R.H. Kemp
R.H. KEMP heads the unit of Tropical Silviculture, Commonwealth Forestry Institute, Oxford University.
The period since the last World Consultation has seen much activity, from the work of individuals and research institutes to the national and particularly the international scale. The developed countries, mainly in temperate regions, have been largely responsible for the elaboration of the philosophy and strategy of genetic conservation, and they have to a large degree the means to explore and conserve their plant genetic resources in their own interests, both severally and in collaboration. It is in the tropics and sub-tropics that the greatest potential exists for development of forest genetic resources, and it is also there that action on conservation is most urgently needed.
NICARAGUAN FORESTER SEARCHING FOR Pinus pseudostrobus SEEDS special skills
The FAO Panel of Experts on Forest Gene Resources initiated in 1968 an action programme of exploration and seed collection, which has been implemented through the work of research institutes and organizations, with a large financial contribution from bilateral aid funds, as well as some from the FAO regular programme (FAO 1969, 1972, 1974a, 1975a, 1976). In addition the Panel has helped to achieve the still growing acceptance that the conservation of forest genetic resources is a matter for global concern and action. This concern received expression at the United Nations Environment Conference at Stockholm, in 1972, and in the Global Programme for Improved Use of Forest Genetic Resources (FAO, 1974b), but action to conserve genetic resources on the scale required and recommended by the Panel has scarcely begun.
The problems of both exploration and conservation are different for a species whose natural range is restricted to a single country from those of a species which extends through many different countries, perhaps on different continents (Koster, 1977). The earliest schemes of international cooperation were between countries equipped and interested to exchange seed from their own sectors of a species' range. Cooperative action through the International Union of Forest Research Organizations (IUFRO) has been very successful and has embraced a further development, whereby one institute may undertake seed collections on behalf of several countries, including its own, which are interested in the species. Such expeditions (e.g., Barner, 1971) have been influential in the development of procedures, in revealing problems and in setting standards.
Some countries, with well-developed forestry services, already concerned with exploration and seed collection, provide material for international distribution, as seen in the work of the Poplar Council and the U.S. Forest Service Tree Seed Service Centre, or the Seed Centre of the Commonwealth Scientific and Industrial Research Organization (CSIRO) in Australia. The programme on eucalypts started by the Forestry and Timber Bureau, and continued by CSIRO (Turnbull, 1977), has been of the greatest importance for many developing countries which otherwise would not have had access to this valuable material. The modest financial contribution from FAO is an example of the most effective use of the limited international funds so far available.
Even for Australia the extent of the exploration required is an immense task, in its demands on the time of skilled staff, as well as the financial cost. An example of cooperation is seen in the seed collection mission of the French Centre technique forestier tropical (CTFT) to Australia, Timor and Indonesia in 1973 (Cossalter, 1974). Although this mission was undertaken primarily for the benefit of the countries where CTFT is involved in research programmes, many other countries may also benefit. As a result of trial plots, and some larger stands, hitherto unavailable provenances of tropical eucalypts have now been established in Africa for the first time, and these few exotic populations will become local sources for wider distribution of reproductive material to other African countries in the future.
AUSTRALIAN FORESTER SHOOTS DOWN BRANCHES FOR SEEDS aiming for high quality
Very few developing countries have the resources to undertake exploration and seed collection outside their own borders, and even within their own countries they have other prior demands on limited staff and funds. In these circumstances the programme recommended by the FAO Panel, and mainly financed and executed by bilateral aid organizations, notably in Denmark and the United Kingdom, has had the greatest success in recent years (Keiding and Kemp, 1977; Wood and Greaves, 1977). As a result of this work nearly 50 countries in the tropics and sub-tropics have been able to establish provenance trials of promising fast-growing species, including many provenances never previously collected. The trials show that some of these are distinctly different from and markedly superior to earlier introductions of the species (Keiding and Kemp, 1977). In order to complete range-wide provenance sampling, and to meet demands for trials and the further needs for gene conservation, several years of seed collection are normally needed. In some cases local seed centres have been established in association with the overseas institutes conducting exploration and seed collections, with bilateral aid funds, and this development offers the best prospects for future supplies of seed.
New thought has been given to the study of genetic variability and its implications for exploration. The theoretical basis of sampling strategies for crop plants has been examined in recent papers (see Hawkes et al., 1976), many of which are relevant to exploration of tree species (see also Stern and Roche, 1974). However, basic information on patterns of variation, flowering and fruiting behaviour and breeding systems is lacking, especially for tropical tree species. For this reason most exploration has been guided by general principles, modified by the practical necessities imposed by local conditions (Kemp, 1976). The explorations made have added greatly to previous knowledge, through collection of ecological and distribution data, herbarium samples and biochemical samples (e.g. pine resin) for taxonomic studies.
The international programmes in which the distribution of seed and coordination of trials are centralized, have opened new possibilities for genecological studies, as well as for the evaluation of provenance and environmental interaction (Burley and Kemp, 1973). They have made necessary the development of computerized systems for data storage and retrieval (e.g. Burley et al., 1973). Advances have also been made in methods for rapid evaluation of large numbers of wood samples from trees under the influence of the environment at the site of introduction (Hughes and Andrew, 1974). Through the collaboration between the institutes conducting and coordinating the research, and the nearly 50 tropical countries carrying out the field trials (see Wood and Greaves, 1977), new possibilities exist for the early recognition of populations deserving priority for further collection and gene conservation.
Recognition of the need to conserve forest genetic resources has widened and much thought has recently been given to the methodology of conservation, both in situ and ex situ (e.g., Fowler and Yeatman, 1971; Roche et al., 1975). Individual countries with adequate means to explore and protect their genetic resources are already taking such action in their own interests, and thereby helping to demonstrate and test the methods used (e.g. Hagman, 1971). In some cases neighbouring countries are cooperating in the compilation of lists of threatened species, and in the conservation of their genetic diversity, for example, Canada, Mexico and the U.S.A. (Barber and Krugman, 1974). Species known to be in serious danger are recorded in the International Union for the Conservation of Nature and Natural Resources (IUCN) Red Data Books, while others not severely endangered as species, but undergoing serious genetic impoverishment in some part of their range, are included in a series of data sheets produced under the auspices of the International Union of Forest Research Organizations (IUFRO) Working Party on Gene Resource Conservation (FAO, 1974 c).
Basic information on patterns of variation, flowering and fruiting behaviour and breeding systems is lacking, especially for tropical tree species.
In many tropical and sub-tropical countries the means are very limited, and international assistance, both financial and operational, has been proposed (Kemp et al., 1972; FAO, 1974b). Effective financial support from international organizations has yet to be secured. However the Governing Council of the United Nations Environment Programme (UNEP) decided to allocate funds to support conservation of genetic resources, during its first biennium, and subsequently UNEP agreed to finance a short term pilot study on "The Methodology of Conservation of Forest Genetic Resources" (Roche et al., 1975) Following the recommendations of this study, which supported recommendations made earlier by the FAO Panel (FAO, 1974b), a new project for the "Conservation of Forest Genetic Resources", submitted by FAO in June 1975, became operational in the second half of 1976.
As noted by Frankel (1977) secondary gene pools of a number of tree species already exist in many countries through the widespread use of exotics in plantations. These are normally managed, and their genetic resources are therefore influenced by production criteria, and the idea of plantations specifically to conserve forest genetic resources is comparatively recent (e g. Bouvarel, 1970). The FAO Panel (FAO, 1974b) recognized the international responsibilities involved and the principles and methods were fully discussed by Guldager (1975).
The project, as accepted by UNEP, constitutes the pilot stage of what is hoped to be an expanding, long term programme, and at present has a duration of only two years. The main emphasis is on action ex situ, through support for seed collection and the establishment of conservation stands in 10 tropical countries. The species proposed by the FAO Panel were chosen partly because of their proven importance and partly because seed of some of the most valuable and endangered provenances was already available from collections made by the Commonwealth Forestry Institute, Oxford (C.F.I.) and the CSIRO. During 1977 the distribution of seed, the selection and preparation of sites, and the raising of nursery stock for the conservation stands will be well advanced in several countries. Limited provision was also made to assist with conservation in situ of forest ecosystems in India and West and East Africa, but in this field dramatic achievements cannot be expected in such a short period.
Another field in which achievements have been made is seed research and seed storage. The seed of many fast growing plantation species can be readily stored for long periods, often for decades, without serious loss of viability or other apparent deleterious effects. However there are other species, particularly in the tropical rain forests, whose seed loses all viability within a few weeks, or even days, after seed fall, under normal conditions. This has been a major constraint on the use of such species in plantations, especially as exotics. Much information has recently become available, particularly in agricultural practice, but often of relevance to forestry (e.g. Harrington, 1970, 1972, Roberts, 1972, 1975; Wang, 1974, 1975; Barner, 1975, Turnbull, 1975).
Frankel (1977) has drawn attention to the new techniques in prospect for the storage of "recalcitrant" seeds (e.g. Villiers, 1975; Sakai and Noshiro, 1975). Although such fundamental studies will no doubt concentrate on agricultural seeds, the results are likely to improve the prospect for long-term storage of forest tree seeds, and may be particularly important for many tropical rain forest species. Little or nothing is known at present about the behaviour of the seed of many such species, and the way it may be affected by the treatment given during collection, handling and storage. The need for more studies of tropical tree seeds was stressed at the IUFRO International Seed Symposium, in 1973 (Simak and Kamra, 1973; Kamra, 1974) and at the International Seed Testing Association (ISTA) Workshop on Forest Tree Seed Testing, 1975.
Important advances have been made in the successful storage of seed which, while they are essentially "orthodox", require special care and treatment during the critical period immediately after seed fall to bring them safely to a suitable condition for low-temperature storage. Once through-this critical period they may be successfully stored for decades, thereby opening new possibilities for conservation and utilization. An example is the work on the seed of Agathis spp. As a result new possibilities exist for plantation trials and genetic conservation of selected populations of this valuable genus (Whitmore, 1977).
Despite some positive achievements in exploration and conservation, the loss of forest genetic resources has increased during the past decade, and is still accelerating. This loss cannot be exactly quantified; indeed lack of precise information on the nature, location and conservation status of valuable genetic resources is itself a major problem. Nevertheless the destruction of natural forest, particularly in the tropics, is so rapid and widespread that there can be no doubt that valuable populations have already been lost in recent years and others are severely threatened. The danger is probably most severe in the tropical rain forests, since these are not well represented in the U.N. List of National Parks and Equivalent Reserves, 1975. Some areas which have hitherto received selective protection within forest reserves are now threatened, as the natural forest is increasingly converted to high-yielding plantations, often using exotic species (Kemp et al., 1975). This process is evident and rapidly increasing in many African countries, while elsewhere areas such as the Amazonian rain forest, which had previously been protected by their inaccessibility, have recently been exposed to exploitation and massive clearance of forest. In South-East Asia it has been estimated that the entire timber resources of the tropical rain forests are unlikely to last beyond the end of the century (Whitmore, 1975). Owing to the great diversity of species in the rain forests, and the almost complete lack of knowledge of their ecology and genetics, their conservation is at present inseparable from the conservation of the ecosystems in which they occur (Roche, 1975b).
The danger of destruction is most severe in the tropical rain forests, even in protected areas, as the natural forest is increasingly converted to high-yielding plantations. It is happening in Africa, in the Amazonian rain forest, in southeast Asia.
Even species and populations already proved to be among the most valuable for-high-yielding plantations are in danger (Turnbull, 1977; Keiding and Kemp, 1977). Frequently the populations most endangered are those at the fringe of the geographical or ecological range (e.g. Kemp, 1975) and these may be among the most valuable for use elsewhere under extreme site conditions (Koster, 1977; Holzer, 1977). There is evidence that such marginal populations are genetically different from those in the centre of the range, due to the different selective pressures for survival (Sziklai and De Vescovi, 1977). For species already being used in plantations there is a danger of loss of genetic integrity within local populations, through contamination from other populations introduced from elsewhere (e.g. Frankel, 1977; Holzer, 1977; Keiding and Kemp, 1977; Turnbull, 1977).
There are at present only twelve tropical species in major international provenance trials, and for some of these the sampling of the natural range is far from complete. With the resources so far devoted to this work there seems little prospect of adding to this number at an average rate of more than perhaps two species a year. At this rate it would take more than a century to explore and collect representative material of those species already recommended by the FAO Panel as in need of such action.
The determination of priorities in the list awaiting action is most important but in this, as in many other aspects of exploration, the lack of information is a major problem (Keiding and Kemp, 1977). Later, at the start of exploration, lack of data on the natural distribution of a species, its variability, flowering and fruiting times throughout the range, and the possibilities for access, travel and transport of material further compounds the practical problems involved (Kemp et al., 1975). Really efficient sampling schemes can only be designed after a comprehensive study of the variation within the species but in many cases seed collection is an essential prerequisite to exploration of the genetic variation (Turnbull, 1977).
For many developing countries the need to select and multiply the most suitable high-yielding varieties, and to initiate further improvement based on these populations is immediate. However, seed of the provenances which have been proved superior in trials may be unobtainable. Even if the original population has not been entirely destroyed it may have been reduced by exploitation or forest clearance to areas of young regeneration, with perhaps a few overmature and unsound trees, or to remnant stands in remote and inaccessible situations. It has been noted that even for species of widespread importance the repro-curement of seed of provenances which have proved superior in trials in quantities sufficient to supply the needs of many developing countries is a major difficulty (Turnbull, 1977). The international programme of exploration and research, with the cooperation of many developing countries, can open the way to the creation of highly productive plantations on a large scale, but only if action is taken to ensure that sufficient quantities of reproductive material of the most appropriate provenances are available when and where they are needed (Keiding and Kemp, 1977).
SORTING A PROVENANCE COLLECTION OF Pinus oocarpa conserving gene pools
Despite the immediate danger, the longer-term prospects for conservation have improved since the last World Consultation, due partly to the awakening of interest in nature conservation, and partly to the development of cooperative action within the proposed Global Programme (FAO, 1974b ). The cooperative organization is still fragmentary and weak, with inadequate funds and staff, but stronger international support is in prospect.
Frankel (1977) has drawn attention to the special affinity between forest tree conservation and nature conservation, and to the possibilities for the establishment of gene pool reserves of threatened species in association with nature reserves (see also Roche et al., 1975). Turnbull (1977) indicates how this approach is developing in Australia, and similar examples could be drawn from other countries. The essential need for in situ conservation of the genetic resources of many tropical rain forest species has been noted earlier and, when 100 to 200 tree species occur in a single hectare, while the number of breeding individuals of a species may be only one or two per hectare, this may demand the setting aside of large areas of forest. However, it should be remembered that the genes, and gene complexes, which control most important characteristics may be safely preserved in relatively few individuals, and it is generally wrong to suggest that vast areas of forest must be set aside to meet the needs of gene conservation.
The integration of nature conservation and genetic resource conservation on the global scale was taken a stage further by the formation of the Expert Panel on MAB Project 8, in 1973, and the resulting final report on this project (Unesco, 1973). A later report on the choice and establishment of biosphere reserves (Unesco, 1974) recommended the provision of international funds to assist countries in the development of reserves, specifically suggesting recourse to the advice, cooperation and assistance of UNEP. Altogether over 200 sites for biosphere reserves have been proposed by forty countries. Most of the proposed biosphere reserves are to tee found within existing national parks or other protected areas and their designation may not in itself alter their previous protective status, or reduce the threat of damage or encroachment.
The International Union for the Conservation of Nature and Natural Resources (IUCN) is carrying out surveys to assess the extent to which representative samples of natural ecosystems are effectively protected in national parks, nature reserves and other protected areas (e.g., Lamprety, 1975; Whitmore, 1976). The conservation review of tropical rain forests, already completed for Asia, is being extended to Africa and America. Through the IUCN association with the World Wildlife Fund (WWF) substantial financial assistance may be secured for conservation in Indonesia and Suriname. A previous example is the creation of the Tiger Reserves in India. These are situated within existing forest reserves and include a "buffer zone", the recommended pattern for in situ conservation (Roche, 1975a). Such action, in view of its very strong financial and political support, offers good prospects for effective long-term conservation of forest genetic resources of tropical forests.
However, this example also illustrates the limitations of nature reserves for this purpose, since both selection and management of the Tiger Reserves are biased toward their primary purpose. Although a well-chosen pattern of strict natural reserves may conserve the genetic resources of the important tree species in some part of their range it may fail to protect particular valuable populations of a species, for example in marginal areas. Moreover many fast-growing plantation trees are pioneers, not members of the climax communities but of seral stages, and to conserve such populations in situ may require precise management of the reserve, or of a sufficient part of it, to that end, for example by a regime of controlled burning. This means that informed forestry opinion must play its part in both the selection and management of the nature reserves.
The establishment of plantations for the conservation of the genetic resources of selected species and provenances ex situ, in a variety of environments, offers new opportunities for the expression of existing genetic diversity, and for the development of new gene combinations, in harmony with the new environments.
CONES FROM A SINGLE PROVENANCE COLLECTION OF Pinus oocarpa nature works through variation
In the wider context of the conservation of plant genetic resources generally, a ' global network ' of genetic resources centres is beginning to emerge (Franker, 1975) and regional centres are being developed in key areas (e.g., Kjellquist, 1975). Regional centres for collection, distribution and conservation of forest tree seed, and for associated research, have long been advocated (e.g., Hagman, 1969) and prototypes exist in several tropical as well as temperate countries, although at present these may function mainly as national rather than regional centres. The evolution of a national centre into a regional one, wherever appropriate, is probably the best method of establishment. A national centre concerned with forest genetic resources is needed in every country, although in its first and simplest form it may be only an information centre, perhaps one man, located within an existing organization, to provide and receive information as part of the international network (Kemp, 1974). In response to a circular letter distributed by FAO in 1975, over 70 countries have so far each nominated a National Coordinator of Forest Genetic Resources Information and these will in future help to identify the danger spots where urgent action is needed, as well as to assist with information needed for exploration and collection. Much information could be gathered during the course of other routine operations, and without the use of extra staff, if simple procedures were instituted. Opportunities exist in the course of forest inventories and surveys, which may have other specific aims, which have ready access to information and material of great value for genetic resource exploration and conservation. All that may be required is some thought in advance, for example by the National Coordinator, and the provision of briefing material, questionnaires, recording forms and collecting apparatus, as appropriate. In some circumstances extra staff, perhaps with special skills, may need to accompany the party, but there would still be a saving in cost and time compared with a separate expedition.
The Consultative Group on International Agricultural Research (CGIAR), after prolonged study, has made available funds for exploration and conservation of plant genetic resources, and is supporting the development of the global network of genetic resources centres. The funds are administered by the International Board for Plant Genetic Resources (IBPGR). Priority has, of course, been given to food crops but in February 1976 the IBPGR agreed unanimously that it should give limited support to exploration and conservation of the genetic resources of a few tree species important in agricultural connections, in addition to actual food trees. (Subject to approval in principle by the CGIAR and based on its recommendations on species made by the FAO Panel at its March 1977 meeting, it is possible that international funds might be secured from the IBPGR.) The CGIAR is also a source of funds for training and documentation in genetic resource work. These activities are urgently required in the forestry sector.
Detailed proposals for action have already been made in the Global Programme (FAO, 1974b) and, given the necessary funds, could be implemented quickly. There are several other main constraints in addition to the financial ones, particularly the lack of information, lack of skilled and experienced staff, and lack of facilities for research. The strength of the Global Programme is that it is an integrated attempt to solve these various problems in parallel, by increased provision for training, information services, research, data storage and retrieval and other related activities, in addition to the field operations of exploration and conservation. Because it is designed to draw its funds from several sources, including both international and bilateral aid, implementation has been partial and uneven, since some major potential donors, such as UNDP and the CGIAR, have yet to provide the support requested.
While every effort must be made to achieve full implementation of the Global Programme, the current economic difficulties faced by many countries and international organizations, and the prior demands for food production, will restrict the possibilities for additional funding of forest genetic resources work. It is significant that the achievements so far made are very largely the result of individual effort and enthusiasm, and the more effective use of existing staff, facilities and funds, rather than any major increase in funds available, particularly at the international level. This is above all due to the coordination achieved through the FAO Panel of Experts on Forest Gene Resources, and the work of the FAO staff who act as the secretariat, in Rome. The appointment of the National Coordinators of information in the member countries could lead to the most efficient use of already existing staff, facilities and opportunities for action in exploration, collection and research. However, most of the individuals involved in this international network have other responsibilities, often in other sectors of work, that make more immediate demands on their time. The effectiveness of the network will depend entirely on the effectiveness of the coordination, and the enthusiasm of the individuals, and these can only be maintained through very frequent and close contacts, by visits as well as by correspondence. The coordination should be provided by the FAO secretariat in Rome, but the present staff resources there are totally inadequate to undertake it effectively
A national centre concerned with forest genetic resources is needed in every country. In its simplest form it can only be one man, located in an existing organization, to provide and receive information as part of the international network.
A strengthening of the FAO staff and activities in genetic resource work had been proposed and approved, but implementation of the proposed increase has been restricted in the current economic crisis.
Regional meetings and workshops can contribute greatly to the exchange of information, the maintenance of interest and enthusiasm, and the provision of necessary training. Excellent examples are the FAO/DANIDA (Danish International Development Agency) courses (e.g. FAO, 1974d; FAO, 1975b). To be most effective these must be held in the regions concerned, and should bring together the individuals most closely concerned with the work in their own countries. When well-staffed regional seed centres exist these can undertake such responsibilities but in the meantime the necessary expertise may be found in the institutes undertaking exploration and seed collection under bilateral aid programmes, for example in Australia, Denmark, France and the U.K. In either case provision of international funds for travel and subsistence of the participants may be needed, and the assistance of staff of the FAO secretariat in the organization of the courses is equally necessary.
Action taken during the next decade in exploration and conservation of forest genetic resources, particularly in tropical regions, will decide the continued existence, or disappearance, of many wild populations. The effectiveness of the coordination, and of the links for information and action between the areas in need and the organizations and institutes with the capability to act, will be decisive.
Note: References are at the back of the magazine, following the last article.
