg. and Vigna juruana (Harms) Verdc. Among the species, 55 % are confined to one phytodistrict and just 20 % are found in four phytodistricts (Table 2). This shows the restricted distribution of the abovementioned species, which should therefore be taken into account for active conservation (Fig. 4). Discussion This study aimed at establishing the first national CWR of a sub-Saharan country and setting priority for conservation of the CWR in Benin. There are 266 CWR species in Benin (about 10 % of the floristic diversity). This diversity seems low compared to the one for other countries such as the United Kingdom (65 % of the floristic diversity) (Maxted et al. 2007) 123 and Portugal (75 % of the floristic diversity) (Brehm et al. 2007) but high compared to the one for Venezuela (about 2 % of the floristic diversity) (Chiara and Crespo 2012). The observation above is explained by the fact that the method used to produce the inventory was based on native food crop gene pools, and that many Benin crops (corn, cashew nuts, pineapple, peanuts, papaya, citrus fruits, spinach, etc.) are of exotic origin (MAEP 2010). Therefore, there is a small proportion of native crops and a corresponding native CWR diversity. Much attention has historically been given to plant conservation in Benin (Codjia et al. 2003; Vodouhê et al. 2010; N’Danikou et al. 2011) and priority plant species have previously been included in conservation plans, but previously none has focused systematically on CWR diversity. The only previous attempt at active CWR conservation tried to conserve some families of wild plants especially Dioscoreaceae and Euphorbiaceae (Adomou et al. 2010). Current results Unknown Medicinal and fodder uses Ipomoea beninensis Akoègninou, Lisowski et Sinsin Sesamum alatum Thonn. Unknown Unknown Unknown Unknown Unknown Cyperus papyrus L. Dioscorea preussii Pax Dioscorea hirtiflora Benth. Dioscorea mangenotiana J. Miège Jatropha neriifolia Müll. Arg. Vigna juruana (Harms) Verdc. CR, critically endangered; EN, endangered; NE, not evaluated Medicinal use and strong poison Unknown Cucumis prophetarum L. Unknown Fodder Pennisetum macrourum Trin. Unknown Fodder Pennisetum glaucum (L.) R.Br. subsp. violaceum (Lam.) Rich. Celosia bonnivairii Schinz Food and medicinal use Blighia welwitschii (Hiern) Radlk. Cajanus kerstingii Harms Food and medicinal use Medicinal use Food and craft use Medicinal use Corchorus trilocularis L. Dioscorea burkilliana J. Miège Dioscorea togoensis Knuth Food, insecticide and medicinal uses Dioscorea praehensilis Benth. 3 phytodistricts Rubber production, ornamental and medicinal use Manihot glaziovii Müll. Arg. Piper guineense Schumach. et Thonn. 1 phytodistrict 1 phytodistrict 2 phytodistricts 4 phytodistricts 2 phytodistricts 1 phytodistrict 1 phytodistrict 1 phytodistrict 4 phytodistricts 1 phytodistrict 4 phytodistricts 1 phytodistrict 1 phytodistrict 1 phytodistrict 4 phytodistricts 2 phytodistricts 1 phytodistrict 1 phytodistrict 2 phytodistricts National distribution Ethnobotanical uses Taxa Table 2 Ethnobotanical uses, national distribution and threats of the most prioritized species NE NE NE NE NE NE NE NE NE NE EN NE NE NE NE NE NE NE EN CR Threats assessment (Adomou et al. 2010) NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE Threats assessment (IUCN (2011)) Genet Resour Crop Evol 123 Genet Resour Crop Evol Fig. 4 Map of the Republic of Benin showing the phytogeographical districts and the occurrence of the most prioritized CWR concur with this attempt in finding the latter families to be among the most prioritized for active conservation in Benin. Immediate conservation action to save the CWR of Benin would be the only way to ensure the availability of these plants for future generations. As shown in Table 2, most of the priority CWR species are currently used by local communities. However, these 123 plants are faced with several threats, such as repeated clearing and wildfires due to agricultural activities, which have resulted in serious degradation of natural forest reserves. Further threat comes from ongoing strategic plan for the agricultural sector in Benin through which, the demand for new land for agriculture (land clearing) is an additional threat to the wild plant species in general, and CWR in particular. In Genet Resour Crop Evol addition, urbanization (which increase the demand on fuel wood, charcoal, building materials, medicinal, etc.), the problems of invasive species, and climate change all result in rapid declines of these species. To date, a detailed threat assessment using IUCN Red List Criteria has only been attempted for a few species (Table 2), yet most species facing human harvesting pressure are often exposed to decline. Therefore, there is a need to undertake ecogeographic, distribution, gap analysis studies and the impact of climate change on the conservation of each of the twenty priority species. As available financial resources are not enough to conserve all species, focus can at least be made on the first two in the list (M. glaziovii and P. guineense). Active conservation of CWR in Benin should be a priority as the country occupies an important part in the ‘Dahomey Gap’ which is a break in the dense forest on the coast of West Africa (Akoègninou et al. 2006). Thus, the flora of Benin shares many species with the flora of the countries covered by dense forest (Liberia, Ivory Coast, Ghana and Nigeria). Consequently, only three new species from Benin (Thunbergia atacorensis Akoègninou et Lisowski (Acanthaceae), I. beninensis Akoègninou, Lisowski et Sinsin (Convolvulaceae) and Kyllinga beninensis Samain, Reynders et Goetghebeur (Cyperaceae) are recently described by science and may be considered as the only endemics (Adomou et al. 2010). It is therefore clear that the flora of Benin is very poor in endemic species. Consequently, conservation of the CWR in Benin will benefit other West African countries with which Benin shares the same species. Such conservation will help to maintain the genetic variability contained in most cultivars to meet future demands. Furthermore, CWR provide traits such as disease resistance, tolerance to extreme temperatures, salt tolerance and drought resistance which are useful for strengthening the genetic make-up of the grown crops. An active conservation of PGR, particularly CWR diversity, requires the establishment of priority within species (Maxted et al. 2006). But there is no single method to develop effective strategies for biodiversity conservation (Maxted et al. 2006). Methodologically, our approach differs from that used by Lawrence et al. (2005), Maraseni (2008) and Vodouhê et al. (2009) who identified the most important Non-Timber Forest Products in Cameroun, Nepal, and northern Benin, respectively. N’Danikou et al. (2011) used an independent scoring of species in value and conservation criteria developed by the community. As such, these authors argued that successful management strategies will then need to consider the criteria that communities use in their species valuation, because strategies that operate exclusively with market-based or conservation-based criteria are likely to overlook communities’ interests. All these methodologies differ from the one used in this study because they do not combine the different criteria (importance and ecological criteria) in different methods. Although N’Danikou et al. (2011) recommend the use of the scoring method for studies of this type, the result obtained by one method should be compared to the one of the others to bring out priority species for conservation. Yet, each of these methods used in the current study can be updated whenever new information is collected. As data were not always available for the IUCN status of species, legislation and conservation strategies, it is strongly recommended that priorities be reassessed and refined when more detailed information is available. Brehm et al. (2010) argue that prioritization of species should be a dynamic process and as noted by UNEP (1995), the success of any method of prioritization will depend, in large part, on the inclusion of results in conservation activities, and especially the support of the international community (hardware and financial) in the preservation of biodiversity. This study highlights CWR diversity in Benin and the priority species for conservation. In view of setting appropriate conservation strategies, further steps should include thorough studies on representation, management and ecological gap analyses, state of traditional management practices and the impacts of climate change on each priority CWR. Acknowledgments We are very grateful to the members of the laboratory of Applied Ecology (University of Abomey Calavi) who were actively involved in the study. We are indebted to the Explorers Club Granting Fund which provided us with financial assistance for the field work. Our appreciation also goes to the Ministry of Agriculture and to the national research institute for their assistance. We thank all reviewers and Ayub Oduor for comments on a previous version. Appendix See Table 3. 123 Genet Resour Crop Evol Table 3 List of the cultivated plants and their relatives in Benin Family Genera Crop Crop wild relatives Alliaceae Allium Allium cepa L. cv. group Aggregatum – Allium cepa L. cv. group Common Onion Allium sativum L. Amaranthaceae Amaranthus Amaranthus cruentus L. Amaranthus blitum L. Amaranthus dubius Mart. ex TheIl. Amaranthus dubius Mart. ex Thell. Amaranthus graecizans L. Amaranthus spinosus L. Celosia argentea L. var. argentea (L.) Schinz Celosia bonnivairii Schinz Celosia argentea L. var. cristata (L.) Kuntze Celosia laxa Schumach. et Thonn. Amaranthus viridis L. Celosia Celosia leptostachya Benth. Celosia trigyna L. Anacardiaceae Apiaceae Apocynaceae Anacardium Anacardium occidentale L. – Mangifera Mangifera indica L. – Centella Centella asiatica (L.) Urb. – Daucus Daucus carota L. subsp. sativus (Hoffm.) Arcang. – Petroselinum Petroselinum crispum (Mill.) A. W. Hill – Nerium Nerium oleander L. – Araceae Colocasia Colocasia esculenta (L.) Schott – Arecaceae Cocos Cocos nucifera L. – Asteraceae Elaeis Elaeis guineensis Jacq. – Lactuca Lactuca sativa L. Lactuca inermis Forssk. var. inermis Vernonia Vernonia amygdalina Delile Vernonia adoensis Sch. Bip. ex Walp. Vernonia ambigua Kotschy et Peyr. Vernonia camporum A. Chev. Vernonia chthonocephala O. Hoffm. Vernonia cinerea (L.) Less. Vernonia colorata (Willd.) Drake Vernonia conferta Benth. Vernonia galamensis (Cass.) Less. Vernonia gerberiformis Oliv. et Hiern subsp. macrocyanus (O. Hoffm.) C. Jeffrey Vernonia glaberrima Welw. ex O. Hoffm. Vernonia guineensis Benth. var. guineensis Vernonia guineensis Benth. var. procera (O. Hoffm.) C.D. Adams Vernonia klingii O. Hoffm. et Muschl. Vernonia migeodii S. Moore Vernonia nestor S. Moore Vernonia nigritiana Olı̈v. et Hiern Vernonia perrottetii Sch. Bip. ex Walp. Vernonia poskeana Vatke et Hildebr. var. elegantula (Hutch. et Dalziel) C. D. Adams Vernonia pumila Kotschy et Peyr. Vernonia purpurea Sch. Bip. ex Walp. Vernonia stellulifera (Benth.) C. Jeffrey Vernonia undulata Oliv. et Hiern Basellaceae Basella Basella alba L. – Bombacaceae Pachira Pachira aquatica Aubl. – Pachira glabra Pasquale Pachira sessilis Benth. 123 Genet Resour Crop Evol Table 3 continued Family Genera Crop Crop wild relatives Bromeliaceae Ananas Ananas comosus (L.) Merr. – Burseraceae Raphanus Raphanus sativus L. – Caricaceae Carica Carica papaya L. – Combretaceae Terminalia Terminalia ivorensis A. Chev. Terminalia avicennioides Guill. et Perr. Terminalia mantaly H. Perrier Terminalia catappa L. Terminalia superba Engl. et Diels Terminalia glaucescens Planch. ex Benth. Terminalia laxiflora Engl. Terminalia macroptera Guill. et Perr. Terminalia mollis M. A. Lawson Convolvulaceae Ipomoea Ipomoea batatas (L.) Lam. Ipomoea acanthocarpa (Hochst. et Choisy) Ascherson et Schweinf. Ipomoea alba L. Ipomoea aquatica Forssk. Ipomoea argentaurata Hallier f. Ipomoea asarifolia (Desr.) Roem. et Schult. Ipomoea barteri Baker var. barteri Ipomoea beninensis Akoègninou, Lisowski et Sinsin Ipomoea blepharophylla Hallier f. Ipomoea cairica (L.) Sweet Ipomoea chrysochaetia Hallier f. var. velutipes (Welw. ex Rendle) Lejoly et Lisowski Ipomoea coptica (L.) Roth. ex Roem. et Schult. Ipomoea coscinosperma Hochst. ex Choisy in DC. Ipomoea eriocarpa R. Br. Ipomoea fistulosa Mart. ex Choisy Ipomoea fulvicaulis (Choisy) Hallier f. Ipomoea hederifolia L. Ipomoea heterotricha F. Didr. Ipomoea imperati (Vahl) Griseb. Ipomoea indica (Burm.f.) Merr. Ipomoea involucrata P. Beauv. Ipomoea kotschyana Hochst. ex Choisy Ipomoea marginata (Desr.) Verdc. Ipomoea mauritiana Jacq. Ipomoea nil (L.) Roth Ipomoea obscura (L.) Ker-Gawl. Ipomoea pes-caprae (L.) R. Br. Ipomoea pes-tigridis L. var. pes-tigridis Ipomoea pyrophila A. Chev. Ipomoea quamoclit L. Ipomoea rubens Choisy Ipomoea triloba L. Ipomoea turbinata Lag. Ipomoea vagans Baker Ipomoea verticillata Forssk. Ipomoea welwitschii Hallier f. 123 Genet Resour Crop Evol Table 3 continued Family Cucurbitaceae Genera Crop Crop wild relatives Citrullus Citrullus lanatus (Thunb.) Matsum. et Nakai Citrullus colocynthis (L.) Schrad. Cucumeropsis Cucumeropsis mannii Naud. Cucumeropsis edulis (Hook. f.) Cogn. Cucumis Cucumis sativus L. Cucumis melo L. subsp. agrestis Cucumis metuliferus E. Mey. ex Naudin Cucumis prophetarum L. Cucurbita Cyperaceae Cucurbita maxima Duchesne Cucurbita moschata Duchesne Cucurbita pepo L. – Lagenaria Lagenaria siceraria (Molina) Standl. – Telfairia Telfairia occidentalis Hook. f. – Cyperus Cyperus esculentus L. Cyperus alopecuroides Rottb. Cyperus amabilis Vahl Cyperus articulatus L. Cyperus buchholzii Boeck. Cyperus compressus L. Cyperus congensis C. B. Clarke Cyperus crassipes Vahl Cyperus cuspidatus Kunth Cyperus cyperoides (L.) Kuntze s. l. Cyperus difformis L. Cyperus dilatatus Schum. et Thonn. Cyperus distans L. f. s. l. Cyperus dubius Rottb. Cyperus exaltatus Retz. Cyperus fenzelianus Steud. Cyperus haspan L. Cyperus imbricatus Retz. Cyperus iria L. Cyperus latifolius Poir. Cyperus ligularis L. Cyperus maculatus Boeck. Cyperus margaritaceus Vahl Cyperus odoratus L. Cyperus papyrus L. Cyperus pectinatus Vahl Cyperus podocarpus Boeck. Cyperus pustulatus Vahl Cyperus reduncus Hochst. ex Boeck. Cyperus rotundus L. Cyperus soyauxii Boeck. Cyperus sphacelatus L. Cyperus squarrosus L. Cyperus submicrolepis Kük. Cyperus tenax Boeck. Cyperus tenuiculmis Boeck. s. l. Cyperus tenuis Sw. s. l. Cyperus tenuispica Steud. Cyperus tisserantii Cherm. Cyperus tonkinensis C. B. Clarke var. baikiei (C. B. Clarke) S. S. Hooper 123 Genet Resour Crop Evol Table 3 continued Family Genera Crop Crop wild relatives Dioscoreacea Dioscorea Dioscorea alata L. Dioscorea abyssinica Hochst. ex Kunth Dioscorea bulbifera L. Dioscorea burkilliana J. Miège Dioscorea dumetorum (Kunth) Pax Dioscorea cayenensis Lam. Dioscorea rotundata Poir. Dioscorea hirtiflora Benth. Dioscorea lecardii De Wild. Dioscorea mangenotiana J. Miège Dioscorea minutiflora Engl. Dioscorea praehensilis Benth. Dioscorea preussii Pax Dioscorea quartiniana A. Rich. Dioscorea sansibarensis Pax Dioscorea smilacifolia De Wild. Dioscorea togoensis Knuth Euphorbiaceae Jatropha Jatropha curcas L. Jatropha integerrima Jacq. Jatropha gossypiifolia L. Jatropha kamerunica Pax et K. Hoffm. var. trochainii Léandri Jatropha multifida L. Jatropha neriifolia Müll. Arg. Jatropha podagrica Hook. Lamiaceae Manihot Manihot esculenta Crantz Manihot glaziovii Müll. Arg. Ocimum Ocimum basilicum L. Ocimum americanum L. Ocimum canum Sims Ocimum americanum L. Lauraceae Persea Persea americana Mill. – LegPapilionoideae Arachis Arachis hypogaea L. – Cajanus Cajanus cajan (L.) Millsp. Cajanus kerstingii Harms Canavalia Canavalia ensiformis (L.) Ce. Canavalia africana Dunn Glycine Glycine max (L.) Merr. Lablab Lablab purpureus (L.) Sweet – Vigna Vigna frutescens A. Rich. Vigna adenantha (G. F. Mey.) Maréchal, Mascherpa et Stainier Vigna radiata (L.) R. Wilczek Vigna comosa Baker Vigna subterranea (L.) Verdc. Vigna falicaulis Hepper Vigna unguiculata (L.) Walp. subsp. unguiculata var. unguiculata Vigna gracilis (Guill. et Perr.) Hook. f. Cajanus scarabaeoides (L.) Thouars var. scarabaeoides Canavalia ensiformis (L.) Ce. – Vigna juruana (Harms) Verdc. Vigna kirkii (Baker) Gillett Vigna laurentii De Wild. Vigna luteola (Jacq.) Benth. Vigna multinervis Hutch. et Dalziel Vigna nigritia Hook. f. Vigna oblongifolia A. Rich. Vigna pseudovenulosa (Maréchal, Mascherpa et Stainier) Pasquet et Maesen Vigna pubigera Baker var. beninensis (Pasquet et Maréchal) Pasquet et Maesen Vigna pubigera Baker var. pubigera Baker Vigna radiata (L.) R. Wilezek var. sublobata (Roxb.) Verdc. Vigna racemosa (G. Don) Hutch. et Dalziel Vigna reticulata Hook. f. Vigna stenophylla Harms 123 Genet Resour Crop Evol Table 3 continued Family Genera Crop Crop wild relatives Vigna trichocarpa (C. Wright) A. Delgado Vigna triphylla (R. Wilezek) Verdc. Vigna unguiculata (L.) Walp. subsp. baoulensis (A. Chev.) Pasquet Vigna unguiculata (L.) Walp. subsp. unguiculata var. spontanea (Schweinf.) Pasquet Vigna venulosa Baker Vigna vexillata (L.) A. Rich. var. angustifolia (Schumach. et Thonn.) Baker Vigna vexillata (L.) A. Rich. var. vexillata Malvaceae Gossypium Gossypium barbadense L. Gossypium arboreum L. Gossypium hirsutum L. Musaceae Musa Musa acuminata L. – Musa balbisiana L. Myrtaceae Psidium Psidium guajava L. – Psidium guineense Sw. Passiffloraceae Passiflora Passiflora edulis Sims Passiflora foetida L. Pedaliaceae Sesamum Sesamum indicum L. Sesamum alatum Thonn. Piperaceae Piper Piper nigrum L. Piper guineense Schumach. et Thonn. Sesamum radiatum Schumach. et Thonn. Poaceae Cymbopogon Cymbopogon citratus (DC.) Stapf Cymbopogon giganteus (Hochst.) Chiov. Digitaria Digitaria exilis Stapf Digitaria argillacea (Hitchc. et Chase) Fern. Digitaria ciliaris (Retz.) Koeler Digitaria debilis (Desf.) Willd. Digitaria delicatula Stapf Digitaria diagonalis (Nees) Stapf var. hirsuta (De Wild. et Th. Dur.) Troupin Digitaria gayana (Kunth) Stapf ex A. Chev. Digitaria horizontalis Willd. var. porrhanta (Steud.) Henr. ex Hubb. et Vaughan Digitaria leptorhachis (Pilger) Stapf Digitaria longiflora (Retz.) Pers. Digitaria nuda Schumach. Digitaria gayana (Kunth) Stapf ex A. Chev. Oryza Oryza glaberrima Steud. Oryza barthii A. Chev. Oryza sativa L. Oryza longistaminata A. Chev. et Roehr. Pennisetum glaucum (L.) R. Br. subsp. glaucum Pennisetum glaucum (L.) R.Br. subsp. sieberianum (Schlecht.) Stapf et C. E. Hubb. Oryza punctata Steud. Pennisetum Pennisetum glaucum (L.) R.Br. subsp. violaceum (Lam.) Rich. Pennisetum hordeoides (Lam.) Steud. Pennisetum macrourum Trin. Pennisetum pedicellatum Trin. Pennisetum polystachion (L.) Schult. subsp. atrichum (Stapf et C. E. Hubb.) Brunken Pennisetum polystachion (L.) Roem. et Schult. subsp. polystachion (L.) Schult. Pennisetum polystachion (L.) Schult. subsp. setosum (Sw.) Brunken Pennisetum purpureum Schumach. Pennisetum subangustum (Schumach.) Stapf et C. E. Hubb. 123 Saccharum Saccharum officinarum L. – Sorghum Sorghum bicolor (L.) Moench Sorghum arundinaceum (Willd.) Stapf Zea Zea mays L. – Genet Resour Crop Evol Table 3 continued Family Genera Crop Crop wild relatives Primulaceae Talinum Talinum portulacifolium (Forssk.) Asch. et Schweinf. Talinum portulacifolium (Forssk.) Asch. et Schweinf. Ixora Ixora coccinea L. Ixora brachypoda ne. Rutaceae Citrus Citrus aurantifolia (Christm. et Panzer) Swingle – Citrus aurantium L. Citrus limon (L.) Burm. f. Citrus maxima (Burm.) Merrill Citrus medica L. Citrus reticulata Blanco Citrus sinensis Osbeck Sapindaceae Blighia Blighia sapida König Blighia unijugata Baker Sapotaceae Synsepalum Synsepalum dulcificum (Schumach. et Thonn.) Daniell Synsepalum brevipes (Baker) T. D. Penn. Capsicum annuum L. (Chinense Group) – Blighia welwitschii (Hiern) Radlk. Synsepalum glycydora Wernham Synsepalum passargei (Engl.) T. D. Penn. Solanaceae Capsicum Capsicum annuum L. (Chillies Group) Capsicum annuum L. (Bird Pepper Group) Capsicum annuum L. (Sweet pepper and paprika Group) Lycopersicon Lycopersicon esculentum Mill. – Nicotiana Nicotiana tabacum L. – Solanum Solanum aethiopicum L. Group Gilo Solanum americanum Mill. Solanum aethiopicum L. Group Shum Solanum anomalum Thonn. Solanum distichum Schumach. Solanum macrocarpon L. Solanum melongena L. Solanum scabrum Mill. Solanum anguivi Lam. 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