Valuation of indigenous farm animal populations and breeds in comparison with imported exotic breeds with a focus on sub-Saharan Africa Henner Simianer (PD Dr) Applied Genetics Network, Pelargusstr 3, D-70180 Stuttgart Tel.: +49-711 6075067 Fax: +49-711 6075066 e-mail: simianer@genetics-network.de February 2000 Compiled for: Deutsche Gesellschaft für Technische Zusammenarbeit GmbH, Eschborn Funded through: Project for Promotion of Postgraduate Education in Agriculture in SADC (Project number 94 2072 001.200) In co-operation with: BMZ/GTZ Sector project: Managing Agrobiodiversity in Rural Areas (Project number 98.2020.0) PD Dr Henner Simianer, applied genetics network, Stuttgart, compiled this study with the assistance of Mr Alexander K. Kahi (MSc), on leave from Egerton University, Kenya, at present PhD student at the University of Hohenheim, Germany. The input of Mr Kahi, especially to sections on poultry, pigs, and unique traits and properties of indigenous breeds of this report, is gratefully acknowledged. Table of contents List of participants Executive Summary Introduction Valuation of farm animal genetic resources Definitions Valuation of biological diversity Biodiversity, species and breeds Economic values of farm animal genetic resources Economic valuation tools Cost-benefit approaches Risk-based approaches Completed and planned research on economic valuation of farm animal genetic resources Factors affecting results on the relative economic superiority of breeds or crosses Unit of comparison Limiting factors Genotype × management level interaction Data editing Performance comparisons of indigenous and exotic breeds and their crosses Cattle Dairy traits Beef traits Small ruminants Sheep Goats Pigs Poultry Chickens Guinea fowls Unique traits and properties of indigenous breeds Disease resistance or tolerance Trypanosomosis Gastro-intestinal parasites Ectoparasites Bursal disease Heat tolerance Cattle, sheep and goats Poultry Camel Adaptation and nutrition Other properties Salt tolerance Scavenging ability Reproductive performance Final remarks Valuation of farm animal genetic resources--prospects and possibilities Crossbreeding with exotic breeds as a threat to farm animal biodiversity Naive crossbreeding schemes Structured crossbreeding programmes Developments of synthetic breeds Decision making Conclusions References Appendix: AnGR valuation method evaluation (Drucker et al. 2001) List of abbreviations List of abbreviations AnGR (Farm) animal genetic resources BMZ Bundesministerium für Wirtschaftliche Zusammenarbeit CABI Commonwealth Agricultural Bureaux International CBD Convention on biological diversity Fi i-th crossbred generation FAO Food and Agriculture Organization of the United Nations FARM-Africa Food and Agricultural Research Management-Africa G × E Genotype by environment (interaction) GTZ Deutsche Gesellschaft für Technische Zusammenarbeit ILCA International Livestock Centre for Africa ILRI International Livestock Research Institute IPR Intellectual property rights KARI Kenya Agricultural Research Institute LW Large White breed Ne effective population size SACCAR Southern African Centre for Co-operation in Agricultural Research and Training SMS Safe minimum standard SSA Sub-Saharan Africa WTP Willingness to pay Executive summary Executive summary This study reviews the different concepts and approaches regarding economic valuation of farm animal genetic resources (AnGR) in the context of programmes for AnGR management to be implemented in sub-Saharan Africa (SSA). The different types of values and methods to estimate them are presented and critically discussed. Decisions related to AnGR management are not exclusively or even primarily of economic in nature, but other aspects, like socio-economic, livelihood or development implications are relevant. Nevertheless, all decisions and activities have economic implications, and it is this aspect the study focuses upon. The relative value of local breeds is observed especially in crossbreeding programmes with exotic, non-adapted breeds, which historically often have led to non-sustainable improvements and complete market failures. The respective studies are reviewed and discussed for the main farm animal species. Values of indigenous breeds are mainly due to unique traits and characteristics (like adaptive capacity, disease tolerance etc.), which are reviewed for the main farm animal species in SSA. Only few (actually two) studies have valued AnGR and few more are underway in developing countries, yet the need for such studies is considered and activities are prioritised. We argued that certain values, which are difficult to assess, have little relevance for conservation decisions. Therefore, and due to the relative lack of knowledge and of experience with respect to valuation methods, we do not recommended to conduct empirical valuation studies using some of the described methods. Alternative approaches have been suggested that focus more on risk of loss and cost of AnGR management, rather than on potential returns. These concepts are extensively discussed in the present study, since they may be a valid and pragmatic alternative to breed valuation studies. Risk-based approaches, like the safe minimum standard approach or the methods Smith (1985); Weitzman (1992) and Weitzman (1993) suggested require the assessment of risk functions and the cost-efficiency of AnGR management, i.e. the risk of extinction has to be quantified for each breed and the return (in terms of risk reduction) of different management activities has to be assessed. These data are generally lacking for developing countries. A further feature to be taken into consideration is the resilience of AnGR management. Respective activities should be designed in a way that even the total failure of some of the components will not lead to a total failure of the whole project, i.e. to the loss of a breed. Taking into account the high requirements of time, labour, money, and logistic input to conduct a sensible set of AnGR valuation studies, we argued that all related activities should be co- ordinated. Focusing on SSA, International Livestock Research Institute (ILRI) has taken the lead to initiate a set of studies based on different valuation approaches and targeting the different species in different regions. If Deutsche Gesellschaft für Technische Zusammenarbeit (GTZ) intends to start major activities in this field, projects should be designed to fit in the overall incomplete matrix of methods, regions and species to be built. We argued that such activities should be planned and carried out as co-ordinated effort in close linkage with other organisations, which are already active in this field. In SSA, ILRI clearly is the organisation with the highest competence and experience in the area of economic valuation of AnGR. Since ILRI activities are mainly based on contingent valuation methods, GTZ studies focusing on risk-based approaches would be a useful complementary effort. Since results of valuation studies are virtually non-existent and consequences of different activities for AnGR conservation are difficult to assess, decision makers could not be able to make specific recommendations. We argued that decision-making with respect to AnGR management activities should not be postponed until sound evidence is available. This could initially be made based on inadequate information by using, e.g. panels of experts to assess the relative values (or conservation priorities) of different breeds. Concerning future activities, the report recommends that highest priority be given to a region- and species-wide framework for decision making and priority setting, which combines information on values, conservation costs, diversity patterns and risk of extinction. Such a framework includes creation of public awareness, training and implementation of economic balancing mechanisms between stakeholders. Introduction Terms of reference Introduction There are numerous national, regional and global initiatives aiming at the management and conservation of farm animal genetic resources (AnGR). Respective activities are within the scope of the Southern African Centre for Co-operation in Agricultural Research and Training (SACCAR), which has, together with the Bundesministerium für Wirtschaftliche Zusammenarbeit/Deutsche Gesellschaft für Technische Zusammenarbeit (BMZ/GTZ) project Managing Agrobiodiversity in Rural Areas, commissioned this study. The latter project aims to contribute to a sustainable use of agrobiodiversity for food security and rural development, and to help reduce agrobiodiversity losses. Terms of reference Aims: Documentation of the actual state of economic valuation of indigenous farm animal populations Development of a model for relevant options of activities, which can be implemented in developing countries (with examples). Working plan: cf. 1: Review of existing data and current research projects or proposals concerning animal genetic resources and economic valuation of indigenous farm animal populations. Evaluation of competitiveness of indigenous vs. imported exotic breeds and crosses. Compilation of unique qualitative characteristics of indigenous populations and breeds (disease tolerance, adaptation etc.) that may determine the value of these breeds. Review of existing approaches for the economic valuation of biodiversity and their relevance for farm animal biodiversity and agrobiodiversity cf 2: Economic values for some farm animal populations have been determined in case studies and consequences for the management of these resources are discussed, taking the risk into account. Sectoral and global strategies are discussed, taking into account different scenarios (market failure, loss of unique characteristics). Based on the framework suggested by Weitzman 1992; Weitzman 1993 a methodological approach should be developed to assess the consequences of different options for farm animal biodiversity. In a world of limited financial resources, it is necessary to develop rules for decision making based on economic arguments, i.e. compare the relation between investments (e.g. for setting up a conservation scheme) and returns (e.g. through the exploitation of the conserved gene pool for some purpose in the future). For objective decisions, it is necessary that both the costs and the returns be quantified. Both due to the considerable time scale (conservation thinks in decades or even centuries, rather than years) and to the vast uncertainty of future changes and events, it is inherently difficult to derive the respective values. Therefore risk-based analyses have been suggested for the purpose of decision-making, which may avoid some of the problems associated with traditional cost-benefit analysis approaches. The different aspects of valuation are described in the first part of the study. The value of indigenous populations under tropical conditions is often most clearly expressed in comparisons with exotic breeds of European origin, both in pure breed comparisons or crosses. On the other side, crossbreeding is one of the major threats of the exotic gene pool, in that uncontrolled mixtures of indigenous and exotic genes displaces the exotic genotypes. In the second part of this report, literature on crossbreeding and breed comparisons in the main farm animal species under tropical condition is reviewed and interpreted from the point of view of genetic conservation. In many cases, indigenous breeds are highly adapted to the conditions they are exposed to, including climate, feed supply, disease challenge etc. The respective traits are in many cases unique and constitute a qualitative difference compared to exotic breeds or crosses, which hardly cope with the respective conditions. Such problems have in many cases led to failures of upgrading programmes, which aimed at improving productivity, but unintended led to a reduction of the adaptive capacity of the generated populations. The third part of this study reviews the main types of unique properties of indigenous breeds across all major farm animal species. The information presented and summarised in this study is intended to serve as a basis for the development of options for effective and efficient activities in AnGR conservation. Knowledge is far from being satisfactory or complete in different areas. Different farm animal species are studied with different intensity: in SSA, by far most information is available on cattle, while information on other important species like pigs, poultry and small ruminants is relatively scarce. Different aspects of methodology are applied with different emphasis: as Ruane (1999) stated molecular distancing seems to be the main activity in AnGR research, while phenotypic description or economic valuation of breeds is done with much less emphasis. While there is a lot of theory about the methodological approaches that can be used to valuate genetic resources, concrete studies on the economic valuation of AnGR are scarce. However, activities in this direction are considered and some pilot projects are started, the state of the discussion and the present and planned activities are discussed in this report. AnGR characterisation is mostly descriptive, while little or no conceptual work is done on how to decide which populations should be maintained, how limited resources should be allocated, and what measures are most effective to reach the often-not-well-defined goal of biodiversity conservation. Methodology development is mainly done for the conditions of the northern world, in that, e.g. conservation concepts are based on the conservation of breeds in the European sense. It must be doubted, whether these concepts can or should be used under the conditions of developing countries, where, e.g. breeds in this sense are often not implemented. It certainly is necessary to develop AnGR conservation strategies that are based on the actual conditions in developing countries. These aspects will be discussed in more detail in the conclusions of this study, taking into account the available information as summarised in the following chapters. Valuation of farm animal genetic resources Valuation of farm animal genetic resources Definitions Valuation of biological diversity Biodiversity, species and breeds Economic values of farm animal genetic resources Economic valuation tools Cost-benefit approaches Risk-based approaches Completed and planned research on economic valuation of farm animal genetic resources Factors affecting results on the relative economic superiority of breeds or crosses Unit of comparison Limiting factors Genotype × management level interaction Data editing Valuation of farm animal genetic resources Definitions Biodiversity can be defined as the diversity of biological units. A biological unit may be an individual or a group of individuals. In the very general sense, diversity1 may focus on very different attributes, like phenotypes (coat colour, body size), production traits (milk yield, growth), behaviour, nutritional aspects etc. Diversity is an attribute of a group. While two units (individuals, breeds etc.) may be different, a whole group (>2 units) is diverse. Genetics have revealed that the genetic code, as deposited in the nucleic and mitochondrial DNA, is the major criterion for the individuality, in other words, the genetic code is the blueprint for an individual, and differences between the DNA sequences of two individuals will to a certain extent lead to differences in the criteria mentioned above.2 Therefore, biodiversity studies prefer to work with genetic data (sequence data, allele frequencies etc.). Biodiversity in non-agricultural applications mostly operates on species, habitats, or ecosystems, but usually not on subgroups within species, like breeds, strains, or lines. Species is a taxonomic term and defines the largest group of individuals that can mate among each other with fertile offspring. Farm animal species are, e.g. horse, goat or chicken, but horse and ass are not the same species, since they can mate, but the offspring usually is not fertile. So, biologists focus their research on between species biodiversity (e.g. between different species of butterflies etc.). Agrobiodiversity, both in the context of farm animals and cultivated plants, usually focuses on within species biodiversity. Farm animal species are subdivided into breeds, which may be further subdivided in strains, lines, local populations etc. Breeds are not at all well defined, and historically, farm animal breeds are a European or at least northern concept in animal breeding, as they have been constituted through breed societies in the early 19th century, first in Great Britain, later in other European countries and in North America (Willham 1987). Lloyd-Jones (1915) already made the point that `... a breed is a group of domestic animals, termed such by common consent of the breeders. There as such is no sharp and objective definition of a breed, but it is a subjective classification.' Köhler-Rollefson (1997) suggested the following criteria for the definition of a breed, which may be applicable not only in the northern hemisphere with its traditional breed concept, but also on a global scale. A group of animals is considered as a breed, if it is subject to a common utilisation pattern shares a common habitat/distribution area represents largely a closed gene pool is regarded as `distinct' by their breeders. The general idea behind within species biodiversity can be demonstrated with the following example: suppose, a species is made up of three breeds, A, B, and C. The whole gene pool of the breed is the sum of all the alleles present in either A, B, or C. Typically, some of the alleles are present in all three breeds, others are represented in a subset of breeds (e.g. only in breed A and B), while some alleles are typical for a single breed and do not exist in the others. This unique set of alleles of, e.g. breed B would be lost if breed B was extinct. As a consequence, the between breed diversity would be reduced by this proportion. Finally, agrobiodiversity tends to use the term `resource' (e.g. AnGR). This is a somewhat unsharp terminology, which reflects the unavailability of a clear definition of breeds, strains, and other units used to substructure species. In an internal proposal, ILRI defines animal genetic resources as `those animal species and populations with them that are used, or may be used, for the production of food and agriculture. These populations within each species can be classified as wild and feral populations, landraces, and primary populations, standardised breeds or strains, selected lines, and any conserved genetic material.' Valuation of biological diversity Conservation of biological diversity, of which agrobiodiversity and even more specifically farm animal biodiversity is an important component, has been formally provided significant global importance and interest in the Convention on Biological Diversity (CBD), signed by 157 countries at the Rio Earth Summit 1992, and put into force in 1993. Among others, the importance of agrobiodiversity for food security, poverty alleviation, and improvement of livelihood are stressed. Valuation of specific segments of agrobiodiversity-in the case of AnGR the most common, though problematic unit, a breed-is an essential component of the whole complex of description of diversity, which is a prerequisite for any rational conservation strategy. However, purely economic considerations are not the only, probably also not the most important aspect in this context. Other aspects, like the contribution of a breed to food security, its ecological role, the insurance function of genetic diversity with respect to eventual future changes etc. may be seen as complementary arguments favouring conservation programmes. Environmental economists however have tried to set up a complete framework of different types of values, claiming that in principle all these values of, e.g. a breed can be quantified in economic terms, so that the whole problem can be discussed in a quantitative economic context. There is some dispute about this approach in the scientific community, and even those in favour of the `economic' approach are forced to admit that not all values can be quantified with sufficient accuracy (Figure 1) to be useful for decision making. Source: Adapted from Virchow 1999. Figure 1. Economic values of genetic resources. Valuing AnGR was preceded by major attempts to value plant genetic resources (PGR), especially crops, so that some methodological issues may be adopted from this field. PGR conservation has certain advantages compared to AnGR conservation, mainly the much less complex biology of plants, their high fecundity, the availability of wild ancestors, the ease and relative inexpensiveness of maintaining long-term seed collections etc. Somewhat simplifying, plant breeding (at least in some of the major crop species) can be seen as sort of experimental combinatorics, i.e. germplasm of different sources is combined and the properties of the hybrids are tested to select the best. Transgenic technology has added efficient tools to introgress single genes across species boundaries. These principal approaches should allow a relatively straightforward valuation of an additional source of germplasm. Despite this claimed relative ease, the economic quantification of the benefits of germplasm to crop development has been described to be still extremely difficult (Evenson et al. 1998). There are some examples of post festum valuation of single genes transferred (through traditional non-transgenic breeding techniques) from exotic resources to cultivated strains. A resistance gene against the yellow dwarf virus extracted from a single Ethiopian barley plant, for instance, has been found to generate a value of US$ 160 million after being transferred to cultivated barley strains (Munasinghe 1992). In the context of plant agrobiodiversity, Swanson (1998) stated that the fundamental nature of the public interest in genetic conservation is supplying those values that the private sector has little or no incentive to pursue. Thus, it is mainly the option value (with a strong emphasis on potential pest resistance) that is seen as the major motivation for plant agrobiodiversity conservation efforts. Another major field of valuation of genetic resources focuses on ecosystems, habitats and wildlife. In this context, some studies have focused on the valuation of large mammals (e.g. elephants) being a major component of safari tourism industry. The usual approach is to use the `willingness to pay', which empirically can be derived, e.g. from travelling costs, people are willing to pay to see elephants. Another approach uses survey based contingent valuation methods, where people are offered, e.g. safaris to parks with or without elephants to derive the marginal added value of elephants. Both types of studies have been successfully conducted (Brown and Henry 1989), and correspondingly resulted in a value/elephant per tourist-contact of US$ 100, leading to an overall economic contribution of elephants to the Kenyan economy of about US$ 25 million/year. Similar studies have been conducted to value entire habitats (Tobias and Mendelsohn 1991). Such approaches especially if they are based on survey techniques offering hypothetical choices, may suffer from biases emanating from discrepancies between claimed and realised willingness to pay. This is especially so, if the good to be valued is of high moral or aesthetical value or if it is thought to be politically correct to choose this good.3 Therefore, methodological approaches are more reliable that are based on empirical data on realised prices from market surveys, rather than on subjective `willingness to pay' data. Another lesson to learn from environmental economics aiming at wildlife and habitat valuation is that species should always be seen and valued as an element of a natural system, and part of the value of a species is derived from the interaction with the whole habitat. An example for such a systematic interaction is the Ronalds day sheep breed with its high copper absorption capacity and salt tolerance that has the ability to feed almost exclusively on seaweed and is an essential factor to maintain the seaweed habitat (Ponzoni 1997). In this case, the value of the breed should reflect the value of the whole habitat, which likely would not continue to exist if the breed was extinct. Similar interactions may be found between indigenous breeds and, e.g. forage plants that are more or less exclusively selected and fed by these animals, so that the feed usage maintains this plant which otherwise would disappear. Of course, there are also close links of indigenous breeds with human cultures, especially in pastoralist societies, so that a holistic approach has to be used to assess the value of a breed in such a system. Biodiversity, species and breeds In farm animal biodiversity studies, the general and widely accepted approach is to focus on breeds as the unit to preserve biodiversity. Under tropical conditions and especially in SSA, we must be aware that the breed concept is not established to a comparable extent for all farm animal species. While for Africa there are 120 cattle breeds on file (Simon 1999), the corresponding number of swine breeds in Africa is only 13 (Hammond and Leitch 1998). Does this mean that there is less genetic diversity in swine than in cattle? Most likely not, it may rather be seen as an indication that even in the non-Islamic African countries the breed concept is less adopted for swine than for cattle. About 50% of genetic diversity of farm animals is between breeds, while the other half of the diversity is within breeds of a respective species (Oldenbroek 1999). This, of course, relies on the assumption that species are stratified into breeds, which not necessarily is the case in less developed regions. As a consequence, a considerably larger share of the within species genetic diversity may be allocated within rather than between the existing breeds, if stratification into breeds is incomplete. Given a sufficient stratification into breeds, the two principal approaches to conservation of AnGR are to preserve breeds and to preserve diversity within breeds. This includes determining which breeds are different enough, so that their maintenance conserves a significant part of the between breed variation. Eding and Laval (1998) established and summarised the methodology to assess phenotypic and genetic diversity. Ruane (1998) suggested criteria for the selection of breeds for conservation. In a critical review, Ruane (1999) argued that biodiversity research focuses too much on distancing, i.e. on evaluating genetic distances between breeds, while too little work is done in the phenotypic description of breeds (including productive and adaptive performance). If breeds to be maintained are below a critical effective population size, measures have to be taken to prevent the loss of this breed or the uncontrolled incline of inbreeding in the breed. Meuwissen (1998) and Toro and Mäki-Tanila (1998) discussed there are multiple options for this task, ranging from cryoconservation to in vivo, in situ schemes. There is, however, a lack of theory and rules for decision making, which breeds should be maintained and when respective activities should start. Using an approach more targeted to species conservation rather than to breed conservation, Weitzman (1993) presented a formal decision-making framework based on a unique concept of diversity (Weitzman 1992). Both Solow et al. (1993) and Thaon d'Arnoldi et al. (1998) discussed a decision-making framework based on the farm animal context. It must be seen that preservation of biodiversity is different from preservation of individual breeds or populations. While each population may have a value as such, the contributions to biodiversity of different units within a species or a taxonomic class may differ. In a world of limited resources, it has to be decided how to allocate the available resources so that the defined goal is achieved with highest probability. As was shown by Weitzman (1993) it may not be the optimum strategy to try and preserve all populations, but it may in some cases be more rational to concentrate resources on the preservation of the major contributors to biodiversity. To adopt Weitzman's (1992) concept to farm animal conservation schemes, it would be necessary to quantify exactly a number of `soft' parameters, e.g. what is the probability p that a certain breed still exists in, e.g. 50 years, and what are the costs of increasing this probability from p to p + _? It should be noted that time horizons for genetic conservation approaches like, e.g. 50 years (as chosen by Weitzman 1992) correspond to quite different equivalents in generation intervals in farm animals, which range from less than one year in small but important species like poultry, rabbits etc. to up to six years in cattle. If, as is often done, the increase in inbreeding per year is seen as the main criterion for genetic erosion within breed, larger effective population sizes have to be maintained for farm animal species with shorter generation intervals. The biodiversity value of farm animal breeds The conventional approach to valuation of biodiversity conceptually uses the same principle to that used to decide, whether a company should continue to produce and sell a certain product. That is, actual and future costs and benefits are assessed, values for the future are discounted to corresponding prices of today, and then the decision is made. The major difference arising from valuating AnGR is that different types of values (Figure 1) have to be considered that it is difficult to attach monetary prices to these values, and that most of them will be realised in the (distant) future and with considerable uncertainty. One of the consequences of discounting is that high economic returns in the distant future have to compete with relatively low (but guaranteed) returns in the present or the foreseeable future. The straightforward application of the respective economic theory may be misleading. Clark (1973) showed that it was economically more profitable, to allow an extinction of blue whales (by selling `whaling licenses' to Japanese companies) and to invest the short-term profit into industries with high economic returns, like the computer industry, than to invest in whale conservation with some expected return in the distant future. While this analysis was principally correct, the result will not be acceptable for most people, not just environmentalists. For plants and wildlife, Ehrenfeld (1981); Ehrenfeld (1986) pointed out that certainly not all, possibly even not the majority of all existing species have a real value, neither at present nor ever in the future. Ehrenfeld (1981); Ehrenfeld (1986) called species without actual or future value non-resources and suggests that, e.g. many of the 600 thousand existing and described species of bugs will belong to this group. The problem, however, is that it is strictly impossible to know, which of the existing variety of species will have a value at some point in the future and which ones will be non-resources. Therefore it is not only necessary to discount future values to today's values, but also to weigh them with the probability that any of the set of species to be conserved will turn out to be a valuable resource. It must, however, be seen that some species potentially have an extremely high value. For the relatively recently discovered Mexican wild grass Zea diploperennis, which is closely related to corn, but is frost resistant, Fisher and Hahnemann (1984) have estimated (under certain assumptions) a potential value of US$ 6.82 billion. Similar returns might be expected from pharmaceuticals derived from natural contents of wild plants. It certainly is difficult to adopt these concepts directly for the situation in farm animal conservation, but some of the arguments deserve further consideration in this context. One major point is that the value is not only attached to any of the breeds in itself, but that there is an additional value, which results from the contribution of the breed to the diversity of the species. Weitzman (1993) gave an illustrative example based on crane conservation that may be translated to the farm animal situation as follows: consider a farm animal species to be a set of N sufficiently distinct breeds. The between breed diversity is to a large part due to two breeds, A and B that are quite similar to each other, but different from all other breeds (the respective phylogenic tree is depicted in Figure 2. a) Complete set b) A missing c) B missing d) A and B missing Let DABR be the diversity for the set of breeds A, B, and R standing for all breeds C to G, and let DÐ BR be the diversity for only B and R etc. Figure 2. Assumed phylogenic tree for seven breeds, and corresponding Weitzman (1992) diversity values Dx for the complete set (a) or for reduced sets (b-d). If, say, breed A is lost for some reason, the reduction of the diversity will be minor, because the genetically very similar breed B is still there. Thus, the relative contribution of A to the diversity of the species is minor, and so is the contribution of B under the assumption that A is still there. If, however, both A and B are lost, there may be a significant reduction of the diversity. This is shown in Figures 2a to 2d, where, as suggested by Weitzman (1992), a diversity measure Dx is used that is equal to the total height of the maximum likelihood phylogenic tree for the set of breeds x. Let us, for example, assume that DABR = 1.00, DÐ BR = 0.98, DA Ð R = 0.98 and DÐ Ð R = 0.80, i.e. diversity is almost complete, if either A or B are present, but is severely reduced, if both of them are lost. Then the relative value of the contribution of A to the whole diversity is (DABR - DÐ BR)/DABR = 0.02 if B is conserved and is (DA Ð R - DÐ Ð R)/DA Ð R = 0.184 if B is extinct. This means that the diversity value of breed A varies-based on what happens to the other breeds (in this example to breed B) in the species. The diversity value of A is almost ten-fold if B is extinct, compared to the situation when B is conserved. As a consequence, it is conceptually impossible, to assign values to individual breeds that reflect their relative impact on within species diversity without considering the genetic structure of all the other breeds in the species at the same time. This, after all, may be one important argument justifying the considerable number of genetic distancing studies for farm animal breeds that have been undertaken in recent years, since reliable distance matrices are an essential prerequisite for assessing a breed's contribution to within species diversity. In the present case, a conservation scheme should be aiming at the preservation of at least A or B. In what Weitzman (1993) called the `crowned-crane paradox' he demonstrated that it may be a more sensible decision to sacrifice one of the two (actually the more endangered one) and to spend all the available resources to ensure the conservation of the less endangered one, rather than to split the money to conserve both with an increased risk of losing both of them. Although the conceptual framework is very clear, applications of Weitzman's (1993) concept are not trivial since a considerable number of parameters have to be specified, like the probability for any one breed to be extinct at the end of the assumed planning horizon (e.g. 50 years from now), the cost of increasing this probability by a certain increment etc. Weitzman's (1992, 1993) theory is also tailored for species rather than for breeds, thus it totally neglects the estimated 50% of the within species diversity that are allocated within breeds, nor does it consider possibilities like genetic exchange between breeds, merging breeds etc. which are not relevant if species conservation is considered. Economic values of farm animal genetic resources While actual breeding programmes operate on costs and benefits related to production under market conditions valid today or in the (relatively) near future (Weller 1994), AnGR economics have a longer time horizon and are not restricted to production values only. Gandini and Oldenbroek (1999) gave the following list of possible objectives for AnGR conservation, which may be considered as a list of actual or future values: opportunities to meet future market demands insurance against future changes in production circumstances socio-economic value cultural and historic value ecological value opportunities for research. Virchow (1999) brought these components of value in a system of use- and non-use values with a suggested decline in quantifiability (Figure 1). Valuation obviously is easier and more reliable, if it refers to actual markets and prices. This implies that a higher degree of uncertainty is involved if valuation aims at future market situations. Valuation becomes even more speculative, if future changes in production circumstances, like environmental changes, changed consumers' wishes or newly arising disease challenges are considered. It must be questioned, whether there is a possibility to derive values for these items that are anything but speculative. A major set of methodologies used for AnGR valuation is based on cost-benefit analyses. In this context, asymmetric incurrence of costs and benefits is an important issue, i.e. which group of stakeholders will have to pay for AnGR management, and which group will have the benefit. Costs are associated with all in vitro and in vivo, ex situ conservation approaches, but also in vivo, in situ approaches will have explicit and/or implicit costs. In the context of developing countries, a cost-related decision affecting conservation matters may not be to replace a local breed through a (presumably) more productive exotic imported breed. The relative inferiority in productive efficiency are costs that are effective now and accrue to the farmer, while the benefit of not loosing the genes of the indigenous population are effective in the long-term and accrue to the `society', or even the `world as a whole' (Drucker et al. 2001). This aspect becomes even more relevant, if property issues are considered, with the key question: who owns AnGR. This question is intensively discussed in the field of plant genetic resources (Zilberman et al. 1998). According to Drucker et al. 2001, the two general market mechanisms in this field are intellectual property rights (IPR) and exploration and extraction contracts. An essential component in both approaches must be an adequate refunding for the initial provision and maintenance of AnGR to the individuals, societies, or states that are providing the actual `raw material', i.e. indigenous breeds with their respective adaptive capacity. Perrings et al. (1992) stated that establishing IPRs for genetic resources could be seen as a means for generating markets, which may lead to incentive effects beneficial for AnGR management. This asymmetry of cost and benefit allocation with respect to time scale (when do they become effective) and target groups (who pays and/or who takes the benefit) must be seen as one major driving force of AnGR loss. Other important driving forces on the macro level are habitat destruction, human population growth, and decision making processes including policy in general and impact of international markets, subsidies and developmental aid (Perrings et al. 1992). Economic valuation tools Cost-benefit approaches Mendelsohn (1999) summarised the set of methodological tools that might be suited to measure values of AnGR; most of them have not yet been used for this purpose, though. This set of methods contains the following approaches.4 The hedonic method is designed to value the observed traits of animals. This approach is based on empirical data on market sales and the characteristics of the sold animals or products. From these empirical data, values of new or alternative products or animals of breeds with specific characteristics can be assessed. Jabbar et al. (1998) reported applications of this approach for cattle breeds in West and central Africa and Jabbar (1998) reported the same applications for sheep and goats in southern Nigeria. Aggregate market analyses are based on information concerning aggregate quantities both on the input and output side of a system following a top-down approach. This information can be derived from empirical time series and/or panel data. The approach takes into account the multiple interdependencies of a production system, so that, in principle, it should be useful to evaluate, e.g. the consequences of a genetic improvement of a breed, taking into account all the corresponding changes like increased food demand, higher management requirements etc. It also reveals to whom the benefits of a potential genetic improvement will accrue. Cross-sectional household studies are based on aggregate demand functions derived from empirical data on consumer's behaviour in different markets. While the aggregate market analysis is based on empirical data from markets differing over time, cross- sectional household studies are based on markets differing geographically. These data may be easier to sample in studies focusing on developing countries. Together with supply functions, derived, e.g. with cross-sectional farm studies, this approach may allow welfare analyses on alternative scenarios. As mentioned before, the cross-sectional farm study allows derivation of supply functions from empirical data on farmer's decisions on price differences in different geographical locations. Contingent valuation methods are based on surveys. Consumers are offered a set of (real or theoretical) products with specific characteristics and are asked for the price they would be willing to pay (in currency or equivalent goods, e.g. sheep or cattle), to rank them or to make pair-wise choices. This approach is especially suited to assess non-use values (e.g. cultural values) or any hypothetic characteristic that is not yet available. Further useful approaches can be to do an economic input-output analysis of breeding programmes, farms or herds involved in animal production. Both are data-intensive and rely on a good quantitative knowledge of the genetic and biological processes underlying the respective systems. It must, however, be seen that examples for economic evaluation of actual breeding programmes are rare even for developed countries (Weller 1994) so that it will not be trivial to assess realistic input and output values for the much more volatile biological processes in developing countries. It must be stressed that the valuation tools that have been briefly introduced are mainly designed to assess direct use values. Most of the other value categories can hardly, if at all, be quantified through these approaches. This is more true the more abstract a characteristic to be valued becomes, the more distant in the future it might arise, and the more it depends on external changes. Some authors stress that there are principal limitations to what can be valued. Norton (1992) puts serious doubts on the question, whether a value for the `insurance' function of a conserved species will ever be anything but speculative as long as it is not clear, which characteristic of the species will be of value under which specific circumstances. Given these limitations, it should be clear that valuation efforts have to be undertaken to gain better insight on the economics of AnGR management. Nevertheless, conservation decisions will always have to be made under considerable uncertainty. Therefore, decision-making processes have to be designed to be robust towards limited or wrong perception of the economic background and consequences of the alternative options to chose from. The cost-benefit analysis tries to assign definitive numeric values to all the components of costs and benefits, and to probabilities for uncertain future events etc. Especially in the context of biodiversity management, the cost-benefit analysis has been criticised to be extremely speculative (Randall 1992) since many of the required quantifications will in most cases suffer extreme uncertainty. To give an example, option values of breeds are based on potential values of these breeds under unforeseeable changes in the future with respect to environmental changes, new diseases, changes in the markets etc. If, for example, a new infectious disease appears, some breeds may be genetically resistant or tolerant against this disease, which gives them an immanent value. However, not all (most likely only a minority) of breeds can be expected to realise their option value, but since we do not know the future conditions, we have to maintain, say, ten breeds to exploit the option value of one of them, not being able to identify which one. Risk-based approaches As an alternative, ecological economists have suggested the concept of `safe minimum standards' (SMS) (Ciriacy-Wantrup 1952), which starts from the assumption that in a certain case, there is biodiversity which is valuable enough not to be acceptable (for the society) and that the respective diversity would be lost. The problem then is to find (and quantify) the costs for the management of this diversity. Of course, it is still necessary to assign values to the required activities, and the objective is to develop a scenario with minimum costs that guarantees the maintenance of the diversity with sufficient probability. This scenario will then be adopted, unless the social costs are unacceptably large (Bishop 1978). Of course, there are some obvious and inherent problems with the SMS concept: e.g. when are social costs `unacceptably' high? And can maintenance of diversity ever be guaranteed by any plan of action? Crowards (1998) pointed out that following the SMS concept does not imply that a quantification of actual or future returns unnecessary. However, the SMS approach has some advantages over the cost-benefit approach in that it poses the null hypothesis on the assumption that diversity (e.g. a set of breeds) should be maintained and this hypothesis only will be rejected, if the society decides that it is not willing to pay the minimum costs required to achieve this goal. SMS therefore is much closer to a decision making tool than cost-benefit analysis, which is sort of vague in its objectives and priorities. The prejudiced approach of SMS, which is a priori in favour of conservation, is justified through the irreversibility of the loss, if diversity is not maintained (Bishop 1979). SMS has been used empirically for some wildlife species, e.g. the preservation of the Californian condor or the Rwanda mountain gorillas, leading to the conclusions that required costs were rather limited for the preservations activities (Bishop 1980). The concept to reduce the risk in the future by maintaining genetic diversity has been discussed by Smith (1985) for the context of farm animal breeding programmes. He introduced an extra discount rate for the remaining uncertainty, which is a function of the number of preserved selection stocks. While this discount rate drops with an increasing number of maintained stocks, the steepness and the shape of the declining curve may vary depending on the considered situation. For the European situation, Smith (1985) concluded that maintaining a sufficient number of reserve stocks is not only economically justified, but will be economically beneficial under a wide range of assumptions, as demonstrated in a sensitivity study. A generalisation of this approach to the AnGR management problem seems possible even though the approach should be based on a genetic valuation rather than on an economic valuation as in the original paper. This means the `currency' should be `useful genes' under actual or future environmental conditions. Let us assume that a given population has a set of Na polymorphic genes that are crucial for fitness and production under the actual environmental and management conditions, but that there are also Nf genes that are polymorphic and may be of some value in the future. Since future production conditions are unknown, it is uncertain which of the Na + Nf genes will be of any value. The objective must therefore be to preserve as large a share of the genetic diversity of these genes as possible under the given budget restrictions. Conceptually, it is possible to define samples of the population, like groups of, say, 100 females with the corresponding males at dispersed geographic spots, which are monitored on a very extensive level. Analogous to the approach of Smith (1985) it then would be necessary to compute the reduction of the risk of losing genetic variation if additional samples are added to the existing ones. To give a simplistic example: if we assume that at a given locus the rarest allele has an allele frequency of 1%, we expect from quantitative genetic theory (Falconer 1984) that with an effective population size of Ne = 50 the probability that this allele is lost in one generation due to genetic drift is 36.6%. The probability that the respective allele is still present after five generations is only 10.2%, that means the vast majority of rare alleles will be lost due to random genetic drift within a few generations. Figure 3 shows how this probability of loss is reduced if more than one sample of size Ne = 50 are maintained. However, the relative benefit of maintaining additional populations is variable depending on the frequency of rare alleles to be conserved. If the focus is on conservation of a very rare allele with p = 0.005, the benefit of an additional sample is almost linear, while for more frequent alleles (e.g. p = 0.02), there are diminishing returns. While the second sample increases the probability of conservation by almost 50% (49.1-74.1%), adding the fifth sample to already maintained four samples increases the conservation probability only from 93.3- 96.6%, i.e. by 3.3%. .Figure 3. Probability that an allele with initial frequency p is conserved over five subsequent generations, as a function of the number of samples with Ne = 50 in each sample. Considering the design of AnGR management programmes in developing countries, high weight should be given to the robustness of the implemented approach, i.e. the prevention of a total failure of the programme if the underlying assumptions are not met. This also has economic implications as may be shown with the following example: It may be decided to establish a conservation scheme for a breed based on the maintenance of four samples of animals of this breed. However, especially under the given conditions in developing countries it never is completely certain that a sample will not be lost within a given time frame. External changes (like environmental or political changes, theft, diseases etc.) and management failures may lead to partial or complete losses with a certain probability. Let us assume that in scenario A, all efforts are made to establish a scheme that the conservation probability over a given time period for any one sample is maximum, which is, say, 95%. In scenario B, much less effort is made to make sure that the samples are conserved over the chosen time period. While this probability for any one sample is only, say, 75%, it is chosen to start with six rather than four samples to balance this reduced individual probability. Figure 4 gives the probabilities for different numbers of conserved samples. While in scenario A the probability that four samples are conserved over the whole period is 81.45%, the probability that four or more samples are conserved in scenario B is 83.06% (sum of probabilities that four, five or six samples are conserved in scenario B). Figure 4. Probabilities for different numbers of conserved samples in the two scenarios. The expected outcome of the two scenarios thus is very similar, and thus the cost of the implementation of the two schemes will determine their relative superiority. In general, it must be assumed that increasing conservation security will be encumbered with over proportionally increasing cost, i.e. increasing the probability of a sample to survive over a given time period from 90-95% will be much more expensive than increasing the same probability from, say, 70- 75%. This is so, because in the latter case, it may be sufficient to improve a traditional management scheme by introducing some health services etc., while in the latter case it may be required to set up a centralised station with professional management staff etc. Figure 5 depicts this increase in costs (on an arbitrary scale) for a range of survival probabilities. Figure 5. Cost and expected return per sample as a function of the sample survival probability. The optimum strategy is to establish a sample survival probability so that the extra costs to conserve an additional sample are equal to the expected return in terms of conserved diversity. In the example shown, this is the case for a rather low sample survival probability of about 65%. As mentioned earlier, risk is another important aspect to be considered when establishing a conservation scheme. Let us consider a situation where we would aim at conserving 4.8 samples of a population. This can be achieved either by conserving eight samples with a sample survival probability of 60% or by conserving six samples with a sample survival probability of 80%. Although the expected number of conserved samples is identical, the strategy to conserve more samples allows a better dispersion of samples, e.g. across regions, environments etc., which reduces the risk of a total failure due to an epidemic, natural catastrophes, drought etc. Completed and planned research on economic valuation of farm animal genetic resources To date, the number of empirical studies that can be interpreted as valuations of AnGR is extremely limited. In fact, only the studies conducted by Jabbar et al. (1998) concerning small ruminants and cattle in West and central Africa can be counted as such. Even these studies initially had been conducted with a different intention of understanding market mechanisms in developing countries. The interpretation from the genetic resources point of view has been imposed a posteriori. Few other studies aiming at the valuation of AnGR are underway, e.g. a study valuing the criolo pig genetic resource in Mexico conducted by Drucker and Anderson (2000, personal communication) that will be finished and published later this year. Nevertheless, the toolbox of potential valuation methods is well described especially in the FAO/ILRI workshop proceedings (Rege 1999) and the survey presented by Drucker et al. (2001). Potential valuation methods including data requirements, data availability, strengths and weaknesses from this study are summarised in the Appendix of this report. n February 2000, ILRI invited an expert panel to a planning workshop to co-ordinate future research activities in this field. The main goal was to conduct a series of valuation studies on different farm animal species at different locations (with emphasis on SSA) and using different valuation tools. In the very intensive discussions, different aspects were seen quite controversial. First of all, the problem of biodiversity can be seen both from a global (species) perspective and from a breed perspective. The species perspective tries to valuate the contribution of a breed (or generally speaking of a group of animals within a species) to the within species biodiversity. Thus, this approach mainly focuses on the option value of a breed, i.e. its eventual `insurance' function against future changes in production environments, market preferences etc. Weitzman 1992; Weitzman 1993 suggested a theoretical framework for this type of perspective, which, however, was applied only once to farm animal data (Thaon d'Arnoldi et al. 1998). The breed perspective tries to validate the actual value of a breed with respect to productivity, non-consumptive and indirect use values etc. The approach is strictly empirical, in that values are derived from actual market data, surveys etc. Within this context, again two principles of valuation can be identified: Subjective valuation approaches based on data what stakeholders (animal breeders and owners, buyers of animals, buyers of products etc.) think a breed, an animal a product, a special characteristic of an animal or product is worth. These subjective valuations can be based on surveys or on empirical market data, which both reflect actual subjective valuations of goods or characteristics. Objective valuation approaches aiming at the identification of real values of certain characteristics (e.g. disease resistance) of an animal with respect to its productivity or of a breed within a production system. These studies are based on system analysis approaches, using farm or production system simulation to study the consequences of breed replacement in such complex systems. Both approaches do have their strengths and limitations. Subjective approaches may fail because the future developments may not be adequately taken into account by today's stakeholder's judgements or market decisions. In Europe, there are quite a few examples where breeds were highly valued by breeders, but subsequently were almost totally replaced within very short time as a consequence of changes in the market or the political system.5 Objective approaches based on farm or production system simulations require a very precise and realistic knowledge of the relevant biological and economic processes involved. Many of these processes are not very well known, especially in the production systems of developing countries, so that wrong assumptions may lead to wrong conclusions. ILRI plans to implement a research focus entitled developing and testing tools for economic valuation of AnGR. This initiative has received an external funding of about US$ 300 thousand to conduct a first phase. The studies planned so far by ILRI for the first phase of the project mainly focus on the breed perspective and are mostly based on subjective approaches. This is mainly due to the fact that results have to be obtained within a rather limited period (until end of 2000). We, therefore, planned to base the majority of the studies on available data on market surveys etc. Developing realistic models for the objective approach is quite demanding and likely not feasible within the given time frame of the first phase. However, there is a strong group stressing the need to also take the global perspective, i.e. to assess the contribution of breeds to the within species biodiversity. We decided to conduct a case study on diversity of African taurine breeds, based on genetic distance data and data on degree of endangerment of breeds available at ILRI. The case study will yield two types of results: A quantification of marginal contributions of the considered breeds to the actual and future within species biodiversity. Estimated marginal benefits of investments to AnGR management and optimal allocation of financial resources to conserve a maximum proportion of within species biodiversity. This case study will also help to identify research priorities to generate empirical data required for the implementation of the global approach. Based on the results of the case study and the integration of all available data, it will be attempted to establish a more systematic approach aiming at the development of a decision making tool for global policy makers. Both approaches, the more global species perspective and the breed perspective are complementary, in that the economic valuation of breeds provides essential data to be used for within species decision making. However, knowing all actual use values of the breeds forming a species will not allow to make rational and optimal conservation decision, since clearly additional information, mainly on the genetic structure of the species, the different degrees of endangerment, and (in the case of limited resources), the respective conservation costs and efficiencies for different breeds have to be taken into account. Therefore both approaches have to be pursued simultaneously. To derive the methodological framework for the global perspective is mostly desk-work and an informal group of scientists (J. Gibson, E. Rege, O. Hanotte, all ILRI; T. Meuwissen, Lelystad, E. van Dusen, Davis, C. Wollny, GTZ; H. Simianer, Stuttgart) have already agreed to go forward in this direction and conduct a case study to develop a project proposal to develop and implement this framework with the respective operational tools. Breed valuation studies have to be planned and carried out systematically with respect to methodological approaches, species and breeds within species considered, and geographical distribution of studies. The optimum design would be a three-dimensional matrix study with methodology, species and geographical region as the three main factors. This goal will not be achieved in the first phase of the ILRI project (which focuses to a large extent on ruminants in SSA plus one study on pigs in Latin America). Since experience with different methods for valuation of AnGR is scarce, it remains to be evaluated if the approaches (especially the survey-based methods) are useful or even feasible in different socio-economic contexts. A survey approach that works well in one situation may entirely fail in another context, since, e.g. the ability to abstract is necessary to value `hypothetical' properties that are present to a different extent in different groups of interviewees. Given the amount of resources required for a sound study (realistically about six to twelve scientist months plus additional personnel, transport etc. for the surveys) it must be doubted that a totally comprehensive matrix approach can be put on the ground. In any case, this is a very convincing argument for a concerted action with a joint and co-ordinated effort. Factors affecting results on the relative economic superiority of breeds or crosses When comparing the relative economic value of breeds or breed crosses under specific conditions, a number of factors have to be taken into account. Unit of comparison First of all, the unit of comparison has to be specified. This can be, e.g. an individual (or even an individual in a given time interval, for instance a lactation period), a herd or a whole production system. As Kahi et al. (1998) pointed out the relative value of imported animals with a higher live weight and a higher absolute productivity per animal will be overestimated as compared to smaller size indigenous breeds if the comparison is made on the individual level. If, however, the comparison is made on a herd level with the main constraint on available feed, light animals will appear more competitive because the ratio of produced output (e.g. milk or meat) to the animal's maintenance requirements are higher for smaller animals. When comparing different types of crossbreeding systems (inter se matings, rotations, crisscross etc.) it must be taken into account that some of these schemes require the continuous availability of purebred males and/or females of the underlying exotic breeds. In this case, the total economic valuation has to take into account the costs of maintaining these animals or alternatively the costs of purchasing semen or live animals to be brought into the system. While in many cases straight F1 animals of a Bos indicus × Bos taurus (e.g. Sahiwal × Friesian) cross have the highest expected performance, the low performance of the pure B. indicus cows that have to be maintained as mothers lowers the productivity of a system based on continuous replacement of F1-genotypes. This was shown in an analytical comparison of different crossbreeding schemes (Kahi et al. 2000c) where a three-breed rotation (Sahiwal, Brown Swiss, Friesian) clearly was the method of choice when the productivity of the entire system was used as criterion, while on an individual base the highest productivity was achieved with F1 Friesian × Sahiwal animals. Limiting factors Kahi et al. (1998) also argued that the most informative comparison of breeds or crosses is made, if productivity is set in relation to the main limiting factor of the production system. If, for example, the availability of concentrates is the limiting factor, the net profit per kg of concentrates should be considered. If, alternatively, available land for pastures is limited, the productivity per unit of land should be taken as the criterion for breed comparison. Kahi et al. (1998) have shown that the application of these different criteria leads to quite different rankings of the genetic groups compared. Genotype × management level interaction Madalena et al. (1990) considered the relative superiority of various grades of B. taurus × B. indicus crosses (Brazilian Guzera × Holstein Friesian) on high and low management levels and observed considerable re-rankings of the crosses. In Figure 6, the production criterion `kg milk per day of calving interval' for the second lactation is given for various proportions (25- 100%) of Holstein genes on two different management levels. In this study, management levels differed both with respect to feed and pasture management and with respect to veterinary supervision and parasite control. The high management group animals were virtually free of ticks, while the low management group animals were not. Source: Madalena et al. 1990. Figure 6. Milk yield (second lactation) per calving interval (kg/day) of different crossbred genotypes under low and high management conditions. On the high management level, pure Holsteins and ¾ or Holsteins perform best and are clearly better than the F1, there is a distinct superiority of the crosses under less advantageous management conditions. Similar significant genotype × management interaction effects were found for all major production criteria in this study. For organisational reasons, breed comparisons are often made on research stations or larger farms with a relative good management and an above average production level. Even the fact to be included in a respective study or survey may positively reflect on the management and the production level during the course of the study. As the results of Madalena et al. (1990) show, the differences found under these conditions do not necessarily apply to lower management and production levels, which may however be more typical for the production system studied. Data editing Madalena (1989) also pointed to the fact that breed or breed cross comparisons may be biased due to selective data editing, e.g. by eliminating extremely short lactations. As a consequence, genotypes that are not able to enter into a normal lactation profile in harsh environments due to insufficient adaptive capacity will benefit from the exclusion of `abnormal' lactations. Rege et al. (1994) reported extreme variability of production traits of exotic crosses, like lactation performances ranging from 5 kg in 9 lactation days to 3059 kg produced over 454 days with a small proportion of cows producing more than 100 kg per lactation in that study. This is not only a result of the genetic capacity of the breed or cross, but again has an interaction with the management, since indigenous breeds tend to go dry within a few weeks if not allowed to suckle. 1. Note that diversity is also relevant for non-biological objects like buildings, languages, landscapes etc. Thus, a major part of `diversity theory' has been developed in non- biological contexts. 2. However, the opposite is not necessarily true, as can be seen with monozygotic twins, which have identical DNA, but still may differ in many ways. This is due to the fact that on average the larger part of phenotypic variability is due to environmental factors. However, sustainable diversity, i.e. diversity that will be passed over to subsequent generations is mainly genetic diversity, and hence, this one is accepted as the major criterion in many diversity studies. 3. Such discrepancies have frequently been reported in market analyses, where potential consumers were asked, how much extra price they would be willing to pay, e.g. for `ecologically produced' goods, which largely overestimated the achievable price in the market (and subsequently often led to market failures). 4. This list is largely a summary of Mendelsohn's (1999) presentation of the topic. 5. See, e.g. the fate of the `SMR' synthetic cattle breed in the former GDR, which was almost entirely replaced through Holstein as soon as the market for animals and concentrates was liberalised. Performance comparisons of indigenous and exotic breeds and their crosses Performance comparisons of indigenous and exotic breeds and their crosses Cattle Dairy traits Beef traits Small ruminants Sheep Goats Pigs Poultry Chickens Guinea fowls Performance comparisons of indigenous and exotic breeds and their crosses Cattle With respect to the general performance of exotic cattle (B. taurus type of European origin) in the tropics, de Vaccaro (1990) summarised that on almost every criterion of survival, European cattle in the tropics did less well than their crosses with Zebu. The impact of heterosis is shown by the stayability data summarised in Figure 7, showing that crosses with 50-62.5% exotic genes achieved on average more than twice the number of lactations (6.6) than purebred European cattle (2.9 lactations). Figure 7. Average herd life (in lactations) of European purebred cattle and different grades of crossbreds with zebu. Another interesting result is that imported purebred exotics produced only 0.74 heifer replacements, i.e. female offspring alive at first calving, while locally born purebred exotics produced 0.98 heifer replacements (de Vaccaro 1990). This indicates that in the experiments studied imported pure-bred exotics could not sustain and continuous importation was necessary to maintain the populations. A relevant alternative to continuous crossbreeding schemes is to establish `new' synthetic breeds with gene contributions from different sources, in most cases mixtures of B. indicus and B. taurus. The great advantage of such synthetics is that they are genetically more stable and uniform and that selection may operate on establishing epistatic gene complexes with desired effects. Developing such a synthetic, however, is a complex and demanding long-term exercise. Most of the relevant synthetic strains thus have been developed in the US (e.g. Santa Gertrudis, Beefmaster, Brangus, Charbray), Latin America (e.g. Indubrasil, Occampo, Jamaica Hope), Australia (e.g. Droughtmaster, Quasar) and South Africa (Bonsmara). There are, however, also risks attached to this strategy. Development of a synthetic breed implies to go through a `genetic bottleneck', which may have consequences for the performance in certain traits, especially related to fitness and reproduction. For Santa Gertrudis cattle (made up from Shorthorn and Zebu) there are reports of poor libido of bulls and long calving intervals in females while Jamaica Hope (½ Jersey, ½ Zebu), suffers from low reproductive efficiency (McDowell and Jones, 1972). Dairy traits In a recent and comprehensive review of the relevant literature, Rege (1998) performed a meta-analysis on the results of 80 reports on crossbreeding in the tropics, including Africa, Asia and Latin America. The least square means for the main traits milk yield, calving interval and age at first calving are depicted in Figure 8. The trait values are expressed relative to the trait value of the F1 (50% exotic genes), which was set to 100. Source: Rege 1998. Figure 8. Relative values (F1 = 100) of the least square means of results from 80 studies on crossbred performance for the traits: calving interval, age at first calving and milk yield. For milk yield, there is a considerable improvement of performance when the percentage of exotic genes is increased from 0-50%, while the average stays approximately on a constant level between 50-100% exotic germplasm. On the individual level, crossbreds with a similar share of exotic genes, but resulting from different crossbreeding schemes (F1, synthetics, rotation) were found to be not much different with respect to milk yield (Kahi et al. 2000a). The situation is similar for the trait age at first calving, where a considerable improvement is observed when increasing the proportion of exotic genes from 0-50%, while the trait remains on a comparable level for higher proportions of exotics. The shortest calving intervals are observed for animals with 50% exotic genes, while calving intervals are longer both for lower or higher grades of crossbreeding. There is, however, a considerable variability between exotic breeds used for the crosses. For the trait milk yield, Brown Swiss crosses outperformed most other crosses over a wide range of crossbreeding grades, while Jersey crosses appeared to be inferior. Crosses with Holstein Friesians and Red Danes were intermediate for intermediate grades of crosses but performed best with high proportions of exotic genes (>). Lactation length is a crucial trait for the overall performance of exotic crosses under tropical conditions. Rege (1998) reviewed that across breeds, lactation length is linearly increased by 6 days with a 10% increase of exotic germplasm in the cross. This, again, is depending on the exotic breeds used. While the increase is similar to the overall picture with Holsteins, Red Danes and Jersey, the average lactation length stays on a low level (260 to 270 days) for Brown Swiss crosses over a wide range of crossbreeding grades. Generally, F1 animals tend to be better than subsequent inter se crosses. While there are numerous reports on the performance of F2 animals, empirical results on F3+ generations are limited. In his literature review, Rege (1998) found considerable average heterosis of up to 40% of the pure parent breed average for Brown Swiss × indigenous breed crosses for the trait milk yield. Sharma and Pirchner (1991) reported 13-18% heterosis for milk yield traits in Sahiwal × Holstein crosses in India, but 30% heterosis for the length of the dry period. Kahi et al. (2000b) found significant heterotic effects of about 15% of the mid-parent value for milk yield in Sahiwal × Brown Swiss crosses, but no positive heterosis for Sahiwal × Ayrshire. Due to recombination loss, this heterotic superiority in the F1 animals is expected to break down in subsequent generations, which is confirmed by findings of Syrstad (1989), who reported considerable declines for all relevant dairy traits when comparing the F2 with the F1 generation. In a literature review, Syrstad (1990) found a comparable development of the relative performance of crossbreds at different grades as Rege (1998) reported (see Figure 9), however when comparing the F2 (inter se crosses of F1 animals) with the F1, he reported a reduction by about 25%. Source: From literature reviews of Syrstad (1990) and Rege (1998). Figure 9. Relative values (F1 = 100) of the least square means for the trait milk yield; F2 average separately indicated. For the trait milk yield of a B. indicus × B. taurus cross, Mackinnon et al. (1996) reported a recombination loss of 679 kg, which exceeds the reported heterosis of 616 kg (20% of the mid-parent value). Similar results were obtained for other dairy traits, e.g. milk yield per day of lactation or per day of calving interval. Both Syrstad (1989); Mackinnon et al. (1996) stated that the superiority of crossbreds is mainly due to the high heterozygosity in combination with dominance effects. This has consequences for the relative benefits and sustainability of crossbreeding schemes. McDowell (1985) stated that `the real challenge is to establish breeding programmes that retain the merits of the first cross'. Madalena et al. (1990) concluded that a continuous F1 heifer replacement strategy has the highest expected benefit, but has also the highest implementation costs. Crisscross schemes (with 50% or 67% exotic genes) are also competitive and may be organisationally less demanding, while continuous upgrading or synthetic breeds through repeated inter se crossing are less advantageous. A continuous production of F1 females through the use of embryo transfer technology, as considered by Teodoro et al. (1996), may be an option for some more developed countries in Latin America. A sustainable implementation of such a scheme must however be doubted in the developing countries in SSA. Beef traits Data on beef production with indigenous breeds vs. exotic crosses under tropical conditions are very limited compared to those on dairy production, especially since performance comparisons of crosses with the pure breeds under the same environmental conditions are scarce (McDowell and Jones, 1972; Mpofu et al. 1998) McDowell (1985) summarised the results of published studies on performance of crossbreds under tropical or subtropical conditions conducted mainly in the southern parts of the US, South America and Northern Australia (Queensland). He reports that superiority of crossbreds over the parent breed average are observed growth rate to 15 month of age (11% heterosis), but no heterosis was observed for carcass traits. He also points to the fact that reciprocal crosses may differ due to the significance of maternal effects. In Queensland, crossbred mothers showed better maternal abilities than imported exotics (75% vs. 56% live calves for crossbreds vs. pure exotic--Shorthorn and Hereford--cows). As a consequence of improved viability, growth rate and reduced cost for tick control, McCarthy and Hodgson (1970) estimated 11.3% higher returns from European × Zebu crosses compared to pure Herefords under Queensland conditions. Chantalakhana (1998) reported experiences with imported cattle and crosses used for beef production in South-East Asia. He refers to factors like tropical disease and parasite challenge, poor management, low quality food and especially marketing and economic problems that pose considerable limitations on the use of upgraded animals. Cattle have multi- purpose functions, being not only used for meat production, but also for work, utilisation of farm wastes and natural pastures. Cattle also play a considerable socio-economic role-being used as long-term savings, and an essential element of ceremonies in some regions (Jabbar et al. 1998). Increasing occurrence of theft and cattle smuggling across national borders has the potential to hinder sound development of profitable (or at least sustainable) beef industry based on upgraded genotypes both in South-East Asia (Chantalakhana 1998) and in the SADC region (Wollny 1999, personal communication). In the semi-arid African subtropics of Botswana, Mpofu et al. (1998) compared performance of crosses of indigenous cows (Tswana or Tuli) with exotic sires of the breeds Charolais, South Devon, Sussex and Santa Gertrudis with purebred Brahman animals. For birth weight, weaning weight and weight at 18 months, all crosses outperformed the Brahman purebreds by up to 25% (Figure 10). Especially for the later weight traits recorded at weaning and at 18 months of age, the Charolais and South Devon crosses appear to be clearly superior compared to the other crosses and compared to the Brahman. The authors suggest that it may be possible that crosses are reared and fattened more intensively and slaughtered at a younger age (than the usual 36 to 48 months) where the required improved management level can be implemented. Source: Mpofu et al. 1998. Figure 10. Beef performance traits for four different crosses relative to pure bred Brahmans in semi-arid Africa. Mpofu et al. (1998) also reported that genotype-environment interaction favoured the smaller, slow growing genotypes on the poorer agro-ecological locations and in dry years. For cow live weight in crosses of Sahiwal with Ayrshire, Brown Swiss raised in the tropical lowlands of Kenya, Kahi et al. (2000b) reported consistently positive, but mostly not significant heterotic effects. The heterotic effect of about 5% of the mid-parent value for Sahiwal × Brown Swiss crosses however is exceeded by an even higher recombination loss. In a comparative study of different indigenous breeds and exotic crosses for beef production traits in semi-arid Zimbabwe, Moyo et al. (1994) found little evidence for a general superiority of exotic crosses over highly adapted indigenous breeds and concluded that indigenous cattle should play a major role in livestock development programmes in the region. Experiences from Northern Australia show that long-term selection in B. taurus type cattle populations may lead to significant improvements in beef production performance (+10-20%) and in adaptation characteristics (tick count reduced by 80%) over a 20 year period (O'Neill and Frisch 1998). Small ruminants For small ruminants, most scientific studies on the effects of crossbreeding have been done with breeds and in locations of temperate zones. If studies were undertaken under tropical conditions, management level was typically high. Data on crossbred performance under smallholder situations are virtually not existent in the scientific literature. In a comprehensive review of crossbreeding studies with sheep in temperate climates, Young et al. (1986) concluded that heterosis retention was significant, but differing with respect to the respective combination of breeds and crossbreeding schemes, so that the optimum scheme has to be determined specifically for any project based on specific combinations. In general, crossbreeding and use of `exotic' breeds is seen as a means to improve production efficiency in temperate zones (Hohenboken 1986). For New Zealand, different economic studies reported high expected returns from a systematic import of exotic breeds (like the Finnsheep), with predicted internal rates of return on public investment between 27% (Sorrensen and Scott 1978) and over 80% (Bushnell and Hutton 1982). Sheep and goat genetic resources account for 62% of the total domesticated ruminant livestock in SSA (Lebbie and Ramsay 1999). Lebbie et al. (1996) listed 61 sheep and 42 goat genotypes for which the real genetic distinctiveness, particularly of the indigenous ones, is not known. In many cases, the breeds are named after ethnic groups (e.g. the Red Maasai sheep) or regions (e.g. West African Dwarf), which says little about there distinctiveness. The role of small ruminants in these agricultural and eco-systems thus cannot be overestimated. Small ruminants both are an indispensable component to environmental and farming systems and in many cases are already or become a threat (through overstocking and overgrazing) of these ecosystems in tropical SSA (El Aich and Waterhouse 1999). Sheep Charray et al. (1992) stressed that genotype-environment interaction with respect to adaptive traits is the main obstacle to successful crossbreeding with exotic sheep breeds. In Ghana, the Nungua Blackhead, a cross of the local West African Dwarf with Blackhead Persian, is known to produce heavier lambs at weaning than the pure-bred Dwarf lambs. However, this advantage is nullified by the fact that litter size of West African Dwarf ewes is larger than that of Nungua ewes. An attempt to cross West African Dwarf sheep with Sahelian sheep in Côte d'Ivoire had to be abandoned in the first year due to high lamb mortality. Similar problems have been encountered with the same cross in Ghana. Osuagwuh et al. (1980) reported a 10% incidence of dystocia and 14% perinatal mortality when West African Dwarf ewes were mated with Permer, Uda or Yankasa rams, while no such problems occurred when the ewes were mated to rams of the same breed. Steele (1983) reported an improved growth rate and wool yield in the Sultanate Oman, when the indigenous Omani breed was crossed with Mediterranean Chios sheep. However, performance of the crossbreds was only acceptable under good management conditions, while poor management conditions led to considerable losses. In South Africa, composites based on Finnsheep × indigenous crosses compared to local Dorper animals were smaller, more fecund and produced faster growing lambs (Schoeman and van der Merwe 1994). The overall efficiency of the composites was 84% above the one achieved with purebred indigenous. However, this and most other results reported in literature refer to performance under station conditions. These may be far away from the conditions given in smallholder farming systems, and it must be doubted whether the results can be generalised in this respect. In Barbados, crosses of Barbados Blackbelly hair sheep with European wool sheep (Suffolk and Dorset) showed low fertility in both sexes, while the carcass traits were similar to the pure-bred indigenous animals (Patterson 1983). In India, about 50% of the variability of wool quality (fibre diameter) could be attributed to the genetic grade level in a crossbred population of indigenous Nali sheep with exotic Russian Merino and Corriedale sheep (Amar et al. 1998). In sheep, repeated inter se matings with selection to create `synthetic' breeds have been exercised, e.g. in India (Avikalin breed) or South Africa (Dorper breed). Again, the development of a new breed or establishing and maintaining a complex mating scheme (such as crisscross or three breed rotation) requires considerable infrastructure and has a high risk of failure. Additionally, populations go through a genetic bottleneck in the formation of synthetics, with a high risk of loss of genes of potential future importance. Like in other farm animal species, F1 animals are reported to show the highest heterosis, which is lost in subsequent inter se matings. Boujenane and Chafik (1994) reported experiences with D'Man × Sardi crosses in Morocco, where an almost 20% heterosis for number of lambs born in the F1 is reduced to 13% and 5% in the F2 and F3, respectively. Indigenous local breeds with special characteristics in one or few traits may be seen as a good resource for improving breeds in the same region through crossbreeding. Mason (1980) suggested using the prolific West African Dwarf sheep to increase the reproductive rate of larger, less prolific breeds in West Africa. Such a cross would have the advantage that the `exotic' breed is adapted to a similar environment and has comparable `hardiness'. In some regions, like in the tsetse-infested rainforest region of western Africa, indigenous trypanotolerant breeds cannot be replaced by exotics, but may act as the base for an upgrading programme with exotic breeds of higher productivity, while the trypanotolerance of the indigenous breeds has to be maintained (Devendra and McLeroy 1982). Indigenous sheep breeds seem to be less threatened than breeds from other farm animal species. In general, sheep use to live in unmodified harsh environments and large-scale genotype replacement is very slow (Gatenby 1986). This, however, is not necessarily true for all regions in SSA. McDowell and Jones (1972) referred to the failure of government breeding programmes in North Africa, resulting from the fact that nomadic shepherds normally are not willing to accept rams that have been `artificially reared'. On the other hand, King et al. (1984) reported crossbreeding with Blackhead Persian and Dorper rams into Maasai flocks, which has been carried out by the stock owners without external encouragement, e.g. through development agencies. Goats According to Shelton (1986), the value obtained from goat milk production exceeds the one derived from other products (meat and fibre) on a world wide scale, although only a small proportion of the world's goat population is milked. This is due to the fact that milk production is a continuous process, while meat production involves slaughtering of the animal and fibre production is done with only a very small proportion of all goats. Most studies in the literature have been conducted in Asia, especially in India, while reports from Africa are scarce. When expressed per unit of input (e.g. per kg dry matter of feed), goats may even outperform cattle, buffaloes and sheep under comparable environmental conditions in the tropics (Knights and Garcia 1997). A review study of Sahni and Chawla (1982) compared native breeds and crosses with exotics (mostly Saanen and Alpine goats). With ½ to exotic crosses, milk production is increased by 100% to almost 150%, with an apparent optimum of ¾ crosses. It should, however, be noted that these results are based on a limited number of studies (26 in total). More detailed results on crossbreeding experiments with Indian Beetal goats and exotic breeds (Saanen and Alpine) are depicted in Figure 11. For milk yield, there is a clear heterotic effect of more than 20% of the mid-parent value, which erodes if the proportion of exotic genes is increased to 75%. For lactation length, however, the ¾ exotic genotypes show the best performance, which is also true for the kidding interval, where the ¾ exotics have the shortest interval. Devendra and Burns (1970) reported similar results for crosses of Malaysian indigenous breeds with Anglo Nubians. Source: Sahni and Chawla 1982. Figure 11. Performance of pure indigenous (Beetal), pure exotic (Alpine and Saanen) and ½ and ¾ crosses for the traits: lactation milk yield, lactation length and kidding interval, expressed as per cent of the pure line average. In SSA, a number of dairy goat crossbreeding schemes are under way or have been conducted with `improver' breeds like the Toggenburg, Alpines, Saanen and Anglo Nubian. These projects utilise the genetic adaptation of indigenous breeds in terms of hardiness, disease tolerance and resistance to exploit the productive superiority of the more disease susceptible and less adapted exotic breeds (Lebbie and Ramsay 1999). Examples for such projects are the FARM-Africa Dairy Goat Development Project in eastern Ethiopia (Peacock et al. 1990), the Ngozi Goat Development Project in Burundi (Rey and Jacob 1991) and the Small Ruminant Collaborative Research Support Program Dual Purpose Goat Initiative in Kenya (Semenye et al. 1989). In other cases, composite or synthetic breeds like the Kenya dual purpose (East African × Toggenburg × Galla × Anglo Nubian) or the Tanzania Blended (Boer × East African × Kamorai) have been successfully developed. The majority of goats are exploited only for meat production. In some tropical regions, however, goat meat is hardly competitive to other sources of meat, due to low dressing percentages, small and bony pieces of meat etc. The major benefit of goats for meat production therefore must be seen in their excellent adaptation to poor environments and the high reproductive efficiency of some breeds at least. Singh and Singar (1981) concluded that while larger breeds are superior when growth rates or body weights at a given age are considered, smaller breeds were more efficient if reproduction was taken into account and meat production per unit of input was measured. A systematic diallel crossbreeding study using the Tennessee stiff-legged, the Spanish and the Nubian goat breeds under hot and dry conditions in the Southwest of the United States revealed no significant improvement of Nubian × Western breed crosses compared to pure- bred Nubians in growth and carcass traits, while Boer goat × Tennessee stiff-legged and Boer × Spanish performed well under these conditions (Dzakuma et al. 1998). There are considerable differences in breeds with respect to the different traits of interest. A crossbreeding scheme to produce F1 animals from highly fertile females (e.g. Black Bengal) with large fleshy males (e.g. Boer or Anglo Nubian) should lead to well adapted and sufficiently productive stocks. However, such a scheme requires considerable infrastructure to maintain the pure lines, which in many cases will be difficult to establish under smallholder management schemes. Pigs As already alluded to, SSA is the home of 13 pig genetic resources (Hammond and Leitch 1998). While pig genetic resources have been described for some of the tropics (Alba 1972; Rigor and Kroeske 1972), there have been few (mostly from Nigeria) detailed and accurate studies on the performance of the SSA's indigenous pig genetic resources (Ilori et al. 1976; Chiboka 1981; Somade 1985; Somade and Makinde 1985). Most of the indigenous pig breeds are kept under village management and are normally for subsistence use. Like other indigenous livestock species, the production potential of these indigenous pig breeds is low. These breeds should not, however, be wholly criticised because they have important characteristics that contribute to their adaptability, which are tolerance to heat and dietary changes. There is clear evidence to indicate that some indigenous West African pig breeds are tolerant to porcine trypanosomosis (Onah 1991). Porcine trypanosomosis has been shown to affect the productivity of exotic pig in Nigeria (Omeke 1989a). While some Chinese breeds are considered prolific (Bidanel 1990), this has not been confirmed in any of the SSA indigenous pig breeds. In some production systems, indigenous genotypes or their crosses with some exotic breeds may in fact be the most suitable. With considerable variability among indigenous pigs and their crosses with exotic breeds, their potential is highly subjective to selection procedures to attain higher levels of performance (Adebambo 1995). In most of SSA, the exotic breeds tend to dominate intensive piggeries because of their genetic superiority over the indigenous breeds although the adverse environmental factors may limit their performance. Therefore, there is increasing indiscriminate crossbreeding between the indigenous and the exotic breeds and even between some of the exotic breeds (Omeke 1989a). The performance of the crosses is determined by the additive genetic values of the two breeds and the amount of heterosis. It should, however, be noted that efforts to improve production at the village level, where the indigenous breeds might continue to be important, are very rare. Since studies comparing the performance of the SSA's indigenous pigs breeds, the exotic and their crosses are few, attempts will be made to review few studies from other tropical areas experiencing climatic conditions similar to those of SSA. However, because of the possible genotype × environment interactions, results from these areas cannot be extrapolated to the African conditions but give an insight on what to be expected. Preweaning traits Reproductive performance of pigs is influenced by complex interplay of environmental factors and physiological processes. In the tropics, climate is the most important environmental factor influencing reproductive performance. Steinbach (1976) discussed the effects of a tropical climate on the fertility of the boar and sow. Reproductive performance is reflected in litter size and weight at birth and at weaning. Several studies on the performance of the indigenous pig breeds, pure-bred exotic breeds and various crosses among them have been made under tropical conditions. In Sri Lanka, Goonewardene et al. (1984) compared preweaning traits of the indigenous pigs, pure-bred Large White and indigenous × Large White crosses and found that the Large White was significantly better than the indigenous in litter sizes, litter weight at birth, birth weight, weaning weight and average daily gain. However, there were no significant differences in weaning weight and average daily gain between the indigenous breed and the Large White-sired crossbred (Table 1). The litter size reported in that study was slightly higher than that reported by Adebambo (1983) (cited in Goonewardene et al. 1984) for the Nigerian indigenous pig. Table 1. Breed cross means for preweaning traits. Breed cross1 Litter size Litter weight (kg) Birth weight (kg) Weaning weight (kg) Average preweaning daily gain (kg/day) I × I 6.38 3.03 0.48 4.50 0.074 I × LW 8.20 8.26 0.97 7.73 0.108 LW × I 9.50 5.47 0.56 4.69 0.074 LW × LW 10.60 14.02 1.30 7.70 0.118 1. I = indigenous pig breed, LW = Large White breed. Breed of sire shown before breed of dam. Source: Goonewardene et al. 1984. In India, the pure-bred Large White Yorkshire breed was superior to it's crossbred with an indigenous Indian pig breed for reproductive performance (Mohanty and Nayak 1986). In southern Nigeria Steinbach (1971) reported significant differences between the Large White and Landrace breeds for birth weights, litter weight at birth and in percentage of stillborn piglets. The Landrace breed had the heavier piglets, heavier litters but the higher percentage of stillborn piglets than the Large White. In a tsetse infected area of Nigeria, Omeke (1989a) showed that the crossbreds between the Large White and Landrace breeds had the bigger litter size and the lower preweaning mortality rates than purebreds and recommended crossbreeding of these breeds in such areas to improve productivity. The performance of exotic breeds is better during the rainy season than during the dry season (Omeke 1989a). There is a significantly higher farrowing rate when exotic sows are mated during the cooler months of the year than during the hotter months (Dan and Summers 1996). Postweaning traits Crossbreeding of the indigenous pig with high productive exotic breeds usually results in heavier weaners and faster growth of fattening stock when compared with their indigenous parents. Usually indigenous pigs have carcasses that have more fat than the exotics. If consumer demand for pork from these pigs is to increase then leaner indigenous pigs will have to be produced. Ilori et al. (1976) compared the performance of indigenous and exotic pigs on restricted feed intake and found that the exotic pigs had leaner carcasses, faster growth rates but lower feed efficiency than the indigenous pigs. In western Samoa, Udo (1982) simulated three different systems of management, namely village, semi-commercial and commercial levels and compared the postweaning performance of the Large White breed, indigenous Samoan pig and crosses between them (Table 2). Under the simulated village management level, crossbreds grew significantly faster and had significantly higher dressing percentage than Large White and indigenous pigs. In contrast to the study of Ilori et al. (1976), there were no significant differences in backfat thickness but carcasses of local pigs were significantly shorter than those of Large White. Based on these results (Table 2), we concluded that there is no need to introduce pure-bred Large White pigs into villages except as breeding boars for crossbreeding purposes. Table 2. Performance data of Large White (LW), LW × Indigenous Samoan crossbred and indigenous Samoan pig (I) under different management regimens. Village Semi-commercial Commercial LW LW × I I LW LW × I I LW LW × I I Initial weight (kg) 10.45 11.59 10.45 11.70 10.71 13.25 10.68 11.36 11.45 End weight (kg) 34.00 44.35 32.04 82.04 80.08 80.80 79.73 81.41 70.41 Fattening period (days) 168 168 168 153 153 153 103 112 168 Daily gain (g/day) 140 195 129 461 453 442 670 625 351 Dressing percentage 63 70 63 71 70 68 68 68 69 Carcass length (cm) 61 70 49 74 72 68 76 75 64 Source: Udo 1982. For the breeding pigs, one of the most important postweaning trait is the age at puberty. It has been shown that the indigenous tropical pig breeds attain puberty earlier than their exotic counterparts. In Nigeria, Chiboka (1981) reported an earlier age at puberty in the indigenous pig breed when compared to the Large White, Duroc or Ukranian White. In contrast to the situation in exotic breeds Young and King (1981) reported the litter size, viability of offspring or normalcy of the parturition process (as measure by frequency of stillborn and milk ejection) were not affected by age at first service in that study. In Sri Lanka, Goonewardene et al. (1984) found that the indigenous gilts were sexually mature at an early age of 4-5 months in spite of the low nutrition and could therefore be bred for the first time when they are four to five months of age. Early mating is not necessarily harmful to the indigenous pig, and that sows bearing normal litters after early mating may be utilised for increasing maturity within these breeds and the exotic through crossbreeding. Post-farrowing traits The total number of piglets weaned by each sow per year determines the production cost and profit margin in a swine enterprise. The number of piglets weaned is in turn determined by the regularity of the successive reproductive cycles of the sow. The most important post-farrowing traits include, the interval from weaning to first oestrus, conception rate, gestation length and farrowing interval. Under conditions in SSA, some of these traits have been estimated in exotic breeds (Omeke 1989a; Omeke 1989b; Omeke 1990) and in indigenous breeds (Chiboka 1981; Somade 1985; Somade and Makinde 1985). Studies to compare exotic breeds, indigenous breeds and crosses between them for these traits in SSA are few. Genotype × environment interactions Genotype × environment (G × E) interactions simply mean that the effect of the environment on different breeds or genotypes is not the same; or in other words, there is a change in ranking of breeds or genotypes when taken from one environment to another environment. Environment might mean a climatic condition, system of management or feeding level. Therefore, there is no universally `best' breed or genotype. In this sense, genotypes selected in developed countries may not be suitable for extensive and semi-intensive production systems in the developing countries (Pathiraja 1986). As shown by Udo (1982) for daily gain (Table 2) in West Samoa, there was a change in ranking of the Large White breed and its cross with the indigenous Samoan pig under village and commercial management. This is clearly represented in Figure 12. The levels of heterosis for this trait were higher under village (45%) than under commercial management (22%). Cunningham (1981) suggested that when there is a substantial difference between the F1 and the local strain, there would be G × E interaction when heterosis heavily influenced production in a poor environment and breed additive effects and small heterotic effects largely determined production in a good environment. Source: Udo 1982. Figure 12. An illustration of genotype × environment (management system) interaction in pigs. Poultry Poultry is the most numerous species of farm animals in Africa with more than 80% of it kept in rural areas (Guèye 1998). Worldwide, poultry out-levels the human population by almost 30% (Horst 1990). Several species namely chickens, guinea fowls, ducks, geese, turkeys, pigeons and quail represent poultry. The ostrich has also been included as a poultry species. The small poultry species are generally kept in conjunction with ruminant animals and equines especially in the agricultural subsystems and in the pastoral system. Despite the substantial contribution of all small poultry species to the SSA's supply of protein, chickens and guinea fowls have received more research attention in recent years than the others. There is also some attention on ostrich under extensive systems especially in the southern part of Africa. However, its significance to the supply of protein to the rest of SSA is yet to be realised. Chickens Of all the poultry species, the domestic fowl (Gallus gallus domesticus) is widely represented in all parts of SSA where different strains or breeds have been known to exist. The indigenous fowls have been variously referred to us the African chickens, bush chickens or runner chickens. However, distinct local varieties have been reported in Egypt, Cameroon, Burkina Faso, Morocco and Sudan (Guèye 1998). Indigenous chickens tend to be robust and are well adapted to harsh environmental conditions such as hot or cold weather, rain and periodic feed shortages. These birds have many advantages such as good egg and meat flavour, hard egg shells, high dressing percentages, and especially low cost with little special care required for production. However, they have some disadvantages arising from the fact that they suffer high rearing mortality (Trail 1962), have slow growth rates and are poor egg layers and attain sexual maturity late. Such birds can be improved genetically through selective breeding or by crossbreeding with exotic stocks depending on the management conditions (Omeje 1986; Mukherjee 1990; Katule 1991; Katule 1992). This section briefly reviews the performance of indigenous chickens and their crosses with exotic breeds under SSA conditions. Growth traits Fast growing chickens have an advantage in that they reach the market weight early hence reducing production costs. It has been shown that crossbreeding the indigenous chickens with the exotic breeds improves the growth traits, which include traits like live weights, daily gains and feed intake (Omeje and Nwosu 1988; Asiedu and Weever 1993). While this is the case, it has also been reported that the indigenous chickens possess the potential to grow well at the early stage of life (Nwosu et al. 1984). Under a good nutritional environment, Katule and Mgheni (1990) reported similar growth rates of indigenous and exotic breeds to 12 weeks of age. In that study, the merits of the indigenous chickens were supported by the fact that the backcross to the indigenous strain grew faster than the exotic backcross. Ihemelandu and Nwosu (1986) also found that the indigenous chicken showed early faster growth rate than the exotic breeds and demonstrated that this early faster growth was also true for other organs (e.g. liver, heart and kidney). This indicates that the growth patterns of some indigenous chickens especially in Nigeria favour that of broilers. Various ecotypes of indigenous chickens produce meat that is in composition, consumer appreciation and acceptability similar to meat from exotic breeds of same age and management (Obanu et al. 1984). Improvement in body size and growth of indigenous chickens is important from economic considerations bordering on the need to increase egg size and to improve the post-lay value of the chickens (Ibe 1995). With well-designed selection programmes, this can easily be achieved in the indigenous chickens because of the appreciable additive genetic variance observed in these breeds (Ebangi and Ibe 1994; Olori 1994). Egg production traits Egg number and weight are major traits of economic interest in commercial egg production. Egg size determines to a large extent the price received in any market. Asuquo et al. (1992) reported that the internal egg quality traits are also important to the consumers especially the bakery industry. In that study, the exotic Isa-Brown breed was superior in all egg quality traits except per cent of yolk to the indigenous chickens. Soltan (1993) compared the egg quality traits of the selected Sinai fowls with those of the Fayoumi and Baladi fowls, which are the standard Egyptian indigenous breeds and found that the differences in these genotypes for these traits were not significant. Essien and Ekanem (1990) reported the effects of the age of layers and the period of oviposition on egg quality produced under humid tropical conditions. Because of the positive genetic correlation between body size and egg size, crossbreeding of the small indigenous chickens with an exotic breed also tends to improve the egg size of the crossbred progeny. In Nigeria, Omeje and Nwosu (1988) compared egg production traits of the indigenous chickens, the Gold-Link exotic breed and crosses between them. Significant differences were observed in the egg weights of the different genotypes. The Gold-Link exotic breed and the Gold-Link-sired backcross laid heavier eggs compared to the first cross, F2 and backcross populations that were sired by the indigenous chickens (Figure 13). The indigenous chickens had the smallest egg size. For total egg mass, a similar trend was observed. In Tanzania, Katule and Mgheni (1990) found that indigenous chickens produced more eggs in early life but were less persistent and laid smaller eggs. Surprisingly, in that study, the F2 generation laid more eggs than any other genotype. This is contrary to the deterioration in performance from F1 to F2 in crosses between tropical and temperate cattle for milk production (Syrstad 1989; Rege 1998). It should be noted that there exists variation in egg production traits within the indigenous chickens in some parts of Africa. In Egypt, for example, comparison of the three indigenous breeds (Sinai fowls, Fayoumi and Baladi) showed that the Sinai fowls produced fewer and smaller eggs than the other two (Soltan and Ahmed 1990). The indigenous Fayoumi and Baladi breeds have been subjected to several generations of selection for these traits and hence their superior performance. Source: Omeje and Nwosu 1988. Figure 13. Some egg production traits for four different genotypes relative to the indigenous chicken in Nigeria. Fertility traits Fertility traits can be classified into those that are associated with the egg, e.g. fertility and hatchability of the egg and those directly associated with the chickens, e.g. age at sexual maturity, which is defined as the age from hatch to the day of first egg. Before the introduction of the exotic chicken breeds for crossbreeding with the indigenous chickens, Trail (1962) carried out an experiment in Uganda to compare the fertility and hatchability of eggs from indigenous and exotic chickens and crosses between them. The fertility and hatchability rates of indigenous eggs were higher than that from the Rhode Island Red. When indigenous hens were crossed with cockerels of the Rhode Island Red, Light Sussex, White Leghorn and Black Australorp breeds, there was little difference between the fertility and hatchability levels of the four breeds of males used. In a study that was necessitated by the need to have information on the level of heterosis in the F1, and that of the residual heterotic effects in the F2, backcross and subsequent generations before starting a meaningful trait-group selection on crossbred populations, Omeje and Nwosu (1988) reported that the indigenous chickens and the indigenous-sired backcross laid the first egg about 10 days earlier than the pure-bred Gold-Link breed and its backcross. The F1 and F2 generations were intermediate. Katule (1992) reported small and non-significant differences in age at sexual maturity between the indigenous and the exotic strain of chickens in Tanzania. In that study the F1 generation matured earlier than any of the parental indicating existence of heterosis for this trait. Normally breeds that attain sexual maturity early end up laying lighter eggs than late maturers (Oni et al. 1991). Genotype × environment interactions Because of the method of rearing chickens in SSA, it would be expected that most investigation of G × E interactions would involve differences in management levels or systems. There are three poultry management systems- intensive, semi-intensive and extensive/scavenging. The intensive system is normally based on specialised breeds and is found mainly in urban areas. The extensive/scavenging is based on indigenous chickens and is mostly found in the villages (Kitalyi 1999). Comparison of breeds or strains of different ability to utilise feed under conditions where there is a scarcity of high quality feed (such as in most villages in SSA) would therefore be expected to favour the breed that is better able to utilise low quality feed. In an experiment to explore the possibility for developing strains of chickens which would perform reasonably well under low input production conditions (a characteristic of most of SSA), Katule (1992) presented evidence of existence of the influence of G × E interaction on both the body size and egg production traits. In that study, the performance of two exotic breeds of chickens (an egg type and a meat type), indigenous breed and crosses between them were compared in two experimental phases. Management (feeding and floor space allowance) was better in the first phase than in the second phase. In the first phase of that study, both the exotic breeds performed well in traits for which they had been developed while in the second phase, the meat and egg breeds performed less satisfactorily than their crosses and backcrosses with respect to body size and egg production traits, respectively. This indicates that genotypes, which have been developed for high performance in good environments, are expected to be more sensitive to environmental changes than those that have not been developed for any specific environment. Guinea fowls In some West African countries, guinea fowls are second only to chicken as a source of poultry meat and eggs (Ayeni 1983). In West Africa, there are two subspecies of guinea fowls. The grey breasted, helmeted guinea fowl (Numida meleagris galeata) is the most common and there are four varieties of this that are domesticated namely Lavender (Ash), Black, Pearl and White (Ayorinde 1991). Ayorinde et al. (1988); Ayorinde (1989a); Ayorinde (1989b); Nwagu et al. (1997) described the external and performance characteristics of these four varieties. The other type is the crested guinea fowl (Guttera edouardi edouardi) and is restricted in distribution to forest and derived savannah-forest edges (Ayeni 1979, cited in Ayorinde 1991). The grey breasted, helmeted guinea fowl is widely raised by peasant farmers extensively. We considered it uneconomical for its low fertility, slow growth and seasonal egg production. However, in France, the uses of light control programmes and selection have resulted in making guinea fowl an all year-round breeder (Nwagu and Alawa 1995). The local species of the guinea fowls is normally characterised with lower performance than the exotic (Ayorinde et al. 1988). Therefore, there has been introduction of exotic species of guinea fowl (Numida meleagris galeata), especially from France, for crossbreeding to improve the overall performance of the indigenous species. Other traits targeted for improvement are egg production and fertility traits. Nwagu et al. (1997) reported that the crossbred guinea fowls were considerably fertile and their eggs more viable than those of any of the local breeds. There was also variation within indigenous species for hatching characteristics. The Pearl variety produced eggs with the highest percentage of fertility and hatchability. We concluded that this variety could be a candidate for commercial hatchery production of a day old kites. There is therefore the need to properly explore the potential of guinea fowls as sources of protein for the ever-increasing population of SSA. Nwagu and Alawa (1995) listed a series of interventions that might help to realise this. Unique traits and properties of indigenous breeds Unique traits and properties of indigenous breeds Disease resistance or tolerance Trypanosomosis Gastro-intestinal parasites Ectoparasites Bursal disease Heat tolerance Cattle, sheep and goats Poultry Camel Adaptation and nutrition Other properties Salt tolerance Scavenging ability Reproductive performance Unique traits and properties of indigenous breeds Different hereditary characteristics of breeds and even types within a breed have resulted in difference in reactions to environmental stimuli. These reactions are intimately associated with anatomical-physiological characteristics, which have developed as the result of natural selection. Indigenous breeds, as the name suggests, have been identified with a particular area or people from time immemorial and are thus considered to be adapted to some of the environmental stresses. The degree of successful adaptation of such animals is accurately reflected in their ability to grow, to reproduce regularly and in their production. Therefore, we can conclude that adaptability and animal's efficiency of production are closely correlated. SSA is the home of roughly 120 indigenous cattle (Simon 1999), 61 sheep, 42 goat (Lebbie and Ramsay 1999) and 13 pig (Hammond and Leitch 1998) breeds.1 There is very little information on the diversity in indigenous African poultry populations, both at phenotypic and genetic levels. Most small-scale farmers and pastoralists owned indigenous breeds for whom they are a source of improved nutrition, income and a secure form of investment. The production systems under which they are produced are normally characterised by low levels of animal husbandry practises coupled with the normal tropical stresses of disease challenges, heat stress and poor nutrition. This demands that an animal must have some degree of resistance or tolerance to these stresses to maintain some degree of production. This section reviews within species documented examples of unique attributes and properties of indigenous breeds that are considered to be the most economically important in SSA. Definitions given were deemed necessary. Disease resistance or tolerance Diseases normally have an impact on welfare and production of a production system. The conventional control measures, such as vaccination and chemotherapy have either been ineffective, unsustainable or uneconomic. The genetic approaches to disease control have been suggested. The genetic basis of disease resistance or tolerance is evident from observations on breed differences and from results of selective breeding experiments and from laboratory studies (Stear and Wakelin 1998). It is important to differentiate between disease resistance and tolerance. Resistance can be defined as the initiation and maintenance of responses provoked in the host to suppress the establishment of parasites and/or eliminate parasite load while tolerance is the ability of the host to survive and produce in the face of parasite challenge (Baker 1998). Trypanosomosis In 1975, the Food and Agriculture Organization of the United Nations (FAO) launched, as a project of high priority, a long-term programme for the control of African animal trypanosomosis (Hoste 1987). This disease is particularly important in Africa and is one of the major constraints on animal production in areas that have the greatest potential for significant increases in domestic livestock population and livestock productivity (D'Ieteren et al. 1998). Cattle Trypanotolerance in cattle particularly in the N'Dama (a West African Longhorn) is well documented (Mortelmans and Kageruka 1976; FAO 1980; Starkey 1984). Other trypanotolerant breeds in SSA include the Baoule (a West African Shorthorn) (FAO 1980) and the Dahomey breed that is found in Zaire (Mortelmans and Kageruka 1976). There are now N'Dama herds in nearly all West and central African countries where they are being used as purebreds or as crossbred with other breeds (FAO 1980). Currently, N'Dama cattle constitute only 5% of the total cattle population in SSA, with an approximate population doubling time of 16 years at current rates of increase (Teale 1993). The N'Dama and Baoule breeds are traditionally small animals but their productivity can match that of trypanosusceptible zebu cattle (B. indicus) in areas where tsetse fly risk is low (ILCA 1979). In high-risk areas most B. indicus breeds require regular treatment however significant differences in resistance to trypanosomosis occur also among various B. indicus breeds (Njogu et al. 1985). Maintenance of exotic breeds even in areas of low tsetse fly risk would require intensive trypanocidal drug therapy and veterinary care. Sheep and goats Studies on the genetic resistance of small ruminants to trypanosomosis are scarce. In West Africa the Small West African Sheep called Djallonké or Fouta Djallon sheep and the West African dwarf goats are the only recognised breeds that are trypanotolerant (FAO 1980). In Kenya, it has been suggested that the Red Maasai and Blackhead Persian sheep and the Galla and Small East African goats have more resistance than imported breeds (Merino sheep and Saanen goats) (Gray et al. 1995). Katunguka-Rwakishaya et al. (1997) reported differences in susceptibility to Trypanosoma congolense infection among three indigenous goat breeds namely, the Kigezi, Mubende and Small East African goat. The Kigezi goat was the most susceptible while the Small East African goat was the least susceptible. This indicates that there is also variation in resistance to trypanosomosis between the indigenous goats breeds. More studies are required to investigate within and between breed genetic variation for trypanotolerance in sheep and goats. Pigs While a lot of attention has been put on trypanosomosis in cattle, little information is available for pigs. Pigs, like other domestic livestock, are infected by several trypanosomes and are therefore hosts for these causative agents that can infect the other domestic species. In West Africa, the long-snouted West African pig breed is less susceptible to porcine trypanosomosis than the exotic (Onah 1991). Most studies are required on this indigenous pig breed to investigate its trypanotolerance status and its possible use both as a purebred and in crossbreeding programmes in the tsetse infested areas of SSA. Camels It is only recently that camels have become the subject of more intensive and systematic interest in connection with increasing the development potential of drylands in Africa and Asia (Baumann and Zessin 1992). The camel is generally considered to be disease-tolerant (Morton 1984). However information on which camel breed is resistant or tolerant to which disease condition is lacking. There exist a large variety of distinct camel breeds that have developed along tribal/ethnic lines (Köhler-Rollefson 1993). For example, classification of camels in Kenya is based on the geographical distribution of the owners and ethnic group. As a result the terms Somali, Rendille, Gabra or Turkana camel, are widely recognised (Kegode 1990). Using this classification, trypanosomosis was perceived as a major disease affecting young stock and adult camels (Kaufmann 1998). However, trypanosomosis was ranked highest in Somali and Rendille breeds. This outcome is not conclusive enough to indicate that the Gabra breed is more trypanotolerant because it is not yet clear whether the camel populations named after the camel keeping ethnic groups differ both in phenotype and genotype. A research programme is underway at Hohenheim University, Germany, in collaboration with ILRI, the Kenya Agricultural Research Institute (KARI) and the Food and Agricultural Research Management (FARM)-Africa to characterise camel genetic resources in dry lands in Africa and Asia. This programme might prove whether these breeds are genetically homogenous or not. Gastro-intestinal parasites Most of the research work on resistance to endoparasites has concentrated on the nematodes and particularly the Trichostronyles (Haemonchus, Ostertagia, Trichostrongylus, Nematodirus and Cooperia) species. In most of the tropics, helminths constitute one of the most important constraints to ruminant production (Fabiyi 1987). The problem is serious, as chemotherapy is becoming less effective due to multiple resistances of parasites to chemicals and prospects for vaccination are not encouraging. Therefore, there has been research emphasis on genetic control of the disease. Cattle Surprisingly, evidence for host genetic resistance to gastro-intestinal nematodes even of African breeds (Frisch and O'Neill 1998) is from Australia. It has generally been observed that B. indicus cattle are more resistant to gastro-intestinal nematode parasites than B. taurus (Turner and Short 1972; Frisch and Vercoe 1984). Comparing beef cattle breeds of African, European and Indian origins, Frisch and O'Neill (1998) showed that the Brahman (Indian Zebu) was more resistant to gastro-intestinal nematodes than the Boran (African Zebu). Among the two African breeds represented in that study, i.e. the Boran and the Tuli, the Boran had higher resistance to worms than the Tuli. The Boran is predominantly B. indicus and originated from Ethiopia while the Tuli is the southern African Sanga and is sometimes referred to as the African B. taurus. Sheep and goats There is more evidence for both between and within breed genetic variation for resistance to gastro-intestinal nematode parasites in sheep and goats than in cattle. Breeding for resistance has attracted considerable research and development attention all over the world. Outside Africa, several indigenous breeds have been identified in the Caribbean (St Croix and Barbados Blackbelly) and North America (Florida Native and Navajo) that are more resistant or tolerant to gastro-intestinal nematodes than Rambouillets or Merinos (Gray et al. 1995), however within breed genetic variation for resistance to gastro-intestinal have been reported in the Merino sheep in Australia (Raadsma et al. 1998). In SSA, the indigenous Red Maasai sheep of East Africa are resistant to naturally acquired and artificial infections with gastro- intestinal nematode parasites and particularly to Haemonchus contortus (Baker et al. 1999). The Red Maasai sheep has the ability to maintain good levels of production when under severe and persistent endoparasite challenge. Based on these results, ILRI has embarked on a programme to identify genetic markers linked to quantitative trait loci that control resistance to infections to gastro-intestinal parasites, H. contortus in particular (Baker et al. 1999). While sheep and goats are considered to share common susceptibility to many diseases, it has been observed that there were significant differences in disease resistance between the two small ruminants. In contrast to sheep, there is less information on resistance to gastro- intestinal nematodes in goats. In East Africa, the Small East African goat has been found to be more resistant than the exotic and the indigenous Galla goat (Shavulimo et al. 1988; Waruiru et al. 1994; Baker 1998). The Galla breed has its origin in the semi-arid areas of East Africa, where gastro-intestinal nematodes are not a major problem. It is therefore probable that the Galla have had little exposure to nematodes and have therefore had no natural selection for resistance. Ectoparasites The most important ectoparasite that imposes severe economic constraints on livestock industries in SSA is tick. Ticks are responsible for losses caused through blood loss, damage to hides, skins and udder, tick worry and the injection of toxins. They also transmit devastating and often fatal diseases such as theileriosis, babesiosis, anaplasmosis and cowdriosis. This brings about mortalities, reduced productivity and direct costs associated with preventive and curative measures (Young et al. 1988). In SSA, there is widespread and indiscriminate application of acaricide to control ticks that has resulted in emergence of acaricide-resistant ticks (Nolan 1990). Therefore utilisation of breeds of livestock that are considered to be resistant to ticks is important to reduce the reliance on chemicals that have negative impacts in animal products and on pastures. Resistance is manifested by reduction in attachment and engorgement of ticks (Fivaz and De Waal 1993). Cattle Mackinnon et al. (1991); Davis (1993); Frisch and O'Neill (1998) (in Australia) and Lemos et al. (1985); Garcia et al. (1989); Gomes et al. (1989) (in South America) established that pure- bred B. indicus cattle and their crosses with B. taurus breeds are more resistant to ticks and tick-borne diseases than pure-bredB. taurus breeds and indicated that there exists both between and within breed genetic variation for resistance to the one-host tick Boophilus microplus. While the indigenous cattle breeds are resistant to ticks, it has been established in the zebu cattle of southern Uganda that its resistance to some tick species, Rhipicephalus appendiculatus in particular, is not strong (Kaiser et al. 1982). An evaluation of the performance of indigenous Nguni, Bonsmara and Hereford in South Africa indicated that the Nguni was more resistant, the Bonsmara being intermediate (Fivaz and De Waal 1993). The Nguni breed belongs to the indigenous Sanga group, which evolved along the east coast of Africa while the Bonsmara is a composite breed developed from crosses between the Sanga Afrikander and exotic Shorthorn and Hereford (Fivaz and De Waal 1993). In Ethiopia, the Boran, Barka and Horro indigenous breeds are more resistant to ticks than their crosses with the Jersey, Friesian and Simmental breeds (Yehulashet et al. 1995). Kiltz and Humke (1986) reported partial tolerance to theileriosis (East Coast fever) in the Ankole cattle in Burundi and Paling et al. (1991) reported the same result in the Ankole cattle in Rwanda. This tolerance is likely to be a result of centuries of natural selection within the Ankole cattle population, surviving in the East Coast fever endemic areas of central Africa. Sheep and goats Although small ruminants are often an important part of the livestock enterprise in tick-endemic areas, there is little information in this class of livestock on resistance to ticks and tick-borne disease. There is however some indication that the Red Maasai sheep in East Africa is resistant to ticks (Ogore 1996). Ogore et al. (1999) reported a simple method of assessing tick burden in sheep. Bursal disease This is an infectious viral disease of poultry caused by the Birnavirus and is a disease of considerable economic importance, particularly affecting birds between three and six weeks of age (Bumstead et al. 1993). However resistance to this disease has been reported in the local chicken breeds in Sudan (Salman et al. 1983) and Nigeria (Okoye et al. 1999). Bumstead et al. (1993) reported within lines or breeds variation to susceptibility to this disease. Heat tolerance SSA is characterised with extreme heat and high humidity in some coastal and inland valleys and basins. Animals under heat stress dissipate heat by increasing their respiration rate and sweating and measurements of these parameters plus pulse rate and rectal temperature have been used to assess the tolerance of ruminants to adverse climatic conditions (Lemerle and Goddard 1986). Heat tolerance is one of the adaptations that contribute to the performance of tropically derived breeds and their crosses in warm environments (Turner 1984). Cattle, sheep and goats Bonsma (1949) recognised differences in adaptability to the effect of heat between tropical and temperate breeds in SSA in the first half of the twentieth century. This has been confirmed in Australia (Turner 1982; Turner 1984), Papua New Guinea (Lemerle and Goddard 1986) and India (Singh and Bhattacharyya 1990). In Nigeria, Amakiri and Funsho (1979) studied the rectal temperature and respiratory rates of German Brown, Friesian, German Brown/Friesian crosses, N'Dama, White Fulani and German Brown/N'Dama crosses. The Friesian and German Brown were found less heat tolerant than the indigenous N'Dama and White Fulani. Use of the sweating rate as an indicator of heat tolerance indicated that the White Fulani and N'Dama breeds were similar for heat tolerance (Amakiri and Onwuka 1980). The superior heat tolerance of the indigenous breeds has been attributed to their coat type and colour, skin thickness and pigmentation, high sweating capacity (which is a measure of the density of sweat glands in the skin), low body heat production as a consequence of their low productivity, body conformation and some physiological aspects (Bonsma 1949; Allen 1962; Allen et al. 1963; Amakiri 1974; Amakiri and Mordi 1975; Turner 1975; Herz and Steinhauf 1978; Amakiri and Onwuka 1980). Most indigenous cattle, sheep and goats breeds in SSA have small bodies, low levels of milk production and slow growth rates and have fat localised in specific areas of the body. For example, the storage of fat in the hump of zebu cattle and camels and in the tails or rumps of many indigenous sheep (e.g. the Red Maasai sheep) leads to more heat loss than an equal amount of fat stored under the skin. The indigenous sheep and goat breeds are smooth- coated, which is in itself an adaptive mechanism because they lose more moisture from the skin as a result of evaporation (Bonsma 1949). Poultry High ambient temperature is regarded as the most important inhibiting factor for poultry production in hot climates (Horst 1990). Under these conditions, chickens cannot dissipate heat produced following meals rapidly enough, leading to reduced feed intake and lower weight gain or egg production (Cahaner and Leenstra 1992). This has lead to systematic comparisons of naked neck and normally feathered chickens at various ambient temperatures (Horst 1988; Mathur and Horst 1990; Cahaner et al. 1993; Ibe 1993). Several breeds exhibiting the naked neck trait have been described, including the `Cou Nu du Forez' from France, the `Malay Game' from Malaysia, the `Shingangadi' from Zaire, the `Trandylvanian naked neck' from Hungary and Romania and the `Peel-Neck' from Belize and Guatemala (Merat 1986; Mallia 1999). Some major genes have been found relevant to the hot tropical production environment. Among these major genes are the feather distribution gene, naked neck (Na) and the feather structure gene, frizzle (F) (Horst 1988). These genes cause a reduction in tropical heat stress by improving the breed's ability for convection, resulting in improved feed conversion and better performance. The Na and F gene confer superiority in some production characters in the tropics (Horst 1988). Horst (1988) and Mathur and Horst (1990) showed that individuals with F and Na genes both singly and in combination were superior to those individuals with normal feathering for egg number, egg mass/weight and forty-week body weight in tropical environments. The Na and F individuals have been shown to mature earlier than individual with normal feathering (Ibe 1992). Cahaner et al. (1993) showed that the heterozygous Na broilers were superior to normal broilers at normal temperatures and were even superior at higher temperatures. In Nigeria, quantitative genetic principles are being employed to explore use of major genes and that of polygenes in bringing about improvement in production characters in synthetic populations of indigenous and exotic strains (Ibe 1990; Ibe 1992). Camel As already alluded to, camels are the most drought-tolerant and efficient converters of feed and water to meat and milk than any indigenous livestock species (Morton 1984). Their unique anatomical and physiological characteristics enable them to exist, reproduce, and produce meat and milk during periods of drought, poor grazing and low management. Camels have begun to receive attention in recent years as far as describing their unique characteristics, physiology and estimating their production capability as a livestock species is concerned (Dolan et al. 1983; Schwartz et al. 1983; Morton 1984; Yagil 1985; Chowdhary 1986; Wilson 1986; Wilson 1988; Khanna et al. 1990; Schwartz 1992; Kaufmann 1998; Hülsebusch 1999). However studies to compare different camel breeds for measure of heat tolerance are lacking. Adaptation and nutrition Another important attribute for surviving and producing in most of the tropics is the ability to utilise lower quality feeds. While studies on the comparison of the indigenous livestock breeds to their exotic counterparts are lacking in SSA, it has been indicated that indigenous zebu (B. indicus) is better able to utilise lower quality feeds than the temperate breeds (Ashton 1962; Moore et al. 1975; Dunkel 1981; Hunter and Siebert 1985a). However, B. taurus cattle have higher intakes than B. indicus of good and medium quality forage diets (Hunter and Siebert 1985b). It has been suggested that the superiority of the B. indicus on low quality feeds could be due to their superior ability to recycle endogenous nitrogen in the rumen (Hunter and Siebert 1985b). Zebus also have higher true digestibility, more extensive ruminal digestion, more efficient protein synthesis and lower metabolic faecal nitrogen excretion than most temperate breeds (Ashton 1962; Moran and Vercoe 1972; Vercoe and Frisch 1972; Kennedy 1982; Hunter and Siebert 1985a; Hunter and Siebert 1985b). The tropical Bedouin goats can maintain a high level of high fibrous feeds compared with the temperate Saanen breed (Silanikove 1986a; Silanikove 1986b; Silanikove and Brosh 1989). This has been attributed to their ability to maintain higher microbial activity in the particulate matter and hence higher total ruminal fermentation rate, which are reflected in the extent and site of absorption of digested end-products (Silanikove et al. 1993). The Bedouin goats have a small body size and are found to thrive in extreme deserts where, with the exception of the camel, no other domestic ruminant lives (Silanikove 1986b). There are reports from Côte d'Ivoire of the indigenous poultry breeds having the ability to use high fibre feeds (Diambra 1990, cited in Sonaiya et al. 1999). Other properties Salt tolerance There might be some relationship between adaptation to native feeds and heat with tolerance to salts. Although not published, reports indicate that the Galla goat found in the arid and semi-arid areas of Kenya has the ability to consume plants that contain high amounts of salts without getting dehydrated. Normally goats have a lower water turnover than sheep and will respond to dehydration by excreting a smaller volume of urine with higher osmolalities (Aganga 1992). Generally in Kenya it has been observed that for grilling, consumers preferred meat from salt tolerant goats because of its characteristic salty taste. Scavenging ability This is an important attribute of the indigenous poultry breeds and is the ability to find feed from the surrounding environment. The land area available for scavenging and the distance a flock can travel to scavenge will depend on many factors such as flock size, feed availability, population density, agricultural activities and predators (Gunaratne 1999). Studies have shown that indigenous birds have better scavenging ability compared to others (e.g. exotic or crossbreds). In Kenya, for example, one of the main reasons for the failure of the National Cockerel Exchange Program was the progeny of the exotic cockerel tended to restrict their scavenging area close to the household where, with no supplementation, is an area with less feed. Therefore, the productivity of the crossbred progeny was low and the farmers refused to give up their indigenous cockerels in favour of the exotics. Roberts (1995) described a simple model for production systems of scavenging village chickens in which the biomass of the village flock is maximised at the capacity of the scavenging feed resource base. Reproductive performance A limited number of indigenous breeds exhibit exceptional reproductive ability. While there is less information on such breeds in SSA, in China for example, the Meishan breeds of pig are very prolific but are characterised by very poor production performance (Bidanel 1990). It has been reported that the Mossi dwarf goat (a local caprine breed reared by smallholders in central and eastern Burkina Faso, West Africa Sudano-Sahelian region) has excellent reproductive performances (Tamboura et al. 1997). Unpublished reports indicate that the twining rate in the Small East African goat is high. 1 . Note the difficulty to define breeds in developing countries as discussed in the section on Valuation of biological diversities. Final remarks Final remarks Valuation of farm animal genetic resources--prospects and possibilities Crossbreeding with exotic breeds as a threat to farm animal biodiversity Naive crossbreeding schemes Structured crossbreeding programmes Developments of synthetic breeds Decision making Final remarks Valuation of farm animal genetic resources--prospects and possibilities Economic valuation of indigenous breeds or populations in tropical countries, especially in SSA, has been done to a very limited extent, so that to date only few results are reported in the literature. There are numerous methodological approaches that can be used and different types of values to be determined, as outlined earlier in this study. For the purpose of decision making with respect to AnGR management, it is necessary that comparable values are determined for different breeds. If, say, breed A has a specific indirect use value in that it fulfils an ecological function, and this breed may be replaced through another breed B that has a higher production value, but presumably is not able to fulfil the ecological function to a comparable extent, the costs and benefits have to be balanced on the same scale. This will be difficult, if, e.g. the ecological value of breed A is determined for the country or region and the production value of breed B is expressed on a farm level. As mentioned earlier, there are serious doubts that an overall economic valuation of breeds will be possible with sufficient accuracy, especially, if indirect and potential future values are considered, the latter being often used as the main argument justifying conservation activities. A comparable, if not greater, lack of knowledge exists with respect to costs of conservation strategies. Breed conservation is not absolute, but is related to extinction probabilities. To give an example: at a given point of time, the probability that a certain breed will be extinct in, say, 10 years, may be 50%. Conservation activities aim at reducing these probabilities. However, it will hardly be possible to reduce it to 0% (i.e. the conservation of the breed is guaranteed), but the extinction probability may be reduced to, say, 30% or 20%. Different activities will result in different extinction probabilities, but will also have different costs. Rational conservation strategies require knowledge of costs per, say, 10% reduction of extinction probability. Since relevant approaches like the `safe minimum standard' concept are based more on quantifications of costs rather than benefits, determination of the costs and the efficiency of conservation strategies is assumed to be more important than valuation of the populations to be conserved. Crossbreeding with exotic breeds as a threat to farm animal biodiversity Crossbreeding appears to be an attractive option if productivity (in terms of output units) is to be increased in short-term. Under tropical conditions, productivity (like milk yield per day or lactation, daily gain, eggs per year etc.) is only one factor among many others determining the overall economic efficiency of livestock and certainly is of much lesser impact than it is in temperate regions. It has been shown in many studies that the price to be paid for increased productivity is reduced adaptation and hardiness, health and fertility problems, increased treatment costs, losses etc. Fair economic valuations of crossbreeding schemes are not available, since in many cases real costs are masked through free provision of exotic genetic material (sperm or live animals) and crossbred animals initially often are kept under station conditions financed by donor organisations. When judging crossbreeding schemes, one has to differentiate between three principal approaches: Naive crossbreeding schemes These schemes are characterised through a more or less unplanned import of exotic genetic material, which is used for one or few generations to `upgrade' local indigenous breeds. Further characteristics are that the exotic genotypes usually are not selected to fit well to the local populations they are merged with. Often, the costs of the import of these exotic genetics are taken over by donors, while the consequences have to be taken by the local farmers. A good example is the frequently practised insemination of indigenous cows with imported Holstein semen, which often is of a quality that cannot be sold in the temperate countries of origin. In the starting phase of such a programme, results are promising due to heterotic superiority of F1-animals. However, recombination loss leads to a breakdown of this superiority in subsequent generations, so that the productive advantage is reduced, while the problems with insufficient adaptation remain. Typically, such schemes lead to continuous indiscriminate and uncontrolled crossbreeding, i.e. after the termination of the project, crossbred animals remain in the country and are disseminated and used without any plan or control. Their different phenotype and (seemingly) higher productivity makes them attractive to the farmers, so that on the long run exotic genotypes replace a significant proportion of the indigenous genotypes. However, the resulting animals will have a lower adaptive capacity and increased disease susceptibility and management and feeding conditions are in many cases not sufficient to allow them to express their eventual productive superiority, so that an overall economic analysis in many cases would not favour such a scheme. On the other hand, this type of indiscriminate dissemination of exotic genes, resulting in a complex mixture of genotypes, is a severe threat to indigenous AnGR. Structured crossbreeding programmes It has been shown that complex crossbreeding schemes like continuous production of F1- animals, crisscrossing, two- or three-breed rotational crosses etc. have the capacity to produce animals with improved productivity and sufficient adaptive capacity to be profitable under the given environmental conditions. Such schemes imply a continuous maintenance of the required infrastructure and logistics, i.e. pure lines have to be maintained or exotic material has to be continuously imported (as semen, embryos or live animals). Establishing and maintaining such schemes requires considerable infrastructure and stable financial and marketing conditions. Therefore, this sort of sustainable crossbreeding schemes will in many cases be limited to special situations, like semi-intensive milk production in peri- urban regions. In this case, the threatening of the biodiversity of the local breed is limited. If, however, such schemes are used to produce `improved' crossbred products to be disseminated as breeding animals to smallholders to be mated to local females, the risk of indiscriminate use and therefore considerable adverse effects on biodiversity are to be expected. Developments of synthetic breeds Synthetic breeds development from a systematic cross of local and exotic breeds has been shown to be a promising strategy to combine desired properties from both sources. If the products are selected under typical environmental conditions, the optimal rate of productivity under the given constraints can be achieved in combination with sufficient robustness and hardiness. Typically, such composite breeds go through a genetic bottleneck since they are derived from a limited number of founder animals. Therefore, within breed diversity is limited and a relatively high level of inbreeding is programmed. Development of synthetics is a multi-generation project, which requires substantial input over a long time period. It is likely for that reason that most existing synthetics in the tropics have been developed in regions like tropical northern Australia, South Africa or Latin America, where both the required infrastructure and the funding are easily accessible than in most of the less developed countries of SSA. Once synthetics are released, they compete on the market with indigenous breeds and may replace them to a certain extent. It must however be realised that these animals have relatively high prices and only perform well under good management and feeding conditions, so the risk of a complete replacement of indigenous genotypes is limited. Crossbreeding certainly is one elementary technique to improve productivity and has its justification to improve the farm animal productivity under tropical conditions. Situations must be avoided, where crossbreeding does not achieve its goal (i.e. improvement of productivity under the given conditions) but still is executed with harmful effects on farm animal biodiversity. This is usually the case, if exotic crosses are made accessible to the farmers without sufficient training, control and monitoring. Already the fact that these animals look different, may be larger, or may perform well on station, makes them attractive, although they or their offspring may not be able to perform well or even to survive under the smallholder conditions. Biodiversity is not static. Breeds have always been newly developed and others have disappeared. It is a matter of the speed of such processes, and it is a matter of rationality, especially it must be avoided that well adapted genotypes are replaced by genotypes that are not competitive under the given conditions. However, it must also be accepted that adapted breeds are improved through well-planned crossing and upgrading. There is a certain trade-off between the goals of conservation and improvement, but many examples have shown that sustainable improvement of breeds is hardly possible without conservation of the most important characteristics of indigenous breeds. Decision making Decisions with respect to AnGR management and conservation have to take into account a variety of implications, e.g. development in the region, effects on the livelihood of stakeholders, socio-economic impacts etc. It was clearly beyond the scope of this study to analyse and discuss all the implications of AnGR management in the different areas, but it is obvious that economic considerations are only a part of the whole framework under which AnGR management has to be discussed. However, all decisions have an immanent economic component, in that present or future costs are involved and/or actual or future values are preserved or sacrificed. It is difficult to assess these economic aspects (in the narrow sense) with sufficient accuracy, so that rational decision making on the base of objective data and economic valuation results under the present conditions in SSA1 must be seen as a major challenge. The problems arising in this context may be illustrated with the following examples: There is no obvious and clear definition of the goals to be achieved through AnGR management activities. The scope of decision making is limited. It is difficult to imagine a negative decision with respect to the conservation of a breed, i.e. to decide that the extinction of a certain breed will be accepted. The real alternative rather is either to become active with respect to certain breeds, or to do nothing about these breeds, which may have the consequence of extinction or replacement for them. Most of the theoretical work on AnGR management assumes well-established breeds and breeding infrastructure. These conditions are not given for most farm animal species in SSA. It must be doubted that this will be changed in the foreseeable future. Scientists questioned the important components of the total economic value, e.g. the option value reflecting the `insurance' function allowing adaptation to future changes, can ever be quantified with sufficient accuracy. Therefore, total economic value comparisons will always be affected by a considerable amount of uncertainty. Apart from the approach suggested by Weitzman (1993), which is aimed at the problem of conservation of biodiversity between species, there is hardly any conceptual work on how to decide which animal groups (breeds, strains etc.) should be preserved. Taking these problems and methodological shortcomings into account, approaches gain attractiveness which are suited to decision making under high levels of uncertainty. One such approach may be to adopt the concept of `safe minimum standards', which mainly requires decision making with respect to finding and implementing the most cost-effective strategy to prevent total extinction of a certain breed or population. This is based on the assumption that a priori every population should be conserved, if possible, since it may have a future value which, however, is largely unknown by definition. In such a case, decisions are required with respect to, e.g. the exact unit to be conserved (definition of the breed or strain, exclusion of crosses etc.); the conservation scheme to be implemented (in situ or ex situ, number of animals, monitoring system etc.); the possibility of combination of this activity with other activities (i.e. improvement of the breed, health services etc.). Of course, there will be costs for the implementation of such schemes, and, given that financial resources are limited, it will in most cases be necessary to set up a priority list of activities and breeds to be conserved. This, again, is a decision-making process that should, in principle, be based on objective data and information. However a fast and inexpensive (but may be suboptimal) decision will in many cases be adequate and tolerable. It may, e.g. in many cases be sufficient to ask experts with good knowledge of the populations in a region to set up a ranked list of breeds with highest priority to be conserved. It also must be doubted, whether a scientifically `sound' description and valuation of the actual biodiversity is always required, which is both expensive and demanding with respect to time and manpower to be invested. Economic resources are limited, and there is a competition between funds to be used either for description or for conservation activities. A considerable proportion of biodiversity may already be lost at the time the respective populations are described and valuated accurately. 1 . In fact, the situation with respect to sufficient knowledge of population structures, breed values and conservation costs is not much better in more developed regions of the world. Conclusions Conclusions Farm animal genetic resources are subject to conservation efforts, because we assumed that they have a value. This value may be of very different nature (productive, ecological, aesthetic etc.) and may be realised at different times (past, present and future). Management of AnGR has to be based on a rational decision-making framework, part of which is the definition of conservation priorities and development of tools for the optimal allocation of resources. For this, it is essential to assess economic values of genetic resources. Environmental economies have suggested different types of values for species or breeds. Use values are related to actual productive and consumptive functions and non-consumptive aspects like aesthetic, cultural or ecological functions. Option values are use values to be realised in the future, e.g. the potential of a breed to satisfy market requirements that may arise in the future, caused, e.g. by climatic changes etc. Non-use values are related, e.g. to the contribution of exotic breeds to the understanding of biological mechanisms. Actual use values are most relevant for the role a breed plays and whether it becomes threatened to be extinct (e.g. if its productivity is clearly inferior to other breeds, which are comparably viable in the given environmental conditions otherwise). Option values are relevant for conservation priorities, since it is most sensible to maintain breeds that will have highest use values in the future. Different types of values can be quantified with different accuracy. It is easier to quantify values that are realised in the present than those values that may be realised in the future, and it is easier to assess values for traits or animals that are traded on markets compared to traits (e.g. ecological functions) for which no market exists. aluation methodologies can be subdivided into empirical approaches and system analysis approaches. Empirical approaches are based either on the analysis of real market data (paid prices) or survey based, trying to assess `willingness to pay' for animals, traits or functions. The latter approach, usually called `contingent valuation' allows assessing values for `hypothetical' traits or for functions, for which no market exists. System analysis approaches try to model a complex system (farm, household, sector, country, market etc.) by defining detailed relations and functions between all relevant elements. This allows to assess the effect of specific changes (e.g. the replacement of one breed through another breed) on the productivity of the studied system. Decision making can be made in a cost-benefit framework, trying to assess all the costs and benefits arising from certain decisions, and maximising the expected economic return. Since it is difficult to quantify all values (and costs) with sufficient accuracy and since especially option values underlie considerable immanent uncertainties, alternative concepts like the safe minimum standard concept have been suggested to avoid these problems. The safe minimum standard approach is based on the assumption that a loss of a breed should be avoided and tries to find a strategy that guarantees conservation with minimum costs. Genetic approaches try to set up least cost conservation schemes that limit the loss of genetic diversity to an acceptable rate. An important and necessary feature of such schemes is robustness, i.e. that they still should prevent a total loss even if the assumptions they are built on are not entirely met. Decision making and priority setting for conservation activities requires well defined objectives. In the AnGR context, these objectives are neither specified nor discussed with sufficient rigidity. There is no uniformly accepted measure of diversity, nor is there any formal concept how to value diversity and how to combine its value with, e.g. production and option values. In environmental economics, Weitzman (1992); Weitzman (1993) suggested a general concept of diversity, which integrates information on structural diversity (in form of a distance matrix), extinction probabilities and cost efficiencies of different conservation activities to an overall operational framework for decision making and priority setting. Adopting this framework for the farm animal situation would require both methodological developments (e.g. Weitzman's approach is designed for species rather than for breeds) and empirical derivation of a large variety of values and costs. However, this approach is the only available formal framework that appears suitable for the desired implementation. There are only very few studies available that have actually derived values of AnGR. ILRI has now initiated a major project to conduct a series of such studies (with focus on SSA) to evaluate the usefulness of different valuation techniques in the farm animal context, since most of these techniques have been developed and used for valuation of non-agricultural species or habitats. These activities will reveal mainly quantifications of actual use values on the basis of empirical studies. This may be useful for situations where, e.g. breeds are threatened due to market distortions and externalities (e.g. due to highly subsidised import of semen of exotic cattle breeds), but it will be of limited use for the assessment of the diversity value of a breed. Diversity is a feature of a set of breeds, not of a single breed (a single breed has a quantifiable contribution to the species' diversity, though). Therefore, decision making at the local level (e.g. on conservation activities for a breed on a national level) has to take into account implications related to the global level (e.g. conservation of the diversity within a species with breeds distributed over a whole continent or worldwide). To achieve this, different aspects have to be considered: A clear framework for priority setting and decision making has to be agreed upon, including the definition of objectives. Information has to be provided and shared to allow the operational implementation of such a framework. Rights and duties of all relevant stakeholders have to be defined, including intellectual property rights (IPR) and ownership issues in relation to animal genetic resources. Balancing mechanisms have to be implemented for situations, where costs and benefits arise asymmetrically, e.g. a national conservation scheme conserves with high cost an actually unproductive breed since this breed contributes highly to the global within species diversity. Education and training is essential especially on the level of decision makers to motivate them to co-operate and to take the `global' view rather than the `local' one. Weitzman (1992), Weitzman (1993) suggested the most promising (and, actually, also the only available) conceptual framework, which, however has to be adapted to the specific conditions given in the farm animal situation. To implement this concept, the following input data are required: Quantification of between and within breed genetic variability and pair-wise breed distances (for African cattle breeds mostly available from ILRI studies, for other species and other regions to be determined). Quantification of the relative risk of extinction of different breeds and development of functional relationship, e.g. to effective population size or inbreeding level. Derivation of costs related to efficiency (in terms of loss of genetic diversity) of different conservation activities under the conditions in SSA. Empirical assessment of values of different breeds and breed characteristics, especially those `unique' characteristics concerning adaptive capacity, disease tolerance or resistance etc., which form a major component of the option values of indigenous breeds. Valuation techniques are available and their usefulness in the farm animal context will be determined. Given the urgency of starting efficient conservation activities now,1 and given the limited resources in this area, research priorities have to be considered and discussed very pragmatically. In many cases, rough estimates that are available today will be more useful than exact scientific figures available in two years time. The main conclusions of this study can be summarised in the following theses: Economic valuation of indigenous breeds is an important part of the description of AnGR. Important components of these values, especially non-use and option values, are extremely difficult to measure and likely cannot be determined with sufficient accuracy. Many of the factors determining the value of indigenous populations are of a qualitative nature and are biologically not well understood, so that it is difficult to quantify their economic impact. The contribution of a breed to the biodiversity within its species (its biodiversity value) may be very different from its actual economic value. Therefore, decision making should not be exclusively based on analytical cost-benefit considerations. Alternative approaches are based on the assumption that as much AnGR as possible should be conserved and focus on safe conservation with minimum cost. Conservation activities should be designed to be robust, i.e. to prevent total loss of a breed even if the assumed conditions the design is based upon are not met. Rational decision making requires empirical knowledge of cost and efficiency of different conservation and management strategies under the conditions in SSA, which is not available to date. Priority should be given to conservation of the most endangered breeds with highest biodiversity value. However, in many cases immediate activity will be more important than accurate quantification of all values, genetic distances etc. Establishing IPR for genetic resources can be a means for generating markets, which may lead to incentive effects beneficial for AnGR management. AnGR management should be an integrated objective of all activities aiming at, or dealing with livestock-related structures in the region. Crossbreeding can be an effective means to improve productivity while maintaining adaptive capacity under certain conditions. However, it poses a considerable threat on indigenous breed biodiversity. Therefore, consequences for AnGR should always be one aspect to be considered in a project introducing exotic genetic material into the region. 1 . According to FAO (Scherf 2000) about one farm animal breed is lost per week on the global scale! References References Adebambo O.A. 1995. Selection response for higher body weight in Nigeria's indigenous crossbred pigs. 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Appendix: Farm animal genetic resources (AnGR) valuation method evaluation Appendix: Farm animal genetic resources (AnGR) valuation method evaluation Valuation methodology Purpose, objective or strength Role in conservation Type of data required Data availability Conceptual weaknesses or difficulties Methodologies for determining the appropriateness of AnGR conservation programme costs Contingency valuation Identify society's WTP for the conservation of AnGR Define upper bound to economically justified conservation programme costs Society preferences expressed in terms of WTP Not normally available. Requires survey Response difficulties when used for `non- charismatic' species and/or chronic genetic erosion Production loss averted Indicate magnitude of potential production losses in the absence of AnGR conservation Justify conservation programme costs of at least this magnitude Estimate of potential production losses (e.g. percentage of herd and market value of animals) Animal market values available for commercial breeds. Potential herd loss must be estimated. Not a consumer/producer surplus measure of value. Ignores substitution effects Opportunity cost Identify cost of maintaining AnGR diversity Define opportunity cost of AnGR conservation programme Household costs of production and net income Not normally available. Requires survey Least cost Identify cost- efficient programme for the conservation of AnGR Define minimum cost of conservation programme Household costs of production and profitability Not normally available. Requires survey Methodologies for determining the actual economic importance of the breed Aggregate demand and supply Identify value of breed to society Value of potential losses associated with AnGR loss Intertemporal or farm-level data Available for commercial breeds. Not normally available for others. Requires survey Requires shadow pricing of domestic labour and forage Cross- sectional farm and household Identify value of breed to society Value of potential losses associated with potential AnGR loss Consumer and producer price differences by location Not normally available. Requires survey Requires shadow pricing of domestic labour and forage Market share Indication of current market value of a given Justify economic importance of a given breed Market value of animal products by breed Generally available but not always by Not a consumer/producer surplus measure of breed breed value. Ignores substitution effects IPR and contracts Market creation and support for `fair and equitable' sharing of AnGR benefits Generate funds and incentives for AnGR conservation Royalty payments or terms of contract Usually available when such arrangements exist although can be commercial secret Limited duration of contracts Methodologies for priority setting in AnGR breeding programmes Evaluation of breeding programme Identify net economic benefits of stock improvements Maximise economic benefits of conserved AnGR Yield effects and input costs Available for commercial breeds. Not normally available for others. Requires survey/research Difficulty in isolating the contribution of genetic resources from other costs of programme Genetic production function Identify net economic benefits of stock improvements Maximise expected economic benefits of conserved AnGR Yield effects and input costs Available for commercial breeds. Not normally available for others. Requires survey/research Hedonic Identify trait values Value potential losses associated with AnGR loss. Understand breed references. Characteristics of animals and market prices Available for commercial breeds. Not normally available for others. Requires survey/research Not a consumer/producer surplus measure of value. Ignores substitution effects Farm simulation model Model improved animal characteristics on farm economics Maximise economic benefits of conserved AnGR Inputs and outputs. Technical coefficients of all main activities Available for commercial breeds. Not normally available for others. Requires survey Correct definition of farm objective function. Aggregation for estimating consumer surplus can also be problematic Source: Drucker et al. (2001).