Second Growth The Promise of Tropical Forest Regeneration in an Age of Deforestation
by Robin L. Chazdon
University of Chicago Press, 2014
Cloth: 978-0-226-11791-1 | Paper: 978-0-226-11807-9 | Electronic: 978-0-226-11810-9
DOI: 10.7208/chicago/9780226118109.001.0001
ABOUT THIS BOOKAUTHOR BIOGRAPHYREVIEWSTABLE OF CONTENTS

ABOUT THIS BOOK

For decades, conservation and research initiatives in tropical forests have focused almost exclusively on old-growth forests because scientists believed that these “pristine” ecosystems housed superior levels of biodiversity. With Second Growth, Robin L. Chazdon reveals those assumptions to be largely false, bringing to the fore the previously overlooked counterpart to old-growth forest: second growth.

Even as human activities result in extensive fragmentation and deforestation, tropical forests demonstrate a great capacity for natural and human-aided regeneration. Although these damaged landscapes can take centuries to regain the characteristics of old growth, Chazdon shows here that regenerating—or second-growth—forests are vital, dynamic reservoirs of biodiversity and environmental services. What is more, they always have been.

With chapters on the roles these forests play in carbon and nutrient cycling, sustaining biodiversity, providing timber and non-timber products, and integrated agriculture, Second Growth not only offers a thorough and wide-ranging overview of successional and restoration pathways, but also underscores the need to conserve, and further study, regenerating tropical forests in an attempt to inspire a new age of local and global stewardship.

AUTHOR BIOGRAPHY

Robin L. Chazdon is professor in the Department of Ecology and Evolutionary Biology at the University of Connecticut and coeditor of Foundations of Tropical Forest Biology and Tropical Forest Plant Ecophysiology. She lives in Storrs, CT.

REVIEWS

“Impressive and very timely given the growing recognition of the importance of secondary forests in conserving biodiversity. Second Growth provides a very thorough review of the human impacts on tropical forests over several centuries and more recently, and on the ecology of forest regeneration after a range of disturbances. It is a well-researched and valuable contribution to the literature on tropical secondary forests.”
— Karen D. Holl, University of California, Santa Cruz

“A tremendous survey. Chazdon draws on paleobotany, ecology, natural history, and policy and forest management to make the case for the potential of forest regeneration. Second Growth is astounding in its breadth and depth.”
— S. Joseph Wright, Smithsonian Tropical Research Institute

“What a wonderful book! Secondary forests have been ignored for too long, and Second Growth brings together a wealth of material from across the Latin American, African, and Asian tropics to provide a synthesis of what we know about how they can develop and what influences their succession. Sadly, even optimists must accept that primary or old-growth forests will continue to shrink. This means secondary forests are likely to replace them in the future as the main repositories of biodiversity and sources of many ecosystem services. This book is destined to become a key text for those protecting and managing these new forests.”
— David Lamb, University of Queensland, Brisbane, Australia

“At last, an authoritative and very readable account of the most neglected aspect of tropical forest ecology: the vast areas of second growth that if restored and managed properly will yield enormous human and conservation benefit. Chazdon’s book fills a yawning gap in tropical ecology and land management. A great and important work, Second Growth will be an enduring scholarly masterpiece.”
— Thomas E. Lovejoy, George Mason University, Senior Fellow, the United Nations Foundation

Second Growth combines an in-depth review with an eloquent case for the importance of understanding, promoting, and managing forest regeneration in contexts ranging from climate change to provision of ecosystem services and biodiversity conservation. In doing so, it may help to meet those challenges by providing evidence to ensure that the value of secondary forests is recognized. It will certainly stimulate the science needed to support practical action.”
— Valerie Kapos, Science

“Chazdon provides a remarkable compilation of our understanding of naturally regenerating forests across the tropics in this book. She draws upon many fields of knowledge—paleoecology, ecosystem ecology, socio-economic and political ecology, forest management—to convey that regenerating tropical forests are socio-ecological systems that play a critical role in human and ecosystem health. The book is rich with case studies and examples from all over the globe, and provides a much-needed synthesis of second growth in the tropics. . . . Each time I open this book I find something new and interesting. For those working in tropical forests, this text will be an essential reference that not only fills an important gap in our understanding of tropical forest regeneration, but also provides a solid grounding for their conservation and future management.”
— Biotropica

Second Growth provides broad coverage of a subject that, despite its importance, has received less popular attention than ‘old growth’ tropical rain forests. Chazdon does an excellent job of synthesizing key ideas and bringing together recent research on tropical forest regeneration. . . . Chazdon’s work is a valuable resource, providing both a solid review for those with knowledge in the field and a starting point for those new to the topic. . . . Highly recommended.”
— B. D. Orr, Michigan Technological University, Choice

“A leading voice in arguing that large-scale forest regrowth can help to solve some of the world’s problems. . . . Decades of watching the Costa Rican forests recover have taught Chazdon that, at least in areas that still have healthy forests nearby to supply seeds, the main thing human beings need to do is just get out of the way. After all, forests were recovering from fires and other natural calamities long before people ever came along to chop them down.”
— Justin Gillis, New York Times

“A very thorough and holistic review of the ecology of secondary tropical forests. For many years, studies of tropical forest biodiversity tend to have been concentrated on primary forest. Here, Chazdon argues well and forcefully that second-growth forests are also an important reservoir of biodiversity and that they are exciting ecosystems in which to do research. . . . A tour de force—absolutely brilliant. Anyone interested in tropical forests, their conservation and their restoration will want a copy of this book.”
— R. H. Marrs, University of Liverpool, Biological Conservation

“A complete review covering topics from ancient forest management to the effect of global economy on the fate of local forests. . . . Chazdon states in the preface that she wants to deliver an urgent message about forests: they are regenerating and this regeneration is malleable and it can be for our benefit. The message is remarkably delivered. This book integrates patterns that occur in the entire gradient from wet to dry forests. . . . Second Growth has answered most of my questions and has raised a thousand more, as good books and research always do. . . . This well-produced book . . . will be enormously useful for students, professors, and practitioners of forest restoration and management.”
— Cristina Martínez-Garza, Universidad Autónoma del Estado de Morelos, Mexico, Ecology

“Throughout the book, Chazdon returns to the central theme that tropical forests are highly resilient ecosystems whose recent histories are inextricably intertwined with people. Contrary to the dichotomous view of ‘primeval’ old-growth versus ‘degraded’ second-growth forests, Chazdon repeatedly and convincingly makes the case that today's tropical forests reflect legacies of human interactions dating back hundreds or even thousands of years. . . . Second Growth is truly extraordinary in its breadth, depth, and synthesis of a voluminous literature. The book covers regenerating forest ecology, dynamics, and socioecological linkages throughout the tropics and is exhaustively researched; there are over 100 pages of references.”
— Nicole L. Michel, University of Saskatchewan, Conservation Biology

“As policy makers come to grips with . . . ecological uncertainty, they are finding Chazdon’s recent book, Second Growth, all the more valuable. Five years in the writing and published last year, the tome is a kind of guide to restoration, synthesizing decades of research and explaining how tropical forests can come back on their own—and what to do if they don’t. ‘It’s an opus; it covers all you would want to know and could imagine you want to know about secondary forests,’ says Thomas Rudel, a rural sociologist at Rutgers University, New Brunswick, in New Jersey. ‘There’s nothing quite like [it].’ The book . . . arrived at a timely moment, just as large-scale forest restoration was gaining momentum.”
— Elizabeth Pennisi, Science

“Chazdon takes us through the ecological stages of forests developing on old agricultural land, selectively logged areas, and land damaged by fire and hurricanes. Then she explores how these regenerating forests change in structure, species composition, and ecosystem properties, and finally what the future holds, all backed up by over a hundred pages of references. I find her message very compelling that these secondary forests have a high intrinsic ecological value. It is not always sugary good news—large mammals can take a long while to return unless there is nearby undisturbed forest, but it’s heartwarming to read this upbeat book that shows that it’s not all doom and gloom in the tropics. Everyone should read this.”
— Bulletin of the British Ecological Society

“A thoroughly researched, authoritative, and comprehensive volume, drawing on hundreds of cited publications. . .  This is an upbeat tour de force, which relentlessly drives home the message that ‘tropical forests are dynamic and resilient.’ It takes a glass-half-full perspective that urges us to see partially disturbed forests not as ‘degraded,’ but as ‘regenerating.’ It should be on the book shelf of anyone starting out or already working on forest dynamics and restoration ecology in the tropics—graduate students designing thesis projects, their supervisors, the planners and implementers of REDD++ projects, managers of protected areas, etc. All in all, an original and powerful work, that is likely to remain the definitive textbook in its field for many years to come. An empty niche has been filled.”
— Kate Hardwick, Royal Botanic Gardens, Kew, and Stephen Elliott, Chiang Mai University, Thailand, Restoration Ecology

“Extremely comprehensive. Though it is not particularly lengthy, running to 316 pages of text, it covers a huge range of topics relating to forest regeneration from traditional knowledge and prehistoric forest transformations by humans to recovery pathways from fire, landslides, volcanic eruptions, logging, and agricultural use. . . . Chazdon masterfully weaves together anthropology, archaeology and ecology in the discussion of prehistorical impacts of humans on tropical forests. . . . If you are interested in the dynamics of forests in any way this book is essential reading. There is no better summary of current thinking on tropical forest succession out there.”
— Ecology for a Crowded Planet

“This book is a comprehensive treatment of our current understanding of the ecology of regrowth of forests after land clearance in the tropics. There is no one better than Chazdon to write such a volume and I suspect she has used this publication as a culmination of her own work over the last 20 years directly studying and observing the dynamics of second growth in Central America. . . . This book is excellent and the first of its kind to really synthesize the literature on regeneration and recovery of second growth forests. For this Chazdon has done a very good job. . . . A definitive textbook and a great effort. Chazdon is to be commended.”
— Mark S. Ashton, Yale University, Quarterly Review of Biology

TABLE OF CONTENTS

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0001
[Deforestation, disturbance, regeneration, regrowth, resilience, shifting cultivation, succession, tropical forest]
This introductory chapter describes tropical forests as dynamic and resilient systems. Indigenous practitioners of shifting cultivation in tropical forests understood the forest regeneration cycle. Forest succession is a process of self-reorganization and reassembly that takes place gradually over decades and centuries. Tropical forest regeneration is influenced by prior land use, initial colonization, climate, soils, and seed dispersal from forests in the surrounding landscape. There is a limit to the resilience of tropical forests, to their ability to recover from disturbances and reassemble all of their components. When the forest regeneration cycle is broken, forests lose their intrinsic capacity to regenerate, requiring human assistance and management to overcome barriers to natural regeneration. Second-growth forests are now the predominant form of forest cover in the tropics worldwide. An understanding of forest regeneration pathways is essential to manage and restore forests, and to forecast forest change. (pages 1 - 11)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0002
[Agriculture, arboriculture, cultivation, Holocene, hunting, tropical forest]
This chapter examines the effects of prehistoric human occupations on tropical forest structure and composition, from the late Pleistocene through the late-Holocene. Early colonization of lowland tropical rain forests led to the development of forest-based subsistence societies, beginning an unprecedented era of landscape management in regions of Africa, South America, New Guinea, and Southeast Asia. Forest dwellers increased the concentration, overall abundance, and geographic ranges of plant species used for food and shelter. The independent origins and spread of agriculture during the Holocene led to increased human populations and to growing demands for food, leading to extensive land clearance and biomass burning in many tropical regions. The conversion of tropical forests to agriculture land beginning 8000 to 10,000 years ago, created patches of forest disturbance and regeneration within human-occupied landscapes and increased the abundance and geographic distributions of species favored by forest disturbance and human use (pages 12 - 32)
This chapter is available at:
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- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0003
[biomass burning, climate variability, earthworks, Holocene, paleoecology, raised fields, terra preta]
This chapter reviews the many ways in which ancient human activities and their aftermath have transformed tropical forest landscapes throughout the world. Archaeological investigations reveal widespread landscape transformations in tropical forest regions as agriculture became more intensified during the late Holocene. These transformations include construction of terraces, raised fields, and canals, forest burning, and soil modification to enhance fertility and permit intensive agriculture. In many regions of South America, extensive raised field complexes were constructed in seasonally flooded savannas to elevate planting surfaces. In Angkor, Cambodia modifications of the natural waterways 800 years ago changed the hydrology of the area in ways that are still evident today. Paleoecological studies provide ample evidence for periods of deforestation followed by spontaneous forest regrowth in many tropical regions. Large and small landscape transformations have left obvious and subtle legacies of prehistoric human impacts on tropical forest composition, soils, and geomorphology. (pages 33 - 54)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0004
[Biomass, disturbance, forest dynamics, patches, regeneration, vegetation]
This chapter focuses on the dynamics of old-growth tropical forests and the legacies of different types of disturbances on their present structure and floristic composition. Natural disturbance regimes interact with anthropogenic disturbance to influence forest composition and structure. Disturbances can be compared based on four major features: spatial extent, frequency, duration, and severity. Disturbance legacies are most pronounced during early stages of forest regeneration, but they may be observed after hundreds of years as remaining individuals from cohorts of long-lived tree species that established early in stand development. Small-scale dynamics are often homogenized and masked by large-scale stability. Rates of recovery of different forest attributes are strongly affected by the spatial and temporal scale of disturbances and by effects on soil properties. Human disturbances that mimic small-scale natural disturbances and minimize soil disturbance will also favor rapid rates of forest regeneration. (pages 55 - 72)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0005
[Chronosequence, disturbance, pioneer, regeneration, stages, succession, trajectory]
This chapter focuses on successional patterns and processes in tropical forests and how they lead to variation in successional trajectories of vegetation structure and composition following human and natural disturbances. Even within a single forest type and region, multiple successional pathways can arise due to local variation in initial conditions. Differences in early species colonization and dominance strongly affect successional trajectories in vegetation structure and composition. Forest successional stages can be defined based on forest structure, age or size structure of tree populations, or species composition. Aspects of vegetation structure, such as tree density and size distributions, tend to change more rapidly than species composition during succession. The shift in dominance by shade-intolerant species to shade-tolerant species is the most generalizable and predictable feature of successional pathways. Beyond this general trend, successional pathways are highly stochastic, reflecting seemingly random factors that affect forest structure and composition. (pages 73 - 96)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0006
[Colonization, disturbance, establishment, facilitation, nitrogen fixation, primary succession, vegetation]
This chapter examines how tropical forests around the world become established on substrates lacking soil, a process known as primary successsion. These disturbances, such as volcanic eruptions, landslides, and riverbank flooding have always been present in the environment of tropical forests. Many pioneer species that colonize early stages of primary succession also play a major role in early stages of secondary succession following forest clearance by human activities. Enrichment of soil by early colonists facilitates establishment of more nutrient-demanding species. During primary and secondary succession regeneration pathways are determined by species availability and resource availability. Facilitation is a key aspect of early colonization during primary succession. Early colonists require nitrogen-fixing capabilities that permit growth on nutrient-poor substrates. Forest succession on newly created substrates is slower than secondary successional pathways. Forest structure can recover rapidly, as observed following the eruption of Krakatau in 1883. (pages 97 - 110)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0007
[arrested succession, colonization, fallow, pasture, resprouting, seed bank, seed dispersal, shifting cultivation]
This chapter focuses on successional pathways in the aftermath of different types of human disturbances, with an emphasis on agricultural land use. The intensity, extent, severity, and duration of land use directly influence the composition of the soil seed bank and newly arriving seeds, soil nutrient availability, remnant vegetation, and conditions for early seedling establishment. The first colonizers of old fields are primarily drawn from established plants, resprouts, or dormant seeds. Repeated burning can eliminate soil seed banks, depleting sources of recruitment during initial stages of succession. Proximity of forest vegetation or remnant trees is particularly important for the arrival of larger-seeded, late successional species. Remnant trees promote seed dispersal following abandonment and potentially well beyond the early stages of succession. Long-term, intensive land use can lead to alternative stable states. Succession can become arrested or deflected due to a variety of factors, including colonization of invasive exotic species. (pages 111 - 126)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0008
[Cyclone, damage, drought, fire regimes, flammability, hurricane, resprouting, tree mortality, typhoon]
This chapter focuses on patterns of regeneration following hurricanes and fires in tropical forests. After these disturbances, some trees remain and produce seeds and resprouts, promoting more rapid recovery of vegetation structure and composition. Fires are a regular feature of the ecology and vegetation dynamics in seasonally dry tropical forests, and dominant tree species are adapted to fire regimes. Fires are rare events in tropical wet forests and occur only following severe drought events that can also directly cause tree mortality. Few tree species in tropical wet or moist forests have characteristics that confer fire resistance. In areas prone to hurricanes and cyclones, tree species have also adapted to this disturbance regime. Successional pathways following cyclones and fire therefore strongly depend on the extent to which forests have been historically exposed to these disturbance regimes and are dominated by “resistant” species. (pages 127 - 145)
This chapter is available at:
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- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0009
[Biodiversity, fire frequency, forest degradation, frugivores, invasive species, land-use synergisms, logging, reduced impact logging]
This chapter summarizes how tropical forests regenerate after logging and how interactions between logging operations and other disturbances can seriously derail regeneration pathways. Selective logging operations occur within an estimated 20.3% of humid tropical forests across the world. Forest logging often sets the stage for a host of other interventions and synergisms that can lead to complete deforestation, forest degradation, or conversion of forest to other land uses. The effects of logging on forest composition and the rate of post-logging regeneration are directly related to the amount of timber removed and to levels of soil and canopy damage. Logging favors the regeneration of timber species and lianas that require high light conditions for establishment. If logged forests are allowed to regenerate without further disturbance, a high fraction of local forest-requiring plant and animal species will return, and biomass and stored carbon will recover their original pre-logging levels. (pages 146 - 166)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0010
[community assembly, competition, environmental filtering, functional groups, functional traits, growth, mortality, pioneer, seed size, shade tolerance]
This chapter examines the life forms, functional traits, and functional groups of plants that predominate during different successional stages, and describes patterns and mechanisms of species turnover during succession. The high photosynthetic and growth capacity of plants that colonize early in succession enables these species to compete effectively for high levels of resource availability. But species with these “fast” traits lose their competitive edge later in succession, when establishment and survival depend more upon “slow” traits that reduce intrinsic rates of growth and enable long-term persistence in shaded understory. Changes in species composition during succession reflect a combination of initial floristic composition and relay floristics, in which species colonize sequentially in response to changing forest conditions. Tree recruitment during later stages of succession favors species that are more functionally and phylogenetically distinct than during earlier stages of succession as biotic interactions become increasingly important drivers of community assembly. (pages 167 - 195)
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- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0011
[Biomass, carbon sink, carbon stocks, hydrology, mycorrhizae, nutrient cycling, nutrient stocks, soil carbon, soil nutrients]
This chapter examines recovery of carbon and nutrient stocks and hydrological functions in regenerating tropical forests. Rates of recovery depend on many factors including soil fertility, soil texture, rainfall and seasonality, frequency of burning, initial species colonization, temperature, and site productivity. Long-term accumulation of biomass and nutrients in forest vegetation ultimately leads to the recovery and retention of carbon and nutrients in the soil and in the entire forest ecosystem. As woody stems grow in height and diameter, basal area increases and woody tissue becomes the dominant component of stand-level biomass. Growth of regenerating forests may be limited by nutrient availability on highly weathered, nutrient poor soils and in areas where soil nutrient stock have been depleted. Colonization of roots by arbuscular mycorrhizae can be important for supporting rapid growth rates of pioneer species. Regrowth of forests can rapidly restore hydrological functions, such as evapotranspiration, infiltration, and stream flow. (pages 196 - 218)
This chapter is available at:
    https://academic.oup.com/chica...

- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0012
[Frugivores, herbivory, interaction networks, mutualism, pollination, seed dispersal, seed predation]
This chapter focuses on changes in animal diversity and composition during forest regeneration and how these changes are linked to interactions between plants and animals. Recovery of animal communities is closely linked to the recovery of plant diversity. Species composition of animal taxa recovers slowly due to the gradual influx of old-growth specialists. Non-flying taxa, rare species, endemic species, and taxa with specialized dietary or nesting requirements are less likely to colonize isolated and small areas of regenerating forests during early stages. Early successional plant species support higher densities of herbivores and sustain greater amounts of herbivory than late successional species. Seed dispersal by frugivores increases during forest regeneration, along with increases in seed size and the proportion of shade-tolerant species. Dispersal limitation and seed predation decrease rates of succession, particularly in isolated pasture sites. Over 90% of the flowering plant species in tropical forests rely on animals for pollination. (pages 219 - 244)
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- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0013
[assisted regeneration, ecological restoration, ecosystem services, enriched fallows, enrichment planting, framework species, nurse species, reforestation]
This chapter focuses on approaches to active reforestation or ecological restoration in the tropics, including three case studies. Active reforestation involves deliberate human intervention aimed at overcoming specific barriers to natural forest regeneration. Reforestation projects use tree planting, soil amelioration, direct seeding, or enrichment planting to accelerate regeneration of native species and to restore natural forest cover. Pioneer tree species are key resources for active reforestation efforts, as these species are adapted to colonize, establish, and grow in conditions where vegetation has been cleared, burned, or washed away. Reforestation decisions and approaches should require active participation of local residents, whose environment and sustenance will be greatly affected by planting schemes and changes in land use. Reforestation efforts are doomed to fail if major local barriers to natural regeneration are poorly understood. Many costly restoration projects have been lost to fire, low seedling survival, and inadequate monitoring and management. (pages 245 - 273)
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- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0014
[agricultural matrix, agroforestry, biological corridor, deforestation, forest cover, forest transition, forestry policy, land-use change, landscape, remittances]
This chapter discusses how tropical forests regenerate within a larger spatial context of human-modified landscapes, where natural or semi-natural habitats coexist within a matrix of different types and extents of agricultural land uses. The spatial distribution of regenerating forests within tropical forest landscapes is not random, in large part because the distribution of prior deforestation, land use, and land abandonment are not random. Deforestation and reforestation pathways emerge from the coupled dynamics of ecological and social systems. In our globalized economy, migration, employment outsourcing, and trade relationships link the fate of forests, agricultural production, and human populations across national borders. Reforestation success in tropical developing countries is affected by socio-economic factors, institutional factors, technical factors and biophysical factors. Forest transitions can proceed as part of a deliberate plan at the local, regional, or national level or can occur as an unintended consequence of economic and political change. (pages 274 - 303)
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- Robin L. Chazdon
DOI: 10.7208/chicago/9780226118109.003.0015
[co-benefits, conservation value, ecosystem services, forestry policies, reforestation, restoration]
Forest regeneration and ecological restoration in tropical regions can have important co-benefits for the billions of people who depend on forests for their livelihoods and wellbeing. Existing old-growth forest areas should be used as leverage in an unprecedented effort to expand forest areas, to enhance biodiversity and ecosystem services on unproductive or unused lands, and to create diverse and multi-functional landscapes. As old-growth forests dwindle, prospects for forest regeneration and ecological restoration become even more challenging, expensive, and risky. The value of regenerating tropical forests for sustaining biodiversity and providing a wide range of ecosystem services is widely underappreciated. Policies regarding secondary forest management, conservation, and land-use should be guided by scientific knowledge and active stakeholder participation regarding the biophysical potential for forest regeneration and the values of biodiversity conservation and ecosystem goods and services that are provided by naturally regenerating forests and native species plantations. (pages 304 - 316)
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