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Conservation of 
genetic resources 
in tropical forest 

Principles and concepts 

Based on the work of 

R.H. Kemp 

with scientific review by 

G. Namkoong 


F.H. Wadsworth 

The designations employed and the presentation of material in this 
pubUcation do not imply the expression of any opinion whatsoever on 
the part of the Food and Agriculture Organization of the United 
Nations concerning the legal status of any country, territory, city or 
area or of its authorities, or concerning the delimitation of Its frontiers 
or boundaries. 

ISBN 92-5-1 03309-9 

AN rights reserved. No part of this publication may be reproduced, stored in a 
retrieval system, or transmitted in any form or by any means, electronic, mechani- 
cat, photocopying or otherwise, without the prior permission of the copyright owner. 
Applications for such permission, with a statement of the purpose and extent of the 
reproduction, should be addressed to the Director, Publications Division, Food and 
Agriculture Organization of the United Nations. Viale delle Terme di Caracalla. 
00100 Rome, Italy. 

FAO 1993 


Growing populations and pressure for social and economic development are 
leading to increasing rates of destruction and degradation of natural habitats, 
including forests and woodlands. The loss of natural resources and the degradation 
of land are already affecting the economies and well-being of the people in many 
countries, especially in the tropics. The loss of habitats are leading to 
accelerated rates of loss of genetic resources, which are fundamentally important 
in the adaptation and improvement of plant species presently under cultivation and 
those whose value is yet to be ascertained. 

It is today widely recognized that the existing Protected Area system on its 
own is not sufficient to provide the necessary geographic and biological coverage 
to conserve either the exceptional diversity of tropical forests or the genetic 
resources of their main component species. It is also recognized that, to succeed, 
conservation must be seen not as a constraint but as an integral part of 

Although forest management interventions will cause more rapid changes in the 
composition of ecosystems than natural forces and, at times, will also accelerate 
or alter successional changes, such interventions can be rendered compatible with 
the conservation of the genetic resources of the species under use. The sustained 
utilization of forests to meet present-day needs coupled with the maintenance of a 
network of areas dedicated to the protection of ecosystems and their functions, 
provides the only solution for lasting, genetic conservation. 

In accordance with its mandate, FAO has for the past 40 years published a 
number of manuals on the sustainable management of tropical forests, complemented 
over the past 20 years by guides on the conservation of forest genetic resources. 
The technical feasibility of both tropical forest management and conservation have 
been stressed repeatedly in these documents. 

The present book constitutes a first step towards a more systematic approach 
to the provision of guidelines for harmonizing sustainable utilization and 
conservation of genetic resources of tropical forest trees. Many presently 
prescribed forest management interventions could with minor adjustment be made less 
harmful to conservation concerns. Conversely, some compromises could be made in 
existing methodologies for the conservation of forest genetic resources which could 
help achieve the main aims of conservation while at the same time meeting pressing, 
present-day needs for the goods and environmental services provided by the forest. 

This document outlines present forest management practices, illustrated by 
case studies from three tropical countries. It briefly reviews available 
strategies and methodologies fof the conservation of forest genetic resources in the 
light of their compatibility with sustainable use of the resources targeted for 
conservation. It is planned to publish, in the near future, a companion volume to 
the present book, in which more specific guidance is given on the management of 
specific forest types and tree species, in situations in which varying degrees of 
priority are given to production and conservation respectively, while at the same 
time meeting at least some of the needs of both of these two complementary aspects. 

f.P. Lanly 


Forest Resources Division 



The present study is based on the work of Mr. R.H. Kemp of the United 
Kingdom and was carried out under the technical guidance of staff of the 
Forest Resources Division of FAO. Professor Gene Namkoong (U.S.A.) and Dr. 
F.H. Wadsworth (U.S.A.) provided scientific review of the document, which also 
benefited from the comments of Dr. J. Wyatt-Smith (U.K.) and colleagues in 
IUCN, Unesco, the Royal Botanic Gardens, Kew (U.K.) and the Oxford Forestry 
Institute (U.K.). Thanks are due to Dr. D. Boshier for providing information 
related to Cordia alliodora (see Appendix 1 ) , and to the Overseas Development 
Administration (ODA, U.K.) for providing information and material related to 
the Case Studies on Ghana and India. Valuable inputs were further made by J.R. 
Palmer, M.E.D. Poore and T.J. Synnott, among many others. 

Greatest credit for achievements in the field on which information in 
this document is based, however, is due to the dedicated efforts of national 
forestry and other staff in the countries concerned. The deep interest of 
professional foresters in the nature, complexity and functioning of the 
natural forests and their openness and willingness to discuss these issues and 
to share their in-depth knowledge for common benefit, is gratefully 



Executive Sumary vii 

Glossary xi 

Key to Abbreviations xiv 


1 . Introduction 1 

2. The Nature of Forest Genetic Resources 5 

2.1 Levels and structure of genetic diversity 5 

2.2 Ecosystem conservation 6 

2.3 Conservation of target species 7 

2.4 Conservation of provenances 7 

2.5 Values of genetic diversity 8 

2.6 Use values and option values 9 

2.7 Precautionary values 9 

2.8 Existence value 10 

2.9 Location of conservation areas 10 

2.10 The link to production forests 11 

2.11 Size of conservation areas 11 

2.12 Dynamic conservation 12 

2.13 Disturbance and succession 12 

2.14 Logging and genetic diversity 13 

3. Impacts of Management in Production Forests 15 

3.1 Continuity and control 15 

3.2 Economic and market influences 16 

3.3 Forest inventory 19 

3.4 Forest dynamics 21 

3 . 5 Regeneration 23 

3.6 Silviculture 25 

3.7 Harvesting 28 

3.8 Non-timber forest products 30 

3.9 Involvement of local people 32 

4. The Future of Tropical Forests 35 

4.1 Population and land use 35 

4.2 Timber demand and international trade 36 

4.3 Tropical forests and environmental concerns 39 

4.4 Protected Area systems 41 

4.5 Buffer Zone forest 43 

5. Strategies for in situ conservation in procuction forests 45 

5.1 National policies 45 

5.2 Management information 48 

5.3 Management systems 49 

5.4 Management plans 52 



6. Ghana 61 

6.1 The economy 61 

6.2 The environment 61 

6.3 Diversity 62 

6.4 Management for timber production 62 

6.5 Policy: linking production and conservation 64 

6.6 Non-timber forest products 64 

6.7 Forest revenue systems 65 

6.8 Forest inventory 67 

6.9 Setting priorities 68 

6.10 Management and harvesting 70 

6.11 Regeneration and silviculture 71 

6.12 Reproductive biology 72 

6.13 Integration and security 72 

7. Brazil: the Amazon forests 73 

7.1 Legal framework 73 

7.2 Setting priorities 75 

7.3 Management options 76 

7.4 Secondary forest and non-timber forest products 77 

7.5 Information, research and coordination 78 

8. India: the Western Ghat forests, Karnataka 79 

8.1 National policy 79 

8.2 Western Ghat forests 80 

8.3 Strategy for integrated development and conservation 81 

Appendix 1. Methodology of a study of the reproductive 

biology and genetics of Cordia alliodora 85 

Box 1. The role of logged forests in the conservation of 

species richness and genetic diversity 14 

Box 1. Management of diversity through 

diversity of management 56 

References 91 

Cover photo; Recently logged forest, Sarawak (Malaysia). 
R.H. Kemp 


General Considerations 

1 . There is today a growing realization at national and international 
levels of the value of forests as a renewable resource and of their role in 
the production of a range of goods and environmental services. The latter 
include the role of the tropical forests as a source of genetic materials for 
the adaptation and improvement of plant species presently under cultivation 
and use, and those whose value is yet to be ascertained. 

2. Increased populations and pressure for social and economic development 
in most tropical countries contribute to a continuing trend of diminishing 
areas under forest cover despite concern for the tropical forest. They also 
severely limit the possibilities for extending existing systems of fully 
protected areas. 

3. While the exceptional biological diversity of tropical forests 
constitutes a unique national and global asset, the extent and integrity of 
this diversity are therefore rapidly diminishing. 

4. The continuing availability of diversity and genetic resources is 
fundamental for the sustainable development of nations. Conservation of 
genetic resources of forest trees and other woody species is closely related 
to all other forms of diversity, and is essential for sustaining the 
productive and protective values of the forest. 

5. In spite of their importance at national as well as local levels, 
forests containing socio-economically valuable tree species have rarely been 
targeted when planning and establishing Protected Areas. Therefore, managed 
production forests play a key role in programmes aimed at the conservation of 
genetic resources of such tree species and are a necessary complement to 
conservation efforts undertaken through Protected Area management. 

6. Management interventions in the forest can be aimed mainly at the 
production of timber, wood and other products, the protection of soil and 
water, or the conservation of biological diversity and genetic resources. 
Productive and protective purposes can be rendered compatible with 
conservation concerns through sound planning and inter-sectoral coordination 
of activities at the national level. 

7. International concern for the conservation of biological diversity is 
likely to lead to increased support to tropical countries through both aid and 
trade channels. Since it is now widely recognized that the Protected Area 
system on its own is insufficient to provide the necessary geographical and 
biological coverage, such assistance may in the future be directed 
increasingly to support forest management carried out with due concern for 
genetic conservation. 

Conservation of Forest Genetic Resources 

8. Genetic resources are associated with the different levels of diversity 
that exist in nature, from ecosystems to species, populations, individuals and 
genes. These levels interact closely and all must be considered when 
conservation objectives are defined and when corresponding action is 


9. Conservation of genetic resources in situ is dependent on maintaining 
the essential functional components of the ecosystem. It implies the planned 
and systematic management of identified target species in a network of 
conservation areas which will include Strictly Protected Areas as well as 
managed forests and multiple-use reserves. 

10. Tropical forests are dynamic and subject to change through natural 
disturbance and succession; the aim of conservation is not to freeze a given 
state but to contain a dynamically evolving system. 

11. Lack of information on population biology, reproductive systems, 
variation and genetics of most tropical tree species limits the possibilities 
for the deliberate management of their genetic resources. Maintaining a broad 
genetic base through the conservation of a range of provenances of target 
species is likely to be the safest available option until more adequate data 
are available. 

12. Conservation efforts must be planned at the national level, but close 
linkages to regional and global efforts are necessary to ensure success. 

Conservation of Genetic Resources and Forest Management 

13. The sustained utilization of forests to meet present-day needs coupled 
with the maintenance of a network of areas dedicated to the protection of 
ecosystems and their functions, provides the only solution for lasting, 
genetic conservation. Harmonizing conservation and management for the 
production of goods and services is especially important in relation to 
tropical tree species which are not included in plantation and domestication 

14. The domination of short-term economic and market forces over ecological 
and technical considerations have frequently been the cause of past failures 
to attain sustainability in natural forest management and conservation of the 
species being utilized in the tropics. 

15. Information on forest composition and growth is critically important for 
both sustainable production and genetic conservation; broadly-based 
inventories, including botanical surveys, regeneration sampling and 
information on non-timber forest products, can be combined to serve both 

16. Some silvicultural operations, including canopy manipulations to favour 
certain species and individuals, can lead to a reduction in the overall 
diversity of tree species in a stand. However, these practices might also be 
skilfully used to maintain or restore diversity, in selected areas. 

17. Logging is at present commonly the only large-scale management 
intervention in the tropical forest. It may either reduce or enhance species 
and intra-specific diversity. It may furthermore contribute to the depletion 
or conservation of the genetic resources of the principal species being 
utilized, depending on the timing, intensity, frequency and discrimination 
employed, and on the effectiveness of protection and management of subsequent 


18. Large scale intensive logging, on short felling cycles, with poor 
harvesting control, may alter the species composition and may damage both 
forest structure and site quality. Logging damage is indiscriminate in its 
impact on genetic resources but heavy logging tends to enhance the opportunity 
for pioneer species. Should this be the case, special attention may have to 
be paid to the conservation of rare populations and species of later 
successional stages. 

19. While harvesting of mature trees of good quality is among the stated 
objectives of forest management aimed at the production of timber, pressing 
market demands coupled with inadequate forest management practices may lead 
to highly selective harvesting having negative (dysgenic) effects on the 
future development of the stand. Silviculture rightly calls for harvesting 
of "the best", but this must not be done without due consideration to 
regeneration potential and the quality of the next generation crop. 

20. The harvesting of non-timber forest products will widen the range of 
management objectives and will diversify management interventions. Hence it 
may increase the possibilities for sustainable management and conservation of 
genetic resources both by broadening the range of species contributing to 
production, and strengthening support of local communities for forest 
conservation through the provision of direct benefits to people living in or 
close to the forest. 

Action Required 

21 . Continuity of management based on adequate information on forest 
composition and dynamics, is essential to meet the production and conservation 
objectives of areas presently under forest. Forest management should be 
carried out within the framework of established plans acceptable to all land 
users and executed by well-trained, informed and motivated staff. 

22. The potential contribution of each area of production forest to the 
conservation of diversity and genetic resources of its component species, and 
the setting of relative priorities between areas in terms of production and 
conservation objectives, must be determined as part of an integrated strategy, 
based on national policies, with appropriate regional and international 

23. Integrated action extends beyond the management of production forests 
and Protected Areas, to aspects of forest industry, marketing and trade, and 
to other sectors in the context of national development policies. 

24. Involvement of local people living in or benefiting directly from the 
forest is fundamental to success of any conservation and management efforts, 
in the short as well as in the long term. 

25. The formulation of a National Strategy for the Conservation of Forest 
Genetic Resources is the most appropriate means to secure the necessary 
integration of action at the national level, and to define appropriate 
institutional mechanisms for its implementation. 

26. The formulation of such a National Strategy for Conservation is an 
essential nucleus for broader action in the conservation of terrestrial 
biological diversity. It will also provide a solid and credible basis for the 
substantial international support which will be required to meet national and 
global conservation objectives. 

27. The scale and complexity of the data needed to meet national 
conservation, protection and production objectives will require coordination 
through some form of a national conservation data centre, which should help 
facilitate and stimulate the collection of related data, make such data easily 
accessible for field implementation and use, and provide international links. 

28. Specific forest management systems must be decided for each forest or 
unit of management in accordance with their specific roles under the National 
Strategy for Conservation. This will help achieve an appropriate balance 
within the national forest estate as a whole, with due weight to ecological 
as well as socio-economic objectives. 

29. Special attention and priority may have to be given to the conservation 
of forest genetic resources in certain areas of the productive forest estate, 
for example at the edges of a species 1 geographical or ecological range, where 
the populations are likely to be genetically distinct, and where they may be 
particularly vulnerable to disruption. 

30. Special attention must further be paid to the generally species-rich 
secondary forests in various stages of recovery, and to the conservation of 
genetic resources of species characteristic of the mature-phase or climax 
forest. This may at times involve the adoption of some constraints on 
management for production, in specific areas of the natural range of selected 
species or communities, at any given time. 

31 . Many practical aspects of concession allocation, duration and operation, 
of critical importance to sustainable forest management for timber and other 
products, also decisively influence possibilities for the conservation of 
genetic resources. Special ecological, taxonomic or other advice maybe needed 
in relation to conservation concerns, for example in the location and 
distribution pattern of seed trees or patches of forest to be left temporarily 
unlogged, to ensure regeneration and maintenance of desirable levels of inter 
and intra-specific diversity. 

32. In all aspects of forest management and conservation the failure to 
comply with prevailing management prescriptions has been to date a common 
cause of unnecessary damage to site, growing stock and regeneration. Strict 
monitoring and control, based on criteria specified in corresponding forest 
management plans and the National Strategy for Conservation drawn up with due 
concern to environmental, technical, social and economic concerns, should be 
carried out on a continuing basis in each management unit and at national 

33. Intensified research into many aspects of taxonomy, forest dynamics and 
the functioning of ecosystems and individual tree species (including 
reproductive biology), are needed to meet both production and conservation 
objectives. Prioritization and coordination of research under the National 
Strategy for Conservation are essential to optimize use of the limited 
scientific resources available. 










Conservation of 
genetic resources 



Ex situ 



Any of the different forms of a gene which may occupy 
the same position (locus) on a chromosome (see also gene). 

Reproduction without fertilization, so that the progeny 
are genetically identical to the parent. 

The ecology of an individual species as opposed to 

The variety of life forms, the ecological roles they 
perform and the genetic diversity they contain. 

Major regional ecological community. 
Terminal stage of ecological succession. 

Threadlike body found within the nucleus consisting 
primarily of DNA (desoxyribonucleic acid) and a protein 
sheath, and containing the genes responsible for most 
hereditary traits. 

Description and delimitation of the distribution of a 
species or other taxon. 

A group of different organisms in close interdependency 
and inhabiting a common environment. 

The management of human use of genetic resourcs so that 
they may yield the greatest sustainable benefit to the 
present generation, while maintaining their potential to 
meet the needs and aspirations of future generations see 
also in situ conservation; ex situ conservation). 

A successional stage maintained short of the true climax 
by e.g. fire. 

A community with its physical environment, interacting 
as a functional system. 

Natural occurrence confined to a particular locality. 

Any conservation method that entails removal of individual 
plants or propagating material (seed, pollen, tissue) from 
its site of natural occurrence, i.e. conservation "off- 
site* 1 in gene banks as seed, tissue or pollen; in 
plantations; or in other live collections, such as ex situ 
conservation stands. 


Basic unit of inheritance; the physical entity being 
transmitted during the reproductive process, and 
influencing hereditary traits among the offspring. Genes 
can exist in different forms or states, called alleles. 


Gene flow 
Gene pool 
Genetic diversity 

Genetic drift 
Genetic resources 


IS situ 


Keystone species 



Exchange of genes between populations owing to the 
dispersal of garnets or zygotes. 

The total sum of genetic material of an interbreeding 

The heritable component of variation. Genetic 
diversity occurs at the gene level, the individual 
level, the population level, the species level and the 
ecosystem level. Genetic diversity is one component of 
biological diversity (see above) 

Random changes occurring by chance in the genetic make- 
up of small, isolated populations. 

The economic, scientific or social value of the heritable 
materials contained within and between species. 

Genetic constitution of an individual (particular set of 

A group of species having similar ecological 
requirements and roles in the community e.g. pioneer 

Mating of closely related individuals. 

Conservation of genetic resources of target species "on- 
site 11 , within the natural or original ecosystem in which 
they occur, or on the site formerly occupied by that 
ecosystem. Although most frequently applied to populations 
regenerated naturally, in situ conservation may include 
artificial regeneration whenever planting or sowing is done 
without conscious selection and in the same area where the 
seed or other reproductive materials were collected. 

Variants of a given enzyme performing the same catalytic 
function but exhibiting the presence of different genes 
(syn. isocnzyme) . 

Organisms which are critical in the functioning of an 
ecosystem (syn. keystone mutualists, pivotal species; 
see also Chapter I, page 6). 

Sudden heritable change in the gene or chromosome 
constitution causing changes in number, structure, size 
or sequence. 

For an indigenous stand of trees the origin is the place 
in which the trees are growing; for a non-indigenous 
stand the origin is the place from which the repro- 
ductive materials were originally introduced (see also 
provenance) . 


Mating of non-related (or distantly related) individuals. 



Secondary forest 




forest management 

Tropical Hoist 

The observable character of an individual resulting from 
interaction of the genotype with the environment. 

A group of interbreeding individuals occupying a 
particular area and usually separated to some degree 
from other similar groups. 

Population of a species referred to by its locality of 
occurrence; the place in which any stand of (indigenous 
or non-indigenous) trees is growing (see also origin). 

Forests which have suffered various degrees of disturbance 
as a result of shifting cultivation, or various intensities 
of logging, as opposed to supposedly virgin forest, or 
mature-phase forest which has achieved more or less the 
climax stage of development. 

One or more populations, the individuals of which can 
interbreed, but which cannot exchange genes with members 
of other species. 

A community of trees possessing sufficient uniformity of 
composition, constitution, age, spatial arrangement or 
condition, to be distinguishable from adjacent 
communities, so forming a silvicultural or management 

Founded on the properties of probability and chance or 
random variation. 

The administrative, economic, social, legal, technical and 
scientific aspects of the conservation and use of forests 
within the framework of a technically sound and politically 
accepted overall land use plan. It implies various degrees 
of human intervention, ranging from action aimed at 
safeguarding and maintaining the forest ecosystem and its 
functions, to favouring given socially or economically 
valuable species or groups of species for the improved 
production of goods and services. The term can only be more 
precisely defined in terms of the management objectives of 
a particular forest, however, it should always incorporate 
the principles of sustainable development, to "meet the 
needs of the present without compromising the ability of 
future generations to meet their own needs". It therefore 
has essential objectives in common with the conservation 
of genetic resources. Both concepts can be most 
effectively and efficiently applied in the context of the 
national forest estate as a whole. 

The closed high forests in the tropics, where the dry 
season is short (4 months) or non-existent. It includes 
both rain forests and monsoon (seasonal) forests. 



ECG Ecosystem Conservation Group (presently includes representatives of 

FAO Food and Agriculture Organization of the United Nations 

IUCN The World Conservation Union (earlier known as the International 
Union for Conservation of Nature and Natural Resources) 

UNDP United Nations Development Programme 
UNEP United Nations Environment Programme 
Unesco United Nations Educational, Scientific and Cultural Organization 

WWF World Wide Fund for Nature (earlier known as World Wildlife Fund 

WRI World Resources Institute 


International concern for tropical forests has risen sharply during the 
past decade. This period has seen the start of the Tropical Forestry Action 
Programme (TFAP) as a framework for action in tropical forestry, and the 
creation of the International Tropical Timber Organisation (ITTO) as a forum 
of consumers and producers of tropical timber moving in international trade, 
and discussion of the problems of deforestation at the highest international 
levels, in the World Commission on Environment and Development, in the UN 
General Assembly, in the Intergovernmental Panel on Climate Change, at summit 
meetings of the group of seven leading industrialised countries, and last but 
not least the United Nations Conference on Environment and Development in Rio 
de Janeiro, Brazil in June 1992. At the same time the FAO Forest Resources 
Assessment 1990 Project has revealed that the rate and scale of tropical 
deforestation in the past decade have been even greater than was previously 
predicted (FAO 1992c). 

Central to any solution to this problem must be the reconciliation of 
the need for development and increased living standards in the tropical 
countries, based on the sustainable utilization of each nation's resources, 
including the natural forests, with the conservation of their capability for 
renewal and adaptation to changing conditions and needs: their genetic 
resources. The problem is complex, being bound up with broader aspects of 
land use and population increase, as well as the impacts of international 
debt, trade and aid relationships. Part of the difficulty in the search for 
quick and effective solutions is the very complexity of the tropical forests 
themselves. In terms not only of the many varieties of dry forests, moist 
forests, hill and coastal formations, but also the different social and 
economic conditions that surround them, there is not one but many thousand 
forest problems, each in need of separate analysis and management. 

One aspect of this complexity of the tropical forests that has now 
attracted international interest is their biological diversity. This brings 
a new appreciation of this unique value of the natural forest ecosystems, in 
addition to their productive and environmental values, which have often proved 
insufficient to ensure their protection. Diversity considerations are 
generally poorly, or not at all, represented in formulae used to determine the 
economic value of an area, and have been discounted in competition with the 
immediate needs for land, food production and revenue from intensive timber 
production, for example through fast-growing plantations, in place of natural 
forest. As human populations continue to increase the opportunity cost of 
setting aside large areas purely to conserve biological diversity for ethical 
reasons and often un-specified "future needs", becomes increasingly difficult 
to defend. At the same time pressures to maximise short-term timber volumes 
and revenue from production forests may increasingly threaten the continued 
existence of natural forests under selective harvesting systems, in which 
funds and revenue are re- invested in care and management of the areas 
harvested. However to the extent that the conservation of genetic diversity 
can be seen as one of the services of natural forest under multiple-use 
management, including appropriate levels of sustainable timber production, the 
reconciliation of developmental and conservation objectives may be easier to 
achieve. The exceptional wealth of the tropical forests in biological 
diversity makes this a strong possibility, given adequate provision for 
effective management. 

In the wake of preferences for other land use options, the progressive 
reduction in size and increasing isolation of the remaining areas of natural 
forest in most countries imposes the need for increased efficiency in their 
management. This is partly to ensure the maximum levels of production, to 
defend the forest against increasing demands of increasing populations, but 
also to the increased stresses on the ecosystem resulting from the loss of the 
"buffering" effects of large areas of surrounding forest. This changed 
situation may have direct impacts on the environmental conditions around the 
forest, and also indirect influence on the breeding and seed dispersal systems 
of tree species dependent on animal vectors. 

The component of the genetic diversity in the forest which is of actual 
or potential use, either for production or to maintain the forest as a 
functioning system, constitutes its genetic resources. The conservation of 
genetic resources is absolutely fundamental to the long-term success of all 
other forms of diversity conservation (Riggs 1990) and is also essential to 
the sustainable and productive management of the forest ecosystem in which 
they occur. To this extent therefore in situ conservation of forest genetic 
resources should reinforce the conservation and management of production 
forests, and vice versa. At the same time the systems of National Parks and 
other fully Protected Areas also have an essential role, particularly in the 
conservation of biological diversity of very uncertain use value or existence 
value, as well as the genetic resources of species of economic importance. 
However the coverage of the tropical forest biome within such systems is 
limited and they will never satisfy the full needs of genetic resource 
conservation. New approaches are therefore needed to integrate national 
conservation activities for maximum effect in both production forests and 
fully Protected Area systems. 

FAO has the leading international role in linking forest management to 
the conservation of forest genetic resources, particularly through the work 
of the FAO Panel of Experts on Forest Gene Resources, established in 1968. 
Folliwing publication in 1975 of the book, " Methodology of Conservation of 
Forest Genetic Resources" (FAO 1975), "Guide to in situ Conservation of 
Genetic Resources of Tropical Woody Species" was published by FAO in 1984 
(Roche and Dourojeanni 1984). These documents reviewed the issues and 
activities involved in genetic resource conservation in forestry. Three case 
studies, from Cameroon, Malaysia and Peru, were published in the later 
document, which further reviewed the role of Protected Area systems, including 
those within production forests, such as the Virgin Jungle Reserves in 
Malaysia, which were discussed in detail. The principles and procedures set 
out in both documents still provide valuable guidance to the subject. Since 
then the compilation and publication of reviews of management practices in 
tropical moist forests in Africa (FAO 1989b), Asia (FAO 1989c) and Latin 
America (FAO 1992a) together with other broader reviews of the subject from 
both the managerial and the ecological viewpoints (e.g. Mergen and Vincent 
1987; Wyatt-Smith 1987a; Poore 1989; Whitmore 1990; Gomez-Pompa et al 1991), 
have provided new opportunities to consider the possible role of production 
forests in relation to the conservation in situ of forest genetic resources. 
The FAO Panel, at its 7th Session in December 1989, endorsed proposals to 
update and expand the guidance given in the 1984 publication on in situ 
conservation of tropical woody species. In doing so it was decided to 
concentrate on the role of tropical forests under management for the 
production of timber or other products and services in the conservation of the 
genetic resources of the woody species. These, and particularly the largest 
and longest-lived trees, are dominant in determining the unique genetic 
potential of the system. 

The diversity of tropical forests is a product not only of the large 
number of species present on a given area but of successional change over 
time, from the colonisation of gaps or cleared areas by pioneer species, 
through complex successional stages, to the mature "climax 11 forest. Economic 
forces and market demands have produced management systems aimed to simplify 
and to truncate the natural complexity and successional stages in the forest, 
to concentrate the growth potential into relatively few species, and short 
cutting cycles, with very limited reinvestment in management activities. This 
situation, in which short-term economic considerations have frequently 
overridden ecological concerns, has been governed by the apparently low value 
of the production per unit area of forest, as expressed in fees and log values 
in the forest. This has been generally incompatible with the level of 
investment in management needed to conserve the genetic resources even of the 
most important economic species, except perhaps and incidentally, those of the 
pioneer and fast-growing gap-opportunist guilds. Even on the basis of the 
very limited knowledge of the species composition and intraspecific variation 
in the forests it is clear that large parts of the useful or potentially 
useful genetic resources are in danger of being lost. Resolution of this 
problem is dependent on a wide range of integrated action, within which 
scientific research and professional forest management are only part of the 

The rapid pace of change in market demands and opportunities has often 
rendered invalid both the initial objectives of management and the programmes 
of data collection or silvicultural action undertaken to achieve them. The 
danger of adopting too narrow and short-term objectives is even greater in 
respect of genetic resource conservation, which has to consider possible 
changes in needs and opportunities over much longer periods than one or two 
rotations. These are likely to include rapid advances in the means to handle, 
manipulate and recombine genetic material. At the same time advances in 
information technology are already transforming the possibilities to handle 
and interpret large and complex arrays of data for the better understanding 
and use of functional relationships in the management of the forest. These 
developments greatly improve the opportunities for multiple-use management, 
including the conservation of genetic resources. A very important aspect of 
such multiple-use approaches, both for the long-term security of the forest 
ecosystem and for the conservation of genetic diversity, is the incorporation 
of non-timber forest products and the interests of local communities within 
the management system. 

Whereas in the past, with more limited resources for managing complexity 
in the system, and narrower management objectives, the response to the 
forest's diversity was to simplify the system, by concentrating action on few 
species, often with unpredictable and unexpected results, much of the current 
and future value of the natural forest is seen to lie in its genetic 
diversity. A more appropriate future response to the management of diversity 
will be greater diversity of management. This may be achieved in various ways 
and at various levels, from multiple-use of the same area of forest, either 
simultaneously or in successional stages, through separate management by 
compartments or working circles, to the integrated and diversified management 
of the entire national forest estate, embracing production forests, protection 
forests, genetic conservation areas, Protected Area networks, and areas 
combining two or more of these functions. 

Even for the management of genetic resources of species whose natural 
range transcends national boundaries the national borders define the ultimate 
unit of effective management, whose freedom and effectiveness to act may be 
further enhanced through international cooperation. It is ultimately the 
national government which holds the power to formulate the necessary policies 
in land and resource use which govern the possibilities both for sustainable 
forest management and for the conservation of the nation's biological 
civersity and genetic resources. 

Each country, and to a large extent each forest area, is unique in terms 
of its genetic resources and of the appropriate strategies at both national 
and local level to manage the forests, both for production and for 
conservation objectives. Attempts at management solutions at the level of the 
individual forest, alone and in isolation from its setting in national 
development policies, not only for forest and land use but embracing forest 
industry, trade and the linkages with other sectors, can only have very 
limited chance of success. 

For this reason a detailed case study of the problems facing the 
conservation of forest genetic resources in one country (Ghana) and of 
approaches to their solution, with shorter contrasting examples from two other 
countries (India, Brazil), form a large part of this overall study of policies 
and activities needed to link conservation with forest production through 
appropriate management. These case studies are presented in Part II of the 
present document. 

In Part I, Capter II brief ly summarises the essential elements of forest 
genetic resources. Chapter III considers the nature and extent of the impacts 
of management for timber production on the genetic resources of the forest, 
and some possibilities for multiple-use management. Chapter IV attempts to 
foresee the future for the natural forests, in terms of both international and 
domestic demand for timber and other products, and the probable influence of 
other factors such as population increase, and environmental concerns, 
including international interest in the conservation of biological diversity. 
Finally in Chapter V suggestions are made for the formulation of National 
Strategies for the Conservation of Forest Genetic Resources. 

Appendix 1 provides an example of the careful research needed to provide 
information on the reproductive biology and genetics of individual target 
species within a national strategy. 

It is hoped to complement the present study in the future with more 
detailed guidelines aimed at harmonising sustainable forest management for 
productive purposes with the conservation in situ of the species, provenances 
and populations under use, for the benefit of these two complementary 


In the same way that the term "forest resources" refers to the 
usefulness of the forests for the production of timber or other products for 
human benefit, the term "genetic resources" implies that elements of the 
genetic variability of the trees and other plants and animals will be used to 
meet human needs and objectives. However as compared with the harvest of 
timber or the collection of other products from the forest, which can be used 
immediately, the benefits from the genetic resources can be used not only in 
current programmes but are essential for future development in the next and 
subsequent generations; and for the continued adaptation of the resources to 
changing environmental conditions and human needs. 

The other important aspect of the genetic resources of natural forests, 
especially the tropical forests, is their great diversity, and this range of 
variation provides the basis for selection and improvement of the products and 
other benefits to meet future needs, so far as they can be foreseen. 
Therefore the greater the uncertainty over future demands, the greater the 
potential value of conserving diversity. Apart from the relatively small 
number of tree species of current economic importance, or domestic importance 
to local communities, which must be the main focus of conservation programmes, 
there are likely to be several hundred or even thousands of other species 
present with lesser or unknown values. Some of these might form an important 
part of future harvests of wood, timber and other forest products, in response 
to changing environmental conditions or market demands. 

2.1 Levels and structure of genetic diversity 

Genetic diversity occurs at various levels of organisation from the 
ecosystem, through its component species, their sub-specific populations 
(provenances), family groups and individual genotypes to the molecular level, 
of the gene. While ecosystem conservation may also achieve the conservation 
of some included species and genotypes others might be lost unless, for 
example in the case of important timber trees, the species and its genetically 
distinct component populations, are also targeted for specific conservation 
measures (FAO 1989a). Within such populations there is likely to be 
substantial variation between individual trees (genotypes). Depending on the 
pattern of distribution of such variation through the stand, as a result of 
the nature of the species' breeding and dispersal systems, highly valuable 
genetic resources at this level may in turn be lost, even if the population 
as a whole survives. In this respect, special care is needed to ensure that 
progeny of the best individuals of desirable species, when logged at maturity, 
is adequately represented in the existing regeneration, to avoid dysgenic 
effects and loss of diversity. 

At the level of the gene allelic differences could be the basis of 
valuable traits, for example in resistance to insect pests or to severe 
environmental stress, of great potential value for adaptation to changing 
environmental conditions and for future use. The rapid advances in genetic 
engineering may permit new combinations of genetic characteristics which could 
transform the possibilities for highly productive or otherwise desirable 
plantations, making best use of deforested and degraded land. The chances of 

recognising such genetic variation at that level in the natural forest are 
clearly remote. However the strong probability of such potentially valuable 
genetic resources occurring in highly diverse populations is a further reason 
to guard against inadvertent loss of diversity. 

It is therefore essential that all levels of genetic diversity be 
considered, and to the extent appropriate and practicable, included in the 
objectives and the activities of a conservation programme (Namkoong 1990). 
Moreover it is the organisation and structure of genetic diversity at the 
various levels that underlie both the functioning of the ecosystem and the 
approaches to the conservation of the genetic resources of individual species 
(Riggs 1990). 

The genetic structure of a species is defined by the way in which 
genetic variation is distributed between and within populations. This 
structure is the result of mutation, migration, selection and gene flow 
between separate populations and is strongly influenced by the genetic system, 
embracing the mating system and dispersal systems for pollen and seed. 
Information on the diversity and distribution of genes within species and 
their local populations is essential to the management and conservation of 
their genetic resources, but such information is very limited or non-existent 
for most tropical tree species. 

2.2 Ecosystem Conservation 

As stressed above, the conservation of the diversity of the natural 
forests is dependent on maintaining all essential functional components of 
the ecosystems in situ. These are likely to involve a range of ecological 
interactions, particularly symbiotic relationships and interdependent 
connections, for example between tree species and their animal pollinators, 
seed dispersers and so forth. Although the objective may in many cases be the 
conservation of particular target species and populations, in practice this 
is likely to involve conserving whole communities, at least until we have a 
more complete understanding of ecosystem dynamics. Such research as has been 
done commonly indicates hidden complexities and interactions, for example, 
among so-called "plant-web"- or "food-web" systems (Gilbert 1980; Terborgh 
1986; Whitmore 1990). A wide variety of animals, including many groups of 
invertebrates, birds, bats and other mammals, are involved in pollen and seed 
dispersal of trees of known economic importance and may in turn be dependent 
for their survival, in the areas of forest where they are needed, on sources 
of food or breeding niches provided by quite different and apparently 
insignificant trees or other plant species. The disappearance of such 
"keystone" or "pivotal" species might then lead to the loss of plant species, 
including timber trees, dependent on the animals for pollination or seed 
dispersal (Howe 1990). 

Although the great majority of species in mixed tropical forests may 
play no essential part in ecosystem function, and may therefore be in that 
sense redundant, the present level of knowledge and understanding is 
inadequate to determine with certainty all the key components of the 
ecosystem. The precautionary principle therefore requires that management and 
harvesting practices should conserve as wide a spectrum of species of as yet 
uncertain linkages and values as is practicable. Such an approach is 
consistent with the growing international concern for the conservation of 
biological diversity in general. Tropical forests and woodlands, in addition 
to the genetic resources of woody species they contain, are the habitat for 

a wealth of other plant and animal species, all of which contribute to the 
total sum of genetic diversity and resources on earth. 

We may therefore distinguish four arbitrary categories of forest genetic 
resources for conservation in situ; (i) the principal economic tree species 
(ii) other trees, plants and animal species of known but lesser value in the 
national economy or to local communities (iii) key functional species for 
sustainability of the ecosystem (iv) other elements in the total biological 
diversity, of as yet uncertain value. 

2.3 Conservation of target species 

Just as the overall objectives of forest management for productive 
purposes will give preference to the desirable trees of the principal economic 
species these species will also be a main target for in situ conservation of 
the forest's genetic resources. This will aim to maintain viable breeding 
populations, and to favour reproduction of the better individual trees, 
insofar as this may be judged from their phenotypic appearance in the forest. 
This objective should be a normal component of all sustainable forest 
management but in practice is likely to increase the cost of harvesting and 
management operations as they are currently conducted in most countries. To 
add further to the costs of management and harvesting, through greater care 
to avoid damage to seed trees and regeneration of additional "non-economic" 
species, would therefore require the imposition of special conditions, through 
incentives or constraints. Nevertheless the maintenance of viable breeding 
populations of "keystone" species and those important for domestic use by 
local communities may be essential for long-term sustainable management of the 
forest for all its values, including the production of timber. 

Some overall loss of biological diversity must be inevitable as a result 
of harvesting and management practices, at least for a period and in the areas 
affected by these operations. A single management system, if consistently 
practised, will affect all stands similarly. The timing, frequency, intensity 
and extent of logging will be the principal factors which determine the 
severity of the impact on overall biological diversity. However the use of 
diverse management systems can increase the variation between management 
units. The impact on the principal target species will depend also on the 
level of understanding of the pattern of variation within the species, both 
within and between different populations in the forests, and of its 
interactions with other species in the ecosystem, and then the willingness and 
capability to adjust management and harvesting operations to minimise losses 
in diversity. 

2.4 Conservation of Provenances 

The practical implications of the need to understand the genetic 
structure include the likelihood that genetically distinct populations of a 
species may exist in different forest areas within the species 1 range (i.e. 
different provenances of the species), as a result of isolation and/or 
adaptation to different environmental conditions. These distinct populations 
may differ substantially in theirsocio-economic value and production 
potential, or offer possibilities for improvement by combination of their 
qualities through breeding programmes. The design of in situ conservation 
strategies is therefore particularly dependent on estimating the likelihood 
and location of such intra-specific variation. 

Another important aspect of the genetic system of individual species in 
determining patterns and levels of diversity within provenances and 
populations is the extent to which pollination takes place between different 
individual trees (outbreeding) or the frequency of self-pollination, or even 
the production of seed without fertilisation (apomixis). Although very few 
species have been adequately studied many scientists who have undertaken such 
investigations have concluded that there is a strong tendency for cross- 
pollination and outbreeding among tropical tree species (Bawa 1974; Bullock 
1985; Bawa et al 1985; Bawa and Krugman 1990; Janzen and Vasquez-Yanes 1990). 
Some authorities believe that the high gene flow between individuals resulting 
from the predominance of cross-pollination contributes to the high levels of 
genetic diversity characteristic of the tropical forests. Others have 
attributed high levels of speciation in tropical rainforests to the isolation 
of the individual trees of a species, with consequent reliance on inbreeding 
or apomixis (for a discussion of these views see Whitmore 1990 and Bawa et al 

The practical implications are that without specific safeguards, based 
on adequate knowledge of the genetic systems and variation patterns of 
important tree species, the effects of logging or extensive deforestation on 
the population, through alterations in the levels and patterns of outcrossing 
or inbreeding, are likely to increase with increased severity of disruption 
of the populations. As a result there could be a lack of adequate fertile 
seed production, or excessive inbreeding, which could endanger the viability 
of the species or the provenance in the longer term. The effects might not 
necessarily be negative, since in some circumstances the removal of closely 
related individuals could encourage wider outcrossing. However the essential 
aspect is that the effects will be unpredictable in the absence of adequate 
information on the species' population biology. 

The absence of such information in respect of almost all tree species 
in most areas of tropical forest must severely limit the possibilities for 
effective management of their genetic resources. The safest practicable 
conservation strategy must be to maintain the broadest possible genetic base 
in the species as a whole by conserving as wide as possible a range of 
provenances over the natural geographical and ecological range. In doing so 
it is also desirable to protect the integrity of each such population against 
genetic pollution which can be caused by the introduction of outside 
provenances of the species, such as might occur for example in the course of 
gap planting or enrichment planting within the forest, or following the 
establishment of plantations using outside, hybridising species or provenances 
in an adjacent area within the range of natural pollen distribution. 

2.5 Values of Genetic Diversity 

The relative value of different individual trees or populations of a 
species is very difficult to judge in natural stands, where uncertainties over 
age and past history are compounded by accidental differences in site quality 
and competition with other vegetation. For the few tropical tree species 
which have been the subject of provenance trials and breeding programmes 
highly significant increases in productivity and related economic and social 
benefits have been achieved, based on selection from the very diverse 
provenances sampled. These gains have been achieved through careful 
comparative trials and the application of the results in fast-growing 
plantations. The genetic resources of other tropical tree species to be 

conserved in situ in the natural forests might provide the basis for similar 
increased productivity if subjected to trials ex situ to determine their 
heritable qualities and performance in given environments. 

2.6 Use Values and Option Values 

Economists recognise two main types of value: use values and non-use 
(existence) values. Use values may be further sub-divided into those 
available for known and immediate uses and those which might become available 
in the future (option values). While the direct use value of the genetic 
diversity in the forests can best be measured in respect of the few most 
marketable species the option value of species not currently in demand may be 
quite high. The latter could become important to adapt to possible climatic 
changes, for example, or to meet changed market demands for timber or non- 
timber forest products (NTFP) . There have been many examples in recent years 
of species earlier regarded as unmarketable or as "weed" species, but which 
are now highly valued. The effect is to increase the options open to the 
forest manager, as a result of having conserved a wider range of genetic 
diversity in the natural forests than those species previously known to be of 
immediate use (see the Ghana case study in Part II). 

2.7 Precautionary Values 

Among the important indirect (non-consumptive) use values of the genetic 
diversity of tropical forests is its possible contribution to the stability 
of the ecosystem particularly in the face of climatic changes at global or 
regional levels. Virtually nothing is known of the functional role of overall 
diversity in ecosystem stability, or of the acceptable levels of species loss 
or the thresholds of irreversible change with decreasing levels of genetic 
diversity in natural forests. Similarly it is impossible to forecast with 
certainty the response of tree species to the likely climatic changes. These 
high levels of uncertainty, combined with the dangers of irreversibility in 
the reduction of genetic diversity place emphasis on following the 
precautionary principle of avoiding unnecessary loss of such diversity and of 
genetic resources. 

The value of conserving the genetic resources of a range of populations 
of a species of established socio-economic importance may also be greater in 
the context of expected climatic changes. Particular interest attaches to the 
edge of a species 1 natural range, where the local populations may be adapted 
to more extreme environmental stresses, for example in the transition zones 
between moist and dry forest types, or between dry savanna woodland and more 
arid thorn scrub or desert formations. Moreover recent developments in 
molecular genetics and genetic engineering may give additional value to such 
populations which, although they may exhibit very slow growth and poor stem 
form or other production limitations, may contain valuable genetic material 
conferring characteristics such as resistance to drought, or to high salt 
levels in the soil. Similar considerations apply to populations of species 
in other transition zones, for example, in the inter-tidal zone of coastal 
forest formations, where the effects of sea-level rise associated with global 
warming could be severe. 


2.8 Existence Value 

The final category of value - existence value - is likely to be 
relatively high for the rarer and more precious tropical tree species most 
vulnerable to genetic impoverishment, or even to extinction, through intensive 
harvesting without adequate management, and without adequate attention to the 
conservation of their genetic resources. It is such existence values, coupled 
with concern for the existence of tropical forest ecosystems as a whole, that 
are uppermost in the minds of the public and the media in many developed 
countries, particularly in regard to tropical rain forests. 

While there may be future benefits from the conservation of a wide 
spectrum of genetic diversity, including some of possible indirect value in 
ecosystem stability, and others of uncertain option value, in respect of as 
yet unidentified market opportunities, there are likely to be immediate 
financial costs. These may be both direct costs of protection and management, 
and also indirect costs of opportunities foregone in the immediate term 
through lower levels of production from the forest as a whole. In the short 
term the stability of the forest ecosystem is unlikely to be affected by a 
reduction in species diversity. Although this could lead to the progressive 
elimination of e.g. some slower-growing cabinet or joinery timber species, 
especially if accompanied by more intensive harvesting systems that tend to 
favour fast-growing pioneer species, the overall levels of total wood 
production may also be largely unaffected by loss of species diversity. 

2.9 Location of conservation areas 

The greatest diversity of tree species is found in the lowland humid 
tropics, for example at Yanamomo, in the Peruvian Amazon, where 283 species 
of trees of 0.1 m in diameter or larger were recorded on a one hectare plot 
(Whitmore 1990). Historical and environmental influences have led to the 
concentration of species diversity in certain areas, as a result of 
evolutionary pressures, related for example to long periods of environmental 
stability or to periodic disturbance, isolation, the removal of barriers 
between populations, migration and other influences. Although the level of 
information on species numbers and distribution patterns is still very sparse 
biogeographic divisions between the world's major terrestrial ecosystems may 
be used as an initial stratification, and with other available data provide 
a basis for selecting areas of high levels of diversity, or of exceptional 
degrees of endemism, or both together, as high priorities for ecosystem 
conservation. These two criteria, together with the degree of depletion or 
of threat to the genetic resources of the area, have been used by various 
authors and organisations to single out countries and locations to be accorded 
high priority (e.g. Myers 1988; Reid and Miller 1989; McNeely 1990). Such 
areas are most appropriate for conservation within fully Protected Area 
systems, such as Nature Reserves or National Parks, rather than areas managed 
for the production of timber or other products. The criteria suggested above 
may be the most appropriate to select a single location of high priority for 
ecosystem conservation but if more than one area can be selected as part of 
an integrated system of reserves the criteria should be extended to include 
a diversity of sites to capture also intra-specific variation of target 
species. Moreover the association of adjacent areas of natural forest under 
productive management as "buffer zone" to the fully Protected Area may provide 
a valuable extension to the range and size of populations of many tree 
species, and to the efficient coverage of their intra-specific variation. 


2.10 The Link to Production Forests 

The role of managed production forests in genetic resource conservation 
is particularly important for the conservation of intra-specif ic variation at 
the population level, of tree species of known or probable value. Ideally the 
location of areas serving such conservation objectives should be determined 
from data on patterns of variation, or of genetic structure and gene flow, 
since the latter should reveal the geographic scale over which populations may 
diverge from each other. In the absence of such information, some reasonable 
assumptions can be made from geographical and ecological data. 

Differentiation between populations may develop in response to selection 
pressures resulting from local environmental conditions. There is thus 
generally a correlation between geographic/ecological factors on the one hand 
and inherent morphological or physiological properties of the local population 
of a species on the other (FAO 1989a). While the fully Protected Area system 
of Nature Reserves, for example, may include some part of the range of a 
species the effective conservation of the gene pool (i.e. the total sum of 
genetic materials) of the species as a whole requires the inclusion of a much 
wider range of populations representative of possible genetic differences 
which can only be surmised from their geographical or ecological situations 
(Frankel 1970). This is likely to require a number of conservation areas 
distributed over the entire natural range of the species and to the extent 
that such a strategy might be practicable in any given case it is likely that 
most of these would have to serve multiple objectives, including production 
of timber as well as other forest products. The desirable number and location 
of such conservation areas must be determined for each species and in the 
absence of specific data it may be assumed that for widespread and strongly 
outcrossing species a few locations in each major ecological/geographical zone 
might be adequate. For strongly inbreeding species, and those exhibiting 
scattered and isolated occurrences, more conservation sites might be needed. 

2.11 Size of Conservation Areas 

Much thought has been given, particularly in respect of populations of 
large animals, to the minimum size of a viable population that would allow 
scope for continued undirected evolution, and related to that the minimum 
conservation area needed. Other calculations have been made based on known 
or probable levels of inbreeding and the size of population needed to minimise 
consequent loss of genetic variability, over a given number of generations. 
A commonly quoted figure is a minimum of 50 breeding adult individuals for 
short-term maintenance of fitness in the population, and 500 to sustain long- 
term genetic adaptability to change (FAO 1989a). A figure of 1 000 indivi- 
duals has been suggested as desirable to maintain "evolutionary potential", 
while at the other extreme it has been estimated that a genetically effective 
size of less than 50 may be adequate for several generations and that nearly 
all the genetic variability of a population can be conserved temporarily in 
only a few breeding individuals (Wilcox 1990). 

For long-lived species such as trees and those under some degree of 
management over much of their natural range, namely those of recognised 
economic or local value, individual population size is less important than the 
distribution of conservation sites to sample likely patterns of diversity 
across the species 1 range. In the absence of information on patterns of 
variation within a species the conservation of populations of a few hundred 
individuals at the extremes of the geographical and ecological ranges is the 


most practical option. However for highly diverse rainforest containing many 
hundreds of tree species each of which may exist normally at very low 
freguency 9 conservation areas of 5 000 ha have been suggested as a "rule of 
the thumb' 1 , based on estimates that this would cover 95% of the species 
(Ashton 1984). 

2-12 Dynamic conservation 

An essential feature of in situ conservation is that it provides for 
"continuing evolution" (Frankel 1981). In this sense both the genetic 
resources themselves and the practice of their conservation are essentially 
dynamic and should not be seen as an attempt to preserve a fixed and finite 
resource. At the same time conservation implies the avoidance of the rapid 
erosion of genetic variability, for example through the extinction of species 
or unique populations, or strong directional change in the genetic composition 
of a population as a result of severe reduction in numbers, causing increased 
inbreeding and/or genetic drift, as a consequence of isolation. As the 
possibilities for setting aside substantial, additional fully Protected Areas 
for conservation are progressively reduced, the necessity for the more 
deliberate management of socio-economically important species must continue 
to increase. 

2.13 Disturbance and Succession 

Changes in species composition by the ecological succession of different 
plant communities on the same area are a common feature in any type of 
vegetation, as the colonising pioneer species and communities give way to 
later stages. Local extinctions of pioneer species are a necessary part of 
this process but provided that the natural or artificial disturbances in the 
climax formations leave opportunities for cyclical recolonisation by pioneer 
species in other areas the overall composition of the forest over large areas 
is not affected. Such natural processes in forest dynamics are generally 
accepted to occur in the most biologically diverse formations, notably in the 
tropical rain forests, where the most species-rich areas are likely to be 
those which include patches of secondary forest in various stages of recovery 
following disturbance, as well as patches of mature-phase forest (Whitmore 

The goal for genetic resource management is thus to maintain a dynamic 
system (Namkoong 1986), which may entail the deliberate removal of trees in 
the later stages of natural succession, as well as the deliberate conservation 
of some elements of the mature-phase forest. Depending on the exact systems 
of management, and the degree of understanding of the forest dynamics on which 
they are based, genetic diversity and specific genetic resources may be 
enhanced or reduced in specific areas of forest, over specific periods of 
time. The lack of active management, for example through the complete 
exclusion of human intervention, may tend to reduce genetic diversity within 
any given area, although in certain circumstances such (usually temporary) 
non-intervention may be a conscious management decision aimed at conserving 
specific genetic resources within the framework of an overall conservation 

2.14 Logging and Genetic Diversity 

Selective logging in mixed tropical forests might, in theory, be managed 
to maintain an optimal balance between the various stages of ecological 
succession to allow for maximum genetic diversity and the conservation of the 
genetic resources of both pioneer and late succession species. This might be 
achieved either by clear-cutting, at very long intervals, to allow each felled 
area in turn to revert eventually to the mature condition, or by careful 
opening of small gaps by the removal of individual trees, or various possible 
intermediate patterns and levels of harvesting that may favour the advanced 
regeneration of different species. However this presupposes not only the 
willingness to subordinate short-term financial gain to long-term ecological 
objectives but also a degree of understanding of forest composition and 

In addition to the immediate effect of logging on the advanced 
regeneration and environmental conditions, such as light, temperature and 
humidity on the forest floor, all of which may affect the regeneration of 
different species in different ways, there will be further effects on the 
density and spacing of populations of the species felled. These changes may 
influence flowering and fruiting patterns and the mating relationships within 
the population. In addition to such direct effects there may be impacts on 
the populations of pollen vectors or seed dispersers of tree species through 
the removal of "keystone" plant species on which these populations depend. 

One often inadvertent and usually severe effect of human intervention 
which may follow logging is increased susceptibility to fire. While some 
forest formations are adapted to survive periodic burning, and may be fire 
disclimax communities with particular qualities of genetic resources of 
potential value associated with their capabilities to colonise fire-prone 
areas, the adverse impact of indiscriminate fire in other more complex forest 
formations can severely reduce the genetic resources of the more valuable tree 
species. In extreme cases whole populations may be lost through fire 
following the felling of all adult trees of a species in the area concerned. 

Nevertheless with adequate control and based on a sufficient 
understanding of the ecological processes involved logging and timber 
extraction can be used to assist the conservation of a wide spectrum of 
genetic resources of the principal tree species. Both the efficiency with 
which this can be achieved, and the security against accidental loss of 
substantial elements of the gene pool, will be dependent on the management of 
a network of conservation sites in both the production forests and the fully 
Protected Area systems, extending across the natural range of the principal 



The role of logged forests In the conservation of 
species richness and genetic diversity 

Most remaining unlogged areas of forest outside the limited extent of 
Protected Area systems will be exploited for timber within the next few decades. 
The logged forests, except where excessive machine operations, fire or illegal 
cultivation have caused intensive damage, still retain much of the original plant 
diversity and are often apt for recolonisation by the major fauna (Johns 1988 
and 1992; Whitmore and Sayer 1992). The effect of the initial logging on the 
forest structure, composition and regeneration have been well documented and 
the procedures to minimize damage and improve cost-effectiveness fully 
described (e.g. Nicholson 1979; Dykstra and Heinrich 1992). Ideally harvesting 
operations in production forests aim to mimic processes of natural gap 
formation in order to provide a sustainable yield of timber without radically 
altering the composition and structure of the forest as a whole. Retention of 
unlogged areas amounting to only 5% of an intensively logged forest may be 
adequate to conserve populations of vertebrate species regarded as intolerant 
of logging (Johns 1992), and a single cycle of togging need not reduce species 
richness among the tree populations (Whitmore and Sayer 1992) provided 
adequate regrowth is present and not severely damaged during logging, or 
seed for subsequent regeneration is available in the soil seed bank or adjacent 

Major impacts on both species richness and the genetic resources of the 
prime timber species, especially those more shade-tolerant and characteristic 
of mature-phase forest, are likely to be associated with the second cycle of 
logging and subsequent harvesting. The length of the cutting cycle will be 
critically important to the maintenance of satisfactory breeding populations of 
the principal tree species. Even when small areas, such as Virgin Jungle 
Reserves may be left unlogged the progressive effects of forest fragmentation, 
and isolation, on gene flow, inbreeding rates, genetic variation, seed fertility etc. 
will decisively affect the genetic resources. There may be impacts on key 
pollinators or seed dispensers, as well as on soil structure and even soil 
chemistry (House and Morrtz 1991). 

Coordinated long-term studies are needed to guide the management of 
logged forest for the conservation of species richness and genetic resources, 
including where necessary remedial management (Ng 1983). 



Although forest management can be defined in various ways (Vanniere 
1975; Philip 1986; FAO 1989b; FAO 1992a), there is general agreement that it 
should consist essentially of taking firm decisions about the future of a 
particular area of forest, planning and implementing action to achieve those 
objectives and monitoring the results. It depends ultimately on the national 
forest policy and its basic components are the definition of objectives, 
planning, control, protection and records. It is also concerned with the 
allocation of resources to meet the defined objectives and it is commonly the 
wide disparity between the objectives and the resources provided to meet them 
that causes the greatest impacts on the forest. As used here it is taken to 
include both the controlled harvesting of timber and associated silvicultural 
operations aimed at the sustainable provision of specified goods and services 
from the forest. An essential aspect is that the land should remain in forest 
use after harvesting and that the operations should be conducive to adequate 
regeneration and to the maintenance of the environmental and social functions 
of the natural ecosystem. This implies involvement and prior consent by all 
land users, control and continuity of purpose, which are also essential 
elements for the conservation of the forest's genetic resources. 

3. 1 Continuity and Control 

Genetic resource management can only be effective if it is an integral 
part of land use management as a whole (FAO 1989a). The objectives of in situ 
conservation may involve various approaches to land use, including multiple 
use production areas and timber production forests as well as fully Protected 
Area systems, in the attempt to reconcile the dual requirements of present day 
demands for revenue and basic human needs (food, wood and other forest 
products), with long-term conservation objectives. Ideally this requires a 
comprehensive national land-use policy and plan, embracing an appropriate 
forest policy and based on a national inventory of the forests, including 
specific attention to plant species of socio-economic importance and 
conservation concern. Failing that a broad zonation of vegetation, following 
the principles of the Biosphere Reserve Programme (Unesco 1984), may be used 
to focus the conservation objectives to the most important areas, which may 
then be given special reserved status, within fully-protected categories of 
land. In reality the pressures of increasing human populations and related 
economic development programmes on the land and natural resources severely 
restrict the areas likely to be set aside and retained as National Parks or 
equivalent, fully protected reserves. The potential contribution of 
production forests to the conservation of genetic resources is therefore 
important and may be high, given a conservative and sustained approach to 
forest management, based on natural regeneration systems. The deliberate 
inclusion of genetic conservation objectives in the management plans of 
production forests may be essential, to secure the necessary range and 
diversity of sites for an efficient national conservation network. 

The initiation of tropical forest management, first in the Indian sub- 
continent, and subsequently in Africa (FAO 1989b; FAO 1989c) included the 
establishment of firm administrative control over the forests, for example 
with the designation and demarcation of a "Permanent Forest Estate 11 whose 
boundaries and use could be changed only by decision at the highest levels of 


national authority. Although this authoritarian approach has sometimes 
provoked local opposition, and there is now general acceptance that the 
security of the forest cannot be assured without the consent and involvement 
of local people dependent on the resource, the establishment of the Permanent 
Forest Estate has been a powerful element in the conservation of genetic 
resources. Although in many cases the objectives of sustainable timber 
production have not been sufficient to ensure the retention of all of the 
reserved forest estate from reallocation to permanent agriculture, or from 
illegal encroachment, in most tropical countries legal protection for 
productive forests has had an important and positive impact on the 
conservation of their genetic resources. However there have also been severe 
negative impacts from the often excessive exploitation (unaccompanied by 
adequate control and management to ensure regeneration) resulting from short- 
term political, financial and economic pressures. 

3.2 Economic and market influences 

Most recent reviews of tropical forest management (Masson 1983; Mergen 
and Vincent 1987; Schmidt 1987; Wyatt-Smith 1987a; FAO 1989b; FAO 1989c;FAO 
1992a; Poore 1989) conclude that management of the tropical forest as a 
renewable and truly sustainable resource is technically possible and that the 
past failures or abandonment of attempts at the sustainable production of 
timber in tropical forests have been due to socio-economic or political 
pressures. These have frequently imposed severe constraints on the resources 
available for the various aspects of management referred to above, resulting 
in part from the failure to recover adequate revenue from the exploitation of 
the resource, and/or failure to reinvest such revenue in the regeneration and 
management of the forest (Repetto and Gillis 1988). 

At the root of the problem is the weakness of the national economy in 
most tropical countries combined with the scarcity of capital to invest in 
development. The forest itself has been treated as a source of finance to 
support development in other sectors of the economy and while in total this 
source of funding has been sufficiently large to make exploitation worthwhile, 
the apparent returns per unit area of forest are commonly too low to secure 
the necessary level of investment for sustainable management, or even to 
retain the land against pressures for alternative land use. Even when the 
arguments for the long term economic benefits from reinvestment in forest 
management may be accepted urgent short-term financial constraints, coupled 
with short-term interests of concessionaires, logging enterprises and other 
concerns, may lead to excessive exploitation without regard to forest 
regeneration. This is encouraged by short-term concession agreements which 
provide no motive for longer-term planning. In order to recover the investment 
in large-scale machinery and road construction the logging companies 
frequently seek maximum returns from harvesting at minimum costs, and with 
minimum concern for environmental impacts. Where market demand is highly 
selective the exclusive concentration on the extraction of the best phenotypes 
of the most valuable species will have negative (dysgenic) effects on 
subsequent generations, if such interventions are carried out without due 
consideration to regeneration potential and the quality of the next generation 

Where a broader range of species, including many lesser-known or lesser- 
used species, are marketable there may be potential for greater revenue, 
providing the opportunity for increased reinvestment in forest management. 
In practice, however, although the more intensive felling has tended to 


increase profits, reinvestment has remained static or even been reduced; the 
forest has in these cases been left to recover naturally without regard to 
species composition. If logging damage to the advance regeneration and to the 
site conditions for seedling establishment were closely controlled the 
harvesting of a wide range of timbers might be more compatible with the 
conservation of a similarly broad range of species, than the very selective 
logging characteristic of earlier exploitation regimes (see for example the 
Ghana case study Part II). However without strict controls over road 
construction, logging plans, timber marking, harvesting and extraction there 
must be a danger of severe ecological impacts on the site capability, and on 
the genetic resources especially of the slower-growing species characteristic 
of the mature-phase "climax" forest. 

The introduction of heavy mechanical equipment for timber exploitation, 
together with increased demands for a wider range of tropical timber species, 
has had an overwhelming impact on the species and genetic diversity of some 
natural forests. The effect is to shift the composition of the forests in the 
felled areas towards relatively few predominantly fast-growing pioneer species 
characteristic of the earlier stages of the ecological succession. Management 
for the in situ conservation of genetic resources must give priority to the 
principal economic species, some of which at least will be fast-growing 
pioneers or gap-opportunist species, which will be favoured by a certain 
degree of canopy opening. However the effect of large-scale and extreme 
reduction of the forest canopy which is associated with heavy mechanical 
logging of a wide range of species is most likely to favour very short-lived 
pioneers with timber low in density and durability. It is certain to 
discriminate, at least in the short term, against the multitude of slower- 
growing shade-tolerant species, including some high quality cabinet woods and 
veneer timbers. The overall impact of these interventions is therefore to 
reduce the range of species of known economic value, uniformly in favour of 
a narrower band of fast-growing, low to medium density species. At the same 
time there is likely to be a reduction in the biological diversity of the 
ecosystem as a whole, both flora and fauna, unless certain areas of the forest 
are excluded from simultaneous, intensive logging regime. Economic pressures 
to maximise the returns from the investment in logging equipment, roads and 
related infrastructure have frequently overridden such ecological 
considerations . 

Volume yields in naturally regenerated tropical moist forest are likely 
to be on average only 2 to 3 m 3 per hectare annually, although in some cases 
silvicultural operations and rational management might increase this by a 
factor of three or four times, where there is ample regeneration of marketable 
species and where site conditions are favourable (Wyatt-Smith 1987a). A 
common feature of management systems aiming to increase the yield of valuable 
species in this way is the elimination, usually by poisoning, of unwanted and 
competing trees. This and other "refining" operations, if maintained 
consistently through successive cutting cycles, will also reduce the diversity 
of species in the areas treated but need not do so over the forest as a whole, 
if selected areas were excluded from such treatments or were treated to favour 
different components. In most existing operations the loss of yield involved 
in such exclusion would be regarded as unacceptable on purely economic 
grounds . 

The impact of the international trade in tropical timber on the genetic 
resources of particular genera and species has been assessed in a recent study 
undertaken on behalf of the International Tropical Timber Organisation (ITTO) 


by the World Conservation Monitoring Centre (WCMC 1991). This study compiled 
data and expert observations on 1 868 tree species, and assigned them to 
conservation categories in accordance with the IUCN system. Of these 304 
species were classified as threatened at a global level and 190 as threatened 
in two or more countries. The reliability of the data may be questionable 
since there is little available information based on detailed forest inventory 
and the assessments are therefore based largely on the opinions of specialists 
in the various geographical and botanical fields. Particular attention was 
given to the family Dipterocarpaceae, for which the study claimed to have 
achieved a relatively complete assessment of the conservation status. Even 
allowing for some uncertainty and possible exaggeration of the degree of 
depletion or threat the study revealed a serious impact on the genetic 
resources of this important family of high quality timbers. The very 
intensive logging now being carried out in several of the countries in which 
species of this genus occur naturally, must give cause for concern unless 
urgent attention is given to their sustainable management within which 
harvesting is only one step; and to genetic conservation measures, carried out 
in parallel with their use. 

The ITTO study covered countries in Africa and Asia in 1991/92. In 
Latin America another aspect of the market impact on forest genetic resources 
has been identified where the demand for mahogany (Swietenia macrophylla) has 
led to highly selective harvesting of good phenotypes of the species over 
large areas without provision for subsequent protection of the regeneration 
and management to ensure its diversity and quality (Monbiot 1991). The danger 
of severe dysgenic effects and extinction of local populations is clearly a 
possible consequence of the market forces in this case. 

In general the effect of economic and market forces has been to impose 
the reduction of species and genetic diversity in the timber-producing 
forests. This has been the result of very narrow, selective markets, and 
failure to re-invest part of the revenue in forest management, thereby 
enforcing low-cost "blanket" management systems with short-term objectives. 
The inadequacies of economic theory and analysis relating to natural 
management systems in tropical mixed forests (Leslie 1987) relate particularly 
to the need to widen the scope of the analysis beyond the short-term benefits 
to the agencies and individuals most directly involved, to include the broader 
interests of the nation and society as a whole, and of future generations. 
This applies particularly to the values attached to the genetic resources of 
the forests. To the extent that the timber production areas can also be 
managed to conserve genetic resources this additional objective should 
reinforce the economic case for natural forest management to be governed by 
ecological as well as economic considerations. 

Even the much narrower range of overall genetic diversity that will 
result from repeated extensive harvesting and refinements of the forests is 
likely to be much greater than if the same area were converted into forest 
plantations, and certainly substantially greater than that which would result 
from most alternative forms of land-use. However, many species and 
populations, especially those characteristic of the primary or mature-phase 
forests, will be lost unless the current severe constraints on the funding of 
natural forest management are reduced. Increased funding (including the re- 
channelling of revenue back into the forest) would allow more deliberate and 
diverse management of the tree populations, possibly at the expense of some 
loss in harvested volume. This loss might be partly offset by an increase in 
unit value from the management of selected areas on longer rotations for 
higher quality timber. 


Whereas rethinking of the economic and financial constraints on natural 
forest management seemed unrealistic earlier, and may still be so for most of 
the production forests, the economic value of genetic diversity is now 
increasingly recognised as of international concern. That value, and 
therefore the costs that might be borne for conservation, will be dependent 
on the location, composition and existing condition of each specific area of 

3.3 Forest Inventory 

The scientific basis for the conservation of species and of their 
genetic resources depends essentially on the study and interpretation of 
taxonomic information on genetically determined differences and affinities, 
their patterns of natural distribution (chorology) and the ecological basis 
for their occurrence. These three interdependent sets of data which should 
form the basis for drawing up sound conservation strategies, are inadequate 
in the tropics and in many cases non-existent. Very often the only data 
available are the result of forest inventories concerned primarily or 
exclusively with stocks of harvestable timber. The techniques for efficient 
surveys of the standing timber resources in the forest have shown considerable 
progress in the past 30 to 40 years (FAO 1989b) and there has been increasing 
emphasis on the need for National Forest Inventories and for detailed and more 
comprehensive inventories of selected areas. However, all too often such 
inventories have provided little of the information needed to plan the long 
term, sustainable management of the forest, but have been confined to the 
determination of the marketable volume of a limited number of so-called 
economic or potentially economic species (Masson 1983). Little regard has 
been paid up to now to the determination of the actual composition of the 
forest or its condition after logging. 

Inadequate information on the species composition of the forest is one 
of the main problems confronting natural forest management, both in terms of 
the forest's full economic value and its regeneration potential (Wyatt-Smith 
1987b). Some of the important questions to be answered in this regard are the 
adequacy of existing levels of seedlings, saplings and advanced growth of the 
marketable species as the basis for the future crop, both before and more 
importantly after logging. Assessment of regeneration was an integral 
component of the earliest attempts at natural forest management, for example 
in Malaysia and Nigeria, but latterly the tendency has been to reduce rather 
than to increase the time and manpower resources devoted to such surveys, and 
thereby to limit the level of detailed information available even in respect 
of the preferred economic species. However in recent years, with the 
development of conservation biology as an applied science, it has become 
increasingly clear that more detailed studies and surveys must be an 
inseparable component of forest management systems to produce an adequate 
scientific basis for conservation action. 

Diagnostic sampling to determine the stocking and silvicultural 
conditions of young stems of "desirable" species below the exploitable girth 
limit is a common practice in the more deliberate attempts at natural forest 
management (FAO 1989b) and there is a tendency for inventory and resource 
assessment in advance of logging to be taken further, to extend beyond 
commercial timber resources to non-timber products, including those of 
interest to local communities (FAO 1989c). Such action is essential to obtain 
a more complete valuation of the forest in support of the conservation of a 
wider spectrum of valuable genetic resources (see also the Ghana case study) . 


The central and traditional component of forest inventory, namely the 
accurate assessment of the allowable cut and sustainable yield of timber, is 
vital to conservation of the resource, to ensure that the rate of harvest 
should not exceed the regenerative and growth capacity of the forest. This 
requires an accurate knowledge of the growing stock, its distribution by 
species, size classes and location, and an understanding of how these change 
with logging and silvicultural treatments. Availability of accurate forest 
inventory and stock maps in reducing logging damage and protecting the 
regeneration is also essential in the conservation of genetic resources. 
However such inventory data are insufficient as a basis for more positive 
action to identify and conserve the most valuable components of genetic 
diversity. Given the high proportion of the cost of inventory operations that 
relates to the actual access to and work in the forest the additional cost of 
collecting a wider range of data and observations in the course of the timber 
survey can be relatively low. Moreover the widespread availability of 
powerful electronic computing facilities for the handling of large and complex 
sets of data has largely removed that constraint on the collection of an 
increased range and volume of field data, in the course of forest inventory. 

In general 10% of the tree species in a tropical forest comprise at 
least 50% of the stand (Ashton 1988). Rapid extensive surveys may give 
guidance to conservation priorities not only in terms of species richness but 
also in respect of the genetic resources of a forest. Recent studies by 
Hawthorn in Ghana indicate the potential value of broad botanical surveys in 
association with forest inventory in developing strategies for the 
conservation of forest genetic resources (see also the Ghana case study) . 

For the forest inventory to serve its full purpose in assisting the 
conservation of forest genetic resources it must attempt to assess the 
relative genetic value of a given area of production forest, for example in 
regard to the range of distribution of selected species or forest types, in 
relation to other managed or reserved areas, including the fully Protected 
Area system of National Parks etc. This information could help determine the 
most efficient combination of the minimum number of sites needed to cover 
species, populations and communities at the minimum level essential for 
conservation of the desirable range of diversity. Given the very large 
numbers even of tree species in the tropical forests, and the need to include 
surveys of some other socio-economically important plant species, or of 
species essential to the functioning of the ecosystem, and therefore to the 
overall management of the resource, it is often necessary to adopt a 
compromise between detailed biological surveys and more general assessments, 
based on variations in landscape or other environmental features. It may be 
assumed, for example, that much intra-specific (provenance) variation, of 
potential economic significance, follows patterns of variation in the 
environment and in the plant community as a whole. 

A key to efficient and cost-effective inventory is at the planning 
stage, to ensure inclusion of an appropriate range of botanical, ecological 
and sociological expertise, for example, both in the design and in the 
execution of the survey. Given the range of expertise and strength of 
interest in many universities and research institutes in both tropical and 
industrialised (often aid donor) countries in such scientific investigations 
in tropical forests, the additional human resources needed may often be 
available at comparatively little extra cost, compared with the expenditure 
on the basic inventory, and in relation to the value of the additional 
information to be derived from such expert involvement. 


A particular aspect of this planning phase which may require expert 
involvement is the planning of data capture, handling and analysis. The 
widespread availability of small but powerful computing capabilities for data 
analysis has also transformed the possibilities for obtaining an understanding 
of forest composition and genetic diversity from limited data. Approaches to 
the development of inventory procedures and growth models for the management 
of tropical forests are advancing rapidly (e.g. Vanclay 1989; Alder 1990) and 
will progressively gain the capacity to incorporate broader information 
relating to the management of the genetic resources of the forest. Simulation 
of the variability and complexities of population distributions is already 
possible using some stochastic models such as those designed to measure the 
average number of occurrences in a given area, and to reveal patterns of their 
variability (Jeffers 1982). This information can be an important aid to 
selecting locations for in situ conservation. 

Sustainability as an objective of forest management demands that forest 
inventories be planned to look far beyond the assessment of marketable volume 
to the establishment of baseline data for the continuous monitoring of forest 
condition, and the implications for the conservation of genetic resources. 
Among these are the retention of sufficient individuals of "keystone" species 
to maintain their own breeding populations and therefore the long-term 
contribution of that species to the functioning of the natural forest 
ecosystem. This requires the identification and recording of such species in 
pre-logging inventories, so that sufficient numbers, with an appropriate 
distribution in the forest, are marked for retention, based on a sufficient 
understanding of the forest dynamics. 

3.4 Forest Dynamics 


The aspect of forest dynamics which has attracted most attention in 
relation to timber exploitation is the growth and yield of the principal 
marketable species. The lack of reliable annual growth rings in the timber 
of most tropical tree species, and therefore of convenient "short cuts" to the 
determination of increment, together with the complexity of the growing 
conditions and species composition in the forests, has made predictions of 
yield difficult (e.g. Kemp and Lowe 1970). Despite the progressive 
development of systems of Permanent Sample Plots (continuous forest inventory 
plots) in representative samples of forest in a number of countries the basis 
for calculations of growth and yield is still generally weak. Probably the 
best level of prediction has been achieved in Queensland, Australia (Vanclay 
1989). However even in those intensively studied forests there is very little 
if any reliable information on the relative importance of genetic and 
environmental factors in determining growth rates of individual trees in the 
natural stands. 

The influence of competition from other vegetation on the relatively 
small proportion of "desirable" stems has been the basis of management 
interventions to effect improvement thinnings to favour the growth of the 
potential final crop trees (Hutchinson 1987; Maitre 1991; FAO 1989c). The 
first step in this treatment, as applied in Sarawak (Hutchinson 1987) is to 
group the species into "wood quality" categories, to arrive at lists of 
desirable species, and then to assess and attempt to predict their response 
to different levels of release from overhead shade and competition resulting 
from commercial logging or subsequent thinning of the stand. For this purpose 
the species are allocated to ecological groups according to their degree of 
shade tolerance and/or capability to increase growth rapidly in response to 


Classical forest management started from attempts to understand and 
utilise such ecological processes and interactions f and particularly the 
natural cyclical changes in the progression from gap colonisation by pioneer 
species to the mature or late successional forest condition. Arbitrary 
divisions into gap-phase, building-phase and mature-phase forest development 
are now commonly recognised (Whitmore 1990). The concept of "forest gap-phase 
dynamics" appears to apply as a general model, possibly with local variants, 
in a wide range of forests in all tropical regions and although it is an 
oversimplification to recognise only "pioneer 11 or "climax" species the 
definition of two broad groups or "guilds" based on their behaviour in 
response to gap creation has proved useful in approaches to natural forest 
management (Whitmore 1991). It is also important in approaches to in situ 
conservation of forest genetic resources, particularly those of the principal 
economic tree species. 

In all tropical rainforest floras there are fewer pioneer than climax 
species (Whitmore 1990) and the pioneer genera are less likely to include 
narrowly endemic member species of very restricted geographic range. Their 
efficient systems of seed dispersal are probably responsible for their 
generally wide range of distribution and reduce the possibilities for the 
development of localised genetically distinct populations. 

The complex interrelationships in pollination, seed dispersal, food web 
and plant web systems described earlier are important both for the attempted 
regeneration and management of timber species, particularly those in the 
"climax" guilds, and for in situ conservation of forest genetic resources. 
Without adequate understanding of the ecological processes and relationships 
the attempts at natural forest management must rely on "blanket" application 
of silvicultural treatments with unpredictable and possibly perverse results. 
However it has latterly proved impossible to justify the necessary expenditure 
on detailed ecological studies in terms of the increased yield of timber alone 
over the period of one or two cutting cycles or short rotations. Nevertheless 
recent reviews of management methods have emphasised the inadequacy of 
"blanket" canopy treatments and the necessity for improved knowledge of the 
autecology of individual species, and not only those with marketable timbers 
(FAO 1989b). 

The field of reproductive biology, embracing studies of pollination, 
seed dispersal and predation, and the dynamics of regeneration banks of seed, 
seedlings and saplings, which are clearly important in genetic resource 
conservation, have also been identified as necessary to the forest manager 
(Palmer 1989). The case study of Cordia alliodora in Appendix 1 shows the 
level and intensity of research needed to provide reliable information for a 
single species. The number of species requiring such study even in a single 
forest, is very large, and the resources of skilled manpower are very limited. 

Seed dispersal biology is directly relevant to natural forest 
management, particularly in respect of the mature phase species, many of which 
have high value timbers, and are characterised by the production of large 
seeds. There is a growing body of evidence that the frequency and density of 
such species in the natural forests is limited by seed predation and that 
chances of survival are improved by dispersal away from the vicinity of the 
parent tree (Terborgh 1990). A good example is the dependence of Virola 
surinamensis on distribution by toucans and other large frugivorous birds, 
without which it would be unable to recruit seedlings, and the species would 
face local extinction (Howe 1990). An understanding of the identity and 


behaviour of animal dispersers, and their possible dependence on other tree 
species for food or nest sites, is an area of common concern to both forest 
managers and forest geneticists concerned with conservation. 

3.5 Regeneration 

The regeneration potential of the desirable species is of key importance 
in the management of the productive forest estate and clearly fundamental also 
to the conservation of their genetic resources. For in situ conservation 
natural regeneration is the clearly preferable strategy, although some aspects 
of artificial regeneration, such as enrichment planting using seed or 
seedlings randomly obtained from the same natural stand, may sometimes be 
acceptable and desirable. The encouragement of natural regeneration is likely 
to be the cheapest option for timber production objectives also, provided that 
it can be easily and confidently obtained. In practice, however, it has 
proved to be one of the most difficult and uncertain aspects of management in 
tropical forests, despite having been the subject of much field study and 
experimentation for over a century. 

The problem is not peculiar to the tropical forests, but is evident from 
the large areas of previously forested and now barren land in the 
Mediterranean and some temperate regions. Grazing and browsing by domestic 
livestock have had a major influence in such areas, and in the more densely 
populated tropical forest regions, for example in India, even in tropical 
forests of high production potential, such as the Sal (Shorea robusta) and 
evergreen forests (FAO 1989c) despite the long history of research and study 
by trained staff. Similar failures have been common elsewhere in Asia, except 
in some of the richer dipterocarp forests, where as a result of the range of 
desirable species whose silvicultural characteristics are favourable to 
promotion by controlled canopy opening, satisfactory, although by no means 
predictable, natural regeneration has been more often achieved. Failure to 
ensure adequate regeneration of the selected economic timber species proved 
a fundamental problem in many African forests (Nwoboshi 1987; Kio and 
Ekwebelan 1987; FAO 1989b) and was a principal reason for the development of 
forest plantations in African countries, as an alternative to natural forest 
management . 

Among the most important aspects of regeneration behaviour are the 
frequency of seed years, the length of seed viability, seedling survival, the 
pattern of distribution and abundance of seedling regeneration on the forest 
floor, the tolerance of shade and response to light and the competitive 
potential of the preferred species under different degrees of canopy opening. 
The complexity of the interactions in the natural forest is such that the 
provision of a set of environmental conditions guaranteed to favour the 
regeneration of a few selected species is virtually impossible, without 
detailed study of the actual state of each limited area of more or less 
uniform conditions within the diverse mosaic of the forest, and of the 
autecology of all the principal species. In practice diagnostic sampling 
often by means of systematic parallel transects through the forest, is used 
to determine the overall adequacy of stocking of young stems of desirable 
species, which is found to survive undamaged after logging is complete, as the 
basis for the next crop. A diversity of sites and silvicultural systems will 
be likely to create increased diversity, thus being an advantage from the 
genetic resources point of view. 


A shift in approach to the management of the regeneration stage, from 
earlier attempts to control the composition and development of the seedling 
population to the acceptance of the largely accidental composition actually 
obtained, is reflected in the procedures for regeneration sampling. For 
example in the earlier development of the Malayan Uniform System (MUS) , which 
was among the first and most successful attempts at management in the tropical 
moist forests, a three-stage linear sampling regime was used to determine 
different stages of regeneration development before logging was undertaken. 
This procedure was later modified by merging the three stages into a single 
linear regeneration sampling operation after logging was completed (Wyatt- 
Smith 1987b). The change represented the acceptance that economic imperatives 
rather than silvicultural considerations, are decisive in determining 
management practices. The MUS as originally designed made provision for 
situations where the level of seedling regeneration was found to be 
inadequate, by proposing deferment of logging until adequate regeneration was 
obtained. However in practice the inadequacy of regeneration stocking before 
felling was not allowed to hinder the progress of exploitation (Ismail 1966). 

In the Malaysian dipterocarp forests the effective fruiting years are 
the massive long-interval ones (Appanah and Salleh 1991), and it is preferable 
to log following a mass fruiting year. Silvicultural operations to kill 
unwanted "weed" species and to liberate juveniles of desirable species can be 
increased following abundant fruiting, for maximum benefit in the improvement 
of future timber production. The abundant regeneration which may be achieved 
by deferment of logging until after a heavy fruiting year offers maximum 
possibilities for effective silvicultural operations to improve the 
composition of the final crop; it will also favour conservation of the genetic 
resources of the species favoured for commercial utilization. It seems 
probable that the small amount of seed of given species produced between mass 
seed years may be provided by a small and possibly distinct segment of the 
population; as the regeneration resulting from such seed will not be 
representative of the population, its value for conservation will be limited, 
unless complemented by seedlots/regeneration over a range of years.. 

The importance of leaving se6d trees of good phenotypic quality at the 
time of logging, particularly if regeneration sampling has revealed low levels 
of established seedlings and advance growth of the desirable species, is a 
further instance of close coincidence of interest between the objectives of 
production and those of genetic resource conservation. In practice, however, 
this vital aspect has been commonly overlooked or overridden by the pressures 
for maximum harvest yield and profit. The retention of a number of large seed 
bearers after the main logging operation does present some disadvantages in 
subsequent management of the stand if they are so numerous as to cause 
depressive shade or competition (Catinot 1986), or if they are subsequently 
harvested, with related extraction damage to the regenerating forest. However 
the effective loss in production is slight in comparison with the dangers of 
progressive deterioration in the genetic quality of the population through 
effective reliance for regeneration on residual, probably less vigorous and 
less desirable phenotypes to complement soil seedbanks and existing seedling 
regeneration, if these latter ones are inadequate. Despite the practical 
difficulties and dangers of attempting to select genetically "superior" 
individual trees in natural tropical forests the real dangers of dysgenic 
effects from the selective, systematic removal from the breeding population 
of the best and most vigorous phenotypes should not be ignored. More 
information based on solid research on this aspect is urgently needed. 


Reliance on natural regeneration systems in production forests 
undoubtedly offers important opportunities for in situ conservation. The 
regrowth that follows heavy logging will favour pioneer (gap-phase) species 
and large-scale clear cutting typically leads to the complete dominance of 
soft-wooded, low density timber trees (Jordan 1986). As primary forests are 
progressively cleared or exploited the populations of many prime hardwood 
species characteristic of climax, mature-phase forests must decline unless 
specific measures are taken to maintain them. The role of the seedling 
populations in the natural regeneration of such species is particularly 
important. For example the Asian dipterocarps generally require at least a 
small gap in the canopy to develop to adult size but the chance of seeds 
naturally being placed in such a gap is low due to poor seed dispersal and 
infrequent seed years (Ashton 1982). This, combined with the lack of seed 
dormancy (Ng 1980) gives exceptional significance to the ability of 
dipterocarp seedlings to survive under the low light intensity on the forest 
floor until an adequate gap in the canopy occurs. The loss of the seedling 
population before such an opportunity occurs can have profound influence on 
the future species composition of the forest. The same is true for other 
species groups of economic importance in some African and neotropical forests 
(Whitmore 1991). An understanding of seed dispersal biology, and seed and 
seedling physiology, at least in so far as they are expressed in the incidence 
and behaviour of seedlings in the forest in relation to light conditions, must 
be the necessary basis for the conscious manipulation of species composition 
for both production and genetic resource conservation objectives. 

Pioneer species typically have seeds that can withstand desiccation and 
may become dormant for long periods in the soil. Wherever a soil seed bank 
has been looked for in lowland tropical rain forest one has been found 
(Whitmore 1990). Some pioneers have small, copious and easily dispersed seed 
which allows for frequent replenishment of the seed bank, or for the rapid 
colonisation of canopy gaps soon after they occur. Generally, therefore, the 
need for management interventions to favour the regeneration and conservation 
of the genetic resources of pioneer species is adequately met by normal 
logging and silvicultural treatments. 

3-6 Silviculture 

Success in silviculture aimed at the production of timber, has been 
defined as the achievement of "a stand of fine-timber trees brought to 
maturity and producing natural regeneration on a site where it has matured 
before, and where the soil shows no sign of deterioration 11 (Dawkins 1988). 
On this definition silviculture in the originally closed tropical forest has 
been more successful over the past century than has been commonly believed, 
not only in Myanmar (Burma), India and Malaysia, but to a limited extent in 
some African and neotropical forests. However in terms of impacts on the 
genetic resources, and especially on the total range of biological diversity 
of plants and animals in the forests, silvicultural operations, particularly 
the use of arbor icides, can have far more persistent and discriminating 
influence than crown manipulation or logging. Insofar as the treatments 
achieve predictable increases in the regeneration, growth and representation 
in the final crop of the principal economic species, they are likely also to 
favour the conservation of their genetic resources. However the treatments 
have often proved to be unpredictable even in respect of the species they were 
intended to favour, and in many cases involved the attempted elimination of 
species which have subsequently proved valuable both for their role in the 
functioning of the forest and for their acceptability in the international 
timber markets. Conversely the success of the silvicultural treatments has 


sometimes been due to their accidental encouragement of the regeneration of 
species which were not at the time considered desirable, but which are now in 
substantial international demand. 

Silvicultural systems in natural tropical forests may be broadly divided 
into two main groups. Monocyclic systems, also known as shelterwood systems, 
aim at a single comprehensive harvest of all marketable timber at the end of 
the rotation, with reliance on seedling regeneration to form the next crop. 
Probably the best developed and best known example is the Malayan Uniform 
System (MUS) under which, as originally designed, the initial logging was 
followed by poison-girdling of virtually all the remaining trees, down to a 
specified minimum girth at breast height, by which means the canopy became 
progressively more open, and conducive to satisfactory growth of the generally 
abundant regeneration of desirable species, principally dipterocarps. Various 
attempts at shelterwood systems have been made in all three tropical regions 
(FAO 1989b; FAO 1989c; FAO 1992a; Schmidt 1991) but problems were commonly 
encountered with severe climber infestations and failure to induce adequate 
regeneration of the principal economic species. The increased demand for a 
wider range of marketable species has made the failure to induce regeneration 
of the few chosen species less critical and the use of heavy mechanical 
harvesting and extraction equipment, coupled with the increased market demand, 
has increased the degree of canopy opening by logging alone, making the need 
for poisoning of residual trees less necessary. The overall ecological effect 
of such monocyclic systems is to favour "desirable" pioneer or near-pioneer 
species, including some with light, pale, general purpose, marketable timber. 
There are likely to be adverse impacts on the breeding populations and genetic 
resources of the slower-growing heavier hardwood species characteristic of 
climax forest not favoured by this silvicultural system and the shorter the 
rotation (or the more restrictive the selection of "desirables"), the more 
severe the impact will be over time. 

Polycyclic systems involve the selective removal of a limited number of 
stems on two or more occasions over the full rotation cycle, thereby 
maintaining a less uniform stand of mixed ages, with reliance on advanced 
regeneration for the next harvest. Such systems are theoretically capable of 
incorporating mature-phase climax species at the expense of accepting rather 
lower volume growth rates, but possibly higher value increments and therefore 
of conserving a broader spectrum of genetic resources in terms of species and 
timber qualities. However there may be some danger of dysgenic effects within 
populations of individual species if the selective felling removes the 
fastest-growing and most desirable individuals, leaving less vigorous and 
possibly defective stems to regenerate, in the absence of adequate, existing 
seedling regeneration and/or soil seed banks. Moreover if the desirable 
species are a small minority of the larger trees in the forest it may be 
necessary to undertake (expensive) operations to favour the immature trees of 
the valuable species, to avoid progressive impoverishment of the stand. 
However, potentially deleterious influences of selection on genetic quality 
of the species or the stand in this silvicultural system can be avoided by 
responsible management and harvesting practices, as were practised for example 
in Queensland, Australia. Under the Queensland Selection System the 
deliberate selection of trees to be retained in the forest, and the 
enforcement of strict logging controls to avoid damage to these selected 
trees, was designed to guard against possible dysgenic effects of selective 
logging. Effective operation of selection systems requires skilled and 
frequent tending of the desirable components of the forests and, especially, 
skilled and responsible logging practices, which help preserve existing 
advanced regeneration from accidental damage. 


It has been suggested that for the first 30 to 40 years after the 
initiation of proposed systems of management the distinction between 
mono eye lie and polycyclic systems lies more in the future intentions and 
expectations than in real and irrevocable differences in practices and in the 
structure of the forest (FAQ 1989b). The essential question in regard to the 
conservation of genetic resources is the extent to which the harvesting 
practices and silvicultural systems allow for the retention of a wide spectrum 
of potentially valuable genetic diversity. This is most likely to be achieved 
if different forests, and different sections within the same production 
forest, are subjected to different systems, based on ecological principles to 
favour the regeneration and bringing to maturity of different elements of the 
main "guilds", including the climax species. Such management systems would 
increase the complexity and cost of harvesting and marketing, and might 
therefore be judged as uneconomic on recent thinking, which has tended to 
favour very low cost and minimum intervention approaches, since the apparently 
low level of financial returns per unit area of natural forest has been 
considered unable to bear the cost of intensive skilled management (see Mergen 
and Vincent 1987). This implies a management strategy restricted by the 
forced acceptance of the fortuitous composition of the residual growing stock 
after logging, with any subsequent tending limited to "blanket" operations 
applied without discrimination across large areas of initially diverse forest. 
If accompanied by the search for additional uses and market outlets for the 
range of timbers actually produced, the resulting balance between revenue and 
expenditure may appear favourable. However in terms of the impact on the 
genetic resources such "blanket" operations are likely to cause a progressive 
loss of over-all diversity, particularly through the impacts on the breeding 
populations of the slower-growing species characteristic of mature-phase 

However, genetic losses are not inevitable since the composition of 
seedling regeneration and advance growth left after a single logging 
operation, provided that it does not remove too high a proportion of the 
growing stock (e.g. 20 to 30 m 3 per hectare on average), is likely to contain 
representatives of all species and guilds. These will probably be adequate 
to permit the restoration of genetic diversity and the deliberate 
encouragement of selected elements through subsequent tending, especially if 
different sets of elements are favoured among the management units. The 
practice of "liberation thinning" (Hutchinson 1987; FAO 1989c; Maitre 1991) 
is an example of the deliberate selection and promotion of individual stems 
to form the final crop, based on the residual advance growth after logging. 
It allows for the encouragement of a range of desirable species, presently 
according to lists based on timber qualities, and in accordance with 
assessment of the ecological requirements of selected species in terms of 
their likely response to treatments aimed to manipulate the overhead canopy 
and competition. Once the release of the selected "leading desirables" to 
form the final crop has been assured this system can permit the retention of 
a wide range of other species and therefore, if applied over large areas of 
diverse forest, can be consistent with the conservation of a broad range of 
species and genetic resources (Hutchinson 1991). Depending on the criteria 
used to select the "leading desirables" the system could be used to achieve 
ecological and conservation objectives, at the expense of some reduction in 
yield of the faster-growing pioneer component of the crop in some areas of the 

Ng (1983) has drawn attention to the increasing need for "remedial 
management" in Malaysia in order to attempt to restore and to maintain the 
structure and composition of areas of mature forest. While current trends to 


maximise production from the timber-producing areas are not conducive to the 
adoption of less intensive harvesting, the growing strength of both the 
national economy and the conservation consciousness in Malaysia may permit 
such ecological considerations to be applied in selected forest areas within 
a decade or so. The possibilities for such "remedial management tf will be 
dependent on research in progress or to be initiated now. 

The case studies in both Ghana and India illustrate other approaches to 
"remedial management" of production forests, to restore and maintain a 
sustainable and productive system. 

While the forest areas successfully managed in this way will certainly 
be different from the mature natural forest, their contribution to the 
conservation of genetic resources, within the context of a National Strategy 
for Conservation (see Section 5.1) which includes a range of forest conditions 
and management systems, must be very high in comparison with any alternative 
form of land use that might realistically be considered. Provided that the 
forest after logging is allowed to regenerate naturally, and is not subject 
to conversion to other land use, the options for conservation in situ will 
remain open. However they will be most heavily influenced by the care 
exercised in the logging, its intensity, and the interval allowed before the 
next harvesting operation. 

3.7 Harvesting 

Timber exploitation in tropical forests was initially highly selective 
and based on a combination of animal-powered extraction and transport by 
river. Where such systems have persisted, as is the case e.g. in many parts 
of Myanmar, they will have been largely compatible with the conservation of 
ecological values and genetic resources. However the increasing, global use 
of heavy mechanical equipment, more demanding in the intensity and width of 
road construction, and more severely damaging to regeneration and the soil, 
has had very severe impacts on the sustainability and functioning of the 
forest ecosystem. Although the effects of intensive logging, and the 
associated environmental damage, may be less discriminating in their de facto 
impact on genetic resources than highly selective harvesting and silvicultural 
"refinement" operations, they tend to revert the forest to a less species-rich 
phase corresponding to the earlier stages of ecological succession. This 
effect is made worse by the severe compaction of the soil resulting from the 
careless use of heavy equipment, which may leave substantial areas of bare 
hardened and eroded surface hostile to seedling development. Coupled with the 
harvesting of a larger number of species, as for example in the dipterocarp 
forests in Malaysia, Indonesia and the Philippines, where the ease of grouping 
of species by timber quality has increased market opportunities, the logging 
damage has sometimes been so severe that both advance growth and seedling 
regeneration of desirable species were virtually eliminated (Masson 1983). 
Since most dipterocarp seedlings do not develop readily on bare exposed soil 
the effect of intensive logging, where up to 40 per cent of the area might be 
bared by careless operation of heavy equipment, was to lose nearly half the 
potential regeneration. 

The intensity of timber harvesting also determines the degree of canopy 
opening, which has strong influence on the successful development and 
composition of the regeneration. Large gaps in the canopy favour pioneer 
species while low intensity selective logging more closely mimics the natural 
processes of forest dynamics and scarcely alters the species composition 
(Whitmore 1990). However too frequent repetition of harvesting even on a 


light selective system may adversely affect the breeding populations of the 
slower-growing species if the number of mature reproductive individuals which 
are present before the subsequent felling cycle is severely reduced. 
Repeated, intensive logging at short intervals may eliminate species 
characteristic of late-stage mature-phase and climax forest and is liable to 
produce a combination of fast-growing pioneer species of low timber value with 
masses of climber tangles and areas of bare soil. Such a mosaic of cleared 
areas and low perennial weed growth may be invaded by fire, particularly in 
semi-deciduous and monsoon forests, with catastrophic effects on the 
regeneration of most timber species and their genetic variation. 

The constraints on investment in forest management referred to earlier 
have led to increasing reliance on the logging operation as the principal 
means of influencing forest composition, structure and development, rather 
than the (expensive) thinning and refinement operations of the more complex 
silvicultural systems. Even when silvicultural operations are carried out 
after logging their effectiveness is determined largely by the state of the 
canopy, soil and regeneration left by the exploitation. Skilful and 
responsible harvesting, undertaken with understanding of ecological principles 
in forest dynamics, can itself serve silvicultural and conservation 
objectives. However all too often such aspects are entirely disregarded by 
logging personnel paid on a task or output basis and concerned to maximise 
rates of extraction regardless of the effect on the forest and the site. 
Further damage may be done by repeated harvesting without allowing sufficient 
time for recovery and regeneration. This has happened where market demand has 
arisen for species considered uneconomic at the time of the initial selective 
logging, and concessionaires have been allowed to re-enter the forest, 
regardless of the impact on the regrowth. 

Failure to adjust the harvesting operation to meet long-term management 
and silvicultural objectives is the greatest and most dangerous weakness in 
existing attempts at tropical forest management (FAO 1989c). Conversely the 
development of a productive partnership between the timber operators and the 
forest managers is the most essential component of a management and 
conservation strategy. Without this the possibilities for the conservation 
of genetic resources in production forests must be severely limited and 
restricted to accidental residual populations. 

Recent studies (Kerruish 1983; Jonkers 1987; FAO 1989b; FAO 1989c; FAO 
1992a: Jonsson and Lindgren 1990) show that much of the damage caused during 
exploitation could be readily avoided at little if any additional cost. 
Although the choice of equipment is important, as powerful tractors and cable 
extraction systems can both cause severe damage, it is above all the lack of 
planning, training, supervision and appropriate incentives to the proper use 
of equipment which are responsible for most damage. Some studies in Sarawak 
have shown that the introduction of orderly logging reduced the area of forest 
severely damaged by 44% and at the same time effected a saving of 20% in the 
logging costs. Similarly in Surinam damage from skidding operations was 
reduced by 40% in properly controlled logging, while overall productivity was 
improved by 20% (Jonsson and Lindgren 1990). 

The amount of damage is broadly related to the number of trees felled 
rather than the total volume of timber extracted (Whitmore 1990). Insofar as 
the effect on the advance regeneration is accidental it tends to be 
distributed over all tree species in an essentially random way, so that in 
terms of impact on the genetic resources it is unselective (Johns 1988). 
However the impact on species already rare and subject to selective logging 


will be potentially severe, if the future breeding populations are thereby 
further reduced. The seedlings of shade tolerant species, which depend for 
successful regeneration on survival for long periods under the forest canopy 
rather than rapid colonisation of gaps or sprouting from seed banks dormant 
in the soil, are particularly vulnerable to damage by heavy logging equipment. 
This compounds the adverse impacts on such species from extensive and sudden 
canopy opening. Since it is the heavy-seeded climax species which are most 
often dependent on animal seed dispersal the extent to which logging disrupts 
animal populations may also further affect these timber species. Stock 
mapping, timber marking, regeneration surveys and operator training, allied 
to careful planning of road making and logging operations, could be designed 
to conserve selected species and populations. Some studies have also 
indicated that quite small areas of logged forest within or adjoining logging 
concessions may be critically important to the survival within the area of 
keystone animal species (Johns 1989). 

Although the determination of the influence of different logging 
practices and intensities on the species composition and genetic resources in 
the forest requires ecological and autecological research there is no doubt 
that the degree of care exercised in the harvesting operation has the most 
profound influence on the future options open to management and conservation 
actions. The nature of the changes needed to control and provide incentives 
for responsible logging is clear in terms of the length and nature of timber 
concession agreements, levels of stumpage fees and so forth. Equally 
important is the interest and involvement of local communities in and around 
the forest, whose activities in the wake of logging operations providing 
increased access to the forest can strongly affect subsequent regeneration. 

3.8 Non-timber Forest Products (NTFP) 

The importance of the many non-timber forest products extracted from 
natural tropical forests is now widely recognised. The term now current for 
such products, NTFP, generally embraces all materials of a biological origin 
excepting timber which is being extracted on an industrial scale. The range 
of products includes foods, spices, medicines, fodder, essential oils, resins, 
gums, latexes, tannins, dyes, rattan, bamboo, fibres, a great variety of 
animal products and ornamental plants. Food and fodder sources in the natural 
forest are particularly important as dietary supplements, to reinforce 
seasonally dependent agricultural systems, and in times of drought or other 
emergency conditions (FAO 1989d). They often represent the highest evidence 
of value of the forest as forest in the eyes of the local people, and are 
therefore an important factor in the conservation of the total resources of 
the forest, including its genetic diversity. 

The NTFP may also represent a major source of economic benefit in the 
national economy. Because those products which are used locally, very often 
the major component of NTFP, do not enter the market in which traded values 
are recorded, it is difficult to quantify properly their actual or potential 
value and therefore this is certainly greatly underrated. The potential value 
of future products which it may be assumed remain to be identified in tropical 
rain forests, for example possible pharmaceutical or cosmetic products, is 
often invoked in the context of the option values of biological diversity. 
However even without allowing for such possible future additional benefits the 
actual quantified value of NTFP is very significant. Southeast Asian sources 
probably account for most of the several billion dollars in annual world trade 
in NTFP (de Beer and McDermott 1989). Available estimates of export values 
indicate that the total figure for Indonesia alone in 1987 was at least 


US$238 million. Probably US$100 million of this figure relates to rattan, of 
which Indonesia supplies 90% of the world demand. However deforestation and 
forest exploitation are eroding the resource base and it has been estimated 
that about one third of the rattan species in Malaysia and Indonesia are under 
threat of extinction (Dransfield 1987). 

Characteristic attributes of NTFP include their great variety and 
relatively high value per unit weight or volume, as compared with most 
tropical timber. Their harvesting is more labour-intensive and requires 
relatively little capital investment. Although the yield per unit area of 
forest is usually low it can be, in the case of some such products, harvested 
annually on a sustainable basis, with little or no disturbance to the soil or 
ecological functioning of the forest. Often the maintenance of a forest 
canopy is a necessary condition for the production of NTFP and therefore the 
possibilities exist for the simultaneous development of both timber and non- 
timber resources, with the latter providing an earlier economic return and a 
continuous source of income for local populations while the timber crop is 
coming to maturity. Selective logging can have a positive effect on some 
NTFP, such as rattan and edible fungi. Rattan grows best in gaps in the 
canopy, which can result from selective timber extraction. 

Insofar as the conservation of biological diversity and genetic 
resources of the tropical forests is dependent on systems of management that 
mimic as closely as possible the natural ecological conditions and processes, 
rather than drastic alteration of the forest condition through intensive 
timber exploitation or clear-felling, the simultaneous harvesting of NTFP may 
therefore be important in bearing the costs of conservation within production 
reserves. However it will be essential to undertake adequate inventories of 
the non-timber resources, in association with normal forest inventories, and 
to specify precise objectives in the management of each area of forest. The 
degree of precision needed in assessing the NTFP resources may be determined 
by the level and methods of harvesting proposed. If the products are to be 
gathered by local people freely and informally the level of information needed 
will relate principally to the regeneration and sustainability of the resource 
and qualitative assessments may be sufficient (see also Ghana case study). 

The interest in NTFP in the context of in situ conservation of forest 
genetic resources is therefore twofold - for their contribution to the 
feasibility of the conservation and management of forest resources; and for 
their genetic resources and their intrinsic value as components of the genetic 
diversity of the ecosystem. This implies the need to develop systems of 
multiple-use management of the forest (FAO 1984; FAO 1985a). Forest trees 
producing edible fruits or other products are frequently distributed widely 
but at low density per hectare and special attention may be needed to maintain 
viable breeding populations. Moreover they are likely to be involved in food- 
web systems, whereby seed dispersers or pollinators of other tree species may 
be dependent on food supplies from the fruit-bearing species. Conversely 
important fruit trees, such as the Brazil nut (Bertholettia excelsa) have 
known dependencies for their satisfactory pollination on certain large nectar- 
gathering bees which in turn are dependent for successful mating behaviour on 
wild orchid species. The loss or scarcity of orchids thus threatens the fruit 
production of Brazil nut trees (Prance 1985). 

Management for timber production as the primary objective should be 
planned to be compatible to the greatest extent possible with the production 
of NTFP (FAO 1989b). This implies not only a wider information base from 
broadly based forest inventories but also much more complete understanding of 


the forest dynamics. High hopes have been raised for the future of so-called 
Extractive Management Reserves in the Brazilian Amazon, based on a range of 
NTFP resources, of which the best established are rubber and Brazil nuts. The 
range of products already identified is certainly large and some studies have 
indicated a high commercial potential (Peters et al 1989). However there are 
considerable uncertainties over the replicability and sustainability of 
extractive management systems and some recent studies indicate that careful 
selective extraction of timber is likely to be a necessary component of 
overall management to secure sufficient levels of income from the forest. 
While the development of NTFP resources may have positive impacts on the 
conservation of genetic resources in a wider sense, the production of NTFP and 
of timber is not necessarily always fully compatible. Favouring NTFP can, at 
times, lead to a marked decrease in levels of timber harvesting, at least in 
the short term, since many timber trees are also sources of fruits or other 
extractives; the cultivation of some high value products such as cardamom 
(Elettaria cardamom) in the forest may hinder the establishment of 
regeneration of timber species in these areas (FAO 1984); etc. 

Nevertheless the possibilities for different systems of multiple use of 
natural forests are of undoubtedly high potential in terms of their 
contribution to conservation of ecosystems and the in situ conservation of 
genetic resources of a variety of species. While it may be difficult and, in 
some cases at least, impossible to combine the different management objectives 
on the same limited area of forest, with the same intensity of timber harvest, 
canopy opening and population refinement, different working circles within the 
same forest may be used to maintain a mosaic of different ecological stages 
or conditions. In some cases working circles may overlap and in others they 
must be kept geographically separate. Such zonation is also compatible with 
the development of "buffer zones" around the forest, and "core zones" devoted 
to strict protection. This is dependent on high levels and intensity of 
management (see also the India case study in Part II). 

3-9 Involvement of local people 

Very little tropical forest is truly virgin forest in the sense of 
never having been inhabited by man (Webb 1982). Most areas have been subject 
to forest clearance and cultivation and are now composed of a mosaic of 
patches in varying stages of development towards the "climax" condition. A 
Nigerian forest described by Jones (1955-56) had apparently not reached a 
steady state after about 250 years (Whitmore 1991). In some areas, 
particularly where clearance was very extensive and prolonged by repeated 
fires, the species composition of the vegetation has suffered a major change, 
for example to a derived savanna woodland, or even grassland, possibly 
maintained by grazing and browsing as well as occasional burning. However in 
the absence of such repeated destruction of the regeneration, tropical forests 
show considerable resilience in recovery after clearance. On the other hand, 
unless given areas are allowed to remain as, and return to, the mature-phase 
(climax) forest, the genetic resources of species characteristic of this phase 
may become threatened due to the long timescale needed to reach it. The 
establishment of forest reserves to be managed for timber production, to the 
exclusion of traditional shifting cultivation, was initially a safeguard 
against such a danger. This has been compromised by the increased intensity 
of timber exploitation and the widespread failure to apply adequate harvesting 
controls and lack of subsequent management interventions to ensure effective 
regeneration in harvested forests. At the same time increasing encroachment 
or illegal logging which have reduced at least parts of the forest in many 
(most) tropical countries to a very degraded condition. As the increasing 


human populations put more pressure on the scarce resources of fertile land 
the threat to the remaining forests has continued to increase (see also India 
case study). 

Recent reviews of forest management recognise that activities must take 
full account of the needs of the rural communities and that no attempted 
management system can be sustainable without the broad approval of local 
people in both planning and implementation (FAO 1989b; FAO 1989c; FAO 1992a). 
Such approval is unlikely to be achieved without the provision of some 
tangible benefits in the short as well as the longer term. As long as the 
perceived benefits of logging are that it allows access to the forest for 
illegal farming and theft of timber, particularly the undersized stems and 
advance growth of valuable species left as a result of selection felling to 
girth or diameter limits, the combined effects of the legal exploitation and 
subsequent depredations must be increasingly damaging to the genetic resources 
of valuable species. However the incorporation of extractive management of 
NTFP, together with the development of small scale rural enterprises based on 
the selective extraction of timber, could provide local employment and income. 
Increased participation of local people, combined with less intensive logging 
regimes, could then help to conserve a wider spectrum of genetic diversity iri 
situ. However the fundamental requirement is the full endorsement by the 
local population that the land should remain permanently under forest. 

Such theoretical models of the positive involvement of local communities 
in fully participatory management of production forests on a sustainable basis 
are still at only a planning or pilot stage in a few countries. The OEPF 
(Organisation de Ejidos Productores Forestales) project in Quint ana Roo, on 
the Yucatan peninsular in Mexico, has been cited as an example of the 
involvement of local communities in the management of timber production 
forests, formerly under concessions to logging companies. Here the 
communities (ejidos) are directly involved in all aspects of forest management 
and share in the profits generated through the sale of forest products. A 
high level of local support for the forest management is reported, related to 
the provision of employment and income (WWF 1991). However technical problems 
related to the regeneration and other silvicultural aspects of the project are 
also reported to require substantial external assistance (WRI 1991). 

Following earlier reviews of possible approaches to multiple-use forest 
management in India and in Ghana (FAO 1985a) new initiatives are now under way 
in both countries, including elements of both the involvement of local people 
and the conservation of biological diversity and genetic resources. In the 
Western Ghat forests in southern India draft proposals for multiple-use 
management are based on the zonation of the forest and surrounding land into 
five management zones, of which the central zone (Zone I) is to be dedicated 
principally to the conservation of biological diversity and genetic resources. 
This approach reflects principles recognised in the "buffer zone" concept 
(Sayer 1991) as well as the participatory approach of social and community 
forestry schemes. 

Another example of ways in which local people, particularly those long 
settled or indigenous to the area, can have positive impacts in the 
conservation of the forest and its genetic resources is through the use of 
local knowledge in taxonomic, ecological and phenological studies. Such 
studies are essential to the adequate understanding of forest dynamics for 
conservation objectives. With appropriate orientation to the lines of 
scientific research needed and some training in the categories of information 
required, the intimate knowledge of the forest and of many of the species 


which is often held by local people provides a valuable basis for taxonomic 
and ecological studies. Many tropical botanists can testify to the skill and 
value of local "tree finders" in taxonomic and ecological studies in all 
tropical regions. 

The scale of the data collection needed, even for a small selection of 
the many thousand species present in most tropical forests, is out of all 
proportion to the scientific manpower and financial resources available within 
each country and internationally. An interesting example is the use of so- 
called "para-taxonomists" in the current programme for a national inventory 
of biological diversity in Costa Rica. This is under the overall direction 
of the Costa Rican National Biodiversity Institute (INBIO) which initiated the 
programme in 1989 with the joint objectives of conservation and the 
exploration of potentially valuable organisms. Data from new collections is 
brought together with earlier information in a computerised database which can 
provide information on important aspects of a species 1 biology and autecology 
linked to taxonomic identifications. The mounting of intensive short courses 
in M para-taxonomy l! has transformed the rate of collection and subsequent 
specialist identification of arthropod species. In addition to the value of 
the information and material collected this approach can provide an important 
link between the local communities and the forest managers (Tangley 1990). 
In the programme in Costa Rica, in contrast to the situation in many African 
and Asian countries, the local population is not indigenous to the region and 
lacks the long history of forest use still found in most other areas in 
Central and South America. The involvement of the para-taxonomists provides 
a link between the communities and the better understanding of the natural 
resources of the country. Since the most damaging impacts on the natural 
forests came from immigrant populations and technologies this improved 
understanding through involvement in gathering data on the biological 
diversity has both direct and indirect benefits. 



When considering possible action and investment in the conservation of 
ecosystems, species and genetic resources some assessment of the future 
demands on the forest, and the land it presently occupies, is essential. The 
existing growing stock and particularly the advance growth at the time of 
harvesting will determine the species composition and possible selection of 
the seed bearers at the end of the cutting cycle, perhaps 15 to 30 years, or 
up to 45 years or more ahead in some cases. However the management and 
conservation of the genetic resources of the timber crop are concerned with 
the influence on the next reproductive phase and on subsequent generations, 
taking into account such factors as the possible effects of inbreeding, 
genetic pollution of locally adapted genepools by outside (introduced) pollen 
sources, or genetic drift on the evolution of the populations. This implies 
consideration of the management and production objectives that may apply 
perhaps 150 or more years in the future. 

Any attempt to predict the socio-economic and environmental situation 
of the production forests so far ahead must be so uncertain as to be of very 
questionable value. Even within the past half century the changed nature of 
the demands and opportunities surrounding the forests has radically altered 
approaches to management. Moreover the rapidity and scale of the changes, 
particularly in aspects of technology relating to the use of timber and wood 
and to the possibilities for manipulating the crop, including aspects of 
genetic engineering, are still apparently increasing. In addition the 
probability of significant change in global and regional climates adds further 
uncertainty, since the exact nature of such changes and therefore the nature 
and extent of their impacts on the forests, and on the environments for future 
production forests, are largely unpredictable. It is thus not to the 
prevailing socio-economic, market and environmental conditions, nor those of 
the past decades, that the genetic resources of the forest must be matched, 
but to those well into the next century. Failure to keep this fact in mind 
could lead to a dangerous limitation of the conservation objectives and of the 
necessary genetic base to adapt to unforeseen demands. 

4. 1 Population and Land Use 

Despite concern for great uncertainty related to future environmental 
conditions and market demands described above, the inevitability of the 
increase in human populations and in the severity of the related impacts on 
natural resources, including natural forests, must be the central concern in 
forward planning. Unless some significant additional improvements beyond 
current expectations can be made in agricultural productivity on the existing 
crop lands, very large areas of additional arable land will be needed each 
year to feed the increasing population. On past and present experience this 
land will be taken largely from existing forests. Economic analysis of the 
productivity and contribution to national and local needs from the natural 
forests, in the face of the increasing demands on land, is likely to conclude 
that the natural forest is unacceptably expensive in its use of land. At the 
same time shortage of capital in most tropical countries, and the high 
discount rates applied to project appraisal in the use of external funds, are 
a strong deterrent to investment in the rehabilitation of the extensive areas 
of already deforested and degraded lands. The effect on the forest "land 


bank" is therefore likely to be as devastating in the near future as in the 
recent past, and as the forests are further reduced the impact on their 
genetic resources is made more severe. Again on past experience this is 
likely to be the situation even if, as is increasingly the case, agricultural 
production on the deforested land becomes unsustainable due to the inherent 
intractability and low fertility of most areas still under forest. The 
eventual costs of attempting to restore productivity to the deforested lands 
are likely to be very high, and the success may well depend on the 
availability of the genetic resources of some of the original perennial 
vegetation, shrubs and trees. 

On the assessment of some agricultural crop geneticists we have already 
come very near to the limits of lands utilizable for agriculture (Hawkes 1990) 
and must look to the more effective use of marginal lands through adaptation 
based on selection and breeding, for the needed increase in food production. 
This underlines the importance of the genetic resources of woody multi-purpose 
species in dry and semi-arid lands, which becomes even more significant in the 
context of global warming and with the likelihood of more extreme climatic 
events and stresses. In this respect the conservation of critically important 
and endangered fragments of dry forest, and of their genetic resources such 
as i..a. those in the Pacific lowlands of Central America, is dependent on the 
recognition of their importance for long-term sustainability of land use. 
Similar considerations apply to many remaining areas of tropical forests and 
rainforests on sites of low inherent fertility, such as the deep sands 
underlying some Amazonian rain forests and the podzols of heath forest 

4.2 Timber Demand and International trade 

While trends in population and land use are clear, predictions of future 
levels and patterns of timber and wood consumption, even a matter of a few 
decades ahead, are more problematical. The long-term predictions required in 
the context of genetic resource conservation must be extremely uncertain. 
Most forecasts of timber trends have a shorter horizon but a recent study by 
Arnold (1991) undertaken for the U.K. Forestry Commission, provided some 
valuable indications of longer term trends. 

In global terms, and also for each of the major industrialised regions, 
the rate of growth in industrial wood consumption has been slowing down - 
worldwide from 3.5% annually during 1950/60 to 2.2% in 1960/70 and 1.1% in 
1970/80 (Sedjo and Lyon 1990 in Arnold 1991). This apparently reflects a 
number of long term trends which seem likely to continue, namely that the uses 
and markets for some applications, such as housing in much of Europe, North 
America and Japan, seem to be approaching maturity, while improvements 
extending the life of wood products, reductions in wasteful uses of wood, and 
increases in the role of substitutes for timber, are having an increasing 
influence on levels of demand. The results of the analysis reported by Arnold 
(1991) show that consumption is rising faster in the developing countries than 
in the industrialised regions and that substantial increases in the demand for 
industrial wood are expected in Africa, Asia and Latin America in the 
remainder of this century. 

The forest-based sector in developing countries experienced rapid 
industrialisation between 1950 and 1980 but increasing domestic demand, linked 
to the expanding populations, absorbed the major part of the increase in 
output (ECE/FAO 1986). Since the domestic markets in tropical timber- 


producing countries are less demanding in terms of quality than their export 
markets they are able to utilise a wider range of species and, as the total 
wood resources from dwindling areas of natural forest diminish, they may 
accept low-grade material of mixed origin, rather than the high quality and 
consistent supplies required by the international markets. At the same time 
the pressures on land have led several tropical countries, especially those 
with high population pressures, to propose the effective replacement of 
natural forest by industrial plantations to meet timber production needs 
(Nwoboshi 1987; Kio and Ekwebelan 1987). 

The implications of these trends for the management of natural tropical 
forests are for the acceptance by the domestic market of mixed, general- 
purpose timber, to be provided at minimum cost with little regard to the 
selection of species, and perhaps accompanied by the progressive reliance on 
artificial regeneration of fast -grow ing species, including exotics, rather 
than natural regeneration of the native forest. Unless active moves are made 
to counteract it, the management of natural forest is likely to tend towards 
the "log and leave 11 approach (Poore 1989) with limited reinvestment in control 
either of logging or of subsequent protection of the regenerating forest. 
This pattern of operation is already followed in many tropical forests, 
reflecting the low-input and low-output levels of funding and revenue. 
Investment may be more readily attracted to industrial plantations and so far 
as these also will be intended primarily to meet domestic needs for wood and 
timber, the species selected will be fast-growing general-purpose ones, 
capable of satisfactory growth on the degraded land not required for food 
production. Pioneer species, such as the tropical pines, are the natural 
choice for such plantations, with the advantage of having already been the 
subject of substantial research into their genetic variation and adaptation 
to a range of sites, through provenance trials (Barnes and Gibson 1984; Gibson 
et al 1989). If this trend were followed to its' logical conclusion the 
conservation of the genetic resources of tropical timber species carried out 
in production forests, would be limited to the pioneer and fast-growing gap- 
phase species capable of maintaining viable populations through repeated 
cycles of largely indiscriminate fellings, at intervals of probably 15 to 20 
years on many sites. 

The international trade in tropical timber has been an important source 
of revenue for many tropical countries, based in the past largely on the first 
cut in natural forest, including a high proportion of high-quality hardwoods 
with exceptional or even unique qualities. A majority of the highest value 
timbers are among the slower-growing late-phase ecological groups, 
characteristic of the mature-phase and climax forest. Some important 
commercial hardwoods, particularly among the Meliaceae (e.g. Cedrela odorata, 
Entandrophragma spp, Swietenia spp, Terminalia ivorensis, T. superba, Milicla 
excelsa Lsyn. Chlorophora excelsaj etc) are natural gap-phase species but are 
unlikely to form a large proportion of the crop under short-cycle "log and 
leave" systems of management. Without availability of continuing supplies of 
substantial volumes of such medium-value hardwood species in future timber 
harvests the prospects for profitable export-oriented management of tropical 
forests seem poor. 

The bulk of the world's current supply of industrial wood comes from the 
forests of the north temperate zone where in total, in contrast to the 
tropical forests, the net increment exceeds the removals and the volume of the 
growing stock is increasing. Moreover the developing trend in this zone to 
remove productive land from agricultural production in response to economic 


and market related pressures, is expanding the potential for further increases 
in both area and productivity of the forests. While there may be some 
constraints on industrial wood production from these temperate forests if 
decisions are made to restrict timber harvesting in favour of amenity and 
recreational objectives, and due to hazards as a result of possible pests, 
diseases or air pollution, the likely effects of global climate change seem 
favourable for increased growth rates in forests in the north temperate zone. 
Moreover the strong concentration of scientific expertise, and the high level 
of forest management generally in this zone, offer the best basis to 
anticipate the effects of climate change, to mitigate the adverse impacts and 
to take advantage of the new opportunities for faster growth. It seems 
likely, therefore, that the future trade in tropical timber will be 
increasingly restricted to the export of higher quality woods to the 
industrialised world, with some continuing and developing trade in general 
purpose timber between developing countries (Arnold 1991). 

Leslie (1987) has drawn attention to the special economic importance of 
the high value timbers which are unique to the natural tropical forests, and 
for which, therefore, there are no adequate substitutes. He points out that 
the market outlook for these timbers is the direct opposite of the weak market 
prospects for most of the products of the alternative agricultural or 
plantation management systems which compete for land with the natural forests. 
In these circumstances the relative economic prospects for natural forest 
management, if based on such high-value timber species, can only improve. 

Some recent studies of the possible incentives to sustainable tropical 
forest management highlight the potential importance of securing or 
transferring a greater proportion of the ultimate value of the products, as 
realised in the market outlets in the industrialised countries, for 
reinvestment in the forests (e.g. OFI 1991). An important aspect of this is 
the development of efficient secondary and further processing of forest 
products in the countries of origin. Exports of processed products have been 
expanding, despite some evident constraints on market opportunities in the 
main user regions. There are dangers in the expansion of industrial capacity 
beyond the properly sustainable levels of the allowable cut in the forests. 
Nevertheless with adequate quality control and marketing arrangements high- 
value secondary and further processing seems the best option to maintain a 
unique market niche for selected tropical timbers, thus providing an incentive 
for sustainable management of natural tropical forests. Reports of species 
such as ebony, teak and rosewood being exceptionally traded at prices between 
US$ 5 000 and $ 7 000 per m 3 (ITTO 1991) indicate the existence of a high 
value market niche which is likely to remain firm as the availability and 
supply of such high quality timbers from the natural forests continues to 

Concern among the public and the media in the industrialised countries 
over deforestation and degradation of the tropical forests, and particularly 
the impact of logging on tropical rainforests, is likely to have an increasing 
impact on international trade. On current trends it seems likely to reduce 
substantially the traditionally strong demand for tropical timber in Germany, 
the Netherlands, the U.K. and the U.S.A., and even in Japan. The 
International Tropical Timber Organisation (ITTO), at its 8th Council Session 
in 1990, adopted the target date of the year 2000 by which to ensure that all 
tropical timber in international trade should come from sustainably managed 
forests. At the same session it approved a set of international guidelines 
for the sustainable management of natural tropical forests (ITTO 1990) 
elaborated by an Expert Group in which individual experts, international 


organizations (FAO) and an NGO (WWF) were represented. At its 10th Session 
in 1991 the ITTO Council initiated action to develop Guidelines for the 
Conservation of Biological Diversity in Production Forests, to complement the 
already existing, general guidelines on sustainable management in natural and 
plantation forests. 

To the extent that the timber producing countries were able to achieve 
the ITTO target of ensuring sustainable management of the production forests 
by the year 2000, and installing internationally accepted certification for 
timber to that effect, the related consumer resistance to tropical timber 
imports might be removed. The criteria for sustainable management seem 
certain to include aspects of ecological and environmental concern, including 
the conservation of biological diversity and of genetic resources of the tree 
species being marketed. Depending on the criteria adopted for judging 
sustainable management, production forests managed to maintain a broad 
spectrum of species and their intra-specif ic variation, and including a range 
of successional stages over the national territory, are more likely to qualify 
for public and political approval than more intensively logged on a short 
cutting cycle, without concern for genetic conservation. 

Countries which are seen to incorporate the conservation of forest 
genetic resources, and concern for the broader range of biological diversity, 
within their management systems in production forests will be best placed to 
secure favourable markets for timber. 

4.3 Tropical Forests and Environmental Concerns 

The link between tropical forests and the stability of local, and 
possibly regional and global climatic conditions is widely accepted, if still 
imperfectly understood. Studies in the Amazon and in West Africa have shown 
the importance of transpiration from tropical forests in influencing local 
rainfall, and their significance in the hydrological cycle (Salati 1987; 
Shuttleworth 1988). It is thought that tropical forests may play a key role 
in the general circulatory systems of the atmosphere, with related influence 
on precipitation patterns. Their role as a major store of carbon is clear, 
although the mature-phase forests are presumed to be in approximate 
equilibrium in their sequestration and discharge of carbon dioxide. In that 
respect the establishment of fast-growing young forests is clearly a 
potentially greater influence on possible global warming, to the extent that 
it may be the result of increasing carbon dioxide concentration in the 

The species composition of tropical forests, except insofar as it may 
be essential to the functioning of the ecosystem on a given site, is unlikely 
to be of critical importance to their role in regional or global climatic 
stability. There is also little evidence for links between species diversity 
and ecosystem function (di Castri and Younes 1990). With the exception of 
certain "keystone" species there is apparently a high level of species 
redundancy in the functioning of the highly diverse tropical forests, and it 
seems likely that well-managed secondary forests, composed largely of 
relatively fast-growing species characteristic of the earlier stages of 
ecological succession, could adequately fulfil the forest's environmental 


However it is also increasingly accepted that the loss of biological 
diversity is itself an environmental problem. The danger for such loss is 
immediate, and therefore as urgently in need of increased international action 
as is the threat of global climate change. 

In 1988 UNEP, together with other members of the Ecosystem Conservation 
Group (then consisting of FAO, Unesco, UNEP, IUCN, and WWF International) 
initiated action towards the preparation of an International Convention on 
Biological Diversity. From the outset the special importance of tropical 
forests in this connection was recognised. The fundamental requirement of the 
in situ conservation of genetic resources and the conservation of ecosystems 
and natural habitats was taken as a central principle, linked to general 
obligations on all parties to a possible Convention to conserve natural 
habitats, species, viable populations and genetic resources in situ. There 
was also widespread recognition of the need to integrate the conservation of 
biological diversity with development, and the possible role of forests 
managed for the production of timber and other products in this connection. 
The various drafts of a convention prepared by the Ecosystem Conservation 
Group, were subsequently discussed by national Governments in the context of 
preparations for the United Nations conference on Environment and Development 
(UNCED) 1 . WRI, IUCN and UNEP, in consultation with FAO and Unesco, also 
prepared a "Biodiversity Strategy and Action Plan 11 , calling for a decade of 
action, and for the necessary financial resources to be made available 
internationally (WRI 1992). 

Discussions of a possible international "umbrella" convention, charter, 
protocol or other agreement aimed at the conservation of forests, as suggested 
by the group of seven leading industrialised countries at their meeting in 
Houston in July 1990 were actively pursued in the FAO Committee on Forestry 
in September 1990, and at the FAO Governing Council meeting in November 1990. 
Subsequently it was agreed to pursue these discussions in the forum of the 
preparatory committees for the UNCED meeting, and this led to the formulation 
of a draft statement of principles for the management, conservation and 
sustainable development of all types of forests 2 . Although not legally 
binding the "Forest Principles" may be used as a basis for a more formal 
international agreement in future. Any such international instrument should 
pay particular attention to the importance of tropical forests in the 
conservation of biological diversity and genetic resources, and to the 
implications for the provision of both financial and technical assistance to 
the tropical countries in that connection. 

While genetic diversity of the tropical forests may not be significant 
in the forest *s climatic function, it is certainly vulnerable to the effects 
of climate change. The effects may be particularly severe in the major 
ecotones, where adjacent biomes meet, for example in the transition zone 

1 The resulting International Framework Convention on Biodiversity was 
signed at the UNCED Conference in Rio de Janeiro (June 1992) by 154 
countries; it will come into force after national, governmental 
ratification by 30 signatory countries (see e.g. FAO 1992b). 

2 "A Non-Legal ly Binding Authoritative Statement of Principles for a 
Global Consensus on the Management, Conservation and Sustainable 
Development of all Types of Forests", was adopted at UNCED, Rio de 
Janeiro (June 1992). 


between closed tropical forest and savanna woodland (Holdgate et al 1989). 
In such situations, where it has been suggested that a change of 3C in 
average temperature would lead to a shift in habitat type of roughly 250 km 
in latitude (MacArthur 1972, in McNeely 1990), each species will respond 
within its own capacity. This will be strongly influenced by the levels of 
genetic diversity between populations and between individual trees within each 
species, and places greater significance on the conservation of the genetic 
resources of the woody species in such transition zones. 

In the tropical forests pioneer species with light, wind-distributed 
seed, or those with highly effective dispersal through birds, bats or other 
animals known to disperse seeds over wide areas, are likely to be able to 
adapt more easily to climatic change than species with large, heavy fruits 
that fall intact, and whose seedlings are adapted to survive under the forest 
canopy. This suggests that the trees characteristic of mature-phase climax 
forest are likely to be more disadvantaged by climate change than the more 
widespread pioneer species, especially if the latter also exhibit a high 
degree of diversity and strongly outbreeding systems. 

4.4 Protected Area Systems 

Increased recognition of the importance of biological diversity, and the 
exceptional wealth of tropical forests in this respect, may improve the 
opportunities for systematic inclusion of aspects of in situ conservation of 
genetic resources in fully Protected Areas, such as National Parks or Native 
Reserves. The principles were already established in the World Conservation 
Strategy (IUCN 1980) which has been widely accepted. Central to the Strategy 
is the recognition of the interdependence of conservation and development. 
This theme is further developed in the 1990 adaptation of the Strategy (IUCN 
1991 a) which calls for a comprehensive system of protected natural forests and 
for expansion of efforts to conserve forest genetic resources. The use of the 
term "Protected Area" covers a variety of approaches to the protection and 
management of natural and semi-natural areas, classified into eight major 
categories. Several of these categories can permit sustainable harvesting of 
forest products within the management objectives and practices, both to 
conserve biological diversity and to provide sustainable benefits to local 
people and to national economies. This applies for example, to Category IV 
(Nature Conservation, Managed Nature Reserves or Wildlife Sanctuaries), 
Category VI (Resource Reserves), Category VII (Natural Biotic Areas or 
Anthropological Reserves, for Traditional Local Harvesting of NTFP) and 
Category VIII (Multiple-Use Management Areas or Managed Resource Areas) (IUCN 
1990). However the fundamental objectives of the Protected Areas are to 
maintain the ecological processes inherent in the natural systems and to 
conserve their genetic diversity and resources for sustainable use. 

Two other categories of conservation sites are recognised 
internationally which may overlay the IUCN categories, namely the Biosphere 
Reserves, under the Unesco Man and the Biosphere (MAB) Programme; and the 
World Heritage Sites, nominated by countries party to Unesco 1 s World Heritage 
Convention. The Unesco MAB Programme works closely with FAO, UNEP and IUCN and 
with the International Council of Scientific Unions (ICSU). All Biosphere 
Reserves are intended to have a scientific as well as a developmental 
objective, and may permit harvesting, including logging, where considered 
appropriate to improve understanding of the scientific basis for sustainable 
management. Areas accepted as World Heritage Sites may qualify for some 
financial assistance through the World Heritage Trust Fund, which might entail 


restrictions on use of the resources. Roche and Dourojeanni (1984) have 
evaluated the various categories of Protected Areas in terms of their 
contribution to forest genetic resource conservation. 

Although Protected Area systems form a central "core" in national and 
international action to conserve biological diversity, the developments in 
conservation biology (e.g. Harris 1984; Sou 14 1986; Wilcox 1990) have revealed 
the limitations of such systems in the conservation of ecosystems, species and 
genetic resources. Most conservation biologists now recognise that the 
Protected Area networks, even at the more optimistic assessments of the areas 
likely to be secured, will not be able to conserve all, or even most, of the 
species and genetic resources desirable (FAO 1989a; FAO 1992c; McNeely et al 
1990). The total area and pattern of distribution needed would far exceed the 
practical possibilities and the willingness of local communities and 
governments to set aside such areas from productive use, or to decisively 
limit such use. The only solution, therefore, is a planned mosaic of 
Protected Areas, integrated with production forests managed in such a way as 
to contribute in a complementary capacity to the overall conservation of 
biological diversity in general, including genetic resources of important 
component species. 

The location of Protected Areas is normally based on considerations of 
natural landscape value, species richness, endemism, degree of threat from 
destruction of the habitat, theories of Pleistocene refugia and attention to 
so called "hot spots" in deciding priorities (Wilson 1988; Myers 1988; Reid 
and Miller 1989; McNeely et al 1990; Wilcox 1990). Their representativeness 
in respect of forest genetic resources is severely limited both by the lack 
of information on patterns of distribution, especially at the intra-specif ic 
level of variation, and by the pressures from other land-use systems. Most 
remaining areas of primary or climax forest are on sites judged unsuitable for 
intensive agriculture or very remote from centres of population, or both. In 
the fertile tropical lowlands, forests have either been heavily modified or 
eliminated. Where the opportunity to select and set aside Protected Areas 
still exists the attempt to conserve potentially valuable populations of 
important species must normally be based on patterns of environmental 
variation or of variation in plant communities, rather than on detailed 
knowledge of the genetics and actual variation patterns of their component 

The size and shape of individual Protected Areas also are strongly 
influenced, and often determined, by outside pressures rather than theoretical 
considerations of minimum population size. However even small areas can make 
important contributions to conservation (Simberloff 1982, 1983) and reserves 
of even less than 10 ha can be effective in conserving viable populations of 
many plant species (McNeely et al 1990). One advantage of small areas is that 
they are more manageable, both for the study of their species composition and 
autecology and for their effective protection, and they offer greater 
possibilities for diversification over the whole of the natural territory. A 
thorough inventory of the tree flora, and desirably other flora and fauna 
(considering occurrence, density and distribution), is essential for efficient 
conservation, and may enable such small areas to be effectively used in 
conjunction with other in situ conservation sites, and with ex situ action 
(see also the Ghana case study). 


The value of Protected Areas for the conservation of genetic resources 
is usually weakened by lack of resources and capability for their control and 
management. Roche and Dourojeanni (1984) have drawn attention to the 
necessity for effective management of Protected Areas, and the key role of the 
Managed Resource Areas (IUCN Category VIII) in this respect, since they permit 
manipulation of populations with a view to the utilization and possibly 
incremental development of their genetic resources. They have also suggested 
a review of the legislation governing other categories of Protected Areas, 
such as National Parks, in order to permit activities such as harvesting of 
seed and other reproductive materials. They point out that the totally 
protected area ("core") within many National Parks is often sufficiently large 
to allow for zonation into different sections, under possibly different 
management objectives and practices. This principle, of the association of 
different systems of management within contiguous forest blocks, is very 
important to the effective integration of conservation and development 
objectives. It is well illustrated by the Virgin Jungle Reserves within 
production forests in Malaysia, and by the concept of "buffer zones" around 
protected forests. 

4.5 Buffer Zone Forest 

Protected Areas, particularly those of less than ideal size and 
location, can only meet their conservation objectives if the land surrounding 
them is under appropriate management compatible with the objectives within the 
Protected Area itself. This may require the establishment of a "buffer zone", 
which can both meet those objectives, and also provide appropriate benefits 
to the local people. Experience has shown that legal protection alone of 
conservation areas is insufficient to prevent encroachment or damaging 
incursions into the forest, particularly where human settlements are located 
close to the boundary (Sayer 1991). Moreover in relatively small reserves the 
tree species which occur naturally at very low densities will be at risk from 
possible inbreeding or inadequate levels of regeneration, which may be 
relieved if their populations are extended into surrounding areas of managed 
forest. Such extension of protection may help conserve a wider range of 
intra-specific variation. 

Some attempts at buffer zone development have proved disappointing 
(Wells et al 1990) but others have been more successful. The benefits, in 
addition to the extension of the effective population size of some species and 
the separation of the "core" protected area from human settlements and 
intensive agriculture, include the development of local support for the 
conservation objectives as a result of beneficial involvement in use of the 
buffer zone, including e.g. hunting areas. 

The definition of the buffer zone concept by MacKinnon (1981) was "areas 
peripheral to National Parks or reserves which have restrictions placed on 
their use to give an added layer of protection to the native reserve itself 
and to compensate villagers for the loss of access to strict reserve areas". 
However Sayer (1991) interprets the concept more widely to incorporate a range 
of possible developmental activities which can deliver benefits to local 
people as well as extending the effect of the Protected Area, at least for 
some animal and plant species. These activities might in some cases include 
agroforestry, tree plantations or a variety of other land uses but in terms 
of the in situ conservation of the forest's genetic resources the most 
appropriate form of management is clearly modified utilization of the natural 
forest. The Corbett National Park in northern India is a long-established 


example of the way in which the maintenance of semi-natural forests of Sal 
(Shorea robusta) have buffered the Protected Area while providing a harvest 
of quality timber to the forest authorities and non-wood forest products to 
the local people. 

The best realistic prospects of maintaining a buffer zone of natural 
forest with relatively little change in its species composition, and therefore 
maximum contribution to in situ conservation, are in the few remaining major 
forest blocks, such as Amazonia, the Zaire-Congo Basin and parts of Indonesia. 
Some possibilities still exist elsewhere in Africa, where population pressures 
around the forest are still not severe, to develop a buffer zone of secondary 
forest, with a higher concentration of pioneer species. However light, 
sustained-yield selective logging or harvesting of NTFP, possibly in 
combination, should be better able to maintain a wide range of total 
diversity. The important thing is for the management of both the Protected 
Area and the adjacent land to be planned and managed as an integrated unit. 
By this means the genetic resources of species of both the late succession 
forest and those of the earlier stages of ecological succession can be 
suitably catered for in separate management zones. Clear objectives and 
priorities must be set for each zone, from the central "core 11 , where 
conservation of over-all biological diversity may be paramount, through 
selectively logged forest, with due attention to the conservation of genetic 
resources of important timber trees, to the outer zones, where production of 
the wood and non-timber forest products may be the highest priority, with 
close involvement of local communities (see India case study). 

A further important role for areas of managed forest outside the main 
Protected Areas is to serve as "corridors 11 for the movement of animal 
populations, particularly in the context of possible climate change. If 
sufficiently broad and secure they might also allow for migration of some 
plant and species, including tree species. However given the probable speed 
of change and the extensive fragmentation of tropical forests the movement of 
genetic resources of important tree species will be more dependent on ex situ 
conservation strategies. 



5.1 National Policies 

Genetic resources are a national asset of the country and therefore 
dependent on clear national policies for their use and conservation. The 
potential contribution of each production forest, or unit of management within 
a forest, to the national objectives for conservation and/or sustainable 
productive use will vary according to location, species composition, size, 
shape, environmental features and many other factors. Its actual contribution 
should be determined by the objectives and quality of management, related to 
its potential conservation value. It is neither possible nor necessary to 
prescribe equal priority and intensity of in situ conservation in all 
production forests. At a minimum level sound sustainable management should 
include provisions for the protection of site conditions, seed trees, seedling 
regeneration and advance growth of desirable species in appropriate 
combinations according to management and silvicultural prescriptions. To the 
forest geneticist a range of possible production management systems may be 
acceptable in terms of their impact on the genetic resources, depending on the 
target populations. Extreme refinement of the composition of the forest to 
favour one or a few species in an originally poly-specific forests, if done 
wi,th full understanding of the dynamics of the ecosystem and of the effect on 
its long-term functioning, may be an acceptable means to conserve the genetic 
resources of the principal species, albeit at the expense of the broader 
(species) diversity of the forest, which should receive due attention in other 
areas of the Forest Estate. Even clearfelling and replacement of the mixed 
forest by planting a single indigenous species, if based on broadly 
representative seed collections from the same site, might qualify as in situ 
conservation. However such approaches, would be exceptional to the general 
rule which would seek conservation through natural regeneration of the 
component species in the ecosystem, targeted for conservation. 

In many cases the need to maximise production of wood and other products 
from limited areas of land requires the concentration of the site capability 
on the growth of one or a few fast-growing species or species groups, most 
likely to be from pioneer or gap-opportunist guilds, and possibly exotics. 
The forest manager might argue that this action supports the allocation of 
other forest areas to in situ conservation, as part of an overall strategy. 
However past experience with the putative development of "compensatory 11 
plantations to safeguard natural forest has indicated the need for a stronger 
and more coherent overall plan to ensure that the conservation objectives are 
also met. As stated earlier the role of individual areas of production forest 
^ or in situ conservation must be determined in relation to patterns of 
distribution and variation of target species and populations, and to the 
contribution of the network of fully Protected Areas, National Parks etc, 
within an overall, national conservation strategy. In addition to using 
production forests to complement the Protected Area system, by filling key 
"gaps" in species ranges, or forest types, there can be additional value in 
the conservation role of production forests sited adjacent to a protected 
area, or forming a "corridor" to other managed or protected areas. 


The wider regional and international interests in the conservation of 
biological diversity and genetic resources related to patterns of distribution 
extending through several countries, and to global recognition of the 
importance of the tropical forests in this regard, presents an opportunity to 
secure additional financial and technical support for conservation actions. 
While these international concerns, particularly when focused on existence 
values of biological diversity, are likely to extend beyond the more immediate 
use values of the forest genetic resources of direct interest to the host 
country, they may be used to reinforce national objectives in the 
conservation of selected forest areas and populations. However to ensure that 
full advantage can be taken of these expanding opportunities, in ways that 
serve the national interest, as well as broader global objectives, a clear and 
coherent national policy on the conservation of its genetic resources is 

As seen in the case studies on Ghana and India, for example, the actions 
needed to secure effective conservation must carefully consider and include 
the interests of local populations in and around the forest, and will also 
extend increasingly outside the forests into aspects of industry and trade; 
and outside the forestry sector as a whole, to areas of responsibility in 
other sectors of government. They may impinge on central policies related to 
the allocation of revenue and expenditure, and to both internal and external 
commercial and trading practices. In these cases the necessary changes to 
management practices in individual forest areas are dependent on central 
policy decisions at a high level, involving various sectors. Review and 
revision of existing laws and regulations may be needed. Although many of the 
actions, such as the review of the level of forest fees, the adoption and 
effective imposition of differential stumpage rates, the systems of 
allocation, duration, size and operation of timber concessions etc, are needed 
as much for the broad objectives of sustainable forest management, as for 
conservation objectives, the latter roust also be kept clearly in view in 
determining overall policy. Similar considerations apply to the changes 
needed in forest management practices, and perhaps in laws and regulations, 
to incorporate NTFP and the involvement of local people in such management 
systems. The potential for this is seen in all three case studies presented 
in Part II of this document. The provision of a coherent system of incentives 
to sustainable management of the forests, at all levels from the local 
populations to other national groups in both the private and the public 
sector, and to international market conditions and investment policies, 
relating for example to trading practices, support for local processing, 
marketing assistance and maximising value of products to the country of 
origin, will be needed to secure greater attention to conservation objectives 
in the production forests. These incentives must be the subject of national 
formulation and may require international assistance (see the Ghana case 
study) . 

An urgent task is the determination of priority species, populations, 
areas and actions for the conservation of forest genetic resources in each 
country. This must take account of possible ex situ as well as in situ 
conservation, within a coherent national programme, in accordance with 
national policies. In a broader context, embracing all natural resource 
sectors, the formulation of National Conservation Strategies has been used as 
an aid to reviewing policies and redefining priorities (Poore and Sayer 1987). 
The breadth of subjects and issues involved in the whole field of biological 
diversity conservation has proved difficult to contain within a single 
comprehensive assessment, and the need for more specific definition of 


policies and practices linking development and conservation has been suggested 
(FAO 1985b; Prescott-Allen 1986). In the narrower context of the conservation 
of forest genetic resources, the multiple-use management of production and 
protection forests for the production of timber, NTFP for local use, 
environmental protection and the conservation of genetic resources provides 
a contribution to exemplify and help define the broader Conservation Strategy. 

For all of the above reasons the formulation of a National Strategy for 
the Conservation of Forest Genetic Resources is essential, to make the best 
use of the land and other resources devoted to production, protection and 
conservation, and to the opportunities for regional and international 
cooperation. An important task under such a National Strategy for 
Conservation will be the prioritization of action and coordination of 
activities, including research, to make the best use of financial and manpower 
resources available both nationally and internationally. Without inclusion 
of research within the over-all strategy, there is a danger that the limited 
scientific expertise will be wasted or inefficiently used, through the choice 
of irrelevant subjects for research, or failure to integrate the work of 
different specialists needed to resolve a particular problem. This has been 
observed particularly in the fields of reproductive biology and genetics of 
tropical trees in relation to their conservation and management (see e.g. Bawa 
and Krugman 1991; Wadsworth 1975). 

Studies of the effects of disturbance on the forest, and the response 
of the principal tree species of socio-economic value to such disturbance at 
various stages of their life cycle, are important for both management and 
conservation objectives. There is a need to coordinate action within a wide 
range of differing biological fields, using varying degrees of expertise, 
experience and technological sophistication. However too often ecological 
and autecological studies have lacked direct relevance to forest management 
activities, e.g. caused by failure to examine the comparative situations in 
both logged and unlogged forest. 

A National Strategy for Conservation could assist in channelling 
research activities to the areas of highest national priority. 

The same main elements and lines of research needed are likely to be 
common to many countries and at least at a regional (or eco-regional) level 
there may be considerable coincidence of interest in the principal species and 
genetic resources requiring study. More effective coordination of action, 
including research at the regional level has frequently been suggested but 
only infrequently achieved. The preparation of national research priorities 
in the context of the National Strategy for Conservation could assist more 
productive regional cooperation. 

A further task in formulating the National Strategy for the Conservation 
of Forest Genetic Resources will be the definition of appropriate 
institutional structures to guide and coordinate subsequent action. The exact 
nature of the institutional arrangements will need to be decided in accordance 
with existing national structures and mechanisms. Action to formulate the 
strategy may best be taken by the national Forestry Department but the need 
to secure strong inter-sectoral cooperation and high level policy review 
indicates close involvement with, if not actually within, central government 
structures. An example is the adoption by the National Planning Commission in 
Nepal of the National Conservation Strategy Implementation Programme in that 
country. This is most likely to ensure the necessary inter-sectoral 


coordination and policy integration at a high level that is essential. 
However as an initial step the establishment of a National Genetic Resource 
Unit, as envisaged under the Tropical Forestry Action Programme, TFAP (FAO 
1985b) may be sufficient, if an appropriate equivalent body does not already 
exist. The TFAP is the most appropriate existing international mechanism to 
assist the national sector review in all aspects, including the more 
appropriate appreciation of the capital value of the natural forest and its 
genetic resources. It is the failure to understand or to accept the use 
values and option values of genetic diversity in the national economy which 
has prevented the necessary investment in genetic conservation. 

5.2 Management Information 

In addition to the undervaluation of the resource, and the associated 
lack of investment in its conservation and sustainable management, the 
principal deficiency in the approaches to management is the lack of 
information on the forest structure and dynamics. For effective sustainable 
management data are needed on the composition of each of the main forest 
types, the silvicultural characteristics of the principal species and of the 
others which may compete with them at various stages of their development, 
their regeneration behaviour, growth rates, and response to canopy opening, 
logging or silvicultural operations. In most cases even the basic knowledge 
of growth and yield of the principal economic species is lacing. Management 
f r in situ conservation of genetic resources requires much of the same basic 
data on ecology and autecology that underlies attempts at silviculture in the 
natural forest, but with greater emphasis on breeding biology and genetic 
structure. Nevertheless a broadly-based forest inventory, as exemplified in 
the Ghana case study, accompanied by surveys of NTFP and of patterns of 
variation in the floristic composition of the forests, is of fundamental 
importance for both production genetic and conservation objectives. The 
inventory can also provide information on species distribution patterns, which 
are a first step in exploring intra-specif ic variation, to be included in in 
situ conservation strategies. The network of Permanent Sample Plots or 
continuous inventory plots required for production management can also be used 
for e.g. phenological studies and more fundamental research, possibly 
involving scientists from universities, both national and international see 
also Section 3.3). 

There is frequently a - wealth of unpublished data available from a 
variety of sources, from herbarium sheets to expedition records and academic 
theses, which could contribute to the information needed on forest 
composition, species distribution, phenology of flowering and fruiting and so 
forth. Modern computer-based approaches to information management can be used 
to store and assist the interpretation of such data, and to guide the 
efficient collection of additional information to fill the most important gaps 
(Jenkins 1988; Davis et al 1990). The use of Geographic Information Systems 
(CIS) can be a powerful aid in the definition and interpretation of species 
distribution patterns in relation to environmental variables and vegetation 
types. Similarly computer-based systems for handling taxonomic data have 
improved the accessibility and usefulness of information on genetic diversity 
in respect of both taxonomic groups (e.g. ILDIS: International Legume Database 
and Information Service) and geographical areas. The ILDIS system 
incorporates information on economic importance and conservation status, as 
well as botanical and vernacular nomenclature, vegetation types, references 
to maps etc (Jury 1991). The Botanical Research and Herbarium Management 
System (BRAHMS) developed at the Oxford Forestry Institute, U.K. has been 


designed specifically to handle data related to a forest genetics research 
programme (Filer 1991). Many organisations in tropical countries use CIS and 
other database management systems on microcomputer. In view of the importance 
of the efficient handling of data needed for in situ conservation a National 
Data Centre for this purpose should be a component in the national strategy. 
This can not only assist but also actively encourage the efficient collection 
of data from many sources and potential collaborators. 

Such a Data Centre would assist coordination of action within the 
country and provide a link to the regional and international data bases such 
as that at the World Conservation Monitoring Centre (WCMC) Cambridge (U.K.), 
supported by IUCN, WWF and UNEP, and to the Forestry Department of FAQ, which 
provides the Secretariat to the FAO Panel of Experts on Forest Gene Resources. 
An important activity of this latter Panel is the preparation and revision at 
its regular meetings, presently held approximately every 3 years, of global 
lists of species, by regions and operational priorities (exploration, 
collection, evaluation, conservation, breeding and use), most in need of 
action. At its Seventh Session, in December 1989, the Panel noted the need 
for more frequent and vigorous review of the priority lists. Close liaison 
and cooperation between the National Data Centres and the FAO Secretariat to 
the Panel would be an important element in this process. 

National Data Centres should also have a key role in regional 
cooperation, for example in the project proposed for the Conservation and 
Rational Use of Central Africa 1 s Forest Ecosystems, linking activities in 
seven countries involved. Similar cooperative action would be valuable in 
West Africa (Tufuor 1990b), Central America and other regions and sub-regions. 

5 3 Management Systems 

The fundamental principle in the selection of the management system for 
a particular area of forest, to take account of the needs for genetic resource 
conservation, is that it should be based on adequate understanding of the 
forest ecology, and the autecology of the principal component tree species of 
socio-economic value. 

A central problem which has plagued attempts to achieve predictable 
results from management interventions in tropical forests has been the failure 
to match the operations to the actual nature and state of the forest and the 
site. This has been the main reason why successful silvicultural experiments, 
carefully executed, have often failed to translate successfully into large- 
scale practice over the diversity of conditions in the forest as a whole. The 
attempt to apply one given management system, either monocyclic or polycyclic, 
throughout the forests, and often to apply the same silvicultural treatments 
as a "blanket" prescription, without adequate attention to variation in forest 
types, associations and stages of development, can never be more than locally, 
partially and to a considerable degree accidentally successful. Except for 
areas where intensive "refinement" of the species composition of the forest 
may be consistently maintained through several felling cycles, the impact on 
the genetic resources of the forest is also likely to be accidental and at 
times irreversible. On the other hand, regeneration studies under a variety 
of systems in different countries have shown a general resilience of the 
forest composition, provided that logging is not succeeded by further 
catastrophic impacts on regeneration, such as fire or illegal cultivation of 
non-forest crops. 


Available information on the genetics and variation of individual 
species, including the principal economic tree species in the tropics and sub- 
tropics, is generally inadequate to predict the effects of logging and 
silvicultural operations on their genetic resources, except in very general 
terms* However, as observed in previous sections, short cycles of logging and 
the indiscriminate use of heavy machinery, are likely to be more damaging to 
both species composition and reproductive systems than longer cycles (70 to 
100 years)* Similarly light selective logging, leaving a well-dispersed 
population of seed bearers of the principal economic species, is likely to be 
preferable from both viewpoints to more severe disruption of the forest* The 
methods to reduce felling damage (topographic and stock mapping, tree marking, 
road planning and construction, directional felling, choice of equipment etc) 
can be cost-effective in economic as well as ecological terms. At the 
simplest level, therefore, careful planning and control of harvesting even 
without more elaborate management prescriptions, as practised in e.g. the 
Unit&s Forestieres d'Amenagement in the Peoples Republic of the Congo (FAO 
1989b), is likely to serve both production and conservation objectives. 

Beyond this level of management the financial and manpower constraints 
related to the low valuation of the natural forest cause increasing divergence 
between the degree of information and precision needed for the management of 
genetic resources, in tailoring operations to the actual forest condition, and 
the level of investment in management permitted by the economic calculations 
of costs, yields and discount rates. In addition to securing more appropriate 
levels of revenue, as in the Ghana case study, and applying them to forest 
management, additional value should be attached to the conservation of genetic 
resources in situ, to reduce the pressures for intensification of yield. Such 
pressures, operating both on the length of cutting cycles and through the 
concentration of growth into a few currently commercial species, necessarily 
conflict with wider conservation objectives, concerned with species richness, 
ecosystem conservation and option values. However the actual level of value 
to be assigned to the genetic resources must be assessed, and the appropriate 
compromise must be struck in respect of each forest or management unit, taking 
account of the National Strategy for Conservation. 

The need for flexibility in management, according to the nature and 
actual state of the forest, is recognised at least in respect of timber 
production, in both the Malayan Uniform System, MUS (though often not 
implemented) and the Malaysian Selective Management System, SMS, which allows 
for the adaptation of management and possible silvicultural operations to the 
state of the populations of young trees eligible to form the next crop, after 
the effects of logging (FAO 1989c). The SMS prescriptions are also intended 
to take account of ecological considerations and these should include the 
conservation of genetic resources, as defined for a particular forest area, 
within a National Strategy for Conservation. Knowledge of the composition, 
ecology and silviculture of the forests, particularly in Peninsular Malaysia, 
is almost certainly better developed than that of any comparable area of 
tropical forest (e.g. Wyatt-Smith 1963; Burgess 1975; Ng 1978 and 1989; Tang 
1987; Whitmore 1990; Appanah and Salleh 1991). The practice of liberation 
thinning to manipulate the performance of selected potential final crop trees, 
as practised initially in Sarawak (Hutchinson 1987), has been used with 
apparent success in several countries, notably in Cote d'lvoire (Maitre 1991) 
and in Surinam (Graaf 1991) at least on an experimental scale. Such 
approaches can be used to favour different species or species groups according 
to the prescriptions made for a particular forest or compartment. This is 
easier to apply in the Malaysian forests, where there is a wide range of 


potentially marketable species, and a strong information base both for their 
management in the forest and their grouping for the market. However the same 
principle may be applied in forests less rich in economic species, if there 
are one or two common general purpose timbers with vigorous regeneration. 
This principle is being explored, for example, in Costa Rica, based on 
Pentaclethra macroloba to support the conservation of the populations of 
known, valuable species such as Cedrela odorata (Finegan pers. comm. 1991 ) 1 . 

The implications of the long-term trends in the international trade in 
timber, discussed in Chapter IV, are that the most secure market for tropical 
timber is likely to be for high quality species for high-priced veneers, 
joinery, furniture, musical instruments or other special applications. 
Although the bulk of such trade may be small prices are likely to be high 
(ITTO 1991). Highly selective logging of species used for such purposes, most 
of which are likely to be slow-growing and characteristic of mature-phase 
forest, possibly in combination with harvesting of NTFP, could be compatible 
with the conservation of their genetic resources and other genetic diversity 
best conserved in mature-phase and climax forests. 

At the other extreme the increasing acceptance, particularly for the 
domestic or regional markets, of a wider range of currently lesser-used 
species, including those with light, non-durable but treatable timbers, offers 
opportunities for other management models, also compatible with the 
conservation of genetic resources. These may include uniform systems, as in 
the strip shelterwood management system, operating on a pilot scale in the 
Palcazu valley in Peru (Hartshorn 1989), where early results indicate that it 
may be possible to retain a high proportion of the original tree species 
within a viable production system, with the support and involvement of local 
communities. The greater use of lesser-known species as an aid to 
conservation of the depleted resources of more valuable species, and the range 
of management actions which may be needed to achieve this, are illustrated in 
the Ghana case study. 

The management of production forests in support of genetic resource 
conservation may be most effective in close conjunction with management of 
Protected Areas, such as National Parks. These provide a reservoir of seed 
and a haven for fauna involved in seed and pollen dispersal. Although as 
explained earlier quite small areas of unlogged forest may serve this purpose, 
for example in supporting adequate populations of frugivorous birds or small 
mammals involved in seed dispersal, in certain cases much larger areas are 
required. This applies e.g. to the role of elephants in natural forest 
ecosystems in Africa in the ecology and distribution of timber trees such as 
Tieghemella heckelii (Martin 1991). The association of the Bia National Park, 
in Ghana, with the adjacent timber production forest is a good example of 
this. At the same time the production forests can provide an effective 
extension of the range of species found in the Protected Area. The valuable 
principle of incorporating protection areas, such as the Virgin Jungle 
Reserves (VJR) in Malaysia, within the production forests is clearly a most 
important component of any National Strategy for the Conservation of Forest 
Genetic Resources. 

B. Finegan, CATIE, Turrialba (Costa Rica). 

5.4 Management Plans 

It is at the level of individual forests, working circles and 
compartments, and through the preparation of detailed management plans, or 
working plans, that the principles of forest management and genetic resource 
conservation must be put into operation. To the extent that working plans for 
the forest are carefully prepared and scrupulously carried out this is a 
further instance of potential benefit to the conservation objective through 
their association with timber production. As stressed earlier many aspects 
of sustainable forest management, from inventory, growth and regeneration 
studies to responsible harvesting, should simultaneously assist genetic 
conservation objectives. This applies particularly to the provision made to 
secure satisfactory regeneration, for example through the timing of 
exploitation in relation to mass fruiting years or the retention of adequate 
numbers and distribution of seed trees in the absence of adequate regrowth of 
desirable species, as well as the avoidance of damage to soil seed banks. 

The collection and validation of data on growth and the accurate 
forecasting of future sustainable timber yields is both essential for forest 
management and also of critical importance to conservation objectives. There 
are many examples of severe threat to the genetic resources of major timber 
species through over-cutting of forests on the basis of over-estimates of 
future levels of harvest, and through repeated "creaming" of the stand. These 
have sometimes been used to justify large scale industrial development which 
then creates an insatiable and destructive demand on the forest resources. 
This is an area where the interests of long-term sustainable forest management 
and the conservation of genetic resources closely coincide. 

There are other aspects of management in which the need for care to 
conserve genetic resources goes beyond, and may at times even be seen to 
conflict with production objectives. As stated earlier it is neither possible 
nor necessary to attempt to impose conservation measures with equal priority 
and stringency in all production forests; to do so would weaken the support 
for the general concept of genetic conservation among forest managers, and 
invite widespread disregard for application of related principles in practice. 
Therefore realistic assessment of the situation in respect of each area of 
forest is essential, so that firm objectives, activities and criteria for 
monitoring their effects can be included in each working plan. Where the 
practices for sustainable forest management and genetic resource conservation 
closely coincide the prescriptions should apply in all production forests, and 
it is only necessary to refer to the additional importance of careful 
attention, for example to the restrictions on harvesting damage to 
regeneration or the importance of subsequent protection from fire, in the 
context of the National Strategy for the Conservation of Forest Genetic 

In certain forest areas to be designated under the National Strategy for 
Conservation, additional measures will be needed. These will relate to the 
assessed importance of the forest in the conservation of the genetic resources 
of the principal species of known socio-economic value, lesser-known species, 
NTFP or particular vegetation or forest types. An important criterion will 
be the location of the forest in the overall species 1 range, and in relation 
to environmental conditions. The importance of conserving a wide range of 
provenances has been established earlier, and this may be achieved to a large 
extent through the application of sustainable forest management throughout 
the Forest Estate. However particular care may be needed where the species 


occurs under more extreme environmental stresses, related to climate, soil, 
altitude etc, or at the edges of its natural range. Not only are these 
populations likely to be genetically distinct, through adaptation to the local 
environments, but they may also be more vulnerable to disruption through 
disturbance, which could radically reduce the long-term viability of such 
populations. Special consideration should be given to these marginal 
populations through limiting the intensity of logging, and through 
intensification of research into their genetic structure and reproductive 

The other main criterion in selecting areas for special consideration 
relates to the production management systems in general use in particular 
forest formations or vegetation zones. If, as is most likely, these tend to 
favour pioneer or early gap-phase species, and thus to reduce the 
opportunities for the maintenance of satisfactory breeding populations of 
mature-phase species, special attention will be needed to conserve the genetic 
resources of this latter stage of the forest's development. The extent to 
which this should be done in production forests will depend on the role and 
de facto contribution to genetic conservation of existing Protected Area 
systems but it is unlikely that production forests alone could provide 
adequate levels of geographical and ecological coverage for species and 
populations concerned. 

Genetic resource conservation should be an important factor in the 
design and operation of working circles within the forest. Hitherto the 
allocation of areas to a protection (conservation) working circle has been 
based almost invariably on considerations of accessibility, steepness of 
slope, adverse geological or drainage condition etc. rather than their value 
for genetic resource conservation. The protection of special sites such as 
streams ides is desirable for several reasons, including conservation of 
biological diversity. However the siting of conservation areas such as Virgin 
Jungel Reserves (VJR 1 s) should attempt to conserve associations representative 
of the diversity and genetic resources of the forest as a whole, or of the 
particular forest type. This is likely to frequently conflict with immediate 
production objectives but the more intensive the form of management in the 
remainder of the forest, the more valuable the diversity and genetic resources 
of the VJR or other Conservation Area may be. From the viewpoint of the 
conservation of the most desirable economic species the practice of leaving 
small pockets of unlogged forest, and well dispersed seed trees, throughout 
the production areas is likely to be needed in addition to the allocation of 
a larger area, such as an entire compartment, as a VJR. 

The real problem commonly lies in reconciling these various objectives 
with the interests of the timber concessionaire. The provisions recommended 
to secure real interest on the part of the timber harvester in the 
regeneration and long-term sustainable management of the resource should also 
be conducive to conservation of the genetic resources of the principal 
economic species. However to conserve desirable option values of the genetic 
diversity, which have no evident, present-day market value, some additional 
incentives will be needed. The most promising concept may be the setting of 
high concession rents, together with clearly defined conservation objectives, 
related for example to the omission of selected "pockets" of forest from the 
harvesting operation. To the extent that the conservation objectives were met 
there could be a refund of a proportion of the fees, or conversely for gross 
violation the imposition of penalties under the contract. It has been 
suggested that ideally concessions should not be granted purely for timber 


extraction, but as a contract between the government and the private sector 
to manage the forest for a range of products and benefits, including timber 
(Johnson et al 1991). 

In all aspects of forest management and conservation the failure to 
comply with the prescriptions and conditions set has been a frequent cause of 
damage to the growing stock, and particularly its capacity for regeneration. 
Close monitoring of operations, both to judge their conformity with 
prescriptions and to assess their effect relative to the declared objectives, 
is in the common interest of both sustainable production and genetic resource 
conservation. It may also be an important aspect of "forest accounting* 1 
designed to monitor the condition of the resource and to evaluate the standard 
of management. This will include aspects involving scientific criteria, for 
example in regard to genetic resource conservation, as well as others related 
to technical and socio-economic objectives. Criteria for monitoring should 
be clearly set in the management plan, taking account of the objectives set 
under the National Strategy for the Conservation of Forest Genetic Resources. 

The genetic resource conservation value of a concession area, as 
determined under the National Strategy for Conservation, should be used in 
assessments of the quality of sustainable forest management. Systems and 
criteria for the assessment of sustainability in forest management will be 
needed at least in respect of forests managed for the export of timber, in 
accordance with the ITTO guidelines and target mentioned above (ITTO 1990). 
There might be positive incentives through the market for conforming to the 
accepted guidelines, which will include consideration of the impacts on 
biological diversity in the forest. In any case it is in the national 
interest to ensure that the objectives set under the National Strategy for 
Conservation in respect of forest genetic resources are achieved. In both the 
assessment of the genetic resource value and objectives in a particular forest 
and the setting of criteria by which to measure the achievement, specialist 
advice of forestry geneticists may be needed. This is most likely to be 
necessary if management action is required to conserve the genetic resources 
of given target species or populations, which may involve considerations of 
population genetics and viable population size. Other specialist advice in 
regard to forest ecology, the role of "keystone" species in the functioning 
of the forest and the genetic systems of economic species may also be 

As indicated earlier, and emphasised in most recent reviews of tropical 
forest management, the involvement of local communities situated in and around 
the forest in aspects of forest management is likely to be increasingly 
essential to long-term conservation objectives. The concept of joint planning 
and management, operated differentially according to the management objectives 
of the different zones, as outlined in the India case study, allows for a 
contractual relationship with local people, without weakening the legal status 
or government control over the forest land. Such a careful balance between 
the long-term security of the genetic resources in those areas requiring 
strict protection, and the sustainable development of a variety of indigenous 
trees and other plants in appropriate management zones, provides wide 
possibilities for genetic resource conservation. However to an even greater 
extent than the management of the natural forest for timber production alone 
such multiple-use systems are likely to require specific formulation in the 
particular ecological and socio-economic situation, and incorporation of the 
full range of concerns in the management plans of the forest concerned* 


The management of areas of production forest acting also as a "buffer 
zone 11 to a National Park or other Protected Area was discussed in Chapter IV. 
The most important aspect is for the management of the two kinds of area to 
be fully integrated and under single effective control within one management 
system. This may require special administrative arrangements if primary 
responsibility for production forest on the one hand and the fully Protected 
Area on the other, come under separate ministries. However the same principle 
of joint planning and management, with clear objectives and management plans 
for each zone, should apply. The efficient management of the genetic 
resources of key economic species represented in a National Park or other 
Protected Area, may require specif ic management interventions, for example the 
collection of seed or other propagating materials for ex situ action, or 
liberation thinning to favour the development of young trees of the species 
targeted for conservation in situ. Such activities may (at times properly) 
be contrary to the general Park regulations. This could be the subject of 
examination under the National Strategy for Conservation and provision could 
then be made by amendments to laws and regulations to permit action for the 
conservation of forest genetic resources, with appropriate safeguards and 
restrictions, in accordance with the National Strategy for Conservation. 

International concern for the tropical forests is certain to continue 
and to deepen, as the forests are further reduced. This concern, and the 
allocation of resources to the extent needed to conserve the existing natural 
ecosystems and their component species, will come too late to retain them 
intact in many areas in many countries. However except where large areas are 
severely altered, for example by fire or continuous intensive cultivation, 
valuable genetic resources are likely to survive under sustained forest 
management systems; such resources are invaluable as the basis for the 
regeneration and/or restoration of functioning natural forests, in various 
forms and for a variety of uses. 


Box 2 

Management of diversity through diversity of management 

The Protected Area network, which must be the core of ecosystem 
conservation, is likely to prove increasingly inadequate to meet long-term 
objectives for the conservation of genetic resources. In addition to its limitations 
in geographical coverage, restricted often arbitrarily by other demands on land, 
it is likely to be further limited in terms of environmental diversity by the size, 
shape and management practices in each reserve area. Dynamic conservation, 
in a world in which future human needs and climatic conditions must be 
increasingly uncertain through successive generations, requires more than the 
attempt to prevent genetic erosion and to retain a fixed current state of diversity. 
It implies conserving the evolutionary process, to advance and develop 
ecosystem diversity, and targeted genetic resources, in directions which seem 
likely to prove most useful, and to retain a wide range of options to continue this 
process into the very distant future. Within this concept management actions, 
including harvesting and siMcultural activities, can be creative of genetic diversity, 
if they are freed from past pressures towards "blanket* uniformity, centred on a 
few species with minimal reinvestment of resources skills and finance. 

Logging operations alone, if they are neither too intensive nor too frequently 
repeated, but follow known principles for good practice (Dykstra and Heinrich 
1992) are likely, in initially diverse tropical forests, to leave levels of complexity in 
species composition and plant associations which, although they may not exactly 
follow previous patterns of distribution, can maintain or even increase overall 
genetic diversity. Given adequate information on the patterns of such diversity 
between compartments (or in critical cases sub-compartments), as well as across 
larger forest management units, and within the National Forest Estate as a whole, 
and given the continued development of advanced systems for handling and 
interpreting data, future management systems could be tailored to serve dynamic 
conservation strategies, at the same time as they meet the needs of industry for 
predictable flows of raw materials. This, however, will require greater 
coordination of management planning at the national level, as well as the 
integration of conservation concerns in the management prescriptions of both 
production forests and the Protected Area network, under a coordinated, 
National Strategy for the Conservation of Forest Genetic Resources. 



fiox_2 continued 

At national level the combination of a comprehensive National Forest 
Inventory with a detailed botanical survey, as exemplified in the Ghana case 
study (see Part II), provides a good basis for such strategic planning, in terms 
of geographically-related data on the distribution of many tree species and plant 
associations. At the level of Forest Management Units diagnostic sampling by 
compartments, designed to collect information to support genetic conservation 
strategies, as well as forecasts of harvestable production of timber and other 
commodities, could provide much information, to adequate levels of precision 
(often presence or absence rather than volumetric or qualitative data), at 
acceptable cost (Hutchinson 1991). 

The Malaysian Selective Management System (FAO 1989c) allows for 
siMcultural operations to be tailored to the actual condition of the forest after 
togging, taking account of environmental objectives. It is likely that, given 
responsible harvesting planning and control, largely accidental influences on 
species composition in the advanced regeneration, superimposed on site-related 
differences, will leave considerable diversity within individual forest reserves, and 
even more over the National Forest Estate as a whole. The coordinated 
management of this existing diversity might go a long way to achieving the 
objectives of dynamic conservation, linked to productive use of the forests. 
However, in some areas, carefully targeted siMcultural operations might be 
needed to conserve the genetic resources of selected populations of trees of 
socio-economic importance, or to maintain given species richness or ecosystem 
diversity. This might involve management decisions to prolong felling cycles or 
leave selected trees or patches of forest unlogged, as well as some silviculture! 
interventions, for example to release the advanced regeneration of selected 
individuals of chosen species in particular areas, as part of a coordinated national 
strategy. The prescriptions for each area to be treated would take account of its 
comparative value in terms both of production and genetic conservation. 

Some areas, particularly in the boundary zones of production forests, 
might be devoted to intensive management of non-timber forest products for 
national use or local community benefit. This could extend the overall diversity 
of management systems, with the introduction of the influence of cultural diversity 
on forest composition, while maintaining the principle of linking conservation with 
productive use. The underlying principle, coordinated in application at the 
national level, must be the management of diversity through planned 
diversification of management. 




The ecological principles underlying approaches to the sustainable 
management of natural forests, briefly reviewed in Part I, have been long 
understood. The same elements, in particular the manipulation of the forest 
canopy in relation to regeneration and growth, form components of a wide 
variety of silvicultural and management systems, from strip-shelterwood clear 
felling operations to highly selective polycyclic management. The factors 
determining the degree of success, particularly in the attempt to move from 
the small scale research or pilot-phase operations to management of entire 
forests, have been fundamentally economic, social and political forces, 
deciding the level of capability for management, including the adequacy of its 
information base, and its freedom of action to interpret and to apply 
ecological principles. 

Similarly the implications of genetic systems and genetic structure for 
the conservation of intraspecif ic variation are sufficiently established even 
though detailed knowledge of their expression in the individual species is 
almost entirely lacking. A central activity in any strategy for genetic 
resource conservation must be the gathering of that information, in respect 
both of patterns of variation across a species range and the actual mating 
system, seed dispersal mechanisms etc operating within the populations. The 
almost incalculable number and variety of investigations needed to cover all 
species of actual or potential economic importance requires rigorous setting 
of objectives and priorities for each forest area or management unit, and in 
accordance with the national Forest Genetic Resource Conservation Strategy. 

In both respects, of management and conservation, every nation and to 
a degree every forest, is unique. The choice of strategy will depend as much 
on socio-economic forces around the forest, and on national policies for land 
use and for development both within the forestry sector and in other sectors, 
as on the basic scientific principles for management and conservation. Key 
factors will include the relative importance of the forests for the export of 
timber and for the needs of local communities; their existing extent and 
distribution in relation to population density and demands for land; the 
importance of their influence on local and regional environments; the national 
and global value of their genetic resources, in terms both of principal 
economic species and of biological diversity in general; and the nature and 
degree of threat to valuable ecosystems, species, genetically distinct 
populations and genes. 

The following three case studies have been chosen to illustrate the 
principles outlined in Part I of this publication, in three contrasting 
situations, and the ways in which sustainable forest management and, more 
recently, genetic conservation, are being approached in each country. The 
starting point in all three is seen to be considerations of national policy 
and land-use but the nature and balance of the activities in progress, and the 
implication for future action and research vary greatly between them. The 
ideal basis for the development of a National Strategy for the Conservation 
of Forest Genetic Resources is a review of the forestry sector, in its wider 
context with other sectors, and the preparation of a national forestry action 
plan. This in effect has been the case in Ghana, which was one of the very 
first countries to be involved in the development of a National Plan, within 
the framework of the Tropical Forestry Action Programme (see e.g. FAO 1985b). 


The Ghana case study illustrates many aspects of the common problem of 
harmonizing genetic resource conservation with the utilization of existing 
resources for development. Timber is important to the Ghanaian economy and 
the forests 1 environmental role is important in sustaining agricultural 
production. At the same time population pressures are rising and demands from 
local communities around the forests are increasing, relative to the 
diminishing resource. The resources of the principal economic timber species 
have been depleted to a critical level in some cases. Responsive action by 
the government has provided the information base for revised policies and 
strategies now being introduced and implemented. Within these the 
conservation of forest genetic resources is seen to be largely dependent on 
national action to increase the diversity of management in both the forest and 
the forest industry sectors, and this action to depend in turn on 
international trading conditions. In view of its wide applicability this 
study has been treated at length, to illustrate the range and interdependence 
of actions. 

In contrast the Brazil case study focuses on a single large region 
within the country, within which the forest biome and its genetic resources 
are still largely intact. As a result of this the possibilities exist for 
integrated programmes of conservation and development, involving planned 
coordination of action in both protected areas and production forests, and 
multiple-use forest management, with involvement of local people. The major 
problems and challenges relate to the scale of action needed, and the 
complexities of coordinating programmes involving many administrative 
divisions and possibly even more technical and scientific bodies, generating 
large quantities of data, as contributions to what should be a single coherent 
strategy for the region. 

The Indian case study focuses down even further to a single unique area 
of natural forest. In contrast to most other tropical countries the forest 
has been under planned protection and management for well over 100 years but 
now as the result of increased population pressures, the survival of the 
natural ecosystem and of viable breeding populations of key species have been 
judged to depend on a new and more active approach to the integrated 
development of the forest resources, with involvement of communities within 
and around the forest. Both this approach and the scientific surveys and 
research needed to provide the basis for efficient conservation strategies, 
are essential elements in all three case studies. 

All three case studies illustrate the emerging principle that the 
conservation of valuable genetic diversity depends on a greater diversity of 
forest management approaches, specifically adapted to the existing diversity 
of ecosystems, species and demands on the forest. They also demonstrate the 
universal need for the rapid and efficient collection and interpretation of 
essential information on the composition and dynamics of the forests, as the 
basis for more exact systems of sustainable forest management compatible with 
genetic conservation concerns. 

To complement the above, the case study of Cordia alliodora (Appendix 
1) describes detailed studies undertaken on the biology and "functioning" of 
an important species of present-day economic importance, to illustrate the 
information needs in this regard and the utilization of data collected in 
development or refinement of genetic conservation strategies. 


6-1 The Economy 

The forestry sector in Ghana is the third highest source of export 
earnings after cocoa and minerals, and accounts for 5 to 6% of the gross 
domestic product (GDP), providing employment to about 70 000 people (Asabere 
1987). Export earnings in 1987 represented 11.4% of the total, with cocoa 
contributing by far the greatest proportion (60%). However timber is seen to 
play a key role in the economy (Frimpong-Mensah 1989). Forests cover about 
11.3 million hectares or 48% of the total area of the country (Ghartey 1990), 
and in addition to meeting all its timber requirements they supply some 75% 
of energy needs. The population of 14 million people is estimated to be 
increasing at 3% a year and population densities range from 17 persons per 
square kilometre to over 150, with an average of 63 (WRI 1990). With a GNP 
per capita of US$ 390 the country has a low-income economy, heavily dependent 
on the agricultural sector. 

The high forest zone, roughly one third of the country, supports two 
thirds of the population. The forest itself, which covered virtually all the 
8.2 million hectares of this zone at the beginning of the century, has been 
reduced to about 1.7 million ha. The principal cause of deforestation has 
been demand for agricultural land and to a considerable extent the successful 
retention of a national estate of reserved forests, in the face of such 
massive conversion, is the result of sound land-use policies. However there 
is now relatively little high forest left outside gazetted forest reserves and 
with the population doubling in 22 years, and projected to reach a total of 
37 million by the year 2 025 (WRI 1990), the area of land under agriculture 
is still increasing. 

6.2 The Environment 

The protective role of the forests was recognised from the outset, and 
particularly its influence on local climate and hydrology; and the dependence 
of the major agricultural crops, especially cocoa, on the maintenance of the 
forest cover (Gent 1929, in FAO 1985a). While the principal concern was for 
the agricultural activities within the high forest zone there is now increased 
understanding of the wider influence of the moist forests on the savanna zones 
to the north, covering 66% of the country. These include four vegetation 
types, progressively drier, from the Derived Savanna, and Southern Guinea 
Savanna zones, to the Northern Guinea and Sudan Savanna. The rainfall ranges 
between 800 and 1,200 mm/ year but has shown a decreasing trend over the past 
30 years. Most significant are the length and severity of the dry season and 
the uncertainty and erratic nature of the rains. Since the main rain-bearing 
winds come from the south west, to displace the severely dry north eastern 
winds, the rainfall in the savanna zones is influenced by the state of the 
vegetation in the south of the country, and particularly the forest cover, 
which through transpiration effectively recycles the precipitation northwards. 
Although the West African data are less complete the situation is believed to 
be similar to that in the Brazilian Amazon in regard to the links between 
forest and the hydrological cycle (Salati 1987). Increasing deforestation in 
the northern part of the country, with annual fires, has already caused 
substantial ecological degradation and increasing population pressure seems 
certain to aggravate the problems. The possibility of global warming is a 
further reason to retain as far as possible the stabilising effect of the 
remaining forest cover. 


6.3 Diversity 

The rain forest flora, both herbs and trees, changes clinally from west 
to east across the country over a distance of 300 km, with increasingly dry 
climate, and changes in soil type (Whitmore 1990). 

Ghana's biogeographic affinities are Guinea-Congo lian in the southwest, 
Sudanian in the north, with the important division of the high forest zone by 
the "Dahomey Gap" to the east, separating to some extent the forests of the 
Upper Guinea from those of the Lower Guinea. Although the forests are not 
rich in biological diversity or end em ism, in comparison for example with those 
of Cameroon, over 2^100 plant species have been recorded in the forests (Hall 
and Swaine 1981), including about 680 tree species (Hawthorn 1990a). 

6.4 Management for timber production 

As a result of an initial survey of the sector in 1908, following the 
enactment of a Timber Protection Ordinance the previous year, the reservation 
of selected areas of forest was proposed but met with opposition from the 
local chiefs (FAO 1985a; Asabere 1987). However by 1939 some 1.6 million ha 
had been reserved in the high forest zone, surveyed, demarcated and 
constituted under the Forest Ordinance. About 70 per cent is made up of 
production reserves, which are the source of future supplies of timber, as the 
unreserved forests still remaining are not expected to survive as timber 
production areas beyond the end of the century. The remainder of the 
permanent forest estate is reserved for protective purposes, a proportion of 
this area being made up of plantations. There are 13 wildlife reserves 
covering a total of nearly 1.2 million ha, of which four are in the high 
forest zone and nine in the savanna. Within the high forest zone the areas 
reserved strictly for protective purposes are those which are most 
inaccessible and which cannot be easily or safely logged (Tufuor 1990a), and 
are not therefore necessarily representative of the range of plant communities 
in the zone. 

The tropical high forest zone, which is the main source of timber, shows 
a wide range of variation 'in plant communities, which have been broadly 
grouped into four ecological types: the Wet Evergreen Forest (Cynometra - 
Lophira - Tarrietia association), the Moist Evergreen Forest, the Moist Semi- 
Deciduous Forest and the Dry Semi-Deciduous Forest (Hall and Swaine 1981). 

Three main approaches to the management of the forests have been made 
over the past 40 to 50 years, namely the Tropical Shelterwood System (TSS), 
enrichment planting, and the Modified Selection System (MSS) which was 
introduced in 1956, and has been the main system employed throughout most of 
the zone, particularly in the Moist Semi-Deciduous Forests, until 1970. The 
main features of the MSS were stock-mapping of all "economic" trees over 7 ft 
(2.1 m) girth and selective felling on a 25-year cycle. The yield was 
regulated by area, with minimum girth limits set according to the species 
class. Felling was based on the stock maps, starting with the largest trees 
and going down until the prescribed yield had been achieved. However trees 
below the minimum girth limits were never permitted to be felled, even if 
there proved to be insufficient trees above the limits to make up the 
prescribed yield. Moreover there was a stipulation that the remaining stock 
should be left well distributed over the compartment, so that in some cases 
a tree well above the limit might be left as the only one in that part of the 
area (Asabere 1987). This relatively simple system of control, although based 


on incomplete data on forest dynamics, was reasonably solid and ecologically 

In the late 1960's the felling cycle of 25 years was increasingly 
criticised by timber concessionaires as being too long and resulting in the 
accumulation of over-mature trees subject to decay. As a result it was 
decided in 1970 to permit the felling of all economic trees above 3.4m girth 
or above 2.1 m girth (depending on species class) and to reduce the felling 
cycle to 15 years. The assumption that all trees over these dimensions were 
over-mature was not based on systematic studies, and such an investigation 
later revealed that the trees in the highest classes with girths of 3.4 to 4.0 
m were far from being over-mature. Rather than reducing waste of timber it 
appears that the blanket application of salvage felling throughout the 
production forests, irrespective of stocking, encouraged more wasteful and 
less efficient practices, with large numbers of logs being left to rot in the 
forest (Asabere 1987). 

The greater intensity and frequency of logging, concentrated on the few 
preferred species, with the removal of all the largest trees (whereas 
previously some had been allowed to remain) was bound in time to have an 
increasingly dysgenic effect on the populations. At the same time the 
tendency to increase the size of gaps in the canopy, as the result of removal 
of all large "economic" trees regardless of their proximity or situation, must 
lead to greater structural changes, and to a stronger shift in species 
composition towards the pioneer species, and with greater danger of dense 
climber tangles developing. There would therefore be a predictable adverse 
impact both on the genetic resources of the species being logged and in terms 
of overall diversity, as the prescriptions applied throughout the forest. 

From the mid-1 970 f s to the early 1980's the forestry sector suffered a 
marked decline both in the production of logs and in the total value of timber 
exports. This accompanied general economic problems, related to the 
overvalued currency (cedi) and linked to the poor state of logging, transport 
and sawmilling equipment, lack of spare parts, inadequate investment in roads 
and infrastructure generally, and inadequate funding of the Forestry 
Department. From 1983 a series of macro-economic and sectoral reforms, 
followed by renewed investment in equipment and infrastructure, assisted by 
multilateral and bilateral funding, led to a progressive recovery in log 
production and export revenue, which had been substantially achieved by the 
end of 1987 (Frimpong-Mensah 1989). 

With the restoration of export-led logging and milling operations there 
was a danger that exploitation would override concerns for sustainable 
management, centred on sustained yield of timber. The initiation of the 
Tropical Forestry Action Plan (TFAP) in 1985 had promoted the concept of joint 
sector reviews, with assistance from several donor agencies, and in Hay 1986 
a TFAP review was started in Ghana, led by the World Bank, with direct 
collaboration and assistance from FAO, CIDA (Canada) and ODA (U.K.). 
Following the studies, project preparation, appraisals and negotiations the 
Forest Resource Management Project was initiated in March 1989, with funding 
from the World Bank, Denmark and the U.K. This provided the opportunity for 
the Government of Ghana to undertake a comprehensive review of the forestry 
sector, linking conservation to production objectives, and related policy 


6.5 Policy; linking production and conservation 

The 1948 forest policy provided a broad basis for both production and 
conservation objectives, if interpreted in that spirit. It included specific 
reference to the maintenance of the permanent forest estate, the protection 
of environmental values, management for sustained yield, research with 
emphasis on ecology and silviculture, public education, staff training and 
optimal land use (Kese 1989). Existing legislation (principally the Forests 
Ordinance (1927) and associated Forest Reserves Regulations; the Concessions 
Act (1962); the Timber Operations Decree (1972); the Forest Protection Decree 
(1974); the Trees and Timber Decree (1974) and the Timber Industry and Ghana 
Timber Marketing Board Amendment Decree (1977) with some provisions of the 
investment Policy Decree) provided a suitable basis for policy application. 
The success of the original forest reservation policy was strikingly revealed 
by Landsat imagery which showed clearly defined reserve boundaries which had 
been generally respected, in contrast to the surrounding deforestation 
(Hawthorn 1989). Revision of the National Forest Policy, completed early in 
1992, further strengthened environmental provisions relating to soil and water 
resources, with specific references to the conservation of flora, fauna and 
biological diversity, and to sustained yield of non-timber as well as timber 
resources. This implied due attention to in situ conservation of genetic 
resources . 

At the same time as the strengthened concern for the environment and 
biological diversity there are the increasing demands on the economy and on 
land resources from the expanding population. Revised and strengthened inter- 
sectoral policies, with appropriate sectoral strategies, are essential to 
secure the appropriate reconciliation of conflicting production and 
conservation demands. Grut (1989) from a study of the forestry sector in 
Ghana and in Guinea, has shown that on an analysis of timber production alone, 
the most profitable option would be to take out all merchantable timber (i.e. 
to well below the normal minimum diameter limits set by management) in one 
felling. Calculations used to reach this conclusion did not consider the 
substantial, but usually non-market, values of non-timber forest products, and 
the severe impacts on forest structure, ecology and species composition that 
might result. 

6.6 Non-Timber Forest Products (NTFP) 

The role of non-timber forest products (NTFP) is critically important 
to the long-term conservation of the forest resources and their genetic 
diversity. A detailed study undertaken in the context of the forest Resource 
Management Project in Ghana (Falconer 1991) shows that NTFP's form the main 
link between the reserved forests and the communities living immediately 
around it. The study revealed the importance of a wide variety of products 
in human nutrition, medicines, materials for house building, household and 
agricultural implements, fuelwood, fodder, and trade and processing activities 
based on NTFP's. For house building materials the most important qualities 
sought were durability and insect resistance, for the principal building poles 
in particular. These qualities are commonly associated with denser and harder 
wood more frequently found in slower-growing species (usually non-pioneer 
species). It was evident that many chiefs and elders in the communities 
linked the value of the forest strongly to their traditional collection of 
materials for house building. 


Forests were also found to be very highly valued for medicines. All the 
people interviewed used plant-based medicines and 80% relied on them 
exclusively. Most of those commonly used were gathered around the village and 
in fallow areas of secondary forest, rather than in the forest reserve. 
Nevertheless the forest itself was seen as an important source of medicines 
by the majority of those interviewed. 

For many households the gathering, processing and trading of forest 
products provided an important source of supplemental income, especially in 
the periods of less agricultural activity or when there is an acute need for 
cash. For some of them NTFP's provided the main source of income, either 
directly or as a source of materials used for production equipment and raw 
materials used in off-farm processing activities. 

Forests were the main source of traded NTFP's in the region although in 
some areas and for some products fallow lands were important sources of 
supply. In many communities people claimed that increased land clearance for 
agriculture, increased bush fires and degradation of the fallow environments 
(secondary forests) had depleted the NTFP resources so that collectors and 
processors had come to rely more heavily on the forest reserves. For many 
products, including bushmeat, the study revealed the importance of the fallow 
areas as sources of supply. The degradation of the secondary forest was seen 
as a great problem by many communities and more and more the forests remaining 
in reserves are likely to become important sources of NTFP's. 

Flexible forest management systems which can meet local needs as well 
as those of the timber industry offer the best prospect for long-term security 
of a wide range of tree and plant species. There is also scope for the 
propagation and cultivation of some species in buffer zones around the forest, 
and provided that the source of seed or selected clonal material is taken from 
the local populations in the forest this could be an acceptable aspect of in 
situ conservation for the species concerned. 

6.7 Forest Revenue Systems 

A major problem in conservation and sustainable management in many 
countries has been the apparently low rate of economic return from the natural 
tropical forest (Mergen and Vincent 1987) despite the evident real value of 
the resource (Leslie 1987). This is frequently due to failure to capture 
appropriate levels of revenue from timber (Repetto and Gillis 1988). Grut 
(1989) shows this clearly in respect of Ghana: both the levels of royalty set 
and the effectiveness of collection of the fees due have hitherto been 
extremely low. Even with the very low level of stumpage demanded the revenue 
actually collected was only one sixth of the amount due (Grut 1989). The very 
low level of royalties on timber in Ghana, and the need for substantial 
increases, particularly for the most valuable and marketable species, had been 
recognised earlier (FAO 1985a). Substantial increases have now been 
introduced and others are expected to follow. Assuming that these are also 
implemented to raise average stumpage fees to about 10% of weighted average 
FOB log export value by 1994, with increased effectiveness in revenue 
collection rate to 50% by the same date, revenue would be sufficient to meet 
the entire costs of the Forestry Department, and not only the costs of 
management of the production forests (Grut 1989). 


Both the size and length of tenure of timber concessions are key factors 
in determining the interest and capability of the concessionaire in sustain** 
able management. The fragmentation of concessions in Ghana was criticised 
earlier (Asabere 1987) and suggestions made in the context of the Forest 
Resource Management Project to set a minimum size of 10 000 ha and duration 
of 50 years are under consideration by the government authorities. Related 
to these suggestions are consideration of more strongly market-determined 
allocation of concessions, to secure the best value possible, and a more 
significant level of concession rent. The latter could be particularly 
influential in securing conservation objectives, including in situ conserva- 
tion of genetic resources. It has been suggested, for example, that 
substantial concession rents would be set, based on a formula derived from 
weighted FOB values, less costs of logging etc and an adequate level of 
profit. The concessionaire could subsequently claim back a proportion of the 
fees paid, in respect of costs incurred in sustainable management and con- 
servation activities. The latter would then be seen by the concessionaire as 
a revenue-earning opportunity (Grut 1990). Such reforms of revenue levels and 
collection can also potentially improve recovery and reduce waste of timber, 
both in the forest and in industrial processing (Asabere 1987; Chachu 1989). 
The Government of Ghana has set up a special Committee to evaluate measures 
to improve forest management and productivity in the timber industry, and the 
Committee is expected to review the size and related aspects of timber 

Ghana had earlier introduced differential royalty rates for 50 timbers. 
The new royalty fee structure being set involves considerably higher increases 
in respect of the species in highest demand, and therefore in danger of 
depletion, than for those currently neglected. This has important implica- 
tions for the conservation of the genetic resources of the main economic 
species. However, such action cannot be effective in isolation, as many 
lesser-known species have lower natural durability and therefore require 
special care both to ensure that the logs are not left lying in the forest and 
for appropriate seasoning and preservation treatments. Others which may have 
more durable but hard and possibly siliceous timber require special processing 
equipment. The securing of an adequate market return for such lesser-known 
species may be assisted by local processing (Asabere 1987; Ofosu-Asiedu and 
Ampong 1990; Parant 1990) for which investment in equipment, infrastructure 
and training may be needed. Ghana is promoting further processing of timber, 
and log exports as a proportion of total export volumes have been falling from 
63Z in 1988, to 53% in 1989 and 42% in 1990. However in 1989 less than 6% by 
value of Ghana's forest products exports were processed beyond air-dried 
timber (ITTO 1990). 

There are also important forest management and silvicultural consider- 
ations, as well as ecological concerns, related to the promotion of harvesting 
of lesser-known species, and these are reviewed below. However the implica- 
tions of the results of the forest inventory carried out clearly indicate the 
need to make maximum sustainable use of the neglected timber species, while 
reducing pressure on the stocks and the genetic resources of the principal 
economic species. This is in line with the new recognition that harvesting 
convenience and the demands of the wood industry alone should not be allowed 
to determine forest management (Chachu 1989) but that royalty rates, girth 
limits and other objectives and intensity controls should reflect the 
ecological differences between species in their regeneration and growth, as 
well as ultimate timber values (Hawthorn 1990b). There is already abundant 
evidence of the impermanence of classifications based solely on market 
acceptability and valuation of timber. 

6.8 Forest Inventory 

The first survey of forest resources in Ghana was made in 1908 and the 
first National Forest Inventory in 1947, covering 1 290 square miles of forest 
(Logan 1947). The main focus of this and subsequent inventories in the period 
1952 to 1973 was the then currently merchantable species, of which only 26 
"economic 11 species were listed compared with 334 "secondary timbers" (FAO 
1985a). In 1985 a new National Forest Inventory was started, primarily also 
to provide an estimate of the total commercial log volume, in view of concerns 
that the demands of the rehabilitated timber industry would exceed the 
sustainable supply from the forest. However broader aims were included, to 
supply the wider information needed for sustainable management, and to assess 
the biological productivity and ecological status of the forest, including the 
collection of data on non-timber forest products, and on non-woody plants and 
fauna (Adlard 1990). 

To meet the primary objective stratified random sampling was done over 
546 000 ha in 43 forest reserves. Stratification was based on the ecological 
survey and classification by Hall and Swaine (1981). In addition to 
measurements of tree diameters, for volume estimations, assessments were made 
of form and quality, together with crown classification, observations on 
logging, burning etc. About 420 tree species were recorded, and grouped into 
three classes according to their current market acceptability, size and 
frequency of occur rence:- 

Class I: Species exported from Ghana at least once during the period 1973- 
88, including all the main economic species plus some lesser- 
known species being actively promoted for export (total 66 
species) . 

Class II: Species attaining 70 cm diameter (marketable size) and present at 
a frequency of at least 1 tree per km 2 , although not previously 
exported (total 58 species). 

Class III: All remaining species, not considered to have potential for 
timber production. 

All the data were stored on computer and constitute an important 
database on species occurrence and distribution (Wong 1989). 

The gross national standing volume of timber of exploitable size was 
estimated as 102 million m 3 , concentrated mainly in the Class I species. 
However the bulk of this volume was found to be made up by the less desirable 
species, while the volume of the traditional economic species was very limited 
and heavily dependent on one mainstay, Triplochiton scleroxylon, which 
although in constant demand is not intrinsically of high value. It appears 
that current exploitation of most "economic" species is unsustainable and that 
they have a very limited resource life (i.e. the period during which the 
existing supply of trees of exploitable size will be exhausted). The valuable 
Meliaceae (Entandrophragma angolense, E. cylindricum, E. utile, Khaya 
grandifoliola, K. ivorensis, K. anthotheca), as well as Milicia (syn. 
Chlorophora) excel sa, are likely to be exhausted within 2 to 3 decades at 
current levels of felling. Among the most valuable species Pericopsis (syn. 
Afrormosia) elata is estimated to have a resource life of 3 years or less 
(Alder 1989). The calculation of the species resource life can only be 
approximate, based on the division of the existing resource by the rate of 


extraction, and if with a diminishing resource the rate of extraction were 
allowed to rise, the resource life would be even less. 

The implication is clear: the future of a viable timber industry in 
Ghana is dependent on a substantial and rapid move into the marketing of 
presently under-utilised species, in Class I. This could be achieved by the 
promotion of perhaps 10 to 15% of the 50 odd presently under-utilised species 
listed, with maximum reliance on those well represented in the forest, 
alongside the sustained logging of Terminalia excelsa. If this were done with 
effective monitoring and controls to limit the logging of the principal 
economic species to sustainable levels, to allow their populations to recover 
between logging cycles, there need be no long-term danger to the genetic 
resources of these species. However their future existence is clearly 
dependent on positive action in this regard. 

Growth data for the estimation of increment rates were based on 
measurement of about 11 000 trees in 256 permanent sample plots. The 
development of a forest growth simulation model (GHAFOSIM) for Ghana forests 
is in progress (Alder 1989; 1990) and will be assisted by the progressive 
accumulation of data from the greatly expanded number of permanent sample 
plots, in which all trees of all species over 10 cm diameter (dbh) are 
measured. This will provide a much more complete understanding of the forest 
dynamics. It is evident that no single management strategy will serve 
adequately for all the forests but that for efficient linking of production 
to conservation the management of individual reserves or concessions will need 
to be decided separately, within an overall picture of the genetic diversity 
in the forests as a whole. 

6.9 Setting Priorities 

Conservation of the principal economic tree species is clearly a high 
priority, and is seen to be dependent on developing the markets for presently 
lesser-used species. However conservation of intra-specific variation in 
these species is also of greatest concern, including the maintenance of viable 
populations of genetically distinct provenances. In the absence of direct 
evidence of provenance differences from field trials and/or isozyme studies 
the best guide is likely to be patterns of environmental variation. These may 
be seen directly from environmental variables or indirectly from patterns of 
vegetation. The way in which population-level variation in individual species 
reflects patterns in the community as a whole is apparent in many cases 
(Okafor 1975; Hall and Swaine 1981). The assumption that this occurs is 
recognised in the botanical survey being carried out in association with the 
on-going National Forest Inventory (Hawthorn 1991). 

The aim of the botanical survey is to develop a computerised database 
of the distribution of vascular plants in the forests of Ghana, thereby 
securing clearer definition of the substantial f loristic variation within each 
of the forest zones recognised by Hall and Swaine (1981). This will reveal 
patterns of intra-specific variation at local as well as national level, and 
the relationship between them, and thereby assist in designing strategies for 
the conservation of economic plant species. 

The survey gives particular attention to patterns of distribution of 
rare plants and trees as a direct aid to their conservation. The importance 
of combining taxonomic, ecological and choro logical information is recognised 
in the study, in order to set priorities for conservation of selected areas. 


Preliminary analysis (Hawthorn 1991) has defined three sets of conditions 
under which rare plants are most likely to be found. One of these is among 
the dry forest species in or near the coastal or Afram plains. Such West 
African Dry Forest is now of very limited distribution and some species are 
unknown elsewhere except for occurrences in East Africa, while others are 
endemic to Ghana. 

Special consideration should also be given to fire prone reserves in the 
Dry Semi-Deciduous zone, where the forest plants are especially at risk. The 
threat of fire to the genetic resources is particularly acute where the 
population of mature seed-bearing individuals has been severely reduced, seed 
years may be infrequent and a fire following seed fall or seedling 
establishment can be catastrophic. This may be the case with Pericopsis 
elata, particularly at the fringe of the species 1 range. 

The botanical survey is also developing an objective rating system for 
the classification of plant species in terms of priority for conservation. 
This will be used to assist the selection of areas, for example within 
production forest reserves, where forest management should take particular 
account of the importance, and the nature, of conservation objectives. The 
classification being developed will be based essentially in the information 
on distribution, derived from the earlier surveys of Hall and Swaine (1981), 
together with the more general observations in the Flora of West Tropical 
Africa for the wider picture, the substantial herbarium collections at Legon 
and Kew (Royal Botanic Gardens) and the new data from the botanical survey. 
This will provide a basis for highlighting the "genetic hot spots" in Ghana's 
forests, as concerns ecosystems, species and genetic resources of species 
targeted for conservation. 

Most categories in the rating system are concerned with degrees of 
rarity, both within Ghana and in the wider contexts of Upper Guinea (the 
region within which the contribution of the Ghanaian flora is likely to be 
most significant), West Africa and Africa as a whole. However for the 
conservation in situ of the genetic resources of species of known economic 
importance a separate category is recognised. This includes species that may 
be fairly common or widespread but heavily exploited, and requiring individual 
attention and control, particularly in regard to the conservation of 
intraspecific variation. This category includes all the heavily utilized 
timber species, canes (rattans) and other species of known economic value 
(e.g. Thaumatococcus daniellii, a source of sweetening agents and other 
potentially commercial organic compounds). Under this classification species 
are allocated a "Red Star" rating if, as in the case of sought-after timber 
trees, a significant proportion of the mature individuals are affected by 
exploitation pressures (e.g. species with a short resource life, as listed 
above); or if it is advisable to "flag" the occurrences of a species in the 
forest for conservation of specific provenances. 

An additional and immediate practical outcome of the botanical survey 
will be a report on the current status of existing Protected Areas and of the 
degree of protection afforded within the production reserves, with 
recommendations regarding forest management in these latter ones, particularly 
in the more sensitive areas and those of exceptional biological value. This 
will assist the development of comprehensive approaches to reconcile the 
production and conservation objectives, within the Forest Management Units 
(FMU) now being established. 

6.10 Management and Harvesting 

The Modified Selection System, described briefly above, had apparently 
succeeded in maintaining satisfactory levels of timber yield but at the 
expense of the stocking of the main economic species. Data from the on-going 
inventory and intensified permanent sample plot network will permit the future 
development of detailed and flexible management plans separately for each 
management unit (single reserve or large concession area) using simulation 
models to be derived from continuation of the work initiated on the Ghana 
Forest Simulation Model (GHAFOSIM). Such an approach, capable of 
incorporating information from the botanical survey, and ecological studies 
on regeneration in relation to logging intensity, pattern and methods, offers 
the best prospect for combining genetic conservation with production, both at 
the local and the national level. The possibilities for effective integration 
of conservation and production objectives in forest management plans have been 
strengthened by the introduction of Forest Management Units (FMU), which group 
the existing production reserves into units of about 50 000 hectares. Each 
FMU will be subject to a management plan, incorporating the recommendations 
on the conservation of biological diversity and genetic resources, emerging 
from the botanical survey. 

As an interim measure a forty year felling cycle has been imposed and 
stock surveys and stock mapping of all compartments are being undertaken to 
determine the permissible yield. This is calculated for each species in each 
compartment by a simple formula which takes into account the number of trees 
above the minimum felling limit, and those in the diameter class immediately 
below, which will form the next crop. The formula also allows for the 30% 
retention of the mature crop and 20% mortality in the residual population 
intended for the next felling cycle (Ghartey 1990). Provided that logging 
controls are enforced to prevent unnecessary damage to advance growth and to 
the conditions for regeneration (see below), and that sufficient seed bearers 
of the important economic species are left where needed, this interim regime 
should be sufficiently conservative to prevent further degradation of the 
genetic resources of the forest. However this is ultimately dependent on 
action to promote the lessor-used species, in order to maintain acceptable 
levels of harvest while conserving the remaining genetic resources of the most 
valuable and most depleted populations. 

In November 1990 the Ghanaian authorities imposed Forest Improvement 
Levies on six species of log exports, complementing export bans already in 
place on 19 other species, including the valuable Meliaceae. The levies are 
intended to conserve the species concerned and to provide logs for domestic 
processing (ITTO 1991). The success of these moves and of plans for greater 
reliance on the lesser-used species will have a major influence on the 
possibilities for future management. 

The impact of increased logging on the genetic resources of presently 
under-utilised species will also need to be considered in respect of each 
forest reserve and management unit, in the preparation of management plans. 
However the inventory shows that overall over 40% of the potential timber 
harvest from species in Class I (i.e. those that have been exported at all) 
is made up by four species still in abundant supply in the forest (Ghartey 
1989). A strategy based on the promotion of these up to the limits of the 
annual allowable cut, together with lesser contributions from some other 
presently neglected species, should be consistent with more balanced 
management of the total genetic resources of the forest, provided that 
satisfactory conditions for regeneration are maintained. 

6.11 Regeneration and Silviculture 

Under the TSS system introduced in 1946 the attempt was made to 
influence regeneration of the relatively few desirable species by manipulation 
of the canopy and cleaning operations but it was judged unsuccessful and 
abandoned in the mid-1960 ! s. Since then, apart from attempts at artificial 
regeneration through enrichment planting, the attempts to influence the crop 
through silvicultural operations under the MSS were directed to favouring the 
young trees in the 1 to 5 ft (0.3 to 1.5 m) girth class by improvement 
thinnings (Asabere 1987). Nevertheless from the viewpoint of long-term 
sustainable management, and the conservation of the genetic resources of the 
principal economic species, an understanding of the effect of logging on the 
species composition of the regeneration is essential. 

In association with the National Forest Inventory, regeneration studies 
were undertaken in logged and adjacent unlogged forest within the Bia South 
reserve (Hawthorn 1990b), to assess the various effects of logging operations 
on the pattern of regeneration of the tree species. A secondary objective was 
to describe the state of the vegetation as a whole following logging, as 
background information on the biotic environment of the developing trees. 
Since the forest had not been exploited previously, and the only control 
exercised had been through the minimum girth limit, with all trees above that 
limit eligible for removal, the logging intensity in some compartments had 
been very high. However this effect was mainly confined to the most desirable 
species such as Khaya ivorensis and Entandrophragma utile. Large trees of 
less desirable species had been left. Logging had been done in the three 
years immediately prior to the study. 

The main subdivisions of the species "guilds 11 recognised were between 
true pioneer species and under storey pioneers (small trees and shrubs) on the 
one hand, and between non-pioneer light demanders and shade-bearers on the 
other, although it was recognised that these divisions may be arbitrary 
separations in what is a continuum of preferences. The report on the study 
(Hawthorn 1990b) is detailed and reveals a complex situation. Nevertheless 
some clear conclusions were drawn regarding the effects on early seedling 
regeneration e.g:-4 

(i) the overall species diversity is not likely to be adversely 
affected by well-controlled logging, although the balance of 
species is expected to change unless special steps are taken to 
prevent this. The trend will be declining levels of regeneration 
of some valuable (non-pioneer) timbers (e.g. Entandrophragma 
spp, Khaya spp) and greater relative abundance of others (e.g. 
Triplochiton scleroxylon and Terminalia spp); 

(ii) large canopy gaps show very poor regeneration of preferred 
species and low species diversity; 

(iii) regeneration of most timber species, particularly non-pioneers, 
requires the retention of parent trees evenly dispersed 
throughout the forest, for example in small unlogged pockets 
within logged compartments; 

(iv) small gaps and skid trails helps to add to overall species 


Clear general guidance can therefore be given regarding strict logging 
controls, to keep road widths and the area of loading bays to a minimum, to 
avoid the creation of large canopy gaps and to ensure the retention of seed 
bearers evenly spread through the forest. To avoid damage to regeneration 
logging should be restricted to a single short period. With such provisions 
there seems little danger of any sudden loss of genetic resources of the main 
economic species, and good possibilities to retain a high level of species 
diversity at least of a wide range of Class I timbers. 

The most serious threat to genetic resources is undoubtedly fire, and 
associated studies of fire damage suggest that logging, especially in the 
drier forest types, may render the forest more liable to fire (Hawthorn 1989). 
The recommendations for controlling the intensity of logging are therefore 
even more important in fire prone areas. 

Although the results of these studies are both valuable and encouraging 
more widespread and comprehensive data will need to be gathered from the 
expanded series of permanent sample plots, which could also be the sites for 
more detailed studies in population dynamics, pollination and seed dispersal 
systems and agents, and other ecological processes (Adlard 1990). These could 
provide a scientific basis for possible future silvicultural action to 
influence the regeneration and development of selected populations maintaining 
high levels of genetic diversity. 

6.12 Reproductive Biology 

While much valuable information on genetic systems may be gained from 
well-documented pheno logical observations, more detailed and systematic 
studies of the key economic species (e.g. Entandrophragma spp, Khaya spp, 
Milicia excelsa, Pericopsis elata) which are in urgent need of action to 
conserve their constituent populations, are strongly advisable. Data on 
reproductive biology, used in conjunction with information on species 
distribution and variation patterns in individual species, can assist in 
determining both the siting of in situ conservation areas and sampling 
strategies for complementary ex situ conservation. The principles involved 
are summarised in e.g. Bawa and Krugman (1991). Their application is also 
illustrated in the case study on the reproductive biology and genetics of 
Cordia alliodora, in Appendix 1 . 

6.13 Integration and Security 

The database developments associated with the forest inventory and the 
botanical survey provide a framework for planning the integrated management 
of genetic resources, both within fully protected areas and in production 
reserves. They also provide a basis for monitoring future changes in the 
forest under the influence of prescribed harvesting and management regimes. 
However there will be a need for high-level decisions on the balance to be 
struck between timber exploitation and the conservation of biological 
diversity, to ensure that the necessary "trade off 11 between short-term 
financial costs of conservation (including revenue and profits which may be 
foregone) and the longer-term social, economic and environmental benefits are 
settled in the national interest, and where appropriate with international 
assistance. This is likely to be needed particularly in enabling the 
essential changes in the timber trade, to largely replace current dependence 
on the few principal economic species, with very limited resource life, by the 
promotion of lesser-used species, with greater reliance on local processing 
and the export of components and finished products. Action in these areas of 


international trade, linked to investments in local provision of physical 
capacity and human resource development, will determine the feasibility of 
forest management systems linking sustainable production and genetic resource 


The Amazon basin as a whole contains the largest single extent of 
tropical forest in the world, of which approximately 60% (some 350 million ha) 
lies within Brazil (Dubois 1991). About 280 million ha are closed forests 
with the potential to produce industrial timber (FAO 1981). At the same time 
the forests are almost certainly the richest in the world in terms of total 
biological diversity, and cover a wide range of forest types. The possible 
contribution of the potential production forests to the conservation of such 
diversity and of forest genetic resources is therefore very high. 

According to Cochrane and Sanchez (1982) some 93% of the soils of the 
Amazon Basin have major fertility deficiencies and although perhaps half of 
this area is underlain by soils of good physical properties, and might be 
capable of sustainable agriculture or pasture use, given substantial inputs 
of nutrients and under careful management, they are susceptible to rapid 
degradation after clearance of existing forest. In addition studies have 
indicated the interdependence of the local hydrological cycle and the forest 
cover (Salati 1987) and it has been suggested that to safeguard local and 
regional climate, until a fuller understanding of this relationship is 
achieved, about 80% of the area should be kept under forest cover (Prance 

Despite some recent immigration into the Amazon region from elsewhere 
in Brazil the human population is still low relative to the total area, with 
over 40% of the Brazilian population concentrated in some 10% of the national 
territory in the south eastern region. Nevertheless the pressures on 
currently forested land in Amazonia will inevitably increase and it is 
essential to develop sustainable systems of land use which will be compatible 
with the need to retain a forest cover over substantial areas for its 
environmental influences. The opportunity still exists to develop patterns 
of land use which optimise the sustainable management of natural forest, with 
the simultaneous conservation of the rich biological diversity of the region. 
This requires the integrated planning and development of the systems of fully 
protected areas and production forest reserves, with both systems contributing 
appropriately to both conservation and development objectives. 

7-1 Legal framework 

The Forestry Law (1965) makes provision for the creation of reserves 
both for protection and for production objectives. Under Article 5 it decrees 
the creation of:- 

(a) National, State and Municipal Parks and Biological Reserves, to 
protect exceptional natural features, flora, fauna and scenic 
values, and to reconcile this protection with educational, 
recreational and scientific objectives. 

(b) National, State and Municipal Forests, for economic, technical or 
social objectives, including possible afforestation. 


In specific reference to the Amazonian forests the Forestry Law calls 
for the utilization of the forests to be placed under technical constraints 
and management plans. 

In addition the Fauna Protection Law of 1967 makes provision for 
National, State and Municipal Biological Reserves, and the Law for a National 
Environmental Policy of 1981 covers the establishment of Ecological Stations 
and Environmental Protection Areas. The Federal Constitution of 1988, in 
reference to the rights of the people to an ecologically stable environment, 
requires the public administration to ensure proper ecological management of 
ecosystems and preservation of the diversity and integrity of the genetic 
heritage of the country. It further declares the Brazilian Amazon forest, the 
Atlantic forest, and three other regions, as a national estate, placing 
limitations on their utilization, to guarantee preservation of the environment 
(Schubart 1990). 

The principal legal category of reserves which offers the best 
possibility to combine management for timber production, and other forest 
products, with the conservation of genetic resources in situ is the National 
Forest. This corresponds approximately to the IUCN Category VIII (Multiple 
Use Management Area/Managed Resource Area) and has as its primary objectives 
the sustainable harvesting and management of the forest, principally for 
timber and other forest products, together with maintaining the protective 
functions of the forest over water resources, and the conservation of 
biological diversity, flora and fauna, insofar as this is compatible with the 
principal production objectives. Appropriate scientific and technological 
research, and environmental monitoring, are also envisaged. The basic 
criteria for the selection of an area as a National Forest are the potential 
for sustainable production of timber and/or other forest products. If rare, 
endemic or endangered species are present that might be adversely affected by 
the production operations it is suggested that another more restrictive 
category of reservation would be more appropriate (Padua 1989). There has 
been a marked increase in the number of National Forests in Brazil in the past 
three years, particularly in the Amazon region, where Schubart (1990) lists 
24 such Forests up to the end of 1990, covering a total area of over 12.6 
million hectares. 

Extractive Reserves are the other major category where direct productive 
uses of the forest may be combined with conservation of genetic resources. 
According to Padua (1989) these are intended to meet the needs of social 
groups who are dependent on the gathering of forest products for their 
survival, and who harvest in a sustainable manner according to traditional 
practices and in conformity with pre-established management plans. Secondary 
objectives include the conservation of biological diversity, and possible 
contributions to scientific and environmental education and monitoring. On 
this interpretation the harvesting of timber is specifically excluded. The 
traditional products of extractive forest management, as the term is used in 
Brazil, have been primarily rubber and Brazil nuts, although a wide variety 
of other fruits, fibres etc may be included. 

To provide a legal basis for the concept the former National Institute 
of Colonization and Agrarian Reform (INCRA) issued a Decree in July 1987 which 
provided guidelines for the settlement of Extractive Reserves as a mode of 
agrarian reform in the Amazon region (Allegretti 1990). The Decree used the 
concept of land use concessions, ceding use of the Reserves from the State to 
the practitioners during a minimum period of 30 years, with specific 


regulations on land use practices. It also established a mechanism by which 
the State could mediate between the inhabitants of the Reserve and outside 
economic interests. Administration of Extractive Reserves on this model is 
by a group elected by the local inhabitants, either in the form of a 
cooperative or an association, thereby avoiding subdivision of the land into 
separate private holdings. According to Schubart (1990) the legal status of 
Extractive Reserves has yet to be finally decided but action has been taken 
under the National Environmental Policy (January 1990) to create Reserves of 
this nature. Insofar as some Extractive Reserves are being created inside 
National Forests the legal basis for combining their traditional practices 
with some selective harvesting of timber may be envisaged. 

The principal categories of reserve forming the fully protected area 
system in Brazil include National Parka (IUCN Category II), Ecological 
Stations, Ecological Reserves, Biological Reserves (all equivalent to IUCN 
Category I), and Environmental Protection Areas (similar objectives to IUCN 
Category V but including specific reference to biological diversity). All of 
these have significance in the conservation of ecosystems, species and genetic 
resources. In addition Wildlife Refuges and Game Reserves might provide 
incidental protection to plant genetic resources. 

7-2 Setting Priorities 

The First International Seminar on Tropical Forest Management in Brazil, 
in 1985, in stressing the "need to maintain and preserve biological diversity 
in any undertaking in the Amazon", drew particular attention to the importance 
of a better understanding of forest dynamics and the interactions in processes 
such as seed dispersal, pollination, regeneration systems etc (Siqueira 1989). 
It also stressed the importance of typological classification and zoning of 
forests for production and conservation objectives. The principles involved 
have been discussed earlier, and in particular the vital necessity for 
gathering data on variation and variation patterns in species and populations, 
and on autecology and breeding biology, as a basis for action towards in situ 
conservation of genetic resources of priority species. Compared to some other 
tropical regions in the New World the Brazilian Amazon has a long history of 
botanical surveys and substantial holdings in several herbaria (Daly and 
Prance 1989). Nevertheless the sheer scale of an inventory needed to 
establish and interpret patterns of distribution of communities, major tree 
species and their probable intraspecif ic variation in relation to the existing 
environmental conditions and past history, is vast. 

The total area of Amazonian forest within Brazil in all of the reserved 
categories is probably around 5% of the total forest. While the location, 
individual size and shape of Protected Areas are more important than overall 
percentage figures there is clearly scope for the selection of a substantial 
number of additional reserves, given the limitations of the soils and the 
acknowledged environmental importance of forest cover. At the same time, for 
long-term security and in the face of increasing population pressures, the 
development of multiple-use management areas (predominantly National Forests 
and Extractive Reserves) that can contribute to the conservation of genetic 
resources in situ and complement action in fully protected areas, will be 
necessary. Since most of the Amazonian forest biome is still largely intact 
the possibility exists to plan the networks of production reserves and fully 
protected areas in an integrated manner, to maximise the effective use of the 
land and the resources devoted to management and protection. This is an 
important aspect of the zonation referred to by Padua (1989). 


Vegetation mapping in Brazil has a long history but has been complicated 
by the use of a variety of classification systems. Knowledge of the Amazon 
was significantly advanced by the use of radar imagery in "Projeto RADAM" in 
the 1970 v s and although the maps produced were purely physiognomic they were 
extremely detailed. This information was used for the most recent vegetation 
map of the Amazon (Prance and Brown 1987) which recognises four main sub-types 
of the rain forest on "terra firma" (about 53% of the region) as well as 
several types of seasonal transition forests, savannas and savanna woodlands, 
forests on white sand soils, several types of inundated forests etc. 

Prance (1977; 1982) has drawn attention to centres of endemism believed 
to have been isolated refuges for tropical moist forest flora during the 
cooler and drier periods coincident with the Pleistocene glaciations. While 
the theory of refuges as an exploration of existing patterns of diversity and 
species distribution may be contested there is general agreement on the 
location of the centres of endemism. 

An important first step towards the preparation of a plan for the 
selection of conservation sites throughout the country were taken in 1982, 
with the publication of the Plan for a System of Conservation Units in Brazil 
(IBDF 1982). This plan has yet to be implemented as a coherent whole, the 
information presented in it is still valid and is still useful. In respect 
of the Amazon region a further major step was taken by the International 
Symposium on Priority Areas for Conservation in the Amazon Basin - Workshop 
90, held in Manaus in January 1990 (Rylands 1990; Prance 1990). This meeting 
produced the initial draft of a map covering the entire Amazon basin, showing 
94 proposed priority areas, in three levels of priority for conservation. The 
final selection of areas was the product of discussion between some 100 
scientists working initially in small specialist groups (plant systematics; 
plant ecology; mammals; ornithology; herpetology; ichthyology; entomology; 
geomorphology and climate; and units of conservation). The botanists and 
zoologists subsequently pooled data from their component specialist groups to 
consolidate the first proposal for selection of areas while the data from 
geomorphology and climatology were used to identify fragile soils and 
ecosystems most in need of protection. Finally a synthesis was made from the 
botanical and zoological priorities to produce the draft map. Much further 
work is needed in the field to select actual sites for protected areas or for 
managed multipurpose production forests, in which conservation concerns are 
fully reflected. This may be progressively assisted by data from an ongoing 
study on the Biological Dynamics of Forest Fragments, north of Manaus. 

Although this initial prioritization is based on limited and unevenly 
distributed sources of information it has the great advantage of being 
regional in coverage, and can provide an initial framework for more detailed 
studies. It also provides some guidance for the allocation of priority to 
conservation objectives in existing production reserves. 

7.3 Management Options 

The earliest attempts at natural forest management for the sustainable 
production of timber in the Brazilian Amazon are comparatively recent. 
Experimental silvicultural work in the tropical moist forest at Curua-Una, 
near Santar&m, Para, began in 1957. It included some experimental logging and 
regeneration studies on a small scale and the trials then established are 
still monitored. In 1972 some management activities started in a second area, 
designated as the Tapajos National Forest (600 000 ha) in 1974 and as a result 


of studies and assistance from FAO a full management plan was prepared, 
envisaging selective harvesting, with a 70 year rotation based on natural 
regeneration, and a substantial component of line planting, to be harvested 
after 35 years (UNDP/FAO 1980). The aim was to allow logging of 1000 ha 
annually but the market conditions, heavily affected by an economic downturn 
in the early 1980 f s and by the availability of logs virtually free from large- 
scale forest clearance in other areas, prevented the implementation of the 
plan. However the forest has been well protected and studies have been 
undertaken which, together with the intensive inventory and studies undertaken 
initially, suggest the adoption of a polycyclic harvesting system, with 
cutting cycles of 30-35 years. Provision was made in the management plan for 
Biological Reserves and phenological studies, as well as 48 sample plots 
(Carvalho et al 1984), and the spatial distribution of 11 major species and 
the regeneration of 106 species were studied. 

Regeneration in the Tapajos forest is generally encouraging and 
frequently abundant, especially of valuable and fast-growing gap opportunist 
species such as Vochysia maxima after some opening of the canopy (Viana 1990). 
The ITTO, in response to a request from the Brazilian authorities to develop 
a demonstration model for forest management in the region, is providing 
assistance for controlled logging in the Tapajos National forest, to allow for 
pilot studies of management systems and silviculture on an operational scale, 
in concert with continued research into forest ecology and dynamics. An 
associated research proposal makes provision for identifying and protecting 
seed production stands of major species in the natural forest, covering a wide 
area in Amazonia, in collaboration with various organisations, as sources for 
future enrichment planting. 

In parallel with the study of forest management systems linked to 
genetic conservation objectives in specific areas, research into the genetic 
diversity and genetic systems of individual species will be needed. This 
applies for example to Swietenia macrophylla, which is apparently endangered 
by dysgenic selective logging outside the Reserved Forests. Other species 
likely to be given high priority for such action include Aniba rosaeodora, 
which has been subject to intensive exploitation for its essential oil 
content, and appears to show some chemical differences between provenances. 
In regard to in situ conservation of forest genetic resources the system of 
Genetic Reserves established in the Jari forest estate, with studies of 
phenology, ecology, regeneration and mortality, should provide valuable 
additional data. 

7,4 Secondary Forest and Non-Timber Forest Products 

Although the scientific approach to natural forest management for timber 
production is relatively recent and limited in scale other models of 
traditional systems of management, including utilization and management of 
timber trees, exist which may be incorporated in a comprehensive system of 
genetic resource conservation. Secondary forests are now widespread in 
recently settled areas of Amazonia and although frequently seen as examples 
of land abandonment following shifting cultivation or pasture degradation they 
are in fact used and managed by a variety of rural communities (Dubois 1990). 
Some of these forests include a high proportion of economic species, although 
selective logging of accessible areas along the rivers and streams has 
seriously depleted the populations, with possible dysgenic effects. Dubois 
(1990) proposes systems of enrichment planting, combined with the management 
of natural regeneration, to intensify productivity of a variety of tree 


species, in association with other crops, in agrof ores try systems dominated 
by the perennial vegetation. With appropriate attention to seed sources and 
management practices such systems could contribute to in situ conservation, 
as well as to the retention of forest cover and sustainable livelihood for the 
local people. 

Undoubtedly the longest history of forest management in Amazonia is that 
of the indigenous communities, harvesting mainly non-timber forest products 
(NTFP's). The Amerindian groups and "caboclos" (residents of the floodplains, 
of mixed descent) have detailed knowledge of the forest resources they depend 
on for survival (Parker et al 1983; Anderson 1990). The indigenous reserves 
may contribute to in situ conservation of some timber species but in addition 
knowledge of local traditional practices in the harvesting of NTFP's might be 
incorporated in management systems in National Forests and Extractive 
Reserves, with the possibility of retaining a wider base of species diversity 
in production systems. 

7,5 Information, Research and Coordination 

Prioritization and coordination of research are essential to focus 
resources where they are most needed, in terms of the areas to be productively 
managed, and the centres of diversity and endemism most endangered. For this 
reason, the association of Biological Reserves and Ecological Stations with 
the new National Forests, with provisions for research in aspects of ecology, 
phenology and forest dynamics relevant to silvicultural and conservation 
objectives, to be conducted in advance of the preparation of management plans 
and actual exploitation, is most likely to support efforts in in situ 
conservation. An example of this strategy is seen in the recent siting of a 
Biological Reserve for scientific research under the Museu Goeldi, Belem, in 
association with the Caxiuana National Forest. 

Equally important is the establishment of efficient, computerised data- 
banks, to facilitate the storage and retrieval of information on species 
distribution, with related environmental and other data bearing on the nature 
and future use of the genetic material. A good example within Brazil, 
although outside the Amazon .region, is the proposed cooperative programme of 
the Projeto Nordeste for a botanical species inventory, to be carried out in 
nine states of north eastern Brazil. This is expected to involve 
collaboration between ten or more Brazilian herbaria, research centres and 
universities, with the involvement of the Royal Botanic Gardens, Kew, U.K., 
within which the Biodiversity Database will form a coordinating link. The 
same principle should enable the progressive development of data needed for 
planning in situ conservation of forest genetic resources in the Amazon 
region. The involvement of major Brazilian centres of scientific expertise 
outside the region, such as the Universities of Sao Paulo and Rio de Janeiro, 
as has been recommended (Daly and Prance 1989), may also be facilitated by 
such coordinating mechanisms. 

Skilled human resources are the most vital requirement to gather and 
interpret the data needed for efficient and coordinated in situ conservation 
programmes. Because of the intrinsic scientific interest of the Amazon region 
it is possible to attract substantial international expertise and funding to 
support expeditions and inventories in this region, to be carried out in 
collaboration with Brazilian scientists. For example the research undertaken 
at the Maraci Ecological Station, Roraima, embracing both rainforest and 
savanna, at the invitation of the Brazilian Secretariat of the Environment, 


in 1987-88, mobilized the resources of 148 scientists and 55 technicians and 
resulted in the collection of data involving many new species, and leading to 
the compilation of the most comprehensive study to date of any forest area in 
northern Amazonia (Hemming 1989). Such collaborative schemes also offer 
opportunities for further training of local scientists, both through direct 
association with other specialists in the field and through postgraduate study 
offered within the programmes. 

The institutional focus to determine the selection of the areas and 
subjects for collaborative research, and to coordinate subsequent action in 
accordance with national priorities, is critically important in securing the 
most efficient use of available resources. In view of the very extensive 
geographical area covered by the Brazilian Amazon, which includes several 
states, and the number of research institutions and other organisations which 
might usefully be involved, such central coordination will be critically 
important to the success of the overall strategy for in situ conservation of 
forest genetic resources in the sub- region. 


8.1 National Policy 

Forest reservation and management, involving detailed working plans 
based on inventory growth and yield data and associated research, has been 
practised in the Indian subcontinent for over 100 years. 

The concept of Protected Areas in India can be traced back even further 
to at least the 4th century BC, when the establishment of Abhayaranyas (forest 
reserves) was proposed (Singh 1985). However in recent times since the 
establishment of the existing network of Forest Reserves the human population 
has more than trebled and the cattle population has grown by a factor of 2.5 
(Shyamsunder and Parameswarappa 1987; in IUCN 1991b). The country now 
contains around 15% of the entire human population, expanding at about 2% 
annually and expected to reach over 1.4 billion by year 2025 (WRI 1990). 
Through increasing population pressures and the demands of economic 
development, particularly in the period 1950 to 1980, India has lost large 
areas of its tropical forests. In 1952 the revised National Forest Policy 
called for the retention of one-third of the total land area as forests and 
this remains an objective in the 1988 Policy revision, which reinforced the 
increasing attention to the role of trees in land-use systems both within and 
outside the reserved forests. Whereas earlier policies and programmes had 
placed emphasis on meeting the raw material requirements of wood-based 
industries the 1988 policy gave priority to environmental conservation and to 
the needs of local people. Improved utilization of the large areas of 
degraded lands resulting from past deforestation and misuse had been made a 
priority action in both national and state government programmes. In 1985 the 
creation of the National Wasteland Development Board provided a central 
coordinating point for the massive reforestation programme (target 
approximately 9 million hectares during the 7th Five Year Plan) to develop 
fuelwood and fodder supplies. Through reorganisation of forestry research and 
training, greater attention was focused on aspects of social and community 
forestry, and to reforestation, in accordance with the revised Forest Policy. 
At the same time the rate of deforestation was substantially reduced, and 
conservation efforts were strengthened. 


Wildlife protection also has a long tradition in India and by 1988 there 
were 65 national parks and 407 wildlife sanctuaries covering an area of 
approximately 131 800 km 2 or 4.4% of the land area (IUCN 1991b), although a 
significant proportion had not yet achieved full legal status. The network 
of 16 Tiger Reserves, covering 26 000 km 2 , including large areas of tropical 
moist forest, has been recognised internationally as a most significant 
conservation success. Nevertheless major problems remain in securing an 
adequate national network of Protected Areas for the conservation of 
biological diversity and genetic resources. To complement such areas, 
considerable reliance will need to be placed on the protection and management 
of ecosystems, species and genetic resources outside the Protected Area 
network, and particularly within Forest Reserves while at the same time taking 
care to not further deepen the already existing conflict between the needs of 
the rural populations surrounding Forest Reserves and Protected Areas, for 
fuelwood, fodder and a range of other forest products. Among the most 
significant areas for special attention in this respect are the surviving 
tropical rain forests, confined to the Anadaman and Nicobar Islands, to areas 
in the north-eastern states, and to the Western Ghat forests in southern 

8-2 Western Ghat Forests 

The Western Ghat Forests occupy the range of high hills extending for 
about 1 600 km from the southern point of India up the western coast, to about 
21N. Three main regions within them have been recognised (Pascal 1982) with 
the largest and best conserved areas of the original forest lying in the 
central region, within Karnataka State. Here the hills range from 700 to 
nearly 1 900 m a.s.l. The rainfall varies along three gradients: south- 
north, east-west (with a marked rain shadow on the eastern side) and 
altitudinally. Annual totals show great variation, but are concentrated 
within a few months (e.g. at Agumbe the mean of 7 000 mm/ year is concentrated 
in 128 days) with a dry season lasting from 3 months (at high altitude in the 
south) to 8 months, depending on locality. All the major rivers of South 
India arise in the Western Ghats and their environmental significance was the 
prime reason for early demarcation, reservation and the protection of 
significant areas of Reserved Forest in this area over the past 120 years, 
since the 1870's. 

Despite the legal status of the Reserved Forests, and the efforts of an 
efficient State Forest Service, large areas were lost to dams, mining and 
cultivation, including substantial illegal encroachment. Nearly 60% of the 
remaining forest is classified as degraded, much of it severely, but there is 
still a significant "core" area of evergreen and semi-evergreen forest (Sinha 
1988), of the greatest ecological importance and arguably of at least equally 
high priority in global terms as any other tropical moist forest in the world. 
The Western Ghat Forests lie between the African and the Indo-Malaysian 
rainforest blocks and are isolated from neighbouring areas of rainforest. 
While the levels of total biological diversity are lower than those present 
in most other tropical rainforests in Asia, they are rich in endemic species. 

Of the 15 000 higher plants recorded in India over 4 000 species are 
found in the Western Ghats (5% of the land area) and of these 1 800 species 
are endemic, mainly found in the remaining evergreen forests (IUCN 1991b). 
These include 130 tree species endemic to the Ghats (Bor 1953) and six endemic 
mammals, including the lion-tailed macaque (Macaca silenus), while 84 of 
India's 112 endemic amphibians are found there (Inger and Dutta 1987). In 
regard to trees and shrubs, both the diversity of species and intraspecific 


variation, adapted to the local environment, are clearly of great potential 
importance. In no other part of the world is evergreen rainforest found under 
such seasonal climates (Rai and Proctor 1986), and the adaptations of plant 
populations, including many valuable timber trees, to the climatic conditions 
may be even more valuable in the context of possible climate change. 

Among the most valuable timbers of international significance are 
Dalbergia latifolia (Indian rosewood), Calophyllum inophyllum, Pterocarpus 
marsupium, Tectona grandis, Terminalia crenulata, T.paniculata and other 
species suitable for furniture, doors, windows, interior construction and 
other high value applications. Many more are of great local importance for 
house construction, bridges, carts, agricultural and household implements etc. 
As a result of over-exploitation and particularly illegal logging, 
overgrazing, fires, encroachment by cultivators and other pressures from the 
increasing populations, both the effective area and the regeneration of the 
forests have been severely reduced. In the surrounding areas there are almost 
as many cattle as people and it has been estimated that at least 10% are 
dependent on free grazing in the forests. Whereas this had formerly been 
regulated within permissible limits such control is no longer possible because 
of the huge numbers involved (IUCN 1991b). The effect on the genetic 
resources of the tree populations from the reduction and local elimination of 
regeneration as a result of these pressures is likely to be increasingly 

8.3 The Strategy for Integrated Development and Conservation 

Despite a clear policy, sound legislation and an efficient Forest 
Service, which had succeeded in protecting the reserved forests for over 100 
years, it became clear that their effective regeneration and therefore long- 
term survival were increasingly threatened. While the intensified programmes 
of reforestation were expected to mitigate the pressure, further action was 
needed to conserve the natural ecosystems and their unique genetic resources. 
In 1988 the Karnataka State Forest Department drafted a proposal for a project 
on the Integrated Development of Forests in the Western Ghats to attempt to 
reconcile satisfactorily the many conflicting demands made on the forests, 
including the central need to conserve ecosystems and genetic resources. The 
central element of this strategy was the recognition of three sets of 
management objectives, namely ecological, economic and social ("local needs 11 ). 
The conservation of the genetic resources of the tree and shrub species 
relates in different ways, and to different degrees, to all three of these 
objectives. However the actual operations needed to achieve conservation will 
vary from strict protection, involving exclusion of other human activity, to 
agro-forestry programmes. The concept involves a multidisciplinary approach, 
and the recognition that local people, who are responsible for much of the 
biotic pressure on the forest, must play a role in the planning, management 
and protection of the forest resources. The means to achieve the 
reconciliation of development and conservation objectives is the Joint Forest 
Planning and Management (JFPM) system, which derives from practices being 
developed by the Karnataka Forest Department (KFD). 

The two essential elements are: 

(a) Joint Planning - a consultative process by which the KFD and 
local people jointly discuss the ecological and environmental 
condition of a specific area of the forest, and the scope for it 
to meet specific needs. 

(b) Joint Management - for certain areas, where the KFD and forest 
users will agree on a division of responsibilities for management 
and a division of the proceeds. The precise arrangements will 
vary from one specific area to another and as appropriate the 
joint management may cover all, or only certain categories of the 
trees and other plants present. 

A key element is the division of the forest into four zones, with an 
additional zone comprising an area immediately outside the Reserved Forest to 
be jointly managed to meet the needs of the adjacent villages. Within that 
a boundary zone of forest, also under joint management with local people, will 
serve similar objectives. The main part of the forest is to be managed for 
production of timber and other products, divided into two zones, depending on 
whether or not there are forest-dwelling people. Finally the "core" zone is 
strictly reserved for conservation objectives. An important aspect of the 
management of all areas where joint responsibilities operate is the 
preparation of contractual forest agreements according to specific micro-level 
plans. The ownership and statutory responsibility for management over the 
entire forest reserve remains with the KFD, even where management 
responsibility may be shared by contractual agreement. 

Future working plans for the management of the forest will be prepared 
by zones. In the "core" zone the plan will be oriented to research, concerned 
with autecology and the interrelationships between species, and with studies 
providing a "control" for comparison with the regeneration, growth and yield 
studies in the production zones, which will be designed to gather the data 
needed for management, silvicultural and harvesting activities. 

The satisfactory harmonization of the different objectives and interests 
is dependent not only on the process of joint planning and management but also 
on the incorporation of new information to be gathered in the course of the 
project into the management process, related to improved understanding of both 
ecological and socio-economic relationships; and on the realisation and 
distribution of benefits from improved management of the resource. In 
addition to the biological research directly related to the conservation and 
management of the forest's genetic resources, aspects of social and economic 
research, and the incorporation of staff training and public education into 
the programme are also needed. 

The conceptual relationship between meeting the needs of the local 
people, restoring and maintaining the productivity of the forest resource and 
the conservation of overall genetic diversity and genetic resources, is 
illustrated below. 



















Efficient data collection, handling and analysis will be an important 
feature of the strategy, using enhanced GIS capabilities, taxonomic databases, 
and when appropriate modelling of growth, yield and other features of the 
production system. Where, because of the severe reduction in the regeneration 
within the forests, and the need to reestablish viable populations, enrichment 
planting and gap planting is planned, careful selection and documentation of 
seed sources will be needed to safeguard the genetic integrity of the local 
populations. In some circumstances, particularly in the outer zones, the use 
of exotic species incapable of hybridizing with indigenous populations, may 
be appropriate. However important opportunities exist to reinforce in situ 
conservation of the genetic resources of the original forest by artificial 
regeneration using local seed sources. For some species the strategy may also 
include aspects of ex situ conservation, possibly in association with the 
Trivandrum Botanic Garden in the adjacent Kerala State, which has an emerging 
programme for the conservation of rare and endemic plants of selected centres 
of endemism within the Western Ghats. 

Appendix 1, 


David Boshier 1 

1 . Introduction 

Sound decisions in genetic conservation and progress in long term breeding 
require detailed knowledge of the taxonomy, population structure, reproductive 
biology, mating systems, etc of the species concerned. For many tropical forest 
tree species, little information is available and managers are faced with 
problems in developing effective strategies. For example, questions that need 
to be answered are:- Are neighbouring trees in natural stands highly related 
and is there inbreeding? What, if any, is the incompatibility mechanism to stop 
inbreeding? What agents effect pollination and how do they affect gene flow and 
population size? What degree of inbreeding is acceptable? 

The case study presented here describes the methodology used in ongoing 
research carried out for one particular species - Cordia alliodora. It is not 
intended that the methodology described here be used as a recipe for studying 
any tropical tree species, but that it may provide ideas on how to approach the 
problem. Techniques used will always depend on the type of tree, its 
distribution, size of flowers, pollinators etc. Previous work with C^ 
alliodora, relevant to the study, is generally of a preliminary nature but 
provides useful information. 

C. alliodora is an important neotropical tree, which combines high quality 
timber and value with fast growth on good quality soils. It is used extensively 
throughout its natural range and its light crown and self pruning habit make it 
particularly suitable for use in various agrof ores try systems, providing 
valuable timber and income to small farmers. The species has a wide 
distribution occurring from northern Mexico through Central and South America 
as far south as Bolivia, Paraguay, southern Brazil and northern Argentina. In 
Paraguay, northern Argentina and parts of southern Brazil the closely related 
C. trichotoma (also classified as C. alliodora var tomentosa) predominates over 
C. alliodora (Gibbs and Taroda 1983). C. alliodora is found on most of the 
Caribbean Islands from Cuba to Trinidad, but is almost certainly not native to 
Jamaica. Through this geographical range the species occurs under a wide 
variety of ecological conditions, varying from very wet (as much as 6 000 mm per 
year) to seasonally dry (as low as 600 mm per year); and from sea level to as 
high as 1 200 m a.s.l. in Central America and 2 000 m a.s.l. at lower latitudes 
in Colombia. In lowland humid tropical regions it is generally a tall, thin, 
lightly crowned tree, reaching heights over 40 m and d.b.h. up to 1 m in mature 
trees, although in the region of 50 cm d.b.h. is more normal. In seasonally dry 
areas it is smaller, and more poorly formed rarely reaching more than 20 m in 

The overall objective of the present study was to gain an understanding 
of the reproductive biology of C. alliodora that will give adequate knowledge 
to make sound long term decisions for in-situ/ex-situ conservation and breeding 
of the species. 

Oxford Forestry Institute, South Parks Road, Oxford, 0X1 3RB, U.K. 


The specific objectives were to: 

1) study in detail the phenology of flowering; 

2) determine the type of pollination and possible incompatibility systems; 

3) study the mating system, gene flow and neighbourhood area found in natural 

2- Flowering and Seed Phenology 

Flowers are hermaphrodite, unspecialized, about 1 cm in length and occur 
in large panicles. The size of panicles varies, with as few as 50 flowers to 
as many as 2 000. The petals are white and persistent turning to brown and 
acting as a parachute in wind dispersal of the seed. The aim of the first phase 
of the phenology study was to detect any patterns of flowering that may exist 
within the tree, branch and inflorescence and provide data on length of the 
flowering and fruiting periods and the proportions of seed reaching maturation. 
It is important to know how a tree flowers and what effect that might have on 
pollinator movements. For example: sequential flower opening from the top to 
the bottom of the crown may mean that early flowers are pollinated by one group 
of pollinators and later flowering by a completely different group of 
pollinators. Conversely, random flowering within the crown could stimulate 
greater movement of pollinators between panicles and different trees. 

2.1 Within inflorescences: individual groups of flowers were identified and 
the stage of flowering observed every three days from flowering to seed fall. 
Classification of the stages of flowering was based on Mendoza (1965), 
distinguishing : 

1 - start of formation of flower buds 

2 - individual flower buds visible, but not open 

3 - individual flower buds full 

4 - white petals emerge from flower buds, but not open 
5a- petals open, stigma receptive 

5b- petals still white, stigma withered 

6 - petals turn brown 

7 - embryo starts to swell. 

It was thus possible to observe at which stage of flowering most losses 
occurred, the length of each stage, any sequential pattern of flowering or 
losses within a panicle, and the effect certain weather conditions might have 
on flowering and fruiting. 

2.2 Within branches and trees: individual inflorescences/branches within a 
number of branches/trees were observed periodically from flowering to seed 
production. The same flowering categories were used as in 2.1 (categories 5a 
and 5b combined) and each panicle/branch classified on the predominant category 
of flowers. 

2.3 Within populations: flowering phenology within a population of trees of 
the same species is of fundamental importance to understanding gene flow, 
genetic structure, seed production and yet few studies exist of individual 
tropical tree species based on large samples. Similarly few studies have looked 
at the flowering of the same population for more than one year. It is important 
to know about periodicity of flowering for the species and how individual 
species flower with respect to one another from year to year. For example, 
because of a synchrony of flowering two adjacent trees may be unable to mate, 
which may be the case in one year but not in the following. Similarly some 
trees may be heavy flowerers in one year and totally sterile the following year. 


To look at flowering within natural populations, in a 26 hectare plot all 
trees (216) of a flowering age/size surrounding three C. alliodora selected 
trees up to a radius of approximately 500 meters were mapped, and labelled. The 
stand, which was also to be used for the mating system and gene flow study (see 
5), was chosen taking into account various factors: (a) the presence of 
selected plus trees would provide direct information on the degree of diversity 
sampled when collecting open-pollinated seed from the plus trees, which are part 
of a breeding population (Boshier and Mesen, 1989); (b) the possibility of 
easily defining a population, there being a natural break between the trees 
under study and nearest trees of the same species; (c) ease of access to allow 
frequent visits. 

The crown of each tree was observed every three days to determine the 
start, peak and end of flowering and the percentage of the total number of 
flowers open by that date. Subjective assessments of flower (scale of 0-5) and 
seed production (scale of 0-3) were also made. The same observations were made 
over three successive flowering seasons (Jan-April) to study year to year 
variation. An index for synchronicity of flowering was calculated, for the 
whole stand as well as for the plus trees in relation to the surrounding trees 
(Augsberger, 1983). To look at the effect population substructure may have on 
flowering synchrony the individual tree and stand index was re-calculated on the 
basis of increasing population size from a base tree (a plus tree). Two other 
measures of population synchrony were made: (1) synchrony of the first day of 
flowering, calculated as one standard deviation around the mean of the first day 
of flowering; and (2) synchrony of the median day of flowering, calculated 
as one standard deviation around the mean of the median flowering day. For one 
flowering season similar phenological observations were carried out in a stand 
in the seasonally dry Pacific region to allow comparison of flowering under 
differing climates. 

2.4 Between populations 

Much information on the general phenology of a species can be gained from 
the examination of herbarium specimens. Even for species for which full 
collections are not available, studying existing specimens can reveal much about 
phenology, distribution and likely regions where the species may occur. 
Herbarium collections of C. alliodora and C. trichotoma were studied and full 
details noted form specimen labels as well as the stage of flowering. It was 
possible to classify well-preserved specimens under one of the flowering 
categories used in 2.2. Logging of the information onto the OFI herbarium 
database, BRAHMS (Filer, 1991) allows mapping of the distribution of the species 
as well as studying gross variation in flowering time over this range. 

3. Breeding System 

Many species of Cordia are heterostylous, indeed in one case failure to 
recognize this originally led to one species being classified as two (C^ 
thaisiana and C. apurensis, Agostini, 1983). Gibbs and Taroda (1983) studied 
the C. alliodora * C. trichotoma complex based on herbarium specimens from South 
America and distinguished the two species on the basis of the pattern of 
heterostyly. In C. alliodora heterostyly appears to have broken down although 
there is variation in style length. Opler et al (1975) studied the pollination 
biology of a number of species of Cordia and found varying degrees of self- 
incompatibility in the trees of C. alliodora they sampled. 


A combination of field and microscope work were used to study stylar 
patterns, both within and between families and their relationship to the 
incompatibility mechanism. On freshly cut inflorescences from an open 
pollinated progeny trial, flowers were emasculated as they opened and before 
anther dehiscence had occurred. Controlled cross, self and related pollinations 
between half-sibs were carried out and the flowers fixed at varying times after 
pollination to allow study of pollen tube development on the stigma and through 
the style. 

The same controlled crosses were carried out in the field to allow 
comparison with the results from the microscope study, checking that crosses 
deemed compatible produce seed and those deemed incompatible failed to do so. 
The flowers on any one panicle open during a period of 4-6 days. The numbers 
of seed produced per cross and empty flowers were counted to compare fertility 
from related and unrelated crosses. The seed was used to mount a nursery trial 
to look at the effects of inbreeding on initial growth rate. Controlled 
pollinations were also carried out in the field to look at length of stigma 
receptivity, and to exclude any possibility of parthenocarpy. It was noticeable 
that in the dry zone the stigma was receptive for only one day at the end of 
which it was withered whereas in the wet zone the stigma could be receptive for 
as long as three days, emphasising the importance of caution in extrapolating 
results from one region to a climatically different one. 

4 Pollinators 

Opler et al (1975) looked at potential pollinators and concluded that 
noctuid and geometrid moths and anthophorid bees were the most likely 
pollinators. Their work was however restricted to the dry Guanacaste region of 
Costa Rica and it is likely that in the wet lowland regions other pollinators 
are of importance. As many insects are seasonal, it is likely that different 
insects act as pollinators at different times during a prolonged flowering 
season. Collections were made of insects visiting flowers during the day and 
night at various times in the flowering season. The insects were mounted, the 
identified to genus and stored for future use as a reference collection. 
Observations were made on several occasions of numbers and types of insects 
visiting flowers, to see which were the more important ones. Other studies such 
as those on the effectiveness of different pollinators in depositing pollen on 
the stigma, or marking and recapturing to look at fidelity to food source could 
also be carried out (Frankie et al 1976). 

Flowers were collected and fixed at different times of the day following 
anthesis (dawn, midday, dusk) from a number of trees. Microscopic observation 
of these flowers for the quantity of pollen on stigma and pollen tube growth can 
provide supportive evidence on the time of pollination and the relative 
importance of certain pollinators. Similar collections were made in the Pacific 
region to compare pollination for the species in differing climates. 

It is important to study variation of nectar flow with time as this can 
give clues as to which vectors are active pollination and when. A number of 
trees should be studied as there can be variation from tree to tree both in time 
of production, quantity, and quality (sugar concentration). Even within an 
inflorescence patterns of nectar production may exist which will tend to move 
pollinators in certain directions. To characterize the availability of rewards 
to pollinators, variation in nectar production was studied on a number of trees 
at intervals during a three day period after flower opening. The process was 
repeated on a number of occasions during the flowering season. 


5. Mating System, Gene Flow and Neighbourhood Area 

Isozymes may be used a genetic markers to provide estimates of the mating 
system, gene flow, neighbourhood area and paternity. The interpretation of such 
data is made more significant when it can be combined with field observations 
of the spatial distribution of the trees and flowering phenology. For this 
reason the same population was chosen for studying mating system as for the 
within population phenology study. Seed was collected during the 1989 season 
from the trees surrounding two plus trees; during 1990 and 1991 seed was re- 
collected from the plus trees to look at year to year variation in outcrossing 
rates and paternity for these two trees. Although sufficient seed could be 
collected from one panicle, seed was collected from the whole crown, to ensure 
that the sample was not biased due to particular pollination events. To look 
at variation of pollination within a tree, at the time of collection, seed from 
one plus tree was divided into three lots representing the upper, middle and 
lower parts of the crown. Seed from three panicles on the same branch on the 
plus tree was also separated as individual lots. 

A total of 163 trees occurred within a 250 m radius of the plus trees, a 
number prohibitively large to allow all to be studied for isozymes. A sub- 
sample of 52 trees was therefore selected on the basis of distance from the plus 
tree and phenological data obtained during field work (see 2.3). The following 
trees were eliminated from the sample: (a) non-flowerers (36 trees); (b) trees 
that flowered completely before or after the plus tree (5); (c) trees 
classified as 1 for quantity of flowering (i.e. six or less panicles) (12); (d) 
trees in which less than six days synchrony in flowering was observed (13); (e) 
trees with scarce flowering (category 2) (21); (f) trees in category 3 for 
flowering and greater than 200 m from the plus tree (24). To aid in determining 
paternity for progeny arrays of specific trees, genotypes were assigned to un- 
sampled trees by assaying leaf material. The leaf material was collected and 
immediately frozen in the field in liquid nitrogen. 

To complement estimates of pollen flow from the isozyme work and allow 
estimates of neighbourhood size and area, seed dispersal was studied for four 
trees in the same population used for phenology and isozymes. A UV fluorescent 
dye was used to spray seed in the canopy and so enable tracing distance of seed 
dispersal from the trees. Seed was collected in 1 m 2 traps, placed at intervals 
along transects, laid out in the direction and reverse of the prevailing wind. 
The dye was still easily visible on the seed after six weeks and probably 
persists much longer. 

6. Variation between Populations 

There are two complementary approaches to studying genetic variation 
between populations. The first, provenance testing, is generally used in 
forestry and studies the performance under uniform environmental conditions in 
one or more planting sites of trees grown from seed collected from different 
populations. Traits studied are usually of a continuously variable nature and 
the observed variation is split into genetic and environmental components. 
Provenance variation for Cordia alliodora has been studied in an international 
trial coordinated by the Oxford Forestry Institute based on collections made in 
the late seventies principally in Central America, with assistance from FAO and 
ODA (Stead, 1980). Over a wide range of sites, provenances from the humid 
Atlantic region of Central America and in particular Honduras and Costa Rica 
showed superior growth and form to Pacific region provenances, even when grown 
in seasonally dry areas (McCarter 1988). The study thus suggested ecotypic 
differentiation of the Pacific and Atlantic populations, but also indicated that 


within these broad bands variation within provenances was greater than between. 
There appears to be great potential for genetic improvement through individual 
tree selection, the provenance assessments showing a 200-3002 difference within 
provenances between the best 10X of trees and the mean. 

The second approach is to study genetic diversity shown as variation in 
enzymes (and more recently DNA) for specific gene loci. The enzyme survey 
carried out in (5) above showed differences in the number of loci staining; and 
variability within loci, between the population under study and a Pacific zone 
population previously studied. Further studies of isoenzyme variation in the 
original OFI provenance collections of C. alliodora, being carried out at 
University of Massachusetts, Boston (U.S.A.), indicate interesting differences 
between provenances. 

7. Chromosome Number 

More information on variation within and between closely related species 
can sometimes be revealed by looking at chromosome variation, both in terms of 
numbers and overall quantities of DNA. The two existing reports of chromosome 
number in C. alliodora; n=15 (Bawa 1973) and 2n=72 (Britton 1951), differ 
greatly. The possibility of gene duplication for a number of enzyme systems in 
the Atlantic population also suggests that some benefit may be gained from 
looking at chromosome number to see if intra-specific variation does exist in 
this trait, and if this bears any relation to the Pacific/Atlantic provenance 
differences encountered in the international provenance trials. 


The above study was financed by the Overseas Development Administration 
of the United Kingdom (R4484 & R4724). The author is grateful to the following 
people; Ing. F. Mesen, CATIE; Ing. F. Lega, Scott Paper, for facilitating the 
field work in Costa Rica; to Dr. K.S. Bawa and Mr. M. Chase, Univ. Mass., Boston 
for collaboration with the isozyme work and Dr. G. Frankie, Univ. California, 
Berkeley, for entomological advice. 



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; 111403 



1 Forest utilization contracts on public land, 1977 30 
(E F S) 31 

2 Planning forest roads and harvesting systems, 32 

3 World liat of forestry schools, 1 977 (E/F/S) 33 
3 Rev. 1 . World liat of forestry schools, 1 98 1 (E/F/S) 34 
3 Rav. 2. World list of forestry schools, 1 986 (E/F/S) 35 
4/1 World pulp and paper demand, supply and trade 36 

-Vol. 1,1977 (EFS) 37 

4/2 World pulp and paper demand, supply and trade 

-Vol.2, 1977 (EFS) 38 

5 The marketing of tropical wood, 1 976 (E S) 39 

6 National parks planning, 1 976 (E F S ) 40 

7 Forestry for local community development, 1978 41 
(Ar EFS) 

8 Establishment techniques for forest plantations, 42 
1978(ArCE* F S) 

9 Wood chips - production, handling, transport, 43 

10/1 Assessment of logging costs from forest 44/1 

inventories in the tropics 

- 1 . Principles and methodology, 1978 (E F S) 44/2 
1 0/2 Assessment of logging costs from forest 

inventories in the tropics 

- 2. Data collection and calculations, 1978 (E F S) 44/3 

1 1 Savanna afforestation in Africa, 1 977 (E F) 

12 China: forestry support for agriculture, 1978 (E) 45 

1 3 Forest products prices 1 960-1 977, 1 979 (E/F/S) 46 

1 4 Mountain forest roads and harvesting, 1 979 (E) 47 

1 4 Rev. 1 . Logging and transport in steep terrain, 1 985 (E) 

1 5 AGRIS forestry - world catalogue of information 48 
and documentation services, 1979 (E/F/S) 49 

16 China: integrated wood processing industries, 1979 

(E F S) 50 

17 Economic analysis of forestry projects, 1979 50/1 

17 Sup. 1 . Economic analysis of forestry projects: 51 /I 

case studies, 1979(ES) ' 

17 Sup. 2. Economic analysis of forestry projects: readings, 52/1 


1 8 Forest products prices 1 960- 1 978, 1 980 (E/F/S) 52/2 
1 9/1 Pulping and paper-making properties of 

fast-growing plantation wood species 53 

-Vol. 1,1 980 (E) 

19/2 Pulping and paper-making properties of 54 

fast-growing plantation wood species 55 

- Vol. 2, 1980 (E) 

20 Forest tree improvement, 1985 (C E F S) 56 
20/2 A guide to forest seed handling, 1 985 (E S ) 57 

21 Impact on soils of fast-growing species in lowland 58 
humid tropics, 1 980 (E F S) 59 

22/1 Forest volume estimation and yield prediction 

- Vol. 1 . Volume estimation, 1980 (C E F S) 60 
22/2 Forest volume estimation and yield prediction 

- Vol. 2. Yield prediction, 1980 (C E F S) 61 

23 Forest products prices 1961-1980, 1981 (E/F/S) 62 

24 Cable logging systems, 1 981 (C E) 

25 Public forestry administrations in Latin America, 63 
1981 (E) 64 

26 Forestry and rural development, 1981 (EFS) 65 

27 Manual of forest inventory, 1 981 (E F) 

28 Smalt and medium sawmills in developing countries, 66 
1981 (ES) 67 

29 World forest products, demand and supply 1 990 

and 2000, 1982 (EFS) 68 

Tropical forest resources, 1982 (E F S) 

Appropriate technology in forestry, 1982 (E) 

Classification and definitions of forest product! 


Logging of mountain forests, 1982 (E F S) 

Fruit-bearing forest trees, 1982 (E F S) 

Forestry in China, 1982 (C E) 

Basic technology in forest operations, 1982 (E 

Conservation and development of tropical fore 

resources, 1982 (EFS) 

Forest products prices 1962-1981, 1982 (E/F< 

Frame saw manual, 1982 (E) 

Circular saw manual, 1983 (E) 

Simple technologies for charcoal making, 198 


Fuel wood supplies in the developing countries 


Forest revenue systems in developing countri* 

1983 (EFS) 

Food and fruit-bearing forest species 

- 1. Examples from eastern Africa, 1983 (E F 
Food and fruit-bearing forest species 

- 2. Examples from southeastern 
Asia, 1984 (EFS) 

Food and fruit-bearing forest species - 3. Exi 

from Latin America, 1986(ES) 

Establishing pulp and paper mills, 1983 (E) 

Forest products prices 1963-1982, 1983 (E/l 

Technical forestry education - design and 

implementation, 1984 (E F S) 

Land evaluation for forestry, 1984 (C E F S) 

Wood extraction with oxen and agricultural 

tractors, 1986 (EFS) 

Changes in shifting cultivation in Africa, 1 98 

Changes in shifting cultivation in Africa 

- seven case-studies, 1 985 (E) 

Studies on the volume and yield of tropical f 

stands - 1 . Dry forest formations, 1 989 (E I 

Cost estimating in sawmilling industries: gui 


Field manual on cost estimation in sawmillin 

industries, 1985 (E) 

Intensive multiple-use forest management ir 

1984 (EFS) 

Planificaci6n del desarrollo forestal, 1984 (S 

Intensive multiple-use forest management ir 

tropics, 1985 (EFS) 

Breeding poplars for disease resistance, 1 9! 

Coconut wood - processing and use, 1 985 

Sawdoctoring manual, 1 985 (E S) 

The ecological effects of eucalyptus, 1985 

(C E F S) 

Monitoring and evaluation of participatory f 

projects, 1985 (EFS) 

Forest products prices 1965-1984, 1985 (! 

World list of institutions engaged in forestr 

forest products research, 1985 (E/F/S) 

Industrial charcoal making, 1985 (E) 

Tree growing by rural people, 1985 (Ar E F 

Forest legislation in selected African count 


Forestry extension organization, 1986 (C E 

Some medicinal forest plants of Africa and 

America, 1986 (E) 

Appropriate forest industries, 1986 (E) 

69 Management of forest Industrie*, 1 986 (E) 

70 Wildland fire management terminology, 1986 

71 World compendium of forestry and forest products 
research institutions, 1986 (E/F/S) 

72 Wood gas as engine fuel, 1 986 (E S) 

73 Forest products: world outlook projections 
1985-2000, 1986 (E/F/S) 

74 Guidelines for forestry information processing, 
1986 (E) 

75 An operational guide to the monitoring and 
evaluation of social forestry in India, 1986 (E) 

76 Wood preservation manual, 1986 (E) 

77 Databook on endangered tree and shrub species 
and provenances, 1986 (E) 

78 Appropriate wood harvesting in plantation forests, 

79 Small-scale forest-based processing enterprises, 

80 Forestry extension methods, 1987 (E) 

81 Guidelines for forest policy formulation, 1987 (C E) 

82 Forest products prices 1967-1986, 1988 (E/F/S) 

83 Trade in forest products: a study of the barriers 
faced by the developing countries, 1 988 (E) 

84 Forest products: world outlook projections 
1987-2000 - product and country tables, 1988 

85 Forestry extension curricula, 1988 (E/F/S) 

86 Forestry policies in Europe, 1988 (E) 

87 Small-scale harvesting operations of wood and 
non-wood forest products involving rural people, 
1988 (EPS) 

88 Management of tropical moist forests in Africa, 

89 Review of forest management systems of tropical 
Asia, 1989 (E) 

90 Forestry and food security, 1989 (Ar E S) 

91 Design manual on basic wood harvesting 
technology, 1989 (EPS) 

(Published only as FAO Training Series, No. 18) 

92 Forestry policies in Europe - an analysis, 1989 (E) 

93 Energy conservation in the mechanical forest 
industries, 1990 (ES) 

94 Manual on sawmill operational maintenance, 

95 Forest products prices 1 969- 1 988, 1 990 (E/F/S) 

96 Planning and managing forestry research: guidelines 
for managers, 1990(E) 

97 Non-wood forest products: the way ahead, 
1991 (ES) 

98 Las plantations a vocation de bois d'osuvre en 
Afrique intertropicale humide, 1991 (F) 

99 Cost control in forest harvesting and road 
construction, 1992 (E) 

100 Introduction to ergonomics in forestry in developing 
countries, 1992(E) 

101 Amtnagement et conservation des forto denses en 
Ame*rique tropicele, 1992 (F) 

102 Research management in forestry, 1991 (E) 

103 Mixed and pure forest plantations in the tropics and 
subtropics, 1992 (E) 

104 Forest products prices 1971-1990, 1992 (E) 

105 Compendium of pulp and paper training and 
research institutions, 1 992 (E) 

106 Economic assessment of forestry project impacts, 

107 Conservation of genetic resources in tropical forest 
management: Principles and concepts, 1 993 (E) 

108 A decade of wood energy activities within the 
Nairobi Programme of Action, 1 993 (E) 

Availability: February 1993 













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