John C. Weber

Genetic variation of Calycophyllum spruceanum in the Peruvian Amazon Basin, revealed by amplified fragment length polymorphism (AFLP) analysis

An understanding of the level, structure and origin of genetic variation within and among populations of tropical trees is essential for devising optimum management strategies for their sustainable utilization and conservation. Here, amplified fragment length polymorphism (AFLP) analysis was used to partition genetic variation within and among nine populations of the predominantly riverine tree, Calycophyllum spruceanum, sampled across a wide geographical range along river tributaries of the Peruvian Amazon Basin. Analysis of molecular variance (AMOVA) employed 65 AFLP markers and revealed most variation among individuals within populations (91%), although variation among populations was highly significant (P < 0.001). Calculation of genetic distances and nested AMOVA indicated a degree of structuring among populations based on geographical proximity, although clustering did not depend on geographical distance alone. No firm evidence was obtained for unidirectional seed dispersal by water playing an important role in determining genetic structure over the geographical range sampled. Implications of data for optimising genetic management of the species are discussed and areas for further study identified.

Managing genetic variation in tropical trees: linking knowledge with action in agroforestry ecosystems for improved conservation and enhanced livelihoods

Tree species in agroforestry ecosystems contribute to the livelihoods of rural communities and play an important role in the conservation of biodiversity. Unless agroforestry landscapes are productive, however, farmers will not maintain or enhance the range and quality of tree species in them, and both income opportunities and biodiversity will be lost. Productivity depends on both tree species diversity and genetic (intra-specific) variation, but research on the latter has until recently not received the recognition it deserves. Worse, when knowledge on tree genetic variation in agroforestry systems has become available, it has not generally been linked in any systematic way with management, indicating a disjunction between research and field-level practice. In this essay, we attempt to bridge this gap by considering three questions: why is genetic diversity important in tree species? What is our current state of knowledge about intra-specific variation in trees in agroforestry systems? And, finally, what practical interventions are possible to support the conservation of this diversity in agricultural landscapes, while enhancing farmers’ livelihoods? A wide genetic base in agroforestry trees is essential to prevent inbreeding depression and allow adaptation to changing environmental conditions and to altering markets for tree products. Recent evidence shows, however, that many species are subject to poor germplasm collection practice, occur at low densities in farmland, and are found in highly aggregated distributions, all of which observations raise concerns about productivity and sustainability. A range of germplasm-access based interventions is necessary to improve current management, including the enhancement of community seed- and seedling-exchange networks, and the development of locally based tree domestication activities. Equally necessary, but more difficult to address, is the development of markets that support genetic diversity in tropical tree species; we discuss approaches by which this may be undertaken.

Do farmers reduce genetic diversity when they domesticate tropical trees? A case study from Amazonia.

Agroforestry ecosystems may be an important resource for conservation and sustainable use of tropical trees, but little is known of the genetic diversity they contain. Inga edulis, a widespread indigenous fruit tree in South America, is used as a model to assess the maintenance of genetic diversity in five planted vs. five natural stands in the Peruvian Amazon. Analysis of five SSR (simple sequence repeat) loci indicated lower allelic variation in planted stands [mean corrected allelic richness 31.3 (planted) and 39.3 (natural), P = 0.009]. Concerns regarding genetic erosion in planted Amazonian tree stands appear valid, although allelic variation on‐farm is still relatively high.

What is the relevance of smallholders’ agroforestry systems for conserving tropical tree species and genetic diversity in circa situm, in situ and ex situ settings? A review

Smallholders’ agroforests may be valuable for conserving tropical trees through three main mechanisms. First, trees planted and/or retained by farmers in agricultural landscapes where wild stands were once found may be circa situm reservoirs of biodiversity. Second, farmland trees may support conservation in situ by providing an alternative source of product to reduce extraction from forest, and by acting as ‘corridors’ or ‘stepping stones’ that connect fragmented wild stands. Third, the additional value that planting assigns to trees may result in greater interest in including them in seed collections, field trials and field ‘genebanks’ that support ex situ conservation. Here, we critically review the evidence for these mechanisms, and highlight areas for research and for intervention so that agroforestry practices can better support conservation in each setting, with an emphasis on often neglected genetic-level considerations. Based on current global challenges to diversity, conservation will need to rely increasingly on a smallholder-farm circa situm approach, but concerns on long-term effectiveness need to be properly quantified and addressed. Connectivity between widely dispersed, low density trees in agricultural landscapes is an important factor determining the success of the circa situm approach, while improving farmers’ access to a diversity of tree germplasm that they are interested in planting is required. The circumstances in which agroforestry plantings can support in situ conservation need to be better defined, and research on the stability of active tree seed collections (how long are species and populations retained in them?) as ex situ reservoirs of biodiversity is needed.

Restoring forests: What constitutes success in the twenty-first century?

Forest loss and degradation is occurring at high rates but humankind is experiencing historical momentum that favors forest restoration. Approaches to restoration may follow various paradigms depending on stakeholder objectives, regional climate, or the degree of site degradation. The vast amount of land requiring restoration implies the need for spatial prioritization of restoration efforts according to cost-benefit analyses that include ecological risks. To design resistant and resilient ecosystems that can adapt to emerging circumstances, an adaptive management approach is needed. Global change, in particular, imparts a high degree of uncertainty about the future ecological and societal conditions of forest ecosystems to be restored, as well as their desired goods and services. We must also reconsider the suite of species incorporated into restoration with the aim of moving toward more stress resistant and competitive combinations in the longer term. Non-native species may serve an important role under some circumstances, e.g., to facilitate reintroduction of native species. Propagation and field establishment techniques must promote survival through seedling stress resistance and site preparation. An improved ability to generalize among plant functional groups in ecological niche adaptations will help to overcome site-limiting factors. The magnitude and velocity of ongoing global change necessitates rapid responses in genetics that cannot be naturally induced at valid temporal and spatial scales. The capacity for new concepts and technologies to be adopted by managers and accepted by society will depend on effective technology transfer and a community-based approach to forest restoration. The many benefits human society gains from forests requires that forest restoration considers multiple objectives and approaches to minimize trade-offs in achieving these objectives.

Climate change and tree genetic resource management: maintaining and enhancing the productivity and value of smallholder tropical agroforestry landscapes. A review

Anthropogenic climate change has significant consequences for the sustainability and productivity of agroforestry ecosystems upon which millions of smallholders in the tropics depend and that provide valuable global services. We here consider the current state of knowledge of the impacts of climate change on tree genetic resources and implications for action in a smallholder setting. Required measures to respond to change include: (1) the facilitated translocation of environmentally-matched germplasm across appropriate geographic scales, (2) the elevation of effective population sizes of tree stands through the promotion of pollinators and other farm management interventions; and (3) the use of a wider range of ‘plastic’ species and populations for planting. Key bottlenecks to response that are discussed here include limitations in the international exchange of tree seed and seedlings, and the absence of well-functioning delivery systems to provide smallholders with better-adapted planting material. Greater research on population-level environmental responses in indigenous tree species is important, and more studies of animal pollinators in farm landscapes are required. The development of well-functioning markets for new products that farmers can grow in order to mitigate and adapt to anthropogenic climate change must also consider genetic resource issues, as we describe.

FARMERS' PREFERENCES FOR TREE FUNCTIONS AND SPECIES IN THE WEST AFRICAN SAHEL

ABSTRACT Surveys were conducted in villages in five regions of Burkina Faso, Mali, Niger and Senegal to determine priority tree functions and species. Villagers listed eight priority functions and 116 important species. The most important functions were essential products (human food, medicines, animal food, wood/energy/fiber) followed by environmental services (soil fertility improvement, soil/water conservation, shade) and sale to generate revenue. The relative importance of most functions and several species differed significantly among some regions. There were significant positive or negative associations between functions. Villagers used significantly more species in Niger, the driest region, in order to diversify options and minimize risk.

Strategies for genetic conservation of trees in the Peruvian Amazon

Forestry practices and high rates of land clearance for agriculture are causing genetic erosion of valuable tree species in the Peruvian Amazon, thereby endangering the economic sustainability of rural communities and limiting Perus opportunities for the development of new timber and non-timber forest products. The potential utility and limitations of six low-input interventions to help forestall further genetic erosion in the region are discussed, with a focus on local community involvement. Improved agroforestry systems may help reduce deforestation by increasing farm productivity, although methods to increase the currently low adoption rate of these technologies need to be developed. Use of strategic tree domestication techniques can also improve farm productivity and prevent inadvertent genetic drift and inbreeding associated with traditional domestication practices, although to have a major impact, current programs need to be extended across the region. Woodlot forestry could supplant selective extraction of timber and offers an attractive opportunity for poverty alleviation if appropriate credit and land tenure policies can be developed. However, it may also result in increased deforestation if activities on public land cannot be controlled. The implementation of improved seed collection systems and simple seed transfer guidelines would help to reduce the collection of seed of poor quality and low genetic diversity, and avoid maladapted plantings, although such programs are difficult to monitor and seed costs may increase. Strategic identification and design of in situ conservation areas would help to ensure the viability of conserved populations, but requires the forfeiture of significant revenue from timber concessions.

SHORT COMMUNICATION: Do farmers reduce genetic diversity when they domesticate tropical trees? A case study from Amazonia

Agroforestry ecosystems may be an important resource for conservation and sustainable use of tropical trees, but little is known of the genetic diversity they contain. Inga edulis, a widespread indigenous fruit tree in South America, is used as a model to assess the maintenance of genetic diversity in five planted vs. five natural stands in the Peruvian Amazon. Analysis of five SSR (simple sequence repeat) loci indicated lower allelic variation in planted stands [mean corrected allelic richness 31.3 (planted) and 39.3 (natural), P = 0.009]. Concerns regarding genetic erosion in planted Amazonian tree stands appear valid, although allelic variation on‐farm is still relatively high.

Origins and genetic conservation of tropical trees in agroforestry systems: a case study from the Peruvian Amazon

Hundreds of native tree species are currently found in extensive agroforestry ecosystems in the Peruvian Amazon, forming an important reservoir of biodiversity. To further promote conservation, farmers are encouraged to supplement intra-specific genetic diversity in these populations with seed collected from local forests. For some tree species, however, this approach may be inappropriate, as stands of these taxa already found on-farm may not be of local origin. Despite this issue being of importance for conservation, little information is available on the history of cultivated trees in the region, a situation that we here rectify for the important fruit tree Inga edulis. Based on nuclear SSR and chloroplast DHPLC analyses of closely geographically matched natural and planted stands at five sites, it appears that cultivated material of I. edulis is primarily of non-local origin, indicating that conservation based on new wide-scale infusions from local wild stands into farms may be inappropriate in the region. Although nuclear and chloroplast diversity were both lower in planted stands, values were still relatively high (∼80 and 70% of natural stands, respectively), indicating that when farmers plant trees, good collection practice of seed from already cultivated I. edulis should be an effective means for ensuring long-term conservation on farms.