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      Habitat Fragmentation, Variable Edge Effects, and the Landscape-Divergence Hypothesis

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          Abstract

          Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This ‘landscape-divergence hypothesis’, if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.

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          Most cited references109

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          Confounding factors in the detection of species responses to habitat fragmentation.

          Habitat loss has pervasive and disruptive impacts on biodiversity in habitat remnants. The magnitude of the ecological impacts of habitat loss can be exacerbated by the spatial arrangement -- or fragmentation -- of remaining habitat. Fragmentation per se is a landscape-level phenomenon in which species that survive in habitat remnants are confronted with a modified environment of reduced area, increased isolation and novel ecological boundaries. The implications of this for individual organisms are many and varied, because species with differing life history strategies are differentially affected by habitat fragmentation. Here, we review the extensive literature on species responses to habitat fragmentation, and detail the numerous ways in which confounding factors have either masked the detection, or prevented the manifestation, of predicted fragmentation effects. Large numbers of empirical studies continue to document changes in species richness with decreasing habitat area, with positive, negative and no relationships regularly reported. The debate surrounding such widely contrasting results is beginning to be resolved by findings that the expected positive species-area relationship can be masked by matrix-derived spatial subsidies of resources to fragment-dwelling species and by the invasion of matrix-dwelling species into habitat edges. Significant advances have been made recently in our understanding of how species interactions are altered at habitat edges as a result of these changes. Interestingly, changes in biotic and abiotic parameters at edges also make ecological processes more variable than in habitat interiors. Individuals are more likely to encounter habitat edges in fragments with convoluted shapes, leading to increased turnover and variability in population size than in fragments that are compact in shape. Habitat isolation in both space and time disrupts species distribution patterns, with consequent effects on metapopulation dynamics and the genetic structure of fragment-dwelling populations. Again, the matrix habitat is a strong determinant of fragmentation effects within remnants because of its role in regulating dispersal and dispersal-related mortality, the provision of spatial subsidies and the potential mediation of edge-related microclimatic gradients. We show that confounding factors can mask many fragmentation effects. For instance, there are multiple ways in which species traits like trophic level, dispersal ability and degree of habitat specialisation influence species-level responses. The temporal scale of investigation may have a strong influence on the results of a study, with short-term crowding effects eventually giving way to long-term extinction debts. Moreover, many fragmentation effects like changes in genetic, morphological or behavioural traits of species require time to appear. By contrast, synergistic interactions of fragmentation with climate change, human-altered disturbance regimes, species interactions and other drivers of population decline may magnify the impacts of fragmentation. To conclude, we emphasise that anthropogenic fragmentation is a recent phenomenon in evolutionary time and suggest that the final, long-term impacts of habitat fragmentation may not yet have shown themselves.
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            Selective logging in the Brazilian Amazon.

            Amazon deforestation has been measured by remote sensing for three decades. In comparison, selective logging has been mostly invisible to satellites. We developed a large-scale, high-resolution, automated remote-sensing analysis of selective logging in the top five timber-producing states of the Brazilian Amazon. Logged areas ranged from 12,075 to 19,823 square kilometers per year (+/-14%) between 1999 and 2002, equivalent to 60 to 123% of previously reported deforestation area. Up to 1200 square kilometers per year of logging were observed on conservation lands. Each year, 27 million to 50 million cubic meters of wood were extracted, and a gross flux of approximately 0.1 billion metric tons of carbon was destined for release to the atmosphere by logging.
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              Positive feedbacks in the fire dynamic of closed canopy tropical forests

              The incidence and importance of fire in the Amazon have increased substantially during the past decade, but the effects of this disturbance force are still poorly understood. The forest fire dynamics in two regions of the eastern Amazon were studied. Accidental fires have affected nearly 50 percent of the remaining forests and have caused more deforestation than has intentional clearing in recent years. Forest fires create positive feedbacks in future fire susceptibility, fuel loading, and fire intensity. Unless current land use and fire use practices are changed, fire has the potential to transform large areas of tropical forest into scrub or savanna.
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                Author and article information

                Contributors
                Role: Academic Editor
                Journal
                PLoS ONE
                plos
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2007
                10 October 2007
                : 2
                : 10
                : e1017
                Affiliations
                [1 ]Smithsonian Tropical Research Institute, Balboa, Republic of Panama
                [2 ]Biological Dynamics of Forest Fragments Project, National Institute for Amazonian Research (INPA), Smithsonian Tropical Research Institute, Manaus, Brazil
                [3 ]Institute of Zoology, Zoological Society of London, London, United Kingdom
                [4 ]Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana, United States of America
                [5 ]Department of Botany, National Institute for Amazonian Research (INPA), Manaus, Brazil
                University of Kent, United Kingdom
                Author notes
                * To whom correspondence should be addressed. E-mail: laurancew@ 123456si.edu

                Conceived and designed the experiments: WL. Performed the experiments: WL SL. Analyzed the data: WL. Contributed reagents/materials/analysis tools: HN SL AA KH JR RL. Wrote the paper: WL. Other: Helped to conceptualize findings: RE.

                Article
                07-PONE-RA-01733R1
                10.1371/journal.pone.0001017
                1995757
                17925865
                b80e427d-9541-4e75-a25c-0d1a64619363
                This is an open-access article distributed under the terms of the Creative Commons Public Domain declaration which stipulates that, once placed in the public domain, this work may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose.
                History
                : 12 July 2007
                : 21 September 2007
                Page count
                Pages: 8
                Categories
                Research Article
                Ecology/Community Ecology and Biodiversity
                Ecology/Conservation and Restoration Ecology
                Ecology/Spatial and Landscape Ecology

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