Robert M. Ewers

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.

Habitat fragmentation and its lasting impact on Earth's ecosystems

Urgent need for conservation and restoration measures to improve landscape connectivity. We conducted an analysis of global forest cover to reveal that 70% of remaining forest is within 1 km of the forest’s edge, subject to the degrading effects of fragmentation. A synthesis of fragmentation experiments spanning multiple biomes and scales, five continents, and 35 years demonstrates that habitat fragmentation reduces biodiversity by 13 to 75% and impairs key ecosystem functions by decreasing biomass and altering nutrient cycles. Effects are greatest in the smallest and most isolated fragments, and they magnify with the passage of time. These findings indicate an urgent need for conservation and restoration measures to improve landscape connectivity, which will reduce extinction rates and help maintain ecosystem services.

Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis

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 worlds 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.

Functional traits, the phylogeny of function, and ecosystem service vulnerability

People depend on benefits provided by ecological systems. Understanding how these ecosystem services – and the ecosystem properties underpinning them – respond to drivers of change is therefore an urgent priority. We address this challenge through developing a novel risk-assessment framework that integrates ecological and evolutionary perspectives on functional traits to determine species’ effects on ecosystems and their tolerance of environmental changes. We define Specific Effect Function (SEF) as the per-gram or per capita capacity of a species to affect an ecosystem property, and Specific Response Function (SRF) as the ability of a species to maintain or enhance its population as the environment changes. Our risk assessment is based on the idea that the security of ecosystem services depends on how effects (SEFs) and tolerances (SRFs) of organisms – which both depend on combinations of functional traits – correlate across species and how they are arranged on the species’ phylogeny. Four extreme situations are theoretically possible, from minimum concern when SEF and SRF are neither correlated nor show a phylogenetic signal, to maximum concern when they are negatively correlated (i.e., the most important species are the least tolerant) and phylogenetically patterned (lacking independent backup). We illustrate the assessment with five case studies, involving both plant and animal examples. However, the extent to which the frequency of the four plausible outcomes, or their intermediates, apply more widely in real-world ecological systems is an open question that needs empirical evidence, and suggests a research agenda at the interface of evolutionary biology and ecosystem ecology.

SYNERGISTIC INTERACTIONS BETWEEN EDGE AND AREA EFFECTS IN A HEAVILY FRAGMENTED LANDSCAPE

Both area and edge effects have a strong influence on ecological processes in fragmented landscapes, but there is little understanding of how these two factors might interact to exacerbate local species declines. To test for synergistic interactions between area and edge effects, we sampled a diverse beetle community in a heavily fragmented landscape in New Zealand. More than 35,000 beetles of approximately 900 species were sampled over large gradients in habitat area (10(-2) 10(6) ha) and distance from patch edge (2(0)-2(10) m from the forest edge into both the forest and adjacent matrix). Using a new approach to partition variance following an ordination analysis, we found that a synergistic interaction between habitat area and distance to edge was a more important determinant of patterns in beetle community composition than direct edge or area effects alone. The strength of edge effects in beetle-species composition increased nonlinearly with increasing fragment area. One important consequence of the synergy is that the slopes of species area (SA) curves constructed from habitat islands depend sensitively on the distance from edge at which sampling is conducted. Surprisingly, we found negative SA curves for communities sampled at intermediate distances from habitat edges, caused by differential edge responses of matrix- vs. forest-specialist species in fragments of increasing area. Our data indicate that distance to habitat edge has a consistently greater impact on beetle community composition than habitat area and that variation in the strength of edge effects may underlie many patterns that are superficially related to habitat area.

Boom-and-bust development patterns across the Amazon deforestation frontier.

Boom and Bust The Brazilian Amazon is renowned for its biodiversity and for its influence on climate regulation and geochemical cycles. It is also one of the countrys poorest regions. For decades, much economic development has been pursued through conversion of forest for agriculture and cattle-ranching. Rodrigues et al. (p. 1435) investigated whether this pattern of land use brings lasting prosperity by analyzing data on the economic development of nearly 300 municipalities across the deforestation frontier. Relative development, in terms of life expectancy, literacy, and standard of living, increases as deforestation begins but then declines again as the frontier passes through. As a result, pre- and postfrontier levels of development are similarly low, indicating a pattern of boom and bust. Rainforest loss in the Amazon is associated with ephemeral increase in people’s relative prosperity. The Brazilian Amazon is globally important for biodiversity, climate, and geochemical cycles, but is also among the least developed regions in Brazil. Economic development is often pursued through forest conversion for cattle ranching and agriculture, mediated by logging. However, on the basis of an assessment of 286 municipalities in different stages of deforestation, we found a boom-and-bust pattern in levels of human development across the deforestation frontier. Relative standards of living, literacy, and life expectancy increase as deforestation begins but then decline as the frontier evolves, so that pre- and postfrontier levels of human development are similarly low. New financial incentives and policies are creating opportunities for a more sustained development trajectory that is not based on the depletion of nature and ecosystem services.

Pervasive impact of large-scale edge effects on a beetle community

Habitat edges are a ubiquitous feature of modern fragmented landscapes, but a tendency for researchers to restrict sampling designs to relatively small spatial scales means that edge effects are known to influence faunal communities over small spatial scales of only 20–250 m. However, we found striking changes in the abundance and community composition of 769 New Zealand beetle species (≈26,000 individuals) across very long edge gradients. We show that almost 90% of species respond significantly to habitat edges and that the abundances of 20% of common species were affected by edges at scales >250 m. Moreover, as many as one in eight common species had edge effects that appeared to penetrate as far as 1 km into habitat patches. Even 1 km inside forest, beetle communities differed in species richness, β-diversity (spatial turnover), and composition from the deep forest interior. Spatially explicit models of fragmented landscapes have shown that such large-scale edge effects can lead to an 80% reduction in the population size of interior forest species in even very large fragments. Moreover, such large-scale edge effects can drive species that inhabit central habitat core—which are among the most threatened species in fragmented landscapes—to local extinction from habitat fragments and protected areas. In a global analysis of protected areas, we show that kilometer-scale edge effects may compromise the ability of more than three-quarters of the worlds forested reserves to conserve the community biostructures that are unique to forest interiors.

Extinction Debt and Windows of Conservation Opportunity in the Brazilian Amazon

Growing Extinction Debt Predicting, and potentially preventing, extinction is a central goal of conservation biology. Wearn et al. (p. 228; see the Perspective by Rangel) describe a mathematical approach for predicting the time lags in extinction following habitat loss. The model was applied to the highly biodiverse Brazilian Amazon region, and used to reconstruct the spatial and temporal patterns of extinction and the accumulation of extinction debt from 1970 through to the present, and to extrapolate to 2050 under four deforestation scenarios. The Amazon basin sits at a critical point: Few species have been driven extinct to date, but an extinction debt is rapidly accumulating, which could lead to an increasing rate of extinction in the next four decades. Deforestation scenarios predict species extinction rates and identify targets of conservation efforts. Predicting when future species extinctions will occur is necessary for directing conservation investments but has proved difficult. We developed a new method for predicting extinctions over time, accounting for the timing and magnitude of habitat loss. We applied this to the Brazilian Amazon, predicting that local extinctions of forest-dependent vertebrate species have thus far been minimal (1% of species by 2008), with more than 80% of extinctions expected to be incurred from historical habitat loss still to come. Realistic deforestation scenarios suggest that local regions will lose an average of nine vertebrate species and have a further 16 committed to extinction by 2050. There is a window of opportunity to dilute the legacy of historical deforestation by concentrating conservation efforts in areas with greatest debt.

A large-scale forest fragmentation experiment: the Stability of Altered Forest Ecosystems Project

Opportunities to conduct large-scale field experiments are rare, but provide a unique opportunity to reveal the complex processes that operate within natural ecosystems. Here, we review the design of existing, large-scale forest fragmentation experiments. Based on this review, we develop a design for the Stability of Altered Forest Ecosystems (SAFE) Project, a new forest fragmentation experiment to be located in the lowland tropical forests of Borneo (Sabah, Malaysia). The SAFE Project represents an advance on existing experiments in that it: (i) allows discrimination of the effects of landscape-level forest cover from patch-level processes; (ii) is designed to facilitate the unification of a wide range of data types on ecological patterns and processes that operate over a wide range of spatial scales; (iii) has greater replication than existing experiments; (iv) incorporates an experimental manipulation of riparian corridors; and (v) embeds the experimentally fragmented landscape within a wider gradient of land-use intensity than do existing projects. The SAFE Project represents an opportunity for ecologists across disciplines to participate in a large initiative designed to generate a broad understanding of the ecological impacts of tropical forest modification.

Fragmentation impairs the microclimate buffering effect of tropical forests.

Background Tropical forest species are among the most sensitive to changing climatic conditions, and the forest they inhabit helps to buffer their microclimate from the variable climatic conditions outside the forest. However, habitat fragmentation and edge effects exposes vegetation to outside microclimatic conditions, thereby reducing the ability of the forest to buffer climatic variation. In this paper, we ask what proportion of forest in a fragmented ecosystem is impacted by altered microclimate conditions driven by edge effects, and extrapolate these results to the whole Atlantic Forest biome, one of the most disturbed biodiversity hotspots. To address these questions, we collected above and below ground temperature for a full year using temperature sensors placed in forest fragments of different sizes, and at different distances from the forest edge. Principal Findings In the Atlantic forests of Brazil, we found that the buffering effect of forests reduced maximum outside temperatures by one third or more at ground level within a forest, with the buffering effect being stronger below-ground than one metre above-ground. The temperature buffering effect of forests was, however, reduced near forest edges with the edge effect extending up to 20 m inside the forest. The heavily fragmented nature of the Brazilian Atlantic forest means that 12% of the remaining biome experiences altered microclimate conditions. Conclusions Our results add further information about the extent of edge effects in the Atlantic Forest, and we suggest that maintaining a low perimeter-to-area ratio may be a judicious method for minimizing the amount of forest area that experiences altered microclimatic conditions in this ecosystem.