Synergistic Interactions between Habitat Fragmentation and Fire in Evergreen Tropical Forests

Abstract: The growing prevalence of fragmentation and fire in tropical forests makes it imperative to quantify changes in these disturbances and to understand the ways in which they interact across the landscape. I used a multitemporal series of Landsat images to study the incidence and coincidence of fire and fragmentation in two areas of Pará state in the eastern Brazilian Amazon: Tailândia and Paragominase. In both areas, deforestation and forest fires were quantified for time series of 6–10 years. The Tailândia study area typifies a landscape with the herringbone pattern of government-settled colonists, and the Paragominas area is dominated by large cattle ranches. In both areas, over 90% of the forests affected by fire were associated with forest edges. Although most burned forest occurred within 500 m of forest edges, some fires occurred in deep forest, several kilometers from any edge. The obvious synergism between forest fragmentation and fire poses serious risks to tropical ecosystems and has important implications for land management.     

Positive Feedbacks among Forest Fragmentation, Drought, and Climate Change in the Amazon

Abstract: The Amazon basin is experiencing rapid forest loss and fragmentation. Fragmented forests are more prone than intact forests to periodic damage from El Niño–Southern Oscillation ( ENSO) droughts, which cause elevated tree mortality, increased litterfall, shifts in plant phenology, and other ecological changes, especially near forest edges. Moreover, positive feedbacks among forest loss, fragmentation, fire, and regional climate change appear increasingly likely. Deforestation reduces plant evapotranspiration, which in turn constrains regional rainfall, increasing the vulnerability of forests to fire. Forest fragments are especially vulnerable because they have dry, fire-prone edges, are logged frequently, and often are adjoined by cattle pastures, which are burned regularly. The net result is that there may be a critical "deforestation threshold" above which Amazonian rainforests can no longer be sustained, particularly in relatively seasonal areas of the basin. Global warming could exacerbate this problem if it promotes drier climates or stronger ENSO droughts. Synergisms among many simultaneous environmental changes are posing unprecedented threats to Amazonian forests.     

Landscape history, fragmentation, and patch occupancy: models for a forest bird with limited dispersal.

We developed and tested patch occupancy models for an endemic understory bird with limited dispersal ability, the Chucao Tapaculo (Scelorchilus rubecula), in two South American temperate rain forest landscapes that differed in levels and duration of forest loss. We assessed cover changes since 1961 in each landscape and surveyed patches for Chucao Tapaculo occupancy. We then developed incidence-based predictive models independently for each landscape and tested each model reciprocally in the alternative study area. We thereby assessed the domain of model applicability and identified those predictor variables with general effects and those that varied between the two landscapes. The two models were consistent regarding variable selection, and predictive accuracy of each model was high in the landscape where training data were collected. However, the models differed substantially in the magnitudes of effects related to patch size, with larger unoccupied patches observed in the landscape with the more advanced stage of fragmentation. Due to this discrepancy, each model performed poorly when applied to the alternative landscape, potentially reflecting the contrasting stages of habitat loss. Although it was impossible to dissociate effects of level and duration of forest loss, we viewed the landscapes as representing two extremes along a continuum of fragmentation, providing insights into potential trajectories for portions of the biome where deforestation is occurring. Further, our data suggest that static equilibrium models developed from distribution patterns in recently fragmented landscapes may overestimate persistence when used as a forecasting tool, or when extrapolated to alternative landscapes where fragmentation is more advanced. Thus, we suggest that landscapes used as standards for model building should be selected with caution. We recommend that distribution patterns be obtained from landscapes where fragmentation is advanced, preferably with histories of fragmentation long enough that time-delayed extinctions already would have occurred.     

Forest Management Patterns in the Floodplain of the Amazon Estuary

In the Amazon basin, few studies have focused on environmentally sound land-use alternatives that are linked to markets. This paper analyzes land uses carried out by traditional inhabitants ( ribeirinhos ) on three islands in the Amazon estuary. Management of the native floodplain forest is the most extensive form of land use on these islands. Such management varies in response to local economic and ecological conditions but invariably targets key nontimber forest resources destined for local markets, requires minimal input of labor and capital, and maintains a considerable stock of biotic resources. While restricted to a specific forest type, this form of land use is associated with relatively high population densities and, if supported by appropriate development policies, could provide an environmentally sound land-use alternative for hundreds of thousands of rural inhabitants of Amazonia.     

Contrasting Colonist and Indigenous Impacts on Amazonian Forests

Abstract: To examine differences in land use and environmental impacts between colonist and indigenous populations in the northern Ecuadorian Amazon, we combined data from household surveys and remotely sensed imagery that was collected from 778 colonist households in 64 colonization sectors, and 499 households from five indigenous groups in 36 communities. Overall, measures of deforestation and forest fragmentation were significantly greater for colonists than indigenous peoples. On average, colonist households had approximately double the area in agriculture and cash crops and 5.5 times the area in pasture as indigenous households. Nevertheless, substantial variation in land‐use patterns existed among the five indigenous groups in measures such as cattle ownership and use of hired agricultural labor. These findings support the potential conservation value of indigenous lands while cautioning against uniform policies that homogenize indigenous ethnic groups.     

Thresholds of species loss in Amazonian deforestation frontier landscapes

In the Brazilian Amazon, private land accounts for the majority of remaining native vegetation. Understanding how land‐use change affects the composition and distribution of biodiversity in farmlands is critical for improving conservation strategies in the face of rapid agricultural expansion. Working across an area exceeding 3 million ha in the southwestern state of Rondônia, we assessed how the extent and configuration of remnant forest in replicate 10,000‐ha landscapes has affected the occurrence of a suite of Amazonian mammals and birds. In each of 31 landscapes, we used field sampling and semistructured interviews with landowners to determine the presence of 28 large and medium sized mammals and birds, as well as a further 7 understory birds. We then combined results of field surveys and interviews with a probabilistic model of deforestation. We found strong evidence for a threshold response of sampled biodiversity to landscape level forest cover; landscapes with <30–40% forest cover hosted markedly fewer species. Results from field surveys and interviews yielded similar thresholds. These results imply that in partially deforested landscapes many species are susceptible to extirpation following relatively small additional reductions in forest area. In the model of deforestation by 2030 the number of 10,000‐ha landscapes under a conservative threshold of 43% forest cover almost doubled, such that only 22% of landscapes would likely to be able to sustain at least 75% of the 35 focal species we sampled. Brazilian law requires rural property owners in the Amazon to retain 80% forest cover, although this is rarely achieved. Prioritizing efforts to ensure that entire landscapes, rather than individual farms, retain at least 50% forest cover may help safeguard native biodiversity in private forest reserves in the Amazon. Umbrales de Pérdida de Especies en los Paisajes Fronterizos de Deforestación en el Amazonas Ochoa‐Quintero     

The distribution of lianas and their change in abundance in temperate forests over the past 45 years

Lianas (woody vines) are an important and dynamic component of many forests throughout the world, and increases in CO2, mean winter temperature, and forest fragmentation may promote their growth and proliferation in temperate forests. In this study, we used a 45-year data set to test the hypothesis that lianas have increased in abundance and basal area in the interiors of 14 deciduous temperate forests in Wisconsin (USA) since 1959. We also censused woody plants along a gradient from the forest edge to the interior in seven of these forests to test the hypothesis that the abundance of lianas declines significantly with increasing distance from the forest edge. We found that lianas did not increase in abundance within the interiors of temperate forests in Wisconsin over the last 45 years. However, relative and absolute liana abundance decreased sharply with increasing distance from forest edges. Our findings suggest that forest fragmentation, not climate change, may be increasing the abundance of lianas in northern deciduous temperate forests, and that lianas may further increase in abundance if the severity of forest fragmentation intensifies.     

The relative influence of habitat loss and fragmentation: do tropical mammals meet the temperate paradigm?

The relative influence of habitat loss vs. habitat fragmentation per se (the breaking apart of habitat) on species distribution and abundance is a topic of debate. Although some theoretical studies predict a strong negative effect of fragmentation, consensus from empirical studies is that habitat fragmentation has weak effects compared with habitat loss and that these effects are as likely to be positive as negative. However, few empirical investigations of this issue have been conducted on tropical or wide-ranging species that may be strongly influenced by changes in patch size and edge that occur with increasing fragmentation. We tested the relative influence of habitat loss and fragmentation by examining occupancy of forest patches by 20 mid- and large-sized Neotropical mammal species in a fragmented landscape of northern Guatemala. We related patch occupancy of mammals to measures of habitat loss and fragmentation and compared the influence of these two factors while controlling for patch-level variables. Species responded strongly to both fragmentation and loss, and response to fragmentation generally was negative. Our findings support previous assumptions that conservation of large mammals in the tropics will require conservation strategies that go beyond prevention of habitat loss to also consider forest cohesion or other aspects of landscape configuration.     

Clearance and Fragmentation of Tropical Deciduous Forest in the Tierras Bajas, Santa Cruz, Bolivia

Abstract: The Tierras Bajas is an area of 20,000 km ² of lowland deciduous forest in eastern Santa Cruz, Bolivia, that has undergone rapid change during the past two decades. As part of the largest remaining area of intact deciduous tropical forest in the world, it has been nominated a priority area for conservation by several environmental organizations. We quantified the spatial and temporal patterns of deforestation in the area by digital processing of high-resolution satellite imagery from 1975 through 1998. The estimated rate of deforestation was among the highest in the world for such a limited area, ranging from 160 km ²/year in the early1980s to almost 1200 km ²/year in the late 1990s. Although most deforestation up to 1984 was in Bolivian peasant and Mennonite colonies, most deforestation after 1984 was in non-Mennonite industrial soybean farms. The level of fragmentation of uncut forest, caused by the spatial patterns of deforestation, also differed among these broad land-use types. Deforestation in planned and spontaneous peasant colonies was complex in shape, forming relatively large areas of edge-affected forest, whereas that in Mennonite and other industrial farms was in large, rectangular increments, creating relatively less edge. But the distribution of these farms and the practice of initially clearing around the peripheries of properties resulted in the isolation of large areas of forest. In 1998 four-fifths of the remaining forest were either within 1 km of a clearance edge or in isolated fragments of <50 km². Compared with deforested areas, the areas of isolated and edge-affected forest were disproportionately large during the early stages of frontier colonization. These results imply that if the fragmentation effects of deforestation are to be minimized, conservation planning must occur at the earliest stages of frontier development.     

The Early Development of Forest Fragmentation Effects on Birds

The early development of forest fragmentation effects on forest organisms is poorly understood partly because most fragmentation studies have been done in agricultural or suburban landscapes, long after the onset of fragmentation. We develop a temporal model of forest fragmentation effects on densities of forest-breeding birds and provide data from an active industrial forest landscape to test the model. The model and our empirical data indicate that densities of several forest-dwelling bird species can increase within a forest stand soon after the onset of fragmentation as a result of displaced individuals packing into remaining habitat. Along with higher densities in the newly formed fragments, pairing success in one species, the Ovenbird ( Seiurus aurocapillus ), was lower in fragments than nonfragments, possibly due to behavioral dysfunction resulting from high densities. Thus, density was inversely related to productivity. The duration and extent of increased densities following onset of fragmentation depends on many factors, including the sensitivity of a species to edge and area effects, the duration and rate of habitat loss and fragmentation, and the proximity of a forest stand to the disturbance. Incipient forest fragmentation may affect populations differently from later stages of fragmentation when the geometry of the landscape has reached a more stable configuration. Our model and data indicate, for reasons unrelated to traditional edge effects, that large tracts of forest can be important because they are relatively free from the variety of plant and animal population dynamics that might take place near new edges, including the encroachment of individuals displaced by habitat loss.     

Effects of Forest Fragmentation and Habitat Degradation on West African Leaf-Litter Frogs

Abstract:  Habitat degradation alters the dynamics and composition of anuran assemblages in tropical forests. The effects of forest fragmentation on the composition of anuran assemblages are so far poorly known. We studied the joint influence of forest fragmentation and degradation on leaf-litter frogs. We specifically asked whether the processes structuring leaf-litter anuran assemblages in fragmented forests are the same as those in continuous forests. We analyzed anuran assemblages with respect to habitat characteristics, including fragmentation and degradation parameters. In comparison with continuous forests, species richness and diversity were lower and assemblage composition was altered in forest fragments. These changes seemed to be mainly caused by habitat degradation rather than forest fragmentation. Availability of aquatic sites for breeding, vegetation structure (including those variables indicating degradation), and leaf-litter cover had the most influence on the presence of single species. The comparatively small impact of fragmentation on anurans might be due to the location of the study area; it still possessed large tracts of continuous forest. These forest blocks may stabilize the regional rainforest climate and thus weaken the effects of fragmentation .     

Tropical Forest Fragmentation and the Local Extinction of Understory Birds in the Eastern Usambara Mountains, Tanzania

Abstract. Small tropical forest fragments in the Eastern Usambara Mountains have lost understory bird species following forest fragmentation. The local extinction of these species is documented through a comparison of species number on nine smaller fores t fragments with species number on a tenth larger control forest site. The number of species captured in the control forest site is significantly larger than the number captured in eight of the nine smaller forest fragments under an equivalent capture effort. Forest-dependent understory species vary greatly in their vulnerability to forest fragmentation. Relatively rare species and forest interior species are the avifauna most adversely affected by forest fragmentation. A concerted effort should be made to protect the existing corridors of native forest that link established forest reserves in the Eastern Usambara Mountains.     

Quantifying forest fragmentation spatial process in the developing State of Selangor,peninsular Malaysia

Forest fragmentation has several phases; thus, the ecological significance of each phase during a particular period of time must be interpreted. To interpret, this study quantifies the magnitude of forest loss and the changes in the temporal pattern of fragmentation in the State of Selangor, peninsular Malaysia. Using the decision tree model of land transformation, five phases of forest fragmentation were identified: perforation, dissection, dissipation, shrinkage and attrition. This analysis showed that the magnitude of forest loss was the highest during the dissipation phase. The patchiness analysis showed that dissipation contributes to the highest environmental uncertainty found for the forest patches. This study can be considered a first step in the exploration of the properties and the behavioural pattern shown by the spatial process of forest fragmentation.     

Understanding Trait‐Dependent Community Disassembly: Dung Beetles,Density Functions,and Forest Fragmentation

Abstract: Anthropogenic disturbances such as fragmentation are rapidly altering biodiversity, yet a lack of attention to species traits and abundance patterns has made the results of most studies difficult to generalize. We determined traits of extinction‐prone species and present a novel strategy for classifying species according to their population‐level response to a gradient of disturbance intensity. We examined the effects of forest fragmentation on dung beetle communities in an archipelago of 33 islands recently created by flooding in Venezuela. Species richness, density, and biomass all declined sharply with decreasing island area and increasing island isolation. Species richness was highly nested, indicating that local extinctions occurred nonrandomly. The most sensitive dung beetle species appeared to require at least 85 ha of forest, more than many large vertebrates. Extinction‐prone species were either large‐bodied, forest specialists, or uncommon. These explanatory variables were unrelated, suggesting at least 3 underlying causes of extirpation. Large species showed high wing loading (body mass/wing area) and a distinct flight strategy that may increase their area requirements. Although forest specificity made most species sensitive to fragmentation, a few persistent habitat generalists dispersed across the matrix. Density functions classified species into 4 response groups on the basis of their change in density with decreasing species richness. Sensitive and persistent species both declined with increasing fragmentation intensity, but persistent species occurred on more islands, which may be due to their higher baseline densities. Compensatory species increased in abundance following the initial loss of sensitive species, but rapidly declined with increasing fragmentation. Supertramp species (widespread habitat generalists) may be poor competitors but strong dispersers; their abundance peaked following the decline of the other 3 groups. Nevertheless, even the least sensitive species were extirpated or rare on the smallest and most isolated islands.     

Contribution of Roads to Forest Fragmentation in the Rocky Mountains

The contribution of roads to forest fragmentation has not been adequately analyzed. We quantified fragmentation due to roads in a 30,213-ha section of the Medicine Bow-Routt National Forest in sout heastern Wyoming with several indices of landscape structure using a geographic information system. The number of patches, mean patch area, mean interior area, mean area of edge influence, mean patch perimeter, total perimeter, and mean patch shape identified patch- and edge-related landscape changes. Shannon-Wiener diversity, dominance, contagion, contrast, and angular second moment indicated effects on landscape diversity and texture. Roads added to forest fragmentation more than clearcuts by dissecting large patches into smaller pieces and by converting forest interior habitat into edge habitat. Edge habitat created by roads was 1.54–1.98 times the edge habitat created by clearcuts. The total landscape area affected by clearcuts and roads was 2.5–3.5 times the actual area occupied by these disturbances. Fragmentation due to roads could be minimized if road construction is minimized or rerouted so that its fragmentation effects are reduced. Geographic information system technology can be used to quantify the potential fragmentation effects of individual roads and the cumulative effects of a road network on landscape structure.     

Sustainable Use of the Tropical Rain Forest: Evidence from the Avifauna in a Shifting-Cultivation Habitat Mosaic in the Colombian Amazon

We carried out a one-year study of the avifauna along a regeneration gradient created by indigenous shifting cultivation in the Colombian Amazon rain forest. Regenerating one-hectare crop fields, from 1 to 17 years after abandonment, and undisturbed understory sites were compared in a sample effort of 4323 net hours. We made 878 captures of 103 species belonging to 11 trophic groups. Differences in species richness and number of individuals captured between regenerating areas and primary forest were low. A comparison of similarity indices among the sites for each habitat type showed no consistent differences, suggesting similar levels of local heterogeneity. Similarity indices calculated between sites of different-aged habitats, however, showed that young secondary growth at less than 10 years of abandonment was least similar to other sites, whereas old secondary growth between 13 to 17 years of abandonment, and forest understories, had the most similar avifauna. Two trophic groups were absent from the samples of the forest interior, and the abundance of the other nine varied among sites of regeneration. We conclude that slash and burn agriculture represents a high-intensity, small-magnitude disturbance to the forest avifauna. Large changes occur when the crop field is established and last until about 10 years after abandonment. Our data suggest that abandoned crop fields older than this age mimic the natural patch dynamics of the forest, making this type of agriculture an example of a potentially sustainable use of tropical forests, at least from the standpoint of its impact on bird communities.     

Effects of small-scale habitat fragmentation on predator–prey interactions in a temperate sea grass system

During the last decades, fragmentation has become an important issue in ecological research. Habitat fragmentation operates on spatial scales ranging over several magnitudes from patches to landscapes. We focus on small-scale fragmentation effects relevant to animal foraging decision making that could ultimately generate distribution patterns. In a controlled experimental environment, we tested small-scale fragmentation effects in artificial sea grass on the feeding behaviour of juvenile cod (Gadus morhua). Moreover, we examined the influence of fragmentation on the distribution of one of the juvenile cod’s main prey resources, the grass shrimp (Palaemon elegans), in association with three levels of risk provided by cod (no cod, cod chemical cues and actively foraging cod). Time spent by cod within sea grass was lower in fragmented landscapes, but total shrimp consumption was not affected. Shrimp utilised vegetation to a greater extent in fragmented treatments in combination with active predation. We suggest that shrimp choose between sand and vegetation habitats to minimize risk of predation according to cod habitat-specific foraging capacities, while cod aim to maximize prey-dependent foraging rates, generating a habitat-choice game between predator and prey. Moreover, aggregating behaviour in grass shrimp was only found in treatments with active predation. Hence, we argue that both aggregation and vegetation use are anti-predator defence strategies applied by shrimp. We therefore stress the importance of considering small-scale behavioural mechanisms when evaluating consequences from habitat fragmentation on trophic processes in coastal environments.     

Deforestation in Brazilian Amazonia: History, Rates, and Consequences

Abstract:  Brazil's Amazon forest remained largely intact until the "modern" era of deforestation began with the inauguration of the Transamazon Highway in 1970. Amazonian deforestation rates have trended upward since 1991, with clearing proceeding at a variable but rapid pace. Although Amazonian forests are cut for various reasons, cattle ranching predominates. The large and medium-sized ranches account for about 70% of clearing activity. Profit from beef cattle is only one of the income sources that make deforestation profitable. Forest degradation results from logging, ground fires (facilitated by logging), and the effects of fragmentation and edge formation. Degradation contributes to forest loss. The impacts of deforestation include loss of biodiversity, reduced water cycling (and rainfall), and contributions to global warming. Strategies to slow deforestation include repression through licensing procedures, monitoring, and fines. The severity of penalties for deforestation needs to be sufficient to deter illegal clearing but not so great as to be unenforceable. Policy reform is also needed to address root causes of deforestation, including the role of clearing in establishing land claims.     

Bird Extinctions in Atlantic Forest Fragments of the Viçosa Region, Southeastern Brazil

Abstract:   We studied the conservation status of Atlantic forest birds in 43 forest fragments ranging in size from 1 to 384 ha in the Viçosa region of southeastern Brazil. We compared data from 15 years of field work with historical records from the region, mainly originating from specimens collected by João Moojen during the 1930s. We used published studies associated with museum data and current field work to assess the decline of forest birds during the last 70 years and to relate their disappearance to forest fragmentation and destruction. At least 28 bird species have become locally extinct, 43 are critically endangered, and 25 are vulnerable, representing 60.7% of the original forest bird community known to exist in the region. Vulnerability to fragmentation differed among guilds, forest strata, and endemicity status. Birds that feed on fruit and seeds, and those that feed on insects, were more threatened than omnivores and carnivores. Nectarivorous species were less threatened than other guilds. Moreover, terrestrial and understory birds or birds using only one forest stratum also were more likely to have been threatened. Finally, Atlantic forest endemics were more likely to have become extinct than nonendemic species. In general, sensitivity to environmental disturbance at the local level was similar to the predicted vulnerability to regional disturbance derived from the literature. Our results indicate that a serious decline of Atlantic forest birds is underway and that many other species of birds, not previously recognized as threatened, are of conservation concern .     

Independent effects of fragmentation on forest songbirds: an organism-based approach.

The degree to which spatial patterns influence the dynamics and distribution of populations is a central question in ecology. This question is even more pressing in the context of rapid habitat loss and fragmentation, which threaten global biodiversity. However, the relative influence of habitat loss and landscape fragmentation, the spatial patterning of remaining habitat, remains unclear. If landscape pattern affects population size, managers may be able to design landscapes that mitigate habitat loss. We present the results of a mensurative experiment designed to test four habitat loss vs. fragmentation hypotheses. Unlike previous studies, we measured landscape structure using quantitative, spatially explicit habitat distribution models previously developed for two species: Blackburnian Warbler (Dendroica fusca) and Ovenbird (Seiurus aurocapilla). We used a stratified sampling design that reduced the confounding of habitat amount and fragmentation variables. Occurrence and reoccurrence of both species were strongly influenced by characteristics at scales greater than the individual territory, indicating little support for the random-sample hypothesis. However, the type and spatial extent of landscape influence differed. Both occurrence and reoccurrence of Blackburnian Warblers were influenced by the amount of poor-quality matrix at 300- and 2000-m spatial extents. The occurrence and reoccurrence of Ovenbirds depended on a landscape pattern variable, patch size, but only in cases when patches were isolated. These results support the hypothesis that landscape pattern is important for some species only when the amount of suitable habitat is low. Although theoretical models have predicted such an interaction between landscape fragmentation and composition, to our knowledge this is the first study to report empirical evidence of such nonlinear fragmentation effects. Defining landscapes quantitatively from an organism-based perspective may increase power to detect fragmentation effects, particularly in forest mosaics where boundaries between patches and matrix are ambiguous. Our results indicate that manipulating landscape pattern may reduce negative impacts of habitat loss for Ovenbird, but not Blackburnian Warbler. We emphasize that most variance in the occurrence of both species was explained by local scale or landscape composition variables rather than variables reflecting landscape pattern.