A global approach to crop wild relative conservation: securingthe gene pool for food and agriculture

Nigel Maxted1, Shelagh Kell1, Álvaro Toledo2, Ehsan Dulloo3, Vernon Heywood4, Toby Hodgkin3,Danny Hunter3, Luigi Guarino5, Andy Jarvis6 & Brian Ford-Lloyd1

Summary. In light of the growing concern over the potentially devastating impacts on biodiversity and foodsecurity of climate change and the massively growing world population, taking action to conserve crop wild rela-tives (CWR), is no longer an option — it is a priority. Crop wild relatives are species closely related to crops,including their progenitors, many of which have the potential to contribute beneficial traits to crops, such as pestor disease resistance, yield improvement or stability. They are a critical component of plant genetic resources forfood and agriculture (PGRFA), have already made major contributions to crop production and are vital for futurefood security; their systematic conservation in ways that ensure their continuing availability for use is thereforeimperative. This is a complex, interdisciplinary, global issue that has been addressed by various national andinternational initiatives. Drawing on the lessons learnt from these initiatives we can now propose a global approachto CWR conservation, the key elements of which are: (1) estimating global CWR numbers, (2) assessment of theglobal importance of CWR diversity, (3) current conservation status, (4) threats to CWR diversity, (5) systematicapproaches to CWR conservation, (6) CWR informatics, and (7) enhancing the use of CWR diversity.

Key Words. conservation, crop diversity, crop wild relatives, genetic diversity, plant genetic resources for food andagriculture.

IntroductionIn light of the growing concern over the predicteddevastating impact of climate change on global biodiver-sity and food security, coupled with a growing worldpopulation, taking action to conserve crop wild relatives(CWR) has become a high priority. Crop wild relatives arespecies with a close genetic similarity to crops and manyof them have the potential or actual ability to contributebeneficial traits to these crops, such as resistance to bioticand abiotic stresses, and higher, more stable yields(Prescott-Allen&Prescott-Allen 1986; Hoyt 1988; Maxtedet al. 1997a; Tanksley & McCouch 1997; Meilleur &Hodgkin 2004; Stolton et al. 2006). A pragmaticapproach not without problems places all species ofthe same genus as a crop in this category (Heywood1994; Maxted et al. 2006). CWR have already mademajor contributions to crop production, and arevital for future food security. The systematic con-servation of this critical component of plant geneticresources for food and agriculture (PGRFA) in ways

that ensure their continuing availability for use istherefore imperative.

Darwin (1868) observed “… it appears strange tome that so many of our cultivated plants should stillbe unknown or only doubtfully known in the wildstate”. It was the Russian botanist N. I. Vavilov whofully recognised and championed the potential ofCWR for crop improvement in the 1920s and 30s,referring to the use of wild Aegilops L., Secale L.,Haynaldia Kanitz and Agropyron Gaertn. species inwheat breeding, for example (Vavilov 1949). CWRwere first routinely used by agricultural scientists toimprove major crops in the 1940s and 50s and by the1960s and 70s this was leading to some majorbreeding improvements (Meilleur & Hodgkin 2004).

There has been increasing interest in CWR con-servation and use in recent years, arising fromincreased recognition of their value as well as increas-ing ease of use. These are complex, interdisciplinary,global issues that have been addressed by various

Accepted for publication December 2010.1 School of Biosciences, University of Birmingham, Birmingham, B15 2TT, UK.2 FAO Commission on Genetic Resources for Food and Agriculture, Viale delle Terme di Caracalla 00153 Rome, Italy.3 Bioversity International, Via dei Tre Denari 472/a, 00057, Maccarese (Fiumicino), Rome, Italy.4 School of Biological Sciences, University of Reading, Reading, RG6 6AS, UK.5 Global Crop Diversity Trust, c/o FAO, Viale delle Terme di Caracalla 00153 Rome, Italy.6 International Centre for Tropical Agriculture, Recta Cali-Palmira, Apartado Aéreo 6713 Cali, Colombia.

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national and international initiatives, including twoGlobal Environment Facility projects (‘In situ Conser-vation of Crop Wild Relatives through EnhancedInformation Management and Field Application’ and‘Design, Testing and Evaluation of Best Practices for insitu Conservation of Economically Important WildSpecies’), the European Community-funded project‘European Crop Wild Relative Diversity Assessmentand Conservation Forum (PGR Forum)’, the FAOcommissioned ‘Establishment of a global network forthe in situ conservation of crop wild relatives: statusand needs’, the IUCN Species Survival CommissionCrop Wild Relative Specialist Group and the European‘In Situ and On Farm Network’. The need to addressCWR conservation is also highlighted in internationaland regional policy instruments, such as the Conven-tion on Biological Diversity (CBD 1992), FAO GlobalPlan of Action for the Conservation and SustainableUtilization of PGRFA (FAO 1996), European Com-munity Biodiversity Strategy (EC 1998), CBD GlobalStrategy for Plant Conservation (CBD 2002), Interna-tional Treaty on Plant Genetic Resources for Food andAgriculture (FAO 2001), European Plant Conserva-tion Strategy (Planta Europa 2001), Global Strategyfor CWR Conservation and Use (Heywood et al.2008), and most recently in the European Strategyfor Plant Conservation (Planta Europa 2008). Thislast recommends the establishment of 25 CWRgenetic reserves in Europe and undertaking gapanalysis of current ex situ CWR holdings, followedby filling of diversity gaps. Drawing on the experi-ence of these initiatives, and to help meet interna-tional and regional obligations, this paper aims tooutline a coherent policy for CWR conservation anduse to be implemented over the next 10 years. CWRconservation and use provides an excellent exemplarof how to address the dual Millennium DevelopmentGoals of combining biodiversity conservation withpoverty alleviation.

Estimating global CWR numberRecent studies have found that using the broaddefinition of a CWR (i.e. all species in the same genusof the crop) the number of CWR species of interestmay be much larger than previously recognised. Forexample, using this definition, Kell et al. (2008) foundthat 17,495 (8,624 of them endemic) out of approx-imately 20,590 species, or 85% of the European flora,comprises crop and CWR species. Practically, withsuch large numbers there is a need for more effectiveprioritisation. Maxted & Kell (2009) calculated thatthe 77 major and minor food crops as defined byGroombridge & Jenkins (2002) contained around10,700 species using the numbers of species per genusfrom Mabberley (2008). However, this number, likethe European estimate, is likely to be inflated by the

inclusion of remote as well as closely related CWRfound in the same genus as the crop. Maxted et al.(2006) argued that a more precise target could beobtained by focusing on the crop gene pool GP1Bor taxon groups TG1b and TG2 alone, whichcontain the closest CWR species. Based on an initialsample of 14 food crop groups (see Table 1) Maxted& Kell (2009) estimated that globally, for the 77major and minor food crops, there are 221 very closewild relatives and 471 close wild relatives. Thus, as aworking estimate, we may need to deal globally witharound 700 closely related CWR species (i.e., lessthan 0.26% of the world flora) in order to ensurethat the highest priority genetic diversity of majorand minor food crops is conserved and madeavailable for use in crop improvement programmesas a contribution to future worldwide food security.In addition, CWR of forage species which alsocontribute to food security will need to be identifiedand conserved. A case can also be made for similaraction for the CWR of fuel and fibre crops andindustrial and ornamental crops although they arenot the primary concern of this paper.

Therefore, a goal over the next ten years is torefine the working estimate of global highest prior-ity CWR for food and agriculture. Once such a list isavailable it would greatly facilitate the targeting of insitu and ex situ conservation actions. In the longer-term, however, it would be unwise to restrict con-servation to these species alone, as we know thosetraits desired by germplasm users are not exclusivelylocated in the closest CWR species, so a secondarypriority would be the genetic conservation of all10,739 CWR species congeneric to the 77 major andminor food crops.

Assessment of global importance of CWR diversityCWR, like other wild species, may be valuablecomponents of ecosystems but many of them alsohave additional specific value as gene donors forplant breeding. Prescott-Allen & Prescott-Allen(1986) calculated that the yield and quality contribu-tion by CWRs to US grown or imported crops wasover US$350 million a year, while Phillips & Meilleur(1998) estimated that potential losses associated withendangered food crop wild relatives were around US$10 billion annually in wholesale farm values. Fur-ther, Pimentel et al. (1997) estimated that thecontribution of genetic resources to yield increase isabout 30% of production, and much of this resultsfrom CWR species, so the introduction of new genesfrom wild relatives contributes approximately $20billion toward increased crop yields per year in theUS and $115 billion worldwide.

Despite their known value as gene donors, Tanksley& McCouch (1997) argued that breeders were notfully exploiting the potential of CWR because, histor-

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ically, they relied on searching for specific beneficialphenotypic traits associated with particular CWR taxa,rather than searching for beneficial genes. Although itwould be very difficult to give a precise estimate ofCWR use by breeders because the data are likely to becommercially sensitive and therefore not readilyavailable, Maxted & Kell (2009) recently reviewedthe use of CWR in crop improvement and cited 291articles reporting the identification and transfer ofuseful traits from 185 CWR taxa into 29 crop species.They found that the degree to which breeders hadused CWR diversity varies markedly between crops,being particularly prominent in barley, cassava, potato,rice, tomato and wheat. Of these, rice and wheat arethe crops in which CWR have been most widely used,both in terms of the number of CWR taxa used andsuccessful attempts to introgress traits from the CWRto the crop (Fig. 1). The most widespread CWR usehas been and remains in the development of diseaseand pest resistance, with references citing diseaseresistance for 39% of inter-specific trait transfers, pestresistance 17%, abiotic stress 13%, yield increase 10%,cytoplasmic male sterility and fertility restorers 4%,quality improvers 11% and husbandry improvement6% of the reported inter-specific trait transfers. It isalso notable that the number of publications detailingthe use of CWR in breeding has increased graduallyover time (Maxted & Kell 2009). The exploitation ofthe potential diversity contained in CWR speciesremains poorly directed as the approach by breedersto CWR use has not been systematic or comprehen-sive; therefore, most CWR diversity that might be usedfor breeding remains untapped. Hajjar & Hodgkin(2007) comment that although quantitative trait locihave been identified in many CWR species, thepotential to exploit them as a breeding resource usingnew molecular technologies has yet to be fully

realised. Although this situation is likely to improvewith time, it does underpin the need for thecontinued availability of a broad range of CWRdiversity, also emphasising the conservation-use link-age and the need for the conservation communityto meet the evolving needs of the users. Therefore,the goal over the next ten years is to take advantageof novel technological advances in trait recognitionand inter-specific breeding to extend the breadth ofCWR use to a broader range of crops and system-atically review the potentially useful diversity inCWR gene pools.

Threats to CWR diversityCWR are biologically no different from other wildplant species, and, like them, many are currentlythreatened with loss of diversity and/or extinction, asa result of anthropogenic influences: habitat destruc-tion and fragmentation, unsustainable resourceexploitation, changes and intensification of landmanagement, and invasive species. In addition tothese factors, and interacting with them, acceleratedclimate change is likely to present a step-shift in termsof extinction and genetic erosion (IPCC 2007) andthis threat requires urgent action to avoid foodinsecurity (Lobell et al. 2008). In Europe, Thuiller etal. (2005) modelled projections of the future distribu-tion of 1350 plant species and found that by 2080more than half of them could be vulnerable orthreatened by climate change. In one of the fewstudies of the effects of climate change on CWR, Jarviset al. (2008) projected that 16 – 22% of wild Arachis L.,Solanum L. and Vigna Savi species would go extinct by2055. Similar modelling of climate change scenarios inMexico by Lira et al. (2009) highlight that most of theeight wild Cucurbitaceae taxa studied are predicted

Table 1. Numbers of primary and secondary CWR species in 14 crop gene pools (Maxted & Kell 2009).

Crop Crop taxonSpeciesin genus

PrimaryCWR species

SecondaryCWR species

% Priorityin genus

Finger millet Eleusine coracana Gaertn. 9 3 3 66.67Barley Hordeum vulgare L. 16 1 1 12.50Sweet potato Ipomoea batatas (L.) Poir. 600 – 700 3 11 2.00Cassava Manihot esculenta Crantz 98 3 13 16.33Banana/plantain Musa acuminata Colla 30 10 15 83.33Rice Oryza sativa L. 23 8 9 73.91Pearl millet Pennisetum glaucum (L.) R. Br. 80 – 140 1 2 2.14Garden pea Pisum sativum L. 3 1 2 100.00Potato Solanum tuberosum L. 1000 6 24 3.00Sorghum Sorghum bicolor (L.) Moench 25 2 2 16.00Wheat Triticum aestivum L. 6 + 22 6 12 64.29Faba bean Vicia faba L. 140 1 0 0.71Cowpea Vigna unguiculata (L.) Walp. 61 1 3 6.56Maize Zea mays L. 4 1 3 100.00

Totals 2117 – 2277 47 100 6.45% 100 2.06 4.39

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not to survive under accepted climate change models.A further threat that is specific to CWR diversity is thatmany CWR of major crops are found in disturbed, pre-climax communities (Jain 1975), which are thehabitats that are likely to be subject to increasinglevels of anthropogenic change and destruction.

The most commonly applied means of assessingthreats to wild taxa is the application of the IUCNRed List criteria (IUCN 2001) but CWR have thus farnot been specifically prioritised for assessment,although it should be noted that a Red Book of CropWild Relatives in Bolivia (Libro Rojo de ParientesSilvestres de Bolivia) has recently been published(Mora et al. 2009). Of the 14 food crop groupsstudied by Maxted & Kell (2009), only two cropgroups had been assessed — one sweet potatorelative, Ipomoea pulcherrima Ooststr. was assessed asbeing vulnerable and 44 potato (Solanum) species wereassessed as being threatened to varying degrees.

Although it is difficult to quantify the loss of geneticdiversity within CWR species, it is likely to be verymuch greater than the loss of species given that mostof the species that are able to survive the threats towhich they are exposed will suffer some geneticerosion (loss of genetic diversity (Maxted et al.1997b). It therefore seems likely that virtually allCWR species are currently suffering some degree ofloss of genetic diversity. Maxted et al. (1997b) esti-mated that 25 – 35% of plant genetic diversity wouldbe lost between the ratification of the CBD in 1993and the 2010 Biodiversity Target date. The magnitudeand rapidity of climate change, coupled with otherthreats, is likely to impose extreme selection pressureon the surviving populations of CWR over the coming

50 – 100 years. Loss of any genetic diversity means thatspecies may not be able to adapt to changingconditions quite so readily in the future, but also thatvital diversity necessary to underpin our future foodsecurity will not be available to breeders.

A goal over the next ten years should be toundertake systematic threat assessment for as wide arange of CWR taxa as possible, using IUCN or nationalcriteria, or both. This is a critical element of the IUCNCWR Species Survival Commission Specialist Group’sOperational Strategy (see http://www.cwrsg.org/). Asa first step, the global priority 700 CWR species shouldbe assessed, then as a secondary priority all 10,739CWR species related to the 77 major and minor foodcrops. The IUCN Red List criteria are not entirelysuitable for assessing the threat to genetic erosion orextinction within species, and so a subsidiary goal willbe the development of a practical means of assessingthreat to genetic diversity within species. Likewise, thecurrent IUCN criteria do not take climate changespecifically into account, although plans are in handto do so.

Another urgent goal should be to undertakebioclimatic modelling of as many CWR as possible, sothat we can obtain as accurate a picture as possible oftheir likely adaptation, migration or loss.

Active CWR ConservationA complementary approach to CWR conservationshould be adopted, encompassing in situ conservationin natural habitats as well as ex situ measures focusedmainly on seed storage. As for other wild species, thepreferred means of conserving CWR is in situ, as living

Fig. 1. The number of references reporting the identification and transfer of useful traits from 185 CWR taxa to 29 crop species,showing the number of CWR taxa used in each crop (Maxted & Kell 2009).

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populations in protected areas or genetic reserves1 oras seed samples dried and stored at sub-zero temper-atures in gene banks (Maxted et al. 1997c; Heywood &Dulloo 2006; Stolton et al. 2006; Iriondo et al. 2008a).Currently, the highest proportion of CWR diversity isactively conserved ex situ, although the coverage is farfrom comprehensive. Although CWR diversityundoubtedly occurs in existing protected areas, theCWR species are rarely actively managed and moni-tored in them, so their conservation status is uncertainand populations or entire species could be at riskwithout the protected area management being aware ofthe threat or consequences.

The First Report of the State of the World’s PGRFA (FAO1998) listed 4% of governmental, 14% of CGIAR and6% of private gene bank holdings as wild species, whilethe Second Report of the State of the World’s PGRFA (FAO2009) listed 10% of gene bank holdings as wild speciesand concluded “Interest in collecting and maintainingcollections of CWR is growing as land-use systems change,concerns about the effects of climate change grow andtechniques for using the material become more powerful andmore readily available”. So it appears that ex situcollections and interest in conserving CWR hasincreased, yet FAO (2009) ultimately concludes that“For several major crops, such as wheat and rice, a large partof the genetic diversity is now represented in collections.However, for many other crops, especially many neglected andunder-utilized species and CWR, comprehensive collections stilldo not exist and considerable gaps remain to be filled”. ThoseCWR collections that do exist, because of the ad hocmanner in which they have largely been collected, areunlikely to constitute genetically representative samplesof global CWR diversity.

As an indicator of the coverage of ex situ collections,analysis of the data in the European crop gene bankportal (EURISCO) revealed that CWR taxa account for5.6% of total germplasm holdings, and that the 1,095CWR species included represent only 6% of the 17,495CWR species found in Europe (O’Regan 2007). Theratio of cultivated to wild species in ex situ collections is12:1, though most diversity is located in wild species(Maxted et al. 2008a). In contrast, analysis of Europeanex situ seed collections held in botanic garden genebanks via the ENSCONET portal revealed that CWRtaxa account for 61.8% of total germplasm holdings,and that the 5756 CWR species included represent 33%of 17,495 CWR species found in Europe. Initially itwould appear that botanic garden gene banks do amuch better job of conserving CWR diversity than cropgene banks. However, care should be taken in drawingtoo crude a conclusion from this stark difference, as it is

likely that the 1,095 CWR species held in crop genebanks are more likely to include high priority CWRspecies from the primary and secondary gene pools.Moreover, many of the botanic garden accessions aresmall and genetically poorly sampled (Heywood 1999).

The effectiveness of in situ CWR conservation is, ifanything, more uncertain than ex situ, and althoughthere has been considerable attention paid to the theoryof design, establishment, management and monitoringof CWR diversity in genetic reserves (see Jain 1975; Hoyt1988; Gadgil et al. 1996; Hawkes et al. 1997; Maxted et al.1997a, c; Safriel et al. 1997; Heywood & Dulloo 2006;Stolton et al. 2006; Iriondo et al. 2008a), full practicalimplementation remains limited (Maxted et al. 1997a, c;Meilleur & Hodgkin 2004). In practice, conservation ofCWR is often planned within existing protected areasbecause: a) by definition they are protected andmanaged (although not always effectively) and alreadyhave an associated long-term conservation ethos and areless prone to short-term management changes, b) insome cases, it is relatively easy to negotiate changes inthe existing site management plan so as to facilitategenetic conservation of CWR species, and c) thecreation of new conservation sites can be avoided alongwith the problems and cost of acquiring new land forconservation (Iriondo et al. 2008a). There are somenotable examples of activities that have made a signifi-cant contribution to the process of conserving CWR insitu (see Table 2), but even in some of these cases thesites are not managed in the most appropriate mannerto conserve CWR genetic diversity.

There are a number of potential approaches tosystematic CWR conservation, but three distinct (thoughcomplementary) approaches may be adopted — indi-vidual, national and global (Maxted & Kell 2009). It isimportant to recognise, however, that conservationstrategies are more likely to be successful if nationalgovernments, on-the-ground agencies and local peopleset the agenda, for it is they who will be responsible fortheir implementation, with international NGOs andIGOs playing a supporting role (Smith et al. 2009). Theindividual approach involves a protected area or genebank manager actively promoting CWR conservationwithin the site or facility that they manage. The nationalapproach requires countries to develop CWR conserva-tion strategies, which when implemented over timewould result in the systematic representation of thenation’s CWR diversity in an in situ network of geneticreserves or other conservation areas, with complemen-tary ex situ storage of genetically representative popula-tion samples in national gene banks (Fig. 2). Thismodel for establishing a national CWR inventory wasrecently tested in the UK and Portugal (see Maxtedet al. 2007; Magos Brehm et al. 2008) and showed that inthe UK 17 sites contain 152 (67.3%) of the priority UKCWR species. However, so far, very few countries havedeveloped a national CWR conservation strategy and

1 Synonymous terms include ‘genetic reserve management units’(GRMUs), ‘gene management zones’ (GMZs), ‘gene sanctua-ries’ or ‘genetic sanctuaries’ and ‘crop reservations’.

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the United Nations Environment Programme (UNEP)/Global Environment Facility (GEF)-supported projectdescribed below is exceptional in including such anational strategy for each of the five countries involvedas one of its outputs. The global approach entails theestablishment of a worldwide network of in situ geneticreserves that is independent of national politicalborders and focuses on worldwide priority crop genepools, again with complementary ex situ storage ofgenetically representative population samples.

A global approach to the in situ gap analysis, thatinvolves the comparison of naturally occurring CWRdiversity with the diversity sampled and conservedeither in situ or ex situ (see Maxted et al. 2008b), for14 globally important food crop groups (finger millet,barley, sweet potato, cassava, banana/plantain, rice,pearl millet, garden pea, potato, sorghum, wheat, fababean, cowpea and maize) was undertaken by Maxted &Kell (2009) and suggested priority locations for CWRgenetic reserve establishment (see Map 1). Althoughfurther crop groups should be added, these prioritysites can be used to begin recommendations forestablishment of the global network. For each foodcrop group the identified priority sites are primarilywithin the boundaries of existing protected areas,although this is not always the case and the results doindicate the need to establish new protected areas. Theresults of the 14 crop complex analyses (with theexception of the Middle East and Eastern Congo) showfew obvious opportunities for conservation of multi-

crop gene pools in single genetic reserves,2 furtherresearch is likely to identify additional potential multi-taxon CWR genetic reserves locations where limitedconservation resources could be effectively targeted.

A complementary global approach to ex situ gapanalysis of 13 globally important food crop groups(chickpea, common bean, barley, cowpea, wheat,maize, sorghum, pearl millet, finger millet, pigeonpea, faba bean, and lentil) has also been undertaken.Jarvis and colleagues (Jarvis et al. 2008) analysed28,751 herbarium and gene bank specimen/accessionoccurrences accessible through the Global BiodiversityInformation Facility (GBIF) for 643 CWR taxa belong-ing to the 13 gene pools (see http://gisweb.ciat.cgiar.org/gapanalysis/). They used a maximum entropyclimate envelope model to create distribution mapsfor each species under current climates using theWorldClim database. The results show the species andgeographic regions where high priority for geneticresource collection exists (see Map 2). The resultsshow that significant gaps still exist in ex situ con-servation, and that protected areas do not currentlyprovide adequate conservation of the species. Prior-ities for collection occur especially in Africa, northern

Table 2. Examples of CWR conserved in protected areas.

CWR Protected Area Country References

Wild emmer wheat (Triticum turgidum subsp. dicoccoides) Ammiad, Galilee Israel Anikster et al. (1997);Safriel et al. (1997)

Teosinte (Zea diploperennis Iltis, Doebley & R. Guzman) MAB Sierra de ManantlánBiosphere Reserve

Mexico Sanchez-Velasquez (1991)

Wild wheats (Triticum turgidum subsp. dicoccoides,T. monococcum L., Aegilops tauschii Coss, A. speltoides Tausch)

Ceylanpinar Turkey Karagöz (1998)

Chestnut (Castanea sativa Mill.), wild plum (Prunus cerasiferaEhrh. var. divaricata)

Kazdağ Turkey Kűçűk et al. (1998)

Medicago L. spp., Vicia L. spp., Trifolium L. spp., Lathyrus L. spp.,Lens Mill. spp., Triticum L. spp., Avena L. spp., Hordeum L. spp.,Aegilops L. spp., Allium L. spp., Amygdalus L. spp., Prunus L. spp.,Pyrus L. spp., Pistacia L. spp. and Olea L. spp.

Abu Taha Lebanon Al-Atawneh et al. (2008)Sale-Rsheida SyriaAjloon JordanWadi Sair Palestinian

TerritoriesWild wheats (Triticum boeoticum Boiss., T. urartu

Thumanjan ex Gandilyan, T. araraticum Jakubz.)Erebuni Armenia Avagyan (2008)

Wild bean populations (Phaseolus L. spp.) Central valley Costa Rica Zoro Bi et al. (2003);Baudoin et al. (2008)

Wild coffee (Coffea mauritiana Lam., C. macrocarpaA. Rich., C. myrtifolia (A. Rich. ex DC.) J.-F.Leroy)

Black River GorgesNational Park

Mauritius Dulloo et al. (1998)

Wild onions (Allium columbianum (Ownbey & Mingrone)P. M. Peterson, Annable & Rieseberg, A. geyeri S. Watson,A. fibrillum M. E. Jones)

Great Basin,Washington State

USA Hannan & Hellierin Pavek et al. (1999);Hellier (2000)

Wild grapevine (Vitis rupestris Scheele, V. shuttleworthiiHouse, V. monticola Buckley)

Witchita Mountains andOuachita NationalForest, Oklahoma,Clifty Creek, Missouri

USA Pavek et al. (2003)

2 It should be noted that both multi-species and single speciesreserves have advantages and disadvantages and much moreexperience is needed to be able to draw conclusions as towhich is preferable (see discussion in Heywood & Dulloo 2006:40 – 41).

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Australia, Central America and the Andes. As many as39 species occur sympatrically in parts of Africa, wheremultiple gaps could be filled with targeted collectingor establishment of genetic reserves.

Over the last 10 years the GEF has supported anumber of projects that seek to enhance the conserva-tion and use of crop wild relatives. One such project isthe UNEP/GEF-supported project, ‘In situ conservation

of crop wild relatives through enhanced informationmanagement and field application’, coordinated byBioversity International. Five countries — Armenia,Bolivia, Madagascar, Sri Lanka and Uzbekistan — areinvolved in the project through their governments andother agencies. This project has expanded substantiallythe previously limited body of knowledge on in situCWR conservation in developing countries. The project

Fig. 2. Model for the development of national CWR strategies (Maxted et al. 2007).

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facilitated the prioritisation of CWR species from 36different genera for ecogeographic assessments andthreat assessment through Red Listing. Over 310 CWRspecies were assessed according to the current IUCNcriteria and Bolivia has produced the first ever nationalRed Book of CWR. This is probably the largest set ofsuch assessments undertaken for CWR and represents amajor contribution to practice. Further, the partner-ship, through the involvement of protected areaauthorities and other relevant stakeholders such aslocal and indigenous communities, has seen thedevelopment of CWR species management plans forimplementation in protected areas, as well as theadaptation of protected area management plans them-selves so as to take into account the management needsfor CWR conservation. Species management andmonitoring plans have been developed to manageCWR diversity in protected areas in each projectcountry (Table 3). In addition the project has identifiedareas outside protected areas for in situ CWR conserva-tion, including those for Oryza nivara S. D. Sharma &Shastry and O. rhizomatis D. A. Vaughan in PuttalamDistrict, Sri Lanka and Malus sieversii M. Roem., Juglansregia L. and Pistacia vera L. in Uzbekistan. Therefore,although significant progress has been made in recentyears, CWR species, as well as the diversity within them,are seriously under-conserved both ex situ and in situ.

It is critical that in the next ten years a more strategicapproach is taken to targeting ex situ CWR conservation.First the initial strategic target should be to conserve exsitu genetically representative samples of the highestpriority 700 globally important CWR food crop-relatedspecies, and second to ensure that there is systematicrepresentation of the 10,739 priority CWR speciesrelated to the major and minor food crops conservedin gene banks. Further, the extension of the gapanalysis of food crop groups by Jarvis and colleagues isstrongly recommended to help prioritise the collectionof the 77 major and minor food crop gene pools.

The establishment of genetic or other kinds ofreserve for CWR in times of rapidly rising humanpopulation, climate change and ecosystem instabilityas well as a global economic crisis is a complex goal,which necessitates a carefully researched strategicapproach. Sites competing for reserve status wouldneed to be assessed and prioritised for their longer-term sustainability in terms of the predicted impact ofclimate change on the site and the development plansassociated with local communities (Brooks et al. 2006).Some CWR populations conserved in vulnerablelocations are unlikely to be able to adapt sufficientlyquickly to climate change or even in doing so may gothrough a genetic bottleneck reducing the long-termviability of sustaining their diversity in situ. On the

Map 1. Global priority genetic reserve locations for wild relatives of 14 food crops (FAO 2009). The ‘centres of crop diversity’(indicated by the enclosed lines) are likely to contain further priority sites for other crop gene pools.

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Map 2. Locations of priority for ex situ and/or in situ conservation based on the identification of gaps in our current ex situconservation system for 13 important crop wild relative gene pools. A species level richness in gaps, and B genus level richness ofgaps, showing sites where gaps exist in multiple gene pools.

Table 3. Examples of CWR conserved in protected areas in Armenia, Bolivia, Madagascar, Sri Lanka and Uzbekistan.

Crop Gene Pool CWR Protected Area Country

Yam Dioscorea maciba Jum. & H. Perrier, D. bemandry Jum. &H. Perrier, D. antaly Jum. & H. Perrier, D. ovinala Bakerand D. bemarivensis Jum. & H. Perrier.

Ankarafantsika National Park Madagascar

Cinnamon-tree Cinnamomum cappara-coronde Blume Kanneliya Forest Reserve Sri LankaAlmond Amygdalus bucharica Korsh. Chatkal Biosphere Reserve UzbekistanWheat Triticum araraticum, T. boeoticum, T. urartu

and Aegilops tauschiiErebuni State Reserve Armenia

Cacao Theobroma L. spp. Parque Nacional y TerritorioIndigena Isiboro-Secure

Bolivia

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other hand, basing a strategy on ex situ action alone isunlikely to be economically feasible or practical given(a) the very large number of species involved (possiblyas many as 229,500 spp.; Maxted & Kell 2009; see alsodiscussion in Heywood 2009a), (b) the need to sampleand conserve ecogeographically and geneticallydiverse populations for each species, and (c) the factthat few species have immediate use potential. Assuch, the strategic establishment of a global network ofin situ CWR genetic reserves to complement targetedex situ collections should be regarded as a priority. Insitu CWR conservation has been primarily associatedwith genetic conservation in protected areas. However,CWR species are just as likely to be found outsideexisting protected areas as within them. In fact, as isargued above, many CWR of major crops are found indisturbed, pre-climax communities and these aremore rarely designated as protected areas. Therefore,another target in the next ten years will be to furtherelaborate methodologies for the in situ conservation ofCWR diversity outside of conventional protected areasthat are still able to meet the objective of maximisinglong-term sustainability of CWR diversity. Variouspublic and private initiatives to provide some degree ofprotection to species outside protected areas exist,especially in Australia, Brazil, China, Costa Rica, Mexico,South Africa, the USA and several European countries(Maxted et al. 2008c; Heywood 2009b). One approach isto promote CWR in situ conservation in less formallydesignated protected areas such as Indigenous andCommunity Conserved Areas (ICCAs) (Pathak et al.2004; Kothari 2006, 2008; see also http://www.iucn.org/about/union/commissions/ceesp/topics/governance/icca/), where indigenous peoples and local commun-ities have for millennia conserved natural environmentsand species for economic as well as cultural, spiritualand aesthetic reasons independent of more formalconservation sector interventions.

CWR Information ManagementCWR conservation requires effective information man-agement. There have been many initiatives to developinformation management systems for PGRFA, one ofwhich specifically addressed CWR information manage-ment. The FP5 funded “European Crop Wild RelativeDiversity Assessment and Conservation Forum (PGRForum)” (Maxted et al. 2008a) developed the CropWild Relative Information System (CWRIS) (Kell et al.2008) with the goal of providing a model for collationand management of CWR conservation and sustainableuse data and a system for accessing this information.This system makes available the PGR Forum Crop WildRelative Catalogue of Crop Wild Relatives for Europeand the Mediterranean (http://www.pgrforum.org/cwris/cwris.asp), containing more than 25,000 speciesrecords and in excess of 273,000 records of taxon

occurrences in 130 geographical units across theregion, with a limited number of detailed case studiesgiving the Catalogue biological depth. The structure ofCWRIS is generic and independent of the exemplardata initially included, and therefore could be used tomeet the information management requirements ofany CWR conservation and sustainable use data. AnXML (Extensible Markup Language) schema corre-sponding to the data model is available (see Mooreet al. 2008). Recently CWRIS has been further devel-oped and extended by the EC Gen Res project “AnIntegrated European In Situ Management Work Plan:Implementing Genetic Reserves and On Farm Concepts(AEGRO)”. CWRIS was used tomanage population leveldata for gene pool based studies of Avena L., Beta L.,Brassica L. and Prunus L. spp. (Frese & Maxted 2009).In the next ten years, CWRIS should be populated withother gene pool data and further developed andextended using data sets from other regions.

The GEF-supported project ‘In situ conservation ofcrop wild relatives through enhanced informationmanagement and field application’, has contributedto CWR information management through the estab-lishment of national information systems on CWR inArmenia, Bolivia, Madagascar, Sri Lanka and Uzbeki-stan. The systems have facilitated the mapping ofCWR distribution and identified priority areas fortheir conservation. In addition to the nationalinformation systems, a global portal (see http://www.cropwildrelatives.org), maintained by BioversityInternational, has been developed through whichnational CWR inventories can be searched.

While progress has been made in CWR informationmanagement this progress has been largely focused onshort-term research-based projects and there remainslittle coherent vision. There is a need to unite the variouselements already existing, such as CWRIS (http://www.pgrforum.org/cwris/cwris.asp for the taxonomic back-bone and EURISCO (http://eurisco.ecpgr.org/) forgene bank holdings, and to globalise their application toform a seamless information system from CWR diversityin thefield, in situ and ex situ conserved diversity, throughto characterised and evaluated CWR diversity, in orderto ensure the highest priority CWR diversity is conservedand available to use by the broad user community.Meeting the needs of the latter is the raison d’être ofCWR diversity conservation, so the system needs to bedeveloped jointly with the germplasm user communityto provide the quality of service that they require.

Enhanced CWR useConservation is not an end in itself. To be effective,conservation should be linked to use. In fact, sustain-able use is seen as the long-term means of sustainingactive conservation. Therefore, to encourage CWRconservation there is an associated need to promote

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use of the conserved diversity. There are numerous waysin which CWR use in breeding can be promoted, buttraditionally this has focused on trying to identify traits ofinterest through phenotypic characterisation and evalu-ation. This has in many cases proved prohibitivelyexpensive. The Report on the State of the World’s PGRFA(FAO 1998) highlights the fact that two thirds ofglobally conserved ex situ germplasm lack basic passportdata, 80% lack characterisation data and 95% lackevaluation data, making the use of such germplasm,including CWR germplasm, much more difficult than itneed be. Since the publication of the first SoW report,the Second Report on the State of the World’s PGRFA (FAO2009) details several new international initiatives thatsupport the increased characterisation and evaluationof germplasm, including the fairly widespread adoptionof core collections that are adequately characterisedand evaluated. However, it still concludes that “Thecountry reports were virtually unanimous in suggesting thatone of the most significant obstacles to a greater use of PGRFAis the lack of adequate characterization and evaluation dataand the capacity to generate and manage such data”.

The situation regarding access to in situ conservedgermplasm is likely to be even less conducive toexploitation — there are currently no known geneticreserves where the conserved species are fully charac-terised and evaluated. The bottleneck over systematiccharacterisation and evaluation has been acknowledgedalmost since the need for their conservation wasrecognised in the late 1960 s and early 1970 s (Frankel& Bennett 1970). It could be argued now that simplyincreasing the amount of ‘traditional’ characterisationand evaluation is unlikely to result in the required stepchange in the exploitation of CWR. This is despite theincreasing knowledge of useful genes that can be foundin CWR and what can be achieved by their introgres-sion into crops (Maxted & Kell 2009). For these reasonsthere is a major incentive to embrace innovativetechnologies to improve CWR utilisation.

‘Next generation technologies’ offer a way of screen-ing thousands of samples of germplasm for thoseinteresting gene variants that are adaptively important,making them available for use in conventional breeding.Given that such useful forms of genes will essentiallyrepresent ‘natural’ genetic variation, this approachcould even help to alleviate the concerns associated withGM technology. There are now many candidate genesshown to be involved in some way in drought tolerance,for instance, and large scale transcriptomics and rese-quencing will allow us to identify these genes and all thevariants that can be found in the CWR related to manycrops. A recent study of wild Brassica nigra (L.) W. D. J.Koch, B. rapa L. and B. oleracea L. populations fromNorthern and Southern Europe (Mitchell 2008)searching for genes associated with climate changeadaptive characters (i.e. rainfall and temperature)located 42 genes which showed differential north-south

expression across all three species (e.g. water channel-like protein, low temperature and salt responsiveprotein, wax synthase-like protein). The clear advant-age of this approach is that rather than select CWRgenetic diversity in general, the potential user can focusmore directly on adaptive capacity. Genomic databasescontaining such information must increasingly belinked to web-enabled databases of ex situ conservedCWR germplasm, such as the European InternetSearch Catalogue of Ex Situ PGR Accessions (EUR-ISCO) (http://eurisco.ecpgr.org/), System-wide Infor-mation Network for Genetic Resources (SINGER)(http://singer.cgiar.org/) and Germplasm ResourcesInformation Network (GRIN) (www.ars-grin.gov).

There is another, complementary approach to enrich-ing data on conserved germplasm, and therefore stim-ulating its use. ‘Predictive characterisation’ is the use ofspatial analysis to predict which germplasm might havedesired traits. For example, GIS can be used to identifygermplasm likely to be drought or frost tolerance byoverlaying environmental data with the locality of pop-ulations (see Pollak & Pham 1989; Chapman & Barreto1996; Guarino et al. 2002; Kaur et al. 2008). Similarapproaches can be adopted for other abiotic traits (e.g.,other climatic variables, salinity, soil mineral excesses ordeficiencies, and day length requirements). For desiredbiotic characteristics, the distribution of the CWR taxoncan be overlaid with the known distribution of pests ordiseases. The CWR populations found to be coincidentwith high levels of pests and diseases are likely to haveevolved resistance over time; therefore, these popula-tions might be predicted to harbour the required pest ordisease resistance. Further, if the pest or diseasedistribution is not precisely known, it is also possible tooverlay the CWR taxon’s distribution with the climaticconditions suitable for that disease. For example,Bhullara et al. (2009) used this Focused Identificationof Germplasm Strategy (FIGS) approach and startingwith 16,089 accessions of bread wheat, reduced thenumber to 1,320 then isolated 7 new resistance alleles topowdery mildew (genePm3) from these accessions.However, caution is necessary, as contrary examples arefound, where resistance alleles are discovered in areawhere the disease is unknown, such as the location ofRhizomania resistance in wild beet populations fromKalundborg Fjord, Denmark (Lothar Frese pers.comm.). It is more difficult to imagine how predictivecharacterisation might be used to predict yield or qualitytraits — therefore, for these traits, traditional character-isation and evaluation will remain necessary. While aliterature review currently reveals limited use of GISanalysis for predictive characterisation, with an increasedemphasis on the need to link in situ CWR conservationto use, these techniques offer innovative opportunities(Guarino et al. 2002).

Ultimately, unless the professionals involved with CWRconservation can ensure that conserved germplasm is

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held in a form better suited for breeders and other usergroups and that there is less of a barrier betweenconservation and utilisation, then the use of conservedCWR is not likely to improve. The rapid advancement ofbiotechnological techniques for transferring traitsbetween species is also likely to mean that inadequatecharacterisation and evaluation is likely to prove an evengreater barrier to CWRuse in the future. Therefore in thenext ten years there is a requirement for collectionmanagers, whether the collections are held in situ or exsitu, to promote the use of their conserved germplasm tothe user community by adopting the approaches out-lined above or other novel technologies associated withpre-breeding. Even though it seems likely that thevarious technical challenges remaining in trait transferbetween CWR and elite breeding material will beovercome in the next ten years or soon after, breedersare likely to still largely obtain the diversity they needfrom crop gene banks. This means that ex situ collectionsare likely to act as a conduit between in situ conserveddiversity and the breeders, so the unification of the insitu and ex situ CWR conservationists must also beseamless if we are to give users what they will demand.

ConclusionThe central proposition of this paper is that climatechange presents a new and rapidly developing threat

to global food security. CWR contain the geneticdiversity that can at least partially mitigate this threat,yet CWR themselves are in turn threatened. Conserva-tion of CWR has not been immune to the ‘research-implementation gap’ between science and real worldaction that has been identified elsewhere for conser-vation assessment. This is a real dilemma that has to beaddressed urgently. We already have the techniquesand experience to conserve and use CWR diversityeffectively for the benefit of humankind — the onus isnow to strengthen the weak existing links betweenbiodiversity and agrobiodiversity conservationists to actsystematically between themselves and in concert withcrop breeders to help guarantee the basis of globalfood security. It is very important, therefore, thatgovernments build on recent knowledge and proto-cols and incorporate responsibility for maintainingand implementing the conservation of CWR into theirnational biodiversity and PGR systems.

Further suggestions on how the systematic conser-vation of CWR diversity might be achieved and howthe conservation-utilisation link might be improvedand expanded are summarised in Appendix 1.Although the points are extensive, they are notexhaustive. It is also likely that they are already beingapplied by many proactive genetic reserve and genebank managers to help meet the germplasm users’demands.

Suggested action for next 10 years

Applicability

In situ Ex situ

Undertake gap analysis — All conservation requires a choice of the conserved populations, so the better the selectionstrategy for CWR taxa and populations, the more likely the conserved material is to be utilised. The process ofundertaking genetic resource gap analysis is discussed by Maxted et al. (2008b). When discussing CWR conservationpriorities it is advisable to consult potential germplasm users because the views of conservationists and end usersregarding the types of germplasm required may differ.

√ √

Undertake bioclimatic modelling — Gap analysis will help identify priority CWR species and reserve sites, but not allspecies or sites will be equally impacted by climate change, therefore bioclimatic modelling of the potential futuredistributions of CWR species and likely impact on protected areas is required.

√ √

Establish genetic reserve(s) and other in situ conservation areas — A gap analysis results in the identification ofpotential genetic reserve sites, but the sites then need to be actually established — too often site location alone isseen to be the goal. Iriondo et al. (2008a) provide guidelines on the how a genetic reserve might be designed,managed and monitored.

√

Write comprehensive reserve management plans — No genetic reserve should be established without carefulconsideration given to the completion of the management plan, one component of which should be a strategy forlinking the conserved diversity to use. Maxted et al. (2008c) provide a background to the writing of managementplans and how the plan might be linked to a conserved diversity utilisation strategy.

√

Carry out efficient population sampling — Where a sample of the conserved population is to be transferred to aremote location the collector must aim to collect a sample that represents the full range of genetic variation foundin the population and is of sufficient size to avoid the need for immediate regeneration. Guarino et al. (1995),Hawkes et al. (2000) and Smith et al. (2003) provide guidelines on efficient germplasm collection.

√

Collect associated materials — When sampling a population for ex situ conservation the use potential can beenhanced by collecting passport data, herbarium voucher specimens and possibly vegetative plants as well as seed.Although the specific data recorded by each collector is dependent on the target taxon and target area, an efficientcollector will always record certain information; see Guarino et al. (1995) and Smith et al. (2003).

√

Manage conserved germplasm effectively — Whether the sample is conserved in situ or ex situ, once the CWR diversityis actively conserved it should be managed to maximise the maintenance of genetic diversity. Samples from mis-managed collections are less likely to be of use and actually used. In situ, the appropriate management regimeneeds to be in place to maximise the maintenance of genetic diversity within the target taxon population. Ex situ,

√ √

Appendix 1. Suggestions for improvement of CWR conservation in the next 10 years.

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Publicise the conserved diversity — The collection manager can also draw attention to the existence of the diversity topotential users by ensuring the material is listed in appropriate databases (either in an institutional databasedirectly or through a link to a meta-database, such as SINGER (www.cgiar.org/singer/index.htm) or EURISCO(http://eurisco.ecpgr.org/). These points are discussed further by Hawkes et al. (2000) and Iriondo et al. (2008a).Following the conservation of germplasm, either in situ or ex situ, the commissioning agency will require some formof report; but in addition, the existence of novel diversity can be signalled to potential users by publishingcollection or protected area reports to increase potential utilisation and thus the value of the conserved diversity.The precise format of the publication will vary, but it may include details of target species ecogeography, siteselection and sampling strategy, the type of material conserved, characterisation and assessment of genetic erosion,and review of future conservation priorities.

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Duplicate the conserved diversity — There are two primary reasons to promote the duplication of conserved diversity:firstly, to provide a safety back-up of diversity in more than one genetic reserve and/or in more than one gene bankas a form of insurance against loss of diversity due to a catastrophic event; secondly, if the diversity is represented inmultiple ex situ collections it is more likely to be available to the national, regional and international usercommunities.

√ √

Carry out characterisation and evaluation — Any characterisation, evaluation or pre-breeding will enhance theutilisation of the conserved diversity. This may take the form of selection based on passport data (e.g., an accessionwith high drought or salt tolerance is likely to be useful in breeding new drought or salt tolerant crop varieties), orby carrying out characterisation followed by selection based on desirable features. More detailed evaluation fordrought or salt tolerance, or the deliberate infection of the material with diseases or pests to screen for particularbiotic resistances will obviously enhance the utilisation of accessions found to have desirable traits. It is advisable toinvolve potential germplasm users in the characterisation and evaluation process.

√ √

Develop core collections — With the increasing size of many collections and the ever-limited funds available forcharacterisation and evaluation, it may be necessary to develop a core collection of representative accessions toassist the potential user in the selection of accessions that will prove most useful. Hodgkin et al. (1995) provide adetailed discussion of the value and procedures involved in establishing a core collection.

√

Provide a quality service to users — The quality of the service provided to the germplasm user community will affectthe potential for utilisation. If requests for information and/or germplasm from the user community are efficientlydealt with, users are more likely to take advantage of the conserved diversity.

√ √

Seek opportunities to bring conservationists and germplasm users together — The people who conserve and utiliseCWR tend to work in two distinct professions and are often located in different institutes. Utilisation of CWR couldbe improved by bringing conservationists and germplasm users together when the opportunities arise.

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