THE STATE
OF THE WORLD’s
FOREST GENETIC RESOURCES
COMMISSION ON GENETIC RESOURCES FOR FOOD AND AGRICULTURE
FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
Rome, 2014
i
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Foreword
F
orests cover nearly one-third of the world’s land area. They provide vital environmental
services such as soil and water protection, regulate the climate and preserve biodiversity,
produce valuable raw materials and food, and sustain the livelihoods of millions of
people.
Forest genetic resources – the heritable materials maintained within and among trees
and other woody plant species – are essential for the adaptation and the evolutionary
processes of forests and trees as well as for improving their resilience and productivity.
The conservation of forest genetic resources is more topical than ever at a time when the
world is increasingly confronted with challenges from increased human population, landuse changes and climate change. These pressures, and related increases in unsustainable
use, wildire, pests and diseases, as documented in the Climate change 2013 report of the
Intergovernmental Panel on Climate Change (IPCC), are causing losses of forest cover and of
forest biodiversity, both among and within species. Lack of information limits the capacity
of many countries and the international community to develop appropriate policy tools
to address the issues or to integrate forest genetic resources management into relevant
cross-cutting sectorial policies.
Reliable data on the status and trends of forest genetic resources are required for decisionmakers and stakeholders to provide adequate support for their sustainable management.
Recognizing this need for information and the urgency of addressing the conservation and
sustainable use of forest genetic resources, the Commission on Genetic Resources for Food
and Agriculture requested and guided the preparation of The State of the World’s Forest
Genetic Resources, and agreed, in response to its indings, on strategic priorities which the
FAO Conference adopted in June 2013 as the Global Plan of Action for the Conservation,
Sustainable Use and Development of Forest Genetic Resources.
This irst ever report on The State of the World’s Forest Genetic Resources constitutes
a milestone in building the information and knowledge base required for action at the
national, regional and international levels. It has been developed through a country-driven
process, building on 86 country reports – representing over 85 percent of global forest
cover – and with the participation of representatives from national institutions and nongovernmental and community-based organizations. Its recommendations are based on
these reports, which indicate that about half of the forest species in reporting countries are
threatened or subject to genetic erosion, and only about one-quarter are actively managed
for their products and/or services. This publication provides the basis for renewed efforts to
realize national and international commitments to improved conservation, sustainable use
and management of forest genetic resources.
As established in its Reviewed Strategic Framework 2010-2019 and in particular through
its Strategic Objective 2, FAO is striving to “increase and improve provision of goods and
services from agriculture, forestry and isheries in a sustainable manner”. Measures include
strengthening its technical support to countries in the area of forest genetic resources
and promoting the integration of forest genetic resources into broader forest resource
management programmes at the national, regional and international levels. This report is
a key ingredient in this effort.
I am conident that the information in The State of the World’s Forest Genetic Resources
will be used as the basis for policy and technical decisions to strengthen national efforts in
conservation and sustainable management of forest genetic resources, efforts that will contribute to
meeting the world’s current and future needs for forest products and environmental services while
enhancing food security.
iv
Contents
Foreword
Acknowledgements
About this publication
Executive summary
Part 1
Overview
CHAPTER 1
CHAPTER 2
CHAPTER 3
CHAPTER 4
Part 2
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xiii
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xxi
BASIC CONCEPTS
3
Deinitions
Characteristics of forest genetic resources
Species diversity
4
8
11
VALUE AND IMPORTANCE OF FOREST GENETIC RESOURCES
19
Economic value
Environmental value, ecosystem services and resilience
Social, cultural, medicinal and scientiic value
Preserving options for future development and adaptation
20
22
24
25
CONSERVATION OF FOREST GENETIC RESOURCES
27
Management systems in the ield (in situ and circa situm
conservation)
Ex situ conservation
Targeted species-based approach
28
32
39
KNOWLEDGE AND INFORMATION ON FOREST GENETIC
RESOURCES
41
What constitutes knowledge of forest genetic resources?
Availability of information on genetic resources
42
46
Drivers of change and trends affecting forest genetic resources
CHAPTER 5
CHAPTER 6
DRIVERS OF CHANGE
51
Forest conversion and expansion of crop land
Demand for energy
Unsustainable harvesting and use
Livestock and browse animals
Climate change
Changed ire regimes
Invasive species
Genetic pollution
51
52
53
54
54
58
59
62
GLOBAL FOREST TRENDS AFFECTING FOREST GENETIC
DIVERSITY
65
Forest trends
Consequences of forest changes for genetic diversity
65
70
Part 3
Current and emerging technologies
CHAPTER 7
CHAPTER 8
CHAPTER 9
Part 4
TRAIT-BASED KNOWLEDGE OF TREE GENETIC RESOURCES
79
Indigenous and traditional knowledge
Classical tree improvement
Participatory tree domestication
79
83
89
MODERN ADVANCES
91
Population genetics based on molecular markers
Genomic advances
Combining molecular tools with tree improvement:
marker-assisted selection
Genetic modiication
91
94
96
98
APPLICATION OF GENETIC KNOWLEDGE IN FOREST
CONSERVATION
101
Combining spatial analysis with genetic markers to prioritize
conservation
Research on climate change and forest genetic resources
Genetic technologies for reducing illegal logging
102
103
104
State of forest genetic resources conservation and
management
CHAPTER 10 HOW COUNTRIES MANAGE AND CONSERVE THEIR FOREST
GENETIC RESOURCES
111
Features of effective and comprehensive FGR conservation and
management systems
Approaches to FGR conservation in relation to biodiversity
conservation strategies
National strategies and programmes for FGR conservation and
management
Prioritizing species for FGR conservation and management
CHAPTER 11 CHARACTERIZATION OF GENETIC VARIABILITY AND
MONITORING OF CHANGE
Characterizing interspeciic variability
Characterizing intraspeciic variation
Monitoring of forest genetic resources
Differences among countries and regions in characterization
of FGR
vi
112
114
116
116
121
123
124
131
133
CHAPTER 12 IN SITU FGR CONSERVATION AND MANAGEMENT
Protected areas
In situ conservation outside protected areas
Formal in situ FGR conservation programmes
Forest restoration and FGR
Opportunities from climate change initiatives: restoration and
connectivity for in situ FGR
In situ conservation through sustainable forest management
CHAPTER 13 EX SITU CONSERVATION
Ex situ conservation activities by region
CHAPTER 14 GENETIC IMPROVEMENT AND BREEDING PROGRAMMES
Improvement approaches
Administration and coordination of breeding and
improvement programmes
Prioritizing uses, traits and species for improvement
The state of tree improvement and species priorities by region
International collaboration and donor programmes
for tree improvement
A cautionary note: potential threats to FGR from breeding
and improvement programmes
CHAPTER 15 GERMPLASM DELIVERY AND DEPLOYMENT
Uses of germplasm and plant materials
Demand for germplasm and planting materials
Actors involved in production, distribution and deployment
Production of germplasm and planting materials
Movement and transfer of genetic material
Information management in delivery and deployment of
germplasm
International assistance
CHAPTER 16 INSTITUTIONAL FRAMEWORK FOR CONSERVATION
AND MANAGEMENT OF FOREST GENETIC RESOURCES
National institutions dealing with forest genetic resources
Legal and policy framework
Education and training
Research
Raising public awareness and communication
Support to forest genetic resources
International and regional collaboration
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138
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152
163
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Part 5
Needs, challenges and required responses for the future
CHAPTER 17 PRACTICES AND TECHNOLOGIES FOR IMPROVED
MANAGEMENT OF FOREST GENETIC RESOURCES
Monitoring
In situ conservation
Ex situ conservation
Domestication, breeding and improvement
Germplasm delivery and deployment
Assisted migration to accelerate adaptation to climate change
215
218
222
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225
CHAPTER 18 POLITICAL AND INSTITUTIONAL RECOMMENDATIONS
227
National polices and institutions
Capacity building
Improving information availability and access
Priority areas for research
Communication and awareness raising
In conclusion: what needs to be done
227
230
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232
234
235
References
Acronyms and abbreviations
viii
215
237
275
BOXES
Box 1.1
Box 1.2
Box 2.1
Box 3.1
Box 3.2
Box 3.3
Box 3.4
Box 4.1
Box 5.1
Box 5.2
Box 5.3
Box 6.1
Box 6.2
Box 7.1
Box 7.2
Box 8.1
Box 10.1
Box 10.2
Box 12.1
Box 12.2
Box 12.3
Box 14.1
Box 14.2
Box 15.1
Box 15.2
Box 16.1
Box 18.1
Box 18.2
Box 18.3
The Commission on Genetic Resources for Food and Agriculture
Examples of some of the oldest known trees and woody shrubs
Conserving distinct and unique tree lineages
Valuing non-wood forest products demand
Application of genetic principles in forest ecosystem restoration
and management
Evolving use of tree germplasm in modern agroforestry in South Paciic islands
Millennium Seed Bank Partnership
Biological models for predicting risk associated with seed storage for
tree species
Filling the knowledge gap in botany: how many tree species are there
on Earth?
Selecting for salt tolerance: one way to address impacts of sea-level rise on
coastal forests
Predicting impacts of climate change on distribution of forest insect pests
Some destructive pathogens in Northern Hemisphere forests
Conservation status of forest species assessed under the Global Trees Campaign
Loss of intraspeciic diversity in valuable species: some examples
Adapted social structures underlie resilient societies
Research organizations historically active in work on forest genetics
Use of genomic tools in Eucalyptus spp.
Contextual features that inluence a country’s system of FGR conservation and
management
Summary: how FGR conservation approaches differ from usual biodiversity
conservation approaches
Community and participatory management
Global Objectives of the Non-Legally Binding Instrument on All Types of Forests
Addressing genetic resources in sustainable forest management plans
Pinus radiata – a species improved outside its native range
Early tree breeding programmes in Canada and the United States of America
Germplasm production, storage and progagation and distribution facilities:
some challenges
Germplasm production and dissemination in Ethiopia
Example of an international FGR network: the European Forest Genetic
Resources Programme (EUFORGEN)
Integrating forest genetic resources in international forest and natural
resource management policy framework
Regional collaboration in FGR conservation and management: joint strategies
and priorities
The state of knowledge on forest genetic resources: a summary
xv
9
16
21
29
31
32
34
43
57
58
62
72
75
82
84
98
113
115
152
153
154
177
185
196
197
211
228
229
233
ix
TABLES
Table 1.1
Table 1.2
Table 2.1
Table 6.1
Table 6.2
Table 7.1
Table 7.2
Table 8.1
Table 8.2
Table 9.1
Table 11.1
Table 13.1
Table 13.2
Table 13.3
Table 17.1
Table 17.2
x
Main types of forest and tree resources management
Life span of some of the longest-lived conifers
Value of removals of plant-based NWFPs (and bee products) by category
and region
Area of primary forest change, 1990−2010
The ten countries with the largest annual net loss of forest area, 1990–2010
Amount of environmental difference needed to show a genetic difference
in some conifers
Evidence from reciprocal transplant studies showing local sources as
optimal or near optimal
Indicative studies of tropical tree species using molecular markers since 1990
Number of published successful transgenic experiments achieving gene
expression or overexpression in transgenic cells, by tree species or genus
and by modiication objective
Examples of the use of DNA and markers to control illegal logging
Characters most frequently assessed in 692 evaluations of genetic variability
reported by countries
Species conserved ex situ, by region
Genera of global priority that are conserved ex situ
Wild relatives of fruit-tree crop species reported by Jordan as present but
understudied in terms of ex situ conservation
Potential local- to global-scale operational indicators of forest genetic
diversity, with veriiers
Some constraints, needs, priorities and opportunities identiied by
countries for in situ FGR conservation and management
5
10
21
66
66
88
88
93
99
105
131
163
164
170
216
218
FIGURES
Figure 1.1
Figure 4.1
Figure 4.2
Figure 5.1
Figure 6.1
Figure 6.2
Figure 6.3
Figure 6.4
Figure 6.5
Figure 8.1
Figure 9.1
Figure 10.1
Figure 10.2
Figure 12.1
Figure 12.2
Figure 12.3
Figure 14.1
Figure 14.2
Figure 15.1
Figure 15.2
Countries reporting for The State of the World’s Forest Genetic Resources
Number of species and subspecies mentioned as actively managed in
country reports, by region
Proportion of the world’s plants in accessible plant lists
Example of a species distribution map: Pinus sylvestris in Europe
Changes in area of cropland, 2000−2010
Characteristics of the world’s forests in 2010
Proportion of planted forest area made up of exotic species
Designated functions of forests reported in the Global Forest Resources
Assessment 2010
Number of species and subspecies mentioned as threatened (at various levels)
in country reports, by region
Main threats to 52 endangered tree species proiled by the Global Trees
Campaign
Major categories of forest biotechnology activities
Genetic reference map for Swietenia macrophylla (mahogany) in
Latin America
Reasons for nominating species for priority for FGR conservation and
management
Most common priority species, by region
Number of species and subspecies conserved in situ and ex situ, by region
Reasons cited by countries for conserving species in situ
Number of species mentioned as actively managed in country reports,
by main management objective
Most common species in tree improvement and conservation programmes
worldwide
Number of species and subspecies in improvement programmes, by region
Most widely planted species in seed orchards
Purposes of germplasm transfer reported by countries
xviii
13
43
47
52
68
69
69
73
73
91
107
117
119
138
147
155
178
179
195
204
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Acknowledgements
T
his report has been made possible thanks to the contribution of time, energy and
expertise of many individuals, and the collaboration and support of governments and
partner institutions. FAO would like to take this opportunity to acknowledge these
contributions.
The report was prepared by the FAO Forest Assessment, Management and Conservation
Division. The core team was coordinated by Oudara Souvannavong and included Albert
Nikiema and Judith Nantongo. The inalization was supervised by Albert Nikiema.
The country reports submitted by governments were the main source of information for
The State of the World’s Forest Genetic Resources. FAO wishes to thank the governments and
all the individuals involved, in particular the National Focal Points, for their contributions
on the status of forest genetic resources in their countries.
The preparation of the report would not have been possible without the generous
inancial support provided by the Governments of Germany, Norway, Spain and Sweden,
and the in-kind contributions from the Governments of Canada and Uruguay.
During the entire report preparation process FAO beneited from the very close
collaboration of the Bioversity International FGR team composed of Judy Loo, Laura Snook
and Jarkko Koskela, and the World Agroforestry Centre (ICRAF) team comprising Ramni
Jamnadass and Ian Dawson.
The development of the guidelines for preparation of country reports, the organization
of training workshops and other international, regional and national consultations and
follow-up workshops involved the following team of FAO Oficers at headquarters and
in decentralized ofices, visiting scientists and experts from regional and international
institutions and networks: Moujahed Achouri, Nuria Alba, Ricardo Alia, Simmathiri
Appanah, Nabil Assaf, Daniel Baskaran, Zohra Bennadji, Nora Berrahmouni, Foday Bojang,
Sairusi Bulai, Rene Czudek, Ian Dawson, Patrick Durst, Claus Eckelmann, Lay-Thong Hong,
Benoit Horemans, Riina Jalonen, Ramni Jamnadass, Edward Kilawe, Judy Loo, Aru Mathias,
Mario Mengarelli, Judith Nantongo, Jean-Claude Nguinguiri, Albert Nikiema, Hivy Ortiz,
Cenon Padolina, José Antonio Prado, Dan Rugabira, Mohamed Saket, Aubain Saya, HeokChoh Sim, Oudara Souvannavong, François Tapsoba, Maarten van Zonneveld, Barbara
Vinceti, Norbert Winkler and Ekrem Yazici.
A series of regional consultation workshops was organized to share preliminary
indings of country reports, discuss regional syntheses and adopt recommendations on
needs and priorities for action. This approach greatly contributed to the preparation of
the report and FAO wishes to acknowledge the participants, including representatives of
82 countries, the National Focal Points, partner institutions and other interested parties.
For the organization of the regional workshops, FAO also wishes to acknowledge the
generous support and the active collaboration of a number of organizations: African
Union Commission; Asia Paciic Association of Forestry Research Institutions (APAFRI); Asia
Paciic Forest Genetic Resources Programme (APFORGEN); Bioversity International; Central
African Forests Commission (COMIFAC); Kenyan Forest Research Institute (KEFRI); Latin
American Forest Genetic Resources Network (LAFORGEN); National Tree Seed Centre of
Burkina Faso (CNSF); Organisation for Economic Co-operation and Development (OECD)
Secretariat; Royal Botanic Gardens, Kew, United Kingdom; Secretariat of the Convention on
Biological Diversity (CBD); Secretariat of the Paciic Community (SPC); State Committee for
Environment Protection of the Republic of Tajikistan; Sub-Saharan African Forest Genetic
xiii
Resources Network (SAFORGEN); Wallonie-Bruxelles International; World Agroforestry
Centre (ICRAF); and World Wide Fund for Nature (WWF).
Regional synthesis reports, which provided useful underpinning for the publication,
were compiled by Fadi Asmar, Natalia Demidova, Jerome Duminil, Riina Jalonen, Judith
Nantongo, Moussa Ouedraogo and José Antonio Prado.
Data entry and analysis of information from country reports was done by Albert Nikiema
with the assistance of Judith Nantongo and Laura D’Aietti. Maps were prepared by Laura
D’Aietti.
The report was written by Tannis Beardmore, Judy Loo, Steve Mathews, Katherine
McGovern, Albert Nikiema, Oudara Souvannavong, Mary Taylor and Lex Thomson.
Comments by members of the Commission on Genetic Resources for Food and Agriculture
are gratefully acknowledged. Contributions to the text were made by Henri-Noël Bouda,
Stephen Cavers, Jerome Duminil, Marius Ekue, Bruno Fady, Hugh Pritchard, Evert Thomas
and Maarten van Zonneveld.
The draft report was reviewed by numerous countries and experts. It is not possible to
name each of them here, but their contribution is deeply appreciated. Thanks are due to
Ian Thompson for carrying out the inal review of the completed report.
The inal editing was achieved by Andrea Perlis and coordinated by Suzanne Lapstun.
Additional editorial assistance was provided by Miriam Jones. Layout and graphic design
were the work of Joanne Morgante with assistance from Roberto Cenciarelli. The inal
publication of the report was supervised by Douglas McGuire.
Throughout the report preparation process, continuous support and encouragement
were received from the Secretariat of the Commission on Genetic Resources for Food and
Agriculture as well as the Directors of the Forest Assessment, Management and Conservation
Division: José Antonio Prado and Eduardo Mansur. Administrative support was provided by
Mustapha Kerdi, Joy Taylor and Sugi Yoo.
Listing every person by name is not easy and carries with it the risk that someone may be
overlooked. Apologies are conveyed to anyone who may have provided assistance whose
name has been inadvertently omitted.
xiv
About this publication
T
he State of the World’s Forest Genetic Resources addresses the conservation,
management and sustainable use of forest tree and other woody plant genetic
resources of actual and potential value for human well-being in the broad range of
management systems.
This report complements two other FAO lagship publications in the ield of forestry, the
annual State of the World’s Forests and the periodic Global Forest Resources Assessment
(FRA). State of the World’s Forests reports on the status of forests, recent major policy
and institutional developments and key issues concerning the forest sector. FRA provides
comprehensive data on forest distribution and status, including on matters inluencing
forest genetic resource (FGR) conservation and management, such as indicators of
sustainable forest management, extent of permanent forest estate and protected areas,
and regeneration methods used. However, forest cover and related data cannot be used as
a surrogate for assessment of the status of FGR. This irst edition of The State of the World’s
Forest Genetic Resources will help to differentiate between the state of the world’s forest
resources and the state of the genetic resources on which they depend for their utility,
adaptability and health.
The State of the World’s Forest Genetic Resources also complements two lagship
publications in the ield of genetic resources for food and agriculture: The State of the
World’s Animal Genetic Resources for Food and Agriculture, published in 2007, and The
Second Report on the State of the World’s Plant Genetic Resources for Food and Agriculture,
published in 2010. The three reports have in common that they have been initiated by and
prepared under the guidance of FAO’s Commission on Genetic Resources for Food and
Agriculture (the Commission) (see Box).
This is the irst synthesis of its kind for FGR and constitutes a baseline against which
future status assessments can be compared. Sources of information include national
reports prepared by countries, regional summaries prepared following regional workshops
and commissioned thematic studies (see section on the reporting and preparatory process
The Commission on Genetic Resources for Food and Agriculture
With its 177 member countries, the
Commission on Genetic Resources for Food
and Agriculture offers an intergovernmental
forum where global consensus can be reached
on policies relevant to biodiversity for food
and agriculture. The main objective of the
Commission is to ensure the conservation and
sustainable use of genetic resources for food
and agriculture and the fair and equitable
sharing of beneits derived from their use, for
present and future generations.
The work of the Commission focuses on
developing and overseeing the implementation
of policies and supporting initiatives that
not only raise awareness but also seek ways
to solve emerging problems. It guides the
preparation of periodic global assessments
of the status and trends of genetic diversity,
the threats facing genetic diversity and
the measures being taken to promote its
conservation and sustainable use. The
Commission also negotiates global action
plans, codes of conduct and other instruments
relevant to the conservation and sustainable
use of genetic resources for food and
agriculture.
xv
below) as well as published literature. Of necessity the treatment of knowledge of FGR in
this report is selective, as volumes would be required to capture all the available knowledge.
Part 1 provides an overview of basic deinitions and concepts in forest genetics, the value
of FGR (recognition of which is essential to ensuring their conservation), FGR conservation
approaches, and the state of knowledge and information in this ield.
Part 2 addresses drivers of change, including forest conversion and expansion of crop
land, demand for wood energy, and unsustainable harvesting and use. Next it presents
the global forest trends affecting FGR, with data on trends in forest cover, biodiversity
conservation and ownership, drawn primarily from FRA 2010. The consequences of these
trends for FGR are also outlined, in terms of loss of ecosystems, tree species and intraspeciic
diversity.
Part 3 examines the current and emerging technologies in the ield of FGR, including
information on indigenous and traditional knowledge and classical tree improvement,
modern advances including the use of molecular markers and genetic modiication, and
FGR conservation-related knowledge. This part also addresses research on climate change
and FGR and genetic technologies for reducing illegal forest harvesting.
Part 4 reviews the state of FGR conservation and management. This part, based on
the reports submitted by countries, addresses countries’ FGR strategies and programmes,
their progress in characterizing their genetic diversity, and the state of in situ and ex
situ conservation and management. It also reviews progress in breeding and genetic
improvement of forest tree species, and the systems for producing and distributing forest
genetic materials for use on farms, in natural and planted forests and in research. Finally, it
presents the state of the institutional framework for FGR conservation and management,
including national institutions dealing with FGR, policies and laws, education and training,
research, communication and public awareness raising, and international and regional
collaboration, including networks.
The concluding section, Part 5, addresses the needs and responses required to improve
FGR conservation and management in the future. It provides recommendations for
improving practices and technologies in FGR conservation and management; and for
enhancing national polices and institutions, capacity building, knowledge and information
availability, and public awareness for improved conservation and management of FGR
worldwide.
The reporting and preparatory process
In considering the status of conservation and use of forest genetic resources, the
Commission, at its eleventh regular session in 2007, emphasized their importance for food
security, poverty alleviation and environmental sustainability and recognized that the lack
of information limits international, regional and local decision-making and action on these
vital resources. The Commission included the preparation of The State of the World’s Forest
Genetic Resources in its Multi-year Programme of Work and requested that FAO begin to
prepare it. The Commission’s request was supported by the FAO Committee on Forestry
(COFO) at its nineteenth session (March 2009). At its twelfth regular session (October 2009),
the Commission endorsed a proposed outline of the report and agreed on an indicative
timeline and the process for country involvement. The Commission also established an
Intergovernmental Technical Working Group on Forest Genetic Resources (ITWG-FGR) to
xvi
address issues relevant to the conservation and sustainable use of FGR, and to advise and
make recommendations on the report preparation process.
In April 2010, following the process established by the Commission, FAO invited countries
to nominate National Focal Points and to prepare and submit country reports, which have
been the main source of information for the preparation of The State of the World’s Forest
Genetic Resources following a country-driven process. FAO provided guidelines for the
preparation of the country reports, including a recommended structure and methodology
(FAO, 2011). The guidelines suggested that the preparation of country reports represented
an opportunity to conduct a national strategic exercise to assess the status of FGR in the
countries and to relect on needs and priorities for their conservation and sustainable use.
A participatory approach, engaging a wide range of stakeholders, was encouraged.
At its twentieth session in October 2010, COFO welcomed the initiative to develop The
State of the World’s Forest Genetic Resources and recommended that FAO continue this
important effort. It also invited the governing bodies of the member organizations of the
Collaborative Partnership on Forests to consider the information and analysis provided by
FRA and this report in their work.
From November 2010 to September 2011, FAO organized regional and subregional
workshops to train National Focal Points and other national and regional experts on the
preparation of country reports following the approach promoted in the guidelines. These
workshops were organized in collaboration with international partners such as Bioversity
International, the World Agroforestry Centre (ICRAF), the Secretariat of the Convention on
Biological Diversity (CBD) and the World Wide Fund for Nature (WWF), as well as regional
institutions such as the Central African Forests Commission (COMIFAC) and the Secretariat
of the Paciic Community (SPC) and regional networks and programmes such as the Asia
Paciic Association of Forestry Research Institutions (APAFRI), the Asia Paciic Forest Genetic
Resources Programme (APFORGEN), the Latin American Forest Genetic Resources Network
(LAFORGEN) and the Sub-Saharan African Forest Genetic Resources Network (SAFORGEN).
The workshops covered 82 countries and gathered 137 experts.
A total of 86 countries submitted reports (see Figure), accounting for 76 percent of
the world’s land area and 85 percent of the global forest area, with good latitudinal and
ecoregional representation.
Five thematic studies were prepared on issues relevant to the conservation and sustainable
use of FGR at the global level:
• Indicators of forest genetic diversity, erosion and vulnerability
• Role of FGR in adaptation to biotic and abiotic factors in a changing climate
• Trees, tree genetic resources and the livelihoods of rural communities in the tropics
• Genetic considerations in ecosystem restoration using native tree species
• Genetic effects of forest management practices
The country reports and the thematic background studies prepared for The State of the
World’s Forest Genetic Resources will be made available on a dedicated page on FAO’s
website.
A draft of the present report was reviewed by ITWG-FGR at its second session in January
2013 and presented to the Commission at its fourteenth regular session in April of the same
year. Countries were invited to provide comments on the inal draft, which were taken into
consideration in the inalization of the report.
xvii
Countries reporting for The State of the World’s Forest Genetic Resources
Africa (31 countries)
Algeria, Benin, Burkina Faso, Burundi,
Cameroon, Central African Republic, Chad,
Democratic Republic of the Congo, Ethiopia,
Gabon, Ghana, Kenya, Lesotho, Madagascar,
Malawi, Mali, Mauritania, Mauritius, Morocco,
Niger, Republic of the Congo, Senegal,
Seychelles, Somalia, South Africa, Sudan,
Swaziland, Tunisia, United Republic of Tanzania,
Zambia, Zimbabwe
Asia (14 countries)
Azerbaijan, China, India, Indonesia, Japan,
Kazakhstan, Kyrgyzstan, Myanmar, Nepal,
Philippines, Republic of Korea, Sri Lanka,
Thailand, Uzbekistan
Latin America and the Caribbean (9 countries)
Argentina, Brazil, Chile, Costa Rica, Ecuador,
Guatemala, Mexico, Panama, Peru
Near East (6 countries)
Egypt, Iran, Iraq, Jordan, Lebanon, Yemen
North America (2 countries)
Canada, United States of America
Oceania (6 countries)
Australia, Cook Islands, Fiji, Papua New Guinea,
Solomon Islands, Vanuatu
Europe (18 countries)
Austria, Bulgaria, Cyprus, Denmark, Estonia,
Finland, France, Germany, Hungary, Ireland,
Netherlands, Norway, Poland, Russian
Federation, Spain, Sweden, Turkey, Ukraine
Note: The region referred to as Oceania – for consistency with data reported in the Global Forest Resources Assessment
and State of the World’s Forests – is synonymous with the Commission’s South West Paciic region.
xviii
Based on the indings of The State of the World’s Forest Genetic Resources, ITWG-FGR
and subsequently the Commission agreed on strategic priorities for forest genetic resources,
adopted by the Conference of FAO at its thirty-eighth session in June 2013 as the Global
Plan of Action for the Conservation, Sustainable Use and Development of Forest Genetic
Resources. This Global Plan of Action identiies 27 strategic priorities grouped into four
areas:
• improving the availability of, and access to, information on FGR;
• conservation of FGR (in situ and ex situ);
• sustainable use, development and management of FGR;
• policies, institutions and capacity building.
Implementation of the Global Plan of Action will strengthen the sustainability of forest
management while contributing towards the Millennium Development Goals, the post2015 development agenda and the Aichi Biodiversity Targets.
xix
Executive summary
F
orests and trees enhance and protect landscapes, ecosystems and production systems.
They provide goods and services which are essential to the survival and well-being of
all humanity. Forest genetic resources (FGR) are the heritable materials maintained
within and among tree and other woody plant species that are of actual or potential
economic, environmental, scientiic or societal value. FGR are essential for the adaptation
and evolutionary processes of forests and trees as well as for improving their productivity.
The world’s current population of 7.2 billion is projected to reach 9.6 billion by 2050.
Along with population growth, the demand for energy and wood products for both
industrial and domestic uses is expected to increase by 40 percent in the next 20 years. The
demand for other forest-related goods (food, medicine, fodder and other commodities) is
also predicted to increase.
A major consequence of population pressure is land-use change. Forest conversion to
crop and pasture land, together with overexploitation, selective harvesting and high tree
mortality due to extreme climatic events, in combination with regeneration failure, can
result in local population extinction and the loss of FGR.
Conservation and sustainable management of FGR is therefore a must to ensure that
present and future generations continue to beneit from forests and trees.
The State of the World’s Forest Genetic Resources
This irst The State of the World’s Forest Genetic Resources constitutes a major step in
building the information and knowledge base required for action towards better
conservation and sustainable management of FGR at national, regional and international
levels.
The report was prepared based on information provided by 86 countries, outcomes from
regional and subregional consultations and information compiled in thematic studies. It
includes:
• an overview of deinitions and concepts related to FGR and a review of their value;
• a description of the main drivers of change;
• the presentation of key emerging technologies;
• an analysis of the current status of FGR conservation, use and related developments;
• recommendations addressing the challenges and needs.
Preparation of the report
Recognizing that the lack of information limits the capacity of decision-makers to determine
the action needed on FGR at the international, regional and local levels, the Commission on
Genetic Resources for Food and Agriculture (the Commission), at its eleventh session (2007),
emphasized the importance of FGR for food security, poverty alleviation and environmental
sustainability. The Commission stressed the urgency of addressing the conservation and
sustainable use of FGR through sustainable forest management, especially those resources
that are under threat at the global level, and requested FAO to prepare a report on the
state of the world’s FGR based on country reports.
To assist countries in their reporting, FAO carried out regional training workshops which
covered 82 countries and gathered 137 experts. A total of 86 countries submitted reports,
accounting for 76 percent of the world’s land area and 85 percent of the global forest area.
The Commission established an Intergovernmental Technical Working Group on Forest
Genetic Resources.
xxi
The state of knowledge of forest genetic resources: A summary
• Knowledge of FGR is reported to be
inadequate for well-informed policy or
management in most countries.
• Studies have described genetic parameters
for less than 1 percent of tree species,
although both the number of studies and the
number of species studied have increased
signiicantly in the past decade.
• Most studies conducted during the past
two decades have been at the molecular
level, either using DNA markers or
genomic technologies to characterize
genetic resources. Molecular information
is accumulating much faster than wholeorganism information, with the consequence
that little of the accumulating knowledge
has direct application in management,
improvement or conservation.
• A few species have been well researched
– through both molecular and quantitative
studies – and genetically characterized;
these mainly comprise temperate conifers,
eucalypts, several acacias, teak and a few
other broadly adapted, widely planted and
rapidly growing species.
• Quantitative genetic knowledge has led
to signiicant productivity gains in a small
number of high-value planted timber species.
• Genomic knowledge of forest trees lags
behind that of model herbaceous crop
species, including the important agricultural
crops, but for several tree species the entire
genome has been or is in the process of
being sequenced, and novel approaches have
been developed to link markers to important
traits. Genomic or marker-assisted selection
is close to being realized, but phenotyping
and data management are the biggest
bottlenecks.
• Many of the species identiied as priorities,
especially for local use, have received little
or no research attention, indicating a need
to associate funding with priority-setting
exercises.
The draft report was reviewed by the working group, the Commission and individual
experts; it was inalized by FAO incorporating the comments received. Based on the indings
of The State of the World’s Forest Genetic Resources, the Commission agreed on strategic
priorities at the national, regional and international levels. In 2013, the Conference of FAO
adopted these priorities as the Global Plan of Action for the Conservation, Sustainable Use
and Development of Forest Genetic Resources.
Key indings
Access to information and knowledge on FGR needs to be improved
Adequate management of FGR requires the availability of accurate knowledge and
information on ecosystems and species. Although a range of 80 000 to 100 000 is the most
widely used estimate for the number of tree species, the range of published estimates is
much wider, from 50 000 to 100 000, indicating the need for further efforts in botanic
assessment to obtain more accurate igures.
The status of botanical knowledge varies from country to country. Very few countries
have detailed tree species checklists that include species characteristics allowing distinction
between different plant life forms, e.g. trees, shrubs, palms and bamboo. Information on
the conservation status of species populations is not available in many countries.
xxii
The country reports mention 8 000 species of trees, s, palms and bamboo; of these,
genetic-level information is available for only 500 to 600 species.
The collaborative development of an FGR database is urgently needed to enhance access
to valuable information and avoid duplication efforts and waste of resources.
Economic value is the main factor in setting management priorities
Priority setting is fundamental to effective FGR conservation and management, given the
vast number of tree and other woody species and the typically considerable intraspeciic
variation across their natural range. Reasons for nominating species as priorities include
their economic value (timber, pulp, food, wood energy, and non-wood forest products),
social and cultural value, conservation value (biodiversity, threatened species, endemic
species, genetic conservation, scientiic value), environmental value (e.g. soil and water
protection, soil fertility and watershed management) and invasiveness.
Results from the country reports indicate economic and conservation value as the two
main reasons for nominating species for priority for FGR conservation and management;
each accounts for two-thirds of species nominations.
Half of the forest species reported by countries are threatened
Loss of plant species or species genetic erosion in forest ecosystems is mostly due to
conversion of forest to other land use types, overexploitation and effects of climate. The
proportion of threatened species reported by the countries varies widely, from 7 percent
in Oceania to 46 percent in North America. However, some countries included threats at
population level, which may account for the great variation in number of threatened
species reported.
8 000 forest species are used and one-third of them actively managed
Of the 8 000 species of trees, shrubs, palms and bamboo cited in country reports, around
2 400 are mentioned as actively managed, in other words managed speciically for their
products and/or services.
The main products and functions targeted through management activities are reported
by the countries as timber (42 percent), non-wood forest products (41 percent) and energy
(mainly fuelwood) (19 percent).
The high number of species used and their multiplicity of products and services indicates
the enormous value of FGR; it suggests their great potential to support agriculture, forestry
and environmental sustainability, as well as food and nutrition security, if better evaluated
and developed.
Species distribution maps are vital, but rarely available
Adequate management of FGR and monitoring of their in situ conservation status requires
reliable baseline information. Development of species distribution maps showing locations
of all populations is an essential step in conservation. However, not many countries have
the resources to include the development of such maps in their conservation strategies.
Mapping at the regional level can make it possible to cover a large portion if not all of a
species’ distribution range.
xxiii
Most species are conserved in situ, in naturally regenerated
and planted forests
FGR management actions are usually undertaken at forest ecosystem, species (interspeciic)
or genetic (intraspeciic) levels. FGR are to a large extent preserved in wild populations
and managed in naturally regenerated forest except for some commercial wood-producing
genera and species undergoing intensive tree breeding (e.g. Acacia spp., Eucalyptus spp.,
Populus spp., Pinus spp. and Tectona grandis).
In many countries plant wild populations and crop wild relatives are conserved in
protected areas and/or in naturally regenerated forest lands. Examples include Malus spp.
in central Asia, Coffea arabica in Ethiopia and Eucalyptus spp. in Australia.
In addition, farmers contribute to the conservation of populations of many tree species
through traditional agroforestry practices. Vitellaria spp. (shea) is an example from semiarid tropical Africa.
Effective ex situ conservation programmes are restricted to limited
species and populations
Ex situ conservation programmes remain conined to some economically important species
undergoing intensive breeding or under serious threat with high inancial implications.
The Millennium Seed Bank Partnership, based in Kew, United Kingdom, hosts the world’s
largest collection of wild plant species in long-term seed storage. It covers 10 percent of
the world’s wild plant species – including many woody species – and aims to conserve
25 percent by 2020.
Of the 2 400 actively managed species, about 700 are managed in planted forests and
approximately the same number is included in tree improvement programmes. In some
countries planted forests and trials contribute to ex situ conservation programmes.
Tree improvement greatly enhances productivity and offers potential
for adaptation to changing climate
In recent decades government agencies and the private sector have subjected a wider range
of tree species to domestication and formal breeding programmes to produce timber, pulp,
fuelwood and non-wood forest products and to provide forest service functions. Tree
breeding programmes have the potential to improve the production of planted forests
and trees in a sustainable way and are necessary to meet growing global demand for
forest products and services. Through tree improvement programmes, productivity can be
increased by 10 to more than 60 percent depending on the targeted products (wood, fruit,
leaves, resins) and the species.
Examples of tree species in countries’ intensive selection and breeding programmes
include Eucalyptus spp., Pinus spp., Populus spp. and Tectona grandis. Hybrid breeding is
used in many countries to produce trees with superior productive capabilities (through
heterosis) and also to introduce genes for disease resistance. Examples include eucalypt
hybrids, Larix and Populus hybrids and Pinus hybrids.
Tree improvement also has an important role in targeting traits suitable for adaptation
to varying environmental conditions, including those associated with climate change.
These efforts rely on improved understanding of the genetic structure within and between
species populations.
xxiv
Emerging technology opens new avenues in FGR management
and conservation
An array of biotechnological tools are contributing to the knowledge of forest genetic
resources. For natural forests, biotechnology contributes to the knowledge of genetic
variation within and between species populations. In tree improvement programmes,
biotechnology tools such as enhanced vegetative propagation techniques and markerassisted tree selection are making signiicant contributions. Genomics is also being used in
forestry as a tool to enhance conservation, for example through the development of DNA
banks. Biotechnology offers innovative means of controlling illegal forest harvesting, with
DNA ingerprints now used in timber tracking. Genetic modiication has been explored to
increase or improve wood production in a few countries. However, no commercial planting
has been reported.
Of the over 700 tree species reported by countries as subject to tree improvement
programmes, 241 species are included in biotechnology research. The development of
large-scale clonal plantations of some economically important species (e.g. Eucalyptus
spp., Tectona grandis) using biotechnology has been reported by a number of countries,
including tropical countries.
Policies and institutional frameworks are insuficient
Because of insuficient awareness on the importance of forest genetic resources in
improving forest production, enhancing ecosystems and improving adaptation of tree
species to changing environmental conditions, national policies and regulatory frameworks
for FGR are, in general, partial, ineffective or non-existent. Most developing countries lack
the funding and the institutional and technical capacities required to address FGR issues.
The institutional and policy framework therefore needs to be improved to address the
constraints related to the conservation, sustainable use and development of FGR. Many
countries identify integration of FGR concerns into broader forest-related policy as a
priority.
What needs to be done?
Improve the availability of, and access to, information on FGR
• Establish and strengthen national FGR assessment, characterization and monitoring
systems.
• Develop national and subnational systems for the assessment and management of
traditional knowledge on FGR.
• Develop international technical standards and protocols for FGR inventory,
characterization and monitoring of trends and risks.
• Promote the establishment and reinforcement of FGR information systems
(databases) to cover available scientiic and traditional knowledge on uses,
distribution, habitats, biology and genetic variation of species and species
populations.
Enhance in situ and ex situ conservation of FGR
• Strengthen the contribution of primary forests and protected areas to in situ
conservation of FGR.
xxv
• Promote the establishment and development of eficient and sustainable ex situ
conservation systems, including in vivo collections and gene banks.
• Support and strengthen the role of indigenous and local communities in the
sustainable management and conservation of FGR.
• Identify priority species for action.
• Harmonize measures for in situ and ex situ conservation, including through regional
cooperation and networking.
Improve sustainable use and management of FGR
• Develop and reinforce national seed programmes to ensure the availability of
genetically appropriate tree seeds in the quantities and of the quality needed for
national plantation programmes.
• Promote restoration and rehabilitation of ecosystems using genetically appropriate
material.
• Support climate change adaptation and mitigation through proper management and
use of FGR.
• Promote good practices and appropriate use of emerging technology to support the
conservation, development and sustainable use of FGR.
• Develop and reinforce research programmes on tree breeding, domestication and
bioprospecting.
• Develop and promote networking and collaboration among concerned countries to
combat invasive species affecting FGR.
Strengthen policies and institutional capacity
• Develop national strategies for in situ and ex situ conservation and sustainable use of
FGR.
• Integrate FGR conservation and management into wider policies, programmes and
frameworks of action at the national, regional and global levels.
• Develop collaboration and promote coordination of national institutions and
programmes related to FGR.
• Establish and strengthen educational and research capacities on FGR.
• Promote the participation of indigenous and local communities in FGR management
in the context of decentralization.
• Promote and apply mechanisms for regional germplasm exchange for research and
development, in agreement with international conventions.
• Reinforce regional and international cooperation, including networking, to support
education, knowledge dissemination, research, and conservation and sustainable
management of FGR.
• Promote public and international awareness of the roles and value of FGR.
• Strengthen efforts to mobilize the necessary resources, including inancing, for the
conservation, sustainable use and development of FGR.
xxvi
Part 1
OVERVIEW
OVERV IE W
Chapter 1
Basic concepts
Genetic diversity provides the fundamental basis
for the evolution of forest tree species and for
their adaptation to change. The enormous range
of goods and services provided by trees and forests
is both a function of and testimony to the genetic
variability contained within them. Conserving
forest genetic resources (FGR) is therefore vital,
as FGR constitute a unique and irreplaceable
resource for the future, including for sustainable
economic growth and progress and environmental
adaption. The sustainable management of forests
and of trees in agroforestry systems requires a
better understanding of the speciic features
of forest trees and their genetic diversity, and
how they can be best conserved, managed and
utilized. Forest tree species are generally long
lived and extremely diverse. One species can
naturally occur in a broad range of ecological
conditions. In addition, many forest species have
evolved under several periods of major climatic
change, and their genetic variability is needed
for adaptation to climatic regimes different from
those in which they have evolved. FGR have
provided the potential for adaptation in the
past, and will continue to play this vital role as
humankind addresses the challenge of mitigating
or adapting to further climate changes.
Forestry practices that maintain genetic diversity
over the long term will be required as an integral
component of sustainable forest management.
In future more proactive management of
FGR may be needed to accelerate adaptation
of forest trees to climate change including
through breeding and deliberate movement and
relocation of germplasm. Much remains to be
discovered concerning how genes function and
are regulated in different tree species and further
research will likely yield indings of immense
economic, social and environmental importance.
As a precautionary principle, until there is an
improved understanding of tree genetics, there is
a need to conserve as much FGR as possible, i.e.
the heritable materials of important, including
locally important, tree and woody plant species.
There is also a need to ensure the survival of the
vast majority of, and preferably all, tree and other
woody plant species likely to have values hitherto
unknown and/or novel products and services which
may be required by future generations. Especially
critical are those tree species in monotypic families
or genera which are genetically more distinctive
and irreplaceable.
This report addresses the conservation,
management and sustainable use of forest tree
and other woody species’ genetic resources of
actual and potential value for human well-being
in the broad range of management systems (see
Table 1.1). The deinition of forest used in FAO’s
Global Forest Resources Assessment 2010 (FRA
2010) is “land spanning more than 0.5 hectares
with trees higher than 5 metres and a canopy
cover of more than 10 percent or trees able to
reach these thresholds in situ”. Palms and bamboo
forests are included if these criteria are met, as
tree-like monocotyledons such as palms, climbing
rattans and bamboos generally are considered as
FGR and fall under the responsibility of forestry
agencies. This report, relecting the national
reports on which it is based, focuses mainly on
tree and larger woody species present in forests,
3
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
both natural and planted. However, it also deals
with tree and other woody species outside
forests which are arboreal components in more
open situations, including agroforestry systems,
woodlands and home gardens.
Forest cover is commonly used as a proxy to
assess FGR conservation. However, it is only a crude
measure, as the type of forest may range from, at
one extreme, well connected, highly biodiverse,
well managed tree-species-rich communities with
limited immediate threats to their integrity, which
conserve substantial amounts of FGR, to, at the
other extreme, clonal monocultures which usually
have a much more limited FGR conservation role.
Primary and naturally managed forest cover may
be more relevant than total forest cover as crude
surrogates of FGR conserved.
At the country level, a major concern is
that most forest loss is occurring in biodiverse
tropical countries (see Part 2 on drivers of
change). Extensive losses in biodiverse forest
cover are likely to be accompanied by loss of
genetic diversity in socio-economically useful and
potentially useful tree species. Gross losses of
undocumented and poorly documented FGR in
many tropical countries are of extreme concern,
and this dire situation requires urgent action
involving inventory and conservation measures
which must be adequately funded and prioritized
at both national and international levels.
Primary forests, which include some of the
most FGR-rich forests, account for 36 percent of
the global forested area, but have decreased by
more than 40 million hectares since 2000 (0.37
percent per annum) (FAO, 2010a) – in most cases
with a permanent loss of associated forest genetic
resources.
Around the globe the area of planted forest
is increasing and now accounts for 7 percent of
total forest area, with the highest proportion
in Asia (almost 20 percent) (FAO, 2010a). These
igures underscore the need to consider carefully
the genetic materials used to establish planted
forests or to assist regeneration. There is a need
to ensure that such forests utilize appropriate,
diverse, adapted (including for predicted new
4
climatic conditions) and useful genetic materials
and that information on their genetic makeup
is well documented. It is noted that for planted
forests that have production of wood or biomass
as their prime objective, production may be
maximized with minimal diversity (e.g. a single
clone in clonal eucalyptus plantations) and in
these cases it is necessary to weigh risk against
productivity. There is also a need for safe
movement of germplasm to ensure that pests
and diseases are not inadvertently introduced,
especially as forest tree species may become
more vulnerable to pests and diseases as climate
changes.
Deinitions
Forest genetic resources (FGR) refers to the
heritable materials maintained within and among
tree and other woody plant species that are of
actual or potential economic, environmental,
scientiic or societal value. Some country reports
included woody shrub species which may be
regarded marginally as FGR because they are
often of low stature when grown in dificult and
arid environments. Fruit- and nut-trees and their
wild ancestors were in general included in the
reporting as they are frequently multipurpose,
providing timber, medicine and services and often
being handled by forestry agencies. The term
FGR is also sometimes used incorrectly to cover
more generally the tree and forest resources and
products themselves.
Forest biodiversity has a broader connotation
than FGR and denotes the variability among
forest-dwelling organisms and the ecological
processes of which they are a part. It includes
variation at forest ecosystem, species and
molecular levels.
FGR comprise one subset of plant genetic
resources for food and agriculture (PGRFA).
PGRFA are deined as any genetic material of
plant origin of actual or potential value for
food and agriculture (which in the UN system is
taken broadly to include forestry). FGR are also
included as a subset of agrobiodiversity, which is
deined as the variety and variability of animals,
OVERV IE W
TABLE 1.1
Main types of forest and tree resources management
Naturally regenerated forests
Planted forests
Seminatural
Primary
Forests of native
species, where
there are no
clearly visible
indications of
human activities
and the ecological
processes are not
directly disturbed
by humans
Modified
natural
Forests of
naturally
regenerated
native species
where there are
clearly visible
indications of
signiicant human
activities
Assisted
natural
regeneration
Silvicultural
practices in
natural forest
by intensive
management:
• weeding
• fertilizing
• thinning
• selective
logging
Plantations
Planted
component
Forests of
native species,
established
through planting
or seeding,
intensively
managed
Productive
Protective
Forests of
introduced and/
or native species
established
through planting
or seeding mainly
for production
of wood or nonwood goods
Forests of
introduced and/
or native species,
established
through planting
or seeding mainly
for provision of
services
Trees outside
forests and
agroforestry
systems
Stands smaller
than 0.5 ha;
tree cover in
agricultural land
(agroforestry
systems, home
gardens,
orchards);
trees in urban
environments; and
scattered along
roads and in
landscapes
Source: Modiied from FAO, 2006.
plants and micro-organisms that are used
directly or indirectly for food and agriculture,
including crops, livestock, forestry and isheries.
Agrobiodiversity includes the diversity of genetic
resources (varieties, breeds) and species used for
food, fodder, ibre, fuel and pharmaceuticals.
It also includes the diversity of non-harvested
species that support production (soil microorganisms, predators, pollinators) and those in the
wider environment that support agro-ecosystems
(agricultural, pastoral, forest and aquatic) as well
as the diversity of the agro-ecosystems (FAO,
1999a). Traditional knowledge of biodiversity or
ethnobiodiversity is increasingly understood to
be an integral component of agrobiodiversity
(Thaman, 2008), and its loss may threaten diversity
at different levels – in ecosystems and within and
among species.
Intraspeciic diversity, or the genetic variation
within species, may be considered from several
perspectives, ranging from formally recognized
taxonomic categories of subspecies and varieties
through to genetic differences between and
within populations. Subspecies are usually
morphologically
or
otherwise
distinctive
entities within a species which have evolved in
geographic and reproductive isolation. If they
continue to be separated for many generations,
subspecies may become distinctive enough from
each other, or develop reproductive barriers,
to become separate species. Ecotypes are an
intraspeciic group having distinctive characters
which result from the selective pressures of the
local environment. Genotypes can be considered
as the sum of the total genetic information in
an individual or the genetic constitution of an
individual with respect to genetic loci under
consideration. Individual long-lived trees of
different species may develop into chimeras of
many genotypes owing to the accumulation
of spontaneous mutations of neutral selective
itness in nuclear genes in bud meristems, but this
topic has been little researched.
An organism’s genome represents its total
genetic material, and in plants comprises three
separate genomes: nuclear (about 50 000–
100 000 genes), chloroplast (about 100–120
genes) and mitochondrial (about 40–50 genes)
(Murray, Young and Boyle, 2000). Understanding
of genetics and the nature of heritable materials
5
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
in trees is rapidly evolving, informed by genomic
studies in economically important forest trees such
as Eucalyptus and Populus (both angiosperms),
woody fruit-trees such as Malus domestica
(apple) and Citrus sinensis (sweet orange), the
ancestral lowering plant Amborella trichopoda
and coniferous families through transcriptome
(ribonucleic acid [RNA]) sequencing (e.g. Lorenz
et al., 2012), as well as genetic research on other
plants and other organisms. Advances in gene
sequencing technologies have made possible the
sequencing of conifer giga-genomes; several such
studies have been completed, e.g. for Picea abies
(Nysted et al., 2013) and Pinus taeda (Neves et al.,
2014), while others are in progress or planned
(see e.g. Mackay et al., 2012).
Genes are nuclear deoxyribonucleic acid (DNA)
sequences to which speciic functions can be
assigned, while alleles are alternative forms
of a gene located on the corresponding loci of
homologous chromosomes. In plants, as well as
other higher organisms, a variable proportion of
the nuclear genome is composed of non-protein
coding, repeat DNA sequences, which have
several origins. Some of these sequences have
speciic regulatory functions and/or may donate
segments of DNA which can become incorporated
into genes. Angiosperms possess genomes with
considerable gene redundancy, much of which
is the result of ancient polyploidization events
(Soltis et al., 2009).
DNA present in cellular organelles, notably
chloroplasts and mitochondria, is a vital
component of a tree’s heritable materials. While
nuclear DNA is always inherited biparentally (i.e.
from both male and female parents), organellar
DNA may have different modes of inheritance.
Chloroplast DNA is usually maternally inherited
in angiosperms (e.g. in poplars [Rajora and
Dancik, 1992] and in eucalypts [Byrne, Moran and
Tibbits, 1993]) but may also be inherited from
both parents (Birky, 1995) or rarely from the male
(Chat, Chalak and Petit, 1999). In gymnosperms,
chloroplast DNA is mainly inherited paternally
or infrequently from both parents (e.g. Neale,
6
Marshall and Sederoff, 1989; Neale and
Sederoff, 1989; White, 1990; Wagner, 1992). The
mitochondrial genome is most often maternally
inherited in angiosperms (e.g. Reboud and
Zeyl, 1994; Vaillancourt, Petty and McKinnon,
2004) but may be maternally, paternally or
biparentally inherited in gymnosperms (e.g.
Neale, Marshall and Sederoff, 1989; Neale and
Sederoff, 1989; Wagner, 1992; Birky, 1995).
Chloroplast DNA is strongly conserved and
therefore useful for evolutionary studies (e.g.
in Eucalyptus spp. [Freeman et al., 2001] and in
Juglans spp. [Bai, Liao and Zhang, 2010]), while
mitochondrial DNA is commonly used as a source
of genetic markers in studies of gene low and
phylogeography. Heritable changes in gene
expression or cellular phenotype may be caused
by several mechanisms which do not involve any
change in the underlying DNA sequence; these
are the realm of the poorly understood science
of epigenetics.
A population of a particular tree species
comprises all the individuals of that species in the
same geographical area and genetically isolated
from other populations of the same species.
In sexually reproducing species the population
comprises a continuous group of interbreeding
individuals. A metapopulation of a forest tree
species comprises a set of spatially separated
local populations or subpopulations, coexisting in
time, which interact infrequently via pollen and
seed dispersal among them.
The term provenance is particularly important
in relation to forest tree germplasm and
refers to the geographic origin of a particular
germplasm source, although it is sometimes
used synonymously and interchangeably with
“population”. The ield performance of seed
sourced from a particular representatively
sampled provenance, if from a rather narrow
geographic area (including same soil type and
little altitudinal variation), will generally be more
consistent than that of a population, which may
vary considerably owing to clinal variation arising
from gradients in selective pressures.
OVERV IE W
In situ conservation refers to the conservation
of ecosystems and natural habitats and the
maintenance and recovery of viable populations
of species in their natural surroundings. In the
case of domesticated or cultivated species it
refers to their conservation in the surroundings
in which they have developed their distinctive
properties (UN, 1992). Circa situm conservation is
a type of conservation that emphasizes the role
of regenerating saplings in linking vegetation
remnants in heavily modiied or fragmented
landscapes such as those of traditional
agroforestry and farming systems (Barrance,
1999). The related term matrix management has
been coined to refer to approaches for conserving
and managing biodiversity in forests outside
protected areas (Lindenmayer and Franklin, 2002);
dynamic conservation of FGR will mainly occur
in the matrix and will involve management of
trees on farms, in forest fragments and especially
in sustainably managed production forests. Ex
situ conservation refers to the conservation of
components of biodiversity outside their natural
habitats, including FGR in planted forests, tree
breeding programmes, ex situ gene conservation
stands or ield gene banks, seed and pollen banks,
in vitro storage and DNA storage (FAO, FLD and
IPGRI, 2004).
Evolutionary or dynamic conservation of FGR
essentially involves a natural system in which the
evolutionary forces and natural selective processes
which gave rise to diversity are allowed to
operate and over time modify allelic frequencies.
The past few decades represent the beginning
of an era of unprecedented change in selective
pressures on almost all trees species. These altered
selective forces include more extreme climatic
events, gradual increases in temperature and
altered rainfall regimes, changed ire regimes,
increased air pollution and elevated atmospheric
CO2 levels, habitat fragmentation, increases in
pest and disease outbreaks and the appearance
of new pest and disease species, competition
with invasive exotic plant species including
transformer species capable of changing the
ecology of entire ecosystems, and the loss of
or changes in pollinators and dispersal agents.
Dynamic in situ conservation allows species
adaptation through continuous “selection of the
ittest”, co-adaptation of host-pathogen systems
and other complex biological interactions (Kjær,
Graudal and Nathan, 2001; Byrne, 2000).
In situ conservation of the FGR associated with
identiied superior provenances of economically
important tree species is vital even when they
may be relatively well conserved ex situ, e.g.
through planting and breeding programmes. This
is because tree breeders may need to resample in
original populations for infusion in later breeding
populations and/or to identify new desired traits
in already well known and adapted populations.
Selective forces may differ in the native and
exotic/planted environments, and this difference
is the basis of the often remarkably swift
evolution of landraces (local varieties of a tree
species undergoing domestication which have
developed through selection and adaptation to
the new environment in which they are growing)
which are much better adapted than the original
introduction after just one or two generations of
selection. Increasingly rapid climate change and
associated extreme climatic events are altering
the selective forces in both the native and
exotic/planted environments and creating new
challenges for FGR conservation.
Static conservation of FGR involves conserving
individual genotypes – for example in the ield
as clonal archives and in vitro in tissue culture
and cryopreserved embryo culture – and groups
of genotypes in long-term seed storage for tree
species with orthodox seed storage behaviour
(Kjær et al., 2004). This approach has generally
been viewed as a complementary approach to
dynamic in situ conservation and more often as a
short-term conservation strategy and a means of
safety duplication in the case of cryopreservation.
Given the unprecedented scale of threats to FGR
and the likely losses of diversity and changes in
selective forces which will drive rapid changes
in the genetic makeup of natural (and artiicial)
7
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
populations of tree species, it might be timely
to reconsider the potential value and cost
effectiveness of static conservation activities.
Characteristics of forest genetic
resources
Trees and other woody species differ from
other organisms in several key respects. Forest
tree species are generally long-lived and have
developed natural mechanisms to maintain
high levels of genetic variation within species.
They include high rates of outcrossing and often
long-distance dispersal of pollen and seed. These
mechanisms, combined with native environments
that are often variable, have enabled forest tree
species to evolve into some of the most genetically
diverse organisms in existence. Forest community
ecosystem processes, including evolution of
species, have been found to be closely related to
the genetic diversity in structurally dominant and
keystone tree species (e.g. Whitham et al., 2006
and references therein). This section describes
some differences between trees and other
organisms.
Chromosomes and DNA
Large and seemingly inexplicable differences
and variations in chromosome number, ploidy
level and genome size occur both among trees,
and between trees and other organisms. The two
major groups of trees – gymnosperms (including
conifers) and angiosperms – appear to have been
separated by more than 290 million years of
independent evolution. Angiosperms have high
levels of genetic diversity – both a high number
of genes, e.g. more than 40 000 for poplar (JGI
and CIG, 2006–2014), and high allelic variation.
Conifers typically have very large genomes, or
giga-genomes, with an order of magnitude more
DNA than other organisms, but with numerous
highly repetitive, non-coding sequences (Ahuja
and Neale, 2005; Rigault et al., 2011; Mackay et
al., 2012). DNA sequencing studies of selected
model plant species in these two groups are
providing different perspectives and insights into
8
plant genome biology and evolution. While large
sets of DNA sequences overlap between conifers
and angiosperms (e.g. Pozo et al., 2011), about
30 percent of conifer genes have little or no
sequence similarity to angiosperm plant genes of
known function (Pavy et al., 2007; Parchman et
al., 2010).
Whole-genome duplication
(polyploidization)
While
polyploidization
or
whole-genome
duplication is rare in animals and conifers,1 it is
now considered ubiquitous in angiosperms and
has occurred frequently through their evolution.
Polyploidization is a mechanism of sympatric
speciation because polyploids are usually unable
to interbreed and produce fertile offspring
with their diploid ancestors. Polyploidization
may involve autopolyploidy (spontaneous
multiplication involving the chromosomes of a
single species) or allopolyploidy (involving more
than one genome or species).
Whole-genome duplication is considered
likely to have led to a dramatic increase in
species richness in several angiosperm lineages
including families with important FGR such
as the legumes (Fabaceae) and to be a major
diversifying force in angiosperms (Soltis et al.,
2009). In animals, aneuploidy is usually lethal and
so is rarely encountered, whereas in angiosperms
the addition or elimination of a small number
of individual chromosomes appears to be better
tolerated. New research has indicated that
aneuploidization may be a leading cause of
genome duplication; Considine et al. (2012) have
found that autotriploidization is important for
speciation in apples (Malus spp.) and that such
polyploidization confers both genetic stability and
diversity and presents heterozygosity, heterosis
and adaptability for evolutionary selection.
The monotypic small tree Strasburgeria robusta
from New Caledonia (France) has an extremely
1
Polyploids in conifers include Sequioa sempervirens (hexaploid)
and Fiztroya cupressoides (tetraploid) (see Ahuja, 2005 for a
review of polyploidy in gymnosperms).
OVERV IE W
Box 1.1
Examples of some of the oldest known trees and woody shrubs
group more than 2 500 years old, as determined
by tree ring samples (Earle, 2013).
• A specimen of Picea abies in Dalarna Province
(Sweden) has been found to be at least 9 550
years old. It has survived by resprouting from
layered stems, rather than underground root
suckering (Umeå University, 2008).
• Three species of bristlecone pines in the United
States of America may live for several thousand
years, with one specimen of Pinus longaeva in
California determined to be about 4 900 years
old (Currey, 1965).
• One clone of Populus tremuloides in central Utah
(United States of America) is estimated to be
80 000 years old (DeWoody et al., 2008); 5 000to 10 000-year-old clones are reputedly common.
• A sterile triploid clone of the woody angiosperm
shrub king Lomatia tasmanica has been
determined to be at least 43 600 years old
(Lynch et al., 1998).
• A colony of Lagarostrobus franklinii trees
covering 1 ha on Mount Read, Tasmania
(Australia) is estimated to be around 10 000
years old, with individual tree stems in this
high ploidy level (20n with n = 25) which may
have enabled it to adapt to an extreme edaphic
environment, i.e. ultramaic soil (Oginuma,
Munzinger and Tobe, 2006).
Longevity
Trees and other woody species are perennial,
often long-lived, organisms. For long-term
survival at a particular site, they need to be able
to endure environmental extremes and changes
and/or to persist in the soil seed bank or regrow
from root suckers and coppice. The high genetic
diversity that characterizes tree populations and
individuals, and associated stress tolerance and
disease resistance mechanisms, help explain their
capacity to persist and thrive for long periods.
Indeed the only organisms with a life span
comparable to that of the oldest trees are corals,
fungal mats and other clonal suckering plants
such as Larrea tridentata (creosote bush).
The life span of trees typically ranges from about
10 to 15 years (for short-lived pioneer species)
to 200 to 300 years (for many larger species and
those found in arid zones). Root suckering clones
provide the oldest known woody species; some
examples are given in Box 1.1.
Almost all of the world’s oldest recorded trees
are conifers (Rocky Mountain Tree-Ring Research,
2013). It is likely that their xylem structure,
which differs from that of angiosperms, and the
associated ability to survive lower conductivities
and drought (Choat et al., 2012) contributes to
their great longevity. Ancient trees occur in all
three orders of conifers: Pinales, Araucariales,
and Cupressales (see Table 1.2).
Species longevity. The capacity of gymnosperms
to persist over millions of years, often almost
unchanged in form, is evidenced by Ginkgo biloba,
recently rediscovered in the wild in southwestern
China where glaciation was relatively weak (Tang
et al., 2012); it was previously only known from
fossils and from cultivation in Japanese and
Chinese temple gardens. Likewise, Wollemia
nobilis, discovered in 1994 in a valley near
Sydney, eastern Australia, is presumed to be the
last remnant of a genus that evolved about 61
million years ago (Liu et al., 2009).2 It includes
less than 100 stems of this tree, all appearing to
be genetically identical, and likely comprising a
single clonal root suckering clump.
2
Wollemia nobilis may die out in the wild because of the recent
introduction of the virulent root rot pathogen Phytophthora
cinnamomi, but it has been well conserved globally ex situ
through a successful campaign to promote its use as an
ornamental.
9
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
TABLE 1.2
Life span of some of the longest-lived conifers
> 1 000 years
> 2 000 years
>3 000 years
> 4 000 years
Xanthocyparis nootkatensis
Juniperus occidentalis
Fitzroya cupressoides
Pinus longaeva
Cryptomeria japonica
Lagarostrobus franklinii
Sequoiadendron giganteum
Picea abies
Juniperus scopulorum
Pinus aristata
Larix lyalli
Pinus balfouriana
Pinus albicaulis
Sequoia sempervirens
Pinus edulis
Pinus lexilis
Pseudotsuga menziesii
Taxodium distichum
Source: Wikipedia, n.d.
Size
Trees include the biggest and tallest organisms
on the planet. Sequoia sempervirens has been
recorded up to 115 m tall and weighing up to
530 tonnes (Preston, 2007). The world’s tallest
and biggest angiosperms are eucalypts in
southeastern Australia, speciically Eucalyptus
regnans from Victoria and Tasmania, with trees
measuring over 100 m; historically, examples
were known with height of at least 114 m and
trunk volumes to 360 m3 (Carder, 1995).
Trees are the dominant structural element in
forests and several other terrestrial ecosystems
(agroforests, woodlands and gardens), intercepting much of the radiant sunlight, dominating
photosynthetic processes and carbon lows and
comprising a large proportion of the biomass.
Populations of large old trees are rapidly
declining in many parts of the world, with
detrimental implications for ecosystem integrity
and biodiversity (Lindenmayer, Laurance and
Franklin, 2012). Throughout the tropics the
biggest forest trees are disappearing, partly as a
result of selective targeting by loggers, but more
recently as a result of forest fragmentation,
climate change and exposure to drought.
Diverse breeding systems
Trees are notable for their diverse breeding and
reproductive systems, which are in turn major
determinants of spatial patterns of tree species
genetic diversity. Most tree species reproduce
10
sexually, although many have a combination of
sexual and asexual reproductive means, while a
few have lost the ability to reproduce sexually
and are maintained as sterile, root-suckering
clones in certain parts of their range, e.g. Acacia
anomala in southwestern Australia (Coates,
1988), Casuarina obesa in western Victoria
(Australia), and Santalum insularis on Mangaia
(Cook Islands). It is possible that a long-distance
pollen or seed dispersal event could cause such
plants to regain a sexual mode of reproduction.
Tree species reproducing by sexual means
have diverse reproductive biologies including
monoecious (with separate male and female
lowers on the same tree), dioecious (with
individual trees bearing either male or female
lowers), hermaphroditic (with functional
bisexual lowers) and polygamous (with male,
female and bisexual lowers on the same tree).
Almost all lower sex combinations are possible;
trees may have male and bisexual lowers; female
and bisexual lowers; and bisexual lowers with
a small number of male and female lowers.
At the global level, populations of lowering
plant species are mainly hermaphroditic (72
percent), with smaller proportions as follows:
monoecious, 4 percent; dioecious, 7 percent;
gynodioecious (having female lowers on some
plants and bisexual lowers on other plants) or
androdioecious (having male lowers on some
plants and bisexual lowers on other plants),
7 percent; and trioecious (with male, female
OVERV IE W
and bisexual lowers in three different plants),
10 percent (Yampolsky and Yampolsky, 1922;
Dellaporta and Calderon-Urrea, 1993). However,
these proportions vary regionally, and they also
vary between trees and other lowering plants,
For example, dioecy, an obligate outcrossing
pollination arrangement, was found to be higher
(>20 percent) in tree species (Bawa, Perry and
Beach, 1985).
Most angiosperm species with hermaphroditic
lowers have preferential outcrossing systems
such that fertilized, viable seeds are generally
derived from outcrossing. Reported outcrossing
rates in tropical angiosperm tree species from
different families, and including those occurring
at low density, were typically in the range of 60
to 100 percent, but with considerable variation
(Byrne, 2008; Butcher, Glaubitz and Moran, 1999;
Gandara, 1996; Hamrick and Murawski, 1990;
Kitamura et al., 1994; Lepsch-Cunha, Gascon
and Kageyama, 2001; Lepsch-Cunha et al., 2001;
Mandal, Ennos and Fagg, 1994; Moran, Muona
and Bell, 1989; Muluvi et al., 2004; Murawski
and Hamrick, 1991; Murawski and Bawa, 1994;
Murawski, Dayanandan and Bawa, 1994;
Murawski, Gunatilleke and Bawa, 1994; Murawski,
1995; Nason and Hamrick, 1997; O’Malley and
Bawa, 1987; Olng’otie, 1991; Sebbenn et al. 2000;
Stacy et al., 1996; Ward et al., 2005).
Outcrossing rates vary within species and
populations and among different lowering
events. For example, tropical acacias from
humid zones in Papua New Guinea and northern
Australia typically have rates of 93 to 100 percent
outcrossing, but lower rates (30 to 80 percent)
have been found in more southerly populations
of Acacia mangium (Moran, Muona and Bell,
1989; Butcher, Glaubitz and Moran, 1999), while
polyploid dry-zone African acacias had low
outcrossing rates of between 35 and 38 percent
(Mandal, Ennos and Fagg, 1994; Olng’otie,
1991). Plasticity in mating systems has also been
observed in response to changes in pollinators.
Ceiba pentanda (kapok), for instance, had a
predominantly self-incompatible system in
situations of high bat pollinator visitation, but
changed to a mixed mating system with high
levels of self-pollination in situations with low
pollinator visitation rates (Lobo, Quesada and
Stoner, 2005).
Conifers are wind pollinated and either
monoecious or dioecious (with obligate
outcrossing). Species in the families Pinaceae
and Cupressaceae are monoecious (with the
exception of Juniperus spp., which are usually
dioecious); Araucariaceae, Podocarpaceae and
Taxaceae may be monoecious or dioecious;
and Cephalotaxaceae are mainly dioecious
but occasionally monoecious (Earle, 2013).
Mating systems in conifers vary in space and
time, mainly owing to variation in self-pollen
availability (Mitton, 1992). Mechanisms to
promote outcrossing have been identiied in
monoecious conifers, e.g. Pinus taeda (Williams,
Zhou and Hall, 2001). The outcrossing rate for
most conifer species is above 80 percent (e.g.
for 52 species reviewed in O’Connell, 2003).
Through their long evolution, plasticity in the
reproductive systems of conifers may have
helped them to survive. For example, Cupressus
dupreziana has evolved a unique reproductive
system of male apomixis whereby the seeds
develop entirely from the genetic content of the
pollen (Pichot, Fady and Hochu, 2000). Sequoia
sempervirens reproduces by both asexual (basal
suckering) and sexual means but with low seed
set (1 to 10 percent) owing to irregular meiosis
and associated with its hexaploid condition; dual
reproductive systems have enabled redwoods to
maintain heterozygosity and adaptability for
survival (Ahuja, 2005).
Species diversity
The future well-being of the human race,
and the health and productivity of various
ecosystems and communities, will often rely on
genetic diversity both within and among tree
species. While this report is mainly concerned
with intraspeciic diversity, it is also appropriate
to consider the economic and other uses of
trees and other woody species provided through
diversity at species level.
11
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
Interspeciic (between species) diversity
Approximately 80 000 to 100 000 tree species
have been described and currently accepted as
valid and unique (Oldield, Lusty and MacKinven,
1998; Turok and Geburek, 2000); together with
larger woody shrubs they likely represent about
50 percent of all vascular plant species.
Considerable research has been undertaken
to understand how tropical forests develop and
maintain their typically vast tree species diversity,
but answers remain elusive (e.g. Denslow, 1987;
Cannon, Peart and Leighton, 1998; Ricklefs
and Renner, 2012). Tree diversity in complex
ecosystems, moist tropical forest gaps and
regeneration niches may have been generated in
part, and may be maintained, by host-pathogen
and host-parasite interactions (Grubb, 1977; Wills
et al., 1997).
Research, development, conservation and
use of tree species, in particular tropical species,
has often been frustrated by insuficient
and inadequate taxonomic knowledge, e.g.
assessment of conservation status of different
species (Newton and Oldield, 2008). An array of
more powerful and eficient genetic technologies
are increasingly available to complement
traditional, morphology-based taxonomy and
ield studies, and these are leading to better
circumscription of tree species and understanding
of their phylogenetic relationships. The nature of
variation in trees is such that species boundaries
will not always be easily deined, as in the
following examples (Whitmore, 1976):
• species existing as morphologically
distinctive and geographically disjunct
populations which rarely exchange genetic
materials and are best considered as
provenances, varieties or subspecies;
• species that are readily discernible in most
of their natural range and have evidently
been reproductively isolated for much of
their recent evolution, but which form
fertile hybrid swarms in small overlapping
contact zones;
• species with polyploid races, often coupled
with apomictic reproduction;
12
• ochlospecies, i.e. species whose complex
variation patterns cannot be satisfactorily
accounted for by conventional taxonomic
categories.
Based on a literature review carried out for
this report, it is conservatively estimated that
more than 34 000 tree species in more than 1 000
genera are of socio-economic, environmental
and scientiic importance and used on a regular
(daily or weekly) basis by people throughout the
world. This number includes large woody shrubs
attaining more than 2 to 3 m in height, given that
the boundary between trees and woody shrubs is
unclear and individual species may exist as either
trees or shrubs depending on environmental
factors. Also included are fruit- and nut-trees
and their wild relatives. The total comprises both
angiosperms (33 500 species in 976 genera and
131 families, including bamboos and palms) and
gymnosperms (530 species in 67 genera and nine
plant families, excluding cycads).
In total nearly 8 000 species and subspecies
were mentioned in country reports, and about
2 360 species and subspecies were mentioned as
being actively managed (i.e. managed speciically
for their products and/or services) in various
systems (Figure 1.1).
In practice, this vast diversity at species level in
trees means that, for a given product or service,
local people and foresters may have a choice
among hundreds of species which are locally
available and/or suitable options in different
ecological conditions. As well as providing
opportunities, this vast species genetic resource
can also present challenges in ascertaining which
species to prioritize for research and development
and for planting.
Area measures of forest tree species richness
are used to deine biodiversity hotspots and
megadiverse countries and have been used to
prioritize conservation efforts both internationally
and within countries. However, the identiication
and protection of areas particularly rich in FGR
should not detract from other efforts to conserve
and manage FGR throughout the world. Some
forests may contain few species in comparison with
OVERV IE W
FIGURE 1.1
Number of species and subspecies mentioned as actively managed in country reports, by region
Africa
Asia
Europe
Latin America and the Caribbean
Near East
North America
Oceania
Total
0
1 000
2 000
3 000
Total species and subspecies reported
other, more loristically diverse areas, but these
few species may be vital for local communities (e.g.
atoll island communities in Oceania) or may be
genetically unique and have extremely high value
for conservation of FGR (as in Seychelles, which has
only 93 highly distinct indigenous tree species with
endemism of more than 50 percent).
Many country reports indicated that conservation of FGR, including the vast diversity at
the tree species level, is seriously hampered by a
lack of taxonomic skills, inventory and knowledge
of species distributions. Accordingly there is an
urgent and ongoing need to strengthen national
FGR assessment, characterization and monitoring
systems.
Intraspeciic (within species) diversity
Intraspeciic diversity or genetic diversity within
tree species is manifested in different ways and
can be characterized at different levels. It can be
assessed at the molecular level through nuclear
4 000
5 000
6 000
7 000
8 000
Species and subspecies actively managed
DNA (e.g. using neutral markers such as random
ampliied polymorphism DNA [RAPD], chloroplast
DNA (especially useful for providing evolutionary
information), direct RNA sequencing (providing
information on gene regulation and proteins)
and enzyme variation (gene products assessed
through isozyme electrophoresis). Genetic
variation is also observed at expressed levels,
for example through quantitative variation in
growth and other traits as assessed through ield
trials and through morphological, physiological,
entomological and pathology studies. Sometimes
variation is discontinuous, giving rise to the
identiication of varieties including chemotypes,
morphotypes and the like. Intraspeciic patterns
of genetic variation in tree species have been
found to vary as a result of factors such as the
evolutionary history of the species; distribution
of populations and connectivity; reproductive
biology and mating system; dispersal of pollen
and seed; introgression and hybridization with
13
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
related species; chance factors; and genetic drift.
Observed patterns of genetic variation can vary
between different genomes of the same tree
species, if these genomes are inherited differently,
with associated differences in dispersal of pollen
and seed, as described by Tomaru et al. (1998) in
Fagus crenata in Japan.
Humans have long been interested in using
and inluencing diversity within tree species,
especially tree species producing edible fruits
and nuts; an example is the domestication and
selection of Juglans regia (walnut) described
in the Azerbaijan country report. Another well
documented case is the selection, translocation
and domestication of tropical nut-trees in
arboricultural systems in Papua New Guinea and
Solomon Islands, dating back more than 3 000
years (Yen, 1974; Lepofsky, 1992; Lepofsky, Kirch
and Lertzman, 1998). However, for the most
part traditional knowledge and activities related
to improvement of FGR are undocumented.
National-level assessments should be made a
priority, especially in tropical countries, before
the information dies out with those who hold it.
The forestry profession has had a long interest
in studying and using variation in trees, including
geographic variation in economically important
planted forest tree species, which was investigated
through ield trials early in the last century.
The International Union of Forest Research
Organizations (IUFRO) coordinated provenance
trials of Pinus sylvestris established in 1907, 1938
and 1939 in Europe and the United States of
America (Wright and Baldwin, 1957; Langlet,
1959; Giertych, 1979). After the hiatus of the
Second World War, provenance ield trial research
recommenced in earnest, with Pinus ponderosa
provenance trials established in the United States
of America in 1947 (Callaham, 1962) and new
P. sylvestris provenance trials in Sweden from 1952
to 1954 (Eiche, 1966; Eriksson et al., 1976). During
the 1960s and 1970s, assessments of genetic
diversity in forest tree species gathered pace and
extended to tropical and Southern Hemisphere
species. These assessments were focused mainly
on morphological attributes including wood traits,
14
adaptiveness, quantitative growth characters,
disease tolerance, and genotype × environment
interaction. These attributes were examined
through series of ield trials, often undertaken in
several countries and referred to as provenance
trials. Some of the tree species studied in these
early investigations included:
• Betula alleghaniensis (Clausen, 1975)
• Cordia alliodora (Sebbenn et al., 2007)
• Eucalyptus camaldulensis (Lacaze, 1978)
• Eucalyptus urophylla (Vercoe and Clarke,
1994)
• Fagus sylvatica (Giertych, 1990)
• Gmelina arborea (Lauridsen, Wellendorf
and Keiding, 1987)
• Pinus kesiya (Barnes and Keiding, 1989)
• Pinus patula (Barnes and Mullin, 1984)
• Pinus radiata (Nicholls and Eldridge, 1980)
• Tectona grandis (Lauridsen, Wellendorf and
Keiding, 1989)
• Terminalia superba (Delaunay, 1978)
Based on the success of the earlier provenance
trials, the provenance trial approach has been
continued and extended, including to national
trials with native species. Some examples include:
• Acacia aneura (Ræbild, Graudal and
Nimbkar, 2003)
• Acacia auriculiformis (Awang et al., 1994)
• Acacia senegal (Ræbild, Graudal and
Ouédraogo, 2003)
• Alnus rubra (Xie, 2008)
• Azadirachta indica (Hansen, Lunde and
Jørgensen, 2000)
• Casuarina equisetifolia (Pinyopusarerk et al.,
2004)
• Chukrasia tabularis (Ratanaporncharern,
2002)
• Endospermum medullosum (Vutilolo et al.,
2005)
• Faidherbia albida (IRBET/CTFT, 1985-1988)
• Pachira quinata (Hodge et al., 2002)
• Parkia biglobosa (Ouédraogo et al., 2012)
• Pinus caribaea (Hodge and Dvorak, 2001)
• Pinus tecunumanii (Hodge and Dvorak,
1999)
• Sterculia apetala (Dvorak et al., 1998)
OVERV IE W
Provenance/progeny trials continue to be one of
the irst steps in domestication and improvement
of wild tree species. The range of attributes
assessed is becoming more diverse according
to the particular and sometimes specialized
end uses envisaged, and can include pulping
and ibre properties and timber uniformity, or
characteristics of wood, essential oils, fruit and
nuts for multipurpose species.
Countries reported on species/provenance
trials, and this information is covered in Chapter
11. Many of the trials are in progress, have not
been reported or are not readily available in the
published scientiic literature.
Internationally coordinated provenance trials of
tree species will become increasingly important in
providing data to assess the modelled impacts of
climate change on productivity of planted forests
and to determine which species or provenances
will be best adapted to new, modiied climates
(e.g. Booth et al., 1999; Leibing et al., 2009).
Provenance trial data can also be used to assist
interpretation of the likely impacts of predicted
climate change on native species and populations,
as has been done for Pinus species in tropical Asia
and the Americas (van Zonneveld et al., 2009a,
2009b) and for Eucalyptus species in Australia,
where minor changes in climate will expose at
least 200 Eucalyptus species to completely new
climatic envelopes (Hughes, Cawsey and Westoby,
1996), for which their adaptation potentials are
unknown.
Increasingly, with gathering momentum over
the past 20 years, detailed genetic information is
being obtained for tree species often selected for
study on the basis of their economic importance
or conservation status or for use as representative
model species. Many of the early molecular
studies of diversity in tree species, in the 1980s
and 1990s, focused on high-priority timber trees
and were mainly undertaken in forest genetic
laboratories in developed countries using
electrophoresis techniques. Detailed genetic
evaluations using DNA and enzyme markers
have now been undertaken for many important
forest tree species in Europe and North America;
for example, Canada reported intraspeciic
genetic studies for 32 Canadian tree species.
Until recently, there have been few detailed
studies of intraspeciic genetic variation in
developing countries. Such studies are needed
for the formulation of scientiically based gene
conservation programmes.
Increasingly, as evidenced in the scientiic
literature and reported in the country reports, the
patterns of genetic diversity for a much greater
range of tree species from throughout the world
are being determined using a wide range of
genetic markers; for example, more than 100 tree
species in China have been characterized over the
past decade.
The planning of speciic and effective
programmes for both conserving and exploiting
the genetic diversity in target forest tree species
requires detailed knowledge of the species’ patterns
of intraspeciic diversity, notably knowledge of
how genetic diversity is distributed between
and among populations (genetic snapshot). This
must be complemented by understanding of the
species’ ecology, especially regeneration ecology,
reproductive biology and relationships with other
species (e.g. pollinators, dispersers, symbionts,
predators, parasites and competitors) – in short,
the selective and evolutionary forces that resulted
in its genetic makeup.
Data from genetic studies are increasingly
being used to inform conservation of FGR in
particular tree species (see also Box 1.2). For
example, Canada’s country report refers to this
application for Xanthocyparis nootkatensis and
Thuja plicata. Other examples in the published
literature include the following.
• Genetic data for four evolutionarily
signiicant populations of Calliandra
calothyrsus in Mexico and Central
America indicated the need to conserve
representative populations (Chamberlain,
1998).
• Data for Caesalpinia echinata in Brazil
indicated the need to conserve different
populations in different geographic areas
(Cardoso et al., 1998).
15
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
• Data for Sclerocarya birrea in Kenya
indicated the need to conserve speciic
populations with high genetic diversity
(Cardoso et al., 1998).
• Genetic studies together with quantitative
variation data from provenance trials have
been used to inform conservation plans for
Pinus maximinoi (Dvorak et al., 2002).
• Genetic and quantitative morphological
data have been used to inform conservation
plans for Paramichelia baillonii in China (Li
et al., 2008).
Various genetic studies are demonstrating the
importance of glacial refugia for conserving tree
species and their diversity, e.g. for broad-leaved
trees such as Quercus spp. in Europe (Iberian
Peninsula, Appenine Peninsula and the Balkans)
(Potyralska and Siwecki, 2000); for Irvingia spp. in
Central and West Africa (Lowe et al., 2000); and
for Cunninghamia spp. in East Asia (Hwang et al.,
2003). In addition, species growing in marginal
environments or at the extremes of their range
(in terms of climate and soils) may contain unique
diversity and speciic adapatations that warrant
special attention for evaluation and conservation.
The increased information being generated
through DNA studies is also being used to
make generalized recommendations on how to
conserve genetic diversity. A review by Newton
et al. (1999) noted that application of molecular
techniques to diversity studies in a variety of
tree species had highlighted a greater degree
of population differentiation than indicated by
previous isozyme analyses; thus in the absence of
detailed information on the genetic structuring of
a species, it may be prudent to conserve as many
Box 1.2
Conserving distinct and unique tree lineages
It is logical that national conservation efforts will
focus on maintaining the genetic diversity and
evolutionary potential of high-priority tree species at
the national level, and that international efforts will
focus on those priority species whose distributions
overlap national boundaries and have greater socioeconomic importance outside the country of origin as
planted exotics. There is also a case to be made, from
both an international and a scientiic viewpoint, for
conserving those tree species (families and genera)
that are genetically most distinctive, e.g. monotypic
families and genera, and those that represent
the most evolutionarily divergent lineages. These
genetically distinctive lineages and assemblages may
later be found to hold genes or combinations of genes
that could be extremely useful to future generations;
they need be considered in prioritizing genera or
species of scientiic importance.
The gymnosperms (cone-bearing plants) are replete
with ancient, separate evolutionary lineages, of which
many are vital FGR. Some examples follow.
16
• The order Ginkophytes comprises one family,
Ginkgoaceae, of which there is only one living
tree species, Ginkgo biloba, a living fossil
apparently almost unchanged in form for nearly
175 million years and an important source of
herbal medicine.
• Cunninghamia lanceolata is in its own
subfamily, Cunninghamhioideae.
• Taiwanioideae consists solely of the species
Taiwania cryptomerioides.
• The subfamily Sequoioideae includes three
renowned monotypic tree genera, Metasequoia,
Sequoia and Sequoiadendron, each with one
living species.
Many other coniferous genera comprise a single
tree species which is highly valued for its timber, nonwood forest products (NWFPs) and/or cultural and/or
environmental uses, e.g. Cathaya, Fitzroya, Fokienia,
Lagarostrobos, Manoao, Nothotsuga, Papuacedrus,
Platycladus, Pilgerodendron, Pseudolarix,
Sundacarpus, Taxodium, Tetraclinis and Thujopsis.
OVERV IE W
Box 1.2 cont.
Conserving distinct and unique tree lineages
Several of these monotypic conifer genera are at high
risk of loss of intraspeciic diversity. Some, including
Neocallitropis pancheri in New Caledonia (France),
are endangered according to the International
Union for Conservation of Nature (IUCN). Many of
the endangered and evolutionarily unique lines of
conifer subfamilies, genera and species are endemic
to China, Viet Nam, New Caledonia (France) and other
countries in the Southern Hemisphere, and a strong
conservation effort for these taxa is important in these
countries.
The most primitive angiosperm or flowering plant
is considered to be Amborella trichopoda; this species
has been placed in its own order, Amborellales, and is
of major scientific importance. While its conservation
status has yet to be assessed, it is likely to be at
risk from climate change and fire in previously
unburnt, wet forest ecosystems in New Caledonia
(France). The monotypic Arillastrum gummiferum
from New Caledonia (France) is important for
forest science as an ancestral genus and species
for eucalypts. Many angiosperm genera comprise
a single tree species which may be endangered or
highly valued for its timber, NWFPs, or cultural or
environmental purposes (or a combination of the
above). These include Antiaris, Aralidium, Argania,
Aphloia, Aucomea, Bagassa, Baillonella, Bertholletia,
Bosqueiopsis, Cantleya, Chloroxylon, Crossopteryx,
Cyclocarya, Deckenia, Delavaya, Elingamita,
Eusideroxylon, Faidherbia, Falcataria, Franklinia,
Gomortega, Gymnostemon, Haldinia, Hartogiella,
Itaya, Ixerba, Jablonskia, Jubaea, Kigelia, Kleinhovia,
Koordersiodendron, Kostermansia, Krugiodendron,
Laguncularia, Limonia, Litchi, Maesopsis, Muntungia,
Neobalanocarpus, Noltea, Ochroma, Olneya,
Oroxylum, Platycarya, Pleiogynium, Rhoiptelea,
Spathodea, Ticodendron, Triplochiton, Umbellularia,
Umtiza, Veillonia, Vitellaria, Xanthoceras and Zombia.
Monotypic wild fruit-tree ancestors, such as Clymenia
polyandra from Melanesia, may hold importance for
future citrus breeding. The monotypic Cordeauxia
edulis, an important multipurpose woody shrub in
Ethiopia and Somalia, is classified as vulnerable
on the IUCN Red List of Threatened Species (www.
iucnredlist.org), and the monotypic Canacomyrica
monitcola, endemic to New Caledonia (France), is
classified as endangered.
Several angiosperm orders and whole families of
woody tree species are represented by one or very few
taxa. The family Barbeyaceae comprises the monotypic
Barbeya oleoides, a small tree with medicinal uses
present in northeastern Africa and the Arabian
Peninsula. The Degeneriaceae include two Fijian
timber tree species, ancestral angiosperms in the
genus Degeneria. The Sladeniaceae include three tree
species in two genera: Ficalhoa laurifolia, a timber
tree from montane forests in East Africa, and two
Chinese tree species of the genus Sladenia which have
potential as sources of novel biochemicals, including
for use in insecticides. The order Trochodendrales and
family Trochodendraceae include two East Asian tree
species each in a monotypic genus, Trochodendron
araloides and Tetracentron sinense. These two tree
species are notable in angiosperms for their absence
of vessel elements in the wood, which is thought to
be a secondarily evolved character and of scientific
interest. The 194 palm species in Madagascar, almost
all endemic, and several monotypic palm genera in
Seychelles include unique and endangered genetic
lineages.
Sources: The Plant List, 2013; ILDIS, 2012; Earle, 2013.
17
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
populations as feasible. In many countries, DNA
research on trees is carried out by institutions such
as universities and research agencies that are not
the same as or well linked with the institutions
tasked with developing and implementing
FGR conservation strategies (e.g. forestry and
environment departments, land managers).
18
Accordingly, improved FGR conservation and
management planning and outcomes will require
closer communication between these two groups,
both in identifying priority species for study and
subsequent planning and in implementation and
monitoring of conservation and management
strategies based on research indings.
OVERV IE W
Chapter 2
Value and importance of
forest genetic resources
“ Sustainable forest management of both natural and planted forests and for timber and nontimber products is essential to achieving sustainable development and is a critical means to
eradicate poverty, signiicantly reduce deforestation, halt the loss of forest biodiversity and
land and resource degradation, and improve food security and access to safe drinking water
and affordable energy… The achievement of sustainable forest management, nationally and
globally, including through partnerships among interested governments and stakeholders,
including the private sector, indigenous and local communities and non-governmental
organizations, is an essential goal of sustainable development…”
Paragraph 45, Plan of Implementation of the World Summit on
Sustainable Development (UN, 2002)
In its 2012 Millennium Development Goals
Report, the UN (2012) estimates that, despite
progress in eradication of extreme poverty,
almost 1 billion people will be living on an
income below USD 1.25 per day in 2015;
and that (citing FAO) 850 million people, or
15.5 percent of the world’s population, were
living in hunger in the period 2006–2008.
In this context of poverty and hunger, the
sustainable use of timber and non-wood
forest products (NWFPs) from forests, without
depletion of the supporting FGR, is becoming
increasingly challenging. Limited options for
economic development and an imperative
to focus on immediate needs are dificulties
that promote short-term perspectives in the
use and management of natural resources,
including forests and FGR. The increasing global
population is placing additional pressures on
forests, especially in the tropical developing
regions (see Chapter 5). It is estimated that
about 60 million people (mainly indigenous
and tribal groups) are almost wholly dependent
on forests for their livelihoods. In 2008, for
the irst time in history, more than half of the
world’s population was living in towns and
cities (UNFPA, 2007), but despite the marked
urbanization trend, the demand has not
diminished for wood and ibre for building,
fuel, paper and NWFPs and agroforestry tree
products (AFTPs) such as herbal medicines and
foods.
This chapter reviews the immense value for
humankind, and more generally for life on
Earth, that FGR represent. It is often dificult
to quantify these contributions. While the
country reports note the importance of FGR
to the formal and informal economies and
their social, cultural and environmental value,
limited attempts were made to assign monetary
value to any of the speciic contributions of
FGR. Nevertheless, and although the absolute
and relative value of forests and trees and their
products and services vary tremendously from
country to country, an underlying and unifying
theme from the country reports is that this
value depends on the continued availability,
access and use of FGR.
19
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
Economic value
Economic importance of forests and
forest industries
The speciic economic beneits arising from
conservation and use of FGR are dificult to
isolate from the economic beneits and impacts
of forest and tree resources and the industries
that rely on them. FAO (2013b) estimates
that close to 1.6 billion people – more than 25
percent of the world’s population – rely on forest
resources for their livelihoods. The key economic
values associated with FGR occur both in formal
sectors, e.g. production, trade and employment
(associated mainly with timber, pulp and paper
industries but also with agriculture, horticulture
and pharmacy), and in informal sectors, which
are often poorly documented, such as local uses
of forest foods, fuelwood and herbal medicines.
Most countries reported the economic value of
forest industries in the formal sector (such as
contribution to GDP, exports, employment), but
could not address with precision the value of
forests and trees in the informal sector and their
contribution to rural livelihoods and poverty
alleviation.
The forest products industry alone is a major
source of economic growth and employment,
with global forest products traded internationally
in the order of USD 255 billion in 2011. Some 40
percent of this value is generated in developing
countries, where forest-based employment
provides 49 million jobs (FAO, 2014). The forest
sector is a major provider of rural employment in
many countries; in Africa, for example, it supports
16 to 18 percent of the workforce in Swaziland
and 20 to 30 percent of the workforce in Gabon.
The forest sector contributes substantially to
exports in many countries. At over USD 16 million,
Canada’s forest balance of trade is the largest in
the world, while in Ghana the sector ranks fourth
in contribution to export earnings. The Solomon
Islands report that the export of roundwood
(1.4 million cubic metres in 2008) provided over
70 percent of export earnings and 18 percent of
total government revenue and is the mainstay of
the economy.
20
In all, several billion people worldwide in the
informal economy depend in some form on wood
products and NWFPs from forests and trees (see
Box 2.1). The World Bank (2002) has estimated
that about 1 billion people worldwide depend
on drugs derived from forest plants for their
medicinal needs. In many developing countries
fuelwood is the primary source of energy,
meeting in some cases as much as 90 percent of
energy requirements (Trossero, 2002). Harvesting
of NWFPs from trees and forests in impoverished
rural areas provides income-earning opportunities
for women in particular, contributing to gender
equality. In rural areas of one state in India, women
obtained 2.5 to 3.5 times as much income from
forests and common lands as men (India country
report). NWFPs may also provide signiicant
additional income to small forest owners and
traditional communities in developed countries;
in Canada, for example, maple syrup products
and Christmas trees generate USD 324 million
and USD 36 million worth of sales, respectively,
each year (Natural Resources Canada, 2011).
The conservation of FGR and the development,
distribution and deployment of improved forest
trees for use by rural communities therefore
offers immense potential to improve and increase
security of livelihoods.
Trees have an extremely important role in
supporting agricultural production, particularly in
developing countries, by providing shelter, shade,
soil structure and fertility improvement; reducing
erosion and mitigating loods; and furnishing
materials such as fencing, processing equipment,
and tools. In Ghana, for example, “the use of nontimber forest products in agriculture technologies
is such that in their absence most farming activities
would be impaired” (Ghana country report).
Trees also provide fodder, which may be critical
during the dry season or in times of drought;
for example, in India nearly 39 percent of cattle
depend partly or fully on forests for fodder (India
country report). Biochar, a non-labile carbon soil
additive obtained from biomass, including forest
biomass, can also be used to increase agricultural
productivity (Dawson et al., 2014).
OVERV IE W
Forests and trees also have an important role
in alleviating poverty in times of hardship and
crop failure, for example by providing fuelwood
when other fuels are inaccessible. Tree food crops
are vital in times of drought, when other annual,
rain-dependent crops may fail. The role of trees
as food sources to help alleviate famine is likely
to increase with predicted negative impact of
climate change and associated environmental
stress on agriculture.
Economic contribution of genetic
diversity
The genetic diversity available in tree species
is often of economic utility. In planted forests,
including agroforests, improved, better adapted
and diverse germplasm directly contributes to
improved economic well-being by increasing
the output of superior forest products for lower
inputs (e.g. labour, water and fertilizer), in a
wider range of conditions and environments,
Box 2.1
Valuing non-wood forest products demand
Non-wood forest products are essential to people’s livelihoods and national economies. Recent estimates (FAO, 2010a)
indicate the global value of NWFPs to be less than USD 17 billion annually (Table 2.1). However, their reported value
remains underestimated because of lack of information and relevant assessment tools at the country level.
The country reports on the state of forest genetic resources identified over 1 000 tree species as actively managed
for NWFPs. However, this number is far below the estimates usually found in publications. For example FAO (2011)
indicates that 75 percent of the overall tropical tee species are used for their NWFP value.
TABLE 2.1
Value of removals of plant-based NWFPs (and bee products) by category and region
NWFP category
Total value
(million
USD)
Share of each category in total value
(%)
World
Europe
Asia
Americas
Oceania
Africa
Food
8 614
51
48
67
23
47
93
Other plant products
2 792
17
3
22
61
3
7
Wild honey and beeswax
n.s.
1 805
11
21
n.s.
n.s.
12
Ornamental plants
984
6
10
1
3
4
0
Exudates
631
4
1
7
5
0
25
Plant materials for
medicines, etc.
628
4
5
2
1
9
18
Material for construction,
tools, etc.
427
3
3
1
3
18
n.s.
Fodder
21
n.s.
n.s.
n.s.
n.s.
0
2
Colourants and dies
18
n.s.
n.s.
n.s.
n.s.
0
n.s.
Source: FAO, 2010a.
Note: n.s. = not signiicant.
21
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
with fewer losses to pests and diseases. FGR are
the basis of tree improvement and improved
forest plantation crops, and countries in all
regions reported signiicant gains in productivity
and utility from improvement programmes and/
or widespread use and adoption of improved
materials. Increased yields of superior forest
products generated at lower cost from genetically
improved trees can reduce the harvest pressure on
natural forests and allow them to be harvested in
a less intensive, more sustainable manner, better
enabling them to fulil service roles.
The use of selected better-performing seed
sources (provenances) will often give increases of
10 to 25 percent, and sometimes several hundred
percent, in wood yield above the mean or the
yield of the prior seed source. Given that seed
accounts for a small proportion (e.g. 0.1 to 3
percent of plantation establishment cost, major
economic beneits accrue from using appropriate
germplasm in plantation establishment and
agroforestry (e.g. FAO, 2002). Economically
important, although often threatened, diversity
is contained in wild tree relatives of fruit- and
nut-tree species. For example, germplasm of
a wild and threatened Central Asian apple
species, Malus sieversii, collected in the 1990s
from Kazakhstan, has shown resistance to apple
scab, ire blight, drought and numerous soil
pathogens and is being used by the United States
Department of Agriculture (USDA) Agricultural
Research Service to improve disease resistance
in current apple cultivars in the United States
of America (Forsline et al., 2003; Pons, 2006) for
industry worth USD 2.7 billion in 2011.
Many NWFP species have a wide genetically
determined variation in the yield and quality of
their products, and indeed some industries are
only possible because of this variation. An example
is “unique manuka factor®” (UMF) honey, a
honey with highly antimicrobial properties
produced only by bees feeding on the nectar of
particular populations of Leptospermum spp.,
such as some populations of Leptospermum
scoparium in New Zealand (Stephens, 2006). In
Vanuatu certain individuals and populations of
22
Santalum austrocaledonicum from the islands of
Malekula and Santo produce a sandalwood oil
that meets the international standard for East
Indian Sandalwood oil and accordingly have much
higher value as seed and wood sources (Page et al.,
2010); future replanting will increasingly be based
on these sources (Stephens, 2006). The rich species
diversity of tropical forests directly contributes
to their provision of a wide range of NWFPs; in
Brazil, for example, honey bees have been found
to produce a new type of medicinal red propolis
through collection of resin from the bark of
Dalbergia ecastaphyllum (Silva et al., 2008).
Environmental value, ecosystem
services and resilience
Trees and forests provide a wide variety of
environmental services. As well as holding a
greater proportion of the world’s biodiversity
than any other terrestrial ecosystem, they have
an increasingly recognized role in environmental
protection and rehabilitation. They contribute
to water catchment management, carbon
sequestration and storage, nutrient cycling,
improvement of soil fertility, erosion management and landscape protection, promotion
of agricultural production, animal habitat,
and maintenance of ecological and ecosystem
processes. All trees and woody plants, whether
planted or of natural origin, fulil ecological and
environmental functions and provide a huge
range of environmental services. Nevertheless,
vital environmental services have traditionally
been undervalued owing to lack of markets for
these services.
Biodiversity
Forest ecosystems are repositories of huge
reservoirs of biodiversity. They support a vast
number and wide range of species, most of
which are forest dependent. Nearly 90 percent
of terrestrial biodiversity is found in the world’s
forests.
The most species-diverse ecosystems on Earth
are moist tropical lowland forests, which are
principally located in developing countries. The
OVERV IE W
vast richness of herbivorous insects in these forests
has recently been shown to be driven by the
phylogenetic diversity of their plant assemblages
(Novotny et al., 2006).
Temperate forests and forest tree species
support and provide habitat for myriad other
life forms. For example, two thirds of Canada’s
140 000 species occur in forest ecosystems. In
the United Kingdom, 285 different species of
phytophageous insect have been found on
Quercus robur (Southwood et al., 2004). In
Australia, 306 species of invertebrates have
been found on Eucalyptus obliqua (Bar-Ness,
Kirkpatrick and McQuillan, 2006). In Australia, a
poster produced by the Conservation Commission
of the Northern Territory has referred to the
native Eucalyptus camaldulensis, widely planted
globally, as “nature’s boarding house” in
recognition of the number of mammals and birds
that use it for food, shelter and nesting sites.
In less tree-species rich temperate forests, the
role of trees in promoting biodiversity is likely
to be more associated with and attributable to
their level of intraspeciic genetic variation (e.g.
Whitham et al., 2006).
Ecosystem function
Trees contribute major photosynthetic inputs.
They drive carbon, water and nutrient cycling,
especially absorbing and returning nutrients from
deeper root zones, mobilizing mineral elements
through associations with mycorrhizal fungi and
ixing atmospheric nitrogen through symbiosis
with bacteria. In addition, they provide diverse
substrates and physical structure to forested
terrestrial ecosystems.
Carbon sequestration, climate change
mitigation and resilience
Climate change poses a major threat to forestry,
biodiversity, agriculture and food security
through extreme climatic events, droughts,
increases in temperature, more frequent and
intense wildires, and increased activity of
pests and diseases. Effective FGR conservation
and management will take on even greater
signiicance against a background of such climate
change impacts and associated changes to forest
structure and composition. It will be increasingly
vital to provide the deepest possible reservoir of
genetic variability on which natural and artiicial
selection can act, facilitating adaptation to
changed conditions.
Under more extreme climatic conditions the
use of trees and forests for food and ibre is likely
to become even more important, for example
because of increased risks of failure of rain-fed
agriculture and annual crops. Trees and forests,
and their genetic resources, will also have an
essential and central role in helping to limit
rises in atmospheric carbon and slow climate
change through sequestration and storage of
atmospheric carbon. Mature, new and planted
forests can sequester substantial amounts of
carbon. Vigorously growing planted forests
sequester vast amounts of carbon; for example,
eucalypt hybrids in Brazil can sequester as much
as about 80 tonnes of CO2 per hectare. Brazil also
remarked on the importance of retaining healthy
natural forests (such as the Amazon forest)
to maintain global climatic conditions, noting
that this would also maintain a competitive
agricultural sector. Estimates put the carbon
storage of boreal forest at 703 gigatonnes,
tropical forests at 375 gigatonnes and temperate
forests at 121 gigatonnes (Kasischke, 2000).
Furthermore, billions of people use woodfuel
rather than burning fossil fuels.
While mature forests are more or less in carbon
balance, their between- and within-species
diversity helps to buffer them against change and
destruction (whether related to climate, biotic
factors, ire or a combination of these) which
might otherwise result in damaging releases of
CO2. Tree breeders will require genetic diversity
to develop faster growing, well adapted trees
for a diverse range of environmental conditions
for carbon sequestration and production of
woodfuels.
Resilience capacities of forest ecosystems are
conferred at multiple scales, through genetic,
species and landscape heterogeneity (Thompson
23
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
et al., 2009, 2012). The abilities of different
species, including tree species and genotypes,
to substitute functions is key to their buffering
of impacts of environmental change and
maintenance of ecosystem functioning (Walker,
1992; Lavorel, 1999; Yachi and Loreau, 1999;
Elmqvist et al., 2003; Hooper et al., 2005; Winfree
and Kremen, 2009; Thompson et al., 2012).
Accordingly, the ability of an individual forest
stand to adapt to and recover from environmental
changes will depend on the number of species,
their diversity, individual adaptive capacities
and abilities to substitute different functions.
The roles of forest genetic diversity in ecosystem
adaptation and resilience are fertile topics for
research, but in-depth scientiic investigations are
still in their infancy.
Social, cultural, medicinal and
scientiic value
Forest genetic resources have major social, cultural
and spiritual values, mainly at tree species level,
with many individual tree species distinguished
and named in local languages. In Fiji, for instance,
Intsia bijuga – a tree that has spiritual signiicance
throughout the Paciic islands – is called “vesi”,
which is also the name reserved for village chiefs.
Various native tree species are intertwined with
local cultures, customs, folklore, stories, poems,
and cultural identity and are integral to the daily
lives of indigenous peoples. Many thousands
of tree species are used in products, customs,
ceremonies and rituals, often developed over
millennia, that help give meaning to and enrich
the lives of hundreds of millions of people. In
many parts of sub-Saharan Africa, for example,
certain trees are considered sacred and are
maintained in sacred groves or church plantings.
In India between 100 000 and 150 000 sacred
groves have been reserved (India country report),
and certain tree species have tremendous social
and cultural importance, e.g. Ficus religiosa in
religious ceremonies, Santalum album in burial
ceremonies and Azadirachta indica in traditional
medicinal culture. In the Russian Federation,
24
7 980 ha of forest were opened to the public in
2012 for religious activities.
There are also numerous examples where
intraspeciic
tree
diversity
has
cultural
importance. For example, in the Paciic islands
there are hundreds of named varieties of
Pandanus tectorius, mostly selected female plants
propagated vegetatively. Different varieties are
used at different times for food, for different
types of leis, and in different types of thatched
mats and other plaited wares (Thomson et al.,
2006). Pandanus tectorius is also important for
construction materials, medicines, decorations,
perfumes, and many other cultural uses. For most
people in Kiribati it is the ancestral tree from
which, according to legend, their progenitors
came (Luomala, 1953).
Country reports noted that medicinal uses
of forest resources are extremely important
in some regions and countries. Many more
trees are utilized for medicine in Africa than in
other regions, with medical use named in 14.4
percent of total reported uses. As an example,
in Zimbabwe over 78 percent of the rural
population uses traditional medicines, mostly
derived from trees and woody plants, at least
once a year for humans and livestock. In the
Paciic island countries, medicinal use accounted
for 8.6 percent of reported uses, and medicinal
uses are also important in Indian Ocean island
countries such as Madagascar and Seychelles.
Medicinal uses of trees were also noted as
important throughout Asia, including China (with
nearly 1 000 medicinal plants used, mainly woody
species), India, Indonesia (with 2 039 medicinal
plants used) and Nepal.
The search for medicinal compounds, or
bioprospecting, has potential to yield dividends
to supplier countries where sound beneitsharing arrangements are in place. Many
tree and shrub species are exploited or being
investigated for medicinal purposes. Among
them are Homalanthus nutans from Oceania,
Prunus africana from the humid tropics of Africa,
Cinchona spp. from Latin America, Emblica
OVERV IE W
officinalis from India and Pinus sylvestris from
Europe, to mention some. In some countries
(including among others India, Madagascar and
Solomon Islands), the chemistry and medicinal
value of the lora are being investigated. Given
the vital importance of FGR for traditional
medicines and the potential beneits from
bioprospecting, there is a vital need for more
research on the medicinal value of forest trees to
help unlock the full potential of FGR.
FGR are of major scientiic value. Intraspeciic
diversity can be used, for example, to help
understand the genetic, biochemical and
physiological basis for resistance to pests and
diseases or environmental stresses such as extreme
climatic events (drought, looding) and edaphic
extremes (salinity, acidity, etc.). It can also be used
to identify biosynthetic pathways for production
of important products and metabolites. A recent
and surprising example of the potential scientiic
importance of a previously little-known tree
species is provided by Amborella trichopoda,
a small understorey tree endemic to the wet
upland forests of New Caledonia (France), which
is endangered by habitat destruction. Amborella
trichopoda appears to have diverged earlier
than other lowering plants (about 130 million
years ago) and lacks vessels in the wood which
are characteristic of other angiosperms. In 2012
the Amborella Genome Project (www.amborella.
org) produced a draft genomic sequence which
will be used as key evidence for understanding
the ancestral state of every gene, gene family,
and protein sequence in lowering plants and
their radiation through the history of lowering
plants. This genomic information may provide
insights into the evolution of lowering and
vessel formation in wood.
Preserving options for future
development and adaptation
One of the most important characteristics of
FGR is that they will be vital for adaptation
to future changes – not only to those that are
becoming evident such as climatic extremes and
new warmer climates brought about by increases
in atmospheric CO2, but also to others of which
little is known. Based on geological records,
the Earth is likely to return to a new period of
glaciation possibly 3 000 to 20 000 years hence,
but the possible long-term impacts of humaninduced global warming on a future glaciation
event are unknown. In the meantime it would
be reprehensible to allow useful tree species and
populations adapted to cooler climates to become
extinct from global warming and other factors
when their germplasm might be conserved safely
and relatively cheaply in cold storage such as the
Svalbard Global Seed Vault in Norway (−18oC) for
several hundred to thousands of years.
The importance of maintaining FGR to preserve
options applies to both natural forests, where
a vital dimension is the capacity to adapt to
changing environments, and planted forests,
which may hold the key to new products and
services while at same time proving resilient. In
the case of planted forest tree species, there is a
need to maintain as much intraspeciic diversity
as possible to allow tree breeders to continue
to select and develop improved and adapted
germplasm to cope with new demands and
growing conditions. Conservation of intraspeciic
diversity will also serve the development of
new wood products and NWFPs, especially
pharmaceuticals and neutriceuticals such as
sources of antioxidants, anti-inlammatories
and other chemoprotective natural compounds.
Novel uses may be elaborated, such as breeding
of trees speciically to sequester carbon or to
recycle plant nutrients from beyond crop root
depth, or “harvesting” of precious minerals
through phytomining. Wood and Grauke (2010),
for example, have found that tetraploid Carya
species accumulate high amounts of rare-earth
metals (almost 0.1 percent dry weight), much more
than diploid Carya species and other tree species.
In New Caledonia (France), Boyd and Jaffré
(2009) recorded several tree species, including
Geissois pruinosa and Homalium kanaliense,
as hypernickelophores, i.e. species that can
25
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
accumulate large amounts (up to 1 percent of
leaf dry mass) of nickel. The differential ability
of plants to accumulate gold is also well known
(e.g. Girling and Peterson, 1980) but has not
been commercially exploited to date. In future,
certain tree species and genotypes might be used
or selected and bred for phytoremediation, i.e.
to remove or neutralize contaminants, say from
polluted soil or water (e.g. Raskin and Ensley,
2000; Pilon-Smits, 2004).
To sum up, it is evident that well characterized
(in terms of growth and adaptability attributes,
including genotype by environment interactions,
and the type and quality of end products and/
or services), genetically diverse wild populations
(or provenances) of different tree species will
provide extremely useful genetic materials both
26
for immediate planting programmes and as
the basis for future selection and improvement
programmes. The diverse values and uses of
forests, trees and FGR identiied in country
reports underscore the need for national FGR
strategies and effective programmes and action
plans to address not only the applications and
requirements of the formal forest and forest
industry sectors, but also the role of FGR in the
informal economy, in alleviating poverty, in social,
cultural and spiritual areas, and in environmental
services and rehabilitation. The major contribution
of FGR to the informal economy highlights the
need to consult with the widest range of forest
users possible when preparing national strategies
and programmes.
OVERV IE W
Chapter 3
Conservation of
forest genetic resources
Individual trees contain genetic variations that
distinguish them from other members of their own
species and other species. Variation is continuously
generated through sexual recombination and
mutations, and natural selection acts on this
background of variability through the process
of evolution, producing new variants that are
better adapted to survive, to compete and to
cope with changing environmental conditions.
Genetic variation provides the basis for selection
of genotypes and varieties better suited to
meeting human needs (i.e. able to provide more
useful products or services in a more eficient
manner) in a wider range of settings and under
changing environmental conditions. Provision
of forest-derived goods and services depends on
the presence of FGR and also has implications for
their survival.
The future value of FGR will be determined
by the way humans manage these resources
and act in their role as the primary agents of
environmental change in today’s world. Humans
inluence the value of FGR even when they are
not aware of doing so, by using trees and forest
resources and altering environmental conditions,
as much as through conscious efforts to better
conserve and manage them. Indeed, the growing
awareness of how human actions, or lack thereof,
have impact on FGR is a recurrent theme in
country reports. However for the purposes of this
report, and given the imperative to recognize
that the future of FGR depends on conscious,
effective human intervention through deliberate
management, the term “management” is used
to describe deliberate planned actions taken to
conserve and protect FGR.
Conservation of forest genetic resources can be
deined as the policies and management actions
taken to assure their continued availability and
existence. Conservation and management of FGR
are inextricably intertwined. Conservation of
FGR requires implementation of well planned,
scientiically
sound
strategies,
including
management of FGR in breeding programmes
and in production populations.
The strategies and methodologies applied
in conservation depend on the nature of the
material, the timescale of concern, and the
speciic objectives and scope of the programme.
Two basic strategies are used for genetic
conservation: in situ (on site) and ex situ (off site,
e.g. in designated conservation stands, ield gene
banks, arboreta and botanic gardens). These two
strategies are generally viewed as complementary
and are best carried out in parallel when the
aim is conservation of species and intraspeciic
genetic variation. Coordination is required
among the various agencies and organizations
concerned, i.e. forestry departments managing
reserved forests and in situ gene conservation
stands; environment departments managing
protected areas; government agencies, private
forestry companies and cooperatives carrying
out tree improvement programmes; government
research agencies, botanical gardens and
universities maintaining gene banks of seed
and tissue cultures; and private landholders
and communities managing privately owned
27
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
managed forests, planted forests, agroforests
and farmlands. Both the general and particular
strategies and programmes to be pursued will
depend on factors such as the available inancial,
human and land resources; human population
and resource use pressures on land, forests
and trees; technological options for particular
species; and the nature and dimensions of the
conservation challenges, e.g. whether the aim is
to conserve a large number of forest species, a
smaller number of rare and endangered species,
or the genetic diversity and evolutionary potential
of a smaller number of high-priority species for
planting programmes. Additional challenges and
opportunities arise in situations where there is an
international dimension, e.g. when the natural
range of a species crosses national borders; when
a species has greater economic importance as a
planted exotic than it has in its native habitat;
and when opportunities for ex situ conservation
in well resourced facilities (such as tree seed banks
or tissue culture facilities) are available abroad.
The management of forest genetic resources
to ensure their conservation, improvement and
sustainable use simultaneously is a complex
technological
and
managerial
challenge.
Fortunately, when simple basic principles are
applied, the production of goods and services
from managed forests forming part of a legislated
permanent forest estate is generally compatible
with the genetic conservation and development
of particular forest tree species, as discussed in
the next section.
Management systems in the
ield (in situ and circa situm
conservation)
Forest genetic resources conservation and
management in the ield should ideally be
considered and integrated into all land uses and
management systems containing trees. The most
important of these are sustainably managed
multiple-use production forests, protected forests
and agroforests, described in the following
sections.
28
Ecosystem- and landscape-based conservation
approaches and management regimes can also
conserve a wide array of forest tree species and
their diversity in situ. These approaches lack a focus
on particular tree species except in the case of
keystone species whose continued existence and
diversity are vital for maintaining the ecosystem’s
health. The ecosystem approach is well suited
to areas with high tree species diversity, such
as lowland moist tropical forests; it can ensure
the continued survival and availability of large
numbers of useful tree species which may have
localized and/or potential importance. However,
in locations where local human populations
rely on a vast number of tree and woody
species to provide diverse products, sustainable
management of multiple-use production forest is
a more effective approach.
Conserving and managing the variability
of FGR in situ provides the basis for selection
and adaptation and will promote continued
ecosystem function and services. Forest genetic
diversity helps ensure healthier, more resilient
forests better able to deliver essential functions.
Adaptation to changing environmental inluences
requires a high degree of genetic diversity in tree
species because of their immobility and perennial,
long-lived life forms. Where forests have been
degraded, the use of appropriate species and
provenances, selected from the pool of natural
variability maintained through effective FGR
conservation and management, can assist forest
restoration (Box 3.1). Increasingly the diversity
within and between tree species is being found to
be critical to promoting and maintaining almost
all other life forms present in forest ecosystems.
Increasingly many socio-economically important
tree species are being conserved through use;
they are often planted for productive and other
purposes in planted forests, agroforests, orchards
and urban landscapes (in home gardens and
parks and along streets). The conservation of
FGR in these cases is often incidental, unplanned,
and suboptimal; an exception is breeding
programmes and seed orchards which are often
OVERV IE W
Box 3.1
Application of genetic principles in forest ecosystem restoration
and management
Forest ecosystem restoration is of growing interest
as a means of mitigating climate change and of
combating its negative impacts, which are associated
with continued deforestation and degradation of
forest ecosystems worldwide. Typically, however, little
attention is paid in restoration initiatives to ensuring
the use of appropriate sources of forest reproductive
material. While geographical origin of planting
material is often considered – i.e. it is commonly
understood that local sources are preferable – other
genetic factors are rarely considered (Bozzano et
al., 2014). Guidelines recommending the number of
source trees needed for provision of reproductive
material are sometimes available, but they are often
inadequate. While tree planting for production is
regulated with regard to provenance seed source
delineation and the use of certified or source-
identified seed stand collections, restoration projects
are not legally required to use certified or identified
seeds.
Examples of knowledge gaps in the genetics of
forest ecosystem restoration include quantification
of the risks associated with genetic mismatching
of source of planting material to site conditions
or narrow genetic base, particularly considering
climate change; thresholds for optimal genetic
diversity in restoration material; and genotype ×
environment interactions. It would also be valuable
to understand the potential for combining species
rescue with ecosystem restoration, i.e. the potential
of individual restoration projects to contribute to
species conservation and serve as future seed sources,
especially for rare, endemic and endangered tree
species.
designed for improvement and to maintain a
balance of diversity. Urban landscapes, farms,
hotel resorts, golf courses, etc. offer opportunities
for conserving substantial tree species diversity
(including within-species diversity) if managers
of these areas are made aware of the importance
of tree conservation activities and are linked into
national FGR programmes. However trees outside
forests in non-agricultural land-use systems such
as urban landscapes are outside the scope of this
report.
Species
selection
and
availability
of
reproductive material is another issue of concern.
Exotic species or seed sources are commonly used
for restoration and in some cases are clearly
justiied. In other cases, however, the use of exotic
species not only can result in wasted efforts, but
can threaten native species if the exotic ones
become invasive. Often the reason for using exotic
species is simply local availability in nurseries and
level of knowledge about nursery production
requirements. Expanded knowledge of native
tree species is needed in order to understand
their potential to achieve diverse restoration
objectives in different states of site degradation
and different ecological, environmental and
socio-economic contexts. It is also important to
understand the trade-offs (again, ecological,
environmental and socio-economic) related to
the use of exotic versus native species, as well as
the factors that currently constrain wider use of
native species.
Sustainably managed multiple-use
production forests
Sustainable forest management involves the
management of forests in a manner that ensures
that their overall capacity to provide environmental
and socio-economic beneits is not diminished over
time. Central to the sustainable development of
forests is the challenge of balancing resource use
and conservation. Sustainable forest management
29
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
and the maintenance of FGR must be considered
as interdependent: Forest owners, custodians and
managers need to understand and appreciate the
essential underpinning role of FGR in forest and
natural resource management practice in order
to implement effective interventions for their
conservation and use (Ratnam et al., 2014). Many
management measures have been developed
to maintain diversity in forest ecosystems and
simultaneously to promote the sustainable use
of this diversity (see e.g. FAO, 1993; Thomson,
2004). What is lacking is their constant application
and monitoring. Furthermore, harmonization of
conservation objectives and utilization practices
in production-oriented, multiple-use native forests
will be essential for conservation of the diversity
of most of the tree species, given that they are
not well represented in protected areas, planted
forests and ex situ collections (Thomson, 2004).
Diverse indigenous forest management systems
and practices have employed technologies for
managing and using native forests in a manner
that does not diminish, and preferably enriches,
their FGR. Forest agencies and private forestry
companies can readily integrate FGR management
practices into modern silvicultural systems.
However, in many parts of the world, tree species
diversity and intraspeciic diversity are declining
because best-practice forest management systems
are not being implemented or are breaking down
for various reasons, often as a result of increased
population pressure and associated unsustainable
use including overharvesting of timber, fuelwood
and NWFPs; reduction in seed sources of pioneer
and early secondary trees; and insuficient
time for deep-rooted perennial vegetation to
replenish soil fertility between shortened fallow
periods. Furthermore the area under production
forests, considered vital for conservation of FGR,
has continued to decline at an increasing rate,
by about 2 million hectares per year during the
1990s and 3 million hectares per year between
2000 and 2010 (FAO, 2010a) (see Part 2 on drivers
of change).
Some 80 percent of the world’s forests
are under public ownership, and 80 percent
30
of publicly owned forests are under public
administration (FAO, 2005a), suggesting that
national governments are in a strong position to
directly inluence and control forest management
practices. However, in the developing tropics,
many production forests are under private
logging concessions, and governments frequently
lack resources to develop sustainable best
practices such as codes of logging practice and
reduced impact logging guidelines, or to enforce
their implementation by private operators. The
problem is compounded where concessions
are issued for a short term or only once, as the
logging concessionaire in such cases is likely to
harvest in a manner that will maximize proits,
possibly with little consideration for regeneration
and subsequent harvests.
Sustainable production of goods from natural
forests is expected to be increasingly challenged
by more extreme climatic events in future
(particularly more intense tropical cyclones,
droughts and associated bushire, intense rainfall
events with landslips and looding, and melting
of permafrost). Interactions of climate change
with existing and new pests, diseases and invasive
weeds, as well as climate change impacts on
pollinators and dispersers, will affect production,
selective forces and the future forest composition.
The genetic diversity contained within and among
tree species will provide essential buffering for
these impacts on many productive and service
functions of forests, but a much greater level
of management intervention and manipulation
may be required, including movement of tree
germplasm to respond to new climates, changed
pests and diseases and new selective pressures.
Sustainable forest management cannot by itself
ensure conservation of all FGR. Some tree species
and populations require special and immediate
attention, and many species that are of no or
little current utilitarian value will probably
receive little attention from forest managers.
Some of these lesser-known or less economically
important species may depend on complicated
ecological interaction and may suffer from what is
believed to be gentle use of the forest resources.
OVERV IE W
Therefore, an integrated approach encompassing
management of natural stands and establishment
of speciic conservation populations is advocated.
Protected areas
Existing national protected area systems are
often a valuable starting point for a network
of conservation stands of a particular species.
Indeed it is likely that several thousand tree
species only occur within the existing protected
area network. However, the security of forest
protected areas remains a major concern,
especially in developing nations, where it is likely
that many face threats to their integrity and
existence in the medium term. Fully protected
areas are only likely to succeed long term in
areas of low population pressure.
On a positive note, FRA 2010 (FAO, 2010a)
found that the area of forest designated for
conservation of biological diversity increased by
about 6.3 million hectares per year during the
decade from 2000 to 2010, and a similar increase
occurred in the area of forest in protected areas.
In both cases the increase is equivalent to nearly
2 percent per year.
Agroforestry systems, including trees on
farms
The development of context-speciic agroforestry
systems, integrating traditional knowledge and
scientiic advances and based on diverse, adapted
tree germplasm – as in the work of the World
Agroforestry Centre (ICRAF) and national and
non-governmental organization (NGO) partners
– offers one of the most promising solutions
for addressing problems of overpopulation and
limited land base. It is estimated that 1.2 billion
people use trees on farms to generate food and
cash (World Bank, 2002), and almost half of the
agricultural land in the world, or more than
1 billion hectares, has a tree cover of more than
10 percent (Zomer et al., 2009). The importance
of using appropriate, matching, diverse and
improved germplasm in agroforestry systems
has been increasingly appreciated over the past
two decades, including the need for appropriate
seed and seedling production and dissemination
systems (see Box 3.2). Many indigenous fruit- and
nut-tree species have been domesticated to
provide a source of nutrition and income for rural
households. ICRAF coined the term “agroforestry
tree product” (AFTP) for these new products. The
research and development and extension efforts
in agroforestry will continue to show results as
long as the genetic diversity on which they rely
is both conserved and accessible. In addition,
knowledge on the importance of germplasm
and on its selection and improvement has spilled
over from conventional plantation forestry to
agroforestry research and development.
Box 3.2
Evolving use of tree germplasm in
modern agroforestry in South Paciic
islands
In Fiji and other island nations of Oceania, through
the mid-1970s and early 1980s the official promotion
of village forestry mainly consisted of distribution
of seedlings of Pinus caribaea, including those left
over from P. caribaea planting programmes. During
the 1980s the emphasis moved to alley cropping
systems with fast-growing nitrogen-fixing exotic
trees, supported by the Fiji-German Forestry Project.
Calliandra calothyrsus was shown to be suitable,
but farmers did not adopt the systems. During the
mid-1990s and 2000s the South Pacific Regional
Initiative on Forest Genetic Resources (SPRIG) project,
funded by Australia, worked with national partners to
develop and domesticate a much broader selection
of native tree species and a few key exotics, such as
Swietenia macrophylla and Tectona grandis, which
are now being incorporated into a diverse range of
agroforestry systems, including modified traditional
polycultural systems. The extremely cyclone tolerant,
multipurpose timber tree Terminalia richii, which had
been reduced to scattered trees by the mid-1980s, is
now being widely planted by smallholder farmers and
tree growers in Samoa.
31
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
Ex situ conservation
The primary aim of ex situ conservation has always
been to ensure the survival of genetic resources
which otherwise would have disappeared. For
forest genetic resources, ex situ conservation
has generally referred to storage as seed, when
practical, usually under conditions of low moisture
content. Having evolved over millennia to ensure
the dispersal of the species genomes, pollen and
seed are ideal starting materials for innovative
conservation programmes. Ex situ approaches of
signiicance include enhancing seed production
through artiicial pollination and treatments
ensuring eficient seed germination to generate
the next cohort of plants (which are beyond the
scope of this review). Such work is underpinned
by knowledge of the species distribution and
the application of horticultural skills to seedling
growth.
Where species are intolerant of seed storage
conditions, it has been necessary to rely on ield or
glasshouse collections. However, such collections
are costly to maintain and are at risk from pest
and disease outbreaks and climate variability and
extremes, and therefore are not as safe as seed
storage for long-term conservation. For these
reasons in vitro technology has been proposed as
an alternative strategy.
To improve the ex situ conservation of forest
genetic resources using seed storage, signiicant
effort has to be spent in developing postharvest
technology for proper handling and identiication
of storage behaviour. Once seeds of a particular
species have been classiied, then strategies can
be developed for their conservation according to
their storage behaviour.
Gene banks
Conventional seed storage is believed to be a safe,
effective and inexpensive method of conservation
for seed-propagated species. Many countries have
national tree seed banks, but these are often active
collections with rapid turnover and use of collected
seedlots, in which conservation is a supplementary
or incidental beneit. Yet large numbers of trees
and woody species can be conserved long term in
32
seed banks. A major international example of this
conservation strategy for trees and woody species
is the Kew Millennium Seed Bank Partnership
(see Box 3.3). As an example of the partnership
in action, the National Forest Seed Centre (CNSF)
in Burkina Faso has 160 tree species in long-term
cold storage (Burkina Faso country report). In the
Russian Federation a notably large number of seed
banks contribute to the collection and storage of
forest seeds; among them is the Vavilov Research
Institute of Plant Industry (www.vir.nw.ru), which
stores some 323 000 samples of 2 169 plant species.
For successful long-term conservation through
seed storage, it is necessary to determine the
factors that regulate seed viability and vigour.
Viability must be monitored continuously, with
recollection or regeneration whenever the
Box 3.3
Millennium Seed Bank Partnership
The Millennium Seed Bank Partnership, based in
the United Kingdom (see www.kew.org/scienceconservation/millennium-seed-bank), is the largest
ex situ conservation project based on seed storage
in the world; the project has banked 10 percent
of the world’s wild plant species, including many
woody species, and aims to conserve 25 percent
of wild plant species by 2020. This expansive
global partnership involves about 50 countries and
agencies such as FAO and Bioversity International.
It was developed in part to enable countries to
meet international conservation objectives set by
the Global Strategy for Plant Conservation of the
Convention on Biological Diversity (CBD) and the UN
Millennium Development Goals (MDGs).
The research being conducted by the project into
the challenges of seed banking, such as postharvest
handling (including seed sensitivity to drying)
will significantly expand existing possibilities for
conservation of forest genetic resources. To date the
seeds of more than 20 important palm species and
around 100 dryland species have been tested for
tolerance to drying.
OVERV IE W
viability drops below an acceptable level. Scientiic
collaboration in plant conservation has led to
substantial innovation in seed storage in recent
years, particularly in diagnosing tree seed storage
behaviour, increasing tree seed longevity in the
dry state and improving storage biotechnology
(Pritchard et al., 2014).
Seeds can be categorized according to their
storage behaviour, which is a relection of
the seed moisture content. The inal moisture
content in the seeds depends on the species
and the external environment. Before the seed
from any species can be considered for storage,
its response to desiccation and chilling must be
determined.
• Orthodox seeds dry out to 5 to 10 percent
moisture during maturation. They are shed
in a highly hydrated condition, endure a
chilling period during maturation and are
therefore adapted to the low temperatures
used for orthodox seed storage. They can
be stored for long periods at seed moisture
contents of 3 to 7 percent (on a fresh
weight basis) at –18°C or below (Theilade
and Petri, 2003).
• In contrast, recalcitrant seeds maintain
relatively high moisture content, generally
greater than 40 to 50 percent, and cannot
be stored in conventional seed banks
because of the sensitivity of the seeds to
desiccation. Many forest tree species from
temperate and especially tropical regions
produce recalcitrant seeds.
– Seeds of temperate recalcitrant
species can be stored at near freezing
temperatures for several years but
are intolerant of drying. For example,
Quercus species can be stored for three
to ive years as long as a high (35 to
40 percent) seed moisture content is
maintained.
– Seeds of tropical recalcitrant species
require the same gas and moisture
levels but are very sensitive to low
temperatures. For example, species from
the genera Shorea, Hopea and several
tropical fruit-trees will lose viability at 10o
to 15oC (Phartyal et al., 2002).
• An intermediate category has been
identiied in which seeds are partly tolerant
to dehydration and cold. The longevity
of these seeds is quite short, which is a
signiicant constraint for conservation in a
number of species, including many tropical
forest trees (Joët et al., 2009).
Seed behaviour is generally considered as a
continuum from orthodox to recalcitrant. The
number of species identiied with non-orthodox
behaviour is increasing, and the basis of such
behaviour is more complex than initially envisaged
(Berjak and Pammenter, 2008). Following
application of new knowledge of seed physiology
and some success with seed drying around the
world, some species once considered recalcitrant
have later been identiied as orthodox. Fagus
sylvatica and two tropical species, Citrus limon
and Elaeis guineensis, for example, fall into this
category (Phartyal et al., 2002). Techniques have
also been developed for extending the viability of
non-orthodox seed.
Some genera, such as Acer (Phartyal et al.,
2002) and Shorea (Theilade and Petri, 2003),
include both orthodox and recalcitrant species.
Infrequently, apparent seed storage behaviour
may vary geographically within the same species,
as in Dipterocarpus alatus, in which populations
from drier zones have more desiccation-tolerant
seeds, and Santalum austrocaledonicum (Thomson,
2006). Behaviour may even vary depending on the
stage of maturation at collection and the storage
and rehydration regimes, as in Azadirachta indica
(Sacandé and Hoekstra, 2003).
It is estimated that approximately 60 percent
of the tree species in the Amazon Basin produce
recalcitrant seeds. In comparative studies of seed
morphology in relation to desiccation tolerance
and other physiological responses in nearly 200
moist tropical forest species on three continents,
from 42 to 62 percent of species were found to
have non-orthodox behaviour, with an overall
average of 51 percent (Hamilton et al., 2013;
Ferraz et al., 2004; Ellis et al., 2007).
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Recalcitrance reduces the effectiveness of seed
banks in areas where threats to tree species and
their populations are greatest. However, during
the past 20 years, considerable progress has been
made in:
• understanding the mechanisms of
desiccation-induced loss of viability on
drying, including the role of reactive
oxygen species and programmed cell death
(Kranner et al., 2006);
• estimating the proportion of the world’s
lora that produces such seeds (diagnosis);
• developing methods that can help conserve
such species in ex situ cryobanks (storage
biotechnology).
Short- to medium-term storage of recalcitrant
seeds can be achieved by maintaining the seeds at
the lowest temperature they will tolerate, under
conditions that do not allow water loss. However,
these conditions will encourage the growth
of micro-organisms which must be managed
through appropriate action, such as fungicide
treatment (Berjak and Pammenter, 2008).
Fungicide treatment was effective in extending
the storage life of Hopea parviflora (Sunilkumar
and Sudhakara, 1998). Problems with seed
handling and storage affect the implementation
of conservation programmes.
Generally a relationship between seed size and
tolerance for desiccation holds (Hamilton et al.,
2013). As Daws and Pritchard (2008) discovered,
working with Acer pseudoplatanus, it is important
to pay attention to phenotypic plasticity in the
seed storage response between plants growing
inside and outside the native distribution range,
as this can be responsible for species appearing to
be in a different seed storage category or being
misclassiied.
The half-life (time for viability to fall from 97.9
to 50 percent) of seed in storage varies greatly,
in part depending on the oil content of the seed
in relation to mass. Estimates range, for example,
from 0.95 for Dipterocarpus alatus to 342 years
for Liquidambar styraciflua (Daws, Garwood
and Pritchard, 2006). The information required
34
to predict half-life is not available for most tree
species, but recent advances have improved the
accuracy of predictions (see also Box 3.4). The
shortest life spans are driven by the need to store
the seeds partially hydrated at 10 to 17 percent
moisture content. In contrast, the longest-lived
Box 3.4
Biological models for predicting risk
associated with seed storage for tree
species
It is important to develop predictive biological models
to indicate risks associated with handling of seed with
particular characteristics because the large number
of tree and other higher plant species, estimated
to be as many as 353 000 (Scotland and Wortley,
2003; Chapman, 2009), renders the physiological
screening of all species unlikely in the foreseeable
future. Early studies revealed broad associations
between heavier seed in the Araucariaceae (Tompsett,
1984) and Dipterocarpaceae with seed desiccation
sensitivity. Hong and Ellis (1998), for example, using
multiple criteria, identified associations between
habitat and desiccation intolerance across a broad
range of vegetation types, finding a low frequency
(about 10 percent or less) in the driest regions of
the world and a high frequency (close to 50 percent)
for tropical moist evergreen forests. For about 70
African tree species, Pritchard et al. (2004) confirmed
predictions that in seasonally dry environments (more
than in areas that are moist year-round), species
that produce recalcitrant seeds disperse their seed
in the rainy season, as they must maintain water
status or else they die. Another ecological prediction
that has proved useful is that recalcitrant seeds
should not need as great a defence mechanism
against consumption, e.g. thick seed coats, since
their germination is relatively quick (Hong and Ellis,
1998). Using this relationship, Daws, Garwood and
Pritchard (2006) developed a predictive model for the
probability of recalcitrant seed among 104 trees in
Panama.
OVERV IE W
seeds can be stored in a much drier environment,
at around 3 percent moisture content. Some
species have very long-lived seed; some seeds of
three woody species of the Cape Flora of South
Africa were germinated after about 200 years
storage under museum or cellar conditions.
Seed storage duration varies across different
plant species, hence the Millennium Seed Bank
Partnership (see Box 3.3) is evaluating the impact
on seed storage duration of various factors such
as structure of the seed embryo and climate
conditions during seed development and ripening.
Baseline data on the desiccation tolerance and
longevity of tree seeds are very limited (Hong,
Linington and Ellis, 1998; Dickie and Pritchard,
2002). The DANIDA Forest Seed Centre (now part
of Forest Landscape Denmark) and Bioversity
International led a global initiative from 1996 to
2002, involving about 20 countries, which screened
recalcitrant and intermediate (with partial
desiccation tolerance but with sensitivity to storage
at –20°C and 0°C) seeds of 52 tropical forest trees
belonging to 27 families (summarized in Sacandé
et al., 2005). The project assessed seed responses to
multiple desiccation states and subsequent storage
at a range of temperatures (Hong and Ellis, 1996)
to understand seed storage behaviour.
An alternative screening approach, called
the 100-seed test as that is the target number
of seeds to use (many less than the previous
protocol), deals only with aspects of the effects
of drying and short-term storage on the initial
moisture content of the seed sample (at receipt or
harvest). This approach gives a good indication of
tolerance to rapid artiicial drying similar to that
used in seed banks; in addition, the moist-stored
control can show reduced germination (rapid loss
of viability and thus short life span), no effect on
germination, or increased germination (evidence
of seed maturity during the storage period). This
method has been adopted by tree seed experts at
the Instituto Nacional de Pesquisas da Amazônia
(INPA), Brazil; the University of Queensland,
Australia; and the University of KwaZulu Natal,
Durban, South Africa.
However, knowledge of the seed biology of
forest tree species is limited, and the resources
that are available are scattered. The University
of Copenhagen, in collaboration with the World
Agroforestry Centre (ICRAF), has published a
series of seed lealets since 2000, which are posted
online together with lealets published by the
DANIDA Forest Seed Centre from 1983 to 1986
(http://sl.ku.dk/rapporter/seed-lealets). A study
of seed from 100 Panamanian tree species has
also generated important information (Sautu et
al., 2006). The Tropical tree seed manual (Vozzo,
2002), which includes 175 tree species, is another
highly useful publication. It would be beneicial
to bring all of the information together on one
portal so that ex situ conservation actions can
be supported. More information is needed on
the control of tree seed germination, including
how dormancy can be alleviated and the factors
inluencing varying degrees of dormancy, such as
time of collection and climatic conditions. Sources
of information should include compendia of
national and regional forest seed programmes.
Under the Millennium Seed Bank Partnership,
a unique seed database has been established
which provides information on a wide range of
functional traits or characters, including, among
others, seed desiccation tolerance, germination
and dormancy, and classiies seeds accordingly. At
the same time the lack of knowledge for tropical
species is acknowledged. ICRAF maintains the
Agroforestree Database, which provides storage
information for 670 agroforestry tree species.
In vitro conservation
Many commercially valuable tropical tree
species are estimated to have recalcitrant or
intermediate seeds (Ouédraogo et al., 1999), for
which long-term conservation using conventional
seed storage is not possible. For this reason
signiicant effort has gone into establishing in
vitro approaches for conserving forest genetic
resources. However, woody species are often
dificult to establish in vitro, with problems
35
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
occurring at any one of the multiple stages of
shoot culture establishment.
The irst stage of establishing cultures derived
from mature forest trees can be challenging
because of high levels of contamination and/
or high secretion of polyphenols and tannins.
A review of the progress made in establishing
tissue cultures of threatened plants (Sarasan et
al., 2006) highlights a range of methods that
have been developed to initiate cultures of
often recalcitrant plants of limited number, as
well as different approaches to managing tissue
and medium browning. Successful initiation of
in vitro cultures is not the only challenge; of key
importance is the establishment of stabilized shoot
cultures to provide a stock of plants that are more
reproducible and stable than those in the ield
or greenhouse. Despite progress in this area, in
vitro shoot growth stabilization – that is, culture
with uniform and continuous shoot growth – is
not well understood (McCown and McCown,
1987). However, rejuvenation is undoubtedly a
major contributing factor; explants derived from
juvenile sources are easier to establish in vitro than
adult plants of the same genotype. The use of
juvenile tissue has been successful in a number of
species, for example, Acacia auriculiformis, Acacia
mangium, Aqualaria malaccense, Azadirachta
excela, Calamus manan, Dyera costulata and
Tectona grandis (Krishnapillay, 2000).
Episodic species, such as many nut-trees and
conifers, are highly problematic for tissue culture
as compared with the sympodial species (such
as many pioneer trees), which show continuous
seasonal shoot growth. Success using in vitro
approaches is generally found in non-episodic
species, for example Eucalyptus and Populus
species (McCown, 2000). Episodic trees tend to
maintain their episodic growth pattern in culture,
so that random lushes of growth are followed
by periods of inactivity during which the cultures
deteriorate. However, two approaches to culturing
highly episodic species have been successful. One
approach uses the generation of shoots de novo,
the actual induction of adventitious meristems
36
being a rejuvenation process in itself. This
approach has been very successful with conifers
(Ahuja, 1993). The second approach focuses on
rejuvenation either of the stock source or of
the tissue-cultured tissues (Greenwood, 1987;
McComb and Bennett, 1982). Multiplication
and rooting of shoot culture systems for tree
species can be demanding, with very speciic
requirements depending on the species and
often the variety. For example, multiplication and
rooting of the endangered tree Ginkgo biloba
was promoted by incorporating the endosperm
from mature seeds of the same species in the
culture medium (Tommasi and Scaramuzzi,
2004). Pijut et al. (2012) reviewed in vitro culture
of tropical hardwood tree species from 2001 to
2011, outlining methods used for a wide range
of species of this commercially important group.
Only once an eficient and effective system for
generating stabilized shoot cultures is established
should there be any attempt to develop an in
vitro storage protocol. In vitro conservation
technology provides two options: restricted or
minimal growth conditions and cryopreservation.
Minimal growth culture. The most popular
methods for minimal growth storage are
modiication of the culture medium and reduction
of the culture temperature or light intensity.
Minimal growth storage has been reported
for several tree species such as Eucalyptus
grandis (Watt et al., 2000), Eucalyptus citriodora
(Mascarenhas and Agrawal, 1991) and Populus
spp. (Hausman et al., 1994). Malik, Chaudhury
and Rajwant (2005) reported in vitro conservation
of Garcinia indica with subculture duration of up
to 11 months after the establishment of cultures
from adventitious bud-derived plantlets.
Minimal growth culture is generally only
considered as a short- to medium-term
conservation approach because of problems
in the management of collections even if the
intervals between transfers are extended, and
also because of concerns of genetic instability
caused by somaclonal variation. In addition, it is
OVERV IE W
generally very dificult to apply one protocol to
conserve genetically diverse material. A study of in
vitro storage of African coffee germplasm, which
included 21 accessions, showed large variability in
the responses: some accessions experienced losses
whereas others were safely conserved (Dussert et
al., 1997). Technical guidelines are available on
establishing and maintaining in vitro germplasm
collections, although not speciically for forest
genetic resources (Reed et al., 2004).
Cryopreservation.
Besides improvements in
seed storage protocols, the main innovation for
recalcitrant seed has been improved techniques
for cryopreservation, which is needed for species
that produce fully hydrated recalcitrant seeds.
Cryopreservation is the storage of biological
material at ultra-low temperatures, usually that
of liquid nitrogen, –196oC. At this temperature
all cellular divisions and metabolic processes are
stopped, and therefore the material can be stored
without alteration or modiication, theoretically
for an unlimited period of time. In addition, cultures
are stored in a small volume, are protected from
contamination and require very little maintenance
(Engelmann, 2004). One of the disadvantages
of minimal growth storage is the possibility of
somaclonal variation. Cryopreservation reduces
this possibility because the metabolism of the
plant cells is suspended and subculturing is not
part of the process. However, the cryoprotocol
does expose plant tissues to physical, chemical
and physiological stresses which can all cause
injury. Although few studies have examined the
risk of genetic and epigenetic alterations, there
is no clear evidence that cryopreservation causes
morphological, cytological or genetic alterations
(Harding, 2004). For example, the genetic idelity
of Melia azedarach after cryopreservation was
conirmed using isoenzyme analysis and RAPD
markers (Scocchi et al., 2004). Cryopreservation
is particularly useful for conserving embryogenic
cultures of conifers whose growth and embryogenic
potential could be affected by regular subculturing
in conventional in vitro storage.
Cryopreservation is also more cost effective
than minimal growth storage. To date studies
on cost effectiveness have only been conducted
on crop plants, but the annual maintenance of
the cassava collection (about 5 000 accessions) at
the International Centre for Tropical Agriculture
(CIAT) is USD 30 000 for slow growth storage
and USD 5 000 for cryopreservation (Engelmann,
2010).
The extent to which the seeds can be safely
cooled is limited by the risk of ice crystal formation
for temperate species and chilling stress for
tropical species. Ice crystals can cause irreparable
damage to cell membranes, destroying their
semipermeability. The primary development of
the past 25 years in plant cryopreservation has
been improvement in methods for vitriication
to avoid crystal formation (Fabre and Dereuddre,
1990) and the use of complex solutions of
cryoprotectants that reduce the risk of ice crystal
formation of partially hydrated tissues during
cooling and rewarming.
Vitriication – the formation of an amorphous
or glassy state (non-crystalline) from an aqueous
state – signiicantly reduces cellular water. For
cells to vitrify, a concentrated cellular solution
and rapid freezing rates are required. Three
categories of explants can be cryopreserved
for woody species: shoot-tips for species that
are vegetatively propagated, seeds or isolated
embryo axes for species that reproduce using
seeds, and embryogenic calluses.
Most of the research has been on species
that are of interest to commercial forestry or
are valuable fruit-trees. Many studies report
successful vitriication of the intracellular
constituents when cooling followed partial
drying in air; for example, the embryos or
embryonic axes of ive citrus species were
cryopreserved after desiccation to about 15
percent moisture content (Malik, Chaudhury and
Pritchard, 2012). In this example, longevity of the
embryos at –20°C was limited to a few months,
while the cryopreserved samples retained high
viability after six to eight years.
37
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Cryopreservation of hardwood trees has become increasingly successful since the introduction
of PVS2 (plant vitriication solution 2), which
contains penetrating and non-penetrating
cryoprotectant solutions. Species for which the
vitriication/one-step freezing protocol using
PVS2 has been successful, with survival rates
higher than 50 percent, include Malus, Pyrus,
Prunus and Populus species (Lambardi and De
Carlo, 2003). Over 90 percent survival rates have
been reported for Cerasus jamasakura (Niino et
al., 1997) and Populus alba (Lambardi, Fabbri
and Caccavale, 2000). Vitriication has proved
successful (71 percent recovery rate) with Betula
pendula, and morphology and RAPD analysis of
regenerated plants in the greenhouse suggests
that the genetic idelity remains unchanged
(Ryynanen and Aronen, 2005). Compared with
results from shoot tips, cryopreservation of
embryogenic calluses and somatic embryos from
hardwood trees has been limited. Success using
the vitriication/one-step freezing protocol has
been achieved with Castanea sativa (Correidoira
et al., 2004) and Quercus suber (Valladares et al.,
2004).
Cryopreservation of embryogenic cultures
of conifers is well advanced; it has been
applied successfully to species in a range of
genera including Abies, Larix, Picea, Pinus and
Pseudotsuga. Over 5 000 genotypes of 14 conifer
species are cryostored in a facility in British
Columbia, Canada (Cyr, 2000). The technique
used is mainly based on slow cooling to –40oC,
which concentrates the intracellular solution
suficiently for its vitriication upon plunging
into liquid nitrogen. Other cryobank collections
of tree species include (Panis and Lambardi,
2005):
• 2 100 accessions of Malus spp. (apple)
(dormant buds) at the National Seed
Storage Laboratory, Fort Collins, United
States of America;
• over 100 accessions of Pyrus spp. (pear)
(shoot-tips) at the National Clonal
Germplasm Repository in Corvallis, Oregon,
United States of America;
38
• over 100 accessions of Ulmus spp. (elm)
(dormant buds) at the Association Forêtcellulose (AFOCEL), France;
• about 50 accessions of Morus spp.
(mulberry) at the National Institute of
Agrobiological Resources, Japan.
In addition, some tropical and subtropical
species are being cryopreserved:
• 80 accessions of Elaeis guineensis (oil
palm) at the Institut de Recherche pour le
Développement (IRD), France (Engelmann,
2004);
• collections of Citrus spp., Artocarpus
heterophyllus (jackfruit), Prunus dulcis
(almond) and Litchi chinensis (lychee) held
at the National Bureau of Plant Genetic
Resources, India (Reed, 2001).
Despite the progress made with cryopreservation, only a limited number of truly
recalcitrant tree species have been successfully
cryopreserved. There are many reasons for this
slow progress. A relatively large number of
species, many of which are wild, have recalcitrant
seeds, and little is known about their biology and
seed storage behaviour. The seeds are dificult to
cryopreserve because they tend to be large and to
have high moisture content when shed. Excised
embryos or embryonic axes can be an option;
however, viable tissue culture protocols needed
to regrow embryos and embryonic axes after
freezing are often lacking or not fully operational.
In addition, provenances, seed lots and successive
harvests often demonstrate signiicant variation
in the moisture content and maturity stage of
seeds and embryos of recalcitrant species, which
makes cryopreservation dificult (Engelmann,
2010). Despite these hurdles, groups throughout
the world are working to improve knowledge of
the mechanisms responsible for seed recalcitrance
and are exploring various technical approaches to
understand and control desiccation sensitivity.
Field gene banks and planted stands
Arguably the most effective way to conserve
long-lived tree species ex situ is by planting them
in speciic ex situ gene conservation stands or
OVERV IE W
ield gene banks, arboreta, clone banks, seed
orchards and operational plantings. Ex situ
ield gene banks maintain sources of variation
of functional traits for direct use in production
through propagation and breeding programmes.
Experience has shown that it can be dificult
to identify the species and provenances or
populations most in need of speciic ex situ
conservation activities. Provenances of Eucalyptus
camaldulensis conserved in FAO-supported ex situ
gene conservation stands at Petford and Lake
Albacutya, Australia, have turned out to be some
of the most widely planted operationally and
therefore have been conserved de facto in such
plantings. Ex situ conservation stands are often
less expensive, more effective and in many ways
more practical for maintaining diversity than
seed storage and in vitro methods, which are
better suited to agricultural crops and short-lived
species with plantings that only last a season.
Other examples of ex situ conservation in planted
ield gene banks, both from South Australia,
include ex situ plantings and grafted trees of
Eucalyptus globulus established by the Southern
Tree Breeding Association at the National Genetic
Resources Centre at Mount Gambier; and the
Currency Creek Arboretum, a largely self-funded
arboretum speciically for eucalypts (genera
Angophora, Corymbia and Eucalyptus) which
has the largest global ex situ collection of living
eucalypt species – over 900 species, subspecies
and varieties and over 8 000 individual plants –
established on a single site.
Morphological and biochemical characterization of conserved accessions provides a wealth
of trait-based knowledge. Such collections are
established for tree species that are considered
important for agroforestry or arboriculture
production and include many fruit-tree species.
Most of these collections are ield gene banks
because in some cases the species are recalcitrant
and because ield gene banks generally facilitate
characterization. For example, an exercise with
national agricultural research institutions in
the Amazon to explore the status of ex situ
conservation, characterization and evaluation
demonstrated a high number of collections
and promising advances in morphological
characterization of prioritized local fruit-tree
species including Bactris gasipaes, Euterpe spp.,
Mauritia flexuosa, Myrciaria dubia, Platonia
insignis, Pouteria caimito and Theobroma
grandiflorum (Scheldeman et al., 2006).
However, ield gene banks are also costly to
maintain (Dawson et al., 2013). Their existence
can only be justiied when gene bank material
can be accessed by users and meets their needs.
The National Genetic Resources Program of
the United States Department of Agriculture,
for example, holds collections of a list of tree
species that are characterized (www.ars-grin.gov/
cgi-bin/npgs/html/croplist.pl). They include many
temperate fruit-tree species, but also tropical
ones such as Bactris gasipaes, Canarium ovatum
and Averrhoa carambola. Data on traits can be
consulted freely online and germplasm can be
requested accordingly.
There is no centralized information system
for tree species collections, however; thus
characterization information, if collected, is
generally not accessible, especially for collections
in developing countries. When germplasm
maintained in characterized collections is not
accessible for potential users, the gene banks lose
their connection with users and their needs.
Thus even though ex situ ield collections
exist for only a limited number of tree species,
a considerable number of these collections
are already morphologically characterized.
Unfortunately, many of these collections are not
known to the general public, and it is dificult to
gain access to existing characterization data. To
enhance the use of gene bank characterization,
such information should be systematized and
made accessible at a central point.
Targeted species-based approach
This conservation approach, typically highly
resource intensive, is used to conserve as much
intraspeciic diversity as reasonably possible for
forest tree species with a high priority rating
(usually given to species with major national
39
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
and/or international economic importance). It
may also be used for endangered tree species;
in these cases the genetic conservation effort is
directed towards maintaining enough diversity,
in preferably more than one population, to
ensure the species’ survival. In the ideal speciesbased conservation plan the distribution of the
species’ intraspeciic diversity and associated
relevant factors will be well known. Populations
for conservation are selected with the aim of
eficiently and securely conserving as much
genetic diversity as possible, including rare alleles
and co-adapted gene complexes of identiied
high value populations or seed sources, in a
network of managed in situ FGR reserves. For
species exhibiting clinal variation, connectivity
and gene low between populations can be
maintained through vegetation corridors and/or
linked by circa situm plantings. In many instances
implementation involves a diverse group of land
managers and interested parties, and in some
cases international collaboration. Ideally, safety
duplication of the material conserved in situ is
also undertaken through ex situ methods such
40
as long-term seed storage banks for species with
orthodox seed storage behaviour or tissue culture
banks and ield gene banks for species with
recalcitrant seed storage behaviour.
Although this approach has major beneits and
has been widely promoted by FAO and forest
geneticists over the past 30 or more years, it has
not been implemented widely. Most examples are
from developed countries in Europe and North
America, for example Picea abies in Finland (Koski,
1996). Only a few cases have been documented
in tropical countries, including Pinus merkusii
in Thailand (Theilade, Graudal and Kjær, 2000)
and Terminalia richii and Manilkara samoensis
in Samoa (Pouli, Alatimu and Thomson, 2002).
Since 2007, the European Information System
on Forest Genetic Resources (EUFGIS), hosted
by Bioversity International, has undertaken
considerable preparatory work for many species
in 36 European countries, including through
creation of a national network of FGR inventories
and development of minimum requirements for
dynamic conservation units of forest trees.
OVERV IE W
Chapter 4
Knowledge and information on
forest genetic resources
Although basic genetic principles are consistent
across plant and animal taxa, forest trees differ
from agricultural crops in signiicant ways, and
the study, management and conservation of
their genetic resources have had to be adapted
accordingly. Thus the knowledge of and focus
on particular genetic technologies also differ
between forest trees and domesticated plants and
animals in some regards. Even the relatively few
forest tree species that are undergoing incipient
domestication typically also exist as large wild,
randomly mating and unstructured populations
(Neale and Kremer, 2011). Many forest tree
species have narrow regional adaptation, so the
number of species planted commercially is much
higher than for food crops (Pautasso, 2009).
Trees are long-lived and their generation times
and juvenile phases are generally long. Many tree
species encompass enormous genetic diversity
(Hamrick et al., 1992); even breeding populations
in improvement programmes represent relatively
large gene pools in comparison to agricultural
crops, and tree genomes may be orders of
magnitude larger than those of cultivated
crops (Neale and Kremer, 2011). Tree species are
predominantly outcrossing, and in fact none
are known to be predominantly seling (Petit
and Hampe, 2006) – a potential evolutionarily
selected pathway of long-lived species against
inbreeding depression (Duminil et al., 2009). With
a few exceptions, tree breeders do not aim to
develop varieties as with agricultural crops, and
tree species that have known or potential value
for forestry number in the tens of thousands.
Thus, forest geneticists and genomicists face
different challenges and require different tools
and techniques than those who work with
agricultural crops.
According to the country reports prepared for
The State of the World’s Forest Genetic Resources,
approximately 2 400 species are actively managed
in forestry. The total count of forest species,
however, remains inconclusive (Box 4.1). Only
approximately 700 tree species are subject
to some level of selection and improvement
globally, and progeny tests have been established
for no more than two-thirds of these species. In
addition, a number of non-planted tree species
have been studied, mainly using molecular
markers. Assuming a total global count of at least
80 000 tree species, little more than 1 percent
of the tree species have been subject to genetic
study, and less than 1 percent have been studied
with the aim of improving resources for human
use. Undoubtedly many not yet studied species
have untapped potential that could be realized
given suficient resources, interest and survival of
suficiently diverse populations.
The tree species that have been most studied
using scientiic approaches (since about 1950)
fall in two categories: those that have been
extensively planted in commercial planted
forests for wood, and species that are valuable
in agriculture for fruit and other livelihood
beneits. Commercial planted forests represent
about 7 percent of the world’s forests but
are responsible for more than 50 percent of
the world’s industrial roundwood production
41
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
(FAO, 2010a). Approximately 30 tree species
from just four genera (Acacia, Eucalyptus,
Pinus and Populus) account for much of the
area planted globally (Carle et al., 2008). Most
of these species have been studied in detail,
including quantitative and molecular and/or
genomic analyses. Some of the most studied
species include: Acacia mangium, Acacia
nilotica, Cunninghamia lanceolata, Eucalyptus
camaldulensis, Eucalyptus globulus, Eucalyptus
grandis, Gmelina arborea, Larix gmelinii,
Larix sibirica, Picea abies, Picea obovata, Picea
sitchensis, Pinus caribaea, Pinus elliottii, Pinus
massoniana, Pinus nigra, Pinus patula, Pinus
pinaster, Pinus radiata, Pinus sibirica, Pinus
sylvestris, Pinus taeda, Populus deltoides, Populus
nigra, Populus tremula, Populus tremuloides,
Populus trichocarpa, Pseudotsuga menzesii and
Tectona grandis.
Tree species producing fruit and other foodrelated crops of global signiicance have a much
longer history of domestication and accumulation
of knowledge. Unlike the tree species that
have been managed for wood production, well
recognized varieties of fruit-tree species have
been developed over centuries to millennia. Yet
unlike most agricultural crops, the original fruittree species still exist as wild populations capable
of sharing genetic material with domesticated
varieties. Some of these populations are under
threat; examples include several globally signiicant
fruit-tree species that originated in Central Asia
such as apple, apricot and other species from the
Rosaceae family (Eastwood et al., 2009).
Tree species are much better characterized
in some regions than others. In Europe, North
America and Australia, at least some genetic
knowledge has been generated for most native
tree species. Species in South and Central
America have received more attention than
those of Asia or Africa, in general, although
there are exceptions at the country level.
Even in countries where signiicant funds and
capacity have been allocated for study of FGR,
the evenness of species coverage is variable,
with key species studied intensively and little or
42
no information for many. At the species level,
approaches vary widely, from single population
studies which are common for tropical species,
to range-wide surveys for temperate species of
broad commercial interest.
What constitutes knowledge of
forest genetic resources?
Since forest genetic resources are deined as
materials of actual or potential economic, environmental, scientiic or societal value (see p. 4), for
the purposes of this overview it is assumed that
any species that has been studied has such value.
Relevant knowledge encompasses quantitative,
molecular and genomic information. In addition
to such genetic information, other types of
information can provide insight into the genetic
variability of a species, such as: knowledge of
species distribution and environmental variability
within the species range; population size and
relative degree of contiguity; and observable
morphological variation (in a natural population).
Traditional or indigenous knowledge predates
published information; it is less well documented
but often valuable.
The concept of FGR is also often used in a
practical sense to designate clones, varieties and
populations (i.e. genetic units) that hold special
interest for conservation or use. Many planted
tree species of high commercial value have been
subject to some degree of selection, testing
and breeding which have generated particular
knowledge. Other highly valuable species are
harvested exclusively in the wild, and very little
may be known about their genetic resources.
Why study forest genetic resources?
Uses of forest resources and FGR, management
practices and priorities for research vary by
type of tree species, region and socio-economic
situation of the users. Much scientiic genetic
research on trees has been devoted to species
having commercial value for timber, and most
of these are temperate species. About 30 species
have been studied intensively, tested and bred
for increased wood production, improved quality
OVERV IE W
Box 4.1
Filling the knowledge gap in botany: how many tree species are there on Earth?
Although knowledge of species and their conservation
status has been improving over time (Figure 4.1), it is
still insufficient, and insufficiently accurate, to provide
adequate support to conservation and sustainable
management of forest genetic resources at the global
level. Estimates of the number of plant species vary
widely from 25 000 to more than 400 000 (Stebbins,
1974; Bramwell, 2002; Miller et al., 2013), and
perhaps more than a quarter of all flowering plants
have not yet been named or discovered (Miller et al.,
2013).
Major challenges to filling the gap in knowledge
on plant species include frequent synonymy,
the difficulty of discriminating certain species
by morphology alone, and the fact that many
undiscovered species are small in size, difficult to
find, or have a small geographic range (Scheffers
et al., 2012). A recent, major effort is the Global
Taxonomy Initiative, which enables the international
community to acknowledge the existence of a
“taxonomic impediment” to the sound management
of biodiversity and to initiate a programme with the
objective of removing or reducing the knowledge
gaps in the taxonomic system.
In response to the CBD’s Global Strategy for the
Conservation of Biodiversity, adopted in 2002, the
Royal Botanic Gardens, Kew, United Kingdom, and
the Missouri Botanical Garden, United States of
America, developed The Plant List (www.theplantlist.
org), intended as a widely accessible working list of
known plant species. It aims to be comprehensive in
coverage at species level for all mosses and liverworts
and their allies (bryophytes) and vascular plants
which include the flowering plants (angiosperms),
conifers, cycads and their allies (gymnosperms) and
the ferns and their allies including horsetails and club
mosses (pteridophytes). The last update (September
2013) includes names of 1 064 035 species in 642
plant families and 17 020 plant genera. Of the total,
350 699 are accepted species names, 470 624 are
synonyms and 242 712 are unresolved (The Plant
List, 2013).
The status of botanical knowledge of plant species
varies from country to country. Few countries have
a relatively accurate estimate of the number of their
vascular plants. Some have completed flora lists, but
very few have a detailed plant species checklist that
includes species characteristics and life forms, which
could make it possible to distinguish forest plants,
e.g. trees, shrubs, palms and bamboos.
Moreover, the answer to the question “how
many trees species are there on Earth?” remains
very rough, varying from 50 000 (National Academy
of Sciences, 1991) to between 80 000 and 100 000
species (Oldfield, Lusty and MacKinven, 1998; Turok
and Geburek, 2000). These estimates are even more
confusing in light of the different definitions of a
tree. This report on The State of the World’s Forest
Genetic Resources includes plant species identified
by countries as part of their forest resources. In the
country reports, FGR are often taken to include trees,
shrubs, palms, bamboos, lianas, cycads and ferns, and
in a few cases some herbaceous plants..
FIGURE 4.1
Proportion of the world’s plants in accessible
plant lists
%
100
90
80
70
60
50
40
30
20
10
0
2002
2007
2010
Source: CBD Secretariat (2009).
43
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
(Neale and Kremer, 2011) and/or resistance to
pests and diseases (Yanchuk and Allard, 2009).
A body of knowledge has also accumulated over
a longer time and in a much more fragmented
way on tree species that are important for nonwood forest products including fruit- and nutbearing trees, species with valuable medicinal
properties, oil- or latex-producing trees, and
species having shade or ornamental value.
Traditional knowledge of phenotypic variation
extends from informally recorded traditional
knowledge to recent scientiic studies. Few
NWFP species have been thoroughly studied
using modern techniques, with the exception
of those having high commercial value; in
those cases, the objectives and the techniques
employed resemble those of genetic research on
agricultural crops.
Conservation
During the last three decades, since the
development and broad use of molecular markers,
many studies have been conducted applying
these tools to inform conservation strategies and
approaches. Neutral genetic markers are used, for
example, to deduce population-level parameters
that are informative about spatial patterns of
genetic diversity, reproductive biology (mating
system, pollen- and seed-mediated gene low),
species evolutionary history and demography
(for example, existence of genetic bottlenecks,
localization of refugia sites, founder populations).
Aims of conservation genetic studies are to
understand levels and patterns of genetic diversity,
impacts of land use changes on intraspeciic
variation, and vulnerability of populations to
threats; and to identify populations having high
conservation priority and design approaches for
their conservation. A combination of neutral
molecular markers and either phenotypic
measurements or genomic markers for adaptive
traits is ideal for deining priority populations,
but in most cases, especially in tropical countries,
resource limitations have resulted in the sole use
of neutral markers.
44
Most tropical timber species are managed in
semi-natural forests – in many cases selectively
harvested – and depend on natural regeneration
for their renewal. In some cases the genetic
resources of such species may be degraded through
dysgenic selection; this can occur when the bestquality trees are cut and only badly shaped trees
are left as contributors to the next generation.
Thus in the absence of an improvement strategy
and planting, selective harvesting could be
expected to have dramatic effects on the resource
sustainability. However, few conclusive studies
have evaluated this hypothesis, and one of the
few that has been published found little evidence
(Cornelius et al., 2005).
In general, little is known about the
sustainability of the genetic resources of
selectively harvested tree species, especially those
in the tropics (Wernsdorfer et al., 2011). Effective
long-term management of these species requires
knowledge of population genetic parameters
such as gene low dynamics and the structure of
genetic variation in economically and adaptively
important traits. Such knowledge is important to
ensure that viable populations are maintained in
harvested areas and that harvest does not have a
dysgenic impact on seed trees.
Genetic markers that can be used effectively
for identifying species and origin of timber are
becoming important as well, to monitor legality
of timber harvest. The Barcode of Life project
attempts to use DNA markers to identify species
(see Chapter 9).
Conservation of evolutionary potential is
important for the sustainable management
of forests, particularly for adapting forests to
environmental change (Lefèvre et al., 2013), as
well as for improvement of valuable traits. This
requires a good understanding of the extent
and patterns of genetic diversity throughout
a species’ range. For many species, studies
have been conducted on limited numbers of
populations, using various molecular markers to
elucidate patterns of diversity. Studies are often
limited by small sample sizes and the applicability
OVERV IE W
of results is thus limited. The use of different
markers for subsequent studies in divergent
geographic locations of the same species leads to
disparate results that may even be contradictory,
with little potential for generating a common
understanding.
In many countries, little concrete action has
been taken on the basis of most conservation
genetic studies, in part because of a gap between
the science and the application of conservation
knowledge (Knight et al., 2008). The data are still
largely insuficient to allow testing of whether
congruent patterns of spatial genetic diversity exist
among species (i.e. zones of genetic endemism
and richness; see e.g. Conord, Gurevich and Fady,
2012 for the Mediterranean). This knowledge
would have important implications in terms of
landscape management and conservation, as
forest reserves are ideally localized on the basis
of genetic information across all species.
Tree improvement
Wood production. Improvement programmes
for wood production have generated knowledge
about productivity and quality traits for most
tree species that are used extensively in planted
forests. The earliest genetic studies on forest trees
were designed to quantify variation in traits of
commercial value for selection and breeding.
Most of the effort has focused on increasing
wood volume, both for timber and for pulp
and paper, and on improving the form of trees
for timber production. Many studies have been
reported on genetic parameters associated with
wood quality, including strength, density and
more recently lignose/cellulose ratio, although
these traits have generally been considered
to be of secondary importance. Breeding for
resistance or tolerance to biotic and abiotic
factors tends to be restricted to more specialized
research programmes (Yanchuk and Allard,
2009). Hence much information is available for
most commercial planted forest species on the
complicated growth rate related traits associated
with quantity and quality of wood production,
but relatively little genetic information has been
generated about other traits. This one-sidedness
may have negative consequences in efforts to
adapt to changing climatic conditions.
Agricultural purposes. Fruit-trees and other tree
species that are important for food or fodder
or that have cultural or religious signiicance
have been domesticated for millennia, without
the beneit of scientiic genetic knowledge
for most of that time. The most widely used
method for capturing improvements has been by
cloning trees having desirable properties. Unlike
commercial timber species, domesticated varieties
of trees important for food and other non-wood
amenities have been developed for centuries,
simultaneously, in many geographic locations.
Cross-continental exchange of planting material
started early in human history and has intensiied
since colonial times. For example, in the
Americas, intercultural contacts with European
settlers some ive centuries ago led to an early
but systematically underestimated (intentional
and accidental) loristic homogenization. Useful
plants from the Old World were actively and
passively distributed over the New World tropics
and subtropics as a consequence of colonial
horticultural endeavours (Bennett and Prance,
2000). By the mid-nineteenth century, exotic fruittrees were fully incorporated into home gardens
along the Amazon (Miller and Nair, 2006).
Taxonomy, phylogeny and
phylogeography
Patterns of genetic differentiation and speciation
have been studied in order to understand the
evolutionary history of species. Petit and Hampe
(2006) reviewed the evolutionary consequences
of “being a tree”, noting that the high diversity
and very high fecundity of most tree species
allow for rapid evolution at the micro scale, but
that long generation time, large size and other
characteristics result in slow macroevolution.
Taxonomic studies rely increasingly on genetic
markers (see Chapter 8) to complement or
45
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 1
replace conventional morphological methods
to determine taxonomic status and understand
phylogenetic relationships. Many tree genera
are incompletely described at the species level, in
part because of hybridization and introgression
among species; an example is the genus Quercus
in Mexico, which has between 135 and 150 species
but the exact number is not known (GonzálezRodríguez et al., 2004). A combination of nuclear
markers and morphological traits is employed to
differentiate species.
Chloroplast DNA (cpDNA) is the genome of
choice for phylogenetic and phylogeographic
studies because of the small size of the genome and
uniparental inheritance, which make it possible
to interpret the spatial pattern of haplotypes as
an estimate of past gene low. Chloroplast DNA
is also easily sequenced and assessed for length
variation using restriction enzymes. In addition,
cpDNA shows neutral differentiation among
divergent populations sooner than nuclear alleles.
If population divergence has occurred relatively
recently, neutral cpDNA variation will be more
likely to show the differentiation than nuclear
polymorphisms (Hamilton et al., 2003). Hamilton
et al. (2003) examined patterns of cpDNA
haplotype variation, searching in particular for
evidence of selection acting on several insertion/
deletion regions of the chloroplast genome
of eight species of Lecythidaceae, which is the
Brazil nut family. They found that the rate of
evolution was highly variable among regions in
the genome, but that the variability seemed to
be related to lineage rather than region. They
concluded that the insertion/deletion markers and
nucleotide variation in the chloroplast genome
were selectively neutral and thus should provide
unbiased estimates of population parameters.
Availability of information on
genetic resources
The availability of, and access to, quality and
up-to-date information on FGR is reported to be
poor in many countries. For example, knowledge
on species distribution remains inaccurate in
the tropics, and distribution maps have been
46
developed for only a small proportion of species.
In contrast, more is known for temperate species.
In Europe, for instance, the European Forest
Genetic Resources Programme (EUFORGEN) has
developed 34 species distribution maps (Figure
4.2); they include population-level information
which is essential for monitoring the dynamics of
the species’ genetic resources.
Most country reports highlight the need to
promote awareness among decision-makers and
the general public of the importance of FGR
and their roles in meeting present and future
development needs. Lack of information limits
the capacity of countries and the international
community to integrate FGR management into
cross-cutting policies. In spite of the efforts of
plant taxonomists and geneticists to characterize
and describe forest plant species and species
populations, many key questions still need to be
answered.
OVERV IE W
FIGURE 4.2
Example of a species distribution map: Pinus sylvestris in Europe
30°W
20°W
10°W
0°
10°E
20°E
30°E
40°E
50°E
60°E
70°E
50°N
40°N
30°N
0
250
500
Km
1,000
Source: EUFORGEN, 2009 (by permission).
47
Part 2
DRIVERS OF CHANGE
AND TRENDS
AFFECTING FOREST
GENETIC RESOURCES
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
Chapter 5
Drivers of change
The main drivers of change causing unprecedented
threats to FGR in recent times are almost exclusively
of human origin. The world’s current population
of 7.2 billion is projected to reach 9.6 billion by
2050. Whereas population in developed countries
is expected to remain largely constant, growth is
expected to be particularly dramatic in the world’s
least developed countries, which are projected to
double in size from 898 million inhabitants in 2013
to 1.8 billion in 2050 and to 2.9 billion in 2100 (UN,
2013). Along with population growth, production
of energy, food and many other commodities is
predicted to increase, with some negative impact
on natural resources including forests and forest
genetic resources.
Many major drivers of change in forestry are
external to the forestry sector. Demographic,
economic, technological and climate changes all
shape forest development. With the exception
of geological events, the categories of threat
to species identiied by IUCN are residential
and commercial development; agriculture and
aquaculture; energy production and mining;
transportation and service corridors; biological
resource use; human intrusions and disturbance;
natural system modiications; invasive and other
problematic species; pollution; geological events;
and climate change and severe weather events.
Major modern-day human impacts on
the environment involve massive changes in
land-use systems (e.g. conversion of forest to
agriculture and other land uses), destruction
and fragmentation of natural habitats, air and
soil pollution, salinization and soil acidiication,
climate change, overexploitation of biological
resources, homogenization of biota and
biodiversity loss. These impacts interact in complex
ways and may result in non-additive cumulative
effects (Yachi and Loreau, 1999). Threats to FGR
that have increased greatly in recent times and
could continue to increase in the future include
forest cover reduction and degradation, climate
change, forest ecosystem modiication, spread of
invasive and ecosystem-transforming species and
interactions of different threat factors.
Forest conversion and expansion
of crop land
The main needs of the world’s growing human
population that have impact on FGR in native
forests are additional land for agriculture and
agroforestry production, infrastructure, human
settlements, mining, and planted forests for wood,
paper and fuel. For example, the demand for
wood products for both industrial and domestic
uses is expected to increase by 40 percent in the
next 20 years (FAO, 2010a).
The growing global demand for land for the
production of agricultural commodities has
resulted in sometimes irreversible changes to
the world’s forest cover. The conversion of forest
lands to agriculture is one of the major threats to
forest genetic resources. Expansion of small-scale
permanent agriculture accounts for 60 percent
of forest conversion in Africa, while large-scale
permanent agriculture involving commodities
such as palm oil and soybean represents the major
cause of forest conversion in Latin America and
Asia (FAO, 2001c). Over the decade 2000−2010,
permanent crop area has increased by 15.7
51
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
percent globally and by 22 percent in the least
developed countries (Figure 5.1). Together
croplands and pastures have become one of the
largest terrestrial biomes on the planet, rivalling
forest cover in extent and occupying about 40
percent of the land surface (World Bank, 2008).
Important drivers of forest ecosystem
degradation also include large-scale plantations
for timber or paper pulp or for oil-palm (Foley et
al., 2005; Kongsager and Reenberg, 2012) which
have replaced many natural forests and cover
190 million hectares worldwide.
Demand for energy
Bioenergy, including fuelwood and charcoal,
currently represents a major portion of the
domestic energy consumption in many developing
countries and is one of the main causes of forest
degradation in these countries, particularly in the
dryland or semi-arid countries in Africa. Fuelwood
represents 90 percent of total wood removals in
Africa, compared with 47 percent in the world
overall (FAO, 2010a). Driven by population growth
and the subsequent demand for more domestic
energy supply, global fuelwood consumption has
increased by 4 percent from 2000 to 2010.
Bioenergy is expected to become an important
component of future renewable energy systems,
and policies are being developed to facilitate
this process. The European Union (EU) currently
imports almost half the energy resources it
consumes, including liquid biofuels and wood
pellets. Already in 2007, many European countries,
such as Belgium, Finland, the Netherlands, Sweden
and the United Kingdom, imported significant
amounts of biomass (between 12 and 43 percent
FIGURE 5.1
Changes in area of cropland, 2000−2010
%
0.30
0.25
0.20
0.15
0.10
0.05
0.0
–0.05
–0.10
–0.15
–0.20
Africa
Source: Data from FAO, 2013a.
52
Asia
Europe
Latin America
and the
Caribbean
North
America
Least
developed
countries
World
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
of their total used for energy purposes). The
EU is now promoting energy policies that have
potentially far-reaching implications for forests
and forest use.
Unsustainable harvesting and use
Many of the country reports provided for this
report detail overexploitation and unsustainable
harvesting threats to FGR. Overharvesting by
itself rarely leads to extinction, but it can seriously
erode genetic diversity, and recovery can be
very slow for species that occur naturally at low
frequency. For narrowly distributed and naturally
rare species, overharvesting can directly lead
to or threaten extinction. Thuja sutchuenensis,
for example, a critically endangered, narrowly
distributed endemic tree in Chongqing
Municipality, China, was driven to the brink of
extinction from overharvesting for its precious
scented wood; in 1999 it was rediscovered
and accorded protection. Chile has identified
overexploitation, along with land use change and
deforestation, as a major threat for 22 priority
tree species (Hechenleitner et al., 2005).
Some
countries
report
selective
overharvesting, much of it illegal, as a major
and increasing problem intertwined with rural
poverty, threatening extinction of the highestvalue species in the forests – including highvalue timbers such as Dalbergia cochinchinensis
in Southeast Asia, Dalbergia melanoxylon in
sub-Saharan Africa and Pterocarpus santalinus
in India; and valuable NWFP species such as
Cryptocarya massoia (whose bark provides
massoia lactones for the food industry) in
New Guinea, Prunus africana (whose bark
is used in treatment of benign prostate
hypertrophy), certain Santalum and Osyris
species
(sandalwoods,
whose
heartwood
provides essential oils) in India, Indonesia, Timor
Leste and the Pacific Islands, and Taxus contorta
(producing taxol, a chemotherapy drug to treat
cancer) in Afghanistan, India and Nepal.
Overharvesting usually concerns highly
valuable species such as ebonies, sandalwoods,
agarwoods and frankincense, but in areas
with high population pressure and poverty,
overharvesting may be associated with lowervalue products such as fuelwood and charcoal.
Even an activity as seemingly innocuous as
harvesting for Christmas trees may threaten
FGR; in Guatemala, for example, uncontrolled
cutting of Abies guatemalensis branches for use
as Christmas trees is reducing the regenerative
capacity of the species, which has now
disappeared from some areas (López, 1999). In
Tonga, harvesting of Santalum yasi saplings for
Christmas trees is limiting recruitment and is one
of the major threats to the species (Tuisese et al.,
2000).
Harvesting of wood resources for fuelwood
and charcoal is often less discriminatory but
can lead to permanent loss of tree species
in locally adapted populations, reducing
options for future natural or human-mediated
recovery from the associated environmental
degradation. Somalia, for example, reports
deforestation and elimination of ecologically
and economically important tree species such as
Acacia spp. as a result of charcoal production for
income generation (including though export);
Boswellia spp. have also been overexploited
for frankincense. In northeastern Thailand,
high rates of drug addiction in some villages
have resulted in increased charcoal production
and unsustainable resin harvesting from
dipterocarps and pines in adjacent forests to pay
for illicit drugs, threatening the efforts of the
Thai Forestry Department to conserve unique
lowland populations of Pinus merkusii.
One remedy to overharvesting can be the
greater involvement of indigenous and local
communities in management of forests and FGR,
especially if this is backed by appropriate technical
support,
including
improved
silvicultural
practices to ensure sustainable production of
desired products and regeneration of preferred
species. Other strategies include better legislative
protection, including compliance monitoring,
and development of alternative sources of wood
and NWFPs such as highly productive plantation
systems and improved agroforestry systems.
53
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
Livestock and browse animals
In some regions, grazing by the growing
livestock population is a serious threat to
many woody forage species and can have a
destructive impact on the genetic resources. In
Latin America, for example, rapid expansion
of cattle ranches accounts for a large portion
of forest loss (FAO, 2007a). In the Brazilian
Amazon region, ranches cover an area of at
least 8.4 million hectares in total (UNEP, 2009).
In the Sahelian and other semi-arid countries
of West Africa, where the cattle and small
ruminant population is estimated to be as
high as 60 million and 160 million respectively
(SWAC-OECD/ECOWAS, 2008), forest reserves
are the main source of fodder for cattle grazing
freely during the dry season when animal feed
becomes scarce. Furthermore, herders who
practice pollarding of fodder trees to feed
their animals can put particular tree species in
danger. Heavy grazing in forest land can cause a
shift from perennial to annual vegetation type;
studies have shown up to 50 percent reduction
of large woody shrub cover in South Africa from
grazing (O’Connor et al., 2011). Other studies
(e.g. Olff and Ritchie, 1998), however, mostly on
grassland conditions, suggest that grazing can
sometimes result in increased species richness.
Excluding grazing animals from ecosystems that
evolved with grazing has been documented
to decrease biodiversity through competitive
exclusion of certain plant species (Nuzum, 2005).
Browsing animals, especially introduced
goats, have wrought havoc on tree vegetation
in many parts of the globe, especially on island
communities. Pinus radiata is among the most
important plantation forestry trees species in
the world, but the unique island population on
Guadalupe Island, Mexico, was until recently
highly threatened with surviving trees being
very old and browsing by goats removing any
regeneration (Spencer, Eldridge and Matheson,
1999). The goats have now been taken away from
the island and the population is being monitored.
While the Guadalupe provenance is secured
through ex situ conservation efforts, a loss of the
54
tree in its natural habitats would have excluded
continued evolution and adaptation in the
environment that has resulted in highly drought
tolerant germplasm.
By 1945 goat predation on Three Kings Island
in northern New Zealand had reduced the
entire population of Pennantia baylisiana to
one individual female tree incapable of sexually
reproducing itself. In 1985 researchers treated
latent pollen with hormones, inducing some
seed, including a self-fertile individual; the
future of this species has now been secured. In
French Polynesia (France), rats have prevented
the natural regeneration of Santalum insulare by
eating more than 99 percent of fruits before their
ripening (Meyer and Butaud, 2009).
Climate change
Since the start of the Industrial Revolution in
Europe in the eighteenth century, atmospheric
pollution has caused damage to forests. However,
its importance as a direct threat to FGR is
diminishing, with most damage likely to result
from stressed trees being more susceptible to
insect pests and diseases. Of greater global concern
for FGR are increasing levels of atmospheric CO2
resulting from human activities such as burning
of fossil fuels and forest destruction over the
past half-century. Deforestation and forest
degradation, with the causes described above,
account for nearly 20 percent of greenhouse gas
emissions.
Elevated levels of CO2 are already contributing
to more extreme climatic events, as predicted
by the Intergovernmental Panel on Climate
Change (IPCC, 2013). Climate alterations and
increased occurrence of extreme climatic events
are mentioned as a threat to FGR in many of the
country reports prepared for this publication.
Many Sahelian zone countries in West Africa, for
example, cited prolonged drought as a threat.
High mortality due to extreme climatic events, in
combination with regeneration failure, can result
in local population extinction and the loss of FGR,
particularly at the receding edge of a species’
distribution.
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
Climate change could alter the frequency and
intensity of forest disturbances such as insect
outbreaks, invasive species, wildfires, and storms.
A greater incidence of intense cyclones, extreme
drought, fires, flooding and landslides has been
observed in tropical forest ecosystems which
have experienced increased temperatures and
more frequent and extreme El Niño–Southern
Oscillation (ENSO) events. Some climate change
models predict substantial dieback in parts of the
Amazon and other moist tropical forests, and the
resulting loss of carbon sinks and storage would
exacerbate global warming (Bernier and Schoene,
2009).
Predictions regarding the impact of climate
change on FGR in natural forests, in planted
forests and on farms vary. Although some authors
(e.g. Hamrick, 2004) consider that many trees
have sufficient phenotypic plasticity and genetic
diversity at the population level to withstand
the negative effects of climate change, others
predict severe impacts (e.g. Mátyás, Vendramin
and Fady, 2009; Rehfeldt et al., 2001). Different
positions relate partly to the types of species
and environments being considered. The more
pessimistic authors often base their views
on tropical trees (Dawson et al., 2011) or on
marginal populations of temperate species
(Mátyás, Vendramin and Fady, 2009), while
the more optimistic authors are often those
discussing temperate and boreal taxa (Lindner
et al., 2010). Current and future climate change
impacts on forests are likely to vary from abrupt
negative impacts to more subtle negative and
positive impacts in some regions or at particular
sites, often only for certain tree species. Many
countries urgently require assistance to cope and
deal with impacts of climate change on FGR and
to promote and use FGR to help with climate
change adaptation and mitigation.
Climate impact on species and
ecosystems
Temperature and precipitation are the two
main climate drivers for forest ecosystems; any
significant changes in either of these will have an
impact on species composition and forest cover.
Impacts can range from extreme disturbances
such as forest fires or pest outbreaks to effects on
physiological processes from more subtle changes
in temperature. The ability of a tree species to
survive the current rapid climate changes will
depend on its capacity to adapt quickly to new
conditions at existing sites, to survive changing
conditions through a high degree of phenotypic
plasticity without any genetic change, and/or
to migrate to an environment with the desired
conditions for that species.
Some forest types are more vulnerable than
others to climate change. For example, in tropical
forests, small changes in climate are likely to
affect the timing and intensity of flowering
and seeding events, which would in turn have
negative impact on forest biodiversity and
ecosystem services. Increased frequency and
intensity of extreme events, such as cyclones,
may result in shifts in species composition.
Mangrove ecosystems are especially vulnerable,
with projected sea-level rises posing the greatest
threat. Mangroves could potentially move inland
to cope with sea-level rise, but such expansion can
be blocked either by infrastructure or by the lack
of necessary sediment, such as in the reef-based
island archipelagos in Melanesia. Temperature
stress will also affect the photosynthetic and
growth rates of mangroves (McLeod and Salm,
2006). Climate change impacts are expected to
be severe in dry, high-temperature regions where
trees are at their adaptive limit (e.g. Lindner et
al., 2010 for Europe) and in confined islands of
moist forest that are surrounded by drier land
(e.g. moist forests in Australia [Williams, Bolitho
and Fox, 2003]).
Forest cover will alter under climate change. The
range of some species will expand, whereas that
of others will diminish. Changing temperature
and precipitation regimes will also cause shifts
in forest types. For example, boreal forests
are expected to shift towards the poles, with
grassland moving into areas formerly occupied by
boreal species. There is evidence of the migration
of keystone ecosystems at the upland and lowland
55
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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treeline of mountainous regions across southern
Siberia, Russian Federation (Soja et al., 2007). For
temperate forests, range reduction is expected to
be more rapid at low elevation and low latitude.
At high elevation and high latitude, temperate
forest species ranges are expected to expand
more than those of boreal forests; as a result the
total area of boreal forests will decline. Thuiller
et al. (2006) have shown that at low latitudes in
Europe climate change will have a greater impact
on species richness and functional diversity than
at high latitudes.
In the subtropical forests of Asia, where
key biodiversity hotspots are found, endemic
species are predicted to decline, with changes
in ecosystem structure and function as a result
(FAO, 2010b). Changes in precipitation may be
more critical than temperature changes for these
species and systems (Dawson et al., 2011).
Changes in water availability are a major
emerging threat to FGR; they will be a key factor
for the survival and growth of forest species. The
response to prolonged droughts will vary among
tree species and also among different varieties of
the same species (Lucier et al., 2009). In arid and
semi-arid lands, increased duration and severity
of drought has increased tree mortality and
resulted in degradation and reduced distribution
of forest ecosystems, including pinyon pinejuniper woodlands in the southwestern United
States of America (Shaw, Steed and DeBlander,
2005) and Cedrus atlantica forests in Algeria and
Morocco (Bernier and Schoene, 2009). Indirect
impacts must also be considered. For example, in
Africa, where drought is limiting the output from
adjoining agricultural land, many communities
with limited economic alternatives are likely to
use forests for crop cultivation, grazing and illicit
harvesting of wood and other forest products,
aggravating the local loss of forest cover (Bernier
and Schoene, 2009).
Choat et al. (2012) found that of 226 forest tree
species from 81 sites worldwide, 70 percent have
narrow safety margins in the event of injurious
levels of drought stress and therefore could
face long-term reductions in productivity and
56
survival if temperature and aridity increase as
predicted. While gymnosperms were found to be
more tolerant of reduced hydraulic conductivity
than angiosperms, safety margins were seen
to be largely independent of mean annual
precipitation, with all forest biomes equally
vulnerable to hydraulic failure and droughtinduced forest decline. These findings help to
explain why drought and increased heat are
resulting in forest dieback across a broad range
of forest and woodland types around the world
(Allen, 2009). These dieback problems have
occurred at a time when increases in temperature
have been relatively modest, which does not
bode well for forests given future temperature
predictions. Under a scenario of a 4oC increase
in global temperature, greater mortality rates
can be expected as well as significant long-term
regional drying in some areas.
Changed hydrological conditions associated
with climate change also include increases in
severity and duration of flooding, which can kill
whole stands of trees. Even inundation-tolerant
species, such as Eucalyptus camaldulensis and
Cocos nucifera, are killed by waterlogging if
the trees have not been regularly exposed to
waterlogging and inundation through their
development. Inundation due to sea-level rise is
beginning to kill vegetation in coastal areas (see
Box 5.1).
In temperate and boreal regions, reduced snow
cover, changed timing of snowmelt and shorter
frost periods are contributing to forest changes
and stresses. Reduced snow cover has been shown
to be responsible for the decline of Xanthocyparis
nootkatensis, a culturally and economically
important tree in southeastern Alaska, United
States of America, and adjacent areas of British
Columbia, Canada. Snow normally protects the
vulnerable shallow roots from freezing damage.
The decline is affecting about 60 to 70 percent of
the 240 000 ha of X. nootkatensis. Coastal Alaska
is expected to experience persistent periodic cold
weather events but less snow in the future, which
may support the spread of dieback (USDA Forest
Service, Pacific Northwest Research Station, 2012).
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
Sensitivity to spring temperatures will affect
fecundity (Clark et al., 2011). In central Spain a
decline in cone production in Pinus pinea over
the past 40 years has been linked to warming,
in particular the hotter summers (Mutke,
Gordo and Gil, 2005). In angiosperms, changes
in the climate could have an impact on seed
production; asynchronous timing between
flower development and the availability of
pollinators could result in low seed production
for outbreeding species that depend on animal
vectors. Pollinators worldwide are being affected
by climate change, and this will likely have a major
detrimental impact on breeding systems and seed
production, with consequences for forest health
and regeneration.
A changing climate also provides the
opportunity for some plant species more
Box 5.1
Selecting for salt tolerance: one way
to address impacts of sea-level rise
on coastal forests
In Kiribati, a single king tide can kill established
Artocarpus altilis (breadfruit) trees. As these trees
harbour seabirds such as terns which are used by
local fisherman to locate schools of fish, their loss has
a major impact on food security and livelihoods.
Given the impacts of sea-level rise in Kiribati,
Tuvalu, and other atoll island nations in Oceania,
development of salt-tolerant breadfruit is an urgent
task. Studies with salt-tolerant non-halophyte trees
(Thomson, Morris and Halloran, 1987; references in
Marcar et al., 1999) have frequently demonstrated
considerable genetically based resistance to salinity.
Given the substantial genetic diversity in breadfruit,
including putative salt tolerance in particular varieties
and natural hybrids between A. altilis and Artocarpus
mariannensis (Morton, 1987; Ragone, 1997), it is
almost certain that salt-tolerant breadfruit can be
selected and further developed – illustrating the
need to conserve and make use of genetic diversity in
multipurpose tree species.
suited to a wide range of climate conditions
to invade new areas (Dukes, 2003). The spread
of Leucaena spp. and Eupatorium spp., for
example, is already known to have had adverse
impacts on biodiversity in subtropical forests in
South Asia. In addition to new species invasions,
changing climates will result in altered patterns
of gene flow and the hybridization of species
and populations. Shifting ecological niches will
increase the risk of invasion by more competitive
tree species that are more precocious or can move
more quickly than the present species. Invasions
of new genes via pollen and seed dispersal may
disrupt local evolutionary processes, but could
also be a welcome source of new adaptive traits
(Hoffmann and Sgro, 2011).
Climate impact on insect pests and
diseases
Changes in temperature and water availability will
also influence the incidence and spread of pests
and diseases. For example, unusually warm winters
– i.e. the absence of consistently low temperatures
over a long period – supported the spread of
mountain pine beetle, Dendroctonus ponderosae,
from an existing outbreak to an attack on some
14 million hectares of montane and boreal forests
in North America (see section on insect pests
below). Spreading into new areas and attacking
pine trees with no resistance, the beetle threatened
the genetic diversity of forest populations.
Trees already weakened by climatic stresses
are more vulnerable to destruction by insect
attack (e.g. bark beetles) (McDowell et al., 2008).
In Finland, the spread of a virulent fungus,
Heterobasidion parviporum, favoured by longer
harvesting periods, increased storm damage and
longer spore production season, is expected to
increase root and bud rots in coniferous forests
(Burton et al., 2010).
A thorough analysis of historical records
and adequate knowledge of insect population
dynamics is needed before outbreak frequencies
can be linked to climate change. The availability
of such information has enabled researchers
to link drought stress due to climate change to
57
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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the extensive damage caused by insects to Pinus
edulis in the southwestern United States of
America (Trotter, Cobb and Whitham, 2008)
Climate change may be facilitating the global
spread of harmful forest pests by allowing
species moved through international trade to
find hospitable habitats (Régnière and SaintAmant, 2008). Significant evidence regarding
insect distributions is accumulating, although the
complexity of insect responses to climate factors
makes predictions difficult (see Box 5.2).
Useful insects such as pollinators, biological
control agents and other plant-associated
organisms can be affected by climate change as
well.
Knowledge of forest genetic diversity, including
pest resistance, will therefore be increasingly
important in forest management.
Changed ire regimes
Forest fires can be a great threat to biodiversity,
and climate change may alter their frequency and
intensity. In Siberia (Russian Federation), Alaska
(United States of America) and Canada, extreme
fire years have become more frequent (Soja et
al., 2007). In recent years, wildfires consumed
more than 2.5 million hectares of forest in Alaska,
assisted by warm temperatures and drought
conditions during early summer (CCSP, 2008). In
2006, fires engulfed more than 4 000 ha in New
Caledonia (France), destroying rare fauna in the
archipelago’s unique tropical forest ecosystems.
Other countries reporting forest fires as a threat
to FGR included Algeria, Burundi and Ethiopia.
Fire may cause variable effects depending on
the fire’s intensity and spacial extent. Increased
fire frequency could result in the erosion or
even elimination of fire-sensitive species from
woodlands and forests. In regions that have not
regularly experienced wildfires in the past, fire
may become the main driver of change, with
a rapid transition from fire-sensitive to fireresistant species.
Severe fire may have the same effect as clearing
a forest, especially where fire creates large patchy
openings. The pattern and size of such openings
in relation to the forest cover influence genetic
diversity. Where mortality among burnt species is
heavy, it results in reduced population sizes and
increased genetic drift. For isolated populations,
the migration rates of seed and pollen exchange
are therefore affected. Sources of migration could
Box 5.2
Predicting impacts of climate change on distribution of forest insect pests
Modelling tools such as BioSIM (Régnière and
Saint-Amant, 2008) have been designed to predict
the geographic range and performance of insects
based on their responses to key climate factors.
The basis of BioSIM is the ability of the insect to
complete its life cycle under a specific climate with
all requirements to sustain that cycle available. The
model predicts distributions by mapping climates
that provide viable seasonality and overlaying the
distribution of resources essential for (or most at
risk from) a particular species. Further refinements
can be achieved by also considering the survival
of that species under extreme climatic conditions.
This approach has been applied to three species
58
of importance to North American forests within a
climate change scenario of a 1 percent rise per year
in atmospheric CO2. One of these species, Lymantria
dispar (gypsy moth), is prevalent in the United States
of America and some parts of Canada; however, its
northern limit in Canada is set by adverse climatic
conditions. The model established for this species
shows that it will be a considerable threat to
hardwood forest resources as climate change allows
for its expansion further north and west in Canada.
It has been estimated that the proportion of forest
at risk from this pest will grow from the current 15
percent to more than 75 percent by 2050 (Logan,
Régnière and Powell, 2003).
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
even be cut off, thus reducing the effectiveness of
pollinators (Kigomo, 2001).
Adverse fire may directly affect biotic dispersal
agents, and this may decrease migration of genes
between populations. Migration may increase if
the migration vectors are abiotic. A devastating
fire may affect traits that could have a direct
bearing on fire-resistant species, resulting in
direct selection that indiscriminately removes all
such genotypes (FAO, 2010a).
The cumulative impact of interacting
disturbances can increase fire risk. For example,
drought often reduces tree vigour, increasing
vulnerability to insect infestations and diseases.
Insect infestations and diseases add to the fuel
available and therefore increase the opportunity
for forest fires, which in turn can support future
infestations by weakening tree defence systems
(Dale et al., 2001).
Invasive species
Invasive species, including plants, insect pests
and microbial pathogens, are increasingly
being identified and noted as major threats
to ecosystem integrity and individual species,
including trees. The United States of America,
for example, reports that 46 percent of all
federally listed threatened and endangered
species are considered at risk primarily because
of competition with or predation by invasive
species, sometimes in interaction with other
threat factors. The spread and impacts of invasive
species are frequently exacerbated by climate
change and/or major environmental disturbances.
National assessments, networking, research and
collaboration among concerned countries and the
International Plant Protection Convention (IPPC)
are key strategies to avoid the further spread of
invasive alien species.
Invasive plants
The main invasive plant threat comes from
“transformer” plant species which have the
capacity to invade natural or slightly disturbed
forest associations, becoming the dominant
canopy species and completely modifying or
displacing entire ecosystems, with the loss of
many of the existing species (trees and others).
An example is the introduced tropical American
tree Prosopis juliflora in East Africa, which is
taking over large swathes of natural forest
and woodlands, with considerable negative
impacts on native tree populations (in terms
of both species and genetic diversity). It is also
damaging local livelihoods in the process (e.g.
Mwangi and Swallow, 2005). Another example
is the Australasian tree Melaleuca quinquenervia
in South Florida (United States of America),
introduced in the early 1900s and later planted
to drain swamps; it has since invaded up to
200 000 ha and transformed various ecosystems
in the Florida Everglades, causing major
environmental and economic damage (CarterFinn et al., 2006).
It appears that even minor climatic changes
can result in native tree species becoming more
invasive, spreading into neighbouring regions
and dramatically changing the forest dynamics,
structure and species composition. The spread
of Pittosporum undulatum and Leptospermum
laevigatum in southeastern Australia, for
example, is a portent of future challenges for
in situ FGR management, as the more extreme
climatic changes that are predicted would favour
disturbance-adapted pioneer and early secondary
tree species.
Island ecosystems are especially vulnerable to
invasive species. In just a few decades, Spathodea
campanulata (African tulip tree), introduced
in Fiji as an ornamental in 1936, has taken over
large areas of secondary forest and abandoned
agricultural fields (Brown and Daigneault, 2014);
it has also become invasive in mesic lowland
tropical moist forest up to 1 000−1 200 m
elevation in much of Oceania, including French
Polynesia (Meyer, 2004), Cook Islands (Meyer,
2000), Papua New Guinea and Australia (where
it has been declared a pest and its propagation
and sale as an ornamental are now prohibited).
The tropical American tree Miconia calvescens
has become one of the world’s most invasive
species and has completely taken over more
59
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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than a quarter of moist tropical forest in Tahiti
(France) (Meyer and Florence, 1996). In island
countries and territories of Oceania, excessive
opening of the forest canopy through intensive
timber harvesting, coupled with major cyclones,
has greatly favoured the spread of light-loving
vines such as Merremia peltata and Mikania
scandens; these vines and creepers have now
taken over large swathes of forest ecosystems,
thickly draping all trees and shrubs (e.g. Maturin,
2013; Kamusoko, 2014).
Insect pests
A global review of forest pests and diseases (FAO,
2009) revealed major and increasing threats to
forests from insect pests, both native and exotic.
Some examples of how exotic pests threaten
FGR and the economic and environmental
contribution of forests are discussed here, and are
mainly derived from the FAO review.
Invasive Sirex noctilio (European wood wasp)
has affected thousands of hectares of planted
pine forests in countries around the globe,
including South Africa, South American countries,
and Australia. It continues to spread and is now
threatening native pine and Douglas fir in North
America.
Heteropsylla cubana (leucaena psyllid) is a
significant pest of Leucaena leucocephala, a
fast-growing multipurpose tree legume native
to Mexico and Central America which has been
widely planted throughout the tropics. In the
mid-1980s, this insect spread across Asia (FAO,
2001b); the spread of the psyllid was especially
rapid because most leucaena plantings consisted
of a very narrow, nearly identical genetic base.
Anoplophora glabripennis (Asian longhorned
beetle) spread in Chinese planted forests as a
result of widespread planting of susceptible
poplar hybrids (EPPO, 1999) and has been
accidentally introduced in North America and
Europe. In China more than 200 million infested
trees have been removed to control outbreaks.
Authorities in the United States of America and
Canada have implemented emergency control
measures whenever the pest has been detected.
60
Strains of Populus nigra resistant to attack by the
Asian longhorned beetle have been developed in
China, through insertion of a Cry1Ac gene from
Bacillus thuringiensis (Hu et al., 2001).
Cinara cupressivori (cypress aphid), originally
from Europe (Greece) and the Near East (Islamic
Republic of Iran), spread to Africa around 1986.
In Kenya it rapidly caused major damage to
Cupressus lusitanica (cypress) plantations, which
constituted half of Kenya’s planted forest estate.
The cypress aphid killed a total of USD 27.5
million worth of trees in 1991 and was causing
a loss in annual growth of around USD 9 million
per year (Murphy, Nair and Sharma, 1996). This
is one example, of many, of the perils and risks
of plantation and farm forestry that is reliant on
a single exotic species, especially when grown in
monocultures. In Malawi the cypress aphid also
attacks and kills the highly endangered conifer
and national tree Widdringtonia nodiflora
(Bayliss et al., 2007); genetic resistance has yet to
be found.
A devastating outbreak of Dendroctonus
ponderosae (mountain pine beetle), a bark
beetle indigenous to western North America
that primarily feeds on Pinus contorta var.
latifolia, began in the 1990s and has affected
about 14 million hectares of forest land in
western Canada (Nealis and Peter, 2008), killing
50 percent of the standing volume in British
Columbia. Increased warming associated with
climate change enabled the beetle to expand
its range; it spread across a mountain range
and into Alberta in 2006, and may eventually
cause large-scale destruction to Pinus banksiana
in boreal forest (Cullingham et al., 2011). The
extent to which jackpine might show genetic
resistance to mountain pine beetle is unknown,
but natural hybrids with lodgepole pine are
expected to display some resistance.
Leptocybe invasa (blue gum chalcid), probably
native to Australia, is a relatively new threat to
planted eucalypt forests in Africa; it was first
reported in Kenya in 2002 and in South Africa in
2007. It has also been reported in Asia, Europe
and the Near East.
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
The severity and frequency of insect pest
outbreaks are projected to increase in concert
with extreme climatic factors. China reports
increased forest pest outbreaks in 2009 following
a major snowstorm in South China and severe
widespread drought in 2008 (China country
report).
Pathogens
Cases of virulent introduced pathogenic fungi
wreaking havoc on economically and environmentally important tree species are particularly
well documented in the Northern Hemisphere
(see examples in Box 5.3).
Over the past decade outbreaks of exotic
pathogens have caused major damage in
forests in the tropics and the Southern
Hemisphere. The reported increase in such
outbreaks may be attributed to movement of
goods and people, together with changing
climate and environmental disturbances. An
example is Fusarium circinatum (pine pitch
canker), introduced perhaps originally from
Mexico, which has been devastating the native
California (United States of America) stands of
Pinus radiata, with more than 90 percent of the
trees likely to succumb to the disease (Devey,
Matheson and Gordon, 1999). Pitch canker
has recently been found on P. radiata in South
Africa (where it seriously threatens the future
of the country’s pine plantation industry, which
comprises 670 000 ha and half the country’s
wood and fibre assets) (Coutinho et al., 2007)
and on Pinus species in Colombia (Steenkamp
et al., 2012). In areas of the world where pitch
canker has spread, management practices
will need to be altered and more pitch canker
resistant Pinus species and provenances will
need to be deployed, such as Pinus tecunumanii
from low-elevation sources and Pinus maximinoi
in Colombia.
Poplar rust was one of the first major exotic
tree diseases to be reported from the Southern
Hemisphere. Two species of poplar rust
(Melampsora medusae and Melampsora laricipopulina) appeared in Australia in 1972/73 and
rapidly spread to New Zealand, devastating
poplar plantations. However, considerable
genetic variation in resistance to poplar rusts
has been found among poplar species and
clones (and alternate conifer hosts), and disease
impacts can be managed by planting mixtures of
more resistant clones. Selection for poplar rust
resistance has been complicated, however, by the
appearance of different races of rust.
Fungal pathogens including another rust
(Atelocauda digitata) are a major concern for the
productivity of the more than 1 million hectares
of Acacia plantations in Asia (see Old et al., 2000).
However, different Acacia provenances show
considerable variation in susceptibility to disease,
which indicates potential for selection of resistant
genotypes.
In 2010 a new pathogen, Puccinia psidii (myrtle
rust or guava rust), originating in South America,
was detected in New South Wales, Australia. Most
species in the family Myrtaceae – which has more
than 2 000 plant species including eucalypts, and
which is Australia’s dominant plant family – have
the potential to become infected to some degree
by this rust species (Morin et al., 2012). It could
alter the composition, function and diversity of
many of Australia’s eucalypt-dominated forest
and woodland ecosystems and could have severe
impact on forest industries. Doran, Lea and
Bush (2012) have recently identified family- and
provenance-level resistance to Puccinia psidii in
Backhousia citriodora, an economically important
essential oil producing plant, through wideranging provenance, family, and clone trials.
Because forest pathogens are continuously
evolving, a combination of management
measures is needed to deal with them, including
deployment of diverse resistant genetic materials
and continuing breeding programmes with
access to genetic diversity. The above examples
and those in Box 5.3 illustrate how variation
in susceptibility to disease can be useful for
selection of resistant genotypes, underscoring the
importance of genetic diversity, its conservation
in native stands, and provenance and family
trials in combating threats from pathogens,
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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especially newly introduced strains and species.
Successful examples of breeding for pathogen
resistance include resistance to Mycosphaerella
pini (red band needle blight) in Pinus radiata in
New Zealand (Carson, 1990) and resistance to
Cronartium ribicola (white pine blister rust) in
Pinus monticola in North America (Sniezko, 2006).
Genetic pollution
A significant but largely unquantified risk to
FGR conservation and use is the uncontrolled
and undocumented mixing of gene pools of
forest tree species. This can occur within species,
whereby genetically diversified local populations,
which may possess valuable attributes, interbreed
with non-local germplasm introduced for
planted forest establishment. Hybridization of
local and introduced gene pools may reduce
local adaptation in subsequent tree generations
(Millar and Libby, 1989; Palmberg-Lerche, 1999).
Mixing of gene pools can also inadvertently
lead to incorporation of undesirable genes,
resulting in a diminished economic value for
production forests and vastly complicating and
increasing the costs of tree breeding in cases
where breeders may need to “unscramble the
omelette”. Interbreeding can also occur when
formerly allopatric (geographically isolated)
Box 5.3
Some destructive pathogens in Northern Hemisphere forests
Cryphonectria parasitica (Asian chestnut blight
fungus), accidentally introduced into the United
States of America early in the twentieth century,
wiped out almost the entire population of Castanea
dentata (American chestnut) including more than
3 billion trees over 70 million hectares (Cox, 1997);
this annihilation was accompanied by the extinction
of other species dependent on chestnuts, including
ten species of moths. Ironically, early salvation
logging may have removed some of the few American
chestnut trees that showed resistance to the disease.
Programmes have been implemented to backcross
surviving American chestnuts with blight-resistant
chestnuts from Asia for reintroduction into the former
natural range of the American chestnut.
Ophiostoma spp. (Dutch elm disease), since its
introduction into North America around 1930, has
killed more than 95 percent of Ulmus americana
(American elm), millions of trees. It is estimated that
only one in 100 000 trees is naturally resistant. A
few recently-cloned resistant individuals in Canada
(Shukla et al., 2012), along with newly identified
resistant diploids and triploids in the United States
of America (Whittemore and Olsen, 2011) and
interspecific hybrids derived from crossing with
resistant Asian Ulmus species, are paving the way
62
for American elms to be reintroduced in North
America. In the late 1960s a virulent strain of Dutch
elm disease (Ophiostoma novo-ulmi) introduced into
the United Kingdom wiped out most of the Ulmus
procera trees, although in the United Kingdom and
continental Europe they often survive as suckers and
in hedgerows (e.g. Forestry Commission, n.d.) Various
selection and breeding programmes with Ulmus spp.
in Europe, including development of interspecific
hybrids, have produced clones that are resistant to
the fungus.
Various pathogenic diseases, many only identified
or found over the past ten years, are now threatening
important tree species in the United Kingdom
(Forestry Commission, 2014). A new bacterial bleeding
canker, Pseudomonas syringae pathovar aesculi,
which was first detected around 2002, now afflicts 70
percent of Aesculus hippocastanum (horse chestnut)
trees and is likely to eventually kill them. Chalara
dieback, a serious disease caused by the introduced
fungus Hymenoscyphus pseudoalbidus, first identified
in 2012, infects Fraxinus species, often resulting in
tree death and spreading throughout Europe. Another
new bacterial disease, acute oak decline, threatens to
wipe out Quercus species in the United Kingdom.
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
related species are brought together. If the taxa
are not fully reproductively isolated and share
the same flowering times and pollinators, then
hybridization is likely; if the resulting progeny
are fertile and are not selected against, then the
eventual outcome can be loss of a species through
assimilation.
Factors
that
threaten
extinction
by
hybridization, such as habitat destruction,
fragmentation and species introductions, are all
increasing and often act synergistically (Rhymer
and Simberloff, 1996). Outbreeding depression
from detrimental gene flow may reduce the
fitness of a locally rare species, making it
vulnerable to extinction. Alternatively, pollen
swamping may result in a rare species’ loss of
genetic integrity, and it may become assimilated
into the gene pool of a more common species
(Potts, Barbour and Hingston, 2001).
FAO’s International Poplar Commission’s
Working Party on Poplar and Willow Genetics,
Conservation and Improvement has drawn
attention to the fact that populations of
some native poplar species have been rapidly
disappearing because of their spontaneous
hybridization with cultivars and/or displacement
by agriculture or other land uses (FAO, n.d.).
Natural stands of Populus nigra have almost
disappeared in Europe, and Populus deltoides
is seriously jeopardized in North America as a
result of interbreeding. However, there are few
documented examples in the literature in which
hybridization has threatened the existence of
rare tree species, although presumably this has
happened often during angiosperm evolution.
The main cited example is Cercocarpus traskiae, a
rare endemic species on Catalina Island, California,
United States of America, which has been
reduced to about seven mature pure individuals
and which hybridizes with the more abundant
Cercocarpus betuloides (Rieseberg et al., 1989).
In Fiji and Tonga, Santalum yasi hybridizes freely
with the introduced Santalum album, producing
more vigorous F1 hybrid offspring (Bulai and
Nataniela, 2005); this may eventually lead to
the disappearance of pure S. yasi due to natural
selective pressures and the commercial choices of
smallholder sandalwood growers (Huish, 2009).
Awareness is increasing in the forestry
profession of the risks that hybridization poses to
local gene pools. Lebanon, for example, in order
to protect the genetic integrity of its national
tree (Lebanon cedar, Cedrus libani), has taken a
ministerial decision that prohibits the import of
Cedrus germplasm into the country (Lebanon
country report). In Australia, Barbour et al. (2008)
have formulated a framework for managing
the risk of gene flow from plantations of nonnative Eucalyptus globulus into native eucalypt
populations in southern areas where E. globulus
does not occur naturally. This framework could
serve as a useful model for other tree genera and
species.
63
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
Chapter 6
Global forest trends affecting
forest genetic diversity
Forest trends
Forest cover
Land-use
activities,
primarily
agricultural
expansion and timber extraction, have caused a
net loss of about 700 to 1 100 million hectares of
forest in the past 300 years (Foley et al., 2005).
In spite of the notable forest restoration and
afforestation activities undertaken to reverse the
trend of forest cover loss (5.7 million hectares
restored or planted annually), 13 million hectares
of forest are still being lost every year. This loss
represents a serious threat to habitats, species
and genetic diversity.
Sustainable forest management is of major
importance for its role in maintaining biological
diversity and global ecological functions while
enabling adequate use of the products derived
from forests to meet growing demand. The
Global Forest Resources Assessment 2010 (FRA
2010) (FAO, 2010a) reported on broad progress
towards sustainable forest management since
1990 and found that at the global level the
situation has remained relatively stable. The 2010
assessment did not include species or populationlevel indicators suitable for a global comparison
of trends over time and therefore did not directly
report on FGR. However, biological diversity
was addressed through reference to the area of
primary forest, areas designated for conservation
of biological diversity and area of forest in
protected areas.
This section on forest trends reports some of
the major findings of FRA 2010 (FAO, 2010a), with
notations on impacts and considerations for FGR.
The world’s total forest area is just over
4 billion hectares, with the five most forestrich countries (the Russian Federation, Brazil,
Canada, the United States of America and
China) accounting for more than half of the
total forest area. Human-mediated forest cover
reduction and forest degradation have been
among the main causes of negative change in
forest genetic resources, including loss of tree
species, over the past few hundred years. Loss
of forest cover will almost certainly continue
to be a problem while the world’s population
continues to rise.
The rate of loss of forest cover – mainly from
conversion of tropical forest to agricultural land
– shows signs of decreasing but is still alarmingly
high. Around 13 million hectares of forest were
converted to other uses or lost through natural
causes each year between 2000 and 2010,
compared with 16 million hectares per year in the
1990s.
In spite of its high importance, the natural
moist tropical forest has continued to diminish
rapidly at the global level. Most forest losses in
the period 2000–2010 occurred in the regions
and countries with more biodiversity-rich tropical
forests (Tables 6.1 and 6.2). Brazil and Indonesia,
two countries with biodiverse and FGR-rich
forests, had the highest net loss of forest in the
1990s but have significantly reduced their rate of
loss, which will also entail a slowing in the rate of
loss of tree species and populations.
65
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
TABLE 6.1
Area of primary forest change, 1990−2010
Region
Area of primary forest
(1 000 ha)
1990
1990−2000
2000−2010
1990−2000
2000−2010
7 024
6 430
−57
−59
−0.78
−0.88
North Africa
15 276
14 098
13 990
−118
−11
−0.80
−0.08
West and Central Africa
37 737
32 540
27 527
−520
−501
−1.47
−1.66
East Asia
28 179
26 456
25 268
−172
−119
−0.63
−0.46
South and Southeast Asia
87 062
83 587
81 235
−348
−235
−0.41
0.29
Western and Central Asia
2 924
3 083
3 201
−16
12
0.53
0.38
Europe
5 183
5 360
5 438
18
8
0.34
0.14
207
206
205
n.s.
n.s.
−0.07
−0.02
Caribbean
Central America
North America
Oceania
South America
World
2010
Annual change
(%)
7 594
East and Southern Africa
2000
Annual change
(1 000 ha)
5 766
5 226
4 482
−54
−74
−0.98
−1.52
274 920
273 795
275 035
−113
124
−0.04
0.05
41 416
39 191
35 493
−222
−370
−0.55
−0.99
684 654
653 691
624 077
−3 096
−2 961
0.46
−0.46
1 190 919
1 144 258
1 102 382
−4 666
−4 188
−040
−0.37
Source: FAO, 2010a.
Note: n.s. = not significant.
TABLE 6.2
The ten countries with the largest annual net
loss of forest area, 1990–2010
Country
Brazil
Net loss of
forest area
(1 000 ha)
% loss
2 642
0.49
Indonesia
498
0.51
Nigeria
410
3.67
United Republic of Tanzania
403
1.13
Zimbabwe
327
1.88
Democratic Republic of the
Congo
311
0.20
Myanmar
310
0.93
Bolivia
290
0.49
Venezuela (Bolivarian Republic of)
288
0.60
Australia
280
0.18
Source: FAO, 2010a.
66
Forest loss in tropical areas was relatively
stable on a percent basis from the period 1990–
2000 to 2000–2010. In West and Central Africa
and in South America the rate of forest loss was
the same for both periods (0.66 and 0.45 percent
per annum, respectively). In East and Southern
Africa the rate of forest loss increased from
0.62 to 0.66 percent per annum over the two
periods, while annual forest loss in South and
Southeast Asia dropped substantially from 0.77
to 0.23 percent. Annual forest loss also declined
substantially in Central America, from 1.56 to
1.19 percent.
At the regional level, the changes in the
period 2000–2010 were as follows.
• South America suffered the largest net loss
of forests – about 4.0 million hectares per
year.
• Africa had the next largest loss of forests,
3.4 million hectares annually.
• Oceania also reported a net loss of
forest (about 700 000 ha per year),
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
mainly because of large losses of forests in
Australia, where severe drought and forest
fires exacerbated the loss of forest during
the reporting period.
• The area of forest in North and Central
America was estimated as almost the same
in 2010 as in 2000.
• The forest area in Europe continued
to expand, although at a slower rate
(700 000 ha per year) than in the 1990s
(900 000 ha per year).
• Asia reported a net gain of forest of
more than 2.2 million hectares per year,
in contrast with the net loss of some
600 000 ha annually in the 1990s. The
gains were primarily due to the large-scale
afforestation reported by China.
Both South America and Africa are rich in
tree species and FGR, and their high forest losses
represent a loss, though poorly documented, of
irreplaceable FGR. Meanwhile the gains of forest
area in Asia mask a loss of valuable FGR (of both
species and especially populations of useful tree
species) from shrinking native forests in many
countries in South and Southeast Asia.
Forest area was increasing or stable in a
number of countries, owing to the establishment
of reserves and protected areas, controls on
logging and overharvesting, implementation
of sustainable forest management, planting of
trees and plantations and natural regeneration
of abandoned farmland. Countries with
constant or increasing forest cover tended to be
developed countries with highly developed forest
management and administration institutions,
although China and India achieved large
increases in the decade, mainly through planting.
Between 1990 and 2010 China planted more than
1.9 million hectares per year. The United States of
America also undertook planting on a large scale
(805 000 ha per year).
It is impossible to estimate accurately the
genetic loss that is resulting from deforestation
and forest degradation, given the generally
poor knowledge of forest genetic resources,
particularly for tropical forest. It should also
be noted that many land-use or forest product
extraction practices (e.g. forest grazing, road
construction, fuelwood, NWFPs) can degrade
forest ecosystems – in particular with regard
to their productivity, biomass, stand structure,
species composition and genetic diversity – even
without changing the forest area (Foley et al.,
2005; Todd and Hoffman, 1999).
Primary forests. Primary forests, defined as “forest
of native species where there are no clearly visible
indications of human activities and the ecological
processes have not been significantly disturbed”
(FAO, 2010a), are among the most species-rich,
diverse terrestrial ecosystems, i.e. those most rich
in FGR. Primary forests make up 36 percent of the
global forested area but have been declining by
4.7 million hectares per year during the 1990s and
4.2 million hectares per year between 2000 and
2010 – with a loss of more than 40 million hectares
since 2000 (0.37 percent per annum).
Containing an estimated 50 percent or more
of all terrestrial biodiversity, primary forests (in
particular tropical moist forests) have an essential
role in conservation of biodiversity (Gibson et al.
2011), and their loss is likely to be accompanied by
loss of genetic diversity. There is no substitute for
primary forest to maintain tropical biodiversity.
Furthermore, the food security, livelihoods and
cultural and spiritual identity of many indigenous
people are often linked to primary forests (CBD
Secretariat, 2009). The pursuit of development
can jeopardize their conservation in countries
where primary forests constitute a large portion
of the forest cover (e.g. 95 percent of total forest
cover in Suriname, 92 percent in Brazil, 91 percent
in Papua New Guinea, 89 percent in Peru and 65
percent in Gabon) and where forest production,
in particular timber, provides an important
contribution to the national economy.
Planted forests. The area of planted forest
has increased over time, amounting to a total
of 264 million hectares in 2010, compared with
67
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
Conservation of forest biodiversity
178 million hectares in 1990. Planted forests
currently represent 7 percent of the total forest
area, with the highest proportion in Asia (almost
20 percent) (Figure 6.1) and provide an essential
contribution to the supply of industrial wood,
wood energy and non-wood forest products as
well as environmental services including soil and
water protection.
It is estimated that less than 400 tree species –
still a high number – are used in planted forests.
In many countries in the temperate and boreal
zone, however, the ten most planted species
represent more than 90 percent of the total
growing stock; while in tropical countries with
high species diversity, they represent less than 20
percent of total growing stock.
Regionally and subregionally there are large
differences in the proportion of planted forest
consisting of exotic species, from 100 percent in
East and Southern Africa to a very low proportion
in North America and arid regions (Figure 6.2).
In 2010, 12 percent of the world’s forest area was
designated for conservation (Figure 6.3), while
legally established protected areas – including
national parks, game reserves, wilderness areas
and others – comprise an estimated 13 percent
of the world’s forests. The main functions of
protected areas usually include the conservation
of biological diversity, the protection of soil
and water resources, and/or the conservation of
cultural heritage.
Globally, the area of forest designated primarily
for conservation of biodiversity has continued to
increase substantially, by 1.92 percent per annum
from 2000−2010. This increase is a result of global
efforts to conserve biological diversity (e.g.
the Aichi Biodiversity Targets under the CBD).
However, even if the area of forest designated
for conservation of biodiversity is growing,
primary forests are unfortunately still increasingly
threatened.
FIGURE 6.1
Characteristics of the world’s forests in 2010 (%)
Africa
Asia
Europe
Latin America
and the Caribbean
North and Central
America
Oceania
Primary
Other naturally regenerated
World
Planted
0
Source: FAO, 2010a.
68
20
40
60
80
100
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
FIGURE 6.2
Proportion of planted forest area made up of exotic species (%)
East and Southern Africa
North Africa
West and Central Africa
East Asia
South and Southeast Asia
Western and Central Asia
Europe
North America
Central America
Caribbean
South America
Oceania
0
20
40
60
80
100
Source: FAO, 2010a.
Note: In some regions (e.g. North America) native species may be underestimated, because some countries do not report planted forests as
plantations, so most of what is reported as plantation is exotic by definition.
In sum, the results for forest biodiversity
conservation were mixed, with the area of primary
forest recording one of the largest negative rates
(in percentage terms) of all measures. The area of
forest designated for conservation of biological
diversity increased by about 6.3 million hectares
per year during the decade 2000−2010 with a
similar increase in the area of forest in protected
areas. The area under production forests,
considered equally vital for conservation of FGR,
has continued to decline at an increasing rate,
by about 2 million hectares per year during the
1990s and 3 million hectares per year between
2000 and 2010.
Trends in ownership
Ownership is a key variable determining forest
management options and sustainability. FAO
reported that in 2005, 80 percent of the world’s
forests were public and 18 percent private
FIGURE 6.3
Designated functions of forests reported in the
Global Forest Resources Assessment 2010
Other
(7%)
Unknown
(16%)
Multiple use
(24%)
Social
services
(4%)
Production
(30%)
Conservation
of biodiversity
(12%)
Protection of
soil and water
(8%)
Source: FAO, 2010a.
69
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
(2 percent other). Of the privately owned forest,
58 percent was owned by individuals, 19 percent
was owned by corporations or institutions and 23
percent was community or indigenous forest. Of
publicly owned forest, 80 percent is under public
administration.
Public ownership remains the predominant
ownership category in all regions. In many
countries decentralization has been having a
large impact on forest management. While
forest management is guided by national and
international legal instruments, decentralization
may allow different subnational administrations
within a country (at the community, province or
state level) to apply different forest management
regulations. Furthermore, the role of local
communities and indigenous people in forest
and forest ecosystem management is increasingly
acknowledged, which is leading to greater
integration of their rights in forest management
plans. Ensuring the consistency of the regulations
at the different levels remains a challenge,
especially in seeking to combine the user rights of
local and indigenous people and the conservation
of endangered species.
Consequences of forest changes
for genetic diversity
The loss of forest cover has dire consequences
for forest genetic diversity, loss of ecosystems,
loss of species and loss of intraspecific diversity.
While globally the rate of forest loss is slowing
(FAO, 2010a), the impacts of further forest loss
on FGR are increasing proportionally because
the losses of forests are affecting a smaller
residual base of native forest, are concentrated
in more biodiversity-rich forests, and are leading
to greater fragmentation, with long-term
impacts on associated animal species, gene flow
and viability of more fragmented species and
populations.
Loss of ecosystems
Loss of FGR due to disappearance or significant
modification of the ecosystems of which they are
a constituent is an increasing threat. For much
70
of the twentieth century and until recently, the
main threat was from conversion of forest to a
different land use, mainly agriculture. Examples
of major forest loss include the following:
• Brazil’s Atlantic forest has been reduced
to 2 to 5 percent of its original extent,
according to the country’s report.
• Forest cover in Ethiopia has diminished from
more than 50 percent in the middle of the
twentieth century to around 3 to 11 percent
(depending on forest cover definition) at
present, with most forest types reduced
to small fractions of their former extent,
including conversion to more open
woodland formations (Ethiopia country
report).
• Eastern Australia’s moist subtropical
lowland forest has been reduced to 7.2
percent of its original extent (Australian
Government, Department of the
Environment, 2011).
• Many smaller countries, such as Haiti
and Samoa, have lost almost their entire
lowland tropical forests.
It is estimated that 20 to 33 percent of the
Brazilian Amazon’s more than 11 000 tree
species, especially rare and narrowly distributed
endemics, will become extinct because of habitat
loss (Hubbell et al., 2008); a broadly similar
situation can be expected for much of the tropics.
Dramatic breakdown of forest ecosystems
and changes in their function and structure are
increasingly attributable to climate change and
associated extreme events such as uncontrolled
wildfires and alien invasive species. Climate
change will have particular impact in isolated
montane forest ecosystems (including cloud
forests) in tropical and subtropical zones
(e.g. tropical Central and South America and
the Caribbean, East and Central Africa, the
Philippines, Malaysia, Indonesia and Papua
New Guinea), especially as these forests often
have a high proportion of unique endemic
associated species which may have no possibility
of migrating to other climatically suitable
habitats. Foster (2001) has described a scenario
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
of complete replacement of many of the cloud
forests with narrow altitude range by loweraltitude ecosystems, as well as the extinction of
cloud forests located on high mountain peaks.
Some forest ecosystem changes are associated
with changes to keystone animal species. In a
recent literature review, Ripple and Beschta
(2012) concluded that predation by large
mammalian carnivores, notably sympatric grey
wolves (Canis lupus) and bears (Ursus spp.),
limits densities of large mammalian herbivores
in boreal and temperate forests of North
America and Eurasia, with impacts on tree and
shrub recruitment. The same authors previously
reported that cougars (Puma concolor) limit mule
deer (Odocoileus hemionus) densities, releasing
woody plants from browsing and maintaining
biodiversity in western North America (Ripple
and Beschta, 2008). Large carnivores such as the
tiger (Panthera tigris) and lion (Panthera leo)
are increasingly threatened and reducing their
natural ranges in many places. Reductions in topof-the-chain predator populations and changes
to other keystone species, such as elephants
(Loxodonta spp.) in Africa, will result in changes,
both major and subtle, to forest and woodland
ecosyems and will alter the FGR contained in
them.
Loss of tree species
Scientific consensus is growing regarding the
view that a new era of mega species extinction
has begun, with current rates of extinction at
least three orders of magnitude more than the
average rate of extinction in the Earth’s geological
and biological history (Pimm and Brooks, 1999).
In late 2012 the IUCN Red List of Threatened
Species (www.iucnredlist.org), widely regarded
as the most comprehensive and objective global
evaluation of the conservation status of plant
and animal species, included 65 518 species, of
which 20 219 are threatened with extinction
and 795 already extinct. IUCN (2012) reported
that of Madagascar’s 192 unique palm species,
a staggering 159 species are threatened with
extinction. Indigenous palm seeds are becoming
an important NWFP export product, and this is a
contributory threat factor for some species.
Thomas et al. (2004) have shown through
modelling that between 18 and 35 percent of
the world’s animal and plant species are on the
path or committed to extinction due to climate
change, and this figure does not take into account
interactions with other threats; these authors
have also shown that the threat to survival of
species from climate change is much greater than
the threat from habitat loss, with some variation
depending on the biome under consideration.
Through the Global Trees Campaign and
under the auspices of IUCN’s Species Survival
Commission, conservation status has recently
been partially or fully assessed for certain
plant groups (including conifers [Coniferae],
cycads [families Boweniaceae, Cycadaceae and
Zamiaceae], Magnoliaceae, Acer spp., Quercus
spp., palms [Arecaceae] and Rhododendron
spp.) and certain areas (including Central Asia,
Guatemala, Ethiopia, Eritrea and Mexican cloud
forests) (see Box 6.1). However, most families and
genera comprising mainly tree and woody species
have yet to be subjected to comprehensive
assessments of their level of endangeredness,
which could help determine where best to direct
conservation effort and resources.
The number of threatened forest species
reported in the country reports is shown in
Figure 6.4, by region. These data have different
sources and their detail and reliability vary.
Developed countries with greater available
government resources, but often less species
diversity, sometimes maintain their own Red Lists.
Sweden, for example, maintains a Red List which
includes Fraxinus excelsior, Tilia platyphyllos,
Ulmus glabra, Ulmus laevis and Ulmus minor; the
main threats are exotic diseases (Sweden country
report). Threat assessments for tree species in
developing countries are usually lacking owing to
a shortage of trained botanists and conservation
biologists and resources to support field surveys.
India lists 261 tree and woody species whose
genetic diversity is threatened, including 94
species in the highest threat category; the
71
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
Box 6.1
Conservation status of forest species assessed under the Global Trees Campaign
Geographic assessments
In Guatemala, 79 endangered tree species were
identified, including 10 critically endangered endemics
(Vivero et al., 2006).
In the floristically rich cloud forests of Mexico,
which are replete with endemics, approximately
60 percent of the 762 tree species in 85 botanical
families were assessed as threatened (GonzálezEspinosa et al., 2011).
Of 96 tree taxa assessed in Central Asian countries,
including Kazakhstan, Kyrgyzstan, Tajikistan,
Turkmenistan and Uzbekistan, 46 percent (or 44 taxa)
were found to be threatened with extinction in the
wild. This status was attributed to a combination of
overexploitation, desertification, pests and diseases,
overgrazing, fires, rural poverty, lack of alternative
energy sources and the lack of institutional capacity
to protect and regulate forests (Eastwood, Lazkov and
Newton, 2009).
A preliminary assessment of 428 endemic and
near endemic woody plants in Ethiopia and Eritrea
determined that 135 species (including 31 trees) were
threatened (Vivero et al., 2011).
identified threat types are almost all related to
forest cover loss, degradation and fragmentation,
including combinations and interactions of these
threats. However, the taxonomic assessment of
many tropical tree genera, including those with
important FGR such as Diospyros, Mangifera,
Syzygium and Terminalia, is often incomplete.
Furthermore, updated taxonomic information
and botanical keys may not be readily available
in the countries where the species naturally occur.
Available threat assessments, where these have
been undertaken, are typically several or many
years old and are in need of updating.
The Global Tree Specialist Group, part of
the IUCN Species Survival Commission, has
identified major challenges for conservation of
individual tree species. The group estimates that
approximately 8 000 tree species are threatened
with extinction, with about 1 000 tree species
critically endangered and likely to become extinct
unless urgent action is taken (Oldfield, Lusty
and MacKinven, 1998; Global Trees Campaign,
2014). Figure 6.5 shows the main threats to
52 endangered tree species in different plant
families and geographic regions profiled at the
Global Trees Campaign website (globaltrees.
org). However, many species are threatened
by a combination of threats and interacting
threats. These data are from a small sample
(about 5 percent of threatened tree species), and
overharvesting is likely to be overrepresented
because of deep concerns about precious timber
tree species.
Overharvesting, including poorly regulated,
unregulated and illegal harvesting, is still
arguably the most important threat factor for
FGR at present, because this activity causes a
loss of genetic diversity and populations in those
72
Plant group assessments
A global assessment of 151 species in the family
Magnoliaceae found approximately 74 percent (or
112 species) to be threatened (Cicuzza, Newton and
Oldfield, 2007).
Approximately 45 percent of Quercus species (79
species) are considered endangered of the 176 species
for which data were available and sufficient for
assessment (Oldfield and Eastwood, 2007).
In assessment of the conservation status of 125
species of maple trees (123 Acer spp. and 2 Dipteronia
spp.) at least 54 taxa (28 percent) were found to be
threatened (Gibbs and Chen, 2009).
Of the approximately 1 018 known Rhododendron
species, mainly woody shrubs, approximately 25
percent (or 316 species) were found to be threatened
(Gibbs, Chamberlain and Argent, 2011).
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
FIGURE 6.4
Number of species and subspecies mentioned as threatened (at various levels) in country reports,
by region
8 000
7 000
6 000
5 000
4 000
3 000
2 000
1 000
0
Total
Africa
Asia
Europe
Total number of species and subspecies reported
FIGURE 6.5
Main threats to 52 endangered tree species
profiled by the Global Trees Campaign
Climate change
and invasives
(6%)
Fire and
overgrazing
(13%)
Overharvesting
(37%)
Habitat loss
and conversion
(21%)
Biological factors,
including being naturally
rare or restricted
(23%)
Source: Global Trees Campaign data at globaltrees.org.
Latin America
North
and the Caribbean America
Near East
Paciic
Species and subspecies threatened
tree species that have the most economic value
and utility. Over the next century, climate change
and interactions with other threats are likely to
become the most important threat for tree species
and populations.
Loss of intraspeciic diversity
The loss of intraspecific diversity in economically
important tree species has been a major concern
of the forestry profession for many decades.
Despite the many continuing and long-term
threats to FGR, a high (although variable) level
of success has been achieved in conserving and
using the genetic diversity of many commercially
important tree species for timber and paper pulp
production. The successes, of which there are
many examples, have often been achieved under
the auspices of tree breeding programmes in
developed countries, increasingly led by privatesector consortia. Genetic resources work on some
73
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 2
major tropical timber and NWFP species has also
been undertaken by national agencies of tropical
developing countries, often with international
support. For example, FAO and the Danish
International Development Agency (DANIDA)
Forest Tree Seed Centre have supported work on
Tectona grandis, Gmelina arborea and Azadirachta
indica; ICRAF, work on African agroforestry tree
product (AFTP) species; the Tropical Agricultural
Research and Higher Education Center (CATIE),
work on Swietenia macrophylla; Camcore (an
international tree breeding organization based
in the United States of America), work on tropical
American pines; and the Australian Centre for
International Agricultural Research (ACIAR) and
the Commonwealth Scientific and Industrial
Research Organization (CSIRO) Australian Tree
Seed Centre, work on Chukrasia tabularis and
Casuarina equisetifolia.
The main threats to intraspecific diversity in
tree species are essentially the same as those that
cause species extinction (see preceding section).
For example, major losses to diversity have
occurred for some high-value species that have
been selectively and heavily harvested for their
timber and NWFPs. Paradoxically, this has meant
that some of the most economically useful tree
species have been the most genetically depleted.
According to its country report, Ethiopia is rich
in FGR, including more than 1 000 woody plant
species and two biodiversity hotspots (the Eastern
Afromontane hotspot and the Horn of Africa),
but the viability of populations of important
woody species is threatened by fragmentation
(reduced gene flows), coupled with utilization
pressures, fire and invasive species which increase
the risk of local extinctions.
The loss of entire populations or genetically
distinctive provenances (for species exhibiting
clinal variation) has both short- and longterm adverse consequences. The short-term
consequences include potential major changes to
ecosystem function and services for native forests
in which these populations or provenances occur,
through loss of documented seed sources of
known performance. Longer-term consequences
74
include species extinction (for which loss of
populations is a well identified precursor)
and loss of vital genetic material for selection
and tree improvement programmes. For trees
introduced into a new environment with a broad
genetic base, better-adapted landraces may often
evolve in only one or two generations, but the
same is not true for recovery of lost diversity. A
study on Pinus resinosa has indicated that very
long periods, possibly on the scale of tens of
thousands of years, are required for long-lived,
long-generation organisms like trees to recover
genetic diversity after it has been lost (Mosseler,
Egger and Hughes, 1992).
The loss of intraspecific diversity in economically
important species has consequences not only for
immediate seed supply for replanting, but also
in terms of reduced opportunities for selection
and breeding, as often only lower-quality or less
desirable phenotypes remain. Dysgenic selection
(perpetuation of defective or undesirable genes
and traits) is most likely if successive regeneration
cycles are derived from only a small residual
number of poor-quality phenotypes (Ledig,
1992). Cornelius et al. (2005) found only a small
(≤5 percent) and rather insignificant maximum
negative dysgenic response to a single selective
logging−mediated phenotypic selection event in
Swietenia macrophylla, but for species with more
heritable traits (e.g. chemotypes) and/or several to
many cycles of selection of superior phenotypes,
dysgenic selection is more problematic. Swietenia
mahogani, native to the Caribbean Islands and
southern tip of Florida (United States of America),
is the most valuable mahogany timber producing
species and has been commercially exploited
for more than 500 years. The small residual
populations are thought to have undergone
dysgenic selection (Styles, 1972). Hybridization
with other Swietenia species (e.g. with
S. macrophylla on Cuba) is another threat to the
species’ genetic integrity and genetic resources.
Box 6.2 lists some examples, from around the
globe, of the many hundreds of valuable tree
species that have already lost, or are at imminent
risk of losing, important intraspecific diversity.
DR I VERS OF C HA NGE A ND T RENDS AFFEC T I NG FOREST GENE T IC RESOU RC ES
Box 6.2
Loss of intraspeciic diversity in valuable species: some examples
Boswellia papyrifera is an economically important
NWFP tree in Ethiopia and Eritrea but is rapidly declining
and predicted to become commercially extinct within
the next 15 to 20 years. The decline is related to resin
tapping, which reduces reproductive and recruitment
potential (Rijkers et al., 2006; Eshete et al., 2012), and
to attack by longhorned beetles, excessive fire and
increased grazing pressure. Seedling recruitment is failing
because of excessive fire and increased grazing pressures,
preventing dying trees from being replaced (Groenendijk
et al., 2012). Forest reduction and degradation and
competition for land use have also been identified as
threatening factors (Ethiopia country report).
Dalbergia cochinchinensis has been heavily and
selectively overharvested throughout its natural range
in Cambodia, the Lao People’s Democratic Republic,
Thailand and Viet Nam and continues to be cut, often
illegally. Its intraspecific variability is highly threatened
(Thailand country report). Good seed sources from native
stands are scarce, as surviving populations are reduced to
scattered and isolated trees of poor phenotypes.
Endospermum medullosum – populations of the
fastest-growing trees, which originate from eastern
and southeastern Espiritu Santo in Vanuatu (Vutilolo
et al., 2005), have almost disappeared as a result of
land use change, absence of regeneration in coconut
plantations and cattle ranches, and harvesting of
remnant trees (Corrigan et al., 2000; Vanuatu country
report). Some efforts to conserve this species in ex situ
plantings are now under way (Doran et al., 2012).
Erythrophleum fordii is a valuable timber tree
threatened by overexploitation in China. It now only
occurs there in small, fragmented and degraded stands
and with greatly diminished genetic diversity (China
country report).
Populus euphratica and Populus ilicifolia are
fast-growing, multipurpose riparian poplars from
the Near East, Central Asia, China and Kenya, with a
remarkable range of tolerance to edaphic and climatic
extremes. However, they are declining and endangered
throughout their range by clearance, overharvesting
and modification to hydrological regimes (Viart, 1988;
Ball, Russo and Thomson, 1996; Cao et al., 2012).
Prunus africana is a keystone afromontane species
important for its bark, which is harvested for use in
treatment of benign prostatic hypertrophy. The species
has been listed by the Convention on International
Trade in Endangered Species of Wild Fauna and Flora
(CITES) (Appendix II) since 1995, but almost all native
populations in central, eastern and southern Africa are
threatened by overharvesting, which often kills the trees,
and also by land-use and climate changes. In South
Africa, close monitoring and controls may provide a
greater level of protection than in other parts of its range
(South Africa country report). Populations of P. africana on
Madagascar, which are morphologically distinct and likely
constitute a different taxon, are similarly threatened.
They are no longer exported because of previous
overharvesting (Madagascar country report).
Pterocarpus santalinus, a highly valuable timber and
NWFP species from the state of Andhra Pradesh in India,
has been overharvested, especially during the 1950s
and 1960s. The species was listed in CITES Appendix II in
1995, but an illegal smuggling trade continues, causing
concern for loss of genetic diversity (MacLachlan and
Gasson, 2010; India country report).
Santalum sp. – an undescribed species of sandalwood,
referred to in the literature as S. macgregorii (Brophy
et al., 2009) – exists in three small populations, each
consisting of only a few individuals, in coastal areas
of Western Province, Papua New Guinea. It has highly
fragrant heartwood with high santalol content and is
at high risk of being harvested, which would cause the
species to become extinct as no actions are being taken
to ensure its natural regeneration or ex situ conservation.
75
Part 3
CURRENT AND
EMERGING
TECHNOLOGIES
CU RREN T A ND EMERG I NG TEC HNOLOG IES
Chapter 7
Trait-based knowledge of
tree genetic resources
Until the advent of biochemical and molecular
methods, the only way to estimate genetic values
or variation was by measuring phenotypes and
using statistical tools to separate genetic effects
from environmental influences. Although field
studies designed to estimate genetic parameters
have declined, such studies are still essential for
understanding genetic control of phenotypic
traits.
Before the development of genetics as a
scientific discipline, trees had been planted for
food, wood, shade and religious purposes for
thousands of years. Knowledge of trait variation
was used in traditional farming and subsistence
systems to select, save and/or cultivate valuable
individual trees on the basis of phenotypic
characteristics. Besides food tree species such as
Citrus sinensis (orange), Malus domestica (apple)
and Olea europaea (olive), other cultivated tree
species of significant importance include Cinchona
lederiana from Bolivia, which was transported
to Europe to combat malaria and then grown
in Asia, and Hevea brasiliensis, the Pará rubber
tree, whose seeds were transported from Brazil
to the United Kingdom and then to Asia in the
late 1800s. All of these species were subject to
selection and breeding on the basis of valuable
phenotypic characteristics.
Indigenous and traditional
knowledge
Traditionally living societies generally maintain
an intimate relationship with the natural
world of their (actual and/or historical) living
environments. They are (or were until recently)
strongly dependent on natural resources for their
livelihoods. Through their intimate relationship
with, and dependence on, the natural world,
traditional societies have developed extensive
knowledge about natural resources, often built
up over generations. For such people, trees
are among the most important and useful life
forms because they are more complex and more
multipurpose than herbaceous plants; apart
from fulfilling human needs for food, medicine,
construction materials and fuel, they provide a
wide array of environmental services (Thomas
and Van Damme, 2010). The importance of tree
species for traditional societies depends to a
large extent on the floristic composition of their
living environments. For example, Clement (1999)
calculated that of the 138 species under cultivation
or management at the time of European arrival
in Amazonia, 68 percent were trees or woody
perennials.
Ethnobiological research on people-plant
interactions has traditionally focused mainly on
the utilitarian (ethnobotanical) dimension of
plant species and far less on ecological aspects.
However, studies on traditional ecological
knowledge (TEK) of plants have demonstrated
deep knowledge of species’ habitat preference,
phenology, pollination systems, seed dispersers,
species associations, intraspecific variation,
pests and diseases, environmental services
provided and behaviour under different types
of management (e.g. Assogbadjo et al., 2008a;
Hmimsa, Aumeeruddy-Thomas and Ater, 2012;
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Parra, Blanca and Casas, 2012; Fraser et al.,
2012). Most studies of traditional knowledge
about plants at subspecies level have focused
on ethnolinguistic and ethnotaxonomic aspects,
i.e. the ways traditional societies name and
classify plants (Berlin, 1992). Ethnotaxonomical
classifications often coincide to a certain extent
with corresponding scientific classifications
and can provide a first approximation of
existing intraspecific variation in plants.
Overdifferentiation − splitting a scientific species
or subtaxon into two or more traditionally named
groups − is primarily encountered with cultivated
varieties for which distinctive scientific names are
often lacking; such cultivated varieties may be
genetically different (Hunn and Brown, 2011) or
not (e.g. Assogbadjo et al., 2008b).
TEK can provide valuable information to inform
scientific research on the ecology, management,
intraspecific variation and conservation of tree
species, as scientific information is still lacking for
many tropical tree species. However, TEK about
tree species is being lost more rapidly than the
respective scientific knowledge is increasing;
thus strengthened efforts to record remaining
knowledge are urgently needed. Participatory
tree breeding and domestication of tree species
is a fairly recent approach which combines TEK
about tree use and management with scientific
advances in germplasm collection, selection and
propagation as well as with market development.
Its ultimate goal is to improve people’s livelihoods
(Dawson et al., 2013). TEK can also be a rich source
of inspiration for designing biological approaches
to tackling current environmental problems
such as the development of renewable energy
resources (Martin et al., 2010) or environmental
restoration (Douterlungne, Thomas and LevyTacher, 2013).
Tree and landscape management
Traditional societies are typically positioned at
the interface of the natural and cultural worlds.
They live in close proximity to natural vegetation
from which they extract livelihood goods, and
at the same time they engage in different types
80
of plant management. Plant management in
traditional societies covers a continuum ranging
from gathering and protecting plants in wild
populations, to deliberately tolerating plants in
human-made habitats (also defined as disturbance
habitats), to cultivating domesticates as well as
non-domesticates. Of all anthropogenic habitats,
home gardens in particular are laboratories
where people have experimented with plant
genetic resources; they contain a combination of
strictly wild plants, camp followers (weeds, which
can be trees, e.g. in Amazonia [Balée, 1994]),
spared and tolerated plants, and cultivated and
domesticated plants. Through ongoing processes
of experimentation and innovation, wild plants
with desirable traits are gradually brought into
the cultural sphere (Bennett, 1992; Miller and
Nair, 2006; Clement, 1999; Thomas and Van
Damme, 2010).
Useful wild tree species may also enter the
cultural sphere when spared during land clearing
for human use, as this increases contact intensity
between people and trees remaining in the
margins of crop fields. Intensity of contact,
salience, accessibility and availability of plant
species are often correlated with their perceived
usefulness to people (Adu-Tutu et al., 1979;
Byg, Vormisto and Balslev, 2006; Thomas, 2009;
Thomas et al., 2009; Thomas, Vandebroek and Van
Damme, 2009; Turner, 1988). Tree management
is not limited to humanized landscapes such as
home gardens and swiddens, but occurs also in
natural vegetation where certain species may
receive protection, e.g. through removal of
competing plants or pests to enhance the target
plants’ chances of survival. More significant,
however, is landscape domestication, which was
initiated by early human societies all around the
world (Chase, 1989; Clement, 1999; Anderson,
2005; Young, 2009; Aumeeruddy-Thomas et al.,
2012; Sheil et al., 2012) and has culminated in the
highly artificial land uses of modern society. The
impact of longstanding landscape domestication
in forest environments is, for example, evidenced
by enrichment in useful species (Wiersum, 1997;
Shepard and Ramirez, 2011; Levis et al., 2012),
CU RREN T A ND EMERG I NG TEC HNOLOG IES
anthropogenic forests (Balée, 1989) or anthrosols
such as black-earth soils containing charcoal
and cultural waste from prehistoric burning and
settlement, which carry distinctive vegetation as
a consequence of their high nutrient content.
In Amazonia, black-earth soils are generally
associated with forests that are enriched with
useful species such as Brazil nut trees (Clement
and Junqueira, 2010; Junqueira et al., 2011).
Longstanding human management often leaves
a mark on the geographical distribution of the
genetic diversity of trees (Vendramin et al., 2008;
Shepard and Ramirez, 2011; Thomas et al., 2012).
Archaeological knowledge about historical tree
use and management can provide a valuable entry
point to understanding of contemporary patterns
in inter- and intraspecific diversity patterns of
trees (e.g. Chepstow-Lusty and Jonsson, 2000;
Goldstein, Castillo Vera and Pay-Pay, 2012).
Risk management
Throughout human history, traditional societies
have been in the firing line of environmental
and climate change. They are aware of the
need to monitor environmental change, often
through the use of indicator plants; an example
is Barbaceniopsis boliviensis, a Bolivian Andes
plant whose leaves are said to turn yellow as
an early warning sign to predict rain (Thomas,
2009). Traditional societies have developed a
plethora of risk management strategies to cope
with the adverse impacts of environmental
fluctuations. Most of the strategies are designed
to make opportunistic use of space, natural
resources, social relations and time. People tend
to invest in a diverse portfolio of options, which
lowers vulnerability and increases resilience and
stability by ensuring the availability and supply of
livelihood goods and services (Frison, Cherfas and
Hodgkin, 2011). A popular strategy is to maximize
the accessibility and use of different ecosystems
in the landscape where people can grow a variety
of different plants and/or extract plant and
animal resources for their livelihoods (Berkes and
Folke, 1994; Ladio and Lozada, 2004; Thomas et
al., 2009). This explains why indigenous groups
are drawn to environments with high ecological
variation, such as ecological edges (Turner,
Davidson-Hunt and O’Flaherty, 2003). Biological
and ecological diversification strategies imply
the need for diversified knowledge, not only
about the ecological conditions of different
environments, but also about a high number
of biotic elements, their useful traits and
management. Risk strategies based on optimal
use of natural resources spread risk in terms of
space and resources; crop failure in one ecological
zone may be offset by more stable harvest in
another, and reduced availability of one biotic
resource may be compensated by use of a variety
of alternative resources.
Traditional societies often complement their
biological and ecological diversification strategies
with equally important social risk-management
strategies (van Oudenhoven, Mijatovic and
Eyzaguirre, 2011) (see Box 7.1). Smith et al.
(2012) recently suggested that resilience entails a
dynamic social process determined in part by the
ability of communities to act collectively and solve
common problems. Systems that spread risk and
innovation in social space, either consciously or
unconsciously, simultaneously stimulate further
diversification of available resources and provide
alternative options in case of unforeseen events
(e.g. poor harvest). A good example is the high
variation in germplasm commonly seen in home
gardens in rural communities, with different tree
species, varieties or genotypes generally occurring
at low densities and frequencies (Padoch and
De Jong, 1991; Ban and Coomes, 2004; PerraultArchambault and Coomes, 2008; Jarvis et al.,
2008; Thomas and Van Damme, 2010; Wezel and
Ohl, 2005; Hmimsa, Aumeeruddy-Thomas and
Ater, 2012). A similar pattern is observed at larger
scales, across the home gardens of different
villages and in predominantly agricultural
landscapes (Dawson et al., 2013). For example,
Van den Eynden (2004) reported that of 214 edible
plants used or known in 42 villages investigated
in southern Ecuador, about 60 percent were used
in one village only. Guarino and Hoogendijk
(2004) postulated that because populations of
81
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
many, if not most, species in home gardens are
often small, they are prone to genetic drift and
rapid genetic divergence or differentiation. This
likelihood, together with the experimentation of
individual garden owners selecting for particular
traits of plant species according to their own
interests, may explain the high intraspecific
variability found in home gardens. Regardless
of the biological and human factors responsible
for the high variability of germplasm found
in different gardens, it is clear that within the
social system uniting different home gardens or
villages, social bonds, relations and interactions
are crucial in allowing individuals to benefit
from the system’s diversification strengths. For
example, it has been suggested that the number
of plant species or varieties found in gardens is
positively related to their owners’ opportunities
for exchange of germplasm through social and
kin networks (Ban and Coomes, 2004; PerraultArchambault and Coomes, 2008).
Box 7.1
Adapted social structures underlie
resilient societies
The success of some ancient societies can at least
partly be related to their social structures. For
example, the rise of the pre-Columbian Inca society
has been related to its verticality, specialization
and reciprocity. Verticality refers to the extraction
of resources and production of goods from multiple
ecological zones along steep mountainsides,
while reciprocity refers to the exchange of these
resources and goods for those produced by people
inhabiting other ecological zones (Murra, 1975).
These characteristics are at the base of Andean
people’s extraordinary knowledge and use of microenvironments, a great range of technological and
agricultural innovations, and formalized systems of
reciprocity (e.g. exchange of labour or agricultural
produce) (Alberti and Mayer, 1974).
82
Preserving traditional knowledge
From the above it is clear that traditional
societies are in many cases the creators and
keepers of an often remarkably diverse and
untapped repository of tree germplasm in
varying stages of domestication that is spread
out over ecological and social space. Traditional
diversification strategies represent a large latent
potential which modern society could exploit
to address human development needs, not
only in terms of tangible resources, but also in
terms of social and ecological management and
organization.
It should also be noted that traditional
knowledge is not evenly distributed across or
within indigenous and local communities, but
is known to vary with ethnicity, age, gender,
social status and numerous other possible factors
(e.g. Thomas, 2012). Who holds the knowledge
about certain tree species also depends on where
they grow in the landscape; for example, home
gardens are generally the domain of women,
whereas men are often more knowledgeable
about trees in the forest. These disparities have
important implications when planning research
with traditional knowledge holders: It is not only
important to identify and work with people that
hold the most knowledge on a topic of interest,
but also to include as many people as possible in
order to be able to access the full spectrum of
often complementary bits of knowledge. Another
important aspect of traditional knowledge is that
it is highly dynamic and adaptive, depending on
the context of use. Indeed practical knowledge
is kept alive, at least in part, through its use. If
the plant is no longer required for a particular
use, related knowledge is likely to disappear
eventually. Indeed as traditional lifestyles become
modernized, people tend to replace traditional
knowledge and plant use with modern knowledge
and/or practices. Hence, unless deliberate efforts
are made to retain knowledge of plant uses that
are no longer applied by or relevant for a society,
whether through written or oral transmission,
the knowledge will be lost.
CU RREN T A ND EMERG I NG TEC HNOLOG IES
Classical tree improvement
Methods from crop and livestock breeding have
been adapted to accommodate the peculiarities
of forest trees. Unlike agronomic crops and
livestock, most forest tree species have not
been domesticated, so the starting point for
tree breeders is typically different from that of
crop or animal breeders. Tree breeders usually
begin working with wild populations instead
of varieties or breeds. This means that the type
of knowledge required and generated through
research differs significantly from that required
for agricultural species.
In classical tree breeding, desired phenotypes
are selected in the wild (plus-tree selection)
and propagules are collected; typically seed
for progeny testing and scions or cuttings for
establishment of seed orchards. Quantitative
genetic data are generated from phenotypic
measurements taken under uniform environmental conditions in a series of progeny or
clonal trials. Statistical analyses are employed
to separate genetic from environmental sources
of variation in measured traits. The ratio of
genetic to phenotypic (including environmental)
variation provides a measure of the heritability of
the trait, and thus the potential for improvement.
Types of knowledge that may be obtained from
field trials associated with tree improvement
include:
• genetic variability and heritability of
traits related to growth, product quality
and quantity, and resistance or tolerance
to insect pests, diseases and adverse
environmental conditions;
• genetic correlations between traits;
• epigenetic effects;
• genotype × environment interactions;
• trait-specific juvenile-mature correlations;
• gene action (additive, dominance) and in
some cases, estimates of numbers of genes
involved in traits of interest.
Tree improvement has existed for many
hundreds of years. However, tree improvement
using genetic theory has been under way for
only a little more than a century (Box 7.2), and
intensive selection and breeding for wood
products, only since the 1930s, beginning
in Austria, Denmark, Germany, Italy, the
Netherlands and Sweden (Hitt, 1952). Methods
for provenance trials to identify the best seed
sources and progeny trials to estimate additive
genetic variation and heritability of valuable
traits were adopted by the mid-twentieth century
in Europe, North America (McKeand et al., 2007),
Australia and New Zealand (Burdon, Carson and
Shelbourne, 2008). Thousands of trials were
established, leading to rapid improvement of
growth rate and stem form and associated broadscale planting of commercial tree species such as
Picea abies, Picea sitchensis, Pinus contorta, Pinus
elliottii, Pseudotsuga menziesii, Pinus pinaster,
Pinus radiata and Pinus taeda. Over the course
of selection, testing, and breeding for improved
traits, a body of quantitative genetic knowledge
has been accumulated for the major plantation
species.
Knowledge gained from early tree improvement trials led to valuable insights about patterns
and extent of genetic variation in forest trees.
Early studies in New Zealand and Australia on
Pinus radiata, for example, demonstrated that
although the species’ native range was small
(Rogers, 2002) and the species did not show
much variation or promising traits in the wild,
there was sufficient heritable variation among
the planted trees to make strong and rapid
improvement in traits of commercial importance
(Burdon, Carson and Shelbourne, 2008). Genetic
gains of as much as 33 percent were reported for
volume at age 15 years from the first generation
of selection of Pinus radiata in New Zealand;
furthermore, a literature review revealed the
average heritability for stem straightness and for
branch angle to be about 0.25 and for branch size
to be just over 0.50, all traits that are important
for wood quality – results indicating potential for
rapid improvement (Wu et al., 2007). Although
populations in North America are small, they are
genetically isolated, and variation is therefore
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Box 7.2
Research organizations historically active in work on forest genetics
The International Union of Forest Research
Organizations (IUFRO) has developed well defined
standards for work on FGR through its working
groups involving national forest research agencies
from many different countries (but predominately
from Europe and North America in the first half of
the twentieth century). Indeed the IUFRO provenance
trials of Pinus silvestris initiated more than 100 years
ago were at the genesis of international action on
FGR.
The work of FAO and its Panel of Experts on Forest
Gene Resources, commencing in 1969, has been
pivotal in developing a globally shared understanding
and appreciation of and modus operandi for FGR
conservation and management.
National forest agencies and institutes in
developed countries, with backing from their
governments, have assisted FGR work in tropical
regions of Africa, Asia and Latin America; examples
include the Commonwealth Scientific and Industrial
Research Organization (CSIRO) Australian Tree Seed
Centre, the Danish International Development Agency
(DANIDA) Forest Seed Centre, the Centre technique
forestier tropical (France) and the Oxford Forestry
Research Institute (United Kingdom).
During the early 1990s, forestry research was
incorporated into the Consultative Group on
International Agricultural Research (CGIAR) with
three centres − Bioversity International (formerly
the International Plant Genetic Resources Institute),
the World Agroforestry Centre (ICRAF, formerly the
International Centre for Research in Agroforestry)
and the Center for International Forestry Research
(CIFOR) − subsequently contributing in significant
and complementary ways to FGR research and
development.
higher than would be likely if the populations
were contiguous.
In spite of the great number of tree species
(80 000 to 100 000) and numerous large
international efforts to generate genetic data,
only a tiny fraction of tree species have been
thoroughly studied for breeding purposes, mainly
temperate conifers and some Eucalyptus species.
In addition a limited number of traits have been
studied, mainly growth-related traits, focused on
increasing production of wood. Considering what
has been learned about trait variation in many
tree species, it is clear that a huge untapped
potential exists for improving product quantity
and quality as well as adaptive traits of many tree
species. This potential is especially important in
light of climate change.
for different locations (Mátyás, 1994). The main
objective was to evaluate which sources had the
best performance, usually measured as survival and
growth under particular growing conditions, often
as the first step in a breeding programme. The first
provenance tests were in plantations established
in France in 1745−1755 by Duhamel, the Inspector
General of the French Navy, to compare Pinus
sylvestris seed sources from the Baltic region,
the present Russian Federation, Central Europe
and Scotland (United Kingdom) in the quest for
suitable mast material for naval vessels (Langlet,
1971). It was already known that planting material
from different sources performed differently at a
given planting site. Unfortunately, the results of
this pioneering experiment did not provide lasting
knowledge because they were not written up and
published. Since then, hundreds of thousands
of trees have been planted in provenance trials
around the world.
Testing provenances involves collecting seed
(or other propagules) from populations of trees
Provenance trials
Provenance trials were originally conceived more
than 250 years ago to identify species-specific
sources of planting material that are suitable
84
CU RREN T A ND EMERG I NG TEC HNOLOG IES
covering a range of environments and growing
them together in experimental field trials
(common garden tests); ideally, a series of trials
is established to cover the range of environments
where the species occurs or may be planted.
Although many provenance tests have not been
intended to characterize adaptive traits, survival
and growth are basic measures of the adaptation
of a provenance to the sites where it is planted
(Mátyás, 1994). Ying and Yanchuk (2006) argued
that height growth is generally a valid surrogate
for fitness, noting that this trait responds quickly
to changing environmental conditions.
An important use of provenance trial results
is the definition of seed zones (or zones of
provenance) (Ying and Yanchuk, 2006). Seed zones
are geographic areas within which seedlings of
the tested species, sourced from anywhere within
the zone, can be planted without loss of local
adaptation. For a given species, a provenance
trial usually includes trees grown from seed
collections from ten or ideally more trees at
each of several or many geographic locations;
however, the planting material may vary from
bulk collections from many trees per population
to individual tree collections, which would allow
combined progeny/provenance testing. Planting
sites may be within or outside the species’ range.
The field trials are usually established following a
randomized block design at several locations to
test performance under different environmental
conditions and to allow assessments of genotype
× environment interactions (Sáenz-Romero et al.,
2011). Provenance trials provide population-level
information intended to identify the sensitivity of
seed sources to varying environmental conditions.
Provenance trials can convey knowledge of:
• intraspecific quantitative variation in
survival and growth among provenances
across variable environmental conditions;
• the degree to which adaptation tracks
environmental gradients;
• existence and type of genotype ×
environment interactions;
• probable responses of different
provenances to climate change.
Many provenance trials could yield much more
information than is often measured or published;
nevertheless, the scientific literature is rich with
published results from such trials. Examples from
many countries are given in Part 4, Chapter 11.
An early example of this type of research,
undertaken in the 1960s and 1970s under
the auspices of FAO, was the investigation of
provenance variation in Eucalyptus camaldulensis,
the most widely naturally distributed Eucalyptus
species and one of the most widely planted
trees in the world (nominated by 17 countries
as a priority species in their country reports,
second only to Tectona grandis with 20 priority
listings). The research demonstrated the
substantial and economically important variation
in performance of different provenances or
seed sources, depending on their origin and
the environmental (climate and soil) conditions
under which the different trials were conducted.
Series of international provenance trials of Acacia
spp., Eucalyptus spp., Gmelina arborea, Pinus
spp. and Tectona grandis, arid-zone species, and
more recently Azadirachta indica and Casuarina
spp. have been remarkably successful, making
immense scientific contributions and leading
directly to or contributing to major forest
plantation developments in the developing
tropics. Hundreds of tree species and provenances
(including many Eucalyptus spp.) were introduced
in multi-location arboreta in African countries
to assess adaptation and potential use. These
arboreta now serve as ex situ conservation stands
in some countries.
Evolution of provenance testing. Globally,
provenance testing is often done outside
species’ native ranges to identify sources of
planting material for commercial forestry.
Understanding of growth parameters of exotic
provenances
performing
under
particular
environmental conditions can also be useful in
matching provenances of those species to novel
conditions resulting from climate change, within
the natural range of the species. Thus applying
the information from provenance trials, where
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
a species is exotic, to restoration of harsh sites
within the species’ natural range is an unplanned
potential value of the trials.
During the past two decades, the establishment
of new provenance tests has declined because
large field trials were generally considered too
expensive, slow to produce results and difficult to
maintain. The days of collecting large quantities
of seed from scores or hundreds of provenances
and planting them in dozens of long-term field
trials may be finished, with a few exceptions –
for example, the project REINFFORCE (IEFC, n.d.)
recently established provenance tests along the
Atlantic shores of Europe from southern Portugal
to northern Scotland (United Kingdom) to test
for local adaptation and plasticity under climate
change.
New approaches have emerged that provide
useful information at lower cost. The United
States of America, for example, reported that in
the northwest of the country most provenance
testing is done through short-term commongarden tests to map genetic variation across
the landscape. These tests are carried out in
nursery environments where site conditions are
uniform, so the observed variation in adaptive
traits such as growth rate, phenology, form and
cold and drought tolerance across the landscape
is known to be due to their genetic composition
or epigenetic effects. If the pattern of genetic
variation tracks the physiographic or climatic
characteristics of the seed-source locations, it
provides evidence of natural selection and may
be important for adaptation.
Genetic field trials have given way to the
sometimes unfulfilled promise of laboratorybased molecular analyses. In recent years it has
been increasingly recognized that the knowledge
gleaned from provenance trials cannot be
substituted by molecular data, and that the gap
between molecular and quantitative knowledge
is especially important in understanding adaptive
traits. A resurgence of interest in field trials is also
explained in large part by recognition of their
value for predicting responses and designing
strategies to counter impacts of climate change,
86
which has led to several attempts to capture
data before they are lost as scientists retire. The
Center for Forest Provenance Data in Oregon,
United States of America, is one such effort (see
cenforgen.forestry.oregonstate.edu); it invites
researchers to submit and use provenance
trial data. Although this initiative is intended
to have global reach, to date only researchers
from western North America have been taking
advantage of the opportunity to house data there.
Data have been uploaded from 25 studies on
eight North American tree species. In Europe, the
European Union (EU) is sponsoring TreeBreedEx
(http://treebreedex.eu) and Trees4Future (www.
trees4future.eu) as components of a projected
virtual EU tree-breeding infrastructure; databases
from genetic field tests (not just provenance
trials) are a main component. However, there too,
few data have been submitted so far.
Use of provenance trials to predict responses to
climate change and restore degraded sites. As
noted above, multilocation provenance trials that
have been established in the past to determine
appropriate seed sources for production at
specific sites can provide information that is
highly valuable for predicting responses to climate
change. Results of such trials can be coupled with
precipitation and temperature data and time
series to observe adaptive genetic variation across
provenances under specific climate conditions
(Sáenz-Romero, Guzmán-Reyna and Rehfeldt,
2006; Schueler, Kapeller and Konrad, 2012). Such
analyses are useful for assessment of climate
change impact on tree species distributions and
productivity in planted forests and agroforestry
systems under future climate projections
according to global climate models (SáenzRomero, Guzmán-Reyna and Rehfeldt, 2006).
Collection of existing provenance trial data for a
given species will help in carrying out systematic
analyses of its vulnerability to climate change in
specific geographic areas.
Provenance trial survival and height growth
data of subtropical pine species native to Mexico
and Central America have been used to validate
CU RREN T A ND EMERG I NG TEC HNOLOG IES
predicted climate change impact on natural
species distributions for 2050 with environmental
envelope modelling (EEM) (van Zonneveld et
al., 2009a). Survival data have been used to
calibrate EEM to make more realistic predictions
(Benito Garzón et al., 2011). EEM can overpredict
climate change impact on species’ distributions
because species can also survive and perform
well outside their existing climate niche (van
Zonneveld et al., 2009a). The EEM estimations
for tree species improve considerably when
provenance performance data are included in
the analyses (Benito Garzón et al., 2011). Trait
data can be coupled with climate data to develop
empirical productivity models to identify the seed
sources that perform well for desired traits under
expected climate change (Leibing et al., 2013).
Provenance trial results have value beyond
production forestry. They can be useful in
restoration of degraded sites because often
the conditions in an area to be restored are
substantially different from those of surrounding
forest; for example, degraded sites may be more
drought prone, may have depleted soil, and may
lack some species that would normally be part
of a functioning forest ecosystem. Often early
successional or pioneer species are needed as a
first stage to reclaim a forest site. Provenance
trials have been conducted mainly for early to
mid-successional species, which tend to be the
fastest growing and the most easily cultivated;
hence these results are useful if data have
been generated on the response of different
provenances to diverse environmental conditions.
Species whose adaptive traits track environmental
conditions closely will have different results
in a provenance trial from species that show
little variation across large distances. In the first
case, applying the results is a simple matter of
matching planting sites to trial sites and choosing
the provenances that perform best. In the second
case, planting material can be chosen from a
greater breadth of sources.
Mátyás (1996) noted that although seldom
recognized as such, provenance research may
be among the most important contributions of
forestry to biological sciences. Of course, this
is only true in regions where there have been
serious efforts to test provenances. In addition,
provenance of planting material is legislated
for forestry purposes. In restoration projects,
however, there is often no legal requirement to
use site-appropriate provenances.
Changing climate in combination with other
causes of site degradation will likely render
much of the area that could benefit from forest
ecosystem restoration substantially different
from the environment in which local remnant
trees became established. Several approaches
have been recommended for planning of
planting projects that may conserve rare and
threatened species (Ledig et al., 2010) or simply
increase the likelihood that planted material will
be adapted to future climatic conditions (Wang,
O’Neill and Aitken, 2010; Hamann, Gylander and
Chen, 2011; Rehfeldt et al. 1999; Beaulieu, Perron
and Bousquet, 2004; Saenz-Romero, GuzmánReyna and Rehfeldt, 2006); some of these have
been tested and are now being employed.
Wang, O’Neill and Aitken (2010) claim that an
approach called “universal response function”,
which integrates environmental and genetic
variables, is more precise than other approaches
in situations in which provenances or test sites
are fewer than desirable. Each of the methods
described has potential value in the planning
phases of restoration projects, for identifying
suitable sources of planting material, but only in
cases where provenance testing has been carried
out.
Perhaps the most valuable knowledge gained
from provenance testing is that all species are
different in their responses to environmental
variation. The United States of America country
report notes that in the northwestern region of
the country – one of the areas where provenance
testing has been the most comprehensive over the
past half century and more – all conifers exhibit
clinal variation in all or part of their ranges,
but the amount and patterns of variation differ
for each species. Conifer species sampled in the
region also exhibit differences in the geographic
87
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
distance on an elevational gradient needed to
detect seed source differences, as well as in the
climatic distance (number of frost-free days)
needed (Table 7.1). Rehfeldt (1994) described
Pinus contorta and Pseudotsuga menziesii as
specialists because their populations appear to be
adapted to relatively narrow niches. The opposite
is true for two generalist species, Thuja plicata
and Pinus monticola. In the southeastern United
States, geographic variation is more complex for
Pinus taeda than for the other southern pines
(Schmidtling, 2001). The fact that evidence has
been documented for clinal and/or ecotypic
variation in all forest tree species examined
indicates the importance of matching seed sources
to environmental conditions. In most cases, local
sources are best adapted to planting sites (Table
7.2), but this may no longer be the rule with
climate change. The evidence that patterns and
amounts of adaptive variation also differ among
all species evaluated implies that there are no
model species and no shortcuts. Some form of
provenance or genecological testing is needed
for all planted tree species.
TABLE 7.1
Amount of environmental difference needed to show a genetic difference in some conifers
Evolutionary mode
Elevation difference to find
genetic difference
(m)
Frost-free days
to find genetic
difference
Pseudotsuga menziesii
200
18
Specialist
Pinus contorta
220
20
Specialist
Picea engelmannii
370
33
Intermediate
Pinus ponderosa
420
38
Intermediate
Larix occidentalis
450
40
Intermediate
Thuja plicata
600
54
Generalist
Pinus monticola
None
90
Generalist
Species
Source: United States country report (from Rehfeldt, 1994).
TABLE 7.2
Evidence from reciprocal transplant studies showing local sources as optimal or near optimal
Family
Genus
Species (reference)
Betulaceae
Alnus
A. rubra (Hamann et al., 2000)
Cupressaceae
Chamaecyparis
C. thyoides (Mylecraine et al., 2005)
Fagaceae
Quercus
Q. rubra (Sork, Stowe and Hochwender, 1993)
Pinaceae
Abies
A. grandis (Xie and Ying, 1993)
Pinaceae
Pinus
P. contorta (Ying and Hunt, 1987; Ying and Liang, 1994; Xie and Ying, 1995;
Wu and Ying, 2004)
P. lambertiana (Harry, Jenkinson and Kinloch, 1983)
P. ponderosa (Squillace and Silen, 1962; Wright, 2007)
P. taeda (Frank, 1951; Wakeley, 1944)
Source: United States country report (from Johnson et al., 2010).
88
CU RREN T A ND EMERG I NG TEC HNOLOG IES
Challenges in classical tree improvement
According to the country reports, more than
700 tree species worldwide are subject to tree
improvement, mostly including provenance and/
or progeny testing. The results of such tests
provide valuable information to determine
sources of planting material that are adapted
for a particular site and the range within which
reproductive material of these species can be
moved without loss of adaptation. In many
cases the full potential of this knowledge is not
realized because:
• provenance trials have not been maintained
and measurements have stopped after an
initial assessment;
• measurement data that have been collected
have not been thoroughly analysed;
• results are not readily available (published
or entered in an electronic database);
• data are lost as scientists retire.
A concerted effort must be made to locate and
use existing information about the species that
have been tested.
Participatory tree domestication
People began selecting and planting trees for
their own purposes thousands of years ago, as
they did with other useful species. However,
impeded by long generation times and the highly
outcrossing mating system of many tree species,
they made little progress towards domestication.
More recently scientists began working with
local people in what is known as participatory
domestication. This collaborative approach was
initially developed in Central Africa with a focus
on domesticating fruit- and nut-tree species
valuable to local people (Leakey et al., 2005). The
objective is to bring together local knowledge
and objectives with scientific knowledge and
theory to speed the process of improvement of
specific traits for particular uses (Tchoundjeu et al.,
2012). The World Agroforestry Centre (ICRAF) has
compiled principles and methods for agroforestry
domestication, in large part involving local
participation (Dawson et al., 2012).
The participatory domestication approach in
Central Africa differs from the tree improvement
methods described above in that multiple species
are typically targeted for improvement at the
same time. The multispecies approach is less
risky for the participating small producers than
banking on a single species (Tchoundjeu et al.,
2012). Progress for a given species is likely to be
slower than for commercially planted species
because the effort is less concentrated and the
range of traits of interest is sometimes greater.
The typical production method for improved
stock is cloning, either by grafting or rooting
cuttings, which allows selection for multiple traits
at the same time (Leakey, 2004).
Most examples of participatory domestication
are in the tropics. In the past two decades the
number of tree species discussed in the literature
on agroforestry domestication has increased from
about 10 to 50 (Leakey et al., 2012). Much of the
recent progress in domestication has combined
local and scientific knowledge. Small-scale village
nurseries have proliferated in West and Central
Africa, in particular, and have become on-farm
enterprises and de facto applied experimental
sites where knowledge about cultivation practices
is acquired for a range of tree species.
89
CU RREN T A ND EMERG I NG TEC HNOLOG IES
Chapter 8
Modern advances
An array of biotechnological tools are contributing
to knowledge of forest genetic resources in
naturally regenerated and planted forests.
For naturally regenerated forest, molecular
markers and genomics are providing important
knowledge on genetic variation within and
between species populations. Biotechnology
further provides important insights into the
nature of the entire tropical forest ecosystem,
including the relationship between forest trees
and the soil microbial communities with which
they interact.
For planted forest, depending on the level of
management intensity and genetic material used,
the biotechnology tools can range from tissue
culture in vegetative propagation to molecular
markers, quantitative trait locus analysis, wholegenome sequencing and genetic modification.
These tools are currently applied for a range of
purposes and involve a varied number of species.
The rapid development of tools (e.g. molecular
markers) for analysing the genetic variability
of forest trees has enabled scientists to better
understand the effects of silvicultural practices on
the long-term evolution of the genetic diversity
of forest trees (Carnus, 2006).
An FAO assessment of biotechnology in forestry
(2004) found that major forest biotechnology
activities had been reported for 142 genera in
over 80 countries, with activities relatively evenly
spread among major categories (Figure 8.1). Of
the over 700 tree species reported by countries
as subject to tree improvement programmes, 241
species were included in biotechnology research.
FIGURE 8.1
Major categories of forest biotechnology
activities
Genetic
modiication
(19%)
Vegetative
propagation or
micropropagation
(34%)
Characterizing
genetic diversity
(26%)
Genomics,
genetic maps and
marker-assisted
selection
(21%)
Source: FAO, 2004.
Population genetics based on
molecular markers
The use of terpenes and especially allozymes
opened new avenues for understanding the
population genetics of forest trees, beginning in
the 1960s (Mitton et al., 1979; Guries and Ledig,
1978). Allozyme analyses were instrumental in
accumulating knowledge on relative amounts
and patterns of genetic diversity, gene flow,
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
inbreeding levels and mating systems for many
tree species (see for example, Hamrick et al.,
1991; Hamrick, Godt and Sherman-Broyles, 1992).
Isozyme analyses have been carried out for
most temperate and many (although far from
most) tropical species, allowing researchers to
test theory and predictions based on the HardyWeinberg principle (i.e. that allelic and genotypic
frequencies remain constant in the absence of
evolutionary influences) with real data. Allozyme
markers are also useful in the conservation of
genetic diversity of forest trees, for example
in designing sampling for conservation, in
quantifying distribution and amount of genetic
variability among and within populations, and
in monitoring changes (reviewed in Millar and
Westfall, 1992).
Many studies have been carried out in Europe
and North America, and fewer in the tropics, to
elucidate the genetic structure of tree populations.
Isozyme analysis of temperate conifers in Mexico,
for example, showed that relict Picea populations
were genetically depauperate, as theory would
predict on the basis of their isolation and small
size (Ledig et al., 1997, 2000; Ledig, Hodgskiss and
Jacob-Cervantes, 2002). Ledig and Conkle (1983)
demonstrated that Pinus torreyana, which occurs
in two small populations in the western United
States of America, had the lowest genetic diversity
of any tree species that had been studied; no
difference was detected within populations, but
several alleles were found only in one of the two
populations. They identified the likely cause to be
genetic drift. Hamrick et al. (1991) and Hamrick,
Godt and Sherman-Broyles (1992) demonstrated
relationships between life history traits and
amount and distribution of genetic diversity.
Duminil et al. (2007) have since demonstrated
that positive correlations among life history traits
and genetic parameters have to be interpreted
with caution when phylogenetic relationships
are not taken into account; nevertheless, the
work of Hamrick and colleagues helped to move
application of population genetics for forest trees
from pure theory to evidence-based hypotheses
92
about patterns of genetic diversity, and it is still
useful in this regard.
Allozymes are usually considered to be neutral
in population genetic analyses, but many of the
enzymes are crucial for metabolic processes, and
the variants are not neutral in all circumstances.
Mitton (1997) described situations in which
variants of particular “housekeeping” enzymes
apparently resulted in selective advantages,
particularly for heterozygous individuals of a
number of species. Such effects were found to
be small but significant in a number of studies
(Bush and Smouse, 1992). DNA markers, which
have a stronger claim to neutrality, gradually
replaced allozymes because they provide direct
information on genetic variation; the number of
markers available can be orders of magnitude
greater than for allozymes, the results may be
more repeatable, and samples are more easily
handled because DNA is more durable than
protein. Although initially the use of DNA
markers was expensive, prone to error and time
consuming, during the past 25 years their use
has dominated population genetic analyses.
Early DNA molecular markers used for forest
trees and especially in tree breeding, such as
restriction fragment length polymorphism (RFLP)
and random amplified polymorphism DNA
(RAPD), had distinct advantages over allozymes,
for example in the vastly greater number of
potential markers (Neale and Williams, 1991;
Neale et al., 1992); however, problems such as
low repeatability of results led to adoption of
other approaches. As more informative but also
more expensive approaches were developed,
microsatellites (simple sequence repeats [SSRs])
became the most frequent markers used in
population genetic studies.
Chloroplast
DNA
markers
have
been
particularly useful in elucidating genetic structure
of populations. Reviewing the use of organelles
compared with nuclear markers, mainly SSRs, in
evaluating population structure of plants (for
138 species, including 37 conifers), Petit et al.
(2005) determined that pollen is generally more
CU RREN T A ND EMERG I NG TEC HNOLOG IES
TABLE 8.1
Indicative studies of tropical tree species using molecular markers since 1990
Region
a
DNA analyses
(No. of species)
Isozyme studies
(No. of species)
No. of studies
Asia
51
22
172
Africa
39a
3
114
Latin America and the Caribbean
63
19
239
Oceania (Australia)
24
7
87
Includes exotics.
important than seed in gene flow and that
although nuclear marker variability is lower in
gymnosperms than angiosperms, the opposite is
true for maternally inherited organelle markers.
For the proportion of total diversity among
versus within populations, no correlation was
found between nuclear and maternally inherited
markers.
Because tropical tree species have been
studied less than temperate species, much less is
known about their population genetic structure
and diversity. Within the tropics, some regions
have had much more attention than others. In
Latin America and the Caribbean, the most well
covered tropical region (in part because of several
major European projects during the 1990s), at
least 239 studies have focused on 63 tree species
since 1990 (Table 8.1). Asia is next with at least
172 studies on about 50 species (including DNA
and isozyme studies). In Africa, 114 studies have
reported on the genetic diversity of about 40 tree
species including a few exotics.
The number and scope of the studies on
tropical forest trees vary greatly by species. Some
relatively important species have been subject to
many studies; for example, at least 21 molecular
studies have been conducted on Araucaria
angustifolia in Latin America and the Caribbean,
and at least 15 studies on Acacia mangium in
Asia. However for most species, even those that
have been the subject of at least one molecular
study, little information is available.
Approaches, techniques and sample sizes
vary widely, from single population studies to
distribution-wide surveys, with markers varying
from a few allozymes to genome scale.
Marker-based understanding of
population genetic processes
Molecular markers have been used to quantify
genetic diversity and its partitioning between
and within populations. They have also been
used to understand population genetic processes
that influence or determine levels and patterns of
genetic diversity, particularly gene flow, genetic
drift and mating systems. The following are
some general results regarding genetic processes
derived from marker studies.
• Gene flow is more influenced by pollen than
by seed (Petit et al., 2005).
• Gene flow covers long distances in northern
and temperate forests where most species
are wind pollinated (Petit and Hampe,
2006).
• Fragmentation can actually lead to greater
gene flow among remnant populations and
breakdown of genetic structure (Young,
Boyle and Brown, 1996), although the
extent varies depending on the population
densities of isolated fragments (Ismail et al.,
2012).
• Gene flow varies with pollinator energy in
tropical species but frequently extends over
several kilometres (Petit and Hampe, 2006).
• Most forest trees are strongly outcrossed
(Petit and Hampe, 2006).
• Mating systems can adapt to accommodate
pollen availability (Lowe et al., 2005).
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
• Many tree species naturally have large
effective population sizes (Petit and Hampe,
2006).
• Genetic drift was detected in small
populations that had been isolated for
many generations (Ledig et al., 1997).
This general knowledge and related speciesspecific information that could only be obtained
through marker analyses has been valuable
for testing hypotheses based on population
genetic theory and for informing management
and conservation decisions. However, because
of the relative ease and availability of these
markers, they have often been overinterpreted
(Holderegger, Kamm and Gugerli, 2006). In
general, neutral markers do not provide reliable
information about genetic variation that is subject
to selection, and they have contributed little
to the understanding of natural selection and
adaptation (González-Martínez, Krutovsky and
Neale, 2006). The need for markers that could be
used beyond selectively neutral processes led to
the development of quantitative trait locus (QTL)
detection and mapping and later to genomic
resources from candidate genes, transcriptomes
and now whole genomes.
Genomic advances
Gene discovery begins with construction of maps,
and this has been more challenging for forest trees
than for agricultural crops because of the large size
of the genome (especially for gymnosperms), the
high heterozygosity of most species, the longevity
of trees and their intolerance to inbreeding
which prevents creation of highly homozygous
lines. The history of genetic mapping for forest
trees began with RFLP, then RAPD and amplified
fragment length polymorphism (AFLP) markers,
all of which were less than ideal, in part because
of their poor reproducibility (Neale and Kremer,
2011). The discovery and development of SSR
markers improved the reproducibility problem,
but their development was too expensive
for broad application for mapping purposes.
Single-nucleotide polymorphism (SNP) markers,
combined with high-throughput technology (using
94
robotics, automated machines and computers for
simultaneous processing of many samples), have
made highly saturated genetic maps possible,
however, and rapid progress has been made
during the past decade. SNP frequency is high
in forest trees, providing unlimited numbers of
markers with no reproducibility problems and at
ever-decreasing cost, so this marker is increasingly
the first choice for many applications.
In genomics as in other genetic technologies,
trees lag behind key agronomic crops, and
tropical tree species have tended to receive
less attention than temperate ones (see e.g.
Neale and Kremer, 2011; Gailing et al., 2009;
Grattapaglia et al., 2009). However, the number
of scientists taking on the challenge of applying
the latest technologies to trees is growing. Neale
and Kremer (2011) noted that despite drawbacks
associated with long generation times, large
genomes, lack of well characterized mutants
for reverse genetic methods, and low funding,
forest biology is now well positioned to make
rapid strides. Gailing et al. (2009) argued that
in using genomic approaches to understanding
the genetic basis of adaptation, forest trees have
some distinct advantages over other plant species
because they generally have high diversity within
populations that are still wild, and forest tree
populations are subject to natural selection.
Genomic advances have opened doors to
understanding the molecular biology of trees.
Next-generation high-throughput sequencing
technologies, in particular, have made the
sequencing and understanding of the large and
complex genomes of tree species an affordable
possibility. Significant accomplishments include
draft genome sequences for several tree species,
most recently Picea abies (Nystedt et al., 2013) and
Picea glauca (Birol et al., 2013), the first conifers
for which such data have been published. Populus
trichocarpa was the first tree for which the entire
genome sequence was published (Tuskan et al.,
2006); its provision of an important reference
genetic map has contributed to its status as a
model species. Myburg et al. (2011) published the
Eucalyptus grandis draft sequence.
CU RREN T A ND EMERG I NG TEC HNOLOG IES
Other tree species for which entire genome
sequencing is under way or nearly completed,
with completed drafts, include Azadirachta
indica, Carica papaya, Castanea mollissima, Citrus
sinensis, Coffea canephora, Larix siberica, Malus
domestica (Velasco et al., 2010), Pinus taeda
(Neale and Kremer, 2011), Pinus lambertiana,
Pinus pinaster, Pinus sylvestris, other Populus spp.,
Prunus persica, Pseudotsuga menziesii, Quercus
robur, Salix purpurea and Theobroma cacao
(Argout et al., 2011). Despite the complexity and
size of conifer genomes, their high economic value
and the investment in such species for commercial
forestry has led to significant sequencing efforts.
Important commodity tree crops, including fruittrees, have received significant attention.
The main factor influencing the choice of
species for sequencing is undoubtedly their
economic value and the importance of identifying
genes for important traits for marker-facilitated
selection, which will reduce time and cost in tree
improvement programmes. However, there are
many knowledge spin-offs. Pavy et al. (2013)
noted that the availability of large SNP databases
allows investigation of polymorphism patterns in
genetically distant species to examine evolutionary
pathways. For example, Nystedt et al. (2013)
obtained evidence that the large genome of
conifers is probably not due to a relatively recent
whole-genome duplication event, but is instead
the result of gradual and continuing genome
expansion over time via the steady accumulation
of long-terminal repeat transposable elements
that are not eliminated, as they are in
angiosperms. They found that whole-genome
duplication probably predated the divergence
between angiosperms and gymnosperms. This is
an example of the knowledge that can be gained
through sequencing of whole genomes.
Knowledge of the genetic basis of
adaptive traits, productive traits and
resistance to pests and diseases
As a result of the combination of rapid advances
and rapidly decreasing costs, genetic maps,
markers and candidate gene sequences have
become available for a range of species, allowing
for investigation of complex questions and the
application of these tools to practical problems in
breeding programmes (Hamanishi and Campbell,
2011).
Much of what is known about genes that
are important for adaptive, productive and
resistance traits was discovered prior to full
genome sequencing, based on genome-wide
analysis techniques such as microarray analysis
(Hamanishi and Campbell, 2011). Genome-wide
techniques can reveal global gene expression
patterns and thus are important for identifying
groups of genes that respond to specific stimuli,
such as drought. Early microarray experiments
used sets of expressed sequence tags (ESTs) from
specific tissues from Populus or Pinus species to
understand gene expression patterns (Sterky
et al., 1998; Hertzberg et al., 2001; Heath et al.,
2002) and identify candidate genes involved in
wood formation and drought tolerance.
For most trees, candidate genes are identified
by transferring information from model species
for which gene function has been elucidated or
by carrying out gene expression studies (GonzálezMartínez, Krutovsky and Neale, 2006). Another
approach is to carry out neutrality tests on
population nucleotide sequence data for individual
genes or groups of genes. Deviation from
neutrality may indicate selection. This approach
has been used to identify genes associated with
stress tolerance and disease resistance.
Next-generation high-throughput sequencing
can shed light on epigenetic effects, which
have profound multigenerational impacts on
plant responses to environmental stimuli such
as drought or cold (Hamanishi and Campbell,
2011). Populus trees vary in their epigenetic
effects, which occur when methylation turns
genes off or on under stress conditions. Using
combinations of EST and SNP markers will help
to elucidate the mechanisms by which DNA
methylation influences gene expression under
stress conditions.
Grattapaglia et al. (2009) noted (and it is
still true) that although recent advances have
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greatly increased the understanding of complex,
interacting mechanisms, it is not yet practical to
apply genomic tools to increase productivity and
growth because of the number of component
traits, each of which is genetically variable.
Combining molecular tools
with tree improvement:
marker-assisted selection
The first applications of molecular genetic
information in tree improvement were for other
purposes than identifying genes responsible
for traits of interest. Early uses of genetic
fingerprinting with molecular markers used in
tree improvement programmes included:
• measuring genetic diversity of breeding
population accessions between indigenous
provenances and naturalized landrace
origins;
• testing paternity contributions to offspring
grown in field tests;
• verifying genetic identity during vegetative
propagation.
As genetic mapping became possible with
molecular markers, interest arose in linking
quantitative traits with markers, launching
numerous QTL projects. For a variety of
reasons – including the rapid decay of linkage
disequilibrium, which requires close association
between markers and genes for traits of
interest (Neale and Kremer, 2011) – QTLs did
not provide the expected insights as quickly as
hoped. Attention turned to other approaches,
including association genetics and diversity
arrays technology (DArT). These approaches are
discussed in this section.
As genomic tools have become increasingly
accessible, in terms of both ease of application
and cost, the bottleneck in linking traits to the
growing knowledge of forest tree genomes has
shifted to the cost and time required for sampling
in the field and phenotyping. There is still a need
to measure the range of phenotypic variation
in traits of interest in order to link the genomic
markers to phenotypic expression.
96
Quantitative trait loci
Quantitatively inherited traits are controlled by
many genes, and most phenotypic traits of interest
in forest trees fall into this category. Each gene
controls a relatively small amount of variance
in a quantitatively inherited trait (Brown et al.,
2003); such genes are known as quantitative trait
loci. Over the past two decades much effort has
gone into identifying linkages between various
molecular markers and QTLs by determining
the number and location of chromosome
regions affecting variation in a trait and finding
statistically significant associations between
markers closely associated with the chromosome
regions and quantitative phenotypic traits in
a segregating population (Brown et al., 2003).
Mapping QTLs allows elucidation of the genetic
structure of complex traits for application to
marker-assisted selection in well studied breeding
populations of tree species that are undergoing
genetic improvement. The main focus of QTL
analysis has been on growth traits, which are the
main target of breeding programmes but have
relatively low heritability and are controlled by
multiple genes (Grattapaglia et al., 2009).
QTL mapping requires highly marker-saturated
linkage maps and phenotypic measurements of
all pedigreed individuals in large segregating
populations (González-Martínez, Krutovsky and
Neale, 2006). Such maps have been successful
in showing the existence of loci accounting for
between 5 and 15 percent of observed variance
(Guevara et al., 2005), in spite of significant
challenges such as the instability of associations
across different environments. In long-lived
organisms the expression of QTL is likely to
change on a seasonal or yearly basis. Factors
influencing the ability to detect QTLs include
sample size, genetic background, environment
and interactions among quantitative gene loci
(Brown et al., 2003).
QTL mapping has been used to generate
genetic maps for many forest trees, and many
QTLs have been identified for drought-related
traits (Hamanishi and Campbell, 2011). Yet
CU RREN T A ND EMERG I NG TEC HNOLOG IES
although this knowledge is useful for tree
breeding, progress is slow because of the time
required for identifying genes in species with
limited genomic sequence availability.
Echt et al. (2011) combined SSR markers
with previously mapped expressed sequence
tag polymorphism (ESTP) and RFLP markers to
produce a map of the loblolly pine genome
that is useful for a variety of population genetic
and germplasm management applications. In
addition, the mapped markers can facilitate
understanding of the evolution of candidate
adaptive trait genes that require unambiguous
identification of parental and clonal genotypes.
Association genetics approaches
Association mapping was developed to overcome
problems with QTL mapping experiments. It uses
linkage disequilibrium mapping to understand
the genetic basis of complex traits, relying on
the association between complex traits and
chromosome regions containing genetic markers.
Unlike QTL mapping, searching for loci using
association genetic approaches does not depend
on pedigreed families. Association genetics uses
linkage disequilibrium between a phenotypic
trait and markers, and it can be applied for any
large sample of a natural segregating population
(Ingvarsson and Street, 2011). It requires many
more markers than have typically been used
in QTL mapping, but the rapid development of
sequencing has made it possible; the hundreds of
thousands of SNP markers needed for association
mapping can be generated rapidly and relatively
inexpensively from any species. The bottlenecks
now are related to the fieldwork required, first to
collect enough samples to ensure that associations
are robust and second to phenotype the sampled
populations. Ideally, replicated field trials will
provide the most precise phenotype information,
but time and cost constitute serious impediments.
Association genetics may start with candidate
genes or a genome-wide approach. Using
candidate genes as markers, individual alleles can
be found that are involved in controlling traits.
Neale and Kremer (2011) summarized association
genetic studies using candidate genes for stem
growth, wood quality, pest resistance, bud
phenology, cold hardiness and drought tolerance
in six genera of forest trees including two conifers
and four angiosperms. Gonzalez-Martinez,
Krutovsky and Neale (2006) tested 18 candidate
genes for association with drought tolerance in
Pinus taeda using this approach; all but two were
found to be selectively neutral. A drawback of
using the candidate gene approach is that as with
QTLs, individual associations between markers
and traits account for only very small proportions
of the genetic variation.
An association mapping approach that has
the potential to be very useful for tree breeding
strategies in the future is the whole-genome scan
(Hamanishi and Campbell, 2011). For example,
in Picea glauca, Namroud et al. (2008) identified
SNPs in expressed genes and used them as
genetic markers for mapping purposes to identify
potential associations between local adaptation
of candidate genes and phenotypic attributes
of populations. This approach can be used for
identifying genes under potential selection for
drought tolerance and other adaptive traits in
non-model tree species.
DArT methods
Intragenus species transferability of markers is
highly desirable, and new methodology known as
diversity arrays technology (DArT) offers exciting
prospects for rapid genome-wide screening
of thousands of polymorphisms (Petroli et al.,
2012) across related species. Because the markers
are gene based, they are useful for genomic
selection and can dramatically enhance breeding
and improvement of forest trees that have been
subject to intensive genomic study (Grattapaglia
and Kirst, 2008).
Among uses that Steane et al. (2011) noted
for DArT markers in Eucalyptus spp. (see Box
8.1) are species differentiation, identification
of interspecific hybrids, and resolution of
biogeographic disjunctions within species.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Box 8.1
Use of genomic tools in
Eucalyptus spp.
Some of the most rapid advances in application of
genomic tools to breeding are focused on Eucalyptus
species. Faria et al. (2010) reported development
of 20 microsatellite markers from ESTs which are
fully transferable across six Eucalyptus species. They
predicted that the usefulness of the markers would
extend to all 300+ species in the subgenus, and they
noted that the markers provide excellent resolution
and potential for use in breeding.
Steane et al. (2011) reported the use of several
Eucalyptus species to create more than 8 000
DArT markers which were used for high-resolution
population genetic and phylogenetic studies.
Resende et al. (2012a) used more than 3 000
DArT markers to evaluate the efficiency and
accuracy of genomic selection in two unrelated
Eucalyptus breeding populations; they reported
that for growth and wood quality traits they were
able to match accuracies attained by conventional
phenotypic selection. However, they cautioned
that in spite of the potential for the approach to
revolutionize tree improvement, experimental
support is required, and in the short term it is
likely that predictive models will be population
specific. Resende et al. (2012b) applied the same
approach to Pinus taeda populations across
multiple ages and found high accuracies across
environments within a given breeding zone, but
not when models generated at early ages were
used to predict phenotype at age six.
Genetic modiication
Genetic modification (GM) of forest trees poses
challenges both biologically and in terms of
public acceptance and policy. Walter and Menzies
(2010) reported that activities related to genetic
modification of forest trees are taking place in at
least 35 countries, and 16 of them have field trials
which are generally small and of short duration.
98
The first genetically modified tree, a poplar, was
produced more than 25 years ago (Fillatti et al.,
1987).
Gene transfer is being tested in many forest
species undergoing intensive breeding activities;
Carnus et al. (2006) reported 24 tree species
involved in GM experimental plantations around
the world. Species frequently mentioned are
Eucalyptus spp., Picea abies, Pinus radiata,
Pinus sylvestris and Populus spp. However,
the number of tree species that have been
successfully transformed remains low, especially
among conifers, and transgenic plants have only
been recovered from a small proportion of the
genotypes in which this has been attempted
(Meilan, Huang and Pilate, 2010). Traits targeted
by transgenic experiments include pest, herbicide
and abiotic stress resistance, hormone regulation,
lignin and cell wall biosynthesis and growth
(McDonnell et al., 2010). Among forest tree
species, by far the most numerous transgenic
experiments have been conducted on Populus
species and hybrids.
Except for a few hundred hectares of
genetically modified Populus species planted
in China, no commercial planting has been
reported.
However,
genetic
modification
protocols have been developed and tested for
traits such as stem shape, herbicide resistance,
flowering characteristics, lignin content and
insect and fungal resistance in many commercially
important planted tree species (McDonnell et al.,
2010) (Table 8.2).
Although most genetic modification is
done with the aim of increasing or improving
wood production, it can also be a tool for conservation, for example in the case of Castanea
dentata (American chestnut). Barakat et al.
(2009) compared canker transcriptomes from
C. dentata and Castanea mollissima (Chinese
chestnut) to identify candidate genes that
may be involved in resistance to Cryphonectria
parasitica (chestnut blight). They identified
several candidate genes for resistance and
gained a better understanding of the resistance
pathway in Chinese chestnut.
CU RREN T A ND EMERG I NG TEC HNOLOG IES
TABLE 8.2
Number of published successful transgenic experiments achieving gene expression or overexpression
in transgenic cells (number of genes used in the various experiments in parentheses, where relevant),
by tree species or genus and by modification objective
Species/genus
Pest
resistance
Herbicide
resistance
Populus spp. and hybrids
8 (>5)
11 (9)
Pinus taeda
1
Pinus radiata
1
Picea glauca
2 (2)
Eucalyptus camaldulensis
Abiotic
stress
resistance
11 (10)
1
1
1
Eucalyptus grandis × urophylla
Hormone
regulation
9 (7)
Lignin
Cell wall
biosynthesis
Growth
26 (10)
6 (6)
8 (7)
1
1
1
1
1
Picea abies
1
Larix decidua
1
1
Larix leptoeuropaea
1
Pinus strobus
1
Source: Summarized from McDonnell et al., 2010.
Transgenic trees are likely to be planted
within crossing distance of wild populations of
the same species. Robledo-Arnuncio, GonzálezMartínez and Smouse (2010) concluded that
it is highly probable that transgenes from
genetically modified trees would move into
conventional forest because of the efficiency of
dispersal systems over the long lifetime of tree
species. Concerns about genetically modified
forest trees dispersing pollen or seed, which
will spread transgenic, selectively advantageous
propagules into natural populations, has led to
a strong focus on sterility mechanisms. Attaining
a stable form of both male and female sterility
in transgenic trees before releasing them is not
a trivial problem. Researchers have encountered
significant obstacles in the search for sterility, but
this work has led to expanded knowledge of the
genetic control of reproductive functions and
floral genomics (reviewed by Brunner et al., 2010).
However, when transformations are intended to
keep a species in the ecosystem (by introducing a
gene conferring resistance to chestnut blight, for
example), the objective is actually to disperse the
new genes into the native population.
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CU RREN T A ND EMERG I NG TEC HNOLOG IES
Chapter 9
Application of
genetic knowledge in
forest conservation
Population genetic knowledge is important
to inform conservation actions. For example,
understanding the distribution of genetic diversity
among and within populations and the degree
to which migration occurs between populations
is necessary for prioritizing populations for
conservation. Many studies have been conducted
to understand mating systems and gene flow
(migration and drift) patterns of forest tree
species. The initial focus was on Europe, North
America and Australia, but more recently the
number of studies targeting tropical species
has steadily increased. There is still considerable
disequilibrium among continents, however, with
more study of tropical American species (e.g.
Ward et al., 2005) than of African and Asian ones.
In any case, the percentage of studied species is
still very low in relation to the high levels of tree
species endemism in tropical regions.
As one of many examples of the growing work
on gene flow within and among populations of
neotropical species, Fuchs and Hamrick (2011)
found that isolated remnant populations of the
endangered tropical tree Guaiacum sanctum
(Zygophyllaceae) maintained high genetic
diversity because of long-distance gene flow,
which indicates that the species has potential to
adapt and expand populations if suitable habitat
is available.
Considering that level of heterozygosity may
be related to fitness, conservation strategies
should be responsive to the heterozygosity of
targeted populations or species. Spielman et al.
(2004) conducted a meta-analysis to compare the
heterozygosity of threatened species with that
of their nearest non-threatened relative. They
found that for paired groups of 15 gymnosperm
species and 6 angiosperms, heterozygosity was
lower in 67 and 81 percent of the threatened
species, respectively. Overall, the difference in
heterozygosity between the threatened and nonthreatened species was on average 35 percent for
both gymnosperms and angiosperms.
Genetic interventions may contribute to in
situ conservation, for example, reintroduction
of lost alleles or gene infusion for genetically
depauperate at-risk populations, or breeding
to introduce resistance to pests or diseases and
allow reintroduction of species to parts of their
range where they have been eliminated.
Conservation actions when species are
threatened by invasive insects or disease
require particular genetic information, such
as range-wide population structure including
distribution of genetic diversity within and
among populations, occurrence of rare alleles
and levels of inbreeding. For example, Potter
et al. (2012) studied Tsuga canadensis, a North
American conifer that is threatened throughout
much of its range by an introduced adelgid, to
inform an ex situ conservation strategy. They
used microsatellite markers to identify locations
of glacial refugia which are of interest because
they typically have high genetic diversity. The
study confirmed a negative relationship between
population isolation and diversity; and a positive
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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relationship between diversity and population
size. This information will be used to refine seed
collection areas to ensure that the patterns of
genetic diversity in the landscape are represented
in the collections, and that areas with high
genetic diversity and unique or rare alleles will be
included.
Research by Fady et al. (2008) on the impact
of natural and anthropomorphic factors on
populations of Cedrus libani, a species of the
eastern Mediterranean mountains that has
long been influenced by human activities,
determined that Lebanon and Turkey constitute
two genetically isolated groups where gene flow
patterns are widely divergent. Whereas gene flow
connects populations in Turkey, differentiation
is strong in Lebanon and populations there
experience significant genetic drift. Using a
combination of chloroplast DNA markers and
allozymes, they showed that human impact does
not easily translate into an identifiable genetic
imprint, although they could identify priority
populations for conservation and appropriate
source populations for assisted gene flow.
Azevedo et al. (2007) reported on the population
genetic structure of an Amazonian tree species,
Manilkara huberi, which is endangered because
of overexploitation for its high-value wood.
Based on seven microsatellite loci, they examined
its mating system and patterns and structure
of genetic diversity, to guide conservation and
management of the species. Manilkara huberi
has limited pollen flow and a highly structured
spatial genetic pattern. The researchers reported
evidence for genetic isolation of populations,
indicating that further fragmentation of
the species’ distribution may result in loss of
subpopulations and their associated genetic
variability. This means that at least several large
populations should be maintained to conserve
the evolutionary potential of the species. The
authors estimated, on the basis of population
genetic parameters, that in order to maintain an
effective population size of 500, seed should be
collected from more than 175 maternal trees.
102
The above examples are but a few of the many
studies carried out over the past three decades
using molecular markers to identify conservation
priorities. Such thorough studies cannot be carried
out for all species that are at risk from increasing
land pressures, overexploitation, climate change
and other causes, but lessons learned in one
species can be applied more broadly.
Genomic approaches will be relevant for
conservation and sustainable management of
natural populations of trees in the near future.
For example, the emerging potential to conduct
association studies in a well defined ecological
and evolutionary context, where correlations
can be estimated between phenotypes and
genotypes at a fine scale (Neale and Kremer,
2011), will facilitate identification of populations
having high conservation value.
Combining spatial analysis with
genetic markers to prioritize
conservation
An understanding of spatial patterns of tree
species genetic diversity can maximize the
effectiveness of in situ conservation strategies
(Petit, El Mousadik and Pons, 1998). Areas of
high genetic diversity should be targets for
in situ conservation, as they are considered
more likely to contain interesting materials
for use and genetic improvement. The recent
development of new powerful molecular tools
that reveal many genome-wide polymorphisms
has created novel opportunities for assessing
genetic diversity. The potential is especially
great when these markers can be linked to key
adaptive traits and are employed in combination
with geospatial methods of geographic and
environmental analysis (e.g. Escudero, Iriondo
and Torres, 2003; Manel et al., 2003; Holderegger
et al., 2010; Chan, Brown and Yoder, 2011).
New methods are now available for prioritizing
populations and geographic areas for in situ
conservation and for monitoring genetic
diversity over time and space, and their use can
improve in situ conservation.
CU RREN T A ND EMERG I NG TEC HNOLOG IES
Geospatial analysis of genetic diversity has
been undertaken for a wide range of tree species
that depend largely on in situ conservation
for maintenance of their genetic resources.
Among recent examples, a geographic gridbased gap analysis for Picea abies in Austria
was used to identify new genetic conservation
units – areas managed specifically for dynamic
in situ conservation of genetic diversity,
maintaining the natural evolutionary processes
– to complement the coverage of mitochondrial
and nuclear molecular marker variation as
well as the adaptive genetic diversity in the
current network of conservation units (Schueler,
Kapeller and Konrad, 2012). In another recent
case study, Prunus africana populations were
prioritized at continental scale on the basis of
nuclear and chloroplast microsatellites, combined
with climate clustering as a proxy for adaptive
variation (Vinceti et al., 2013).
One effective method to describe genetic
diversity in the geographic space is circular
neighbourhood-type analysis. This approach
is especially effective when working with
georeferenced individuals rather than with
populations (van Zonneveld et al., 2012). It has
been used to identify genetic diversity hotspots
for the in situ conservation of a number of
important tree species, including the high-value
timber species Cedrela balansae in northern
Argentina (Soldati et al., 2013), Theobroma cacao
(cacao) in its Latin American centres of origin and
domestication (Thomas et al., 2012), the fruit-tree
Annona cherimola (cherimoya) in the Andes (van
Zonneveld et al., 2012) and Irvingia gabonensis
(bush mango) and Irvingia tenuinucleata in
Central Africa (Lowe et al., 2000).
In addition to these large-scale studies to
map genetic diversity, many other studies have
assessed geographic patterns of genetic diversity
at a smaller scale. Most have been carried out in
temperate and boreal zones, and more studies
are required in the biodiversity-rich tropical
regions (Pautasso, 2009). However, the number of
molecular studies is increasing, even in the tropics.
The results can be used in meta-analyses to detect
overall geographic patterns of genetic diversity
for species with similar life history traits or other
analogies (Conord, Gurevich and Fady, 2012); they
can also be extrapolated to provide conservation
recommendations for other tree species that
share common ecological features but for which
no genetic studies have been carried out.
One approach that can be used to extrapolate
patterns from these analyses and to prioritize areas
for maximum capture of tree genetic resources is
to identify Pleistocene refugia and converging
postglacial migration routes. These areas harbour
high interspecific and intraspecific diversity (Petit
et al., 2003). Georeferenced observation points
from herbaria and gene banks can be used to
predict Pleistocene species distributions on the
basis of past climate data (Waltari et al., 2007).
Such data are freely available from the website
of the Paleoclimate Modelling Intercomparison
Project Phase II (www.pmip2.cnrs-gif.fr), although
they still need to be downscaled. Georeferenced
plant data and climate models are increasingly
available through online platforms such as the
Global Biodiversity Information Facility (www.
gbif.org) and WorldClim (www.worldclim.org),
respectively. These data, where available and
when they are of reasonable quality, can be
fed into environmental envelope models to
predict past species distributions and reconstruct
potential Pleistocene refugia (Waltari et al., 2007;
Thomas et al., 2012). However, it should be noted
that species diversity and genetic diversity are not
always congruent and that broad-scale patterns
of species diversity and endemism cannot always
accurately predict areas of high or threateningly
low genetic diversity (Conord, Gurevich and Fady,
2012).
Research on climate change and
forest genetic resources
Predictions of impacts of climate change on forest
populations and species have galvanized a great
deal of research. The impact of specific climatic
changes on tree species will vary with biological,
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genetic and distributional properties of the species
and of populations within the species. When
confronted with significant climatic changes,
populations of native tree species face three
possible outcomes (Aitken et al., 2008): They may
be extirpated, with resulting loss of unique genes
or gene combinations; they may survive in place,
as a result of phenotypic plasticity, adaptation or
a combination of the two; or they may migrate
following the changing climate to establish in
new locations having climatic conditions for
which they are adapted. However, migration by
seed is likely to be too slow for many tree species
if climate change is rapid. Trees, given their long
generation time, are of particular concern.
Examples of impacts of already changing
climate on tree species and their genetic resources
are adding up but are still not readily available in
the published literature. Based on available data
and deduction, climate change is likely to have
impacts on FGR through several processes which
may include: loss of populations and their unique
genetic variation as a result of extreme climatic
events and regeneration failure, especially at
the receding end of distributions; more severe
pest and disease attack in some areas; altered
fecundity of some tree species; pollination failure
because of asynchronicity between flowers and
pollinators or loss of pollinators; decline or loss
of fire-sensitive species because of increased fire
frequency; changes in competitive relationships
resulting in new species invasions and potential
hybridization (Loo et al., 2011) (see Chapter 5).
Past climate-driven demographic events
have left some signatures in the genomes of
species. Such signatures can be traced back using
molecular markers. Phylogeographic methods
have thus made it possible to retrace the impact
of past climate changes on the evolutionary
demographic history of plants (Hewitt, 2004;
Heuertz et al., 2006; Lowe et al., 2010a; Petit et
al., 2002). Modelling of the impact of past climate
changes on species diversity provides useful
information to predict the future evolution of
species.
104
Spatial modelling using geographic information
system (GIS) mapping tools is increasingly used
to examine vulnerability of genetic resources
to impacts of climate change. For example, van
Zonneveld et al. (2014) modelled the expected
impacts of climate change on populations of
Annona cherimola, a species of the Andean
foothills in Latin America, and mapped their
genetic diversity (van Zonneveld et al., 2012) to
assess vulnerability of the populations. Vinceti et
al. (2013) carried out a similar analysis for Prunus
africana, a widely distributed but ecologically
restricted species found in all of the Afromontane
regions. On the basis of climate models, they
predicted that by 2050, the climate will no longer
be suitable for the species over about half of its
current distribution.
The major challenge facing conservation
genetics is the linkage of traits that are important
for adaptation to changing climates with
molecular markers. Technologies that are being
developed for breeding and improvement are
relevant in this regard, particularly the wholegenome association genetic mapping approaches;
these approaches do not require pedigrees, but
they still require more knowledge of ecological
and phenotypic variation than is currently
available for most species of conservation
concern, especially in the tropics.
Genetic technologies for reducing
illegal logging
Unsustainable and illegal logging is a driver of
deforestation and forest degradation worldwide.
Commercial timber extraction and logging
activities account for more than 70 percent of
forest degradation in Latin America and the
Caribbean and in Asia (Kissinger, Herold and
De Sy, 2012). It is estimated that more than 50
percent of wood exported from the Amazon,
Central Africa, Southeast Asia and the Russian
Federation is illegally harvested, resulting in
annual losses of revenues and assets valuing
between USD 10 billion and 15 billion (Goncalves
et al., 2012). Timber-producing countries will
CU RREN T A ND EMERG I NG TEC HNOLOG IES
continue to lose valuable resources and income
until such unsustainable and illegal practices are
stopped.
Examples of common practices associated with
illegal logging are false declaration of:
• species if harvested wood is from an
endangered species or a species excluded
from legal harvest in a particular country or
region;
• country of origin when harvest of a
particular species is allowed in one country
but not another;
• timber that has been harvested outside of a
concession or inside a protected area.
New policy instruments in Europe, the United
States of America and Australia prohibit the sale
of illegally harvested wood and wood products
and require operators to provide proof of the
identity of the species traded and the origin of
their products. Accurate species identification and
tracking of the geographic origin of timber along
the chain of custody are therefore necessary
to control the flow of illegal wood and wood
products. However, there is a mismatch between
the legislated requirements and the capacity
of importers to comply fully because existing
methods for documenting species identity (wood
anatomy and chemistry) and origin (mostly paperbased documentation, tagging) are insufficient,
ambiguous and easily falsifiable.
Advances over the past decade have made
possible the use of new technologies, based
on DNA markers, to provide timber companies
and timber traders a high level of accuracy
in identifying timber species and origin. DNA
provides a scientific, truly independent and
infallible platform to distinguish species, validate the chain-of-custody documentation and
eliminate fraud.
Significant efforts during the past ten years
have been focused on extracting DNA from wood
samples; it is now feasible to use DNA markers
to complement existing tools, both to identify
species and to track the origin of timber along the
supply chain (Lowe and Cross, 2011) (Table 9.1).
Information obtained from studies carried out
TABLE 9.1
Examples of the use of DNA and markers to control illegal logging
Level of verification
Species
Range
References
Species identity
Multiple species from the Meliaceae
family
Worldwide
Höltken et al., 2012
Declared region or country
of origin
Cedrela odorata
Neotropics
Cavers et al., 2013
Neobalanocarpus heimii
Peninsular Malaysia
Tnah et al., 2009; Tnah et al., 2010
Swietenia macrophylla
Americas
Degen et al., 2013; Lemes et al.,
2003; Novick et al., 2003; Lemes et
al., 2010
Brosimum alicastrum
Central America
Poelchau and Hamrick, 2013
Carapa guianensis, Carapa
surinamensis
Amazonia
Scotti-Saintagne et al., 2013
Shorea spp.
Southeast Asia
Tsumura et al., 2011
Milicia excelsa
Central Africa
Daïnou et al., 2010; Bizoux et al.,
2009
Pterocarpus oficinalis
Caribbean
Muller et al., 2009
Quercus spp.
Europe
Deguilloux, Pemonge and Petit, 2004
Declared concession of origin
Entandrophragma cylindricum
Cameroon
Jolivet and Degen, 2012
Individual log tracking
Intsia palembanica
Papua New Guinea, Indonesia
Lowe et al., 2010b
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
in the development of timber tracking methods
also feed into the growing body of molecular
knowledge of forest trees.
The development of DNA technology offers
great opportunities for management of forest
genetic resources, including:
• better enforcement of forest laws and
regulations by improved verification and
monitoring procedures;
• the development of genetic (and isotopic)
reference databases for tracking traded
timber species;
• improved tools to control the trade of
species protected by the Convention on
International Trade in Endangered Species
of Wild Fauna and Flora (CITES) and species
that could be confused with them;
• transfer of expertise and capacity building
in timber producer and timber transit
countries.
Species identiication
For species that are difficult to distinguish by
comparing morphological or wood anatomical
traits, genetic differentiation remains the most
efficient method for species identification.
The Barcode of Life project (www.barcodeoflife.
org) (Stockle and Hebert, 2008) uses DNA
sequences that vary among but not within species
to differentiate species. The concept has been
more useful for fauna than flora, however; plants
require two or more sequences for confident
identification to the genus or species level.
Barcodes using the two core sequences matK
and rbcL are expected to distinguish at least 50
percent of plant species. For a broad selection
of plant species, the addition of the nuclear
ribosomal DNA internal transcribed spacer (ITS)
sequence increases this proportion to about
80 percent (Hollingsworth, 2011). If needed,
additional specific sequences may be used to
increase the resolution and likelihood of correct
identification for timber species. However, for
the highly degraded DNA of timber, the genetic
species identification needs to be based on short
DNA sequences of less than 200 base pairs. Thus
106
the application of DNA barcoding to timber
needs further development (Höltken et al., 2012).
Core barcodes have been developed for only
about half of the 800 commercial timber species.
Some species for which conventional barcodes
(i.e. commonly used target DNA sequences) have
been developed are represented in the Barcode
of Life database by few individuals sampled
from just a part of the distribution range. For
such species, the intraspecific variation and
the effect of geographical scale of sampling on
DNA barcoding are still problematic (Bergsten
et al., 2012; Lou and Golding, 2012). Additional
sequences are used (e.g. for the mahogany family
[Muellner, Schaefer and Lahaye, 2011]) when
the core barcodes are not universal, have bad
sequence quality and lack discriminatory power
(Hollingsworth, 2011).
Tracking of origin
For control of illegal logging, the ability to
determine the origin of wood is as important
as the identification of species. If spatial genetic
structuring is strong, the geographic origin of
a log may be determined with a high degree
of precision (to within less than 10 km). An
alternative approach is to obtain wood samples
from each standing tree prior to harvest, which
makes it possible to track individual logs by
matching fingerprints, if necessary, at any point
along the supply chain (Lowe, 2010b). A genetic
inventory of high-value trees before felling will
not only be a first step in a chain-of-custody
security system, but will also help avoid felling of
the wrong tree species.
Several studies (e.g. Craft, Owens and Ashley,
2007; Tnah et al., 2009; Degen et al., 2013;
Jolivet and Degen, 2012) have demonstrated the
efficiency of DNA technology in differentiating
trees coming from different locations. Genetic
reference maps to trace the origin of timber are,
or soon will be, available for about 50 species
for at least a portion of their natural range (see
example in Figure 9.1).
With the new-generation sequencing techniques, progress in molecular marker development
CU RREN T A ND EMERG I NG TEC HNOLOG IES
for many species is advancing quickly and costs
are decreasing. The most immediate need is
widespread sampling to cover the range of
as many timber species as possible to have a
complete and robust tracking system.
Forensic testing and analysis
The Global Timber Tracking Network (www.
globaltimbertrackingnetwork.org) led by Bioversity
International is working with scientists and other
stakeholders to define international standards
for genetic labs that will conduct forensic testing.
The network is creating a reference database
of DNA fingerprints for traded species to help
identify species and track the origin of wood and
wood products along the supply chain.
The practical application of forensic DNA
analysis in the timber trade requires adaptation
so the methods will work with wood and
wood product samples that have degraded or
low-quantity DNA. In many timber producing
countries where the risk of illegal logging is
high, law enforcement agencies and national
laboratories are poorly equipped to enforce
forest laws before wood and wood products
are shipped overseas. Therefore the laboratory
procedures must also be simplified for use in small
laboratories, without the need for sequencing or
capillary electrophoresis techniques.
This simplification is possible with the use of
polymerase chain reaction (PCR) RFLPs after SNP
detection and has been successfully developed for
FIGURE 9.1
Genetic reference map for Swietenia macrophylla (mahogany) in Latin America
Mexico
Belize
Honduras
Guatemala
Nicaragua
Panama
Costa Rica
Brazil
Bolivia
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 3
species protected by CITES (Höltken et al., 2012),
but it remains to be done for many commercial
species.
Capacity
building
in
timber-producing
countries, including training in molecular
techniques and ensuring availability of basic
laboratory equipment, is essential to foster
routine use of DNA as a forensic tool.
Existing data are available mainly for timber
species traded in the global market. But timber
108
from illegal logging activities is also used within
producing countries. For example, 85 percent of
the timber production in Brazil is for domestic
consumption. Therefore it is important to invest
also in developing national capacity for regional
and local species (those that are not traded
internationally), to reduce illegal logging within
countries. In general, the level of knowledge is
lower for such species, and data needed to track
wood origin are lacking.
Part 4
STATE OF FOREST
GENETIC RESOURCES
CONSERVATION AND
MANAGEMENT
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Chapter 10
How countries manage
and conserve their
forest genetic resources
The national reports describe a vast array of
actions by countries to recognize, understand,
document, manage, and conserve their FGR
against a backdrop of diverse biological,
environmental, geographic, economic, political,
administrative, social and cultural contexts.
Strategies addressed in the country reports fall
into two main categories. First, there are the
strategies that countries develop at the national
level which set broad overarching directions
for the conservation and management of FGR.
This is the level at which national agendas are
harmonized with regional and international
objectives, and opportunities are identified for
coordination and integration among the various
sectors within government, the economy and
the community that have impact on or inluence
FGR conservation and management. Second,
there are the technical strategies or approaches
that are used to achieve FGR conservation and
management. These may involve in situ, ex situ,
and circa situm conservation, sustainable forest
management
and
community/participatory
approaches to conservation and management
of FGR. Comprehensive and mutually reinforcing
strategies need to be employed across the entire
range of activities relevant to FGR conservation
and management and may be implemented by
the public sector, private sector or community
sector, or any combination of these. Public-sector
strategies include the use of regulations and/or
incentives.
Strategies for FGR conservation and management may include policy and legislation, research,
sustainable forest management, private-sector
planted-forest development, community management of FGR and establishment of genetic
conservation reserves. For certain planted
species, breeding and tree improvement are
central to FGR conservation, to improve the
performance of selected species or forests for
economic, social, community, environmental,
conservation or other purposes; for species that
are not planted, including many providing highvalue timber, management in wild populations
is essential. Countries adopt a suite of different
strategies or approaches that best suit their
particular conservation and management needs,
consistent with the resources available, their
own development goals and the requirements
of their economies, communities and particular
biogeographical and socio-economic contexts.
Countries also noted many and diverse activities
that contribute to maintaining FGR even when
FGR conservation is not consciously or explicitly
identiied as the goal.
Rates of progress have depended on political
understanding and will and on the resources
made available. Collectively the reported actions
describe a developing global movement towards
the conscious stewardship and sustainable use of
these precious resources, as well as the protection
and maintenance of the evolutionary processes
that have produced this irreplaceable legacy. The
country reports represent a vital contribution to
global understanding and appreciation of FGR,
and the identiied strategic priorities will help
guide future international action, led by FAO with
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
guidance from its Panel of Experts on Forest Gene
Resources since 1969 (Palmberg-Lerche, 2007).
Features of effective and
comprehensive FGR conservation
and management systems
Inluences on the effectiveness of a country’s
FGR conservation and management include the
level of economic development, the nature of
the forest resource (natural, managed natural or
planted forest), the nature of the forest industry,
patterns of forest use, patterns of landownership,
the type and quality of governance applied to
the management of forests, biodiversity assets
and natural resources more generally, and the
resources available to undertake the task –
economic, technical, institutional and logistical.
The country reports reveal a number of contextual
features and characteristics that inluence and
shape a country’s system of FGR conservation and
management; the most important are shown in
Box 10.1.
European and North American countries, with
well established formal institutions for natural
resource management and conservation, carry
out and implement the most detailed planning.
A number of key functions and features would
constitute an effective system for conservation
and management of FGR, as follows:
• a national strategy for FGR conservation
and management, with a process for review
and updating;
• national strategies in other related and
relevant areas (e.g. biodiversity, forestry,
agriculture, land use, planning, economic
development) incorporating goals and
objectives of international agreements,
including the Millennium Development
Goals (MDGs), the Convention on Biological
Diversity (CBD) and the World Trade
Organization (WTO), harmonized with
the FGR conservation and management
strategy;
• participation in international agreements
that affect FGR conservation and
management;
112
• legislation enacting the FGR strategy and
international obligations;
• a national programme for FGR conservation
and management to implement the
strategy, with adequately funded
subprogrammes, including a process
for identiication and prioritization
of assets, analysis of constraints and
barriers to effective FGR conservation and
management, and the identiication of
issues, areas, approaches and programmes
likely most eficiently to deliver
improvement in FGR conservation and
management;
• administrative infrastructure and capacity to
implement and administer the strategy and
programmes, with appropriate budgetary
allocations;
• country-wide, comprehensive inventory of
FGR assets in naturally regenerated and
planted forests and in trees outside forests
including agroforestry systems, and in
dedicated FGR programmes and facilities,
including assessment of threats to FGR and
trends in the status of genetic variability;
• a process for prioritization of assets
identiied in the inventory, consistent
with the national FGR conservation
and management strategy, national
development goals and international
agreements;
• established principles and practices for
monitoring, evaluating, reporting, and
improving activities and programmes;
• coordination, including integration and
harmonization of strategies, programmes,
administrations and sectors with relevance
to FGR conservation and management
including forestry, agriculture, biodiversity
conservation, national development,
industry, research and education;
• participation in regional and international
FGR networks and donor programmes;
• consultation with all relevant actors and
sectors of the economy and community in
identiication of priorities and development
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
BOX 10.1
Contextual features that influence a country’s system of FGR conservation and
management
Biogeography
• Biological resources – the nature of the forest
genetic resources, for example, species, breeding
systems, patterns and levels of diversity, and
type, location, and distribution of forests, trees,
species and diversity
Economy, industry and population
• Human population – size, density, growth
trajectory, income, and distribution and location
with respect to FGR
• Level of economic development – infrastructure,
income, poverty level, industrial versus rural
development models, maturity and diversity of
economy
• Economic importance of the forestry sector and
demand for forest products (wood products and
NWFPs) – population, income, level of economic
development, standards of living, expectations,
preferences, use patterns
• Nature of the formal forestry sector – level of
development, type and level of investment,
planted versus natural forests, scale of activities
(industrial versus local production)
• Involvement of the private sector – research,
breeding, planted forest development, use and
management of natural forests
• Nature of land-based industries including
agriculture and forestry – product types,
methods, mechanization
Political system, policy framework and
administration
• Political system – centrally planned or
democratic market economies, relative strength
of the State, provincial, local and community
governments, governance systems
• Legislative and policy systems – maturity,
complexity, efficiency, level of integration
• Development goals and trajectories –
urbanization, industrialization, type of industries
and economic activities desired, employment
• Biodiversity and natural resource management
legislation and policy
• Administrative system – maturity of natural
resource management agencies, degree
of complexity, efficiency, transparency,
accountability
• Attitudes towards forests and forest
conservation, protection of biodiversity
and natural resource management among
politicians, policy-makers, administrators, the
private sector, communities, individuals, media,
and educators
• Patterns of landownership – size of holding,
public, private and communal/traditional
ownership
• System of land use planning and allocation
to different uses, extent to which the State is
willing and can regulate activities and use of
non-State-owned lands
Research and education
• Education and training system – availability and
quality of education for various aspects of FGR
conservation and management, development of
expertise
• Forest research capacity – number and quality of
public, academic and private research institutes
and the level of resources available to them in
key areas of FGR conservation and management
Society and community
• Community traditions, customary law relating to
resource use and conservation
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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of FGR conservation and management
strategies and programmes;
• use of a number of mutually reinforcing
approaches for the FGR conservation and
management strategy, programmes and
implementation.
Approaches to FGR conservation
in relation to biodiversity
conservation strategies
Landscape-level biodiversity conservation focuses
on conserving the whole spectrum of biological
diversity in situ, from genes to organisms through
to ecological processes, and this simultaneously
helps maintain and protect the variability
of trees and forests within those conserved
areas. Global activities to conserve biodiversity
through protected areas and implementation of
the sustainable forest management principles
catalysed by the 1992 Convention on Biodiversity
(CBD) make an immense contribution to the
conservation of genetic diversity in trees and other
woody species, including species of potential
importance or those that are lesser known or
unknown and undescribed. Several countries
(e.g. Estonia) acknowledge the contribution of
these activities, supported by the 1994 Montréal
Process3 and Forest Europe.
While forest research programmes in a number
of countries focus on an ecosystem approach,
several countries note important differences in
their approaches to conservation of biological
diversity and of FGR (summarized in Box 10.2).
For example, biodiversity conservation policy
emphasizes ecosystem and habitat protection
rather than focusing on individual species.
Canada, for example, reports that biodiversity
programmes tend to focus on interspecific
3
The Montréal Process countries are Argentina, Australia,
Canada, Chile, China, Japan, Mexico, New Zealand, the Republic
of Korea, the Russian Federation, the United States of America
and Uruguay. These countries contain 83 percent of the world’s
temperate and boreal forests, 49 percent of the world’s forests
and 33 percent of the world’s population, and they are the source
of 40 percent of the world’s wood production.
114
variation, while for FGR conservation and
management, “changes below the species level
can be critical for ensuring that the adaptive
potential of the species is maintained… this is
particularly important when considering threats
such as climate change, invasive pests and
pathogens, and the ability of species to adapt
to these changing conditions”. The introduction
of external genetic material into forests is one
way to confer adaptability to climate change;
this practice has not been widely considered in
biodiversity conservation outside of enriching
genetically impoverished inbred populations
of threatened species. Furthermore, breeding
and genetic improvement programmes in FGR
conservation and management generally focus
on particular economically important traits,
which is much less common in habitat-oriented
biodiversity conservation. For breeding and
deployment of improved stock for commercial
forestry, genetic variability in some traits will
need to be deliberately reduced to achieve a
consistency in genetic makeup and phenotypic
expression of the desired characters, while for
biodiversity conservation the emphasis is on
maintaining processes (particularly evolutionary
processes) likely to favour maximum diversity.
The management requirements and regulations
for strict protected areas (e.g. conservation
reserves) may preclude some essential FGR
conservation and management activities. FGR
conservation has generally not played an explicit
part in countries’ nature conservation measures,
and in management plans for strict nature
conservation areas it is usually not possible to
include genetic aspects.
Several countries remark that to ensure
adequate attention to FGR conservation and
management in law, policy, budgetary allocations
and management, it is important to communicate
the specific requirements of FGR conservation and
management (where these differ from standard
biodiversity conservation activities) to legislators,
policy-makers, managers and communities
involved in biodiversity conservation and forest
and land management.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
The maintenance of genetic and species diversity
implicit in FGR conservation complements the
habitat and ecosystem protection of biodiversity
conservation. A high degree of integration and
coordination between the two activities, in
terms of strategies and programmes, is essential.
Mutual advancement and reinforcement of FGR
in biodiversity conservation can be promoted
by strengthening their alignment in legislation,
policy, budgets and programme support and by
coordinating complementary activities.
The CBD (UN, 1992) notes in article 8(c)
that countries are required to “regulate or
manage biological resources important for the
conservation of biological diversity whether
within or outside protected areas, with a view
to ensuring their conservation and sustainable
use…”. This requirement highlights the need to
strengthen the contribution of primary forests
and protected areas to in situ conservation of FGR.
FGR conservation and management objectives
need to be explicitly incorporated into national
biodiversity conservation strategies and action
plans, and opportunities for complementarities
between FGR and biodiversity conservation need
to be identified and explored.
Box 10.2
Summary: how FGR conservation approaches differ from usual biodiversity
conservation approaches
• Activities for in situ conservation of FGR can
be integrated into biodiversity conservation
strategies. Most of the differences are in ex
situ conservation and genetic improvement
programmes. FGR conservation focuses on
intraspecific diversity in a smaller number of
economically important or threatened tree
species. The use of living gene banks such as
planted gene conservation stands is greater
in FGR conservation and management;
biodiversity conservation focuses more on in
situ approaches.
• For FGR some in situ conservation requirements
may be satisfied by conservation of a small
number of individuals of the target species in a
small area; this contrasts with the ecosystem,
habitat and landscape-scale protection approach
of most biodiversity conservation efforts.
• Genetic improvement for commercial and
productive outcomes is a major component of
FGR conservation and management, employing
a wide range of technical and financial resources
in activities such as breeding and provenance
and progeny trials, in which the private sector
has a significant role. Conservation of biological
diversity is largely a public-sector activity, as it
involves public goods for which markets are as
yet poorly developed and much of the natural
biological estate is on public lands – although
interest in harnessing private-sector finance for
biodiversity conservation is increasing, including
through NGOs such as the Nature Conservancy
and Conservation International.
• Circa situm conservation is generally regarded
as having a greater role in FGR conservation
and management than in usual biodiversity
conservation. Forest remnants in cleared
agricultural landscapes may be extremely
important as breeding stock. For example, in
parts of Thailand where teak has been almost
completely cleared for agriculture, remnant
trees may contain important genetic variability
adapted to local landscapes. Biodiversity
conservation, however, is increasingly focused
on landscape approaches and on reducing
fragmentation by linking protected areas
with vegetation corridors through agricultural
landscapes.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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National strategies and
programmes for FGR conservation
and management
The strategies for FGR conservation and
management considered in this chapter are
those that address conservation, improvement
or breeding from the biological, ecogeographical
or technical points of view. The country reports
also describe strategies for the setting of policy
objectives, directions, approaches and agendas in
the development of high-level public policy; that
type of strategy is addressed in Chapter 16.
Planning, information and technical input
requirements for effective national FGR
conservation and management programmes
include:
• inventory and characterization of
priority species’ FGR (national, provincial,
population, species or group of species,
ecogeographic surveys and traditional
knowledge) based on technical standards
and protocols;
• information management systems, including
databases and GIS for inventory and
monitoring;
• prioritization for conservation and
management of FGR assets falling
within the programme scope, including
identification of populations at the limit of
their range (see next section);
• in situ FGR conservation and management,
including strategies to identify and promote
FGR conservation in primary forests and
protected areas;
• circa situm FGR conservation and
management, including identification of
options and potentials and development
of methodologies for improved on-farm
management;
• sustainable forest management approaches
to maintain FGR while optimizing
production of goods and services;
• community-based, participatory approaches
to sustainable forest management and FGR
conservation and management, including
technical support for management by
116
indigenous and local communities;
• ex situ FGR conservation and management,
including review of options and promotion
of feasible ex situ strategies and
technologies as a back-up or complement to
other approaches;
• incorporation of gene conservation
objectives into breeding and genetic
improvement programmes;
• development of national seed programmes
to enhance their role in dissemination
of genetically appropriate and improved
germplasm;
• roles for genetically appropriate and
climatically adapted germplasm in
replanting and forest restoration
programmes, including for predicted new
climates;
• review and promotion of appropriate
biotechnologies for FGR conservation and
management;
• regional and international networks to
conserve diversity in priority FGR species
and to provide access to germplasm for
important planted exotics.
Prioritizing species for FGR
conservation and management
Priority setting is fundamental to effective FGR
conservation and management. The types of
value for which FGR require conservation and
management at national and local levels must first
be identified and prioritized, taking into account
international agreements that countries have
signed and ratified. The process for developing
national FGR strategies provides the context for
identifying and prioritizing value and then for
prioritizing species. Prioritizing FGR assets for
conservation, management and improvement
facilitates allocation of scarce resources to the
most important assets and programmes.
Country reports list 2 260 tree species that are
considered national priorities for FGR conservation
and management. They also identify the uses
of the main trees managed for human utility,
including those providing environmental services.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
The prioritization is generally consistent with the
guidance set out in Article 7 of the CBD (UN, 1992)
for information and monitoring, which specifies
“components of biological diversity important
for its conservation and sustainable use, paying
particular attention to those requiring urgent
conservation measures and those which offer
the greatest potential for sustainable use”. The
country reports usually demonstrate a greater
interest in economic outcomes than conservation.
Economic value
Countries cite economic value (including value of
timber, pulp, food, wood energy, and NWFPs) as
one of the main reasons for nominating species for
priority for FGR conservation and management;
economic value accounts for two-thirds of
species nominations, equivalent in importance to
conservation (see below) (Figure 10.1). However,
as the figure shows, some regional differences are
observed.
Countries emphasize tree species suitable for
development of forest industries and planted
forests in their priority listings. Most of these
species are well researched, widely planted,
globalized, industrial forestry species whose
wide appeal lies largely in their proved and
documented ability to perform in a variety of
environmental conditions, the high level of
genetic and performance information available
and the relative ease of obtaining germplasm.
Among the most widely used and prioritized
FIGURE 10.1
Reasons for nominating species for priority for FGR conservation and management (percentage of
species nominations)
%
Economic value 100
90
80
70
60
50
40
Conservation,
including
biodiversity
Socio-cultural
value
30
20
10
0
Invasiveness
Africa
Asia
Oceania
Environmental uses
Latin America
and the Caribbean
Europe
Near East
World
Note: North American countries did not report on reasons for prioritizing species.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
species are Eucalyptus and Pinus species (Figure
10.2). Increasingly planted high-value tropical
trees include Tectona grandis (teak), Swietenia
spp. and Khaya spp. (mahogany), Azadirachta
indica (neem), Dalbergia spp. and Pterocarpus
spp. (rosewoods), Santalum spp. (sandalwood)
and Aquilaria spp.and Gyrinops spp. (agarwood).
Among 1 451 species used in plantations whose
origin was identified, 85 percent were exotic
and only 15 percent native, demonstrating
the paramount importance of exotic, widely
planted, economically important “global”
forestry species. However, the native species may
be underestimated, because some countries,
e.g. Canada, do not report planted forests as
plantations, so most of what they report as
plantation is exotic by definition. In nominating
priority species, many countries cite objectives
such as meeting local demand for timber,
wood products and food, import replacement,
facilitating the development of forest industries,
fostering exports, providing employment, and
providing alternatives to unsustainable or illegal
forest harvesting by rural communities and
others.
Countries tend to prioritize species with
uses and potential applications recognized in
their formal economy and forest sector. India,
for example, notes that in prioritizing species
for ex situ conservation, “the efforts must be
proportional to the present knowledge on the
utility of the species”. However, a heavy emphasis
on a small number of exotic commercial species
important to the formal forest sector entails
a risk of overlooking and underestimating
the contribution of many native tree species
to national well-being, particularly in rural
communities. Where a focus on priority species is
at the expense of exploring the potential of local
trees and conserving their genetic variability,
there is a risk of losing opportunities for
development of highly adapted, productive trees
that are also important ecosystem components.
Several country reports list a myriad of tree
species used for a multitude of purposes in rural
areas, often by many millions of people. The
118
United Republic of Tanzania, for example, notes
that a focus on “charismatic” species may draw
conservation and development effort away from
less recognized indigenous species which help
maintain ecosystems and also display excellent
growth characteristics.
In some instances a discontinuity is seen
between the economic species nominated by
a country for priority in FGR conservation and
management and the country’s reported pattern
of use of trees and forests. For example, wood
energy (fuelwood or charcoal) is a primary forest
value in many developing countries, particularly
in Africa, and is also recorded in Europe.
However, the importance of these uses is not
relected in the country lists of priority species
for FGR conservation and management.
An illustration of the tendency to emphasize
commercial
timber
and
planted
forest
establishment at the expense of other uses and
values is reported by Ghana. Energy use now
dominates demand for wood products in the
country: Wood provides 86 percent of urban
energy and more than 95 percent of energy
consumption in rural areas, and it accounts for 91
percent of roundwood consumption. However,
previous forest policy emphasized high-value
timber species, leading to the establishment of
timber plantations that do not address the wood
energy demand. To date, 260 000 ha of planted
forest have been established under various
government-led programmes. Cedrela odorata,
Gmelina arborea and Tectona grandis constitute
90 percent of the total plantings.
Thus in some countries there appear to be
opportunities for closer alignment of species
prioritized for FGR conservation and management
with patterns of existing domestic demand. In
prioritizing species it is important for countries
to address a wide range of needs in both the
informal and formal sectors. It is also important
to harmonize FGR strategies with other national
objectives such as development goals – potentially
through more direct and wider consultation and
participation of communities in setting priorities
for FGR.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
FIGURE 10.2
Most common priority species, by region
Tectona grandis
Eucalyptus camaldulensis
Populus alba
Picea abies
Leucaena leucocephala
Tamarindus indica
Pinus sylvestris
Casuarina equisetifolia
Quercus robur
Khaya senegalensis
Juglans regia
Swietenia macrophylla
Fraxinus excelsior
Betula pendula
Azadirachta indica
Populus nigra
Pinus caribaea
Gmelina arborea
Eucalyptus grandis
Acacia nilotica
0
5
10
15
20
25
Number of countries reporting the species as a priority for FGR conservation and management
Africa
Asia
Europe
Latin America
and the Caribbean
Conservation and environmental value
As mentioned above, in country tabulations
of priority species nominations, conservation
purposes (biodiversity, threatened species,
endemic species, genetic conservation, scientific
value) account for about 66 percent of the
nominations (Figure 10.1).
Environmental uses, including soil and
water protection, soil fertility and watershed
management, account for 37 percent of species
nominations for priority. A number of countries
Near East
North America
Oceania
include aesthetic, cultural and religious values in
the category of “environmental uses”.
The vast majority of species used for
environmental purposes are native (84 percent)
– a contrast with the prioritized economic
species, of which 85 percent are exotic. Some of
the most frequently reported species used for
environmental purposes include Pinus sylvestris,
Alnus glutinosa, Quercus spp., Betula pendula,
Fraxinus excelsior, Ulmus glabra and Populus alba
mainly in temperate regions, Tamarindus indica,
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
Leucaena leucocephala and Faidherbia albida in
tropical regions and Albizia lebbeck, Cupressus
sempervirens and Casuarina equisetifolia in both
temperate and tropical regions.
It is clear that these figures grossly underestimate both the contribution of forests to
environmental services and the number of tree
species contributing.
Assessment of the rate of genetic erosion and
vulnerabilities of different species, subspecies,
varieties and populations is important for
determining their priority for conservation and
management. Countries vary greatly in the
number of tree species under threat (see Figure
6.4 in Part 2). Canada’s analysis of criteria of
rarity, habitat under threat from alternative uses,
decrease in range and lack of viable seed sources
determined that, although extinction risk is very
low at the species level, populations of 52 percent
of all Canadian tree species require some form of
in situ or ex situ conservation. Biodiverse countries
experiencing high rates of forest loss – e.g. Brazil,
Ecuador, Ethiopia, Indonesia, Madagascar, Papua
New Guinea, the Philippines and the United
Republic of Tanzania – also often report the
presence of large numbers of threatened species.
Social and cultural value
Social, cultural, recreational, ornamental and
gardening purposes are cited as the reasons for
41 percent of species nominations for priority
listing (Figure 10.1). Traditional medicinal uses
are included in this category. Social values may
be especially important in certain countries.
For example, sacred and religious values are
reported as important in Burkina Faso, Ghana,
India and Zimbabwe. Most of the species in this
category are native, relecting the close cultural
afinities for native species that people develop
over millennia, which help to shape national and
cultural identity.
Invasive species
Invasive tree species pose a signiicant threat to
the integrity and conservation of FGR, mainly
through their capacity to transform ecosystems.
120
Small island ecosystems are especially at risk.
Understanding the genetic makeup and variability
of invasive trees and shrubs can be crucial to
developing effective control and management
strategies.
About 500 species were nominated as a
priority for management (mainly by African
and European countries) because of their
invasiveness,
with
implications
for
FGR
conservation and management. Most invasive
tree species have been introduced for ornamental
purposes, although several species introduced
in afforestation or as plantation species have
become seriously invasive. Examples include
Prosopis juliflora in several countries and Acacia
mearnsii, Acacia melanoxylon, Pinus patula and
Populus canescens in Zimbabwe. The priority
given to these species in FGR management
highlights the need to consider the potential
invasiveness of species promoted for planting and
to ensure that this risk is minimized.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Chapter 11
Characterization of genetic
variability and monitoring
of change
Countries
recognize
that
for
effective
conservation and management, it is essential
to understand and describe their FGR at both
broad (geographic) and fine (among and within
forest tree species) scales. Countries characterize
genetic variability for two main purposes. The
first is conservation planning and management,
including sustainable forest management, for
which it may be necessary to:
• identify areas, forests, species or populations
with high levels of variability for genetic
conservation (in situ or ex situ) and/or those
whose variability is at risk;
• identify populations or individuals with rare
alleles for conservation (in situ or ex situ) or
with high levels of variability for enriching
genetically depauperate populations;
• characterize the variability within areas,
forests, populations or stands to guide forest
management and silvicultural practice;
• characterize relationships between genetic
variability and environmental parameters to
establish “genecological” or seed-transfer
zones within which transfer of genetic
materials is considered most appropriate;
• monitor the trend in genetic variability
in species, populations or particular
areas or stands, for example in response
to silvicultural and harvesting regimes,
environmental changes or threats, in
order to help guide conservation and
management.
The second purpose is breeding and improvement, for which it may be necessary to:
• identify species and populations with
the greatest potential for commercial
development;
• characterize desirable productive, service
or adaptive traits in priority species and
relatives for further development;
• identify individual trees with desirable
characteristics for breeding and
improvement;
• identify genetic markers, develop linkage
maps and ascribe function to genes,
especially for characters conferring adaptive
advantage or otherwise desirable traits;
• identify stands and individuals for provision
of propagation materials (seeds and
vegetative materials).
Article 7 of the CBD (UN, 1992) details priorities
for information and data collection; it requires the
identification and monitoring of components of
biological diversity important for its conservation
and sustainable use, as well as processes and
activities that have impact on this value.
Recognition of the economic and ecological
importance of genetic variability in tree species has
steadily increased during the past 60 years and has
driven efforts to study, characterize and document
such variation (described in Chapter 7). A number
of methods are available for characterizing genetic
variability, both quantitative and molecular.
Approaches used by countries for characterizing
variability include investigation of morphological
characteristics and the use of various biochemical
and DNA markers through field-based studies,
provenance and progeny trials, and laboratory-
121
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
based investigations. Use of these methods
varies according to the nature of the country’s
genetic resources, whether the information
sought is at the interspecific or intraspecific
level, the country’s priorities and objectives
(conservation, management, improvement), the
resources and technology available for the task,
the degree of advancement of the country’s
FGR conservation and management system and
the organization undertaking the work. For
example, documentation of raw FGR in large
protected areas may require taxonomic surveys to
provide a broad assessment of variability at the
interspecific level, while for detailed conservation
planning or breeding programmes, information
about the genetic variability between and among
populations, provenances or progeny may be
sought, involving investigation at the molecular
level. Measurement of a variety of parameters
is often needed to provide the information
necessary for conservation and management of
species and their genetic resources.
The different methods of characterization
require different levels and types of resources,
including funding, technical expertise, equipment,
facilities and even land, and they are deployed by
different countries in a manner consistent with
the technical, financial and personnel resources
available. Most methods of characterizing genetic
variability require substantial commitment of
resources, and as the number of tree species and
populations to study is considered impossibly
high, priority must be assigned to a limited
number of the most important or model species.
High-value, widely planted commercial species
are generally accorded the highest priority, as
the economic benefits from improvement can be
substantial: Countries rated commercial value as
the most important factor in prioritizing species,
as discussed in Chapter 10. Economic returns
from improvement programmes also make it
possible to allocate significant resources to the
characterization of the variability of high-value
species – including field investigation, provenance
and progeny trials and investigation of end-
122
product quality considerations (e.g. timber and
pulping properties, charcoal-making properties,
fodder properties, fruit and nut nutrient content
for edible species, medicinal properties and
essential oil profiles).
The focus on economically important species
has produced a great depth of information
on genetic variability, and its inluence on
expression of desirable characteristics, for highvalue globalized species and hybrids such as
Acacia mangium, Acacia nilotica, Cunninghamia
lanceolata, Eucalyptus spp. (E. camaldulensis,
E. grandis and E. globulus), Hevea brasiliensis,
Pseudotsuga menziesii, Pinus spp. (P. caribaea,
P. elliottii, P. massoniana, P. patula, P. radiata,
P. sylvestris and P. taeda), Populus spp. and hybrids
and Tectona grandis, which are among the 30
most widely planted trees in the world (Carle, Ball
and del Lungo, 2009). Most of the species that
have been studied in depth are planted mainly
as exotics (see Chapter 10). Historically, countries
have expended less effort on characterizing their
indigenous FGR, except in cases where the species
are of high economic value and widely planted
(e.g. Cunninghamia lanceolata, Pinus massoniana
and Pinus tabuliformis in China and Pinus taeda
and Pseudotsuga menziesii in North America) or
are threatened and the subject of conservation
management interest.
Where appropriate, countries make use
of existing information sources for their FGR
programmes. For example, some countries with
advanced FGR conservation and management
systems had not undertaken systematic or special
inventories of FGR, rather making use of existing
inventories and databases of forest and biological
resources.
Rare and threatened species requiring
conservation are also considered to have high
priority for characterization of variability. They
have been the focus of genetic investigations,
especially those tree species that are also
commercially valuable (e.g. Eucalyptus benthamii
in Australia, Baillonella toxisperma in Gabon and
Dalbergia cochinchinensis in Thailand).
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Assessment of genetic variability has a vital role
in conservation planning and guides decisionmaking and management. Germany, for example,
remarks, “…knowledge will be used in decisions
on natural and artificial regeneration of forest
stands, in the provenance controls of forest
reproductive material and in the choice of gene
conservation forests”.
Characterizing interspecific
variability
Characterization of diversity at the species level is
a key element and priority for FGR conservation
and management. It involves the identification
of tree species and mapping of their distributions
through nationwide inventories. Environmental
and
biogeographic
information
is
also
important for understanding and interpreting
morphological
differences.
Characterization
of interspecific diversity requires botanical,
taxonomic
and
biogeographic
expertise,
field survey, and GIS and data management
systems for mapping, recording, storing and
sharing information. Much of the genetic
characterization at this level is captured through
biological and forestry inventories undertaken
in the course of other resource management
activities such as biodiversity conservation and
forest management. Such surveys often fail to
capture and document the extensive genetic
resources held in circa situm environments. ICRAF
and national partners in Africa have documented
information for important agroforestry tree
product (AFTP) species through participatory
rural surveys (e.g. Leakey, Schreckenberg and
Tchoundjeu, 2003), and CSIRO’s South Pacific
Regional Initiative on Forest Genetic Resources
(SPRIG) has done likewise in five island nations of
Oceania (e.g. Thaman et al., 2000).
Countries with well established biological and
natural resource management administrations
and infrastructure, including herbaria and well
trained taxonomists, have better knowledge
of their tree species diversity and distribution
than countries with fewer resources, which
have a greater need for further characterization
at the species level. Completion of biological
inventories including tree species resources is
more challenging for countries with extensive
areas of highly diverse tree lora distributed over
a wide range of heterogeneous environments,
particularly if forests are inaccessible and poorly
known or if conlict in or near forest areas raises
security challenges.
Interspeciic diversity reported by countries
varies greatly, from as few as 20 to almost
8 000 species, with major implications for
FGR conservation and management systems.
Countries with more species will generally have
greater reserves of genetic variability, and the
task of documentation and characterization is
therefore more dificult. Country reports suggest
that developed countries with lower interspeciic
variability generally have detailed knowledge
of a higher proportion of their indigenous
tree species, especially where these species are
important commercially.
Where botanical inventories are lacking,
the area of forest or vegetation cover may be
used as an approximate surrogate measure of
genetic variability. As many countries do not
have detailed information on variation among
and within species, and are unlikely to have it
in the near future, forested area is often used as
a primary measure of diversity of FGR and as a
means of monitoring their trajectory or trends
(see also Chapter 6). For example, Ghana quotes
the loss of most of its dry semi-deciduous forest
as representing a serious threat to the genetic
diversity of the woody species contained within
it. Many countries, including Ethiopia, Indonesia,
Madagascar and Thailand, report risk from
signiicant deforestation rates. Madagascar lost
4.5 percent of its forest cover between 2000 and
2010 (FAO, 2010a) and reports a risk that “many
forest species may disappear forever, without
ever being discovered”. Ethiopia recognizes
that “the most important threats to genetic
diversity come from deforestation and forest
fragmentation, which can result in total loss
123
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
of genetic information and disturbance in the
genetic structure”.
Where area of forest cover is used as a
surrogate for diversity, estimates of species
richness and distribution per unit of area are
required to establish the relationship between
forest or tree loss and the loss of diversity that
this represents. Ground truthing of GIS data may
assist in this process.
For countries that lack comprehensive
inventories of their forest species, completion
of inventories is a high priority. For example,
the Madagascar country report states that it
is indispensable to complete knowledge of
the forests and the species they contain by
undertaking new loristic inventories across the
country. For assessing changes in forest cover, the
use of GIS is vital and constitutes one of the most
important methods for broadly characterizing
genetic variability, particularly in the context of
less wealthy, biodiverse nations subject to high
levels of forest loss and degradation. International
donor and partner assistance in forest inventories
can have an important role in assessing diversity
and monitoring forest changes.
Characterizing intraspecific
variation
Environmental heterogeneity, breeding systems,
the degree of biogeographic isolation from
other populations or individuals, and the species
evolutionary history all inluence the pattern and
level of intraspeciic variation (see section on
species diversity in Chapter 1). Its characterization
and documentation is widely appreciated
as a central component of conservation and
management of individual species, including
breeding for economic, genetic conservation
and environmental applications; identifying
provenances and seed-transfer or genecological
zones; and enabling selection of germplasm best
adapted to local conditions for use in planting
programmes.
Reporting countries recognize that a thorough
understanding of intraspeciic variation is funda-
124
mental to the sustainable management of
FGR, including in forests managed for multiple
purposes; it is particularly important for forest
types, species and populations containing
valuable genetic resources of narrow or limited
distribution which need to be monitored at the
intraspeciic level. Despite the great value of
knowledge of intraspeciic genetic variability,
the United States of America notes that, with
current resourcing, there are “too many tree
species” to assess genetic variability effectively
at this level. China has genetic information on
100 species, a very high number compared to
many other countries, but these represent only
about 5 percent of the country’s tree lora –
highlighting the need to prioritize species for
further investigation.
As plantation forestry in most parts of the
world is focused on improvement of a small
number of highly productive commercial species,
genetic characterization has similarly focused
on these species. For example, considerable
information is available on the variability of the
most commercially important species in four
of the most widely planted genera globally:
Acacia, Eucalyptus, Populus and Pinus. Efforts to
characterize species that are less widely planted
but important locally or in naturally regenerated
forests are lagging and in urgent need of study.
Sharing of information on intraspeciic variability is essential for effective FGR conservation
and management, and is particularly important
for developing countries that lack the resources
to undertake studies on the exotic species on
which some countries’ forestry industries depend.
Several key international networks, for example
for the genera Eucalyptus, Pinus, Populus, Salix
and Tectona and for Azadirachta indica (neem),
Bambusa spp. (bamboo) and Calamus spp.
(rattan), are reported by developing countries
as being particularly important for knowledge
sharing. Of like importance is the publication of
public-good FGR research in accessible formats,
such as in free or open-access online journals and
websites.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Methods used by countries to assess
intraspecific genetic variation include well
established techniques such as identification of
morphological differences in the field; provenance
testing; progeny testing; genecological studies to
examine the variation of adaptive traits (i.e. those
that are likely to impart an adaptive advantage)
across the landscape, which may assist in
delineating appropriate seed-transfer zones; and
increasingly laboratory-based approaches based
on biochemical and DNA markers. Each method
has its particular applications and advantages in
different country contexts and applications. Of
the examples for which testing methods were
cited in country reports, DNA markers were used
for 58 percent of the species, biochemical markers
for 38 percent, and studies of morphological
characteristics for 4 percent.
Individual species vary greatly in degree of
variability for different traits, both within and
between populations. A species’ biogeographical
and genecological distribution may be simple
or complex, and it may overlap provincial and
national boundaries, which means that efforts to
characterize the species may require cooperation
between agencies in different jurisdictions and/or
countries.
Countries’ approaches to intraspecific characterization are described briely here.
Use of morphological traits
Morphological evaluation or phenotypic selection
is one of the main characterization methods used
by developing countries, even though its use is
not recorded often in tabulations of countries’
characterization methods.
Morphological characters that may be assessed
include bole form, branching pattern, height,
wood and leaf characteristics and growth traits,
as well as structures that show limited phenotypic
variation, particularly reproductive parts such
as seeds and fruits. The low cost and ease of
morphological assessment relative to laboratorybased approaches has led to its widespread
application. However, observed differences
cannot be attributed to genetic differences with
certainty until further testing is undertaken, most
commonly through provenance and/or progeny
testing (see below), increasingly coupled with
molecular marker studies.
Phenotypically based selections and characters
of trees in wild stands typically have very low
heritabilities, especially for those traits showing
continuous variation. Nevertheless, selected
individual trees or seed stands showing superior
expression of desired traits (“plus trees”) are
widely used for propagation materials for
provenance, progeny or other trials and to
produce seed or nursery stock for production
plantings.
Morphological assessment is sometimes also
used to identify variants and guide selection of
more diverse plant materials for conservation
management when establishing ex situ genetic
conservation stands for threatened species.
However, random sampling of gene pools,
involving a large number of selections (e.g.
>30 trees) of widely spaced and presumably
unrelated individuals, is generally preferred
for both provenance trials and conservation
measures. The irst step in many traditional
breeding programmes is to assess morphological
and growth traits on same-aged plants growing
under common environmental conditions, so that
differences can be attributed to genetic effects
rather than environmental variability, yielding
higher heritabilities.
Provenance, progeny and clonal testing
Provenance testing involves growing trees
selected from different locations (provenances)
in the same ield environment so that observed
variation among populations or individuals
can be attributed to genetic differences. These
approaches have a long history and continue to
be used widely in tree breeding and improvement
programmes (see Chapter 7). The country
reports tabulated species/provenance trials,
often extensive, which have been undertaken.
However, many of these trials are in progress or
125
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
not yet mature, have not been reported or are
not readily available in the published scientific
literature.
The results of provenance tests are typically
only valid after half of the projected rotation
age, and some characteristics of interest may not
express themselves for many years. Accordingly
provenance testing is time consuming and rather
expensive and may be subject to high levels of risk
associated with natural disasters such as drought,
fire, cyclones, and social, political and economic
disruption. However, as provenance testing does
not require high levels of technical infrastructure
or facilities, it is used widely in tropical countries,
where trees often have fast growth rates and
shorter rotation periods, meaning that valid
selections may be obtained as soon as five to ten
years after planting for short-rotation species.
Provenance testing, including genotype ×
environment interaction studies and reciprocal
transplant trials, help identify provenances
adapted to particular environmental conditions,
including climate, drought and fire – an
increasingly important task given the predicted
climate changes which are already having
observable effects. Many molecular marker tests,
in contrast, help evaluate genetic differences
or similarities but cannot necessarily be used to
identify genes conferring adaptive or productive
advantage. Several countries give a priority to
research to identify provenances better adapted
to new climate change-induced conditions. More
generally, the United States of America notes:
“Long-term provenance trials test different
provenance collections over a variety
of planting locations, and… in addition
to documenting intraspecies variation,
can provide reliable information for
determining the limits of seed movement
and discern which seed sources are suitable
for planting locations because they
evaluate seed sources over a long period
of time... the wealth of provenance trials
have demonstrated intraspecific variation
for practically all timber species….”
126
Genecology studies use provenance testing
or environmental surrogates to examine the
variation in adaptive characteristics related to
traits such as growth rate, phenology, form,
cold and drought tolerance across a landscape
gradient to delineate seed-transfer zones.
This approach has been used to define seedtransfer zones for conifers in the southeastern
and northwestern United States, for teak and
Pinus merkusii in Thailand and for forest tree
species in Denmark. It allows the best-adapted
plant materials to be matched with appropriate
locations and environmental conditions.
Trials in temperate regions. Large and well
documented provenance testing efforts include
the International Union of Forest Research
Organizations (IUFRO) Picea abies provenance
trial established in 1968, which comprised 1 100
provenances collected from throughout the
range of the species and planted at 20 locations
in 13 countries (Krutzsch, 1974).
Similarly, Russian provenance trials were
established with Pinus sylvestris from 1974 to 1976
throughout the former Soviet Union, including
113 provenances at 33 planting sites (Shutyaev
and Giertych, 1997).
Canada reports 988 provenance tests,
comprising 7 573 provenances, that have been
established for at least 41 forest tree species and
hybrids. Extensive provenance testing has been
carried out for 34 native species and at least
seven exotics, mostly conifers. In recognition of
their wide planting in reforestation programmes,
six native species have been extensively tested
both nationally and subnationally: Picea glauca,
Picea mariana, Pinus banksiana, Pinus contorta
var. latifolia, Pseudotsuga menziesii var. menziesii
and Tsuga heterophylla.
In the 1950s, an ambitious programme with
Pinus resinosa sent seed to other parts of North
America and Europe for testing, in addition to
establishing many experiments in Ontario. The
programme was abandoned when it became
apparent that genetic variation in Pinus resinosa
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
was lower than in most other conifers (Stiell,
1994). In eastern Canada the primary focus for
early exploration and testing was Picea spp.;
range-wide provenance trials were established
for P. glauca, P. mariana and P. rubens. Provenance
trials including native and exotic Larix species
were also established in eastern Canada. Pinus
contorta provenance trials established in Canada
in 1974 in the province of British Columbia and
in Yukon Territory included 140 provenances from
throughout the species’ range in western North
America, tested at 62 locations (Wang et al., 2006;
Wang, O’Neill and Aitken, 2010).
A test in the southern United States of America
in 1926 included four provenances of Pinus taeda
(Rogers and Ledig, 1996). Wakeley (1955) also
initiated a large study of geographic variation of
four southern Pinus species (P. taeda, P. elliottii,
P. palustris and P. echinata), which were planted
in 66 test plantations across the southeastern
United States in 1951. This study set the stage for
most of the later genetics work with these species
(Zobel, 2005).
In Germany, provenance testing is being carried
out for 34 tree species, including several North
American species, by multiple Länder (states) and
institutions. Germany has used this approach since
as long ago as the second half of the nineteenth
century; the first German provenance test for
Pinus sylvestris was established in 1879.
Bulgaria has 57 provenance trials, focused on
38 tree species.
Trials in other regions. While provenances of
many tree species that are planted for wood
production in Europe, North America and
Australia have been well studied, often both
within and outside their natural ranges, in
general fewer species native to the tropics have
been studied and information from existing trials
is not easy to find. In addition, many tropical
tree species have recalcitrant seeds and lower
sparsely; these characteristics are a constraint
to testing because it is dificult to accumulate
the materials to establish a provenance test in a
single planting season. The following are some of
the known provenance trials in Africa and Asia.
In West Africa, 25 provenances from throughout
the range of the multipurpose species Parkia
biglobosa were established in 1995 at two test
locations in Burkina Faso that receive contrasting
amounts of mean rainfall (Ouedraogo et al.,
2012). Results are now available to guide selection
of planting material for locations in Burkina Faso.
Provenance trials have also been used to test for
resistance of Milicia excelsa to a Phytolyma psyllid
(Ofori, Cobbinah and Appiah-Kwarteng, 2001).
Provenance trials on indigenous species have
also been set up for Aucoumea klaineana; 13
representative provenances from throughout
the species distribution were planted in 1967
in M’Voum reserve, Gabon. In the Republic of
the Congo and in Côte d’Ivoire, provenances of
Terminalia superba were established to select
the more productive lineages for plantations.
Some provenance trials have recently been set
up for Baillonella toxisperma, Distemonanthus
benthamianus and Erythrophleum suaveolens/E.
ivorense in different forest gaps in Gabon and
Cameroon on each side of the Equator (J.-L.
Doucet, personal communication). Provenances
come from different populations from each part
of the climatic hinge (seasonal inversion line) in
Cameroon, Gabon and the Republic of the Congo.
The country reports mention provenance
trials with at least eight other species in West
Africa: Acacia senegalensis, Adansonia digitata,
Allanblackia parviflora, Khaya senegalensis,
Tamarinda
indica,
Terminalia
ivoriensis,
Triplochiton scleroxylon and Vitellaria paradoxa.
Other regions of Africa report fewer tested
native species. Substantial effort has been
expended on testing provenances and developing
tree improvement programmes in southern
Africa, but most of the plantation species are
exotics. In Madagascar, provenance trials have
been undertaken for important and promising
forest plantation species, mainly exotics: Acacia
spp., Cupressus lusitanica, Eucalyptus spp., Khaya
madagascariensis, Liquidambar styraciflua, Pinus
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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spp. and Tectona grandis. Zimbabwe’s provenance
testing programmes for indigenous species date
back to the 1980s, and also form the basis of the
country’s ex situ conservation programme.
In Asia, country reports indicate that about
12 native species have been evaluated at the
provenance and or progeny level in Southeast
Asia, 17 in South Asia and 12 in China, Japan and
the Republic of Korea. Central Asian countries
provided little information about provenances,
but Haloxylon aphyllum sources have been tested
and drought- and pest-resistant variants have
been identified. Abies sibirica has been tested in
Kazakhstan.
China commenced provenance trials in the
early 1980s and has now conducted trials for more
than 70 important planted species, including the
following as well as various key exotic species:
Betula platyphylla, Cunninghamia lanceolata,
Larix gmelinii, Larix principis-rupprechtii, Picea
koraiensis, Pinus armandii, Pinus koraiensis,
Pinus massoniana, Pinus tabuliformis, Pinus
yunnanensis, Platycladus orientalis, Populus
tomentosa,
Sassafras
tzumu,
Taiwania
cryptomerioides and Ulmus pumila.
Molecular markers
Molecular marker approaches employ laboratorybased techniques to identify and describe
genetic variation. Greatly reduced costs of gene
sequencing and increases in computer processing
speed and power have led to a proliferation of
DNA studies, including whole genome sequencing
and rapid progress in identifying the location and
function of specific genes.
Entire genomes are now being sequenced
for both angiosperms (including the important
commercial forest trees Populus trichocarpa
and Eucalyptus grandis; four fruit-tree species,
Carica papaya, Citrus sinensis, Malus domestica
and Prunus persica; and the most primitive
angiosperm,
Amborella
trichopoda)
and
gymnosperms (Larix sibirica, Picea abies, Picea
glauca, Pinus lambertiana, Pinus pinaster, Pinus
sylvestris, Pinus taeda and Pseudotsuga menziesii).
128
The sequencing of coniferous genomes, which are
typically an order of magnitude larger than those
of other organisms, has only been made possible
through the introduction of new sequencing
technologies and dramatic reduction in costs4
and has been facilitated by collaboration among
different laboratories and research groups.
These sequencing studies are complemented
by gene mapping studies and elucidation of gene
function and expression for traits such as growth,
wood properties and response to biotic and
abiotic stresses in seven economically important
tree genera: Castanea, Eucalyptus, Picea, Pinus,
Populus, Pseudotsuga and Quercus (Neale and
Kremer 2011). Novel approaches have been
developed to link markers to important traits,
with genomic and marker-assisted selection close
to being realized in several tree species. Molecular
geneticists are increasingly realizing that they
will not be able to use the rapidly expanding
gene-level information without whole-organism
information from the field.
Isozyme and/or allozyme markers were widely
used as molecular markers up to the 1990s to
evaluate genetic diversity and breeding systems
in trees, including for examination of variation
within and between populations, using over 20
enzyme systems; the earliest research was mainly
focused on high-priority plantation genera and
species for production of timber and pulp, such as
Eucalyptus, Pinus and Populus species. However,
since the advent of more informative, accessible
and cost-effective DNA-based approaches,
including direct DNA sequencing, single
nucleotide polymorphisms and microsatellites
(see Chapter 8), these isozyme studies have fallen
out of favour, although they are still useful for
some purposes. For example, isozyme markers
still have a role in low-cost assessment of diversity
and breeding systems.
4
Sequencing costs have declined from more than USD 5 000 per
raw megabase of DNA sequence in 2001 to about USD 0.05 in
2014, with the most dramatic decline having occurred since 2007
(National Human Genome Research Institute, 2014).
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
The country reports relect the trend noted
by Mexico that, apart from ield testing using
quantitative means, “molecular markers have
been the most popular [method of characterizing
intraspeciic variability] for forest species [in]
the last ten years”. Of 409 reported uses of
markers to evaluate intraspeciic variation, 58
percent were DNA markers, compared to 38
percent for biochemical markers. The use of
morphological markers was reported in only 4
percent of evaluations. Of the analyses using
biochemical markers, 65 percent were in Europe
and 21 percent in Asia; while 46 percent of the
evaluations using DNA markers were in Asia and
44 percent in Europe. Only 3 percent of DNA
marker uses were in Africa, possibly relecting
the limited resources available for applying these
techniques.
Enzyme electrophoresis and neutral DNA
markers (RAPD) have been used in Burkina Faso;
species characterized using these methods or
simple description of morphological characters
include Acacia senegal, Adansonia digitata,
Borassus
aethiopium,
Parkia
biglobosa,
Sclerocarya birrea, Tamarindus indica and
Vitellaria paradoxa subsp. paradoxa.
While quantitative techniques are still the
dominant method used to produce improved
germplasm for most commercial forest species,
molecular techniques have promise for rapid
identiication of genetic variability. These
methods can circumvent the long time periods and
risks involved in provenance and progeny testing,
which are a constraint on the development and
production of improved germplasm for general
use. Germany points out that the time available to
develop climate change-adapted planting stock is
extremely limited, highlighting the importance
of developing techniques that can deliver results
more quickly than traditional methods.
Although molecular methods are highly
effective in identifying variability and evaluating
similarity or differences between individuals
or populations, many of these techniques use
neutral markers, i.e. markers with which the
genetic variation identiied does not necessarily
confer an adaptive advantage or contribute to
improvements in productivity, performance or
utility. Many countries involved in research at this
level identify development of molecular markers
for adaptive and productive traits as a high and
urgent priority. For example, Germany states:
“It is essential that future international research
projects also provide more information about the
genetic variation to adaptation-relevant gene
loci. This would provide important information on
the adaptation potentials of tree populations.”
Developing countries with limited resources
face constraints in using molecular markers,
including cost (as noted by Zimbabwe, for
example) and lack of expertise, equipment and
facilities. Nonetheless, it is recognized that these
techniques offer great potential. Ghana, for
example, identiies development of expertise in
biotechnological approaches and upgrading of
existing facilities as key capacity requirements
for advancing its characterization agenda.
Further, many developing countries already use
these techniques, even if to a limited extent.
Cooperative arrangements with international
or regional partners and donors could provide
opportunities for molecular characterization by
developing countries or assist in the development
of in-country facilities and expertise.
In some countries, research facilities with
the capacity to undertake these studies may be
located in universities, agricultural institutes,
research organizations or the private sector,
i.e. outside the institutions charged with FGR
conservation and management. In these cases
effective collaboration can deliver mutually
beneicial outcomes. In the United States of
America, for example, much breeding and
improvement work involving molecular research
takes place through cooperative arrangements
among universities, public land management
agencies and private companies. Burkina Faso
outsources its characterizations based on
molecular markers through partnership with
universities in Denmark, France, the Netherlands
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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and the United Kingdom. Germany notes that
some government departments offer consulting
services in FGR characterization; adoption of this
approach more widely could make it possible to
procure skills, expertise and access to facilities on
an “as needed” basis.
On the other hand, development of these
capacities within countries can offer advantages,
both in facilitating the integration of national
strategic priorities for FGR with research
capacity and function, and in allowing the
country to pursue its own FGR conservation
and management interests without having to
accommodate external demands.
Ethiopia notes that it has facilities engaged
in molecular characterization of agricultural
crops but no dedicated facilities for such
research on FGR; in the few intraspecific studies
of trees, molecular characterization (using
inter-simple sequence repeats [ISSRs], AFLP and
chloroplast microsatellites) has been outsourced
internationally. This example suggests the
potential benefits from coordination and
cooperation in areas of common interest and
underscores the importance of integrating FGR
objectives with strategies and programmes in
related fields through harmonization at the
national level.
Analytic methods
The national reports rarely refer to the methods
used to analyse the data collected in studies using
the techniques described above. However, China
and Mexico detail the analytic methods used in
evaluation of their FGR data.
In analyses of interpopulation variation,
China reported the use of variance components,
genetic distance and phenotypic differentiation
coefficients. Intrapopulation variation has been
evaluated using standard deviation, coefficient
of variation, variance and Shannon information
index. Common parameters used for isozyme and
DNA analyses include allele frequencies and their
distribution, variance of genotypic frequencies,
average number of alleles per loci, effective
number of alleles, percentage of polymorphic loci,
130
Wright’s inbreeding coefficient and Nei’s diversity
index, Shannon information index, coefficient of
genetic differentiation and genetic distance.
In studies of the variability of Mexican tree
species, genetic diversity was inferred through
calculation of expected heterozygosity, observed
heterozygosity, number of alleles per locus and
percentage of polymorphic loci.
Characters investigated in studies of
intraspecific genetic variability
As noted earlier, investigations of variability
for commercial plantation forestry (focused on
economic characters such as growth rate, wood
characteristics or industrial processing qualities)
differ somewhat from those carried out in
the design of effective genetic conservation
programmes (focusing on level and distribution
of variability within a species). Provenance
and progeny testing may, however, be used to
establish variability in conservation management,
for example when selecting materials of
threatened species for circa situm conservation
measures. Characterization of variability that
can contribute to adaptation and survival under
future environmental regimes (involving climate
change, human modification of landscapes and
spread of invasive species), athough recognized
as crucial by some countries, needs further work.
Countries reported the individual characters
that were assessed in the course of evaluating
genetic variability. Of the 27 characters reported
as used in 692 characterizations, the most
studied are shown in Table 11.1; around 15
other characters were evaluated less often.
These data indicate that purely morphological
characters remain widely used in the evaluation
of variability, despite the increasing focus on
molecular markers. They also highlight the
importance that countries place on identifying
trees and genotypes for breeding for pest and
disease resistance. For example, Ghana notes that
although its intraspecific evaluation programme
is limited, “all objectives and priorities for
understanding intraspecific variation are geared
towards identification of planting stocks resistant
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
TABLE 11.1
Characters most frequently assessed in 692 evaluations of genetic variability reported by countries
Character
Type of character
Characters least subject to phenotypic variation, i.e. seed,
fruits, cones and pods
Morphological
17.5
Disease and pest resistance
Adaptive/productive
13
Leaf anatomy
Morphological
Bole/stem diameter
Productive
7
Growth rate
Productive
5.5
Biomass/fodder productivity
Productive
5
Height
Productive
5.5
Drought resistance
Adaptive/productive
5
Phenology
Adaptive
5
Bark
Morphological
5
Chemistry/exudates
Biochemical
3
to insect and disease infestation under forest
plantation conditions”.
Mexico notes that the variables and measures
used in evaluating genetic diversity are indicators
that are often used to represent and infer
the general variability of target organisms.
This process relies on assumptions about the
relationship between the variation observed
in the variables tested and the variation in the
target character or organism under study. Several
countries observe the need for continued research
on the methods for characterizing diversity. As
mentioned above, for example, Germany places
priority on identifying molecular markers for
adaptive characters (e.g. drought tolerance,
fire and wind resistance and pest and disease
resistance) and productive characters (e.g. growth
rate, form and wood processing qualities).
Identification of markers for adaptive traits is
considered especially significant with respect to
climate change, which is widely recognized in
country reports as the major challenge to the
integrity of forest ecosystems and the survival
of individual tree species. A number of countries
point to the need to identify breeding stock for
both productive and environmental purposes
% of total evaluations assessing
this character
7
that is better adapted to the expected conditions,
for example with respect to phenological
responses
(reproductive
phenology
and
deciduousness) or drought, fire, pest or disease
resistance. Germany stresses the high priority for
identifying markers for characters that confer
the ability to survive under the altered climate
regimes predicted by climate models, to facilitate
the selection of climate change-adapted plant
materials.
Monitoring of forest genetic
resources
Monitoring the state and trends of a country’s
genetic resources is an essential requirement for
effective FGR conservation and management and
decision-making. Monitoring assists in identifying
the extent, severity, location and nature of
genetic erosion of species and forests as well as
in evaluating conservation and management
actions. Article 7 of the CBD (UN, 1992) requires
signatory countries to “monitor, through
sampling and other techniques, the components
of biological diversity… paying particular
attention to those requiring urgent conservation
measures and those which offer the greatest
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
potential for sustainable use”. The rapid rate
of forest loss, high levels of genetic erosion and
impending impacts of climate change highlight
the urgency of establishing effective monitoring
programmes.
A number of countries recognize in their reports
the need for an effective forest genetic resources
monitoring and evaluation system as required
by the CBD. Canada notes that monitoring of
inter- and intraspecific variation is a priority for
tracking FGR status of species including threat
status, vulnerability and level of genetic erosion.
For such monitoring, Canada notes the need for
continuing investment in field and laboratory
personnel and information management, as
well as a need to ensure consistency across
jurisdictions.
As it is impossible to measure the genetic
variation and to monitor changes in all or most
tree species, two approaches may be applied.
The first involves measuring and monitoring the
genetic variability in only the highest priority or
model species. Germany, for example, monitors
genetic variation of five species in response to
forest management regimes and silvicultural
practices. The second approach involves
identifying and monitoring surrogates for FGR –
for example, particular species or populations, or
the area of forest or tree cover, in combination
with GIS, ground truthing, biogeographical
interpretation and monitoring of species-rich
area. Effective monitoring of genetic variability
will often require the use and assessment of
several measures in combination.
The current level of FGR monitoring varies
enormously among countries. For example,
Thailand has a network of 1 285 permanent plots
as part of its national forest resources monitoring
information system; sampling commenced in
2008 and is expected to provide valuable input
for updating information on forest cover,
genetic resources and deforestation. Thailand
also has a strategic framework for surveys and
database establishment for biodiversity and FGR
in protected areas. At the other end of spectrum,
132
Solomon Islands currently has no system in place
to monitor or report on FGR erosion.
Monitoring needs to be a requirement of all
FGR conservation and management programmes
and must be included in national FGR strategies
with endorsement at the national level helping
to secure budget allocations. Several countries
point out the need for increased and consistent
monitoring and for its harmonization across
jurisdictions, regions and national boundaries.
Cooperative administrative arrangements, where
they exist, can provide a vehicle for integrating
this FGR conservation and management function
across jurisdictions. For example, European
countries are currently negotiating a European
forest convention addressing sustainable forest
management, which may provide a suitable
avenue for harmonization.
Developed countries with well established
national forest inventories and monitoring
systems, such as Finland and Germany, are better
placed to document and describe changes in FGR
than most developing countries. However, genetic
monitoring of forests is at a very early stage, with
so far only a small number of pilot studies (e.g. in
Germany and in the framework of EUFORGEN).
It is expected that forest genetic diversity can
be better maintained through prevention
of overcutting and forest loss; through the
application of silvicultural techniques favouring
multipurpose mixed hardwood stands established
from a wider genetic base over monospecific
coniferous plantings; and through an increase in
forest area, including the targeted introduction
of rare tree species into natural forest ecosystems.
However, more monitoring and repeat inventories
are needed to confirm such assumptions.
Developed, temperate and boreal zone
countries may be able to maintain and expand
forest cover and genetic variability more easily
because they tend to have fewer tree species,
more effective forest administration and less unregulated harvesting than developing countries.
They nonetheless offer examples of how it
is possible to maintain and expand FGR, for
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
instance through sustainable forest management,
silvicultural techniques and policy mechanisms such
as incentives for sustainable management of FGR.
Differences among countries and
regions in characterization of FGR
As already noted, knowledge of genetic variability
varies widely among countries. From the country
reports, the degree to which FGR have been
characterized and the methods employed appear
to vary with:
• the level of economic development and
resources available for characterization
(financial, technical, institutional and
personnel);
• the level of development of organizations
dealing with conservation and forestry and
of associated information and management
systems;
• the importance of the forest sector in the
economy, and the methods of production
and forest management (e.g. dependence
on planted forests versus naturally
regenerated forests);
• the nature and structure of the forest
industry, i.e. the production profile with
respect to output types and the extent of
development of private-sector capacity in
genetic improvement;
• the degree of engagement with regional
and international networks;
• the support and contribution of
international donors;
• which organization in the country has
prime responsibility for conservation and
management of FGR;
• the involvement and input of relevant
research organizations, including institutes
and universities;
• the presence or absence of supporting
legislation and regulatory frameworks
that give value to FGR conservation and
management;
• the presence or absence of a national FGR
strategy or programme;
• the area of natural forest remaining, the
degree of diversity within the country’s lora
and the heterogeneity of its genecological
zones;
• the number of priority tree species
identiied for conservation and
management.
Developing countries thus face several
challenges causing them to lag behind developed countries in characterization of genetic
variability. First, developing countries, particularly
in tropical or subtropical areas, often have much
higher levels of tree species diversity and FGR
variability, and these countries (especially the 17
megadiverse countries) require greater survey
effort to document their FGR, i.e. to identify
and map distribution of species, to identify areas
of high interspecies diversity and to investigate
genecological relationships. Second, developing
countries generally have fewer resources
available for survey, characterization and data
management. Third, developing countries
generally experience much higher rates of
uncontrolled forest clearing, meaning they have
a greater requirement for inventories of FGR
variability at the species level. Fourth, tropical
developing countries rely heavily on the use
of exotic species for plantation development,
for which extensive characterization work has
already been undertaken but for which access to
germplasm remains critical for future breeding
and improvement. Given the resource constraints
in these countries, remote sensing (coupled with
an appropriate level of ground truthing and GIS)
is an extremely important tool for assessment and
monitoring.
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STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Chapter 12
In situ FGR conservation
and management
In situ conservation of FGR is often considered
to be the core activity of FGR conservation and
management, as it maintains the existing natural
pool of genetic variability while at the same
time permitting natural selection processes to
operate. It also maintains the typically wide
range of variation required for effective selection
for breeding and genetic improvement of trees
with high commercial or service value. Thailand
recounts the challenges of successful longterm in situ conservation in reference to an
initiative begun with assistance from the Danish
Government during the 1970s:
“The natural stands [of Pinus merkusii],
especially in the northeast of Thailand,
have been heavily exploited by local
communities, primarily as a source of resin
and fuelwood. In addition, many good
stands are fragmented and declining as
a result of the widespread conversion of
forest to farmland and frequent fires.
The lowland stands that showed the best
performance in provenance trials are even
threatened with extinction; [conserving]
genetic variation within the species by
selecting a number of populations from
different parts of the distribution area
will serve as a source for protection,
management and maintenance of genetic
resources by providing a basis for future
selection and breeding activities as well
as for seed sources with a broad genetic
base.”
In situ conservation is often considered the
first course of action in conservation of both
FGR and other forms of biodiversity; alternative
methods such as ex situ measures are normally
only considered when it has been established
that in situ conservation is not feasible, or
when species are at serious risk of extinction
in the wild, or for safety duplication purposes.
Advantages of in situ conservation are that it
permits conservation of ecological, aesthetic,
ethical and cultural value at the same time, and
that large amounts of FGR may be efficiently
conserved through simultaneous conservation of
the diversity of multiple species. In indigenous
production forests where sustainable forest
management is practised and FGR variability
is maintained, in situ conservation is fully
compatible with harvesting of timber and forest
products. The security of conservation tenure
and the type and level of management are major
factors in determining conservation outcomes
for in situ FGR.
In situ conservation ensures that in the
absence of catastrophe and genetic bottlenecks,
the genetic variability contained in the target
species is maintained at a high level and serves
as the foundation on which selection pressures
can direct adaptation to new conditions. In situ
conservation therefore allows for the genetic
variation contained in a population or species to
change over time; European countries therefore
refer to it as dynamic. Ex situ conservation, in
contrast, is predominantly static, preserving a
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
“snapshot” of the variability present at the time
of conservation of the germplasm.
Although most in situ conservation assets are
managed as open, dynamic breeding systems,
a small number may be subjected to controlled
pollination or other reproductive manipulations
for specific breeding outcomes.
In situ conservation involves a wide range of
activities, and every in situ programme will include
a combination of these. Measures mentioned in
country reports include:
• a process for prioritizing areas, species and
populations for in situ conservation action;
• research to understand the nature and
distribution of genetic variability within the
area, species or population and to identify
threats to this variability and management
actions to protect it – for example,
determining the location and number of
populations and individuals and the area
of reserve required to maintain variability
at the desired level – to guide conservation
programme design;
• protection of an area containing target
or priority FGR (species, population or
individual) through its dedication as a
protected area or reserve, with restrictions
on activities that threaten FGR;
• legislation and/or regulations enabling
conservation of the area or species,
including control of access and use, for
example by gazetting or listing it on an
official “threatened” list;
• enforcement of legislation and regulations
and corresponding action to control
threatening activities;
• preparation of a management plan for
a forest or species, involving control
of activities that degrade its genetic
resources (e.g. by managing access, use
and harvesting) and maintenance of the
conditions necessary for its survival and
regeneration (e.g. by maintaining ecological
processes, controlling invasive plants and
136
animals, managing wildfire and maintaining
pollinators and dispersers);
• preparation and implementation of a
sustainable forest management plan that
ensures that genetic variability is not
diminished in areas subject to harvesting
and use;
• promotion of the participation of forest
users and adjoining communities in
sustainable forest management including
access and benefit sharing, e.g. through
incentive payments for stewardship or
employment based on activities undertaken
in accordance with sustainable forest
management principles;
• education and awareness raising for forestusing communities and industries, regarding
appropriate uses and activities and practices
that minimize impacts on FGR;
• preparation and/or implementation of
guidelines or codes of practice governing
activities that may be permitted in reserves
or areas providing FGR benefits, to minimize
impacts on variability – for example, a
code of forest practice or guidelines for
reduced-impact logging or for harvesting of
fuelwood or NWFPs;
• provision of alternative livelihood
opportunities for forest users who may be
displaced or disadvantaged by change in
land use (for example rural, traditional or
subsistence communities that rely on forest
products for fuel, housing materials, food,
medicines and income), including forest
plantation or better forest management
to counter any shortfalls in supply of forest
products.
In their country reports, different countries
interpret in situ conservation of forest genetic
resources in different ways, which complicates
interpretation. Countries report conservation in
situ in a wide range of forest reserve categories
and ownership types, ranging from strictly
protected areas to forests used for wild harvest
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
and timber production to private property. The
strict definition of in situ conservation of genetic
resources implies reserves established specifically
for conserving genetic diversity of targeted tree
species. Many European countries have such
reserves and report on this basis. The Russian
Federation, for example, has designated 205 501
hectares as genetic reserves for 21 tree species.
Ownership of natural forest lands inluences a
country’s approach to in situ conservation of FGR;
the ability to make land-use decisions depends
largely on whether the land is under public,
private or communal ownership. In Finland the
area of privately owned forests is 15 million
hectares, more than double the area of publicly
owned forests; while in Canada, which has
10 percent of the world’s forest area, only
7 percent of the forest land is privately owned.
Where signiicant forested areas are publicly
owned, the State can create protected areas
and reserves for in situ conservation of FGR,
consistent with national strategies and priorities.
Governments are generally less able to inluence
land use and protect FGR on the private estate.
Regulations governing protection of FGR on
private land are more effective in countries
where State power is strongest, administration is
effective, incentives are available and community
support for conservation is well accepted.
The numbers of species and subspecies
conserved in situ and ex situ, by region, according
to the country reports, are presented in Figure
12.1. (Ex situ conservation is covered in Chapter
13.) It is important to note that the igures relect
the different reporting approaches taken by the
countries. For example, most of the tree species
in North America are represented in protected
areas, but the areas are not designated speciically
for genetic conservation so the reported number
is low.
Although country reports detail the vast areas
and amounts of FGR conserved in situ in protected
areas and other public lands, many countries note
that they have few if any formal, designated in
situ conservation reserves for priority species. For
example, India points out that nearly 16 million
hectares are conserved in protected areas (almost
5 percent of the land area) but lists only 18 481 ha
in dedicated reserves for in situ conservation of
target species.
Countries that do not have such reserves may
still have effective in situ conservation of FGR. In
British Columbia, Canada, for example, secure in
situ protection is in place for most species across
much of their natural distribution. Genetic gap
analysis shows that all tree species are represented
with adequate population size in existing
protected areas; however, across the species’
ranges gaps remain in some of the biogeoclimatic
zones (Krakowski et al., 2009; Chourmouzis et al.,
2009). Some countries that do not have reserves
designated strictly for conservation of FGR report
in situ conservation in protected areas designated
for a variety of purposes, but without the beneit
of genetic gap analysis it is not possible to know the
degree to which genetic resources are protected.
Integration of FGR objectives into a wide
range of land-use designations and regulations
governing the use and management of forested
land should be considered for both public and
private land. Where legislation and regulation
relating to reserves providing in situ conservation
are inadequate, Germany suggests increasing
the legislative backing for FGR conservation
management actions by including the protection
purpose “conservation and sustainable utilization
of forest genetic resources” in the relevant legal
provisions for designated in situ gene conservation
assets. This could be extended to other public land
reserve categories and designations providing in
situ conservation beneits.
A further example of the need to address
speciically the requirements of in situ FGR
conservation is provided by the many countries
that list their natural or near-natural seed stands
and seed production areas as in situ conservation
areas and reserves. While seed stands clearly have
in situ conservation value, several countries note
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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FIGURE 12.1
Number of species and subspecies conserved in situ and ex situ, by region
8 000
7 000
6 000
5 000
4 000
3 000
2 000
1 000
0
Total
Africa
Asia
Europe
Species and subspecies conserved ex situ
Latin
America
North
America
Near
East
Oceania
Species and subspecies conserved in situ
Total number of species and subspecies reported
Note: Numbers for Europe are deceptively high because the region includes a number of territories in tropical regions.
that they are rarely selected by a formal in situ
conservation programme designed to provide
in situ FGR conservation outcomes. Their value
as in situ conservation resources for priority
species may be correspondingly limited. This
again demonstrates the need for addressing
the particular requirements of in situ FGR
conservation and integrating them more widely
into management practices for all categories of
forested lands. However, for countries lacking the
resources for identification and management of
formal in situ conservation reserves, seed stands
are vital components of their in situ conservation
efforts.
138
Protected areas
Protected areas provide significant in situ
conservation in most countries. For the past 20
years, protected areas have mainly been created
for biodiversity conservation to meet country
obligations under the CBD (UN, 1992), which
states: “…the fundamental requirement for
the conservation of biological diversity is the
in situ conservation of ecosystems and natural
habitats and the maintenance and recovery of
viable populations of species in their natural
surroundings”. The contribution of the CBD to
the conservation of forest genetic resources is
inestimable.s
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Brazil, with 286 million hectares of permanent,
public forest estate, reports that protected areas
play a central part in in situ conservation. In
Canada 9.8 percent of the land area is protected.
China reports that 149 million hectares or 15
percent of the country is managed as some form
of nature reserve providing in situ conservation
of FGR. Seychelles notes that 48 percent of its
land is protected area.
To assess the contribution of protected areas
to in situ FGR conservation, countries were
asked whether they had evaluated the genetic
conservation of tree species within their protected
areas. A standard response was to list the number
of species and threatened species and to detail
any specific programmes for priority species. In
general, little information is available on the
actual FGR benefits of protected areas in relation
to populations, genecological zones represented
and intraspecific diversity conserved. This is
indicative of the immense difficulty of obtaining
the detailed information required to evaluate the
effectiveness of in situ FGR conservation when a
country has many hundreds or even thousands of
tree species distributed over vast areas.
Solomon Islands, with 80 percent forest cover
amounting to 2.24 million hectares, has a tiny
fraction of forest (0.017 percent) under formal
protection in two locations. The country report
notes that the lack of protected areas is of concern
given that logging is occurring at four times the
sustainable rate with serious impacts on FGR:
“The latest update on logging concession
areas provides evidence of forest cover
loss on logged-over areas which [is] also
associated with significant loss of natural
and ecological value…. Some endemic
forest species that are unable to adapt
to new environments face possible
extinction…. While most agree that the
creation of a conservation estate would
be in the national interest there is no
functioning institutional framework for
its advocacy, creation or management.
Even if such a framework existed then
there would be problems in funding it. For
these reasons none of the conservation
areas identified… have been reserved and
in fact many have already been logged.”
The country report from Brazil expresses
the value of the protected area approach to
conservation of FGR as follows:
“In situ conservation of genetic resources
is the most effective strategy, especially
when the main goal is the conservation
of entire communities of tree species,
as in the Brazilian tropical forests. In
these cases, trees of other species than
the target ones must be included in the
genetic conservation scheme, as well
as their pollinators, seed dispersers and
predators… The conservation of forest
genetic resources in Brazil involves a largescale in situ scheme, and for that purpose
a national-scale strategy had to be
implemented. [This involved] the creation
of a significant amount of conservation
units, as well spread over the national
territory as possible, synchronized with a
national strategy for biological diversity.”
Similarly, India notes that “when the whole
habitat or ecosystems are protected, whole plant
genetic resources also enjoy the protection.”
In situ conservation of FGR in forests under
some form of protection involves diverse
designations and categories associated with
widely differing regulatory regimes. The wide
range of protected area designations providing
FGR conservation value includes dedicated FGR
conservation reserves, protected areas, nature
reserves, protection forests, national parks, game
management areas, bird sanctuaries, Ramsar
Sites, scenic parks, ecoparks, forest reserves,
watersheds, mangrove forests and United
Nations Educational, Scientific and Cultural
Organization (UNESCO) Biosphere Reserves.
International Union for Conservation of Nature
(IUCN) protected area categories are useful in
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standardizing country reserve categories, and
some countries adapt their designations to allow
comparison with IUCN categories. Although
the designations vary significantly among
countries, systems tend to be shared in regions
with historical and political affinities. Brazil lists
12 categories of protected area providing in
situ conservation benefits. Canada notes that it
has “numerous categories of protected areas
established by multiple organizations at the
federal and jurisdictional levels and through nongovernmental organizations that either directly
or indirectly have the intent to conserve tree
species in situ.”
Regulations governing permitted activities
and guiding management vary widely and differ
in the extent to which they are compatible with
and/or facilitate in situ conservation of FGR.
Thailand notes that its forest reserves provide
in situ conservation benefits for FGR, but the
laws and regulations governing them are less
strict than those of protected areas. In addition,
in many countries large areas have been legally
designated as protected areas but the legislation
has little or no enforcement. This situation often
coincides with outdated inventory of tree species
and populations (or no inventory at all). In such
situations a tally of the extent of protected
areas does not provide much useful information
regarding the species currently found within the
protected areas or the effectiveness of the in situ
conservation of FGR.
In many countries the multiplicity of designations and regulations presents a significant
challenge for harmonization and coordination of
in situ conservation objectives and requirements.
Closer integration of the requirements for in
situ FGR conservation with those for biodiversity
conservation and with those of other reserve
and public land categories, including productive
designations, could achieve significant synergies.
An important point regarding the protected
area approach for in situ conservation of FGR
is that although countries named a wide range
of protected areas, reserves and land categories
140
as their primary in situ conservation initiatives,
most of these areas were not designated or
selected for the conservation of priority FGR
and did not have management plans specific
to their particular needs. While landscape-scale
biodiversity conservation as currently practised
fulfils many requirements of in situ conservation
of FGR, failure to consider the particular needs
of formal in situ FGR conservation protocols will
inevitably result in losses of genetic variability,
especially for those species that depend most
on ecological disturbance. Canada’s country
report points out that although programmes for
the protection and management of threatened
species also provide in situ FGR conservation, they
do not necessarily address “silent” extinctions
which are “associated with loss of genetically
distinct populations, or loss of locally adapted
gene complexes, [which] are not considered in
[threatened species] legislation, yet may have
devastating consequences for tree species faced
with increasing environmental change”. It is
therefore important to integrate requirements
for in situ conservation of FGR into the full range
of a country’s biodiversity conservation initiatives,
to maximize outcomes for both.
In situ conservation outside
protected areas
Most in situ conservation of FGR takes place
outside protected areas on a range of public,
private and traditionally owned lands, especially
in multiple-use forests and forests primarily
designated for wood production.
Forests on public land used for production of
timber or other forest products may provide major
in situ conservation benefits, depending on the
intensity of use and the management approach:
Forests under sustainable management regimes
that take full account of FGR management
principles and practices will help conserve FGR,
while forests that are heavily exploited and/
or subject to uncontrolled extraction will not.
Sustainably managed natural production and
multiple-use forests are central to the in situ FGR
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
conservation efforts of a number of countries,
especially where native tree species are also
major commercial timbers.
Private and communally owned and managed
forest land may also provide important in situ
conservation of FGR, especially where important
FGR occur outside of protected areas. For example,
Brazil’s landscape-scale in situ programme is
guided by the identification of Priority Areas,
and many of these fall outside the public estate.
In some biomes, such as Mata Atlantica, Pantanal
and Pampa, almost all the Priority Areas and
almost all the important genetic resources occur
on private land. In these cases conservation of
FGR relies on the actions of private landholders.
The conservation regulations applied to
private and customary lands vary. In Zimbabwe,
for example, different categories of private and
communal landownership are subject to different
laws and levels of control. Government regulation
ranges from strong to practically non-existent
both within and among countries.
Achieving in situ FGR conservation on private
and customary land can be problematic, as it may
be difficult to secure long-term or permanent
conservation tenure over the land and to
achieve an adequate level of FGR conservation
management. Securing in situ conservation of
FGR in these contexts may involve a combination
of regulations, sustainable forest management,
education, provision of income and employment
opportunities from sustainable forest-based
industries, and incentives to reward landholders
for stewardship of FGR. Ex situ conservation will
often be required in cases where FGR assets are
outside publicly protected and managed areas.
On private land
Large areas of private forest land are held under
strict conservation tenure and management
regimes;
examples
include
land
under
conservation covenants held by NGOs such as the
Nature Conservancy; land managed according
to sustainable forest management principles
or dedicated as reserves; and other private
land subject to conservation management or
vegetation retention regulations.
The following example from Sweden, where
over 75 percent of productive forest is privately
owned, demonstrates the potential of private
ownership for achieving in situ FGR conservation
outcomes:
“Sveaskog, the largest forest company
in Sweden, owns 3.3 million hectares
of productive forest land, which is 14
percent of Sweden’s total productive
forest land. Sveaskog´s nature conservation strategy includes the ambition
to focus on conservation on 20 percent
of the company´s productive forest
land; 650 000 ha are assigned to
nature conservation using production
forests, nature conservation forests and
ecoparks.”
The use of legislation and regulation to
control land use and activities leading to losses
of FGR have been noted above. However, while
legislation and regulation are important, they
are limited in their effectiveness, particularly
in the private estate. Many countries note that
regulation has failed to control activities leading
to forest clearance. Under the Brazilian Forest
Code, forest in the private estate, if located along
rivers or on hills or slopes, must be preserved
as permanent protection areas; in addition,
minimum percentages must be maintained under
native vegetation (as legal reserves) depending
on the biome: 80 percent of rural properties in
forest areas in the Legal Amazon; 35 percent of
rural properties in savannah areas in the Legal
Amazon; 20 percent of rural properties in forest
or under other vegetation in other regions; and
20 percent in native grasslands in any region.
Legal reserves may be harvested for timber
and other products under sustainable forest
management plans. Permanent protection areas
and legal reserves cover 12 and 30 percent of
Brazil, respectively – twice the area of designated
protected areas on public land. However, 42
percent of the permanent protection areas
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and 16.5 percent of the legal reserves have
been subject to illegal deforestation, as have
3 percent of protected areas and indigenous
lands, and the extent to which the sustainable
forest management plans are adhered to is
unknown.
To counteract illegal clearance of FGR on
private land and encourage protection and active
conservation management of FGR, some countries
(e.g. Brazil, Finland and Sweden) offer incentives
and stewardship payments. In the United States
of America, where 57 percent of forests are under
private ownership (with regional variation),
37 million hectares in the private estate are
protected through voluntary conservation
covenants entered into by individuals, land trusts
and NGOs; these arrangements significantly
supplement the public reserve system. Such
voluntary private land conservation initiatives are
oriented towards the protection of areas, taxa
(species, varieties) and ecological processes. Their
contribution to in situ conservation of FGR would
be substantially enhanced if FGR conservation
and management objectives were incorporated
in reserve selection criteria and subsequent
management.
of its forests as they see fit would be
regarded by them as interfering with
their rights of private ownership. The
Land and Titles Act and other statutes,
including the proposed new Forests
Act, have mechanisms that provide for
the State to impose limits on the use of
private land; however where this is in the
national interest the understanding is that
the owners will be compensated for any
rights foregone. In the current economic
conditions the State is in no position to
make such compensation.”
Where lands under indigenous or customary
ownership and/or management are managed
under sustainable forest management principles,
e.g. as protected areas or under customary
regimes consistent with FGR conservation, they
can provide significant in situ conservation
benefits. In Vanuatu, all forests are under
customary ownership, with beneits lowing from
the direct involvement of landowners in forest
use and reforestation. Shortcomings, however,
include the convoluted arrangements required
for approving government management plans
and disruptions from landownership disputes.
On community or customary lands
Formal in situ FGR conservation
programmes
Indigenous or traditional ownership, use rights
and management may have a significant role
in conservation and management of in situ
FGR in many countries. For example in Canada,
aboriginal people own or control around 3 million
hectares of forested land and play a major part
in the management of FGR; other countries
where indigenous peoples feature importantly in
ownership, use and management of FGR include
India and Brazil. Customary tenure dominates
landownership profiles in many countries in
Oceania and parts of Africa. In Zambia, for
example, 46 million hectares (61 percent of the
country’s area) are under traditional ownership.
Solomon Islands reports the difficulties of FGR
conservation on customary lands:
“In the Solomons any moves to limit a
landowning group’s ability to dispose
142
In well protected areas with effective habitat
conservation enforcement there may be a need
for management actions to maintain genetic
resources that are not allowed by protected area
legislation. In such cases conservation of genetic
resources of tree species must be an explicit
objective of conservation areas.
The irst step in preparing an in situ
conservation programme is to specify and clearly
deine the conservation objectives to be achieved.
Breeding programmes aimed at improving a
species’ commercial or environmental value
may seek to conserve adaptive, productive and
quality traits (e.g. growth rate or form, pest and
disease resistance and adaptation to climatic
extremes) in populations or stands identiied for
conservation. Programmes designed to maintain
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
genetic variability for biodiversity conservation
purposes (e.g. threatened species management)
or general conservation of FGR will seek to
preserve the widest range of variation possible
with the available resources. Many countries
stress the importance of maintaining the widest
genetic base possible through in situ and
dynamic conservation measures to facilitate
adaptation to climate change for all forestry
and tree planting programmes, whatever their
objectives.
In contrast to the multispecies landscape-scale
protected area approach to in situ conservation,
dedicated programmes for in situ conservation
of the genetic diversity of priority trees require
specific provisions. Finland, for example, notes
that “valuable genetic resources exist also
in strict nature conservation areas, but these
areas are not considered to be part of the gene
conservation programme” because of differences
in management approach.
A dedicated programme for FGR conservation
of a priority species may specify the proportion of
variability to be preserved, the number of trees to
be protected, the area required for conservation,
and which populations need to be represented.
The most variability can generally be preserved
by maintaining a selection of genetically
representative populations, although some loss
of low-frequency or rare alleles may occur (as
reported by the United States of America). To
maintain rare alleles it is necessary to conserve
additional populations and/or many more trees
over a wider area, and the additional expense
may be difficult to justify. Measures to protect the
population’s variability and viability by managing
threats and ensuring regeneration of the stand
must also be specified (e.g. disturbance regimes
such as fire management).
Knowledge required for a scientifically sound
FGR conservation programme for a particular
species includes the targeted species’ pattern of
genetic variability, including variability within and
between populations across the species’ range;
its distribution; its ecological and regenerative
requirements; its physiological tolerances; its
genecological zones; its reproductive biology; and
associated pollinators, dispersers and symbionts.
The Thailand report states that “genetic resources
must be selected mainly based on the available
knowledge of spatial patterns of genetic
variation…. The combination of marker-aided
population genetic analysis and information
about adaptive and quantitative traits as well as
forest ecosystems would allow comprehensive
conservation programmes for individual species
in each forest type”. While some countries have
sufficiently detailed information to guide in situ
conservation planning for priority species, concern
is widely expressed about the lack of genetic or
other pertinent information for most species,
with some countries having little information on
which to base in situ conservation programmes.
Even well-off countries note the impracticality of
surveying all populations of all tree species; for
poorer countries with high levels of tree diversity
distributed over wide areas the task is impossible
with current technologies.
Ideally, priority species must first be thoroughly
investigated for genetic and ecological
diversity; then genecological zones need to
be identified and delineated, with conserved
populations selected to represent the full suite
of genecological zones across the entire range,
resources permitting. Thailand’s conservation of
Pinus merkusii, for example, includes all eight
identified genecological zones; similarly, the
country’s in situ conservation programme for teak
covers all five genecological zones identified in 15
locations on the basis of topography, climate and
vegetation. China describes a similar process for
designing in situ FGR conservation for priority
species: “The number of populations or stands
of target species, the size of area and effective
number of trees for in situ conservation were
determined according to the result of genetic
diversity analysis, combined with data obtained
from field surveys.”
Where detailed knowledge is lacking, general
guidelines based on established principles may be
used to design in situ conservation programmes
(e.g. FAO 1993; FAO, DFSC and IPGRI, 2001).
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Parameters that may be considered in the
preparation of such guidelines include:
• breeding system – e.g. obligate outcrossing
versus self-fertility; vegetative propagation;
animal vector versus wind pollination (with
the latter assumed to have a more even
distribution of alleles);
• range – localized or widespread, disjunct or
continuous (level of fragmentation), edge
of range;
• isolating factors – greater or less breeding
isolation between populations;
• environment – homogeneous versus
heterogeneous environmental factors (e.g.
geology, soils, aspect, moisture, climate);
• population size – small in number and
limited in area versus numerous and
extensive;
• level of natural variability occurring within
the species and its populations – high or low
diversity within and among populations;
• patterns of distribution – e.g. clustered
versus dispersed, continuous versus disjunct.
Guidelines for establishing in situ conservation
reserves for particular species have been prepared
in some countries and regions. For example, China
has developed technical standards and codes for
in situ conservation sites; these address selection
of species, number of populations, area, number
of trees and management requirements.
The European Information System on Forest
Genetic Resources (EUFGIS) provides information
for use in defining the minimum number of
individuals required for in situ conservation
of a species in European countries. Regional
guidelines are particularly useful for shared
species and environmental conditions, and
where conservation and management priorities
are similar. The use of the EUFGIS guidelines
is demonstrated by the following description
of minimum in situ conservation programme
requirements from Sweden’s country report:
“Five hundred trees per gene conservation
unit are sufficient for species with large
and continuous populations which
144
are extensively used in forestry. These
species are subjected to extensive forest
tree breeding and import of forest
reproductive material. In Sweden, Pinus
sylvestris, Picea abies and perhaps also
Betula pendula and Betula pubescens
belong to this category.
“Fifty trees per gene conservation
unit are sufficient for species with
populations of varying size and structures
and with no or limited use in forestry. In
Sweden, several tree species such as Acer
platanoides, … Fagus sylvatica, Fraxinus
excelsior, Populus tremula, … Quercus
robur, Salix caprea … and Ulmus glabra
likely belong to this category.
“Fifteen trees per gene conservation
unit are sufficient for species with (very)
small and isolated populations, which
may be situated at the edge of the
species geographic distribution area.
These species may be red-listed or their
populations may have recently decreased
owing to forest damage. In Sweden,
for instance, Acer campestre, Carpinus
betulus, Juniperus communis, Prunus
avium and Ulmus minor belong to this
category of tree species.”
Also in Europe, EUFORGEN (www.euforgen.
org) has developed technical guidelines for FGR
conservation and use for 36 tree species and
species groups. EUFORGEN has facilitated the
development of gene conservation reserves
in many countries. EUFORGEN’s standards
for effective in situ conservation include
periodic monitoring of the FGR contained
within the reserves. The information collected
through monitoring will be extremely useful
for understanding factors that inluence the
effectiveness of conservation.
Finland also lists some general rules:
“A basic requirement for a gene reserve
forest is that it is of local origin and has
been either naturally regenerated or
regenerated artiicially with the original
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
local seed source. The general objective
is that a gene reserve forest of a windpollinated species should cover an area of
at least 100 ha, in order to secure sufficient
pollination within the stand, but smaller
units have been approved in particular
for birch species. The in situ units for rare
broadleaves are much smaller as a rule.”
The Thailand report offers these guidelines
for outcrossing species: “After the genetic
variation within and among populations of any
species has been investigated, the most variable
populations with relatively high outcrossing rates
(for outcrossing species) should be chosen as the
sources for gene conservation.”
Although general principles can be used
to design in situ conservation programmes,
guidelines are best prepared with reference to the
species and conditions existing in the particular
country or region. Guidelines may already exist,
or at least a high level of information may be
available, for well studied genera and species.
Regional associations or networks focused on
particular taxa may provide opportunities to
update and/or prepare guidelines. To facilitate in
situ conservation, the preparation of guidelines
should be considered for countries, regions,
genera and species that lack them; the work of
EUFGIS and EUFORGEN provides a useful model
for other regions.
Dedicated FGR conservation programmes
for priority species, whether based on detailed
species knowledge or guidelines, generally seek
the most efficient design to conserve the optimal
amount of genetic variability, for example
through identifying the number and location
of populations, the number of individuals and
the area required to conserve the most genetic
variation in the target species. The guidelines of
FAO, DFSC and IPGRI (2001) suggest, for example,
that between 150 and 500 interbreeding
individuals are required for each population to
be conserved in situ.
To establish, manage and monitor in situ
conservation units at the fine scale necessary is
resource intensive and expensive. Accordingly,
in situ conservation programmes strive to
conserve the minimum number of individuals
and populations and the smallest possible area
consistent with the conservation outcomes
sought. Thailand’s in situ conservation of Pinus
merkusii includes two reserves of 100 and 960 ha;
Denmark conserves 56 species on 2 880 ha of
genetic conservation reserves; and the average
size of in situ conservation units in Bulgaria is
6.3 to 6.8 ha for conifers and hardwoods. While
these reserves meet the genetic requirements
for in situ conservation, it must be noted that
reserves of limited size and individuals will
rarely if ever be adequate alone to conserve
the full range of ecological and evolutionary
processes needed for the long-term viability
of the population. To remain viable, conserved
populations need to be embedded in a healthy
landscape matrix that includes the pollinators,
seed dispersers, microbial associations, and
myriad of other organisms and processes
that comprise a viable ecosystem. The matrix
and its ecological processes must be properly
managed together with the designated in situ
conservation units.
Consistent with this approach, target
conservation populations of priority species are
almost always located in existing designated
protected areas, forest reserves or production
forests. These areas may be subject to further
specialized conservation and management
measures such as monitoring, stricter control
of use or access or stimulation of regeneration.
Therefore a landscape-scale approach to
conservation of FGR that addresses ecological
processes must be implemented together with
management of smaller, well sited and dedicated
reserves for priority tree species.
The provision of biological corridors to
facilitate gene low may help reduce risks of
inbreeding and genetic drift in situations where
in situ conservation units are functionally and
reproductively isolated. Examples include the
Greater Mekong Subregion Core Environment
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Programme and Biodiversity Conservation
Corridors Initiative and plans to link fragmented
landscapes in Australia and Sri Lanka. In situations
where FGR in situ conservation value is extremely
high and the conservation of reproductively
isolated, small populations is considered essential,
then population viability analysis is needed to
assess the feasibility of such conservation and to
inform management options for it.
Selecting areas for in situ FGR
conservation
Countries use a variety of approaches to identify
and prioritize areas for in situ FGR conservation.
The importance of the landscape-scale, protectedarea approach to in situ FGR conservation has
been noted above; areas may be selected as a
means of preserving the entire range of species
contained within them (including trees) as
well as ecological processes and many other
functions. Conserving large areas of forest in
reserves serves well in the absence of detailed
genetic information on priority species. Criteria
for selecting areas may include high levels of
tree species diversity; the presence of tree species
with a high level of endemism or threatened
status; a high threat level for the particular forest
association/vegetation community; the ability to
support ecological processes; and the viability of
populations and processes.
Brazil reports that since its FGR conservation
involves a large-scale in situ scheme (as described
above), “a national-scale strategy had to be
implemented”. An extensive country-wide survey
identified 3 190 priority areas for biodiversity
conservation and sustainable use, using the
criteria of representativeness, environmental
persistence and vulnerability.
Ethiopia has defined 58 National Priority
Forest Areas and five classes of vegetation
containing priority species; in conjunction with
other protected areas, these are the basis for the
in situ conservation programme, which covers
about 14 percent of the country’s area. Ethiopia’s
Forest Genetic Resources Conservation Strategy
146
sets general criteria for establishment of in situ
conservation sites, including:
• the number of priority species in the forest;
• the presence of unique, endangered and
endemic species within the population;
• the accessibility of the forest;
• the degree or threat of forest disturbance;
• the species richness of the site or
population;
• the attitude of the local people or
community towards conservation.
The Republic of Korea has a vast system of 432
FGR reserves covering a total area of 126 868 ha.
These are areas warranting special conservation
measures. They are classified in seven categories:
primeval forest, rare plant natural habitat, rare
forest type, useful plant original habitat, alpine
plant area, wetland forest and valley stream, and
natural ecosystem conservation.
Zimbabwe uses the level of threat to the
genetic diversity of economic species as a
criterion for establishing “strict natural reserves”,
which are areas containing “commercially
harvestable indigenous hardwoods whose
genetic integrity could potentially be altered by
overexploitation”.
To identify areas where genetic diversity may
be eroding most seriously, the United States
of America assesses range contractions among
forest species as a surrogate for genetic loss. The
northeastern region (from the mid-Atlantic to
New England) shows the greatest loss of forestassociated vascular plants. Sweden also notes the
importance of range criteria for conservation,
especially with respect to adaptation to climate
change in situations where there is clinal variation
along a climate gradient, e.g. latitude, altitude,
aspect and rainfall. Gap studies and analyses are
used to identify FGR that are not adequately
conserved through existing measures, to identify
vulnerabilities and to guide conservation and
management decision-making.
The distribution of natural forests serving as
repositories of FGR may vary significantly across
a country. The most extensive forest clearing has
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
occurred in areas of longest settlement, highest
population density and highest agricultural
productivity, such as Brazil’s Atlantic Forest biome.
In Thailand, as well, much of the teak resources
have been lost to agriculture. The high rates of
clearing in agricultural areas result in the loss of
species and the associated variability adapted to
these highly productive areas, suggesting that a
high priority should be placed on the conservation
of any remnant FGR remaining in sites of high
productivity and fertility. While highly degraded,
overexploited forests in rural areas may provide
opportunities for replacement with more
productive planted forests (as pointed out by
Ghana), it is important to consider the current and
future value of the forests as potential sources of
genetic variability for conservation, and not to
overlook the development of improved varieties
for use and planting in these areas.
Priority species for in situ conservation
While the process of prioritizing species for in
situ conservation is consistent with prioritization
of species for conservation and management of
FGR in general (discussed in Chapter 10), several
particulars are worth noting.
Many countries list priority species for in situ
conservation in their reports; they add up to
743 species globally. The main reasons for this
prioritization are shown in Figure 12.2, although
species are sometimes prioritized for in situ
conservation for several reasons simultaneously.
For example, Dalbergia cochinchinensis is
prioritized in Thailand for both its commercial
value and its threat status.
It is interesting to note the discrepancy
between the relatively small number of tree
species nominated for in situ conservation and
the protected-area approach that many countries
report as their primary means of conserving FGR
in situ. The number of species and populations
maintained in protected areas and other forested
lands far outweighs those in dedicated in situ
conservation programmes for priority species.
This is likely to be due to:
• the high cost and difficulties of undertaking
the research, planning, implementation
and management of dedicated in situ
conservation programmes for priority
species;
• the lack of information available for priority
species on which to base dedicated in situ
conservation programmes;
• the efficiency of the protected area
approach (which is the only option
for countries lacking the resources for
dedicated in situ FGR programmes) in
conserving the FGR of a wide range
of species and ecological processes
simultaneously.
As an example, Sweden states it “has hitherto
very few in situ genetic resources for the
approximately 30 native forest trees to enter
into the EUFGIS Portal. Obviously, there is a clear
need to improve the in situ conservation of forest
genetic resources in Sweden.” On the other hand,
Sweden has 4.7 million hectares in nature reserves
FIGURE 12.2
Reasons cited by countries for conserving species
in situ
Speciically for
genetic conservation
(10%)
Seed production
(34%)
Threatened
species
(16%)
Conservation for
non-speciied reserve
or conservation purposes
(24%)
Economic
reasons
(16%)
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and national parks which provide at least some
level of in situ FGR conservation.
Only 16 percent of priority species for in situ
conservation were nominated for economic
reasons, contrasting with 46 percent of priority
species for FGR conservation and management
in general (see Chapter 10). This difference can
be attributed to the fact that 85 percent of the
species reported as used in plantation forestry
globally are exotic species and are therefore
ineligible for in situ conservation in the countries
where they are widely planted as exotics. These
globally important, exotic industrial plantation
forestry species are usually well documented and
conserved through in situ FGR programmes in
their countries of origin (as well as in ex situ FGR
conservation programmes around the world). In
this regard countries widely acknowledge the
leadership and coordination by FAO since the
1960s, coupled with the work of internationally
active agencies and programmes such as the
DANIDA Forest Seed Centre (DFSC), CIRAD-Forêt
(the forest arm of the French Agricultural Research
Centre for International Development) and its
predecessors, CSIRO’s Australian Tree Seed Centre,
Camcore (a non-profit, international tree breeding
organization established by North Carolina State
University, United States of America) and donor
programmes of Australia, Canada, the United
States of America and several European countries.
Naturally, in countries where commercial
species are indigenous they have been nominated
for in situ FGR conservation; for example, in
Thailand both Tectona grandis and Dalbergia
cochinchinensis are the subject of in situ FGR
conservation programmes to ensure that
high-quality genetic material is available for
improvement. In Germany, important commercial
species account for 70 percent of designated
in situ conservation stands (Fagus sylvatica, 37
percent; Quercus robur and Quercus petraea,
15 percent; Picea abies, 11 percent; and Pinus
sylvestris, 7 percent); 29 other tree species account
for 22 percent of in situ conservation stands, with
shrubs accounting for the remaining 8 percent.
148
In several reports it is apparent that indigenous
tree lora have been poorly investigated for their
economic and productive potential; the United
Republic of Tanzania, for example, notes: “the
[indigenous] gene pool has more to offer…”. This
lack of research is partly being rectiied by ICRAF
and its national partners and by others such as
the Australia-funded South Paciic Regional
Initiative on Forest Genetic Resources (SPRIG)
project. From 1996 to 2005, SPRIG assisted several
countries in Oceania in research, conservation
and development of their indigenous species; this
effort has resulted in much greater planting of
native species such as Santalum species in Fiji and
Vanuatu, Endospermum medullosum in Vanuatu
and Terminalia richii in Samoa.
While 50 percent of the responses cite
conservation as the reason for nominating priority
species for in situ FGR conservation, only 10
percent speciically mention gene conservation,
while 16 percent cite conservation of threatened
species. It appears that some countries correlate
threatened species conservation with in situ
FGR conservation, whereas one seldom implies
the other. Improved understanding and
promotion of the methodology for formal in
situ FGR conservation is required to increase its
application.
Seed production purposes account for 34
percent of priority species nominations for in
situ conservation. However, as noted above,
selection criteria for in situ seed production
stands may differ from those used for in situ
conservation of FGR; for example, ease of
access may inluence seed-stand selection
but is not generally a consideration in formal
in situ FGR conservation programmes; and
the conservation of populations growing in
marginal and extreme environments and/or
uncommon alleles may be considerations for in
situ FGR conservation of a species but are less
important for seed stands providing germplasm
for commercial forestry.
A further point is that priority species
nominations only relect existing knowledge of
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
economic value and threatened status; where the
potential economic value or the threat status is
unknown, species are neither nominated nor dealt
with as priorities. Landscape-scale approaches
such as protected areas and sustainable
management of multiple-use forests, on the other
hand, provide the benefit of maintaining a large
number of species and populations regardless of
the level of knowledge, and for this reason are
more appropriate where species information is
scant or lacking.
Nevertheless, it is essential to prioritize species
for in situ conservation, because the ecosystem
conservation approach alone is unlikely to fulfil
the particular requirements of dedicated in situ
FGR conservation programmes. Both coarse-grain,
landscape scale and fine-grain, species-specific in
situ FGR programme approaches are required and
must be treated as complementary.
Some country reports describe the criteria
used to set species priorities for in situ
conservation of FGR. In China, “priority species
for in situ conservation are determined by
the existing quantity of the species, the socioeconomic value and the depletion of FGR”, while
Thailand’s three priorities are: “species with
socio-economic importance, both commercial
importance and importance for maintaining
ecosystem functions and services; species with
higher levels of genetic diversity; and species
with populations at risk or under threat
from any cause, e.g. critically endangered,
endangered or vulnerable species”.
Management of in situ FGR
conservation areas
In situ conservation of FGR involves more than
legislating for a protected area. Populations
conserved in situ constitute part of an ecosystem,
which must be managed to maintain both intraand interspecific diversity at appropriate levels
over time. Country reports, particularly from
developing countries, note serious losses of FGR
resulting from inadequate management, lack
of oversight and failure to enforce regulations
in protected areas, conservation reserves and
dedicated FGR in situ conservation stands; these
areas accordingly suffer from illegal logging
and harvesting, mining, clearing, poaching,
growth of invasive species and damage from
unmanaged wildlife, uncontrolled wildfire and
pests and diseases, often amplified through
interactions with climate change. The negative
impacts sometimes occur in the last remaining
natural stands of high-value timber species (e.g.
Tectona grandis and Dalbergia cochinchinensis
stands in Thailand) or threatened or otherwise
significant species with extremely valuable
genetic resources.
Countries all note the paramount importance
of managing in situ FGR conservation sites to
ensure the maintenance of the genetic resources,
species and ecological processes contained
therein. The Islamic Republic of Iran remarks the
requirement for “management plans to monitor
the changes in target populations over time and
ensure their continued survival”. Approaches
identified and implemented by countries include
improved and strengthened forest management;
better legislation and regulation; more effective
enforcement; preparation of management plans
and guidelines; better funding; inventories
including conservation trajectories of threatened
species and vegetation communities; expertise;
community education; incentives for improved
management of FGR; and means of generating
financial returns from sustainable management
such as creation and management of wildlife
parks, development of game-based tourism and
sustainable harvest and marketing of forest
products by local communities.
Canada notes the need for better understanding
of the risks faced by individual species and of how
to manage them – requiring research on species’
vulnerability, sensitivity and adaptive capacity; on
their habitat, physiology, phenology, and biotic
interactions; on their exposure to and ability to
respond to threats such as climate change, for
example by adapting in place or migrating; and
on at-risk ecosystems.
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Several countries note the importance of
maintaining genetic fitness and regeneration
of conserved stands. Sri Lanka notes that for
outbreeding species, large areas of contiguous
forest are required to avoid loss of diversity
through genetic drift and inbreeding. Sri Lanka
also notes the value of re-establishing “gene
corridors” to reduce the risks to fragmented,
isolated populations, and proposes conversion
or enrichment of monoculture plantations
surrounding isolated natural forest patches using
mixed indigenous species with appropriate levels
of variability.
Many countries identify well considered and
comprehensive management plans for in situ
FGR conservation areas as essential. Thailand,
for example, reports that after selection of
natural stands in different zones, “conservation
measures and management options were
proposed specifically for each zone based on
the available information on its population
size, legal protection, social aspects, commercial
interest, and management costs”. Thailand
notes that for teak, a “conservation plan
comprising a number of activities [has] been
recommended: field survey and selection of
the populations; demarcation and protection;
monitoring; and management guidelines”.
Thailand also observes the need to ensure that
regeneration occurs in the conserved stands:
“Silvicultural practices and management are
essential to promote the natural regeneration
of the existing conservation areas.” It may be
necessary to research the particular regeneration
requirements of threatened species.
This subject is addressed in more detail in
the section on sustainable forest management,
below.
Forest restoration and FGR
Many
significant
FGR
assets,
including
threatened, high-value timber and NWFP species
and forests with high genetic diversity, have been
seriously degraded through failure to regulate
extraction and prevent overharvesting. These
150
areas must be brought under management
and rehabilitated at the earliest opportunity to
prevent further decline and to improve their
condition, viability and economic utility. This
view is consistent with CBD Article 8(f) (UN, 1992)
which refers to the need to “rehabilitate and
restore degraded ecosystems and promote the
recovery of threatened species… through the
development and implementation of plans or
other management strategies”. Reafforestation
and restoration initiatives are needed especially
urgently in low forest cover countries (i.e.
those with less than 10 percent forest cover).
As recognized by China, knowledge of species
requirements and in situ restoration techniques
for endangered populations is essential. In the
Islamic Republic of Iran, degraded forests are
restored through plantation of native pioneer
species; “the main objective of rehabilitation is to
achieve ecosystem sustainability in forest area and
increased biological diversity”. Ghana undertakes
restoration of degraded “convalescent forests”
by ceasing exploitation and undertaking
management.
In northeastern Thailand, a framework species
selection approach was adopted to identify a small
number of local tree species for revegetation from
more than 350 local tree species, so as to most
efficiently restore forest cover and catalyse return
of biodiversity and regeneration of hundreds of
other tree species (Elliott et al., 2003).
Enrichment planting, whereby threatened tree
species are propagated from local germplasm
and reintroduced into areas in which they have
become depleted, is an important element of in
situ conservation consistent with Article 8(f) of
the CBD. Reintroduction into the wild subjects
the species to natural selection, allowing the
continuation of evolutionary processes. It is vital
to ensure that an appropriately high level of
genetic diversity is represented in enrichment
plantings. Zimbabwe notes that some species
believed to be extinct in the wild are conserved
in ex situ and circa situm contexts (e.g. home
gardens) and could potentially be reintroduced
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
back into natural settings. Ghana uses artificially
propagated stock to enrich forests that have been
depleted by overexploitation and lack adequate
natural regeneration. Denmark has re-established
the previously locally extinct Pinus sylvestris to its
former range in the country, using germplasm
from adjacent countries. These examples
demonstrate the value of adopting multiple,
integrated and complementary approaches to
FGR conservation, successfully combining in situ,
ex situ and circa situm conservation and site
management.
Enrichment and reintroduction plantings
require a supply of plant materials, and for
some species with particular reproductive
requirements, including a number of threatened
species, propagation may be difficult. China
notes the importance of developing propagation
techniques for threatened species, especially
those that are resistant to standard techniques,
for in situ enrichment plantings as well as for ex
situ and circa situm conservation; research may be
required to identify the best and most efficient
propagation techniques for less well known
species. Ethiopia notes a need to promote tissue
culture for mass propagation, which may assist in
situ enrichment plantings.
Opportunities from climate
change initiatives: restoration and
connectivity for in situ FGR
Several countries note opportunities for
simultaneous conservation of FGR, reduction of
carbon emissions and income generation through
REDD+ (reducing emissions from deforestation
and forest degradation in developing countries,
including the role of conservation, sustainable
management of forests and enhancement of
forest carbon stocks). A number of country
reports refer to programmes for planting and
regenerating trees over extensive areas as a
means of sequestering atmospheric carbon and
mitigating climate change. With thoughtful
planning, these programmes may also offer
significant opportunities to improve the
viability of fragmented landscapes suffering
reproductive isolation and genetic erosion, and
to restore local vegetation to its former range.
The following are some programmes cited in the
country reports.
• India’s National Mission for a Green India,
under the National Action Plan on Climate
Change, aims “to double India’s afforested
areas by 2020, adding an additional
10 million hectares”. India’s Sustainable
Landscapes programme is another example.
• Indonesia notes, “40 REDD+ pilot or
demonstration projects across Indonesia
[are] being implemented for ecosystem
restoration concessions for carbon
sequestration and emission reduction…
[and a] massive campaign programme
to plant 1 billion trees nationwide
annually [has been] launched for greening
Indonesia”.
• In Australia, federal and state governments
and NGOs (e.g. Bush Heritage Australia,
Greening Australia and Landcare Australia)
are supporting multipurpose, biodiverse and
carbon-sequestration planting programmes
that also restore and promote ecological
and evolutionary processes by reconnecting
fragmented landscapes. With appropriate
selection of species and planting materials,
these programmes could provide significant
FGR conservation benefits, for example
by using locally sourced planting stock
and incorporating depleted or threatened
species or genetic variability.
• In the Niger, a programme for farmermanaged natural regeneration has restored
over 5 million hectares of barren land into
productive agroforests through protection
of tree coppice and seedling regrowth,
simultaneously sequestering vast amounts
of carbon and restoring degraded FGR
assets and farm productivity and resilience
(Tougiani, Guero and Rinaudo, 2009). This
programme has also given farmers the
confidence to protect trees, and it respects
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the integrity and value of trees as the
property of the landowner.
Where these plantings involve reintroduction
of trees grown from local, genecologically
appropriate genetic sources back into their
former range, they are referred to as “inter situ”
plantings, meaning that germplasm is collected
and re-established in field trials or planted forests
located within the same geographical area,
which allows it to continue to undergo natural
selection under prevailing climatic conditions.
There plantings have particular benefits when
they extend into or connect with existing natural
vegetation, reducing fragmentation, increasing
gene low, supporting ecological processes and
increasing the viability of remnant forest patches
or isolated FGR assets. At the same time, these
plantings may be used to provide a range of
timber and NWFPs and may offer employment
to rural and traditional communities, especially
where access to forests is reduced through
conservation management.
In situ conservation through
sustainable forest management
As mentioned above, most in situ conservation of
FGR takes place outside protected areas, especially
in multiple-use forests and forests primarily
designated for wood production. It is vital that these
forests be managed through sustainable forest
management (SFM) principles and practices. While
almost all extant forests and trees provide some
level of in situ conservation of FGR, these beneits
may be quickly lost or diminished through forest
destruction or degradation. Countries employ
numerous SFM strategies, approaches, initiatives
and programmes to ensure that adverse impacts
on FGR are minimized while forest productivity
and other uses and services are maintained if not
enhanced. These include certiication schemes
for sustainable forest operations, identifying
and protecting stands of high conservation
value, promoting the use of indigenous species,
ensuring that natural and artiicially regenerated
stands contain suficient genetic variability and
Box 12.1
Community and participatory management
Reporting countries identify a suite of causes of
deforestation related to the failure of conservation
management to address indigenous or traditional
ownership, knowledge and practices of forest use and
management, including:
• uncontrolled access to protected areas and in
situ FGR conservation sites by neighbouring
rural communities seeking fuel, food, housing
materials and other forest products;
• a decrease in the value placed on trees and
sustainable forest management owing to the
loss of traditional or indigenous knowledge
about the uses of forest foods and plants and
traditional management systems;
• a poverty-driven increase in use pressure,
amplified by climate change and resulting in
152
encroachment of agriculture and pastoralism on
forest lands;
• conflict caused by loss of access to forests
designated as protected areas, resulting in loss
of informal control over illegal harvesting and
uncontrolled and unsustainable use.
Community or participatory management –
which involves harnessing traditional valuation
and knowledge of the use of traditional forest trees
– is increasingly used to address such causes of
deforestation and genetic erosion. In Madagascar, for
example, local communities are increasingly directly
involved in management and use of natural forests
through licensing and agreed management and
harvesting plans.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
promoting community participation in forest
management, thereby maintaining traditional
valuation, knowledge and management of forests
(Box 12.1). Some SFM certification schemes also
promote replacement of monocultures with
mixed species plantations, although it should be
noted that in certain situations this may reduce
the conservation of FGR at the landscape level,
as highly productive short-rotation monocultures
can remove harvesting pressures from indigenous
forests.
Secure in situ conservation of FGR requires
security of appropriate tenure over the area,
for example through gazettal as a protected
area, conservation reserve or production forest
permanently subject to FGR-appropriate SFM.
It also requires continuing application of
dynamic management regimes that address and
maintain genetic diversity of the tree species.
The importance of SFM is illustrated by the
observation that forests subject to harvesting
and use under SFM conserve FGR more effectively
than protected areas that are poorly managed,
uncontrolled or subject to threats such as illegal
incursions, unsustainable harvesting, wildfire,
grazing or invasive species.
Prior to the modern era and its unprecedented
population growth, many forests under
traditional and customary management were
used sustainably. Forest management has been
progressively developed and institutionalized
over the past three centuries; it was described
in German forestry texts as early as 1713 and
introduced in central Europe in 1742. SFM is now
widely recognized in many national policies and
programmes and in international processes such
as the CBD (e.g. Aichi Biodiversity Targets 5 and
7) and the Non-Legally Binding Instrument on All
Types of Forests (NLBI) (UN, 2008) as a means of
achieving conservation outcomes simultaneously
with multiple productive uses. The requirement
for the sustainable use of forests is enshrined
in CBD Articles 8(c) (in situ conservation) and
10 (sustainable use of components of biological
diversity) (UN, 1992).
Box 12.2
Global Objectives of the Non-Legally
Binding Instrument on All Types
of Forests
Global Objective 1: Reverse the loss of forest cover
worldwide through sustainable forest management,
including protection, restoration, afforestation and
reforestation, and increase efforts to prevent forest
degradation.
Global Objective 2: Enhance forest-based
economic, social and environmental benefits,
including by improving the livelihoods of forestdependent people.
Global Objective 3: Increase significantly the
area of protected forests worldwide and other
areas of sustainably managed forests, as well as
the proportion of forest products from sustainably
managed forests.
Global Objective 4: Reverse the decline in official
development assistance for sustainable forest
management and mobilize significantly increased,
new and additional financial resources from all
sources for the implementation of sustainable forest
management.
Source: UN, 2008.
As defined in the NLBI, “sustainable forest
management, as a dynamic and evolving concept,
aims to maintain and enhance the economic,
social and environmental values of all types of
forests, for the benefit of present and future
generations” (UN, 2008). FGR conservation and
management are crucial to the achievement of
these aims and need to be explicitly recognized
and steadfastly incorporated into SFM. The NLBI
sets out general objectives for SFM (Box 12.2)
which are also relevant to in situ FGR conservation.
SFM applies to all activities and uses of forests
and trees, both extractive (e.g. harvesting of
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Box 12.3
Addressing genetic resources in sustainable forest management plans
FGR conservation and management are primary
components of SFM and must be incorporated in
SFM plans. Information is required on the genetic
resources and associated evolutionary processes
and on how these are affected by use. Appropriate
SFM approaches must then be selected to minimize
negative impacts. An example might be to adopt a
system for valuing FGR and to provide payments or
other rewards for FGR stewardship or conservation.
Written plans provide objectives and benchmarks
against which outcomes and performance can be
measured, thus facilitating accountability. Plans
are vital for monitoring, auditing, evaluation and
improvement and provide a documented frame of
reference for future adaptive management of FGR.
timber and NWFPs) and non-extractive (e.g.
forest-based tourism). To achieve effective in situ
conservation of FGR, SFM plans incorporating
FGR goals and objectives need to be prepared
and implemented for all forests subject to use
(Box 12.3).
Production forestry remains a major focus
of SFM (Figure 12.3). FRA 2010 reports that 39
percent of the world’s forests are primarily used
for production of wood and NWFPs; an additional
24 percent are designated for multiple uses that
usually include the production of wood and/or
NWFPs (FAO, 2010a). The proportion of natural
forests subject to timber harvesting varies greatly
among regions and countries. While a few
countries ban harvesting in natural forests or limit
it to small areas, other countries maintain large
areas of natural forest for production (notably in
Europe and Africa) or multiple use which often
has timber production as a major objective (e.g.
in North America). In Indonesia, 60 percent of
total forest area is managed as production forest.
Germany manages most of its forests jointly for
production forestry and conservation.
154
Some countries comment that lack of funding,
organizational capacity, political support and/
or jurisdiction over forests under customary or
private tenure makes it difficult to implement FGR
conservation and management through their SFM
plans. Adequate finance is vital, whether allocated
from government budgets or raised from forestry
activities (e.g. licence or royalty payments). Capacity
building for incorporation of FGR concerns in SFM
programmes is essential but is often lacking in
developing countries. Some countries (e.g. the
Islamic Republic of Iran) report support to community
participation in the preparation of SFM plans to
facilitate sustainable outcomes.
While protected areas make an essential
contribution to FGR, they comprise less than 13
percent of forests worldwide (see Chapter 6).
Productive uses may occur in various categories
of protected areas. Many countries’ protected
areas include multiple land use designations
that allow for a range of uses and activities,
such as limited timber extraction, harvesting of
NWFPs, grazing, hunting and tourism. A number
of countries also note that protected areas are
subject to encroachment and illegal occupation.
SFM involves both ensuring that approved
activities within protected areas are conducted in
such a way as to minimize impacts on the genetic
resources, and preventing (or, if prevention is not
possible, at least reducing) uncontrolled, illegal
uses.
Forest management, as sometimes currently
practised, may not always adequately address
the requirements of in situ conservation of FGR,
for example in situations where maximizing
timber production is emphasized at the expense
of genetic and ecological outcomes, which are
important components of SFM. It is essential
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
FIGURE 12.3
Number of species mentioned as actively
managed in country reports, by main
management objective
2 500
2 000
1 500
1 000
500
0
Total species
managed
For timber
For non-wood
forest products
For energy
to ensure that SFM protocols address the
requirements for conservation of FGR, both for
major commercial timber species and for other
forest tree species whose continued presence
and variability may be affected by timber
extraction, especially those providing NWFPs.
Nonetheless, for countries where forests are
currently harvested with inadequate controls and
at unsustainable rates, applying the basic forest
management principle of sustainable yield can
contribute significantly to FGR conservation.
China expresses the concern that population
growth, increasing wealth and consumer demand
for timber and wood products will result in a
significant shortfall between national demand
and supply, and the same may hold for other
fast-growing economies in Asia such as India
and Thailand. Some traditional timber-exporting
countries in Africa (e.g. Ghana) have experienced
major declines in exports and may eventually
become net importers of wood products. The
long-term challenge of meeting rising global
demand for wood products will necessitate
adoption of SFM, including in situ conservation of
FGR, in all forms of production forestry. Demand
for NWFPs and environmental services of forests
is also increasing; with proper management to
avoid rapid depletion of the resources, these
uses can usually be fully compatible with FGR
conservation. SFM plans require regular updating
to meet new circumstances including changes in
demand and production for diverse NWFPs (e.g.
cork, honey, mushrooms, wild game, essential oils
and bamboo) and environmental services (e.g.
biodiversity conservation, carbon sequestration
and restoration of degraded soils).
Customary management practices are often
consistent with sustainable management of
FGR. When nearby impoverished communities
are excluded from protected forests that they
previously lived in and used following customary
and sustainable management regimes, the result
is often uncontrolled harvesting and use. Where
SFM makes productive uses possible with minimal
impact on the environment, including FGR and
associated processes, it may be appropriate to
permit productive activities in protected areas
or conservation reserves. Providing controlled
access and shared management responsibilities
to communities under appropriate SFM plans
can reduce negative impacts on FGR and even
provide positive conservation benefits. For
example, involving neighbouring communities in
informal surveillance can help reduce illegal and
detrimental activities.
The following sections outline SFM approaches
and initiatives that contribute to the maintenance
of genetic variability and evolutionary processes.
Reducing use and harvest pressure on
natural forests
The application of SFM to increase the production
of timber and NWFPs from natural forests can
reduce the harvest pressure on other forests
conserved for biodiversity and FGR purposes and
free up production forest for other uses. SFM
can deliver increased timber production from
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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the same or reduced area of natural production
forest. Indeed, a number of countries report
slight increases in forest area, increment of forests
and/or standing volume of timber in production
forests through the application of SFM principles
and associated legislation, including for FGR.
Sweden’s standing wood volume, for example,
has increased by 86 percent over the past 90 years,
with the annual increase in standing volume
doubling over the same period. Although the
forest area in Finland has remained constant, the
volume of timber is 40 percent greater than 50
years ago, and 45 percent more volume is added
annually than is harvested. Standing volume has
also increased in Germany. The United States of
America reports an increase in timber production
with a fairly constant forest base and increasing
standing volume.
Sustainable increases in timber and NWFP
production may be achieved through the
application of SFM practices such as selected seed
sources and improved genetic materials, altering
rotation length, species-specific minimumdiameter cutting limits, enrichment plantings of
preferred timber species, thinning operations,
retention of mother or seed trees, retention of
preferred NWFP species during clearing, and
applying the principle of sustainable yield, where
the amount harvested does not exceed the
annual increment.
To increase production from existing natural
forests through better management, Ghana has
prioritized increasing investment in natural forest
management to relieve the utilization pressures
on forests that have FGR and conservation value.
Sweden remarks that more intensive forestry
to increase forest growth requires an increased
effort in gene conservation to balance production
and environmental objectives. In Japan, national
forests that are not protection forests are also
managed sustainably through long-term regional
forest management plans and contribute to FGR
conservation.
Another strategy is to increase the area of
forest available for productive uses and FGR
156
conservation through planting and facilitation
of regeneration. A number of countries
recorded large annual net gains in forest
area between 1990 and 2010 (FAO, 2010a)
(see Chapter 6). In some cases, especially in
Asia, these gains were the result of deliberate
reforestation programmes, while in southern
Europe and the United States of America
the gains may be attributed in large part to
natural regeneration on abandoned, marginal
agricultural lands. Solomon Islands identifies an
urgent need to rehabilitate harvested areas that
have failed to regenerate, as a step to restoring
the productivity of such areas. This strategy
may extend to increasing farm and landscape
plantings and expanding planted forest areas
to meet the demand for forest products and
services to avoid adding to the use pressures on
natural forests; these plantings may include tree
species and populations under pressure in situ,
thereby providing circa situm FGR conservation
benefits.
Increasing planting to reduce pressure on
natural forests is an important adjunct to
SFM strategies. Ghana proposes intensive
planted forest development for this purpose.
Indonesia is one of several countries adopting
this strategy: “To reduce pressure from natural
forest exploitation, the Ministry of Forests has
increased permits for industrial plantation forest
from 4.5 million hectares in 2000 to 8.97 million
hectares in 2010.” Myanmar, Solomon Islands
and Vanuatu similarly identify this approach
as vital to reducing unsustainable harvesting
of natural forests, including particular highvalue species. Plantations enable Brazil to be
extremely efficient at producing pulpwood for
manufacture of paper. Its 1.1 million hectares of
pulpwood plantations represent just 0.2 percent
of the country’s area. Brazil states that the wood
harvested from 100 000 ha produces 1 million
tonnes of pulp annually in Brazil, while in
Scandinavia and the Iberian Peninsula, 720 000
and 300 000 ha, respectively, would be necessary
to obtain the same amount.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
From an FGR conservation and management
and long-term sustainability viewpoint, it is
essential that existing, genetically diverse natural
forests not be replaced with monocultures of
limited genetic variation. In many locations,
abandoned or marginal agricultural land may be
suitable for planted forest establishment, thereby
avoiding loss of natural forest. Other strategic
measures adopted by countries to facilitate
planting include provision of low-interest
loans for planting programmes (particularly of
multipurpose tree species) and for forest-related
cooperatives; and ensuring that land for planted
forest establishment is affordable.
Application of sustainable yield principles can
contribute to sustainable forest management
and to conservation and management of FGR.
Implementation of sustainable yield principles
requires guidelines, inventory and FGR survey,
planning and effective administration. In
Solomon Islands – where roundwood accounts
for over 70 percent of export earnings, provides
18 percent of total government revenue and
is a major source of income and employment
in rural areas as well as for landowners –
unsustainable extraction threatens to deplete the
resource substantially by 2018, which could have
destabilizing effects on the economy and society.
Institution of a sustained yield regime would be
a critical step in improving forest management
and would provide the basis for conservation of
FGR. In Vanuatu, sustainable forest management
is enshrined in the constitution and backed by
a Forestry Act, National Forest Policy and Codes
of Logging Practice, but its implementation is
hampered by lack of land use planning, out-ofdate forest inventory, inadequate government
resources and a gross imbalance between
extraction and reforestation.
Another important approach in relieving
harvesting pressure on natural forests is to
establish alternative uses and income generation
opportunities for forests, such as ecotourism,
conservation, water provision and environmental
protection. Seychelles has incorporated this
strategy and the alternative uses into high-level
government policies and programmes to ensure
that they are adequately addressed at the field
level. This approach is consistent with Part V,
Paragraph 6(j) of the NLBI (UN, 2008), which
suggests encouragement of “recognition of the
range of values derived from goods and services
provided by all types of forests and trees outside
forests, as well as ways to relect such values in the
marketplace, consistent with relevant national
legislation and policies”. Markets for alternative
goods and services may already be available
through mechanisms such as forest certiication
schemes, stewardship payments, payments for
carbon sequestration or credits for biodiversity,
land or environmental management.
A number of countries seek to provide
alternative goods and services to replace natural
forest products – for example, alternative energy
sources to lessen the demand for fuelwood and
charcoal, which often contributes to unsustainable
harvesting. Seychelles reports that “the demand
for fuelwood appears to be rapidly dwindling
due to rural electriication and expanding use of
kerosene also in the rural areas”. Zambia proposes
rural electriication and more eficient technology
for burning wood and charcoal as strategies for
reducing the demand for charcoal, which is at
present partially responsible for loss of FGR. The
Philippines mentions efforts to extend the life of
construction materials to reduce the requirement
for wood for repairs and replacement. Countries
with less pressure on the forests and/or an
expanding production forest estate, on the other
hand, may promote the use of wood because it has
a lower negative environmental impact than other
products (e.g. Norway).
Value addition to forest products is increasingly
used as a strategy to counter unsustainable highvolume/low-return forestry based on extraction
and export of roundwood. Gabon has prohibited
roundwood exports since 2009 to encourage
domestic processing, while Papua New Guinea
and Solomon Islands have developed domestic
processing targets and projects to add value to
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logs in order to provide greater employment and
increase financial returns to the population as
well as government revenues. Solomon Islands
has a policy that 20 percent of timber logged
under licence must undergo further processing,
implemented as a condition of logging licences,
to facilitate value addition.
Zoning: identifying and protecting
important areas for increased protection
and management
As with more targeted in situ conservation
approaches, SFM for maintenance of FGR
requires that key tree populations or areas
containing priority or threatened tree species or
other essential elements for maintaining FGR be
identified, demarcated and given extra protection
and appropriate management. This approach is
practised by many countries, including Finland,
which identifies and protects valuable habitat
areas within production and other forests; and
Thailand, which in 1989 gazetted National Forest
Reserves as three zones (conservation, economic
and agricultural). Nonetheless, some countries
(e.g. the Philippines) report failure to protect
significant areas for conservation of FGR and
biodiversity once they are identified, even when
protocols have been set out in codes of practice
and other programmes, because of administrative
failure, lack of political will or inadequate
funding.
A number of countries apply land-use planning
to allocate production forestry and plantation
expansion activities to areas where they will
deliver the maximum benefits with the least
impacts, as well as to identify areas suitable
for agricultural expansion. It is essential that
conservation of FGR be taken into account during
land-use planning.
Multiple-use approach to production
forests
Adopting an explicitly multiple-use approach to
production forests facilitates the incorporation
of FGR conservation and management objectives
into their management. Multiple uses may include
158
timber and NWFP harvesting, conservation of
genetic resources and biodiversity, catchment
management and water provision, environmental
protection, CO2 sequestration and recreation.
The various uses must be compatible, i.e. no use
should have negative impact on the others in the
area. In many cases management for multiple
uses is not only complementary to, but may also
assist, FGR conservation and management. For
example, Seychelles reports that allowing honey
production in protected areas helps prevent
damaging fires as beekeepers protect their hives
and nectar sources. Germany’s multiple-use forest
management approach is particularly directed
towards simultaneous production and genetic and
biodiversity conservation, based on “protection
through utilization”. FGR conservation in
multiple-use forests requires detailed information
about the genetic resources and the requisites for
their management requirements, as well as high
organizational capacity.
FGR conservation can also be integrated
with plantation management, for example
through protection of forest buffers along
watercourses or planting of corridors of native
forest species to link retained natural forests and
minimize fragmentation. Indonesia notes that
elimination of regeneration on new plantation
sites prior to planting results in failure to
meet FGR conservation objectives. By contrast,
establishment of mahogany through enrichment
planting in logged-over forest in Fiji has allowed
regeneration and continued growth of many
native tree and understory species, including
species important for FGR conservation such
as Endospermum macrophyllum and Atuna
racemosa.
“Close-to-nature” forestry practices
Adopting management techniques that rely on
or mimic natural forest processes is an important
SFM approach. Natural regeneration of harvested
blocks is one such approach: The countries
reporting on regeneration methods for this report
predominantly regenerate their commonly used
species using natural regeneration (61 percent,
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
as compared with 30 percent regenerated
through planted forest establishment). Natural
regeneration can be lower in countries with
forest production based on indigenous species;
in Canada, for example, about 32.8 percent of
logged forest is regenerated naturally.
European
countries
are
increasingly
applying “close-to-nature” approaches to
forest management that emphasize natural
regeneration, and the approach is also under
investigation elsewhere. Denmark’s 2002 National
Forest Programme recommends adoption of
the close-to-nature approach, and the Forest
Act of 2004 provides the legal framework for
this transition. Germany proposes that close-tonature forestry is the most effective approach for
simultaneous forest production and conservation;
the country’s report notes that “[owing] to the
strict legal provisions and largely practised closeto-nature forestry, overexploitation and clearcutting are no real threat to the state of the
forests’ genetic diversity in Germany”.
Codes of practice
Codes of practice provide guidance for conducting
operations according to SFM principles and,
where relevant, for complying with government
legislation or regulations. They may apply to
the operations of State, parastatal and private
companies or individuals engaged in forest use
on public land, and their use may extend through
legislation to timber extraction activities on
private land. They may be imposed as a condition
of access to public forest resources or government
licences for their use, thereby forming part of the
regulatory system, or they may be developed by
forest industry for voluntary use. Some codes may
be developed as part of certification schemes or to
demonstrate best practices in the marketplace. It
is vital that codes of practice for SFM incorporate
measures to ensure the maintenance of FGR and
the ecological processes that support them. In
Australia, for example, several state jurisdictions
have codes of practice governing forest operations
on both private and public land; these include
provisions for maintaining ecological assets and
conserving non-target species.
For codes of practice to be successful in
promoting SFM and FGR conservation and
management, effective forest and natural
resource
administrations
are
required.
Mechanisms must be in place to ensure
compliance, such as enforcement of penalties
for breaches, implementation of provisions for
damage, lodgement and potential forfeiture of
bonds, fines, cancellation of licences or permits to
operate or denial of access to the forest resource.
Furthermore, the compliance of forest operators
must be monitored. Codes must have legislative
or legal backing to be enforceable.
Some countries report instances of failure
to implement SFM and FGR conservation and
management protocols contained in codes of
practice because of lack of funding or human
resources, insufficient administrative and technical
capacities or lack of jurisdiction over important
forest resources requiring conservation. The
effectiveness of codes of practice in the private
forest estate may depend on the government
having jurisdiction over forest protection
activities and harvesting on private land, for
example through granting of permits. Where
codes of practice exist for activities that can have
impact on FGR but lie beyond the normal purview
of forest operations, these codes must also
address the requirements of FGR conservation
and management. Harmonizing a national FGR
strategy with relevant cross-sectoral programmes
will facilitate the integration of FGR concerns into
codes of practice for related activities.
Harvesting
To conserve and manage FGR effectively,
sustainable harvesting regimes must address the
genetic characteristics of any species, including
but not limited to target or priority species, that
could potentially suffer negative impact from
harvesting, as well as associated evolutionary
and ecological processes. Areas to be conserved
for their FGR value must be identified and,
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if necessary, protected from harvesting. The
harvesting intensity and techniques must
minimize impact on non-target species, and
the harvesting and regeneration practices
must ensure that the genetic variability of the
target species is retained as much as possible.
Monitoring the impacts of forest utilization and
applying appropriate management are essential
elements of SFM.
A number of countries note negative impacts of
unsustainable harvesting regimes and techniques
on their FGR, most significantly the depletion or
extermination of the target species and/or others.
The Islamic Republic of Iran reports a research
programme to investigate damaging silvicultural
practices, while some countries (e.g. Papua New
Guinea, Solomon Islands, the Sudan, Thailand and
Viet Nam) have banned harvesting of selected
endangered and economically important timber
tree species.
Maintaining germplasm variability in
regenerated production-forest stands
Ensuring that harvested stands are regenerated
adequately and contain sufficient variability for
adaptation and continued evolution is crucial
to SFM for FGR conservation and management.
This is particularly important for countries with
significant forest industries that rely heavily on
natural forest and with limited strictly protected
areas. Most commercial timber tree species
regenerate naturally and maintain sufficient
variability as long as enough trees contribute
to the reproductive materials used for stand
regeneration.
Regeneration must be monitored and may need
to be supplemented with artificial seeding or
planting if stocking rates are inadequate. Where
artificial regeneration is the preferred method of
restocking, then genetically diverse germplasm
is essential. For example, Germany mandates the
minimum number of seed trees to be used; with
the large number of seed stands available and
the system of private certification of germplasm
suppliers, “sufficient amounts of reproductive
160
material with high genetic diversity are available
for artificial regeneration of most provenances”.
Germany also notes that incorporating noncommercial tree species into natural regeneration
regimes and cultivating “adapted populations
with great genetic diversity” are particularly
beneficial for in situ conservation.
In order to maintain the variability of
germplasm in the 15 percent of production
forests in Canada that are artificially regenerated,
the country’s criteria and indicators for SFM state
that:
“Under Criterion 1, biological diversity,
Indicator 1.3.1, genetic diversity of
reforestation seed lot, addresses the
variation of genes within a species by
ensuring that seed used to regenerate
harvested areas has sufficient genetic
diversity to respond to changing
environmental conditions… The genetic
diversity of seed used for reforestation
is a result of both the number of areas
where seed is collected and the parental
composition of those areas. Most of the
seed used in reforestation programmes
across Canada is collected from natural
stands where the number of parent trees
is typically in the hundreds to thousands.
This seed likely has genetic variation that is
representative of the natural populations
where it was collected. [However] in some
jurisdictions, a significant portion of the
seed for reforestation… comes from seed
orchards.”
Ideally seed is collected beforehand from the
area to be harvested and regenerated. However,
where this is not possible, germplasm of local
provenance or from the same genecological
zone may be used to maintain variability and
ensure that the stock is well adapted to local
conditions. Guidelines based on knowledge of
the target (or closely related) species may be used
to guide collection and use of germplasm for
artificial regeneration to maximize variability in
situations where genetic variation patterns are
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
known partially or not at all. Germany points out
the need to monitor the genetic impacts of its
silvicultural practices on its priority species, and
is developing a methodology to facilitate this
monitoring. Early work suggests that losses to
date are not excessive. Solomon Islands notes a
failure of natural regeneration and a failure to
undertake remedial measures, which comprise a
threat to the survival of the country’s forests, FGR,
and sustainable forest industry.
Countries increasingly recognize the need to
provide higher genetic variability (both more
species and greater intraspecific diversity) in
production forestry, especially for traits associated
with adaptability to changing climatic and
environmental conditions. Some countries have
identified this as a priority for their breeding
and forest management programmes; and some
seek to incorporate variability that can confer
adaptation to climate change into germplasm
used for regeneration of harvested areas as well
as different types of planted forest.
Using local species
Country reports indicate that there is a global
trend of increasing use of indigenous tree species
for afforestation and environmental services. The
use of locally indigenous tree species in plantings
for timber production, fuelwood and afforestation
can assist in maintaining genetic variation,
especially where planting stock is developed from
genecologically appropriate and genetically variable
germplasm and management regimes do not
excessively deplete variability in non-target species.
The use of indigenous species as an alternative to
commonly planted exotics can yield major FGR
benefits. Nonetheless, consideration must be given
to reducing the risk of loss of local genetic variability
which can occur if genetic material from improved
indigenous species enters local populations of the
same or hybridizing species.
Using local tree species has the added benefit
of raising awareness of their value. The United
Republic of Tanzania recommends further use
of its valuable, high-performing indigenous
species such as Antiaris usambarensis and Khaya
anthotheca, whose growth rates “compare
closely with [those of] industrial plantation
species raised in the country”, in afforestation
programmes. However, where high-performing
exotic species yield a more valuable crop than
local species with minimal impact on FGR, their
use may not only be commercially preferable but
can also lower pressures on a country’s natural
forests. This approach may form part of a highlevel policy and strategy for SFM that addresses
FGR conservation and management.
Increasing variability in establishment of
planted forests
As reported by Indonesia, land preparation
for forest plantation or agroforestry in cleared
forest areas often involves removal of all residual
species and regrowth. Instead, partial retention
of advanced growth and natural regeneration,
especially of economically valuable local tree
species, can assist in maintaining diversity with
minimal economic losses.
Several countries are considering or have
undertaken conversion of monoculture plantations to mixed-species plantings to increase
variability for a variety of reasons, for example to
provide opportunities for adaptation to climate
change or to increase genetic connectivity among
isolated remnants. Germany notes that climate
change may necessitate the transformation
of pure spruce stands in many parts of the
country to mixed stands of putatively adapted
tree species. Mixed-species plantations serving
multiple purposes are increasingly finding favour,
for example under initiatives for reafforestation
in the Islamic Republic of Iran.
Other practices to minimize losses of FGR
during planted forest development may include
documenting and conserving ex situ any threatened
local FGR, avoiding conversion of natural forest
to planted forest, and ensuring genetic diversity
within planted stock across a range of adaptive
traits while maintaining uniformity for desired
production and quality traits.
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Certification: increasing market share for
sustainably produced products
Forest certification schemes specify management protocols that are in accordance with
accepted SFM principles. To obtain certification,
organizations involved in forest management
and harvesting, whether public or private, must
provide proof of compliance with the protocols
and undergo regular auditing to ensure ongoing
compliance. Certification of a product may confer
a marketing advantage. This approach to SFM is
consistent with the NLBI, Part V, Paragraph 6(x),
which encourages “the private sector, civil society
162
organizations and forest owners to develop,
promote and implement… voluntary instruments,
such as voluntary certification systems or other
appropriate mechanisms, through which to
develop and promote forest products from
sustainably managed forests…” (UN, 2008).
Principle 9 of the predominant certification
system, the Forest Stewardship Council (FSC),
is “Maintenance of high conservation value
forests”. The standards specify that management
activities in high conservation value forests must
maintain or enhance the attributes that define
them.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Chapter 13
Ex situ conservation
Internationally,
governments
and
nongovernmental agencies and organizations
have made strong commitments to ex situ
conservation. The CBD Global Strategy for Plant
Conservation 2011-2020 promotes efforts at all
levels – local, national, regional and global – to
understand, conserve and use sustainably the
world’s immense wealth of plant diversity while
promoting awareness and building the necessary
capacities. The 2011-2020 strategy includes 16
outcome-oriented global targets for 2020; Target
8 specifically addresses ex situ conservation: “At
least 75 percent of threatened plant species in ex
situ collections, preferably in the country of origin,
and at least 20 percent available for recovery and
restoration programmes” (CBD, 2013).
Global infrastructure commitments can be best
depicted by, for example, the Svalbard Global
Seed Vault in Norway, which was built in 2007 to
provide insurance against the loss of germplasm
from seed banks as well as a refuge for seed in
the case of long-term regional or global crisis
(Global Crop Diversity Trust, 2013), and the
Millennium Seed Bank Partnership of the Royal
Botanic Gardens, Kew, United Kingdom (see Box
3.3 in Part 1), which both store tree seed.
Increased emphasis in recent years on expanding collections in botanical gardens, especially
in China and other highly diverse countries,
has contributed significantly to knowledge of
cultivation of tropical tree species, particularly
endemics. The global network Botanical Gardens
Conservation International (BGCI) has a mission
“to ensure the world-wide conservation of
threatened plants, the continued existence of
which are intrinsically linked to global issues
including poverty, human well-being and climate
change” (BGCI, 2013). The BGCI network currently
has 700 members in approximately 118 countries.
BGCI supports ex situ conservation through many
of its activities, in particular the establishment of
regional networks that strengthen and support
botanical gardens such as the African Botanic
Gardens Network (ABGN).
Based on data obtained from the country
reports, the total number of species conserved
ex situ is 1 800. The number of species conserved
by region is shown in Table 13.1; see Figure 12.1
in Chapter 12 for a comparison with species
conserved in situ. Many of these are in botanical
gardens. Some 95 percent of the species are
native in the countries where they are conserved;
8 percent are exotic. (The total exceeds 100
percent because some species are present both in
countries where they are native and in countries
where they have been introduced.) In all regions
the majority of species conserved ex situ are
native species.
TABLE 13.1
Species conserved ex situ, by region
Region
Africa
No. of species
1 025
Asia
389
Europe
401
Latin America and the Caribbean
372
Near East
102
North America
265
Oceania
57
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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TABLE 13.2
Genera of global priority that are conserved
ex situ
Genus
No. of species conserved
Pinus
65
Eucalyptus
28
Albizia
24
Acer
15
Quercus
13
Acacia
10
Terminalia
10
Of the 2 260 species listed as priorities in country
reports (see Chapter 10), 626 are represented in
some form of ex situ conservation, and 135 of
them are conserved in more than one country.
Considering genera of global priority, Pinus
has the largest number of species conserved ex
situ (Table 13.2). Globally the total number of
accessions reported is 159 579, and for some
species there are multiple accessions. Most
accessions are as field collections, including clone
banks and provenance trials; far fewer are in seed
or in vitro collections.
Ex situ conservation activities by
region
Africa
Ex situ conservation in African countries is
primarily achieved through arboreta and
botanical gardens, but other means such as
provenance trials, plantations, seed orchards and
seed banks are also used.
Short- and long-term storage of germplasm is
not available in all African countries, and where
that capability exists the capacity varies. Burkina
Faso has one of the most operational seed centres,
established in 1983, which serves as a seed bank
for medium- and long-term storage; it is a partner
in the Millennium Seed Bank Partnership, focusing
on research, seed and herbarium specimen
collection and conservation of duplicates from
164
its collections. Burundi has capacity for mediumand long-term cold storage of seed. Madagascar
has one seed bank, functional since 1986, which
collects seed from rare, threatened and valuable
tree species. Morocco has four facilities for
production and storage of tree germplasm;
these facilities are equipped for conditioning,
conserving, organizing and managing seed lots.
Tunisia has two gene banks; the largest has
the capacity to store 200 000 accessions. The
Zimbabwe Tree Seed Centre has approximately
23 000 accessions. Burkina Faso and Cameroon
have infrastructure for storage but report that
frequent power outages and lack of funds for
maintenance present major challenges. In Malawi,
the Millennium Seed Bank assists in backing up
the ex situ conservation collections.
The tree species conserved in African countries
include indigenous and exotic species from many
genera and families. The main genera conserved
and propagated for multiple reasons are Acacia
spp., Eucalyptus spp. and Pinus spp. Some
countries (e.g. Benin and Burundi) report that
they conserve species valued for medicinal uses.
Mauritania’s main driver for ex situ conservation is
utility for mitigating the impacts of desertification
and restoring degraded ecosystems.
In addition to traditional ex situ conservation
methods such as seed and live field collections
of all kinds, Burkina Faso, Cameroon, Ethiopia,
Gabon, Kenya, South Africa and Zimbabwe
report that they have genetic improvement
programmes to select plus trees for different
purposes including seed orchards. Others (Ghana
and the United Republic of Tanzania) note that
their improvement programmes are moribund or
in decline because of lack of funding. In many subSaharan African countries, ICRAF has partnered
with national government agencies, universities
and others to implement domestication and
improvement programmes for indigenous
multipurpose tree species (see Chapter 14).
A few countries, including European countries
and some developing countries (e.g. Algeria),
have tried in vitro propagation methods such
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
as somatic embryogenesis and axillary bud
inductions for some of their priority species.
Limitations and constraints are numerous,
primarily lack of monetary and human resources,
proper equipment or facilities for effective
storage and management of germplasm for
short- and long-term safekeeping. Another
commonly reported limitation is a lack of
technical knowledge and training for the proper
management of ex situ conservation programmes,
for example relating to reproduction, propagation
and storage, data logging and management
strategies. Future priorities reported by many
African countries include the development of a
national strategy and research programmes for
the management of ex situ conservation and
nomination of priority FGR; and identification
of ways to fund, develop, and maintain proper
infrastructure and collections for the safekeeping
of FGR.
Species conserved. Two African countries,
Ethiopia and Burkina Faso, report details
pertaining to species conserved ex situ. Both
countries have ex situ conservation collections
for native Acacia spp. and Tamarindus indica and
for exotic Eucalyptus spp. (Eucalyptus globulus in
Ethiopia and Eucalyptus camaldulensis in Burkina
Faso).
In Burkina Faso, six native species (Acacia
nilotica var. adansonii, Acacia senegal, Faidherbia
albida, Khaya senegalensis, Parkia biglobosa and
Tamarindus indica) and four exotics (Eucalyptus
camaldulensis, Leucaena leucocephala, Prosopis
chilensis and Prosopis juliflora) are conserved
in field collections, including clone banks, and
in seed collections. The native species with the
largest number of field stands and accessions are
P. biglobosa and F. albida. Eucalyptus camaldulensis
is represented by 101 clones in two clone banks.
In Ethiopia, 92 native species and one exotic
are conserved in multiple field and seed bank
collections. The accessions from all native species
include one or more field stand and ex situ
seed bank collections. Phytolaca dodecandra is
the native species with the largest number of
collections – 59 accessions over 19 field stands and
59 accessions represented in three seed banks.
Five native species are represented by more than
20 accessions over multiple field stands: Acacia
etbaica, Cordia africana, Morinaga stenopetala,
Oxytenanthera
abyssinica
and
Phytolaca
dodecandra. The only exotic species conserved ex
situ, Eucalyptus globulus, is represented by ten
accessions available from one stand; these ten
accessions are also conserved in one seed bank.
Asia
Reported ex situ conservation activities in Asia
include provenance trials, seed orchards, clonal
repositories, botanical gardens and arboreta, and
seed and pollen gene banks targeting multiple
native and exotic species.
Community-based
ex
situ
conservation
programmes are emerging in a number of
countries in the region. For example, in Nepal
the government has endeavoured to shift
responsibility for managing seed orchards to local
communities by providing a mechanism through
which the communities receive direct benefits
from the resources. However, adequate benefits
have not yet been generated, reportedly because
of poor market linkages.
As recorded by Kyrgyzstan, studies under
Bioversity International, the United Nations
Environment Programme (UNEP) and the Global
Environment Facility (GEF) are establishing
demonstration sites and forest reserves in
Central Asian countries (Kazakhstan, Kyrgyzstan,
Tajikistan, Turkmenistan and Uzbekistan) and
conserving wild fruit-tree species in accordance
with State legislation. These efforts aim to
support local farmers through on-farm ex situ
conservation and use of local tree species,
primarily fruit- and nut-trees.
Commonly reported constraints include a
lack of resources to support ex situ conservation
activities, especially limited infrastructure and
lack of trained personnel. Country priorities for
future ex situ conservation activities include the
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development of national cooperatives to promote
FGR conservation and to enhance countries’ selfsufficiency with regard to seed supply.
Species conserved. In Asia, no significant trends
were evident among countries concerning which
native species were conserved ex situ. In both
India and Nepal Dalbergia sisso, a deciduous
rosewood tree, is a main species for ex situ
conservation in field collections (e.g. provenance
and progeny trials). A few Central Asian
countries have ex situ collections that focus on
agroforestry species. For example, Kazakhstan’s
largest ex situ collection is for Malus sieversii,
the primary wild ancestor to most cultivars
of domestic apples, and Uzbekistan’s largest
collections are for nut-bearing trees, Juglans
regia and Pistacia vera.
In China, field collections for ex situ conservation
include progeny and provenance tests for native
and exotic tree species. Four native tree species
(Cunninghamia lanceolata, Larix olgensis, Pinus
massoniana and Pinus sylvestris var. mongolica)
are conserved in over 1 200 accessions in one
or more stands for each species as well as in
extensive clone banks. Of 16 native tree species
stored in seed banks, six (Melia azedarach, Pinus
bungeana, Pinus massoniana, Pinus sylvestris var.
mongolica, Pinus tabulaeformis and Sophora
japonica) each have over 1 000 accessions. For
exotic species, from 62 to 646 accessions are held
for each of ten species (nine Eucalyptus species
and Tectona grandis) in field stands.
India has numerous species in ex situ
conservation field collections. Most are native
(e.g. Acacia nilotica, Azadirachta indica,
Dalbergia sissoo, Tectona grandis and bamboos),
but India also has a few extensive collections
of exotic species. For example, 3 122 accessions
of Acacia auriculiformis are conserved in field
stands and 4 548 accessions of Hevea brasiliensis
in clone banks. India’s ex situ germplasm storage
collections also focus primarily on native species,
and these collections are less extensive than
the ex situ field banks. The largest seed storage
collection is for a native species, Prosopis
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cineraria, with 453 accessions, while the largest in
vitro collection is for an exotic species, Jatropha
curcas, with 145 accessions.
Japan’s ex situ conservation activities are for
native species and focus primarily on clone banks
and seed storage. Collections for some species
are extensive: 7 812 clones in field collections and
2 298 seed accessions of Cryptomeria japonica;
2 452 field clones and 1 515 seed accessions of
Chamaecyparis obtusa; and 2 450 accessions
in clone banks and 508 seed accessions of Larix
kaempferi.
In Kazakhstan, seven native species (Aflatunia
ulmifolis, Alnus glutinosa, Berberis karkaralensis,
Corylus avellana, Juniperus seravschanica, Malus
sieversii and Quercus robur) are conserved ex situ
in field collections, and ex situ seed stores are also
kept for these species.
Nepal identifies 36 native species under some
form of ex situ conservation. Of these, 24 species
are in field stands and 14 species are maintained
in in vitro collections. No ex situ seed collections
are reported for any of the 36 species. Under ex
situ field collections, Dalbergia sissoo has seven
stands, while Cinnamomum tamala and Leucaena
leucocephala each have three stands. The
remainder of the species have two or one stands
per species.
Sri Lanka’s ex situ collections focus on field
collections for exotic species such as Eucalyptus
grandis, Eucalyptus microcorys, Khaya senegalensis
and Tectona grandis. Tectona grandis has the
largest number of field collections, including 200
accessions in five stands. Sri Lanka reports no ex
situ collections for native tree species.
Uzbekistan has four native species in ex situ
field collections (Amygdalus spp., Haloxylon
aphyllum, Juglans regia and Pistacia vera) and has
no other ex situ collections.
Europe
European countries report a range of ex situ
conservation activities; some countries have
extensive activities (including provenance trials,
seed orchards, clonal repositories, botanical
gardens and arboreta, and seed and pollen gene
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
banks) while others have none. Some European
countries (e.g. Bulgaria and Sweden) note that
ex situ conservation is of secondary importance
compared to in situ conservation because it is a
static approach and therefore does not provide
for adaptation to a changing environment.
Northern European countries identify a need
for long-term seed storage, as seed years are
scarce at northern latitudes and high altitudes.
In Norway, for example, the Nordic Genetic
Resource Centre (NordGen), a meeting place for
researchers, managers and practitioners from
Nordic countries working on forest genetics,
seeds, plants and regeneration methods, is
assessing the possibility of using the Svalbard
Global Seed Vault in Norway as a long-term seed
storage option for forest seed banking activities.
Cyprus is considering the establishment of a
national seed banks for forest species.
Ex situ conservation priorities identified by
European countries include the conservation of
rare and endangered species, populations that
are genetically unique, and species of ecological
and economic importance. Many European
countries report that ex situ conservation
activities have been negatively affected by the
fiscal and economic slump of recent years.
Species conserved. Significant diversity is seen
in the native species conserved ex situ in the 15
European countries reporting data. A number
of countries have large accessions of Pinus
sylvestris, to a lesser extent Picea abies, and many
hardwoods including Quercus spp. and Populus
spp. In seven countries, Pseudotsuga menziesii is
a common exotic species conserved ex situ.
Bulgaria has identified 36 native and exotic
tree species conserved ex situ primarily in field
collections and in a few seed bank accessions.
The largest ex situ field stands for native species
are for Quercus petraea, Q. frainetto and Fagus
sylvatica, each with at least 49 accessions, while
Populus spp. and Pinus sylvestris have the largest
number of field stands and accessions. For exotic
species, Pseudotsuga menziesii has the largest
number of accessions, with 55 accessions in one
field stand. Seed is stored ex situ for two species
(Picea abies and Pinus sylvestris).
Cyprus reports that the majority of ex situ
collections are for stored seed; seed is stored for
16 native species, mostly trees, and the number
of accessions ranges from 3 to 15. The largest
accession is of Astragalus macrocarpus subsp.
lefkarensis, a small shrub present in evergreen
mixed forests (IUCN, 2013).
In Denmark, ex situ conservation is in the
form of seed storage, with accessions of ten tree
species. Abies nordmanniana has the largest
number of accessions, followed by the native
species Pinus sylvestris and the exotic species
Pseudotsuga menziesii.
Estonia’s ex situ conservation activities are
also in the form of seed storage, with two native
species, Populus tremula and Populus tremula f.
gigas, and the exotic species Populus × wettsteinii
having multiple accessions.
Finland has ex situ field collections for eight
native tree species (Acer platanoides, Fraxinus
excelsior, Juniperus communis, Quercus robur,
Sorbus aucuparia, Tilia cordata, Ulmus glabra
and Ulmus laevis). Germany has 58 species
represented in ex situ field collections, with the
largest collections for the native species Taxus
baccata, Picea abies and Fagus sylvatica and for
the exotic species Pseudotsuga menziesii.
Hungary has large ex situ field collections for 23
species, mostly native. The largest collections are
for Populus nigra and Picea abies, with over 1 000
accessions each, then Castanea sativa, followed
by species such as Ulmus laevis, Ulmus minor and
Ulmus pumila, each with over 300 accessions.
Ireland has 11 species conserved ex situ,
primarily in field collections; only one species,
Pinus sylvestris, is native. The collections for this
species are extensive: 52 stands, 619 accessions
and three clone banks with a total of 562 clones,
as well as 100 in vitro and 75 seed accessions.
Most of the Netherlands’ ex situ collections
are in clone banks, with 59 native species
represented. The largest collections are for
Crataegus monogyna and Juniperus communis,
with 333 and 284 clones, respectively.
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Norway has approximately 19 species in ex situ
collections, of which eight are native. Most of the
collections are field stands or long-term trials.
Two native species are conserved in clone banks
(Picea abies and Pinus sylvestris). Norway has a
large number of field collections for Picea abies,
with 260 stands or trials and 6 000 accessions.
In Poland, 33 species are conserved ex situ, the
majority native. Seed storage is the main form
of ex situ conservation; the largest collection of
4 764 accessions is for Pinus sylvestris, followed by
836 accessions for Picea abies.
Spain has ex situ collections, primarily in
the form of clone banks, for approximately
27 species, of which most are native species.
The largest collection is for four native species
(Populus tremula, P. nigra, Ulmus minor and
U. glabra). Spain also has ex situ seed collections
for five native species (Arbutus canariensis, Pinus
pinaster, P. pinea, Pinus uncinata and Populus
alba).
Sweden’s reported ex situ conservation
collections are primarily field collections; species
include Alnus glutinosa, Betula pendula, Fagus
sylvatica and Quercus robur. Sweden also has
seed collections for a few species and a few clone
banks.
In Ukraine, ex situ collections comprise field
collections, which include a few clone banks for
native and exotic species. The largest collection
is for Pinus sylvestris, with 95 stands representing
1 148 accessions and 38 clone banks with 1 092
accessions. The next largest collection is for
Quercus robur, with 30 stands representing 539
accessions and 16 clone banks with 540 accessions.
The Russian Federation has extensive ex situ
tree seed collections. The Russian Forest Seed
Warehouse in Pishkino, Moscow Region, for
example, currently stores 10 tonnes of seed,
focusing on Larix spp., Picea spp. and Pinus spp.
Latin America and the Caribbean
Argentina, Brazil, Chile, Costa Rica, Ecuador,
Mexico and Peru have similar ex situ conservation
168
activities; all have multiple seed banks, botanical
gardens or arboreta and seed orchards, and most
have the capacity for germplasm storage in other
forms such as DNA, pollen and cryopreserved
tissue. Germplasm sources range from non-native
commercial species with economic impact to
native species with unique medicinal and other
uses. Gene banks have been set up as networks
within some countries and are therefore
controlled mainly at local level. All of the
countries have storage infrastructure – facilities,
technology and equipment.
Mexico has 54 forest gene banks with
temporary to medium-term storage; their
objective is to supply nurseries for government
reforestation programmes, but some are used
for conservation. Mexico is working with 74
tree species, mostly native and equally divided
between conifers and hardwoods. Conservation
activities in field collections and clone and seed
banks are conducted for 55 native and 12 exotic
species. Seed banks hold over 6 900 accessions
from 36 species; the number of accessions per
species ranges from 1 to 3 665. Mexico has 57
arboreta and botanical gardens which harbour
germplasm for scientific research; nutritional,
medicinal and ornamental uses; and conservation
of at-risk species.
Most countries with ex situ conservation
programmes have identified priorities to guide
their respective programmes. These include
conserving and improving FGR important to the
country, contributing to their sustainable use and
promoting the value of FGR conservation among
scientists and the general population. Ex situ
conservation of a species entails determining an
appropriate representative sample of the species
based on geographic or genetic variation, and
identifying the future uses of the material (e.g.
in breeding, planting, research and development,
and conservation programmes).
The countries in the region report similar
constraints, including a lack of permanent
financing for long-term projects. The knowledge
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
and in-depth research needed to characterize
the material being conserved are often lacking.
Thus a critical area of research that needs to be
addressed is the nature of recalcitrant seed and
protocols for handling seeds of species with this
storage behaviour. Countries also cited a need to
have clear strategies and policies for conservation.
Species conserved. Only Costa Rica, Ecuador and
Mexico report on the species in ex situ collections.
In Costa Rica, four native species and one of
uncertain origin are conserved ex situ in field
collections. The four species, of which three are
native (Dipteryx panamensis, Sacloglottis sp. and
Schlerolobium sp.) and one is of uncertain origin
(Himenolobium parahybum), are represented
in 26 accessions in field stands. Sacloglottis sp.
and Himenolobium parahybum have the most
accessions, 12 and 9 respectively. Swietenia
macrophylla, a native species, is the only
species present in clone banks, with 600 clones
established.
Ecuador has 114 species conserved ex situ, all
in field collections; 64 of these species are native,
49 are exotic and one is of uncertain origin.
The total number of accessions for all species
is approximately 166, in multiple field stands.
Multiple species are conserved in three genera:
Eucalyptus (17 species), Acacia (13 species) and
Erythrina (6 species).
Only one species, Swietenia macrophylla, is
conserved ex situ in all three reporting countries.
This species, a commercially important mahogany,
is native to these countries and is identified by
IUCN as vulnerable (IUCN 1998).
In Mexico, extensive effort is directed to the
genus Pinus (26 species and seven varieties); over
320 accessions of Pinus greggii var. greggii and
P. patula are held in 28 field stands, while just
over 1 000 accessions of P. greggii and P. patula
are represented in 16 clone banks. Considerable
effort has been expended in establishing field
stands of other species such as Calophyllum
brasiliensis,
Cedrela
odorata,
Cupressus
guadalupensis, Eucalyptus grandis, Eucalyptus
urophylla, Guaciacum coulteri and Platymiscum
lasiocarpum, with almost 3 500 accessions
represented in 39 stands. Of these species,
E. grandis and E. urophylla have 1 300 accessions
in five clone banks. The most represented species
in seed banks are Pinus patula and Toona ciliata,
with 700 and 3 665 accessions respectively.
Near East
The reporting countries from the Near East (Egypt,
Islamic Republic of Iran, Iraq, Jordan, Lebanon,
the Sudan and Yemen) cite similar priorities and
constraints in ex situ FGR conservation. All want to
establish, enhance or continue scientific research
and education on aspects of FGR conservation
such as propagation of trees and protocols for
storing seed. The countries report a clear need
to improve their infrastructure and technical
capacity with respect to provenance tests, seed
orchards, arboreta, gene banks and seed centres.
They also cite a need to promote partnerships with
forest-neighbouring communities to manage the
resources and improve livelihoods.
Lack of funding, trained personnel, research and
equipment are reported as constraints. Climate
change is expected to have severe impact on this
region given the current stage of desertification.
Habitat degradation and depletion from logging,
grazing and fuelwood harvesting highlights the
importance of protecting the forest through
establishment of reserves, botanical gardens and
arboreta. National policy to support and create a
strategy for FGR conservation and management is
urgently needed in most countries in the region.
Some countries (e.g. Iraq and Jordan) report
the lack of capacity to develop comprehensive
GIS surveys to identify areas and species that
require conservation or protective management
measures.
Wild relatives of fruit-tree crops are widespread
in the Near East but have been neglected in terms
of ex situ conservation, and little is known about
their distributions (Table 13.3).
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PART 4
TABLE 13.3
Wild relatives of fruit-tree crop species reported
by Jordan as present but understudied in terms
of ex situ conservation
Tree crop
Neglected wild relatives
Almond
Amygdalus spp.
(5 species and many interspecific hybrids)
Apple
Malus spp.
Cherry
Cerasuse microcarpa
Olive
Olea europea subsp. oleaster
Pear
Pyrus syriaca
Pistachio
Pistacia spp.
Species conserved. The Islamic Republic of Iran,
the only country in the region reporting data
on species in ex situ collections, lists 18 Populus
species conserved in clone banks with a total of
258 clones. Populus × euramericana of uncertain
origin and Populus nigra, a species native to the
Islamic Republic of Iran, have the most clones
established, 59 and 91 respectively.
North America
Canada conserves FGR ex situ in trials, plantations
and clone banks; 28 native and five exotic species
are established in over 510 field sites, and 14
native and one exotic species are represented in
37 clone banks (excluding provenance trials and
clone banks for breeding programmes). Five seed
banks – one national and four provincial – hold
15 000 accessions representing 75 species (all
native except one).
Conservation activities in the United States of
America are conducted for many species that are
native not to the continental United States but
rather to its tropical islands; 48 of the 57 listed
species are native to the state of Hawaii or the
territories of Puerto Rico and the United States
Virgin Islands. The United States has multiple ex
situ field and seed collections, predominately
for native species. Ex situ conservation activities
are conducted for 77 species of trees and shrubs
by over 80 arboreta and botanic gardens, and
national and regional seed banks store over
170
120 000 seedlots from over 250 species. Over 200
tree and shrub species are conserved ex situ in seed
collections of the United States Department of
Agriculture (USDA) Forest Service and Agricultural
Research Service. Conifers are the best conserved,
since they have large reforestation programmes.
The USDA Forest Service maintains family seedlots
for 44 reforestation species, totalling over 80 000
families. Breeding cooperatives in the United
States maintain large breeding population sizes
(in the hundreds) for each of their breeding
zones, and their hundreds of progeny trials
represent hundreds of field sites providing gene
conservation as a secondary objective.
Future priorities include conservation and
deployment strategies for mitigation of climate
change impacts; prioritization of species
identified at the federal and provincial levels and
those at risk from invasive alien species; a focus
on non-commercial coniferous and deciduous
species; gap analyses to optimize sampling;
and promotion of the sustainable use of forest
genetic resources.
Species conserved. In Canada, key conifer species
with 550 to 770 accessions in clone banks are
Picea glauca, P. glauca × engelmannii and Pinus
contorta var. latifolia. For five hardwood species
(Fraxinus pennsylvanica, Prunus virginiana var.
virginiana, Quercus macrocarpa, Sherpherdia
argentea and Symphoricarpus occidentalis), the
number of accessions in clone banks ranges from
1 900 to 6 600. In seed banks, 1 000 accessions
each are held for Picea glauca, P. glauca ×
engelmannii and Pinus contorta var. latifolia.
The three most predominant species in ex situ
conservation in the United States of America
are Pinus lambertiana with over 26 000 family
collections, Pseudotsuga menziesii with over
20 000 family collections, and Pinus ponderosa
with over 13 000 family collections.
Oceania
Ex situ conservation activities have been
conducted for about 60 species in the countries
of Oceania that submitted reports: Australia,
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Cook Islands, Fiji, France (represented by three
territories – French Polynesia, New Caledonia and
Wallis and Futuna), Papua New Guinea, Solomon
Islands and Vanuatu. The conservation activities
include species and provenance trials, clonal seed
orchards, seed production areas, clone banks and
cryogenic storage. In 2011, the Secretariat of the
Pacific Community developed the Pacific Islands
Tree Seed Centre to help research, conserve
and disseminate seeds of socio-economically
important tree species for its 22 member countries
and territories.
Australia has more than 1 000 species conserved
ex situ in seed banks and field plantings (clonal
archives, seed orchards, arboreta). The Australian
Seed Bank Partnership (www.seedpartnership.
org.au), which developed from the Millennium
Seed Bank Partnership, has a mission to safeguard
Australia’s plant populations and communities
through a national network of conservation seed
banks. This partnership unites the expertise of
14 institutions, including universities, herbaria,
botanic gardens, NGOs and environmental
agencies. The Threatened Flora Seed Centre
of the Western Australian Department of
Environment and Conservation (one of the
members of the partnership), established to
safeguard a geographically diverse range of seeds
from threatened plant species, has successfully
stored seeds from three-quarters of Western
Australia’s threatened plant species – many of
them trees and other woody species. The centre
has also reintroduced more than 50 threatened
species back into the wild. CSIRO’s Australian Tree
Seed Centre maintains a national ex situ seed
collection of more than 900 tree species, while the
Southern Tree Breeding Association contributes
significantly to ex situ conservation through
provenance and progeny trials for multiple tree
species.
New Caledonia (France) has developed a small
number of clonal archives and seed orchards of
unique endemic species of Araucariaceae.
Papua New Guinea has a national tree seed
centre that stores seed for research, reforestation
and export.
Solomon Islands reports that 30 000 plant
specimens transferred from its National
Herbarium to Fiji during civil unrest in 1999–2000
have not yet been returned.
Constraints for ex situ conservation in the
region include limitations in, or lack of, research,
national policies and strategies, funding,
facilities, public education and training for staff,
as well as land tenure issues. Future priorities
identified in the country reports include staff
training, involvement and engagement of rural
communities; funding commitments; external
collaboration with funding agencies; assessment
of the state of endangered species; development
and upgrading of facilities; and expansion of
collections and field trials.
Species conserved. Only Australia and Papua
New Guinea report data on species under ex
situ conservation. In Australia, well represented
genera and species include eucalypts in the
genera Angophora, Corymbia and Eucalyptus
(900 species in seed banks and arboreta), Acacia
auriculiformis (780 accessions in seed banks),
Araucaria cunninghamii (800 clones and 400
families planted in field), Khaya senegalensis (150
clones and 80 provenances in seed banks) and
Pinus radiata (916 clones and 772 seed accessions).
In Papua New Guinea, 200 field stands
containing 107 accessions have been established
for five native species (Acacia crassicarpa, Acacia
mangium, Araucaria cunninghamii, Araucaria
hunsteinii and Eucalyptus deglupta) and one
exotic species (Tectona grandis). Seven clone
banks contain 114 clones of these species.
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STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Chapter 14
Genetic improvement and
breeding programmes
Tree breeding programmes have the potential to
improve the production of planted forests and
trees in a sustainable way and are necessary to
meet growing global demand for forest products
and services. More extensive development
of forest industries based on planted forests
using diverse, improved tree germplasm has
the potential to meet a large proportion of
the world’s wood requirements and relieve
pressures on natural forests. For example, timber
plantations growing at 10 to 20 m3 per hectare
per year and covering an area equivalent to
2.5 to 5 percent of the world’s forests would
be capable of producing 2 billion cubic metres,
meeting much of the expected global demand
for roundwood projected for 2030 (Carle and
Holmgren, 2008).
At the national level, ongoing tree breeding is
needed to support planted forest development to
help meet the export and local demands for forest
products. This is especially the case in countries
where harvesting in natural forests is occurring
at unsustainable rates or is highly restricted or
banned, and in those countries where natural
production forests are limited in area and/or low
yielding. Tree breeding is also essential to address
new challenges associated with climate change
and emerging pests and diseases.
Trees have been the subject of informal
selection, breeding and movement for centuries
if not millennia, with traditional domestication
efforts mainly focused on edible fruit- and nuttrees. Informally improved trees have contributed
in a significant way to livelihoods in many
countries, although the benefit attributable to
improvement is difficult to quantify. The past two
to three decades have seen growing interest in
domestication and less intensive improvement
programmes for a wider range of multipurpose
and food-tree species, often of native origin. For
example, Brazil describes a successful programme
in which wild races of Bactris gasipaes (peach
palm) were bred and distributed to over 80
percent of the Latin American farms that grow this
high-value product. ICRAF’s tree domestication
programmes with national partners in subSaharan Africa, Southeast Asia and Latin America
focus on improvement of agroforestry tree
products (AFTPs) such as charcoal, fodder, fruits
and nuts, oils and medicines.
For much of the second half of the twentieth
century, tree breeding and genetic improvement
focused on species for commercial production
of timber and pulpwood, and on improving
a relatively small number of traits that would
maximize economic gains (including growth
rate, volume, form, processing and product
quality). The focus on commercial species was
related to the high costs involved in undertaking
a comprehensive breeding rogramme, the
specialist expertise needed, and the typically
long periods before benefits are realized (of
the order of several decades in cooler climates).
The productivity gains and increased commercial
returns of plantations established with improved
germplasm, however, more than compensated
for the high costs involved in tree breeding
programmes.
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In recent decades government agencies and
the private sector have subjected a wider range
of tree species to domestication and formal
breeding programmes to produce myriad goods
including timber, pulp, fuelwood, fruit, nuts,
oils, traditional medicines, dyes, resins, thatch,
and other NWFPs, and to provide forest service
functions. In addition, tree breeding efforts are
increasingly focused on adaptability-related traits,
for adaptation to predicted new climate regimes;
for resistance to drought, fire, pests and diseases
(see FAO, 2013c); and for use in forest restoration
programmes. These breeding programmes are
primarily initiated by public agencies. The United
States of America reports several programmes
of tree breeding for conservation and forest
restoration; for example, to reintroduce species
eliminated by disease, breeding programmes
seek to confer disease resistance from selected
wild individuals (see section on North American
tree improvement activities below). In breeding
for resistance, valuable knowledge is gained
about the behaviour of introduced diseases and
insect pests and about genetic options for control
(Schlarbaum et al., 1997; Loo, 2009). Within
identified genecological zones, inclusion of the
broadest available range of genetic variability
in direct-seeded or planting stock maximizes the
opportunity for a species to reoccupy its former
range, perform vital ecological services, and
adapt for the future.
The change in the scope and role of tree
selection and breeding programmes is a response
to several factors, including:
• the scale and unpredictability of
environmental change (including climate
change, especially extreme climate events
and interactions with pests and diseases);
• new demands and requirements for
trees for food and nutritional security,
environmental restoration and carbon
sequestration to mitigate climate change.
It is increasingly recognized that selected
improved tree germplasm needs to be generated
and deployed with multiple objectives in
mind, including human food, biofuels and
174
environmental
and
ecological
purposes.
Adaptation characteristics that will be more
sought after in some breeding programmes
include improved drought resistance, resistance to
and recovery from fire, and an ability to withstand
hurricane-force winds at all stages during the
development and life of a planted forest. It is
further anticipated that the introduction of more
diverse (in both inter- and intraspecific terms)
and a priori better-adapted genetic material
will increasingly be required in both natural and
planted forests, because the rapid rate of climate
change may exceed the ability of indigenous tree
species and populations to respond to change
through natural selection and/or migration.
Sweden notes that the success of ongoing
and future breeding activities will in large
part depend on the natural variability of FGR
contained in wild populations and in existing
breeding programmes and plantations; this
underscores the importance of FGR conservation.
Breeding programmes can also contribute to ex
situ conservation; the United States of America
remarks that “breeding programmes, by default,
have ex situ conservation plantings in their seed
orchards [and] progeny tests in addition to any
seed stores”. However, breeding activities by
themselves cannot be regarded as secure form of
long-term ex situ conservation, since the facilities
may be abandoned once the testing or breeding
programme has been completed.
Almost ten years ago FAO reported that “The
scale of forest genetics and tree improvement
in the tropics is entirely inadequate, both in
geographical distribution and species coverage,
and bears no relation to its potential value and
importance” (FAO, 2005b). This situation has not
substantially changed, although native species
are being gradually brought into improvement
programmes in all regions.
Improvement approaches
Successful and efficient tree breeding requires
genetic characterization, whether through the use
of growth and morphological characterizations
(principally through field trials) or through the use
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
of molecular markers and DNA characterization,
as discussed in Chapter 11. The improvement
process, described in Chapter 7, typically involves
seed collection of a large number of individuals
from native stands. Collection may be range wide
for exotic tree species or more geographically
focused to match climate and/or soil or if
superior sources have already been identified.
Seed collection is followed by provenance and
progeny trials, from which phenotypic selections
of plus trees are made and/or breeding values
determined, followed by breeding, crossing and/
or selection programmes based on different
strategies depending on species, breeding
objectives and resources available.
In tropical regions, Eucalyptus spp., Tectona
grandis and several other species have been
subjected to intensive selection and breeding and
are now primarily clonally propagated. Eucalyptus
breeding programmes in the tropics often focus
on producing high-value hybrid clones either
between species or between highly selected lines.
The commercially important temperate eucalypt
species are usually managed as seedling-based,
pure species breeding populations, as clonal
forestry is difficult in most temperate eucalypts.
Most teak field trials are clonal because of the low
seed production and the difficulty of conducting
controlled crosses.
Increasingly sophisticated approaches and
technologies are being applied to tree breeding
to generate faster rates of gain, for example
more statistically powerful trial and mating
designs; combined index-based selection taking
into account the economic weight, heritabilities
and covariances of different traits; prediction of
breeding values based on best linear unbiased
prediction; improved management of breeding
populations and maintenance of sublines to
reduce risks of inbreeding; and promotion of
earlier lowering and seed production. The
complementary use of molecular markers and
quantitative data is an important emerging trend
(see Chapter 8).
Hybrid breeding, involving interspeciic hybrids
and wide provenance crosses, is used in many
countries to produce trees with superior productive
capabilities (through heterosis) and also to
introduce genes for disease resistance. Examples
include eucalypt hybrids, Larix and Populus hybrids,
Pinus hybrids and increasingly hybrids for diverse
tree genera such as Acacia, Casuarina, Fraxinus,
Liriodendron, Prosopis and Santalum.
As noted in Chapter 8, at the global level,
relatively few attempts have been made to
develop genetically modiied or transgenic trees,
mainly because of community concerns and
associated legal restrictions and impediments
to their development and use in Europe, the
United States of America and elsewhere. In China
transgenic poplars have been produced through
incorporation of genes for insect resistance into
hybrid poplars.
Some 15 years ago the FAO Panel of Experts
on Forest Gene Resources (FAO, 1999b, 2001a)
noted with concern the widening gap between
science and practice, stressing that successful
application of scientiic knowledge was at risk if
the science was too advanced to be understood
and implemented at the operational level. This is
increasingly the case today, with entire genomes
now being sequenced for trees. Many developing
countries lack skilled tree breeders who can
understand and use the information generated
by forest geneticists and ensure its application
in practical, large-scale research, breeding and
planting programmes.
Administration and coordination
of breeding and improvement
programmes
Tree improvement is undertaken by different
types of organizations in different countries. In
Finland and Norway – countries with a major/and
or diversiied forest sector and market economy
– national tree breeding programmes are
funded by the government as a public service. In
Germany, the government-funded forest research
units of the states (Länder) are the main actors in
forest tree breeding. In other countries breeding
is undertaken by private forestry interests and
independent academic and research institutions.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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The benefits of tree improvement programmes
accrue over very long periods; thus to realize
these benefits, national and local security is
vital, as is a favourable investment environment
and stable global and national economies with
assured demand for forest products. National
governments have essential roles in these
areas. Where tree improvement programmes
are publicly funded, broad political support is
necessary to ensure that they reach fruition
and are either maintained by the government
or adopted by the private sector. For a private
company to engage in improvement work, it
must have a sufficient planting programme to
justify the required long-term investment. Private
companies and others need to be able to protect
the intellectual property resulting from their
breeding work, through patents and plant variety
rights. As reported by Germany, however, patents
do not play a significant role for FGR. Legally
binding and firm agreements for benefit sharing
are necessary (see section on benefit sharing in
Chapter 15).
Coordination and documentation of tree
breeding efforts are important at the national
level. Brazil, for example, reports that the Instituto
de Pesquisas e Estudos Florestais (Forestry Science
and Research Institute, IPEF), associated with the
University of São Paulo, “is leading an effort to
‘rescue’ most of the information scattered all
over the country regarding genetic improvement
programmes for Eucalyptus, pine and other
exotic tree species” – programmes that have been
carried out since the 1970s.
Collaborative tree breeding programmes
are most well developed in North America. The
United States of America, for example, has 43
cooperative breeding programmes working with
31 species. Camcore, a tree breeding organization
based in the United States, also operates
internationally. Collaboration is also strong for
commercial species in Australia and New Zealand.
With the decline of public funding, where private
companies once participated in coordinated
national programmes, they may now be likely to
participate in breeding cooperatives.
176
In most countries, full realization of the
benefits of tree improvement will require
national information systems and better
coordination among all actors – among and
within government agencies and departments
(especially departments of forestry, agriculture
and environment), research institutes and
universities and the private sector. A national FGR
strategy has a paramount role in coordinating
these actors; furthermore, national FGR strategies
need to be coordinated with strategies in forestry,
agriculture and development.
It is evident from several country reports that
governments tend to allocate more resources to
breeding of conventional agricultural crops than
to breeding of forest trees. With few exceptions,
tree breeding is administratively separate from
agricultural breeding. However, forest foods and
tree crops are extremely important for many
people around the world, often providing vital
nutrition and sustenance when other crops fail
because of drought, other environmental stress or
sociopolitical disruption. A number of countries
in Central Asia and the Near East harbour
progenitors or wild relatives of important foodtree species, and these are vital FGR for food
security. Closer cooperation between agriculture
and forestry might thus provide opportunities for
breeding improved trees for a wide range of uses,
particularly on farms and in rural communities,
to alleviate rural poverty and hunger and to
increase food security. There is considerable scope
for greater collaboration among individual tree
farmers practising improvement for edible fruit
and nut bearing trees, through better networking
and sharing of germplasm, information,
technologies and other resources.
A major avenue for international collaboration
in tree improvement is through sharing of genetic
resources, especially for those species whose
distributions cut across national boundaries.
In certain situations it may also be possible to
contract out tree improvement activities to
other countries (breeding cooperatives and/or
companies), as long as genotype × environment
interactions are demonstrated to be limited. A
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
country must have a high level of confidence
to entrust the development of its unique
genetic resources to another country, because
of perceived or real risks of losing control of
the process and not receiving a fair share of
the beneits lowing from improvement. This
underscores the vital role of agreements that
are irm, enforceable and fair between parties
engaging in such arrangements (see section on
beneit sharing in Chapter 15).
Prioritizing uses, traits and species
for improvement
The selection of traits and species for improvement
should relect the demand for forest products,
the needs of their users and the country’s rural
communities, and the goals set out in the
national development and forestry strategies
and of course the national FGR strategy. Where
countries lack these documents or the objectives
are not explicit, these decisions may be made
within government departments or research and
tree improvement institutes. Changes in demand
for forest products must also be considered;
China notes the importance of responding to
market signals. The extremely long time frame
of breeding programmes can be problematic,
however, when it comes to responding to changes
in demand for forest products and services. To
respond to the increasing demand for wood as
a sustainable energy source to mitigate climate
change, for example, Germany has developed
intensive improvement programmes to breed
trees for energy production, with a particular
focus on rapidly growing species harvested on
short rotation.
With shrinking public funding for breeding
programmes, private companies are increasingly
pursuing their own research priorities; the
private sector tends to be guided by potential
for commercial gain and is not bound by tests of
public beneit in its selection of traits and species
for improvement.
Many countries import improved genetic
materials for evaluation and further breeding,
which may involve adapting them to local
conditions. At least 25 North American tree
species have been introduced, tested and
improved in other countries for commercial
use (Rogers and Ledig, 1996) (see example in
Box 14.1). Imported materials are usually exotic
species but may include species indigenous to the
importing country (particularly if the species is
widely distributed).
A relatively small number of commercially
important plantation species, e.g. acacias,
eucalypts, pines, poplars and teak, are used
widely around the globe and are the focus of
many breeding programmes internationally
(Figure 14.1).
Box 14.1
Pinus radiata – a species improved outside its native range
The North American species Pinus radiata occurs
only as five small populations in its natural range. It
has not been planted commercially in North America
but has become a highly important timber species in
other countries (Cope, 1993). In its native range it is
used for ornamental purposes, erosion control and
fuelwood, but it is not considered to have commercial
value for timber because of its typical poor crown
form.
Pinus radiata was first introduced into Australia
for ornamental plantings around 1857 (Wu et al.,
2007), and its rapid growth led to its use in planted
forests beginning in the 1920s. Through selection and
breeding programmes in countries where the species
has been introduced, its productivity and stem and
crown form have been significantly improved. Indeed
P. radiata has become one of the most widely planted
pine species for timber.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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FIGURE 14.1
Most common species in tree improvement and conservation programmes worldwide
Populus alba
Eucalyptus camaldulensis
Under tree Improvement
Under tree Improvement
Under in situ conservation
Under in situ conservation
Under in situ conservation
Environmental
parameters
changing
in
response to climate change will require
planting stock adapted to new conditions.
Several countries observe that their breeding
programmes will increasingly need to focus on
survivability, drought and fire resistance, and
resistance to pests and diseases that may become
more prevalent under climate change. Breeding
for adaptation to climate change is increasingly
considered a high priority.
Many developing countries have not yet
properly explored their indigenous tree lora
for domestication and improvement; the United
Republic of Tanzania, for example, reports that
indigenous species have generally been neglected
in favour of tried and tested exotic species (as
178
Tectona grandis
Under tree Improvement
cited in Chapters 10 and 12). Likewise, Seychelles
has an extremely rich endemic tree lora, with
many excellent timber species that have not
been investigated for their forestry potential;
the country has focused instead on importing
exotic species. Many developing countries
contain large areas of natural forest providing
extensive reservoirs of tree genetic diversity
with potential for development for human use.
National forest agencies are increasingly keen
to explore such diversity through domestication
and improvement of local or indigenous tree
species. International donors and research
partners are also increasingly supporting this
approach, desiring to promote indigenous
species that are already adapted and well
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
known by local communities and to contribute
to biodiversity conservation objectives while
avoiding risks associated with possible invasive
behaviour of exotic species.
Domestication and improvement programmes
can deliver long-term economic and other
benefits and when appropriately designed
can help to ensure conservation of genetic
diversity. However, smaller countries often lack
the resources or capacity to conduct their own
domestication and improvement programmes
even for important local tree species. Some
countries have limited or no local forest
industries and rely instead on imports or minor
local harvesting to meet their needs. Cyprus,
for example, has no forest industry because of
low growth rates; its forests are conserved for
environmental protection. Countries lacking
improvement programmes include almost all
of the 52 small island developing States (SIDS)
in the Caribbean, Indian and Pacific Oceans –
although some SIDS in Oceania have or have had
collaborative tree improvement or domestication
programmes, e.g. with CIRAD-Forêt, CSIRO/ACIAR
and the Australian SPRIG project. Seychelles,
with its small population and limited resources
in terms of expertise, infrastructure and financial
support, has no breeding programme for trees
but rather conducts “adaptive screening” of
imported materials (e.g. mango and avocado).
Several other countries without domestication or
breeding programmes import improved material,
generally high-performing, high-value exotic
industrial forestry species for which germplasm is
readily available.
The state of tree improvement
and species priorities by region
More than 700 species are the subject of
improvement efforts around the world (Figure
14.2). Improvement programmes described in the
country reports are presented here.
FIGURE 14.2
Number of species and subspecies in improvement programmes, by region
Africa
Asia
Europe
Latin America and
the Caribbean
Near East
North America
Oceania
Total
0
200
400
600
800
Note: Numbers for Europe may be high owing to the inclusion of a number of territories in tropical regions.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Africa
Tree improvement has a long history in some
African countries, including Kenya, Morocco,
South Africa and Zimbabwe. One of the world’s
earliest breeding programmes for a broadleaved
tree was for Acacia mearnsii in South Africa; it
was initiated more than 60 years ago (Dunlop et
al., 2003).
The objectives of African programmes were
initially to improve tannin yields, growth and
gummosis resistance, then to improve wood
properties and most recently to breed for sterility
in future plantations to reduce invasiveness risk
(with a 2015 target). A number of the earliest tree
improvement activities were undertaken with
the assistance of the United Kingdom (official
development assistance and the Oxford Forestry
Institute) and France (Centre technique forestier
tropical [CTFT]); these programmes mainly
focused on exotic timber species to be developed
for industrial forestry plantations. Eucalyptus and
Pinus species dominated, and still do today.
Tree improvement typically requires sustained
support and often considerable capital injections;
consequently some improvement programmes
initiated in Africa many decades ago with European
support are no longer operational, while smaller
countries such as Burundi and Seychelles have no
tree improvement programmes. Zimbabwe’s tree
improvement programmes for exotic industrial
species, which had delivered impressive gains
(e.g. cumulative volume gain of up to 45 percent
in third-generation selections of Pinus patula over
the original wild material), are being reactivated
owing to improved economic circumstances.
Most tree improvement programmes in Africa
use traditional and modern breeding approaches
involving provenance testing, plus-tree selection,
family and/or progeny trials, open pollination
and/or hand cross-pollination, recurrent selection
and multiple breeding populations and breeding
indexes for multiple-trait improvement.
Over the past two decades many sub-Saharan
African countries have increased their focus on
improving many and diverse local multipurpose
180
species, often traditional food trees. These
programmes often involve collaboration between
national agencies and ICRAF and may involve
local communities in a participatory process of
selection and breeding.
Some species under improvement in more than
one African country are as follows.
• Eleven African countries have improvement
programmes for Eucalyptus species, notably
E. camaldulensis and E. grandis in eight and
seven countries respectively, as well as
E. globulus, E. tereticornis and E. urophylla.
• Pinus improvement is reported in eight
countries; the main species are P. caribaea,
P. elliottii, P. oocarpa, P. brutia and
P. halepensis.
• Other exotic timber-producing species
undergoing improvement in several African
countries include Tectona grandis, Cupressus
spp., Acacia auriculiformis, Azadirachta
indica and Grevillea robusta.
• Native timber trees under improvement in
more than one country include Khaya spp.,
Milicia excelsa and Terminalia superba.
• Multipurpose and NWFP trees (including
those providing food and medicinal
products) under improvement in more
than one country include Acacia senegal,
Adansonia digitata, Detarium spp., Irvingia
gabonensis, Parkia biglobosa, Sclerocarya
birrea, Tamarindus indica, Vitellaria
paradoxa and Ziziphus mauritiana.
Asia
The Asia region has a great diversity of tree
improvement programmes and species for
improvement which vary depending on the
subregion, level of development and other
factors. China, India, Japan and Thailand have
well developed and comprehensive improvement
programmes.
China has a vast tree improvement programme
including more than 100 mainly native species,
principally being improved for wood production.
An example is the improvement of the native
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
conifer Cunninghamia lanceolata which has
involved trials of more than 200 provenances in
nine regions and resulted in an average gain in
wood production of 16 percent. China reports:
“Significant gains have been achieved due
to the use of genetically improved plant
materials in plantations, achieving an
average growth gain of 10 to 30 percent
for timber trees and an average yield gain
of 15 to 68 percent for fruit-trees. … The
average growth gain of improved timber
trees was more than 10 percent, and the
average yield gain of improved economic
trees was more than 15 percent.”
India has ongoing programmes to improve
more than 140 species, mainly native. However,
most plantations are established with Acacia
spp., Casuarina spp., Cedrus deodara, Eucalyptus
spp., Gmelina arborea, Grevillea robusta, Pinus
roxburghii, Pinus wallichiana, Shorea robusta and
Tectona grandis. While the focus for most species
in improvement programmes is wood production,
a substantial number of species are bred for fuel
production, multiple uses and NWFPs.
Japan’s tree breeding programmes commenced
more than 60 ago and aim to increase
productivity and sawn timber quality for the
major planted native conifers: Abies sachalinensis,
Chamaecyparis obtusa, Cryptomeria japonica,
Larix kaempferi and Picea glehni. The largest
programme is for C. japonica, with breeding
conducted in four regions and based on 500 to
1 000 selected individuals per region.
The Republic of Korea’s tree improvement
activities have mainly focused on plus-tree
selection (2 724 trees of 28 species) and seedorchard development (59 species, including 16
gymnosperms covering 734 ha). The main focus
has been on native species, particularly the
major planted species, Pinus densiflora (now into
the second generation of improvement), Pinus
koraiensis and P. thunbergia.
Thailand also has a long and successful
history of tree breeding developed through
collaboration with Denmark (the DANIDA Forest
Seed Centre) for improvement of teak and pines.
Over the past three decades Thailand has also
collaborated with Australia (the CSIRO Australian
Tree Seed Centre) for improvement of Acacia,
Casuarina, Chukrasia and Eucalyptus species.
Major improvement effort has been on the two
main planted timber trees, teak (Tectona grandis)
and Eucalyptus camaldulensis. Improvement of
the latter species, and its hybrids, has involved
improving form and growth rates, then disease
resistance and most recently pulpwood traits,
increasingly with involvement of the private
sector and clonal registration.
Eucalyptus is an important genus for
improvement in the region, with programmes
reported in eight countries involving many
species including E. camaldulensis, E. globulus,
E. pellita, E. tereticornis and E. urophylla.
Pinus species are also widely planted in Asia,
with more than ten different species undergoing
improvement in seven Asian countries.
Tectona grandis improvement is being
undertaken in seven countries in the region,
including three in which it is native (India,
Myanmar and Thailand).
Other species undergoing improvement in
several Asian countries include:
• several high-value timber species in
the genera Pterocarpus and Dalbergia,
undergoing improvement in five and three
countries, respectively;
• Gmelina arborea and Acacia species
(especially A. auriculiformis, A. mangium
and their hybrid), important industrial
plantation timber species that are in
improvement programmes in four countries
each;
• mainly Asian timber and NWFP species:
Albizia spp., Azadirachta indica, Casuarina
spp., Magnolia spp., Phyllanthus emblica
and Santalum album.
Populus species and hybrids are being improved
in China, India and Kazakhstan, both for wood
and for bioenergy. Populus euphratica has
notable climatic and ecological amplitude which
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
is increasingly being exploited for restoration
programmes. In China and Uzbekistan certain
salt-tolerant Haloxylon species are also being
improved for restoration plantings.
Improvement programmes in Central Asia
are often focused on fruit- and nut-trees and
their wild relatives in the genera Juglans, Malus,
Pistacia, Prunus and Pyrus.
Europe
With the exception of Switzerland, all reporting
European countries have tree improvement
programmes, although in some cases they are
limited or only recently initiated. Northern
European countries have the most advanced
breeding programmes; the most comprehensive
programmes are detailed by Finland, France,
Germany and Sweden. These activities are mainly
for timber and pulpwood species. The main
species undergoing improvement in Northern
and Central Europe are Pinus sylvestris and Picea
abies; 11 or more countries report programmes
for each of these conifers. According to country
reports, at least 25 other native conifer and
broadleaf species are subject to improvement,
but some of the species are in the beginning
stages of selection and breeding. Most of
the efforts are on improving productivity,
wood quality and environmental adaptation,
particularly in view of climate change. Only a
few European countries mention breeding for
pest resistance; the Netherlands, for example,
notes that in addition to selection for adaptation
to site conditions, considering survival and bud
burst, trees are also selected for resistance to
pests and diseases.
Several European countries have breeding
programmes for Christmas trees and other NWFPs.
In addition to improving trees for timber and pulp,
almost half the countries in the region report
improvement programmes for tree species used
for energy. Genotypes of several species are also
being tested in the Netherlands for non-forestry
purposes such as their performance as roadside
trees. The existence of breeding programmes
focusing on such a diverse range of species and
182
traits implies a substantial body of knowledge
about genetic variation and heritability.
In Finland, breeding is managed by the
Finnish Forest Research Institute. Activities are
carried out in six regions and focus on improving
productivity, wood quality and improved climatic
adaptation in the two main native timber species,
Pinus sylvestris and Picea abies. Together these
two species comprise over 90 percent of Finland’s
annual reforestation area.
The Russian Federation adopted State
programmes
and
guidelines
on
forest
improvement through breeding from the 1960s
to the 1980s. The Central Research Institute of
Forest Genetics and Breeding was established in
1971 and has subsequently directed the country’s
breeding and genetic studies and coordinated
the work carried out by research institutions in
different regions of the country. Provenance trials
(mean age of 40 years) and many years of research
have contributed to a broad body of knowledge.
Pine trees, in particular Pinus sibirica, have been
selected and bred for resin productivity and nut
yield.
Sweden’s comprehensive breeding programmes
involve
intensive
plus-tree
selection
in
collaboration with forest owners, large-scale
controlled crossings and evaluations, with 7 to
24 breeding populations for each major species.
The large impact of the tree improvement work
is indicated by the projected annual increase of
10 million cubic metres of wood from planted
improved germplasm of Picea abies and Pinus
sylvestris.
Other coniferous species included in breeding
programmes in at least two or three countries
in Europe include Larix spp. (five countries), the
North American species Pseudotsuga menziesii
(six countries), Abies spp., Cedrus spp. and Taxus
baccata. Species in many broadleaf genera are
also being improved in several countries, mainly
for increased timber production but also for
bioenergy and environmental services. Betula
spp. (mainly B. pendula), Fagus spp. (mainly
F. sylvatica), Populus spp. (especially interspecific
hybrids) and Quercus spp. (mainly Q. petraea),
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
each in six countries; Prunus avium in five
countries; and Alnus glutinosa, Fraxinus excelsior,
Juglans regia and Ulmus spp., each in four
countries.
A different suite of species is under improvement
in Mediterranean countries, including more
drought-tolerant trees in the genus Pinus
(P. brutia, P. halepensis and P. pinaster) and
Quercus suber.
Most of the tree species included in breeding
programmes in Europe are native to Europe and
to the countries where breeding efforts are under
way; however, about one-third of the tree species
under improvement are exotic in the reporting
countries. In some cases these species originated
from North America or other parts of Europe.
Knowledge gained about the performance of
these species outside their native range may also
be useful within their native range, particularly
in matching seed sources to novel environmental
conditions.
France’s tree improvement programmes
include long-distance interspecific hybrids in
three genera, Larix (hybrids between European
and Japanese species), Populus (hybrids between
European and North American species), and
Juglans (hybrids between European and North
American species); these activities highlight the
continuing importance of germplasm exchange
for tree improvement programmes in the
Northern Hemisphere.
Apart from these activities in the region, it
should also be noted that the contributions of the
DANIDA Forest Seed Centre in Denmark and of
donors, tree breeders and geneticists in European
countries (especially Finland, France, Germany
and the United Kingdom) have greatly assisted
development of tree improvement programmes
throughout the developing tropics over the past
40 years.
Latin America and the Caribbean
Well developed tree improvement programmes
are found throughout Latin America, especially
for species in the industrial plantation genera
Eucalyptus and Pinus. Several smaller countries do
not have improvement programmes and may rely
on importation of improved genetic materials,
for example for species in the genera Eucalyptus,
Pinus and Tectona.
Brazil’s
improvement
programme
for
Eucalyptus species, based on considerable
species and provenance selection work, family
evaluation and hybrid development, is especially
noteworthy. This work has resulted in some of
the fastest-growing plantation trees in the world:
Individual clones of E. urophylla × E. grandis grow
more than 100 m3 per hectare per year.
Genetic improvement programmes underpin
Brazil’s 5 million hectare eucalypt plantation
industry, and the commercial benefits of this
improved material are immense:
“Brazil is … one of the main pulp and
paper producers in the world and a
sector reference in terms of sustainable
pulpwood production, which is 100
percent harvested from planted forests,
mainly eucalyptus and pine. The
productivity of these planted forests is the
highest among all pulp producers in the
market, with an annual average growth of
41 m3 per hectare per year for eucalyptus
and 35 m3 per hectare per year for pine
plantations. This is the result of 30 years
of a successful research development and
transfer process in a country where the
climate is very favourable and private
research institutes work… integrated
with researchers in universities to
generate genetically improved material
and advanced silvicultural treatments.”
Several other countries in the region also
have Eucalyptus improvement programmes,
not only for E. grandis and E. urophylla but also
for E. camaldulensis, E. globulus, E. nitens,
E. tereticornis and hybrid combinations as well
as for several other eucalypt species such as
Corymbia maculata in Peru and Eucalyptus dunnii
in Argentina.
Several Central and South American countries
have major improvement programmes for Pinus
species. The largest programme is for Pinus
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
radiata in Chile, initiated over 40 years ago and
now into its third or fourth generation, involving
more than 1 300 trials (including other Pinus
species, notably P. ponderosa). Argentina’s pine
breeding efforts are focused on the two main
planted species, P. elliottii and P. taeda, and their
hybrids. Germplasm collected in Guatemala from
several Pinus spp. (P. caribaea var. hondurensis,
P. maximinoi, P. oocarpa and P. tecunumanii)
has been subjected to breeding by Camcore
outside Guatemala; this material and associated
information are now being used for pine breeding
in Guatemala.
For species in other genera, the main focus
in tropical Latin America has been on fastgrowing exotic timber trees such as Acacia
mangium, Gmelina arborea, Grevillea robusta,
Hevea brasiliensis (in Mexico), Tectona grandis
and Terminalia spp. Some countries (Argentina,
Chile) also have improvement programmes for
Populus species and hybrids; Salix and Populus
species are tested for pulp and paper. Some
Latin American countries employ only exotic tree
species for commercial production. In such cases,
breeding programmes are designed to increase
the adaptedness of selected genotypes to local
environmental conditions and improve their
productivity.
A more recent trend has been the development
of improvement programmes for diverse native
species. A host of tropical American timber species
now feature in improvement programmes in Costa
Rica, Ecuador, Guatemala and Peru. These include:
Alchorneoides hieronyma, Alnus acuminata,
Cabralea canjerana, Calycophyllum spruceanum,
Cedrelinga cateniformis, Cordia alliodora,
Dipteryx panamensis, Guazuma crinita, Jacaranda
copaia, Ochroma pyramidale, Parkia multijuga,
Roseodendron donnell-smithii, Schyzolobium
spp., Swietenia macrophylla, Terminalia spp.
(including T. amazonica), Virola spp. and Vochysia
spp. (including V. guatemalensis). Mexico’s
tree improvement programmes (which are
substantial but mainly in the early stages) are
based on the country’s high forest biodiversity
and focus mainly on native timber species:
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Cedrela odorata, Cupressus lusitanica, Jatropha
platyphylla, six Pinus species (P. douglasiana,
P. greggii, P. leiophylla, P. oocarpa, P. patula and
P. pseudostrobus), Swietenia macrophylla and
Taxus globosa. In Chile four Nothofagus species
and Laurelia sempervirens are undergoing
improvement, while in Argentina the focus for
improvement of native timber species is again
Nothofagus spp. (N. nervosa and N. obliqua) as
well as Prosopis spp. (P. chilensis and P. flexuosa),
the latter not only for timber but also for recovery
of degraded lands.
In Brazil and Peru one focus has been on
indigenous fruit-trees – especially in the families
Myrtaceae (e.g. Acca spp., Campomanesia spp.,
Eugenia spp., Myrcianthes spp., Myrciaria dubia
and Psidium spp.) and Arecaceae (e.g. Bactris
gasipaes and Butia spp.) – but also including
legumes (e.g. Caesalpinia spinosa and Inga spp.),
custard apples (e.g. Rollinia spp.) and stone fruits
(Prunus serotina).
Mexico reports the following uses of tree
species that are undergoing improvement: food,
essential oils, forage, gums and resins, Christmas
trees, medicines, conservation and restoration.
Near East
Tree breeding in the Near East region has been
limited, which is surprising given the need and
potential to improve environmental adaptability
to arid environments. In the Islamic Republic of
Iran improvement work is being conducted on
Fagus orientalis, Populus nigra and its hybrids and
Quercus castanifolia for various product traits (for
timber, pulp and NWFP uses) and for resistance
to environmental stresses such as drought and
salinity.
Other species under improvement in the Near
East include local acacias (Acacia senegal, Acacia
nilotica, Faidherbia albida) and the exotics
Eucalyptus camaldulensis for wood production
and Jatropha curcas for biofuel.
North America
North America has a long history of tree
improvement (see Box 14.2) and highly developed
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
programmes. Approximately 85 species are under
some form of tree improvement at various stages.
However, the most intensive breeding efforts
have focused on just a few species, notably Pinus
taeda and Psuedotsuga menziezii.
Canada has major active tree improvement and
breeding programmes in nine of its ten provinces
covering 40 species (of which 33 are native) in 13
genera and hybrid Larix and Populus. Key target
species for improvement are native conifers for
timber and pulp production including Callitropis
nootkatensis, Larix spp., Pinus spp. (P. banksiana,
P. contorta and P. strobus), Picea spp. (P. glauca,
P. mariana and P. sitchensis), Thuja plicata and
Tsuga heterophylla. First-generation programmes
comprise more than 50 000 plus trees selected
predominantly from the natural forest. Secondgeneration programmes are in place for 13
species, and breeding populations contain more
than 9 000 selections from progeny tests and
other tests. Third-generation selections have
been made for Pseudotsuga menziesii.
Box 14.2
Early tree breeding programmes in Canada and the United States of America
Tree breeding programmes in North America began
generating knowledge about a range of timber species
in the mid-1900s. Exploration, testing and breeding of
Pseudotsuga menziesii began in the 1950s in British
Columbia, Canada (Orr-Ewing, 1962). Beginning in
1958, a breeding population was established with
more than 200 Pseudotsuga menziesii sources from
throughout its range in Canada and the United States
of America as well as Mexico. The objective was to
provide as wide a gene pool as possible for a breeding
programme (Orr-Ewing, 1973). Samples of the other
five Pseudotsuga species were established in the
collection as well.
Early efforts in the United States also focused on
Pseudotsuga menziesii. The first seed source studies
were initiated in the northwest in 1912 in Wind River
Experimental Forest, southern Oregon (Duffield,
1959); the tree improvement programmes began in
the 1960s. Because of the topography and associated
high environmental variation over short geographic
distances, many small breeding programmes were
initiated in North America. By the 1980s there were
125 separate breeding programmes for Pseudotsuga
menziesii in Oregon and Washington in the United
States and British Columbia in Canada (Johnson, 2000).
Species covered in subsequent testing and tree
improvement projects in the Pacific Northwest of both
countries include Larix occidentalis, Picea spp., Pinus
contorta, Pinus ponderosa, Thuja plicata and Tsuga
heterophylla, and to some degree Chamaecyparis
lawsoniana, Pinus lambertiana and Pinus monticola.
Elsewhere in the United States, the foundation for
tree improvement was laid in the 1920s and 1930s.
The Placerville Institute of Forest Genetics in California
was initiated in 1925 to improve forest growth through
breeding. This private research institute was first
focused on pines, and 49 Pinus species from many
countries were planted in 1926. The most complete
pine arboretum in the world was established there in
1931. In 1935, the institute was donated to the United
States Forest Service, and it has been an important
knowledge base for forest genetics research since that
time (US Forest Service, n.d.).
In the early 1950s a tree improvement programme
was initiated in eastern Texas, focusing on drought
resistance and wood properties (Zobel, 2005). Tree
improvement cooperatives were established in Texas
in 1952, Florida in 1954 and North Carolina in 1956,
and programmes developed rapidly during the rest of
the century.
Before 1960, no genetically improved seed was
available and all seedlings for planting were produced
from seed collected in the forest, with little control over
quality (Dorman, 1974). By the mid-1970s, much of
the seed used for tree planting in the southern United
States was from genetically improved seed orchards.
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The United States of America has an
exceptionally well developed tree improvement
programme with at least 150 public or cooperative
breeding programmes representing over 70
species. However, more than 50 percent of the
improved seedlings are just two Pinus species:
P. elliotti and P. taeda (McKeand et al., 2007).
Some 66 of the country’s breeding programmes
involve 14 Pinus species; those appearing most
frequently (in from four to eleven programmes
each) are P. elliottii, P. palustris, P. ponderosa, P.
rigida, P. strobus and P. taeda). Other coniferous
species with many improvement programmes
(three to seven each) include Abies spp., Larix
spp., Picea spp. and Pseudotsuga menziesii.
Among broadleaves, the most breeding effort
is focused on Quercus spp. (18 programmes for
seven species, including six programmes on
Q. rubra) and Juglans spp. (a total of 12
programmes for J. cinerea and J. nigra).
Many breeding programmes are part of the
United States Department of Agriculture (USDA)
Forest Service or are based in universities, which
often lead the cooperative breeding programmes.
These tree improvement programmes involve
traditional techniques, often pioneered in the
United States, and the whole array of modern
breeding approaches and biotechnologies,
including genetic engineering. Some of the most
advanced tree improvement programmes are in the
United States; for example, Neale and Ingvarsson
(2008) reported that more than 11.5 million
progeny from more than 41 000 parent trees had
been tested in just four conifer programmes, of
which two were in the third generation of testing
and breeding at the time of their report.
The United States also has breeding
programmes focused on conservation and forest
restoration for several species. For example,
Castanea dentata and Ulmus americana have
both been severely depleted by introduced
diseases – chestnut blight and Dutch elm disease,
respectively (see Box 5.3 in Chapter 5). In each
case the primary breeding objective is to develop
disease-resistant genotypes. Wild individuals
resistant to these diseases are identified, selected,
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and used in breeding programmes to introduce
the genes conferring resistance.
Oceania
In Australia, tree improvement and propagation
programmes are highly developed and variously
undertaken and managed by government
agencies (CSIRO and State departments),
forest industries and/or cooperatives (e.g. the
Southern Tree Breeding Association). These
improvement programmes mostly rely on
traditional methods of selection, breeding,
improvement and propagation. Molecular
markers are being developed and associated
with traits of interest to accelerate selection
of preferred varieties. Species currently under
improvement include:
• native species for timber, poles and
pulpwood, especially eucalypts (23 species
of Corymbia and Eucalyptus and various
hybrid combinations), native conifers
(Araucaria cunninghamii), acacias
(A. crassicarpa and A. mangium) and
Grevillea robusta;
• essential oil species: Backhousia citriodora,
Eucalyptus polybractea and other oil
mallees, Melaleuca alternifolia and
Santalum album;
• exotic timber trees: Khaya senegalensis,
Pinus species (P. brutia, P. caribaea var.
hondurensis, P. elliottii, P. pinaster and P.
radiata) and Tectona grandis.
The larger island nations and territories in
the region have established tree improvement
programmes, often with assistance from
Australian institutions (CSIRO and universities)
and donors, and in the case of New Caledonia
(France) from French institutes (CTFT and
CIRAD). The species under improvement vary
but are mainly highly valuable timbers such as
the exotic Swietenia macrophllya in Fiji, native
sandalwoods in Fiji, New Caledonia and Vanuatu,
and multipurpose nut- and timber trees such as
Canarium spp. and Terminalia catappa in Solomon
Islands and Vanuatu. Fast-growing industrial
species under improvement include Ochroma
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
pyramidale (balsa) in Papua New Guinea and
Pinus caribaea var. hondurensis in Fiji and New
Caledonia.
A worthy example is the spectacular
improvement of Tectona grandis that has taken
place in Solomon Islands in a short period
through intensive screening of teak stands in the
early 1990s, grafting of 50 selected phenotypes
and use of seed from this clonal archive for
production purposes. This teak material is now
being widely used globally, including through
selections supplied to Malaysia.
Solomon Islands reports that through the
Australia-funded SPRIG project, improvement
activities have extended to promising indigenous
timber species including Cordia subcordata,
Gmelina
moluccana,
Pterocarpus
indicus,
Terminalia catappa and Vitex cofassus. In Papua
New Guinea, collaboration with international
and regional agencies and donor support (e.g.
an ACIAR funded collaborative project between
the Papua New Guinea Forest Research Institute
[PNGFRI] and CSIRO) have been vital for seed
collection, provenance trials, seed orchards and
tree improvement.
International collaboration and
donor programmes for tree
improvement
International collaboration includes networks,
commercial exchange, partnership with academic
and research institutes and donor assistance.
Donor programmes may assist establishment
of improvement programmes within a country;
this support may be especially beneficial where
improvement is targeted at rural communities
and others whom the private sector may not
serve adequately.
Regional coordination and cooperation is
especially vital where species and biogeographic
and genecological zones are shared. Many
countries note the central role of international
species networks (for poplars, teak, neem and
casuarina) coordinated by FAO and IUFRO.
Several countries acknowledge the vital role
of Camcore in the development of improved
exotic plantation species through the sharing
of germplasm. Camcore has been a leader in
establishing trials and generating data on conifers
from Central America, with an initial focus on
Pinus species that were of interest for planting
in other regions. Examples include P. caribea,
P. patula and P. tecunumanii (Dvorak, Donahue
and Hodge, 1996). Many of the trials, which also
serve ex situ conservation, are located in tropical
or subtropical countries. The knowledge gained
from these testing efforts is now also useful in
selecting provenances or families for planting in
the species’ natural range.
A cautionary note: potential
threats to FGR from breeding and
improvement programmes
While the benefits of commercial genetic
improvement programmes are profound and vital
for meeting the ever-increasing global demand for
forest products, breeding programmes also have
potential to affect FGR in a negative way if they
are not managed well. The most significant issue
is the loss of genetic variation in improved tree
stock which is then widely distributed, potentially
around the globe. Sweden notes that “forest tree
breeding may in the long run result in decreased
genetic variation in the production forest. Even
though single stands may have somewhat higher
genetic variation, genetic diversity will likely on
the landscape level be lower than in conspecific
natural populations.”
Clonal plantation forestry is regarded as the
most genetically impoverished industrial forest
option, especially where the entire plantation is
derived from only one or a very small number
of clones. Concentration on one or a few
economically important trees in plantations (as in
Ghana, where 90 percent of plantations comprise
only three species) can also contribute to loss of
FGR. Lack of diversity, especially through clonal
forestry, also exposes both planted and natural
forests to pest and disease epidemics, as the
homogenous feedstock provides an opportunity
for pests and pathogens to become specialized
and dramatically increase in numbers.
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Chapter 15
Germplasm delivery and
deployment
The production, distribution and deployment of
germplasm and planting materials are critical to
the continued global supply of FGR-derived goods,
to the development of forest industries and to
programmes for FGR, biodiversity conservation
and environmental restoration. It is estimated
that planted forests account for 7 percent of the
world’s forest area yet produce over 50 percent of
the world’s industrial timber (FAO, 2010a); further
forest plantings, particularly using improved
germplasm, will help satisfy burgeoning global
demand while reducing harvest pressure on
natural forests. As noted by China in its country
report, “the ultimate goal of FGR conservation is
to utilize these resources, and to bring economic,
ecological and social benefits”; the United States
of America similarly remarks “if germplasm is not
readily available for use, resources expended to
preserve it will be wasted”.
Uses of germplasm and plant
materials
Genetically appropriate germplasm is used as a
base for propagating planting materials for the
wide range of uses of planted forests and onfarm and other circa situm plantings, including
timber and pulp, agroforestry products, fodder,
food and fuel. The deployment of germplasm
with appropriate levels of variability is also an
essential component of many FGR and biodiversity
conservation programmes, as well as in
environmental restoration programmes, research
and development. Germplasm is transferred
and exchanged within and between countries
not only for planting, but also for propagation,
research, genetic improvement, breeding, and
conservation of FGR.
Improved or unimproved germplasm
The germplasm and plant materials produced
for deployment may be either genetically
improved or not. For productive purposes,
improved material from genetic improvement
and selection programmes enables the delivery
of larger amounts of desired benefits with fewer
inputs, or better growth under less favourable
environmental conditions. Programmes for
FGR and biodiversity conservation and for
environmental restoration may also benefit
from intervention and breeding to enrich
genetically impoverished remnants, or to
introduce characteristics vital to the survival
of threatened species; an example is the
introduction of resistance to chestnut blight into
North American chestnut (see section on North
American improvement programmes in Chapter
14). By contrast, many programmes aimed at
conservation of FGR require the inclusion of
materials representing the fullest range of local
genetic variability (while excluding importing
germplasm from more distant provenances or
other seed-transfer or genecological zones).
Plant materials used for commercial and utility
plantings and plantations are largely derived from
high-quality, improved, source-identifed seed and
plant materials. In Chile, for example, about 95
percent of planted forests of exotic species are
derived from improved material, while less than
2 percent of natural forests have some degree of
genetic improvement. For the private sector, the
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use of improved planting material is driven by the
need to obtain an adequate return on investment
by obtaining the highest yield for the lowest cost.
Countries with well developed genetic
improvement programmes generally have
relatively well developed methods of distributing
and deploying their improved materials, whether
in the private or public sector.
Forms of germplasm
The form in which planting materials are
delivered to end-users depends on the purpose
of the planting programme, the characteristics
of the germplasm, the nature of the planting
situation, the method of propagation most suited
to the species, the capacity for production of
plant materials, and the capacity of the people or
organization undertaking the planting.
The country reports indicate that material
available for transfer, both within and between
countries, is strongly dominated by seed (66
percent of the total). Seed is generally the most
convenient and safest form for germplasm
transfer; however, for some species seed is not
suitable for collection, storage, propagation and/
or distribution. Reasons include inaccessibility
of collecting materials, intermittent seed
production, lag time between improvement
and seed production, difficulties in ensuring
that progeny are true to type, limited storability
(recalcitrant seeds), and lack of knowledge
on propagation methods. Macro vegetative
propagation methods (cuttings and micro
cuttings) are commonly used for producing clones,
and micropropagation techniques (tissue culture)
are increasingly applied, although the cost and
technical requirements of micropropagation
are prohibitive for some countries. Nonetheless,
industrial-scale production by micropropagation is
being undertaken for some species of commercial
interest or for large restoration purposes despite
the high costs.
The predominance of seed as the most available
form of germplasm suggests that distribution,
deployment and perhaps even research and
improvement may be skewed towards species that
190
produce orthodox seed that can be transferred
most conveniently and reliably – perhaps at the
expense of other potentially useful species or
species that require conservation.
Demand for germplasm and
planting materials
Countries vary greatly in the amount of germplasm
and planting materials deployed, ranging from
negligible (e.g. in many SIDS) to extremely high
in countries with major planting programmes
such as Brazil, China and Indonesia. Brazil, for
example, planted 330 000 ha per year between
2005 and 2010 and has a huge demand for
planting stock. Some countries note a paucity of
information on demand for germplasm, although
projected increases in demand for wood products
invariably imply an increase in requirements
and a shortfall in supply at current stocking and
planting rates. National programmes requiring
planting stock include planting programmes for
forestry production, combating desertification,
landscape-scale climate change adaptation and
mitigation, environmental restoration and forest
restoration in the wake of major infrastructure
projects.
The approach to management of a country’s
production forests in large part determines the
demand for germplasm and planting materials;
for example, a forest industry based on plantings
will deploy more germplasm and plants than one
reliant on naturally regenerated forests. Several
countries, especially in Asia (e.g. Samoa, Sri Lanka
and Thailand), refrain completely or in large
degree from harvesting their natural forests,
deriving local timber from planted sources. In
these countries, the demand for adapted and
improved germplasm and planting materials
is relatively high. Many countries, particularly
in temperate and boreal regions, adopt hybrid
management models involving a mixture of
artificial and natural regeneration of natural
forests as well as planted forests. For example,
in Canada – the world’s second largest exporter
of wood products in 2012, behind China (Natural
Resources Canada, 2013) – 349 000 ha of land
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
were planted in 2011 and another 11 000 ha
were seeded. In other countries, the availability
of low-cost, high-quality timber resources in
natural forests has served as a disincentive to
invest in development of improved germplasm
and planted forests; these countries have lower
demand for germplasm and planting materials.
Actors involved in production,
distribution and deployment
In many countries vast amounts of tree seed
and seedlings are produced from seed stands,
seed orchards, nurseries and other facilities
on public forest or land, with public-sector
agencies and seed centres key in the exchange,
delivery and deployment of germplasm. In
some contexts the private sector is preeminent,
generating large amounts of germplasm and
plant materials for use in corporate plantation
forestry. In most developing countries and some
developed countries, small-scale seed collectors
and producers, including community or villagelevel entities and individual landholders or
farmers, also have important roles, producing
tree germplasm and planting stock for their own
use or for sale.
Public sector
Public departments, agencies and corporations
with remits in forestry, conservation and natural
resource and land management may have a
central role in the production, distribution and
deployment of FGR materials. Country reports
detail the involvement of the public sector in
these activities for purposes of:
• conservation of FGR, including genetic
diversity in threatened or potentially
threatened species;
• research and tree improvement by local,
national and international organizations
(both public and private);
• supplying planting materials to government
agencies, corporations, NGOs and
individuals;
• forest plantation development on public
land;
• environmental restoration programmes;
• advancing national development and other
policy goals – e.g. poverty alleviation,
food security, climate change mitigation,
biodiversity conservation, environmental
protection and both small- and large-scale
industry development.
Public-sector involvement is thus consistent
with the view expressed in China’s report that
the “collection and conservation of FGR is a
basic, long-term, public welfare and strategic
work”. Public-sector involvement – including the
ownership of forest resources and land used for
collection of germplasm, the creation of seed
orchards and the establishment of plantations –
is especially important in developing countries
where markets are poorly developed or do not
function effectively (whether because of the type
of political and economic system; the inability
of commercial operators to generate income or
capture rewards from the activity; the lack of
regulatory, legal and financial infrastructure; or
the lack of capital).
Foremost among the public agencies involved
in germplasm delivery and deployment are
the national tree seed centres (NTSCs) or their
equivalents, which in many countries serve
as primary agents for the collection, storage
and distribution of forest tree germplasm to
government forest and conservation agencies and
to the private sector (including nurseries, forestry
companies, NGOs, communities and farmers).
NTSCs are also typically involved in regional
and international transfer and exchange of
germplasm. In addition, NTSCs may have central
roles in ex situ conservation and improvement of
FGR.
A number of countries report that public
nursery facilities, either associated with NTSCs or
attached to other public bodies, are significant
producers of plant materials. NTSCs and public
nurseries may also generate funds through local,
regional or international sale of germplasm or
plants, consultations and/or supply of contracted
services to the private sector. For example,
Madagascar’s National Tree Seed Centre sells
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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60 percent of its production to the private sector
and NGOs, and the rest to government agencies.
Countries reporting that NTSCs currently have a
major role in germplasm delivery and deployment
include Australia, Burkina Faso, Ethiopia, Kenya,
Madagascar, Nepal, Papua New Guinea, Sri Lanka
and Zimbabwe. Guatemala’s NTSC, on the other
hand, cannot produce germplasm for commercial
purposes under existing legislation, and many
countries (including much of Africa and most
SIDS) report that their NTSCs have inadequate
facilities and support. Countries often report
the need to increase the productive capacity of
tree seed centres to meet existing and projected
demand for germplasm.
Many public agricultural agencies have
longstanding involvement in the production
and distribution of improved plant materials –
particularly in countries where native FGR include
many progenitors of widely used fruit and nut
trees, as in Western and Central Asia, or where
food- or crop-bearing trees are important as
cash crops or as contributors to food security.
In some countries these agencies also develop
and propagate trees to be used in agricultural
applications, and they may also have a major
role in the ex situ conservation of FGR. In some
countries with meagre resources, these activities
provide at least some level of ex situ conservation,
storage and distribution of forest germplasm.
The public sector also has a vital role in
developing
policy
governing
germplasm
development and deployment, for example:
• enacting laws and regulations governing
germplasm collection, delivery and
deployment;
• developing, implementing and enforcing
standards, guidelines and protocols;
• mandating and overseeing certification
schemes for the production, movement and
exchange of forest germplasm and planting
materials;
• facilitating and regulating privatesector involvement in FGR delivery
and deployment, including facilitating
192
and participating in organizations and
associations concerned with germplasm
delivery and deployment.
Finally, the public sector often has a role in
increasing adoption of improved germplasm. In
many countries the public bodies undertaking
breeding and improvement, such as the NTSCs,
research institutions and forest and agriculture
departments, undertake activities in promotion,
distribution, education and extension to ensure
the deployment of their improved materials.
Private sector
The private sector – including companies, from
local to multinational; small-scale, village and
community enterprises; farmers and landowners
– is increasingly the primary actor in germplasm
delivery and deployment. Private producers
propagate and deploy vast amounts of germplasm
and plant materials, generally for their own use
or to supply contracts. In some instances the
materials may be developed for sale on the open
market; if the species are rare or threatened (e.g.
some ornamental plants), their commercialization
can have both positive and negative implications.
Private-sector participation in germplasm
delivery and deployment can complement public
efforts where the public sector has limited funding,
political will, institutional capacity or remit or
where it has difficulties engaging with market
processes. Where markets operate effectively, the
private sector can respond efficiently to market
signals and demand for goods and services.
Larger, particularly multinational, companies may
have access to capital, technology and expertise
that may be unavailable in developing countries;
if managed appropriately, their activities can help
advance national forestry, FGR conservation and
other development agendas.
The private sector exercises a major inluence
on germplasm delivery and deployment through
landownership. In many countries private
corporations, small enterprises, communities,
villages, families and individuals own or control
signiicant proportions of the total land area.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Private land may be used for establishment
of forest plantings of various kinds, in pursuit
of commercial, environmental or biodiversity
conservation outcomes. In other situations
the private sector may access public land for
collection of germplasm and establishment of
plantations (e.g. in Australia and Indonesia).
The landownership pattern helps determine the
balance between public and private involvement
in germplasm delivery and deployment and the
manner in which these activities are conducted.
The private sector also contributes to uptake
of improved germplasm. In cases where private
enterprises undertake improvement and breeding
activities for sale, they have significant financial
incentive to promote the use of their products.
The following issues concerning privatesector involvement in germplasm delivery and
deployment are gleaned from the country
reports.
• The private sector must be responsive to
market demand. However, where demand
is internationally driven, private-sector
activities may not always be consistent
with national strategies or programmes in
the areas of forests, FGR conservation and
management and national development.
• Adequate protection of intellectual
property and reward (e.g. through sales) are
required to encourage sharing of improved
germplasm.
• A stable investment environment is
required.
• The private sector’s ability to deliver public
goods and benefits may be limited if
markets for them are poorly developed or
non-existent.
• The private sector tends to address
ecological and social objectives only as
required by regulation.
• In the interest of maintaining commercial
advantage, companies may refrain from
sharing improved materials that they deem
proprietary. Sharing may occur through
sales but may thus be limited where markets
do not exist or are limited (e.g. because of
lack of capital).
• Information on private-sector germplasm
delivery and deployment activities may be
difficult to collect because of confidentiality
issues and lack of a process for collecting
and collating these data.
Informal sector
In many countries, small-scale production of
germplasm and planting materials, and their
use in forest and farm plantings, provide
rural employment and income which assists in
alleviating poverty. For example, traditional
home-garden agroforestry practices in Ethiopia
provide employment for many people, and
propagation and exchange of plant materials
are part of the fabric of rural life. Home gardens
produce 42 percent of the timber and 27 percent
of the fuelwood in Sri Lanka, while forest
plantations generate only 11 and 4 percent,
respectively; this decentralized, informal system
of wood production also produces much of its
own planting materials. However, smallholder
farmers and other small-scale producers in the
informal sector often have little or no access to
appropriate germplasm and plant materials,
hampered by lack of money, lack of networks
or information on the availability of germplasm
and its benefits, logistical barriers to delivery or
failure of markets.
Informal small-scale collection and propagation
of germplasm by farmers and forest-adjacent
communities (for use on farms, in home gardens
or for sale) is often consistent with centuriesold tradition and practice. Selection, if any, is
usually based solely on phenotype, for example
harvested from a limited number of parent
trees which may be related (half-siblings) and/or
relatively isolated with high levels of inbreeding.
Such germplasm tends to be of relatively poor
genetic quality and to have little variability. The
Philippines and Madagascar report that smallscale seed producers or individuals generally
collect material from few plants (e.g. less than
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ten), which leads to loss of genetic variability.
These practices limit opportunities for genetic
improvement and maintenance of variability of
FGR. Where collection practices do not conform
to the requirements of national, regional and
international certification schemes (discussed
below), producers will have limited ability to
sell into germplasm markets that demand this
certification.
Despite the difficulties of ensuring adequate
variability, ensuring quality and selecting
appropriate material, opportunities exist for
developing production and use of improved
germplasm in the informal sector. These producers
require access to affordable and appropriate
planting materials, information and market
assistance; in this area NTSCs and extension
services may have an important role, especially
in developing countries. Other strategies
include boosting community involvement in
germplasm production, for example through
participatory models of selection, improvement
and plant production. The United Republic of
Tanzania recommends “strengthening farmer
seed systems” and increasing their access to
information to enhance their contribution to
conservation and promotion of diversity. Sri
Lanka recommends incentives for increased tree
planting in home gardens and development of
a partnership approach for production of highquality planting materials.
The germplasm collection and propagation
activities of the informal sector may also be
supported and improved through establishment
of grower cooperatives and associations. In
the Philippines, for example, the Agroforestry
Tree Seed Association of Lantapan, a farmer
association established in 1998, has educated
many small seed producers in correct techniques
to meet standards and obtain an assured market
for their seeds.
Engaging community volunteers is another
avenue for production, distribution and
deployment of planting materials for conservation
purposes in both developed and developing
countries alike. In the United States of America,
194
for example, “friends” groups help obtain native
plant donations for forest restoration projects.
The “Seeds of Success” programme in the United
States also mobilizes volunteers. Seychelles reports
that the Division of Environment plans to mobilize
people to participate in conservation of endemic
tree species, first by growing indigenous trees.
Public−private partnerships
With the complex interplay of complementary,
overlapping and competing roles of the public
and private sectors in germplasm collection,
storage, propagation, delivery and deployment,
coordination and collaboration are essential.
Countries mention several joint approaches to
delivery and deployment.
Some governments offer incentives to private
companies for the production of high-quality
germplasm and planting materials, particularly
for activities consistent with national goals, e.g.
for conservation of forests, FGR and biodiversity.
In
some
countries,
associations
and
organizations with joint public and private
sector membership assist in data collecting,
standardization, and preparation of guidelines
and policy related to FGR, including matters
relating to exchange and deployment. Examples
include:
• the Canadian Forest Genetics Association,
which promotes information exchange and
sound practice and policy;
• the National Plant Germplasm System in the
United States of America, which has both
government and industry involvement and
includes accessions of 87 percent of the tree
genera in the country;
• in Chile, the Cooperativa de Mejoramiento
Genético, a joint enterprise involving public
agencies, the private sector and a university,
which regulates, certifies and documents
seed produced by its members;
• the cooperative tree improvement
organization GENFORES in Costa Rica, led
by the Technological Institute of Costa Rica
and involving 11 reforestation companies
and local NGOs.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Sometimes initiatives are devised and
implemented jointly. For example, in Germany
private seed certification schemes have been
developed with input from private organizations
and state and federal governments.
In some cases different aspects of the delivery
and deployment system may be shared and/or
divided between the public and private sectors. In
Germany, seed harvest and storage is undertaken
by both private and public organizations, but
the collection and long-term gene bank storage
is undertaken by the state governments. The
Islamic Republic of Iran reports that government
agencies may directly import and distribute
seed or provide oversight of the private sector;
“all the seed imports and distribution is done
either directly by the Ministry of Agriculture or
by private companies after receiving permission
from the Ministry”.
Production of germplasm and
planting materials
Germplasm used by countries as base material for
propagation is selected or improved to varying
degrees and held in a variety of propagative
resources and facilities, including:
• unimproved but selected forest seed stands;
• seed, cutting and clone orchards of
phenotypically selected or genetically
improved genetic materials (Figure 15.1);
• seed and vegetative materials from
individual trees in circa situm situations.
FIGURE 15.1
Most widely planted species in seed orchards
Pinus sylvestris
Picea abies
Tectona grandis
Pseudotsuga menziesii
Fraxinus excelsior
Eucalyptus camaldulensis
Pinus patula
Betula pendula
Alnus glutinosa
Quercus rubra
Prunus avium
Larix kaempferi
Larix decidua
Eucalyptus grandis
Pinus caribaea
Picea sitchensis
Larix x eurolepis
Grevillea robusta
Acer pseudoplatanus
0
5
10
15
20
Number of countries reporting the species’ presence in seed orchards
Africa
Asia
Europe
Latin America
and the Caribbean
Note: Countries of the Near East did not provide data on seed orchards.
North America
Oceania
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In the Philippines, seed stands “are identified
and delineated in natural stands or plantations
with a high frequency of phenotypically good
planting materials”, while seed orchards are
plantations of selected trees, clones or progenies
“which are isolated or managed to avoid or
reduce pollination from genetically inferior
sources outside the orchard, and intensively
managed to produce frequent, abundant, and
easily harvested crops of seeds; a seed orchard
can also be regarded as a breeding population
as a basis for further tree improvement”.
Vegetative materials for propagation from
grafts, stem cuttings and micro cuttings may
be sourced from selected clones or seedlings.
Cloned materials are favoured for plantations
because of their ability to produce true-to-
type, genetically uniform plants from improved
material; they are especially useful for the
heterosis in F1 interspecific hybrids. In many
countries, small-scale or individual producers
may select seed informally, both for use in their
own enterprises or for sale. The implications
of small-scale informal selection are discussed
below.
Country infrastructure for the production
of tree germplasm and planting materials not
only includes the above-mentioned propagative
resources, but also the organizations, enterprises
and facilities dedicated to germplasm collection,
storage and propagation (Box 15.1), such as
government agencies, national tree seed centres
(NTSCs), private corporations, public and private
nurseries, community associations, villages and
Box 15.1
Germplasm production, storage and progagation and distribution facilities:
some challenges
Germplasm production and storage facilities require
committed, ongoing management to maintain and
operate effectively. They require adequate funding,
which can come from government or industry support
and/or the sale of germplasm and planting materials.
Dry/cold seed storage facilities are subject to
humidity, temperature fluctuations, floods, fires, insect
attack and potentially catastrophic power failures
(in the absence of backup generators). The need to
replace or repair failing equipment is a significant
impediment to the success and expansion of ex situ
programmes and the operation of many national tree
seed centres (NTSCs).
Zambia mentions that its seed is stored in deep
freezers, “as all the cold rooms need rehabilitation.
A standby generator may need to be installed to
minimize effects of power disruptions.” Zimbabwe
notes intermittent power supply to cold storage rooms
as a major challenge to its NTSC. Newer freezers
196
with defrosting cycling have fluctuating temperatures
which considerably shorten seed lifespan, but some
people working with tree seeds may not be aware of
this information.
Seed and clone orchards also require ongoing
maintenance and management. They may be subject
to threats that include neglect, poaching, illegal
harvesting and encroachment of agricultural and
other activities. Several countries, including Cameroon
and Zimbabwe, note decline or disruption in their
orchard operations because of such factors.
Political, social and economic stability are required
to ensure continuing maintenance and operation of
these facilities. Because of the risks, it is not advisable
to have a single focal point for the production and
distribution of germplasm. Strategies to encourage
safety duplication with widespread production by
multiple organizations, companies or individuals can
reduce the risks.
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
individuals. Other crucial components include
human and organizational resources – expertise,
personnel, funding and political support.
Country reports describe great variation in
germplasm and plant production capacity, as
illustrated in the following examples.
Country examples
Africa. South Africa produces 200 million plants
from 37 nurseries, almost all Pinus, Eucalyptus and
Acacia species, for its commercial forest industries.
In Ethiopia, the government is the sole supplier of
tested seeds in the country, distributing 7.2 tonnes
of seed annually, capable of producing 570 to 880
million seedlings; however, the informal sector is
also a source of germplasm, which is less reliable
(see Box 15.2). In many other African countries
seedling production is much lower; for example,
Madagascar’s NTSC nursery produces 250 000
plants per year. Most small developing tropical
countries typically have less than five or ten
nurseries producing tree seedlings for production
and environmental plantings.
Asia. China, with its immense area of plantings,
has a vast demand for planting materials of
many types and reports a remarkably extensive
decentralized network of 336 000 tree seedling
nurseries covering 668 000 ha. Tree nurseries
in China source their propagation materials
from:
• 19 600 ha of improved seed orchards
comprising progeny-tested superior families,
selected plus trees and introduced superior
families, yielding an average of 0.71 million
kilograms of seed per year;
• 18 100 ha of improved cutting orchards
of bred and introduced superior clones of
various species, yielding an average of
1.8 billion seedlings annually;
• 146 000 ha of superior seed stands
yielding 1.67 million kilograms of seed
per year;
• 630 000 ha of “seed collection bases” of
superior provenances of several species and
genera, yielding 9.3 million kilograms of
seed per year.
Box 15.2
Germplasm production and dissemination in Ethiopia
The scenario of germplasm production and use
described by Ethiopia demonstrates elements that are
common to many developing countries, particularly
those with large informal-sector involvement and a
multiplicity of actors, including the public sector.
The federal Forest Research Center (FRC, an arm
of the Ethiopian Institute of Agricultural Research)
is the only supplier of tested forest tree seeds in
the country. FRC collects forest germplasm from
identified and established stands. It sells the forest
germplasm collected from these stands to government
organizations, mainly bureaus of agriculture, and to
NGOs and private seed growers.
Many smallholder farmers, youth, women and
private seed dealers and nursery operators are
also engaged in the forest germplasm business.
Seeds provided by farmers, private seed dealers and
other informal sources are often of low quality and
quantity. Movement of forest germplasm within
the country is unrestricted and involves a range of
stakeholders, including government, NGOs, local
people, private seed dealers and nursery operators.
Local communities, especially young people, benefit
from the production and sale of seedlings of certain
forest species.
Smallholder farmers collect and plant seedlings or
sow seeds of various trees for their own use in a variety
of agroforestry and on-farm plantings, or for sale.
Source: Ethiopia country report
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China reports that its major forestry
programmes increasingly need to use improved
genetic materials, but that most of the seed
orchards and stands are first generation and low
in quality.
Other Asian countries reporting significant
productive capacity include Indonesia and
India. Indonesia, through the Department
of Land Rehabilitation and Social Forestry,
produces 29 million kilograms of seed annually
from 104 000 ha. India produces 3.8 million
kilograms of seed of 27 species and plants 102
million seedlings annually. India has 396 seed
orchards covering 7 090 ha (of which 64 percent
is teak).
Europe. Germany has 425 forest tree nurseries
producing 150 million to 185 million plants
annually and 800 ha of seed orchards for trees
and shrubs, including 215 seed orchards for
tree species. In Poland about 90 percent of the
currently established planted forests come from
planting or sowing; Poland imports large amounts
of tree seed (in the order of 10 tonnes per year),
but exports tree seedlings. Several European
countries, including the Russian Federation, rely
heavily on natural regeneration of their forests.
Meeting increased demand for
germplasm and planting materials
As mentioned above (see section on demand),
nearly all countries expect increases in demand
for planting materials and a shortfall in supply,
and many country reports recognize that to fill
the gap it will be necessary to boost production.
Ethiopia, for example, identifies a need for
“…strengthening the tree seed production−
supply system for satisfying needs for quality seeds
and collection and conservation of germplasm”.
The Philippines remarks that “the fundamental
problem to be addressed at this point is the lack
of supply of improved planting materials for
production purposes, and of planting materials
for conservation of endangered indigenous and
other forest genetic resources”. Madagascar
198
similarly notes the need for large-scale, low-cost
production of planting materials for forestry and
mining restoration projects.
Zambia attributes a 38 percent reduction in
plantation area since 1982 in part to “reduced
national capacity to produce suficient good
quality Pinus seed for plantation establishment”.
Because of this critical shortage of high-quality
seed, “seed is at present being imported from
Asia and other places, and used for plantation
establishment without any screening in
provenance or progeny trials….” Zambia notes
that the quality of exotic tree seeds of Pinus
and Eucalyptus species collected annually by the
Forestry Department has declined and that the
quantity has decreased from 250 kg of seed in the
1970s to 25 kg in 2010.
To meet shortfalls in supply of planting
materials, a number of countries express an
interest in developing their capacity to use
advanced propagation techniques. Cloning,
for example, offers the prospect of producing
large numbers of genetically identical improved
planting materials at a low cost. Other techniques
such as in vitro micropropagation and somatic
embryogenesis may assist a range of FGR
conservation programmes, particularly where
the species involved are dificult to propagate or
where mass propagation of superior seedlots is
sought.
A cost-effective, low-technology approach
to boosting the supply of plant materials, both
from seed and through vegetative means,
involves encouraging production by small
producers, communities, villages or farmers using
participatory methodologies.
The time required for seed orchards to produce
improved seed is a constraint on the delivery
of improved germplasm for deployment, often
necessitating the continued use of unimproved
seed. A number of advanced techniques in
breeding and propagation may shorten the time
between improvement and deployment, including
for example increasingly powerful selection
based on QTLs and marker-aided selection,
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
grafting, micropropagation improvements (e.g.
the use of bioreactors), somatic embryogenesis,
and application of hormones to promote early
lowering in seed orchards.
Movement and transfer of genetic
material
The transfer of forest reproductive material has
been a common practice for several centuries. Field
trials established with introduced material have
provided valuable insight on the performance of
different tree species and their provenances, in
turn inluencing requests for further germplasm
transfer and for the opportunity to test new
material.
As discussed above in the section on the actors
involved, both the public and private sectors carry
out activities to promote and deliver improved
material (in the latter case for commercial
gain). Countries with well developed genetic
improvement programmes generally have
relatively well developed methods of distributing
and deploying their improved materials, whether
in the private or public sector.
However, the high collection costs and
dificulties of access to FGR have made it
increasingly dificult to move forest reproductive
material for research purposes. As a result the
large international programmes carried out in
the past to assess systematically the performance
of forest reproductive material would not be
possible today.
The transfer of forest genetic materials for use
in research and tree improvement contributes
to global economic progress by facilitating the
production of better adapted trees with superior
performance that can produce goods and services
more eficiently and cheaply. International
transfer and exchange of germplasm allows
countries that lack the capacity to undertake
their own improvement programmes to partner
with or outsource to countries or external
organizations with the relevant expertise
and resources. Through international trade in
germplasm and plant materials, countries are also
able to purchase improved planting materials to
supply their forest planting and development
programmes.
International trade in germplasm and plant
materials is an important commercial activity in
its own right. China, for example, exports over
300 000 kg of seed and several hundred thousand
seedlings annually for over 400 species. Australia
also has a major international seed export trade,
mainly in Acacia and Eucalyptus species.
Control mechanisms are required to ensure that
the transfer and use of FGR is safe, appropriate
and fairly compensated. Brazil, for example,
reports: “Any intended utilization of genetic
material, native and exotic alike, must comply
with speciic laws and regulations. The import,
export, research and improvement of plant
genetic resources are regulated by phytosanitary,
environmental, access, beneit-sharing and
intellectual property legislation.”
Prioritizing for delivery and deployment
Priorities for the delivery and deployment of
FGR and improved genetic materials are best
established at the national level, in national
strategies for forests and FGR conservation and
management, coordinated with other relevant
strategies. Priority objectives include addressing
commercial or forest service goals, FGR and
biodiversity conservation goals, and development,
social and economic goals. Focal areas include
development of plantation industries; developing
local, small-scale industries and employment;
assisting rural communities; contributing to
poverty alleviation; reducing harvest pressure on
natural forests; promoting environmental and
restoration plantings; combating desertiication;
and conserving genetic variability and threatened
species.
Other priorities to be considered for delivery
and deployment programmes include desirable
genetic traits and production characteristics,
genotypes, populations, species, improved
varieties and planting locations, as well as the
sectors of the economy and society identiied
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for development or assistance. The private
sector mainly prioritizes species, locations and
management regimes for deployment on the
basis of financial returns, which are in turn
determined by market conditions, structure and
demand, although government regulations,
incentives and support may also have a role.
In developing countries it is important that
delivery and deployment of plant materials be
matched with the areas of most need, particularly
given the importance of trees in alleviating
rural poverty. For example, in countries where
fuelwood is a major source of energy, the
improvement, production, distribution and
deployment of multipurpose and fuelwood
species may be considered a priority. Several
country reports remark on the need to match
planting materials more effectively with
their intended purpose. China notes that
market signals may help in the deployment of
appropriate plant materials.
A number of countries, often in collaboration
with ICRAF, adopt a participatory approach,
engaging rural communities, farmers and villages
to facilitate alignment of selection, improvement,
production and deployment of FGR with
community needs. This type of approach is
especially advantageous where markets are
lacking, poorly developed or poorly functioning,
for example because of poverty, lack of purchasing
power, lack of market infrastructure or lack of
market signals and information.
Ensuring the quality of germplasm and
planting materials
To give surety to purchasers of germplasm and
obtain access to markets that require guarantees
of quality, vendors may seek to certify the quality
of their seed and plant materials. International,
regional and national certification schemes
involve adherence to procedural guidelines and
protocols. Quality parameters may include the
source (e.g. geographic location or biogeographic
or genecological zone), whether the material
is improved stock, the breeding generation,
200
the level of natural variability represented,
improvement in performance of desired
characteristics, the collection protocols followed,
phytosanitary criteria, and the quality and health
of the seed lot or plant material batch, including
germination rates.
Other means of assuring, increasing and
monitoring quality include regulation, voluntary
codes of practice, the use of guidelines and
producer education. As mentioned above, in a
number of countries private−public collaborations
or smallholder, farmer-operated associations (e.g.
the previously mentioned Lantapan cooperative
in the Philippines) are involved in development
of guidelines, quality control, certiication and
documentation of seed.
Several regional and international mechanisms
exist for quality control and certiication. The
OECD Forest Seed and Plant Scheme (OECD, 2013),
for example, governs international exchange
of certiied material and aims to have “a major
role in helping world forests adapt to changing
climatic conditions”. This scheme emphasizes
“preserving species diversity, and ensuring high
genetic diversity within species and seed lots
thereby enhancing the adaptive potential of
forest reproductive material for [the] future”.
The scheme now speciies six categories of “basic
material” from which reproductive material can
be selected, i.e. seed source, stand, seed orchards,
parents of family/ies, clone and clonal mixture.
The components of the basic material will have
been selected at the individual level and tested;
the superiority of the reproductive material must
be demonstrated by comparative testing or an
estimate of its superiority calculated from the
genetic evaluation of the components of the
basic material.
Within the European Union, germplasm is
freely exchanged in accordance with EU Directive
1999/105/EC (EU, 1999), which governs particular
species and also provides a standard classiicatory
system for reproductive material. EU regulations
allow members to import forest reproductive
materials in four categories – source identiied,
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
selected, qualified or tested – from EU-approved
external third parties.
Various EU member countries have enacted
laws to implement EU directives on germplasm
production and transfer. EU members may
impose regulations if these exceed the quality
requirements of EU Directive 1999/105/EC (EU,
1999). Germany’s Act on Forest Reproductive
Material governs activities relevant to production
and exchange of germplasm of tree species, for
example by encouraging the improvement of
seed quality through certification. It specifies
categories to describe forest reproductive
material for exchange, and it regulates
commercial production and marketing as well
as imports and exports of forest reproductive
materials. Production and sale of these materials
is restricted to registered seed and plant material
producers, and all materials must be approved.
International forest certification schemes
such as the Forest Stewardship Council and
the Programme for the Endorsement of Forest
Certification (PEFC), which have been adopted
by a number of countries, include requirements
related to the quality, nature and genetics of
forest germplasm and planting materials.
China has implemented a set of rules requiring
and governing the use of improved forest trees
in major forestry programmes; however, uptake
has not been as high as hoped owing to the lack
of incentives for seed producers and users and
geographic limitations.
In Denmark the promotion of improved genetic
material requires approval by law; approval is
granted if the material is deemed above average.
Madagascar categorizes the sources used to
produce seed in line with OECD guidelines,
i.e. assessed sources, selected stands and seed
orchards; seed is tested following International
Seed Testing Association (ISTA) standards. Other
countries encourage the use of improved or
preferred germplasm rather than regulating it,
particularly where small landholders are the main
users; for example, in the Islamic Republic of Iran,
“although the use of recommended varieties
by the farmers [has] been encouraged by the
government, there is not any legal prohibition
preventing them from using a farmer’s variety”.
Identifying regions of provenance and
genecological and seed-transfer zones
The identification of regions of provenance
and of a country’s seed- or germplasm-transfer
or genecological zones (i.e. areas possessing
consistent biogeographical characteristics to
which local populations have adapted) facilitates
the selection and deployment of appropriately
adapted plant materials best suited to local
conditions. The deployment of germplasm from
different, non-conforming genecological zones
may lead to the loss of adaptive advantage
(including through potential genetic impacts on
existing populations) as well as poor performance
or, at worst, complete failure of the introduced
material. Knowledge of the biogeographical
origin of germplasm is also essential in
international transfer for purposes of research
and genetic improvement.
Both the OECD Forest Seed and Plant Scheme
(OECD, 2013) and EU Directive 1999/105/EC (EU,
1999) (see above) address matching of tree
germplasm to planting sites, in requiring the
identification and delineation of provenance
regions. The EU directive cites research showing
that “it is necessary to use reproductive material
that is genetically and phenotypically suited
to the site”; that “demarcations of regions of
provenance are fundamental to selection”; and
that “native species and local provenances that
are well adapted to site conditions should be
preferred”.
Some countries have identified seed-transfer
or genecological zones (fully or partly) and have
focused their selection and tree improvement
programmes on materials adapted to these
zones. The zones are also used as the basis for
determining the movement and transfer of
germplasm.
On the other hand, many developing countries
have not yet defined their genecological zones
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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and have neither identified nor developed
appropriately adapted species and improved
materials for planting in different zones.
Ethiopia identifies the need for “establishing
and strengthening a system for the provision
of indigenous and exotic tree species and seed
inputs that are suitable for the different agroecological zones”. India identified 147 seed zones
in 1978, with the intention of obtaining legislative
support to implement a zoning system; although
at the time the initiative suffered from lack of
support, a bill to enforce the zoning system is
now pending before the Indian Parliament.
A number of countries (e.g. Germany) adopt
strict controls on the movement of genetic
materials across genecological boundaries
or seed-transfer zones; other countries focus
on identifying, selecting and developing
FGR that will perform well in particular
zones. The implementation of seed-transfer
zones encourages tree breeders to develop
appropriately adapted materials by ensuring a
market for them. China, however, notes that too
narrow a geographic focus for development of
improved materials may restrict their application
and deployment.
Some countries that apply the seed zone
concept have yet to develop national guidelines
for transfer within their borders. For example,
Canada notes that provinces develop their own
propagation materials based on seed zones,
often using information from provenance trials,
but the country has no national legislation or
guidelines regarding transfer within the country.
Poland, on the other hand, has strict rules for
the movement of forest reproductive materials
within its borders, for example through the
Forest Reproductive Material Act of 2001. These
rules cover movement not only between regions,
but also between altitudinal zones.
Several countries (e.g. Germany, Sweden
and the United States of America) note that
with climate change, species and germplasm
identified for adaptation to new and rapidly
changing conditions may differ from the
materials identified to date as appropriate for
202
existing genecological zones. Germany, for
example, is currently increasing its use of climatechange adapted species such as the introduced
Pseudotsuga menziesii. The United Republic
of Tanzania remarks that increased transfer of
germplasm may be needed to select and breed
trees better adapted to changes in climate, to
enrich the variability of local FGR to facilitate
their adaptation, and perhaps to assist migration
of certain species to ensure their survival.
The use of transferred germplasm in this way
may run counter to the strict enforcement of
genecological zoning approaches; lexibility will
be required to accommodate changes, particularly
while appropriately adapted species and genetic
materials are being identiied and developed.
Managing risks in transfer and exchange
of germplasm
Phytosanitary, pest and invasive species risks
accompany the transfer and exchange of
germplasm and plant materials. To minimize these
risks, a number of organizations, including FAO,
Bioversity International and the DANIDA Forest
Tree Seed Centre, have developed guidelines for
the safe movement of tree germplasm (see FAO,
2007b). Various mechanisms and regulations exist
at international, regional and national levels to
manage the risks, although they are sometimes
limited in their application. Where regulations and
standards differ between jurisdictions (within a
country, regionally or internationally), movement
of germplasm and tree planting materials from
one jurisdiction to another may be problematic.
Indeed, a lack of harmonization in regulations,
standards and transfer protocols creates barriers
to the eficient exchange of genetic materials;
resultant delays in the supply system may cause
loss of viability in orthodox seeds and mortality
in recalcitrant, dessication-sensitive or short-lived
seeds.
Benefit sharing
Firm, adequate and enforceable beneit sharing
arrangements are essential to ensure that the
interests of the owners of any materials exchanged
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
are duly recognized and that owners receive an
appropriate share of the benefits. Benefit sharing
mechanisms are required to ensure that owners
of germplasm, or of information contributing to
the improvement of that germplasm, are treated
equitably. They allow developers of improved
varieties and other suppliers of services to capture
income generated from their investments and
activities, thereby providing incentives for further
improvement work. Such arrangements are
similarly required when a country contributes
genetic material to a collaborative improvement
programme.
The dialogue on access and benefit sharing
issues for forest genetic resources has thus far
been rather limited in most countries; however,
these issues are increasingly being considered,
following the example of the agricultural sector.
Benefit sharing arrangements require a
legislative and regulatory framework and
an effective administrative infrastructure for
their implementation and enforcement. China
emphasizes the need for an improved benefit
sharing mechanism to enhance development,
delivery and uptake of improved varieties: “The
lack of policies and regulations for protection
of intellectual properties related to genetic
resources, and the lack of effective mechanisms
of sharing responsibilities, rights and interests
between suppliers and users of the genetic
resources, have led to [a situation where] the
suppliers cannot benefit from the exploitation
and utilization of the genetic resources whereas
the users cannot get use right of the genetic
resources.”
Several international agreements address
this subject, such as the CBD, the International
Treaty on Plant Genetic Resources for Food
and Agriculture (which mainly deals with nontree agricultural crops) and more recently the
Nagoya Protocol on Access to Genetic Resources
and the Fair and Equitable Sharing of Benefits
Arising from their Utilization, adopted by the
CBD in 2010. Most countries are signatories to
these agreements. Benefit sharing may also be
conducted under provisions of patent regulations
where applicable. Most countries report that they
have patent laws in place; however, they are not
commonly applied to FGR.
Benefit sharing may involve profit sharing,
access to any improved materials, technology
exchange, contribution to in situ conservation
projects or assistance with other programmes.
However, not all agreements adequately
protect the rights of both parties in commercial
arrangements. To protect the rights of
germplasm owners and suppliers (including
countries, provinces, companies, communities
and traditional owners), strict and legally binding
Material Transfer Agreements (MTAs) should
govern all exchanges both between and within
countries. The Canadian province of British
Columbia, for example, has an MTA governing
transfer of seed and breeding material; it ensures
that ownership or custodianship is recognized and
confers limited use rights, for example for seed
production. ICRAF uses a standard agreement for
the collection of tree germplasm; however, apart
from these examples no standard MTA is used.
Papua New Guinea has a history of exchanging
and supplying seeds of indigenous and exotic
tree species, both for research and commercial
purposes. However, while FGR are recognized
under the CBD as the property of sovereign
nations, in Papua New Guinea they are considered
under the Constitution to be the property of
customary landowners, who own 97 percent
of the land. These landowners are increasingly
preventing access to FGR for research purposes,
as they seek additional benefits through
mechanisms such as an MTA with the Secretariat
of the Pacific Community and others.
Magnitude of germplasm transfer
Country reports indicate that germplasm transfer
within countries is predominantly for commercial
purposes, whereas international transfer is more
often for research (Figure 15.2).
By region, Europe reports by far the highest
availability of germplasm for within-country
transfers for commercial purposes, at 87 percent
of the global total. This suggests that European
203
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
FIGURE 15.2
Purposes of germplasm transfer reported by
countries
%
100
75
50
25
0
International
transfers
Commercial
National
transfers
Research
countries have well developed internal markets
for germplasm and/or that they have an effective
system for documenting germplasm exchange,
incorporating the private sector.
Globally, countries report a total of 412
different species available for national and
international transfer, although this is likely to
be an underestimate as much exchange of tree
germplasm (especially for ornamentals such as
palms) occurs through the private sector and is
not necessarily recorded in official figures. Latin
America and the Caribbean reports an average
of 28 species available for transfer per country,
Asia 20 species per country, and North America
16 species per country. These data are consistent
with the focus on a small number of priority
species of high commercial value as noted earlier
in this report.
Countries report transferring seed and
vegetative material internationally (import
and export) for a total of 534 different species.
The African region reports the most species
transferred (398), followed by Europe (338).
204
Information management in
delivery and deployment of
germplasm
The production and transfer of germplasm for
research, breeding and propagation requires
excellent documentation for quality control,
source identification and the management
and monitoring of activities. An efficient,
comprehensive and integrated information
system is essential. Some countries note that
a lack of information on the availability and
performance of both non-improved and improved
germplasm held by NTSCs and similar institutions
is a significant barrier to wider deployment of
germplasm held in collections.
Detailed, accurate information is required
for certification of forest reproductive material,
for example under the OECD Forest Seed and
Plant Scheme and EU Directive 1999/105/EC (see
above). Information requirements of different
schemes vary, and may include, for example,
seed source and origin, provenance, region,
provenance region/genecological/seed-transfer
zone, performance with respect to desirable
traits, number of parents, amount of variability
represented, germination percentage and
adherence to standards.
Adequate documentation allows purchasers
and users to verify that the material they acquire
is true to type, fit for purpose, and putatively
well adapted for the conditions into which it will
be planted. Proper documentation also permits
genetic material to be tracked, allowing:
• collation of data on performance of
improved varieties for evaluation and
research;
• use of plants as a source of reproductive
material of known origin at a later date
– a particularly important feature for
conservation plantings;
• assessment of any impacts on native
vegetation at the host site.
The adequacy of documentation varies among
countries and regions. For example, Germany’s
Act on Forest Reproductive Material requires the
German states (Länder) to register and maintain
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
databases of seed stands, orchards and clones,
thereby providing both a degree of oversight
and ease of access to site-appropriate, approved
sources of germplasm. Poland has an extensive
information management system documenting
forest reproductive materials around the
country. The Islamic Republic of Iran notes that
safe storage of data is vital for maintaining and
sharing germplasm; the country has a dedicated
unit for data management and for dealing with
germplasm requests. Some Asian countries gather
and disseminate FGR data and information
through the websites and portals of relevant
government agencies.
On the other hand, a number of other countries
identify their documentation as inadequate and
recognize the need to improve their information
management systems in this area. In developing
countries where resources are scarce and much
of the germplasm production and exchange is
informal, documentation of forest germplasm is
largely non-existent.
Some countries report that they have information
on public-sector breeding, delivery and deployment
but lack information on the activities in the private
sector, indicating a need for integrated and
harmonized documentation and data gathering
activities between the sectors. European countries,
under EU Directive 1999/105/EC, have developed
a regional system to document the origins and
track the movement of FGR – an example of a
harmonized, integrated system that facilitates
international and regional transfer of germplasm.
deployment programmes, from DANIDA,
Australia’s CSIRO and ACIAR, and the
International Tropical Timber Organization
(ITTO); some programmes date back to the
1960s.
• Guatemala’s Forest Guild and Guate Group
received technical assistance from Camcore
in establishing seed orchards for several
Pinus species.
• The United Republic of Tanzania has
received assistance from the Gatsby
Charitable Foundation in improving
germplasm for plantations through cloning,
with Camcore as a partner.
Unfortunately, in many countries, interventions
undertaken internationally to improve the
exchange of forest reproductive material are
likely to have only a limited impact on the material
available for smallholders to plant. In developing
countries, formal suppliers are able to provide
only a small proportion of the material cultivated
by smallholders, and most farmers indicate lack of
access to germplasm as a major constraint. There
is a need to rethink the operational means by
which tree germplasm reaches smallholders.
International assistance
Many developing countries have received
assistance from international partners, especially
ICRAF, in building up their production, delivery
and deployment systems. Other examples include
the following.
• The Mindanao Tree Seed Centre in the
Philippines was established with asssistance
from the Australian Overseas Aid
Programme.
• Thailand has received assistance in breeding
programmes, feeding into delivery and
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STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Chapter 16
Institutional framework for
conservation and management of
forest genetic resources
The institutional framework for FGR refers
to the national and international forest
programmes, policies and legislation governing
FGR development, conservation, management
and use; the structures supporting education,
research and awareness raising; and the networks,
agreements and other mechanisms for promoting
and supporting collaboration at the national,
regional and global levels.
National institutions dealing with
forest genetic resources
Many different institutions participate in FGR
management, both public and private.
Public responsibility for FGR and related policy
falls under different levels of government in
different countries. FGR may be integrated in
national forest programmes and managed by
national government institutions such as the
Ministry of Agriculture, Forests or Environment.
If the country has a decentralized administration,
FGR conservation may be managed under
different regions or states.
Most countries have institutions working for
the conservation, management and use of forest
genetic resources. Their work may include applied
research on fast-growing forest tree species,
forest seed testing, registration and control of
forest reproductive materials, sustainable use of
forest resources, in situ and ex situ conservation,
tree improvement, use of improved materials and
experimental stations. The national department
responsible for forests is often the institution
most actively involved in the conservation and use
of FGR. Some collaboration usually takes place
among different institutions, but a coordination
arrangement for FGR-related activities is often
lacking.
Especially in developed countries, institutions
actively engaged in FGR conservation and
management may be numerous, including
universities and colleges, federal and provincial
departments, research institutes, NGOs and tree
improvement councils and programmes.
Regional
governments
are
generally
responsible for managing forests within their
boundaries. Under this mandate they conduct
field and laboratory work. Industry is also often
involved in field and laboratory work concerning
FGR (for example, research related to biodiversity
and ecosystem health). Some jurisdictions have
tree improvement councils or cooperatives that
are responsible for managing and ensuring the
sustainability of FGR, and these groups often
support or engage in field and laboratory work.
Over the past decade, many countries have
successfully formulated national strategies
and programmes for FGR conservation and
management and incorporated FGR protection
into national action plans – from the banning
of harvesting in natural forests to the
establishment and protection of nature reserves
of key species. These are covered in detail in
Chapter 10. However, most countries have
no national programme for FGR. National
forest programmes have general measures for
conserving forest ecosystems, but in most cases
there are no specific provisions concerning FGR.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Legal and policy framework
Many countries have no specific laws on genetic
resources or have outdated FGR-related policies
and management tools. Some have legislative
or policy provisions relevant to, if not specific
for, forest genetic resources. The objectives are
mainly the conservation and protection of the
national forests, and less often the national
forest genetic resources. Most countries have
no legal framework for FGR strategies, plans or
programmes, although there may be provisions in
other national legislation.
Education and training
Many universities around the world offer
bachelor’s degrees in forestry; some also award
advanced degrees (master’s degree or PhD).
Some colleges have recognized technical forestry
programmes that award a diploma upon
completion of a two- to four-year programme.
Few universities address FGR as an independent
discipline or consider it as a thesis subject for
graduate students. However, a number of
programmes in forestry or natural resource
management address forest genetic resources in
their courses.
Some universities, colleges, and institutes
offer specific courses in forest genetic resources
conservation. These may have field and/or
laboratory components and may be delivered
through extension programmes or as part of
certified academic programmes. Some smaller
countries note that their courses on FGR are
insufficient, but that many students study forest
genetics abroad and return after graduation to
work in forestry.
In some countries, the provincial or central
forestry authorities organize FGR training
workshops at different scales. Training on
relevant laws, regulations and policies has
increased understanding of the importance of
FGR and strongly promoted their protection and
sustainable use.
Nevertheless, there has been a general
worldwide decline in enrolment in forestry
programmes over the past several years.
208
Universities and colleges are examining new
ways to entice students into their programmes.
Some ideas are to rebrand and transform their
programmes (e.g. from timber-oriented forestry
to sustainable forest management), develop
new programmes (e.g. international forestry)
and establish new partnerships (e.g. with forest
industry and research organizations).
Research
As touched upon in previous chapters, FGR
research and development (including studies
on collection, evaluation, conservation and
sustainable use of FGR) is generally carried
out, often collaboratively, by forest research
institutes, universities, and companies or
institutions concerned with forest production and
management. Some countries, such as Indonesia,
have research and development programmes on
forest biotechnology and genetic resources under
the Ministry of Agriculture.
In some countries, special research projects
have been set up to catalogue and document
forest genetic resources, establish and network
with FGR conservation banks, assess forest
genetic diversity, develop information platforms
and share FGR information.
Many countries express a need for faster
development
of
scientific
infrastructure,
advanced methodologies, and modern laboratory
equipment; this is related to a need for improved
funding, as sustained and stable financial support
is often lacking.
China, a country with a vast territory and rich
FGR with high genetic diversity, identifies a need
for specialized FGR research and for a nationallevel institution to coordinate FGR collection for
research use and provide technical support to
government departments for the formulation of
relevant policies.
Raising public awareness and
communication
Many countries report a need to raise public
awareness on forest genetic resources, as
the general population is hardly aware of
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
their function and importance. For example,
Germany reports the results of a survey in which
many respondents agreed with the statement
“Biological diversity should be preserved and
passed on to our children and future generations”
but could not explain the meaning of the term
“biological diversity”; only 12 percent were
aware that biological diversity involves genetic
diversity within species.
Furthermore, much of the public is sceptical
about genetic engineering. In some developed
countries, genetic engineering and genetics
are often considered one and the same thing.
Genetics and all related terms often have negative
connotations. This bias greatly hinders promotion
of the importance of forest genetic resources.
Some countries report surveys showing that the
public and NGOs have the lowest awareness of
the roles and values of forest genetic resources,
while industry and government rate much higher.
Some large Asian countries note that with an
increase in education on forest ecology, public
awareness about forest protection has increased
continuously over the past ten years. Furthermore,
the International Year of Biodiversity, 2010,
probably increased public awareness about
biological diversity. However, around the world
the public generally lacks understanding of forest
management and more specifically the planting of
productive provenances and forest plant breeding.
Most countries do not have specific programmes
for creating awareness of forest genetic resources.
Institutions that conduct public relations about
forests, forestry and nature conservation do not
generally make a chief priority of promoting the
importance of forest genetic resources.
In some cases, public awareness concerning the
value of forests and the species within them is
enhanced through programmes and activities of
diverse groups including the federal government,
botanical gardens, small woodlot partnership
programmes, environmental NGOs and forest or
tree-specific conservation groups. Federal and
jurisdictional in situ conservation areas have
also raised public awareness of the forest and its
genetic resources.
Support to forest genetic
resources
Over the past decade, funding for forest genetic
resources has generally decreased slightly. In only
a few countries have budgets slightly increased
over the past ten years.
Little precise and reliable information is
available about budgets for forest genetic
resources because the institutions involved may
be subordinated to many different ministries.
However, the proportion of forestry budget
allocated to FGR is often not more than 1 percent.
It is also difficult to estimate the funding
apportioned to FGR research, since government
budget allocations may fall under a number of
different ministries and government agencies at
both the national and subnational levels; and
universities, colleges and various organizations
may also allocate part of their budgets to FGR
research. In developing countries the national
research budget for FGR is usually especially
poor, particularly where there is no national
research institute, university or school dealing
with forestry. These countries may occasionally
be involved in small-scale research carried out
by forestry schools or institutes abroad or in the
frame of small projects.
Since domestic funding is often limited,
capacity-building activities are sometimes carried
out through bilateral cooperation in the form of
project-based technical assistance programmes
and research grants from international agencies.
The projects normally include funding provisions
for training and postgraduate studies overseas.
International and regional
collaboration
The distribution of forest genetic resources does
not correspond to political borders, and this is an
important basis for cooperation and coordination
on issues related to FGR management. Many
drivers of change affecting FGR, including climate
change, also span political borders.
International
forestry
cooperation
and
exchange are developing rapidly. Many countries
collaborate with other countries and international
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 4
organizations to conserve their forest genetic
resources. Collaborative efforts also assist in
improving countries’ capacity to conserve FGR
and manage them sustainably. The main forms
of regional and international cooperation are
international networks, bilateral and multilateral
cooperation, and international conventions.
These activites are carried out at both regional
and global levels. Many countries, however, do
not have any specific budget for international
gene conservation purposes.
The paramount objective of international
forest policy is to halt further deforestation and
forest degradation, thus contributing to the
internationally agreed Millennium Development
Goals to protect the climate, conserve biological
diversity, combat desertification and alleviate
poverty, especially in rural regions. To this
end, over the past 35 years the number of
international, regional and national institutions,
mechanisms and discussion fora concerned with
forests and forest biological diversity has greatly
increased.
Through
membership
in
international
institutions such as FAO and IUFRO, countries and
institutions participate in global decisions and
guidance concerning conservation, management
and use of FGR and related research. In addition,
in recent years the Ministries of Agriculture
and Forestry in many European countries have
considerably reinforced their roles in international
forestry.
Although
many
collaborative
activities
in forestry do not have the direct purpose
of conserving forest genetic resources, they
safeguard forest habitats and indirectly contribute
to the conservation and sustainable use of FGR.
Many international agreements are relevant to
the sustainable management, use, development
and conservation of forest genetic resources,
addressing such issues as access, benefit sharing,
biosecurity, intellectual property rights and illegal
trade of natural resources.
International cooperation programmes and
information exchange efforts such as shared
databases, joint research and publications,
210
technical guidelines and germplasm exchange
also address forest genetic resources; some do
so indirectly by addressing stresses to the forest
(e.g. the United Nations Framework Convention
on Climate Change [UNFCCC]). International
movement of forest reproductive material is
discussed in detail in Chapter 15.
International and regional FGR networks
Many countries and institutions participate in
global or regional cooperation and exchange
networks related to forest genetic resources.
Network activities – including information
exchange, database development, sharing of
conservation strategies and seed exchange –
promote sharing of FGR information, help expand
research capacity, and help improve technical
standards. Networks that share data on outbreaks
of alien invasive forest pests, for example, can
help researchers and forest managers to develop
proactive responses to future outbreaks in other
regions. Some networks focus on particular
tree species, and some focus specifically on
conservation in situ or ex situ.
Networks may encourage more intensive
communication among the countries in a region.
Similarities among countries, for example in
forest tree species, ecosystems and sociocultural
environments, provide entry points for network
development. Some networks reach out to
existing regional organizations to gain support
from a broader range of stakeholders.
One example of an active network is the
European Forest Genetic Resources Programme
(EUFORGEN) (Box 16.1). Some developing Asian
countries report that they use international
networks to share information on the status
of research and development of forest genetic
resources and to gather relevant inputs for FGR
conservation and management.
In the past ten years some African countries
that had been members of FGR networks and
networking organizations – e.g. Camcore, the
Southern African Development Community
(SADC) Tree Seed Centres Network, IUFRO and
the Global Forest Information Service (GFIS) –
STATE OF FOREST GENE T IC RESOU RC ES CONSERVAT I ON A ND MA NAGEMEN T
Box 16.1
Example of an international FGR network:
the European Forest Genetic Resources Programme (EUFORGEN)
The overall goal of EUFORGEN is to promote the
conservation and appropriate use of forest genetic
resources as an integral part of sustainable forest
management in Europe. It was established in October
1994 as a pan-European implementation mechanism
of Strasbourg Resolution 2 (Conservation of forest
genetic resources) of the first Ministerial Conference
on the Protection of Forests in Europe (now known as
Forest Europe). EUFORGEN is funded by its member
countries and overseen by a steering committee
consisting of national coordinators from all member
countries. The work is coordinated by Bioversity
International in technical collaboration with FAO.
EUFORGEN brings together European experts to
exchange information and experiences and to develop
tools and methods for better management of forest
genetic resources. EUFORGEN has produced numerous
outputs such as genetic conservation strategies,
technical guidelines, distribution maps of European
forest trees, databases, and various publications and
reports.
EUFORGEN had a crucial role in the establishment
of the European Information System on Forest Genetic
Resources (EUFGIS), which provides georeferenced
and harmonized data on dynamic conservation
units of forest trees in Europe. Since its launch in
2010, the EUFGIS Portal has been maintained by
EUFORGEN. EUFORGEN working groups have used
the EUFGIS data, provided by national focal points,
to develop a pan-European genetic conservation
strategy for forest trees and a genetic monitoring
scheme for selected conservation units. Recently, other
EUFORGEN working groups have focused on the use
of forest reproductive material, policies relevant to the
conservation and use of FGR, and the implications of
climate change on FGR conservation.
have let their membership lapse. As these African
economies stabilize they may wish to consider
rejoining these networks and organizations for
the benefits they can provide to scientists working
on forest tree breeding, forest management, and
forest and gene conservation.
Not all networks are successful, however. Some
developing countries say they have not reaped
many benefits from the regional and subregional
networks in which they participate, except in some
cases where networking has assisted technology
development and information sharing. Where
networks are ineffective, the cause is often a
lack of coordination between the government
and the network; a realistic plan of action is
required and must be compatible with national
priorities. In addition, some governments lack
sufficient skilled human resources to participate
effectively in networks and benefit fully from
them. Inadequate conservation infrastructure
can also be an impediment to obtaining benefits
from networking.
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Part 5
NEEDS, CHALLENGES
AND REQUIRED
RESPONSES FOR
THE FUTURE
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
Chapter 17
Practices and technologies for
improved management of forest
genetic resources
The challenge of achieving food security for all
and environmental sustainability in the context of
the combined effects of climate change and the
increasing human pressure on forests is greater
now than it has ever been. More efficient use
and management of available forest resources
is therefore needed, especially in tropical and
less-developed countries, in order to meet the
growing demand for forest goods and services.
Managing FGR involves developing overall
strategies, applying specific methodologies,
developing and applying new technologies, and
coordinating local, national, regional and global
efforts.
FGR conservation and management considerations need to be better integrated into SFM
planning and practices, including undertaking an
inventory of FGR, prioritizing FGR for conservation
and management, characterizing their variability,
defining a management strategy for maintenance
of variability, integrating this into utilization
protocols and plans, monitoring the impacts
on variability, and adjusting the SFM regime as
appropriate. FGR value should be mentioned
explicitly in assessment of high conservation value
forests, and forest certification schemes need to
include requirements for effective conservation
and management of FGR.
The current growing concern about climate
change and its effects on ecosystems and on
performance of forest production systems
challenges stakeholders in FGR management to
better understand forest species’ mechanisms for
adaptation to current and future climate changes.
Genetic diversity is needed in order to ensure that
species can adapt, as well as to allow for artificial
selection and breeding for improved productivity.
Thus genetic diversity, including diversity among
species, is the key to the resilience of forest
ecosystems and the adaptation of forest species
to climate change. Countries should therefore
endeavour to support climate change adaptation
and mitigation through proper management and
use of FGR.
Marginal and/or range-limit populations may
be vital for tree species’ adaptation to the novel
environmental extremes that are expected to
occur as a result of rapid climatic change. It is
necessary to understand the dynamics of marginal
populations through adequate examination
of adaptive genetic variation in quantitative
traits. Furthermore, conservation efforts in the
current context must consider the range of
future climatic projections, and appropriate
conservation measures must be developed using
the principles of risk management. Modelling of
species distribution dynamics needs to account
for changes in species’ distribution areas and in
those of their associated environment correlates
(e.g. pollinators) and also the possible inluences
of interactions with other plant or animal species.
Monitoring
Monitoring of forest genetic resources and biotic
stressors (for example, invasive alien species) can
support FGR conservation and management at the
regional level. It is highly beneicial for developing
effective long-term strategies not only for
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 5
conserving the resources, but also for minimizing
the impacts of the stressors and/or for developing
scale-appropriate mitigation strategies. Research
priorities identified by countries for improving the
monitoring of genetic erosion and for assessing
species’ vulnerability include:
• assessment and monitoring of particular
species’ genetic diversity and their adaptive
potential in regard to various stressors;
• identification of the resistance of native
tree species to high-impact stressors.
Monitoring forest biological diversity and
managing FGR require reliable information on
the status and trends of these resources. There
are no common standard methods for measuring
changes in the status of FGR in relation to
sustainable forest management as undertaken in
most countries. Parameters commonly included in
national and global forest resource assessments,
such as forest area, species occurrence and
richness and forest fragmentation, do not,
on their own, provide information on FGR.
Adequate and commonly agreed indicators are
needed and should be developed and integrated
into the national forest assessment policies and
monitoring tools.
TABLE 17.1
Potential local- to global-scale operational indicators of forest genetic diversity, with verifiers
Operational indicator
Verifiable indicator
Verifier (direct or proxy)
Primary scale
of measure
and indicator
State–Pressure
Trends in species and population
distribution pattern of selected species
Trends in population condition
1
Number of species with known
distribution for which allelic
diversity is declining
1
Number of species with known
distribution for which distribution is
declining
2
Natural distribution range
2
Geographic and climatic range
3
Distribution pattern within
the natural distribution range
where appropriate
3
Geographic, climatic and
ecogeographic distribution of
populations
4
Representation within the
natural range
4
Number of populations relative
to their potential genecological
distribution
5
Number of populations, their
area and density (abundance)
5
Area and density of populations
6
Demographic condition of
selected populations (diversity
in adaptive traits/genes)
6.1
6.2
6.3
6.4
6.5
6.6
Age/size class distribution
Number of reproducing trees
Abundance of regeneration
Environmental heterogeneity
Number of filled seeds
Percentage of germination
7
Genetic condition of selected
populations (population
genetic structure where
appropriate)
7.1
7.2
7.3
7.4
7.5
Effective population size
Allelic richness
Outcrossing/inbreeding rate
Spatial genetic structure
Hybridization/introgression
8
Hectares planted by species/
provenance either locally or as
an exotic
8
Hectares planted by species/
provenance either locally or as an
exotic
9
Profit from breeding vs. loss
from ill-adapted plantations
9.1 Seed source performance (growth and
survival)
9.2 Realized genetic gain and profit
National
Regional
Global
Local
Beneit
Trends in plantation performance of
selected species
216
Local
National
Regional
Global
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
Relevant indicators for monitoring genetic
diversity have lagged in development perhaps
more than any other type of biodiversity indicator
(Laikre et al., 2010); recognizing this, the CBD’s
Strategic Plan for Biodiversity 2011−2020
allows for improved coverage. Nevertheless,
identiication and implementation of indicators
of genetic diversity, including tree genetic
diversity, remains a major challenge.
The dificulties in deining sound and
realistic indicators arise from the fact that they
should be policy relevant, scientiically sound,
understandable, feasible to obtain and sensitive
to changes over time. The use of a hierarchical
approach, introducing the proportion of coverage
as a measure of progress on selected indicators at
different levels, offers a possibility for general use
within the state-pressure-beneit-response (SPBR)
framework. FAO has recently developed a set of
indicators that span geographic scales from local
to global and include state, pressure, beneit and
response measures (Table 17.1) (Graudal et al.,
2014).
In
situ
conservation
often
comprises
maintenance of ecosystem functions and
species interactions rather than conservation of
TABLE 17.1 cont.
Operational indicator
Verifiable indicator
Verifier (direct or proxy)
Primary scale
of measure
and indicator
Response−Benefit
Trends in knowledge of genetic diversity
of species
Trends in education and awareness
Trends in sustainable use of tree genetic
resources
Trends in genetic conservation
10
Increase in number of species
that are described for which
distribution and/or genetic
parameters are known
10
Increase in number of species that are
described for which distribution and/
or genetic parameters are known
11
Number of species with
mapped genecological
variation
11.1 Increase in number of articles on
genetic diversity by species
11.2 Number of species with mapped
genecological variation
12
Among-population genetic
diversity (of selected species)
12
Parameters of genetic differentiation
among populations
13
Change in number of tree
geneticists and tree breeders
13
Number of university courses or
training courses offered in forest
genetics related subjects
14
Existence of networks
14
FGR networks (function/operation)
15
Use in national forest
institutions and programmes
15
Species presence in national forest
institutions and/or programmes
16
Number of tree species for
which regulation of use of
forest reproductive material
exists
16
Number of tree species for
which regulation of use of forest
reproductive material exists
17
Number and type of improved
seed sources traded/
exchanged (status of genetic
improvement)
17.1 Number and type of improved seed
sources traded/exchanged
17.2 Seed source performance (growth and
survival)
18
Guidelines/regulations for
matching seed source and
planting site
18.1 Certification scheme in place
18.2 Use of adapted seed sources
19
Guidelines/regulations for
composition and harvest
of seed sources (number of
mother trees)
19
Use of diverse seed sources
20
Number of tree species directly
targeted in conservation
programmes
20
Number of tree species directly
targeted in conservation programmes
Local
National
Regional
Global
National
Regional
Global
National
Regional
Global
National
Regional
Global
Source: Graudal et al., 2014.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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individual tree species. In addition, forests have
a number of native trees and shrubs that may be
of minor interest to forest managers, but may be
highly valuable in terms of genetic resources and
future use. It is therefore important that forest
and tree management include indicators related
to sustainable FGR management in regular
monitoring protocols.
In situ conservation
In the current context of increasing pressure
on forest land and forest resources, primary
forests and protected areas remain refuges for
threatened FGR. An important proportion of
wild and/or endemic plants occur only in primary
forests and protected forest areas. The genetic
structure of species natural populations is best
TABLE 17.2
Some constraints, needs, priorities and opportunities identified by countries for in situ FGR
conservation and management
Area
Land use pressures,
encroachment and
landownership
Constraints
High exploitation, land clearing
and deforestation (from poverty,
expansion of agriculture, market
demand for timber and fuel, illegal
harvesting)
Needs and priorities
Reduce pressure on in situ
conservation areas
Improve production forestry (identify
areas for forest development, improve
germplasm quality)
Encroachment on in situ conservation
areas
Improve natural resource
management and forest restoration
Private or customary ownership of
land, with limited control (Cyprus,
Solomon Islands), small fragmented
parcels (Japan), inability to ensure
long-term conservation (Finland)
Resourcing and
capacity
Lack of expertise and capacity
Insufficient funds and resources
for training, survey, identification,
management and monitoring
Opportunities
Develop renewable energy sources
alternative to fuelwood from natural
forests (including energy plantations)
Involve private sector in in situ and
FGR reserve systems, funded through
environmental service payments or
government assistance; incorporate
these lands into public estate through
purchase
Increased, more secure, long-term
funding
Capacity building – including
education, training, resources
Involve and better coordinate efforts
of universities, training schools,
NGOs and others in conduct of FGR
inventories
Increase involvement of donors and
NGOs in in situ conservation
Enhance information exchange
and participation in regional
and international networks and
collaborations
Public awareness
and support
Resistance to expansion of protected
areas and conservation reserves
(community concern over loss of use
and access and increased human–
wildlife conflicts)
Loss of traditional knowledge
and beliefs leading to decrease in
valuing of traditional species, forest
management and conservation
Low or no community participation
and lack of benefit sharing
arrangements, resulting in conflict
over forest resources
Lack of public awareness, interest
and/or support
Lack of political awareness and/or
support
Lack of alternative livelihood options
for local populations
218
Sustainable livelihood security, access
and benefit sharing arrangements
Expanded use to increase valuation by
community of indigenous FGR
Promotion of indigenous and
traditional species in research,
conservation and local use
Payments to compensate traditional
forest users where access to forest
revenues is denied; incentives for
stewardship of FGR
Extension, education and awareness
raising for forest owners, users, youth
and general public
Communication strategy, including
media campaigns
Greater involvement of local
communities in management
Sustainable and equitable use of
natural resources; benefit sharing for
local communities
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
conserved in these forests. Natural processes
involved in the dynamics of FGR are better
assessed and understood in protected natural
forests, which remain the best laboratories for
studying species ecology and biology.
Almost all country reports identify constraints,
needs and priorities, and opportunities for in
situ conservation of FGR. Those most commonly
identified by countries are listed in Table 17.2.
The following summarizes the main constraints
identified by countries:
• high levels of land clearing and
deforestation (resulting from a variety
of causes including poverty arising from
TABLE 17.2 cont.
Area
Policy, legislation
and enforcement
Constraints
Lack of coordination among policies,
laws, government departments and
sectors
Insufficient policy support
Lack of adequate legal framework and
legislative protection for designated
areas; inadequate enforcement
Needs and priorities
Strengthened legal framework for in
situ FGR conservation
More effective enforcement of
regulations and laws
Poverty alleviation and employment
creation strategies
Lack of knowledge of relevant
policies, laws and regulations by
stakeholders (including those charged
with law enforcement)
Opportunities
Review, revise or develop supportive
policy and legislation for protected
areas and for FGR currently on
unprotected lands
Provide legislative protection for
threatened, ecological keystone and
culturally important species, including
iconic trees and populations
Improve land-use planning and
policies and tenure systems
Lack of policy on how to increase
benefits of trees on farms and circa
situm conservation
Concentration on charismatic,
priority species at expense of other
indigenous species; undue emphasis
on a few rare species
Technical and
operational issues
Lack of FGR strategy, coordinated
national plans and integrated
approaches linking in situ and ex situ
strategies
Underrepresentation of important
ecosystems in protected area
networks (e.g. lowland primary forests
in Indonesia, plains forests in Nepal,
lowland dipterocarp forests in the
Philippines)
Lack of inventory and knowledge
on natural distributions and
genecological zones, genetic resources
and processes
Management issues of in situ
conservation areas, e.g. invasive
species, fire management
Close-to-nature management
approaches in strictly protected areas
limiting opportunities to conserve
light-demanding or disturbancedependent species
Climate change issues: reduced
regeneration of conserved species,
range shifts of species and vegetation
communities
Better planning
Guidelines for designation and
management of FGR conservation
reserves and for preparation of in situ
conservation strategies and action
plans
Rehabilitation and restoration of
degraded ecosystems and recovery of
threatened species
Use gap analysis to identify priority
areas for FGR conservation
Translate research findings into
practical conservation plans and
actions
Assess existing forest reserves
for synergetic use as in situ FGR
conservation units for main tree
species
Assessment and monitoring of
existing FGR conserved in situ
Protection of high-priority species
(including rare species) and
endangered populations
Conservation of marginal-range
populations
Better management of in situ sites
(including silviculture, regeneration,
protected areas)
Better knowledge sharing,
coordination and networks at
national level (government agencies,
researchers, park managers, forest and
local authorities, farmers)
Improved protected area network
connectivity and reduced
fragmentation
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 5
population growth, change in land use
owing to expansion of agriculture, and
increasing market demand for timber and
other wood products);
• climate change, leading to reduced
regeneration of conserved species and
range shifts of species and vegetation
communities because of shifts in climatic
zones;
• lack of knowledge on genetic diversity,
genetic processes and genecological zones;
• lack of or insufficient integration of FGR
issues into current wider national policies
and laws;
• lack of knowledge of relevant policies,
laws and regulations on the part of
stakeholders, including those charged with
law enforcement.
Adequate in situ conservation measures
are needed to preserve the natural growing
conditions of the tree species in order to study and
better understand their evolutionary process and
adaptation to changes. Information from in situ
conservation activities for marginal and/or rangelimit populations will be essential in providing
options for adaptation to climate change.
Enhancing the role of protected areas for
in situ FGR conservation
In many countries protected areas contribute to
the conservation of viable forest tree populations
of diverse species and of representative ecosystem
samples, as well as maintaining vital ecosystem
services. Their primary objective is ecosystem and
biodiversity conservation; they serve as a refuge
for forest species that are unable to survive in
intensely managed landscapes. However, the
status and role of protected areas differ among
countries, as does their impact on conservation.
The extent of protected areas has been
increasing over the past decades as a result of
national and international efforts to conserve
biodiversity. Protection is implemented under
many management types and categories
including strict nature reserves, national parks,
habitat or species management areas, protected
220
landscapes and protected areas with sustainable
use of natural resources. National programmes
for sustainable use and management of FGR
should therefore take the contribution of
protected areas into account, even if most of
them were initially designed for other purposes
such as wildlife protection, recreation and various
ecosystem services.
Protected areas also have an important role
in enhancing scientific knowledge on FGR. A
substantial proportion of wild and/or endemic
plants occur only in primary forests and protected
forest areas. Only in those forest ecosystems is the
natural population’s genetic structure conserved.
Natural processes involved in the dynamics of FGR
are best assessed and understood in protected
natural forests; they remain the best laboratories
for studying species’ ecology and biology. The
contribution of primary forests and protected
areas to the development of knowledge on plant
species therefore needs to be promoted along
with their contribution to FGR conservation.
On-farm management of FGR
On-farm management of trees, including
agroforestry systems, contributes to in situ
conservation of FGR, particularly for domesticated
or semi-domesticated tree species (Dawson
et al., 2013). The agroforestry parkland, for
example, a traditional land-use system in West
Africa, has been shown to contribute to onfarm conservation of species diversity (Nikiema,
2005). Many priority species identified in country
reports from semi-arid zones are trees growing
on farmlands, often in agroforestry systems.
Most of them are indigenous species that farmers
have managed traditionally for centuries. Tree
diversity in farmland varies from a few species in
some countries to more than 100 in some others.
Some of these species are semi-domesticated
species that occur only in agroforestry systems
(e.g. Acacia senegal and Vitellaria paradoxa).
Agroforestry systems must therefore be managed
sustainably to conserve the genetic resources of
the species. In situ conservation efforts should
also include conservation of natural populations
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
of the species and their wild relatives in order to
preserve the variation needed for adaptation to
future threats or to improve production.
Multiple-use forest management and
ecosystem approach
Naturally regenerated forests across the tropics
provide a wide range of products, ecosystem
services and social and economic opportunities
and potentially can be managed to meet multiple
objectives (Sabogal et al., 2013). Multiple-use
forest management refers to forest management
that combines objectives such as production
of wood, habitat for wildlife, soil and water
protection, recreation, and the supply of a
range of NWFPs (e.g. food, fodder, medicines).
This approach enhances sustainable forest
management by taking into consideration the
concerns of the stakeholders.
The ecosystem approach is a way to manage
entire ecosystems in a holistic manner without
excluding other management and conservation
approaches such as area-based management
tools and single-species conservation practices.
Ideally all these approaches should be integrated,
through regional networks when appropriate.
•
•
•
•
•
•
•
of the forest gene bank concept, whereby
genetically diverse material of priority
economic and threatened species is
planted as a mixture in one (or more)
locations;
use of these areas to raise the living
standards of local people;
ecotourism to generate income from nonconsumptive uses of in situ conservation areas;
enrichment planting and regeneration
of high-priority and threatened FGR in
protected areas, including restoration of
degraded areas;
restoration of the connectivity of protected
forest fragments;
promotion of on-farm or circa situm
conservation of priority FGR, including fruittrees and their wild relatives;
payments for environmental services such as
carbon sequestration (e.g. through REDD+),
watershed protection and biodiversity (e.g.
through bioprospecting licences);
new employment opportunities from
expanded markets for forest products
(including NWFPs such as specialized organic
forest products).
Species and thematic networks
Research on in situ conservation
Regional collaboration through species or
thematic networks should play an important part
in implementation of in situ FGR conservation
strategies and monitoring of progress. While
addressing in situ FGR conservation, such
collaborations should also consider the use of the
ecosystem approach, different forest and tree
management types and different levels of genetic
conservation.
Countries identify a number of specific research
priorities for in situ conservation:
• increased knowledge and monitoring of
genetic variation and its distribution in
priority FGR, including threatened species,
breeding systems and levels of outcrossing,
gene lows between conserved populations
and introduced/planted materials, and
population viability;
• inventory and GIS mapping of FGR in
protected areas (including to identify
centres of diversity) and in non-protected
areas;
• more knowledge of effective in situ
conservation techniques including selection,
establishment, monitoring, restoration and
rehabilitation and management;
• prioritization of species;
Opportunities for adding value to in situ
conservation
Countries identify the following opportunities
for adding value to in situ conservation and/
or improving the functionality of natural forest
ecosystems:
• increased use of these areas as seed stands
for priority FGR, including development
221
THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 5
• research on species importance in
maintaining ecosystem function and
services, as a focus for in situ conservation;
• research on autoecology (the ecology of
individual species) and species genetic
diversity, including monitoring, to inform
and assess effectiveness of FGR conservation
strategies and activities;
• development of low-cost technologies for
better control of invasive species;
• identification of methods to improve
participatory forest and FGR management,
including to reduce deforestation;
• research into socio-economic aspects of in
situ conservation;
• study of the impact of predicted climate
change on the effectiveness of in situ
conservation reserves.
Ex situ conservation
The genetic diversity of forest trees and shrubs,
both adaptive and neutral (i.e. in which variants
have no direct effect on fitness; see Holderegger,
Kamm and Gugerli, 2006), can be maintained
through in situ conservation methods based
on individual
tree
species
distribution
ranges. Although it is regarded as the most
appropriate, sustainable and cost effective way
of conserving FGR, in situ conservation can be
insufficient, difficult or impossible for some
species or species populations under threat
because of exotic pests or disease outbreaks,
extreme environmental conditions due to
climatic changes or loss of habitat. For these
species or particular populations of species, ex
situ conservation is an essential complementary
conservation tool.
Ex situ conservation may involve management
of seed banks, gene banks or field collections.
However, countries often lack adequate policies
and the necessary means to address the needs of
ex situ FGR conservation. Because of the high cost
involved in ex situ conservation programmes and
activities, priority should be given to populations
of endangered species or taxa that are likely to
222
become extinct. In some cases global or regional
initiatives are needed for efficiency.
Needs for ensuring adequate contribution of
ex situ conservation to overall FGR conservation
include:
• good access to ex situ conservation data and
information on FGR;
• improved capacity for ex situ conservation
at all levels (national, regional, global);
• appropriate, efficient and economically
accessible technologies for the conservation
of seed, especially recalcitrant seed;
• expansion of the scientific knowledge base
on tree seed physiology and conservation
techniques.
With many countries reporting negative trends
of overexploitation, land use changes and climate
change effects, and consequently increasing
loss of inter- and intraspecific diversity, ex situ
conservation is warranted as a component of
conservation strategies at the national, regional
and global levels.
Domestication, breeding and
improvement
Tree domestication and improvement can
substantially
contribute
to
sustainable
development through diversification in food
and other commodities that are important to
local communities and national economies,
such as timber and medicinal plants (used by a
large portion of the population in developing
countries). Free grazing is still a common practice
in many developing countries, and forests are
often an essential source of fodder. These various
resources are still harvested from wild plants in
forest lands which in some cases are threatened
by overexploitation. Domestication of such
plants will improve the supply of the targeted
products while reducing the vulnerability
of their genetic resources. Many countries,
particularly in the tropics, underline the need to
develop domestication programmes to improve
the supply of various forest products, including
NWFPs.
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
Tree improvement activities have mostly
been limited to a small number of economically
important, widely planted tree species, because
of financial constraints and because of the
specific biological characteristics of trees;
most trees are long-lived perennial species
with long rotations (usually more than ten
years except for pulpwood and biofuels), long
regeneration cycles and late sexual maturity.
Because of these characteristics, improvement
and breeding research in tree species often
requires many years (more than for other crops)
and considerable resources (trained personnel,
finances, land and laboratories). Accordingly
there is a need to develop and promote the
use of new technologies – e.g. biotechnology,
genomics and micropropagation – to accelerate
the tree improvement process and help unlock
the huge potential of planted forest trees.
These new technologies have proved useful
for understanding forest ecosystem dynamics
and species genetic diversity and processes.
They can provide options for practical measures
for sustainable conservation, management,
restoration and rehabilitation, especially when
there is sufficient scientific evidence of the
relation between phenotype and genotype.
However, as funding and interest have switched
to molecular approaches, many countries have
abandoned progeny and provenance trials that
had been established for many species. Existing
but dispersed data from these trials should be
assembled, maintained and evaluated for their
potential to inform seed zone delineation, plans
for assisting gene low in response to climate
change, and identiication of propagation
material for restoration and conservation of highvalue populations.
Selection and breeding of trees to
respond to climate change
Traditional breeding programmes will need to be
modiied to consider plasticity and adaptation
to increased drought, a substantial change from
current practice. Climate change related traits
need to be included in selection criteria; this is
still rarely done worldwide.
Provenance trials that have been established at
multiple locations using germplasm sourced from
a variety of ecological conditions demonstrate
that variation in adaptive traits is almost always
present within tree species. Not only is genetic
diversity in important adaptive traits expressed
across regions and provenances, but it is also
abundant within populations, reinforcing an
optimistic view that climate change challenges
may be met by standing genetic variation in such
species (Hoffmann and Sgro, 2011).
However, many provenance trials were
established before the response to largescale anthropogenic environmental change
was considered an important research issue,
so trials often have not measured the most
important traits from the perspective of
adaptation to climate change. Nevertheless,
these older multilocational trials provide
insight into the performance of provenances
from different climatic regions and make it
possible to identify sources of locally adapted
material. Survival and growth are considered
good proxies for itness (e.g. Ouedraogo et
al., 2012). New trials speciically established to
assess explicit responses to climate change are
being established in a number of countries,
for example under the Treebreedex project in
Europe (http://treebreedex.eu).
Some important traits needed for adaptation
to different climatic conditions but not often
considered in breeding programmes are pest
resistance, drought resistance, ire resistance or
tolerance, cyclone resistance, salt tolerance and
phenotypic plasticity.
Germplasm delivery and
deployment
Countries report that large plantation areas
are being established to serve many purposes,
including the production of timber, biofuel
and ibre and the provision of environmental
services such as soil and water management
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 5
and reclamation of degraded land. However,
many countries lack adequate forest seed supply
systems and therefore face difficulties in getting
the quantities and quality of forest reproductive
material needed to implement their plantation
programmes. Collaboration among tree seed
centres should be enhanced to encourage
development and use of common seed quality
standards, to facilitate forest reproductive
material exchange within regions and to support
national afforestation programmes. In this
regard the International Seed Testing Association
(ISTA) standards for seed germination are a
widely used reference for the international tree
seed market.
Seed is the most available form of germplasm,
and distribution and deployment tend to be
skewed towards species that produce orthodox
seed, since they can be transferred most
conveniently and reliably. Since other species may
also be useful or require conservation, there is a
need for further research and development on
storage, propagation and transfer techniques for
species with recalcitrant or short-lived orthodox
seeds.
In some countries, public agricultural agencies
have a longstanding involvement in ex situ
conservation and in production, storage and
distribution of forest germplasm (particularly in
relation to food- or crop-bearing trees. Greater
cooperation and harmonization of effort
between agricultural and forest agencies might
prove beneficial in this respect, especially in
countries where resources for FGR conservation
are limited.
Governments may offer incentives to private
companies for the production of high-quality
germplasm and planting materials, particularly
for activities consistent with national goals, e.g.
for conservation of forests, FGR and biodiversity.
The global movement of forest reproductive
material should be facilitated. Priority matters
reported by countries in the area of germplasm
delivery and deployment, including identified
strategic priorities, include:
224
• coordination of public and private sector
activities;
• possible centralized organization to
coordinate exchange of germplasm and
related data collection;
• identification of opportunities for
coordinating germplasm collection, storage
and deployment with ex situ conservation
activities, e.g. through tree seed centres;
• clear benefit sharing arrangements and
use rights to promote production and
deployment of improved trees;
• incentives for adoption and use of improved
varieties;
• increased access to improved varieties for
farmers, rural communities and others in
the informal sector;
• increased capacity for producing adequate
quantities of improved planting materials to
meet demand;
• closer alignment of delivery and
deployment with the needs of communities
and market demand through better
consultation, coordination, participation
of communities setting effective
priorities, development of markets (where
appropriate) and more effective response to
market signals;
• development and expanded use of
consistent standards for collection and
storage of germplasm for exchange,
distribution and deployment at national,
regional and international levels, and
their harmonization with existing
programmes (e.g. the OECD Forest Seed
and Plant Scheme [OECD, 2013] and EU
Directive 1999/105/EC [EU, 1999]), including
promotion of common language and
terminology;
• promotion of production and exchange
under preferred schemes and guidelines
with appropriate rewards, while retaining
the vigour of informal germplasm
production and exchange systems in
developing countries;
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
• development and promotion of appropriate
certification systems;
• use of improved materials in informal
germplasm exchange and production
systems;
• more effective integration of databases,
including accessible information about
deployment in the private sector;
• promotion of improved fuelwood
plantations to provide carbon-neutral
energy and reduce degradation of natural
forests and FGR, particularly for countries
where wood is currently a major energy
source;
• maintenance of genetic variability in the
distribution and deployment process.
Assisted migration to accelerate
adaptation to climate change
Trees and tree populations are amenable to
“facilitated translocation” or “assisted migration”
which involves the movement (by people) of
reproductive materials (seeds, seedlings and
vegetative parts) from existing ranges to sites
expected to experience analogous environmental
conditions in the future (Guariguata et al., 2008;
McLachlan, Hellmann and Schwartz, 2007). The
movement could be latitudinal or altitudinal.
The objective of such intentional movement is
to reduce climate change-related extinction risks
(Heller and Zavaleta, 2009; Marris, 2009; Millar,
Stephenson and Stephens, 2007). Species or
populations that are unable to migrate to new
locations or adapt through natural selection can
be intentionally moved to a region where stresses
are less severe.
Assisted migration can include translocation
over long distances (assisted long-distance
migration), translocation just beyond the range
limit (assisted range expansion) and translocation
of genotypes within the existing range (assisted
population migration) (Alfaro et al., 2014;
Winder, Nelson and Beardmore, 2011). Under
certain interpretations, assisted migration
could include the introduction of new species
to maintain ecological services, such as wood
production and carbon sequestration. Some
guidelines for reforesting harvested sites require
the use of seeds from neighbouring sources that
are already adapted for expected future climates
(e.g. seed from sources south of the area to be
planted, in Northern Hemisphere forests). In
practice, implementation of such guidelines could
be a gradual form of assisted migration.
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NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
Chapter 18
Political and institutional
recommendations
Forest genetic resources offer major opportunities
for humankind to cope with important challenges
such as climate change and the increasing
demand for food, energy, wood products and
environmental services from forests. To realize
their value fully, countries require political
commitment, effective institutions and relevant
policies and legislation in order to respond
to pressing and increasingly varied needs in
conservation and FGR management. Staffing
and capacity must be strengthened through FGR
education, research, and training. Long-term
stable funding support mechanisms should be
established, including for FGR research.
Information on the status, trends and
characteristics of FGR is needed in order to identify
priorities for actions for their sustainable use
and conservation as well as for the development
of tree domestication and improvement
programmes. Awareness building at all levels is a
key prerequisite for mobilizing popular support
and international collaboration to improve the
conservation and management status of FGR.
Appropriate advocacy tools need to be developed
and used to ensure effective communication and
information sharing related to sustainable FGR
management and use.
In the context of scarce resources and a great
risk of duplication of effort, efforts should be
made to promote collaboration, partnership
and coordination at the national, regional and
international levels and to mobilize the funding
required to ensure that the major needs and
priorities on FGR identified by countries are
adequately addressed by stakeholders.
National polices and institutions
Commitment at national and local levels to
specified objectives and priorities is a prerequisite
for the implementation of sustainable FGR
conservation programmes.
In many countries, national policies and
regulatory frameworks for FGR are currently
partial, ineffective or inexistent, partly because
FGR are not well understood in these countries.
In most countries, forest policies fail to address
concerns of sustainable FGR management
(e.g. in situ conservation of species and species
populations), or address them inadequately.
Given the large number of stakeholders
involved in using, developing and managing FGR
at the national level, countries should develop
national strategies and programmes to provide
an appropriate framework of action given the
national context. For example, demand for forest
products including roundwood, fuelwood and
NWFPs is increasing in many countries, and in
some countries the value of NWFPs is higher than
that of roundwood and fuelwood (FAO, 2010a).
Sound social and economic policies are needed to
ensure the integration of FGR in wider national
forest policy frameworks.
For the countries that do not have a national
programme for managing and conserving
FGR, the main challenge is to develop such a
programme with multistakeholder participation.
These countries also need to develop national
forest genetic resources networks or join regional
networks. Networking and institutional twinning,
which have long traditions in forestry, should be
vigorously promoted.
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Countries developing national forestry action
plans should be encouraged specifically to include
genetic-level responses to climate change in their
plans.
Governments ensured broad ownership of the
country reporting process for The State of the
World’s Forest Genetic Resources by organizing
stakeholder workshops to review and validate
their reports. This participatory process needs
to continue, to ensure commitment among all
stakeholders in the country.
Many countries note that FGR legislation and
regulations need to be improved and expanded
and the gaps filled. Requirements for reporting
and sanctions for non-compliance need to be
addressed. Cooperation among national and
subnational authorities and other stakeholders
concerned with FGR should be enhanced. In
some cases, where cooperation among national
authorities does not exist, a permanent national
commission for conservation and management of
forest genetic resources should be established.
Institutional strengthening, training and
support to research are needed for countries
to be able to respond to pressing and
increasingly varied needs in conservation and
FGR management. National research systems,
including tree seed centres, have a crucial role
in this context, as does their support by relevant
international programmes and initiatives.
Countries should particularly aim at creating
synergy
with
FGR-related
international
programmes and conventions (Box 18.1),
coordinated by different national authorities, to
enable efficient information sharing and resource
use and better support of the national FGR
priorities identified.
In addition, most countries, in their reports and
during the regional consultations, highlighted the
need to promote thematic networking to facilitate
linkage among stakeholders and enhance
institutional development and capacity building.
Decentralization
Many developing countries have or are shifting to
a decentralized administration for management
of natural resources, including FGR; the objective
228
Box 18.1
Integrating forest genetic resources
in international forest and natural
resource management policy
framework
A number of international agreements progressively
being implemented under the CBD will serve as
useful international frameworks for promoting the
sustainable use, management and conservation of
FGR at the global and national levels. The Cartagena
Protocol on Biosafety is in force, and the Nagoya
Protocol on Access to Genetic Resources and the Fair
and Equitable Sharing of Benefits Arising from their
Utilization is now being ratified by member countries.
REDD+, in countries where it is implemented, can
be another important mechanism for developing FGR
conservation activities – since REDD+ goes beyond
rewarding carbon storage and also compensates
conservation and sustainable management of forests
in order to encourage developing countries to
contribute to climate change mitigation.
International harmonization of action is necessary
to ensure that wasteful duplication of efforts is
avoided, important FGR issues are not inadvertently
neglected, the reporting burden on countries is
minimized, and provision of data and information
is consistent across sectors, thus facilitating crosssectoral linkages.
is to improve equitable access to and sustainable
management of natural resources by indigenous
and local people who rely on them for their
livelihood.
In some countries regulatory measures are
decided at province or state level. There is
therefore a need to provide appropriate technical
support to decentralized administrations to ensure
that their policy tools provide for sustainable
use and management of FGR, for instance by
retaining customary use by indigenous and local
communities.
To take advantage of the opportunities that
decentralization offers, it is important to be aware
of the political and environmental limitations
of decentralized natural resource management
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
programmes. It is likewise important to be more
strategic in building sustainability based on
the sociocultural, economic and environmental
context of the specific target area, be it a state,
a province, a district or a village. Well planned
decentralization initiatives are enhanced by
people’s participation at local level and ownership
of management programmes by stakeholders.
It is commonly accepted that in situ conservation
of FGR has a better chance of success if local or
indigenous communities living in or near the
forest are responsible for or strongly involved in
implementing the conservation programmes. In
this regard decentralization represents a useful
policy framework for in situ FGR conservation in
many countries.
Prioritizing species at the national level
The number of species mentioned by countries
as priority species varies from less than 10 to
nearly 300. Given the high number of tree species
recorded as priority worldwide – about 2 300
recorded from 86 country reports – it is clear that
prioritization of the many alternative species
should be encouraged for more efficient action
at the national, regional or international levels
(Box 18.2). Priority setting is complicated greatly
by the lack of basic information on the variation,
variation patterns and potential of many tree
species. A species approach is regarded as an
adequate and useful option for understanding
and developing FGR. Updated information
on the country’s forest species, their uses and
their conservation status is a good basis for
sound identification of country priority species
for action. Priority species can be identified at
the national or subnational level and shared
in existing regional and international fora to
enhance focus and efficiency of resource use.
The general aim of priority setting is to compare
the consequences and trade-offs of a range of
actions. It implies that some areas, species or
Box 18.2
Regional collaboration in FGR conservation and management:
joint strategies and priorities
Regional strategies for FGR conservation, including
regional networks of in situ genetic conservation
units and corridors of priority species, are needed
to ensure the dynamic conservation of key forest
genetic resources and their evolutionary ability for
the future. Investment in joint regional activities
may often be more efficient and cost effective than
dealing with common issues at the national level,
which can entail duplication of activities. Definition
and implementation of regional FGR conservation
strategies provide a good justification for coordination
and collaboration at the regional level.
Regional priorities for action were identified during
the regional consultations held as part of the process
for The State of the World’s Forest Genetic Resources,
and some regional priority species were discussed;
in many cases, the same species have been identified
as priorities by existing regional FGR networks.
This process needs to continue in order to define
detailed actions for each species and to allocate
responsibilities among actors and partners at the
national, regional and international levels.
Most of the regional consultation workshops,
furthermore, recommended the promotion of
regional mechanisms to facilitate access to forest
reproductive material for scientific work. It was noted
that regulation of transfer and exchange of forest
reproductive material under international agreements
can sometimes limit access to proper quality genetic
material, even restricting transfer among countries
sharing the same ecological conditions. This constraint
can prevent research programmes from delivering
results that will have real impact. The regional
consultations encouraged regional networking
for exchange of FGR material, in compliance with
national legislation and applicable international
regulations.
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genetic resources will be given lower priority than
others. When different stakeholders have similar
priorities, concerted action on the part of these
stakeholders is possible. When their priorities are
dissimilar, independent but harmonized action
is more likely to succeed. Governmental, nongovernmental and international organizations
active in forest biological diversity and genetic
conservation will often differ substantially in their
priorities as well as in their ability to implement
various management techniques. Where such
differences exist, it is necessary to form coalitions
for action, which must operate under coherent
frameworks and at appropriate levels.
Capacity building
Many countries report weak technical and
scientific capacity in FGR-related fields.
University training curricula on issues such as FGR
conservation, tree breeding and management of
NWFPs are rarely available in those countries. In
most countries (and particularly in developing
countries and countries in economic transition),
research and education need to be strengthened
in all areas of FGR management and at both
technical and professional levels. Establishing,
strengthening
and
maintaining
research
and education institutions is key to building
national capacity to plan and implement priority
activities for sustainable use, development and
conservation of FGR.
Technical training should be designed to help
countries capitalize on recent developments in
forest inventory (remote sensing, GIS) and forest
genetics (traditional and more recent molecular
marker technologies and other biotechnological
tools, as applicable).
In particular, many countries report a weakness
in policy and institutional capacity related to
the conservation, sustainable management and
development of FGR. To fill this gap, the following
needs should be addressed. Those relating to
information and public awareness are discussed
further below.
• FGR conservation and management needs
must be updated and integrated into
230
•
•
•
•
•
•
•
•
wider national policies, programmes and
frameworks for action at the national,
regional and global levels.
Coordination and collaboration among
national institutions and programmes
related to FGR must be promoted.
National capacity to manage FGR
should be assessed, and educational and
research capacity strengthened to ensure
adequate technical support to FGR-related
development programmes.
The participation of indigenous and
local communities in FGR management
should be promoted in the context of
decentralization.
Regional mechanisms for exchange of forest
reproductive material for research and
development, consistent with applicable
international conventions, need to be
promoted and applied.
Regional and international cooperation
should be reinforced in support of
education, knowledge dissemination,
research, conservation and sustainable
management of FGR.
Network activities for information sharing
on FGR research, management and
conservation should be encouraged and
developed.
Public and international awareness of the
role and value of FGR should be promoted.
The necessary resources need to be
mobilized.
Improving information availability
and access
With the increasing pressure on forest resources
and the high cost of land-use change in terms of
forest conservation, the need for reliable data
on FGR status and trends has become acute.
Reliable data are required at all levels to support
decision-making that will enable sustainable
management of forest resources, including FGR.
This publication on The State of the World’s
Forest Genetic Resources provides the first global
overview of the diversity, status and trends of FGR
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
and of national regional and global capacity to
manage these resources.
Many
country
reports
indicate
that
information at the country level is incomplete,
scattered and difficult to obtain, although
some progress has been made during the past
decade. In some countries information is barely
available.
Reported gaps in information and data needed
to support adequate FGR management relate to:
• availability of updated country-level species
lists;
• availability of appropriate indicators that
can be easily used to evaluate the status of
forest genetic resources or to measure the
impact of factors such as changes in land
use and overexploitation on diversity within
and among species;
• knowledge of reproductive and
development characteristics of forest
species, required for ex situ conservation,
production of seedlings, planting and
development of species outside their
original habitats;
• documentation of traditional knowledge
and beliefs related to FGR use and
management.
These deficiencies complicate global monitoring of the status and trends of FGR and limit
capacity for effective decision- making and action
at national and international levels. Global forest
information initiatives such as FAO’s Global
Forest Resources Assessment (FRA) and national
forest inventory and monitoring programmes are
important sources of information; however, the
available information largely relates to forest
resources in general rather than to forest diversity
and variation within tree species. Information
relevant to FGR management needs to be
incorporated in such initiatives. In the absence
of scientifically assessed intraspecific genetic
information, a reasonable short-term alternative
would be to maintain a range of populations for
the targeted species selected from throughout
its natural distribution and covering different
climatic zones and soil types.
National FGR assessment,
characterization and monitoring systems
To improve access to information on FGR for all
stakeholders, establishment and strengthening
of FGR information systems are urgently needed,
including databases to store and share knowledge
(both scientific and, where available, traditional)
on uses, distribution, habitats, biology, and
morphological and genetic variation of species
and species populations. The use of common
protocols for FGR inventories, characterization
and monitoring should be promoted to ensure
that data collected from different countries are
comparable. However, each country may need to
adapt the protocols to its specific constraints and
frameworks.
As mentioned above, national forest
inventories do not usually include the relevant
concerns for the planning and sustainable
management of FGR. Forest inventory activities
should include updating of species lists and
development of species distribution maps, as
this information is necessary for development of
plans for conservation, sustainable management
and development of forest genetic resources.
However, because of the limited number of plant
taxonomists and forest geneticists and the lack of
adequate technical infrastructure and financial
resources, many developing countries have
limited capacity to conduct botanical inventories,
to keep updated species lists and to create species
distribution maps, as well as to manage national
herbaria and gene banks. This problem should be
addressed by including activities such as updating
of country forest species checklists and monitoring
and assessment of populations of important
forest species in national programmes where
appropriate. Countries with limited resources
will need to build synergy among programmes
and institutions at the national, regional and
international levels to minimize the constraints.
Some countries report that they have
information from FGR-related activities in the
public sector but lack information from the
private sector; this disparity indicates the need for
integrated and harmonized documentation and
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 5
data gathering activities between the sectors.
Industry organizations with comprehensive
membership in both the public and private
sectors, as exist for example in Canada, might
assist in promoting this practice.
Networks
Regional FGR networks should be strengthened
and expanded. Regional networks are particularly
important for addressing needs for information,
research and strategy development for species
that cross multiple national borders.
Collaboration among international networks
provides a means to amalgamate FGR knowledge
and data hosted by various agencies and
institutions, which is beneficial for the design
of conservation and management strategies.
Shared national forest resource inventories could
include forest ecosystem maps and disturbance
databases. The opportunity to further strengthen
collaborative relationships and cross-border
studies will become more apparent as gaps in
FGR-related knowledge are studied.
Preserving and enhancing use of
traditional knowledge
Traditional knowledge represents a source of
valuable information that must be adequately
considered in development of national, regional
and global programmes on FGR conservation,
sustainable use and development.
Traditional farming systems and practices for
using and managing natural resources (including
forest genetic resources), based on longestablished knowledge, help to ensure food and
agricultural diversity, livelihoods, food security
and the maintenance of valuable ecosystems.
However, traditional livelihoods and indigenous
plant varieties and landraces are now increasingly
endangered by large-scale commercialization of
agriculture and forestry, land-use and land-cover
changes and the impacts of climate change.
In many countries, traditional knowledge and
FGR use and management are closely linked,
for example in agroforestry farming systems.
232
Traditional knowledge on the use of trees
and tree products contributes to the welfare
of indigenous and local communities in many
countries (in terms of food, medicine, shelter),
while also representing a tremendous asset for
industrial and trade development in such sectors
as cosmetics, pharmacy, food technology and
biopesticides. Most country reports acknowledge
that FGR-related traditional knowledge is an
important asset for improving the contribution
of forests to national economies as well as
people’s livelihoods. Furthermore, it contributes
to sustainable development through practices
such as local conservation and sustainable use
of plants; it can also contribute to mitigating
serious global problems such as climate change,
desertification and land and water degradation.
Traditional knowledge of FGR is increasingly
under threat, however, as a consequence of
FGR degradation and changes in land use and
sociocultural practices; thus the need to preserve
this knowledge is becoming acute. Policies on
FGR information management should consider
traditional knowledge as an important source
of information and an essential asset, and this
viewpoint needs to be adequately relected in
national assessments, technical programmes and
policy documents.
Priority areas for research
In many ways, understanding of genetic diversity
in trees is advancing rapidly, yet the scale of the
task ahead remains enormous, especially for
tropical species (Box 18.3). With highly diverse
forest ecosystems under extreme pressure from
human exploitation, changing climate and
shifting ecology, the need to improve the state
of knowledge is acute. For most species, even
basic data are lacking on the standing resources,
the mechanisms that maintain them, and
their dynamics. It is a vast challenge to gather
and interpret available data to discover how
contemporary genetic resources were formed
and to predict how they may respond to changes
in the future.
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
The
following
research
priorities
are
recommended as an entry point for addressing
this challenge. It is likely that these efforts would
be most effective if targeted at the priority
species identified by regional forest genetic
resources networks, but other species should not
be excluded.
• Improve taxonomic knowledge, particularly
for tropical tree species.
• Improve understanding of the genetic
diversity of important or priority species
and the processes that determine the
structure of the genetic diversity within
and between populations. Identification
and understanding of the genetic traits
that enhance species’ adaptation will be
particularly useful in tackling challenges
related to climate change, while also
providing opportunities to boost the
functions of trees and forests.
• Map tropical forest genetic resources. Assess
key priority species across their complete
range, i.e. map their genetic variation, to
provide a context for FGR prioritization.
Map layers could include neutral genetic
data, genomic data and experimentally
assessed phenotypic variation. Existing
mapping initiatives such as MAPFORGEN
(www.mapforgen.org) provide a lead.
• Promote meta-analysis on key aspects
of forest genetic resources for better
understanding of the link between
characteristics of diversity at species
and genetic levels and the processes
determining the state of genetic diversity of
priority or important forest species.
Box 18.3
The state of knowledge on forest genetic resources: a summary
• Knowledge of FGR is reported to be inadequate
for well-informed policy or management in most
countries.
• Among the 80 000 to 100 000 tree species,
studies have described genetic parameters for
less than 1 percent, although both the number
of studies and the number of species studied
have increased significantly in the past decade.
• Most studies conducted during the past two
decades have been at the molecular level, either
using DNA markers or genomic technologies
to characterize genetic resources. Molecular
information is accumulating much faster
than whole-organism information, with the
consequence that little of the accumulating
knowledge has direct application in
management, improvement or conservation.
• A few species have been well researched –
through both molecular and quantitative studies
– and genetically characterized; these mainly
comprise temperate conifers, eucalypts, several
acacias, teak and a few other broadly adapted,
widely planted and rapidly growing species.
• Quantitative genetic knowledge has led to
significant productivity gains in a small number
of high-value planted timber species.
• Genomic knowledge of forest trees lags behind
that of model herbaceous species, including
the important agricultural crops, but for several
tree species the entire genome has been or
is in the process of being sequenced, and
novel approaches have been developed to link
markers to important traits. Genomic or markerassisted selection is close to being realized, but
phenotyping and data management are the
biggest bottlenecks.
• Many of the species identified as priorities,
especially for local use, have received little or
no research attention, indicating a need to
associate funding with priority-setting exercises.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
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Communication and awareness
raising
Most country reports mention inadequate public
awareness of FGR, highlighting the need to
promote awareness among decision-makers and
the general public of the importance of FGR
and their role in meeting present and future
development needs. Lack of information limits
the capacity of countries and the international
community to integrate FGR management into
cross-cutting policies. Furthermore, lack of general
understanding of the importance of FGR is an
impediment to generation of new knowledge.
Most countries do not have specific programmes
for creating public awareness of forest genetic
resources. Throughout the world the public is
more aware of threats such as climate change
and changes in land use and their implications for
forests, particularly boreal and other old-growth
forests, as these topics are prevalent in the
media. Information about the importance of FGR
is not one of the chief priorities of institutions
conducting public relations about forests, forestry
and nature conservation. In today’s informationlooded society, forests in general and forest
genetic resources in particular compete for
public attention with many other subjects. Even
for those people who are interested in and seek
information about forests, the many stakeholders
each with their respective interests create a rather
confusing picture.
Much of the public is sceptical about genetic
engineering. In some developed countries, genetic
engineering and genetics are often considered
one and the same. The word “genetics” and
all related terms have often acquired negative
connotations. Such preconceptions greatly hinder
efforts to convey knowledge of the importance of
forest genetic resources.
The increasing urbanization of the population
and the lack of knowledge about nature
are impediments to public awareness. Forest
genetic resources generally have no reference
234
in people’s private lives. Only a small fraction of
the population works in or derives income from
the forest. As a consequence, understanding
of forest management is dropping; the forests
are perceived most often as natural assets
for protection or as the green backdrop of
recreational activities. The lack of understanding
of forest management is accompanied by a lack
of understanding about productive provenances
or forest plant breeding.
Some surveys reported by countries showed
that the public and NGOs have the lowest
awareness of the roles and value of forest genetic
resources. Industry and government rated much
higher. In general, the value of forest genetic
resources has not been widely communicated at
the national level.
Awareness raising initiatives or programmes
should be created for greater visibility of forest
genetic resources. The following are some speciic
recommendations.
• Countries should develop a genetic
resources communication strategy and make
forest genetic resources information more
accessible.
• Countries should provide training and
education on forest genetic resources to
improve understanding of their beneits
and value.
• Training is required to sensitize policymakers about the responsibilities and
advantages of FGR management action in
the short and longer term.
• Awareness can be strengthened by using
television, newspapers and other media
to inform the public about forest genetic
resources and their protection.
• Institutions that can usefully contribute
to raising FGR awareness include parks,
reserves, territories of regional forestry
boards, state forests and game enterprises,
educational and practical forest enterprises,
and specialized forest schools.
NEEDS, C HALLENGES A ND REQU I RED RESP ONSES FOR THE FUTU RE
In conclusion: what needs to be
done
Improve the availability and accessibility of
knowledge and information on species and
their genetic diversity, forest ecosystems and
related traditional knowledge, to facilitate
and enable decision-making on sustainable use
and management of FGR and to enhance their
contribution to solving serious global problems such
as food shortage, land and water degradation, the
effects of climate change, and increased demand
for various forest products and services:
• Establish and strengthen national
FGR assessment, characterization and
monitoring systems.
• Develop national and subnational systems
for the assessment and management of
traditional knowledge on FGR.
• Develop international technical standards
and protocols for FGR inventory,
characterization and monitoring of trends
and risks.
• Promote the establishment and
reinforcement of FGR information systems
(databases) to cover available scientific and
traditional knowledge on uses, distribution,
habitats, biology and genetic variation of
species and species populations.
Enhance in situ and ex situ conservation of FGR,
to maintain genetic diversity and the evolutionary
processes of forest tree species:
• Strengthen the contribution of primary
forests and protected areas to in situ
conservation of FGR.
• Promote the establishment and
development of efficient and sustainable ex
situ conservation systems, including in vivo
collections and gene banks.
• Support and strengthen the role of
indigenous and local communities in the
sustainable management and conservation
of FGR.
• Identify priority species for action.
• Harmonize measures for in situ and ex situ
conservation, including through regional
cooperation and networking.
Enhance the sustainable use, development
and management of FGR to contribute to
environmental sustainability, food security and
poverty alleviation:
• Develop and reinforce national seed
programmes to ensure the availability of
genetically appropriate tree seeds in the
quantities and of the quality needed for
national plantation programmes.
• Promote restoration and rehabilitation of
ecosystems using genetically appropriate
material.
• Support climate change adaptation and
mitigation through proper management
and use of FGR.
• Promote good practices and appropriate
use of emerging technology to support the
conservation, development and sustainable
use of FGR.
• Develop and reinforce research programmes
on tree breeding, domestication and
bioprospecting.
• Develop and promote networking and
collaboration among concerned countries to
combat invasive species affecting FGR.
Strengthen policies and institutional capacity to
address major issues related to sustainable FGR
management and enable successful medium- and
long-term planning for long-term sustainable
use, management and conservation of FGR:
• Develop national strategies for in situ and
ex situ conservation and sustainable use of
FGR.
• Integrate FGR conservation and
management into wider policies,
programmes and frameworks of action at
the national, regional and global levels.
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THE STATE OF THE WORLD' S FOREST GENE T IC RESOU RC ES
PART 5
• Develop collaboration and promote
coordination of national institutions and
programmes related to FGR.
• Establish and strengthen educational and
research capacities on FGR.
• Promote the participation of indigenous
and local communities in FGR management
in the context of decentralization.
• Promote and apply mechanisms for
regional germplasm exchange for research
and development, in agreement with
international conventions.
236
• Reinforce regional and international
cooperation, including networking,
to support education, knowledge
dissemination, research, and conservation
and sustainable management of FGR.
• Promote public and international awareness
of the roles and value of FGR.
• Strengthen efforts to mobilize the
necessary resources, including financing,
for the conservation, sustainable use and
development of FGR.
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Acronyms and abbreviations
ACIAR
AFLP
AFTP
CATIE
CBD
CIAT
CITES
COFO
cpDNA
CSIRO
CTFT
DANIDA
DArT
DFSC
DNA
EEM
EST
ESTP
EU
EUFGIS
EUFORGEN
FGR
FRA
FSC
GEF
GFIS
GIS
GM
ICRAF
INPA
IPCC
IPGRI
IPPC
ISSR
ISTA
ITS
ITTO
ITWG-FGR
IUCN
IUFRO
MDGs
MTA
NGO
NLBI
NTSC
Australian Centre for International Agricultural Research
amplified fragment length polymorphism
agroforestry tree product
Tropical Agricultural Research and Higher Education Center
Convention on Biological Diversity
International Centre for Tropical Agriculture
Convention on International Trade in Endangered Species of Wild Fauna and
Flora
FAO Committee on Forestry
chloroplast DNA
Commonwealth Scientific and Industrial Research Organization (Australia)
Centre technique forestier tropical (France)
Danish International Development Agency
diversity arrays technology
DANIDA Forest Seed Centre
deoxyribonucleic acid
environmental envelope modelling
expressed sequence tag
expressed sequence tag polymorphism
European Union
European Information System on Forest Genetic Resources
European Forest Genetic Resources Programme
forest genetic resource(s)
Global Forest Resources Assessment
Forest Stewardship Council
Global Environment Facility
Global Forest Information Service
geographic information system
genetic modification/genetically modified
World Agroforestry Centre
Instituto Nacional de Pesquisas da Amazônia (Brazil)
Intergovernmental Panel on Climate Change
International Plant Genetic Resources Institute (now Bioversity International)
International Plant Protection Convention
inter-simple sequence repeat
International Seed Testing Association
internal transcribed spacer
International Tropical Timber Organization
Intergovernmental Technical Working Group on Forest Genetic Resources
International Union for Conservation of Nature
International Union of Forest Research Organizations
Millennium Development Goals
Material Transfer Agreement
non-governmental organization
Non-Legally Binding Instrument on All Types of Forests
national tree seed centre
275
NWFP
OECD
PCR
PEFC
PGRFA
PNGFRI
QTL
RAPD
REDD+
RFLP
RNA
SADC
SFM
SIDS
SNP
SPRIG
SSR
TEK
UNEP
UNESCO
UNFCCC
USDA
WTO
276
non-wood forest product
Organisation for Economic Co-operation and Development
polymerase chain reaction
Programme for the Endorsement of Forest Certification
plant genetic resources for food and agriculture
Papua New Guinea Forest Research Institute
quantitative trait locus
random amplified polymorphism DNA
reducing emissions from deforestation and forest degradation in developing
countries (including the role of conservation, sustainable management of
forests and enhancement of forest carbon stocks)
restricted fragment length polymorphism
ribonucleic acid
Southern African Development Community
sustainable forest management
small island developing States
single-nucleotide polymorphism
South Pacific Regional Initiative on Forest Genetic Resources
simple sequence repeat
traditional ecological knowledge
United Nations Environment Programme
United Nations Educational, Scientific and Cultural Organization
United Nations Framework Convention on Climate Change
United States Department of Agriculture
World Trade Organization
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