Body size, niche breadth, and ecologically scaled responses to habitat fragmentation: mammalian predators in an agricultural landscape

The ability to make a priori assessments of a species' response to fragmentation, based on its distribution in the landscape, would serve as a valuable conservation and management tool. During 1997-1999, we monitored 717 scent stations to examine seasonal use of forest patches, corridors, and crop fields by coyotes (Canis latrans), domestic cats (Felis catus), foxes (Vulpes vulpes and Urocyon cinereoargenteus), raccoons (Procyon lotor), striped skunks (Mephitis mephitis), opossums (Didelphis virginiana), and long-tailed weasels (Mustela frenata). For each species we developed landscape-based ecologically scaled landscape indices (ELSI), and we modeled species spatial distribution across three spatial scales (landscape-level, element-level, and local habitat-level). Our results suggest that these predators view landscape fragmentation at different spatial scales and demonstrate strong interspecific differences in their response to elements of the landscape. All species except coyotes and domestic cats avoided agricultural fields. In general, predator species that were more mobile (i.e. high ESLI for landscape connectivity; coyotes) were characterized by landscape- and element-based logistic models. In contrast, models including local habitat features generally were most appropriate for less mobile or more stenophagous predators (e.g. long-tailed weasels). Our analysis extends the application of the ESLI concept to species assemblages that do not appear to function as metapopulations, and it highlights the importance of examining spatial scale and species-specific responses to habitat fragmentation. We discuss the relevance of these findings for defining ecological landscapes, understanding predator-prey interactions at multiple spatial scales, and conserving predator and prey populations in fragmented landscapes.     

Bird conservation in fragmented Mediterranean forests of Spain: effects of geographical location, habitat and landscape degradation

The effects of habitat fragmentation on forest bird assemblages were analysed in 214 holm oak (Quercus ilex) remnants spread across the northern and southern plateaux of central Spain. Bird richness was highly dependent on fragment area for all species regardless of isolation, and barely affected by habitat traits. Geographical location was associated with high differences in richness of bird assemblages, which included 17 species exclusive to northern remnants and one exclusive to southern remnants. This supports the hypothesis that habitat suitability deteriorates sharply from north to south for forest birds in Spain. The species-area relationships of bird assemblages sampled in fragmented forests along a broad continental gradient (from Norway to southern Spain) showed that true forest birds only nest in woodlands >100 ha in southern Spain, whereas the full complement of forest species occurs in much smaller fragments in central-western Europe. Loss of species that are particularly sensitive to habitat fragmentation accounts for these differences between dry Spanish and mesic European woodlands. These results are explained by the low habitat suitability of Spanish woodlands, associated with the restrictive conditions for plant regeneration in the Mediterranean climate and long-standing human usage. There is, therefore, a particular need to develop management strategies that conserve birds, and probably other forest organisms, in Mediterranean regions by preventing habitat deterioration and decreases in fragment size, and by conserving all woods >100 ha.     

Bats in a fragmented landscape: Species composition, diversity and habitat interactions in savannas of Santarém, Central Amazonia, Brazil

Species response to forest fragmentation may strongly vary according to ecological requirements, shape, spatial configuration and connectivity of fragments, and the structure of the surrounding matrix. Bats are natural candidates for studies on forest fragmentation. However, the conclusions of such studies are often contradictory. We present the results of a study on the effects of forest fragmentation on bat communities in Alter do Chão, Santarém area, in Central Amazonia, Brazil, an area composed by forests and fragments surrounded by savanna-like vegetation. We compared the species composition between habitats, tested the effect of size, shape and density of trees of forested areas on the presence species and their relative frequency of captures, and investigated the savannas as an ecological barrier for the dispersion of bats. With an effort of 5678 mistnet-hours, we captured 3740 bats (64 species). Multidimensional Scaling indicated no strong separation between the habitats sampled, however, savanna sites were grouped distinctly of fragments and forest sites. Multiple linear regressions indicated no significant correlation between the number of bat species recorded and the size, shape or tree density in forest sites and fragments. There was a significant correlation between the number of captures and the variables tested, explained by the shape of the sites, but not by size or tree density. In general, sites with bigger shape indices had fewer captures. Ten of the 20 most captured species showed no significant difference between the three habitats. The bat communities in Alter do Chão were not strongly affected by forest fragmentation, and the savannas did not appear to act as an ecological barrier to bats.     

Wader nest distribution and predation in relation to man-made structures on coastal pastures

Coastal pastures and other wet grasslands are important but decreasing breeding habitats for many waders (Charadrii). Since loss of suitable habitat is a major reason for population declines, protection and restoration of these habitats is crucial. Reduction of the often high rate of nest predation is a potentially important tool in future conservation work. Here, we focus on predators’ use of raised structures in the landscape when searching for prey. Hooded crows (Corvus corone cornix) use man-made structures such as stone walls and barbed wire fences when foraging on coastal pastures in SW Sweden. However, few studies have examined wader breeding success in relation to man-made structures, and the extent to which such structures are used by searching nest predators. We measured the spatial distribution and rate of predation on wader nests in relation to such structures. Crows spent more time at or near man-made structures than expected by chance, but we found no significant difference in nest predation relative to distance from man-made structures. However, wader nests were placed farther away from man-made structures than expected by chance in two out of three years. Waders thus tend to avoid breeding close to man-made structures, which therefore reduce the suitable breeding area and probably also the local wader population size.     

Amphibian distribution in a traditionally managed rural landscape of Eastern Europe: Probing the effect of landscape composition

A massive decline of biodiversity is caused by land-use changes. Efforts must therefore be made to better understand the factors that govern organismal distribution, especially for countries where traditional management is about to be intensified such as in Romania. We here document the spatial distribution of amphibians from a Romanian rural landscape where land-use is still largely traditional. We related the occurrence of nine amphibian species and species richness to measures of composition and configuration of the landscape surrounding 54 ponds at three spatial scales: circular areas of 400, 600 and 800 m radii. Busy roads most severely impacted single species and amphibian richness whereas landscape composition measures, such as cover of urban areas, agricultural areas, pastures, forests and wetlands were of little importance. We suggest that the relative unimportance of landscape compositional measures on amphibians is a consequence of the traditional management of these landscapes that keep the environmental conditions favorable for most species.     

Do regional gradients in land-use influence richness, composition and turnover of bird assemblages in small woods?

Small patches of natural or semi-natural habitat have an important role in the conservation of biodiversity in human-dominated environments. The values of such areas are determined by attributes of the patch as well as its context in the surrounding land mosaic. There is a need for better understanding of the ways in which assemblages are influenced by patch context and the scale over which this occurs. Here we examine the influence of regional environmental gradients on the richness, annual turnover and composition of breeding bird species in small woods in south-eastern England. Regional gradients were defined independently of woods by an ordination of attributes for 5 km × 5 km landscape units across a 2100 km² region. Patch-level attributes, particularly area, were the most important predictors for most bird variables. For woodland migrants and woodland-dependent species, variables representing the context of each wood, either at a local or regional scale, explained significant additional variance in species richness after accounting for wood area, but did not do so for species turnover. Significant context effects for woodland-dependent species related to the extent of hedges and woodland cover in the local vicinity (<1 km radius), whereas for woodland seasonal migrants the best predictors of richness after patch area were two regional environmental gradients. The initial cue to settlement for migrants may be at a coarse regional scale, with selection for suitable landscapes that have a greater extent of woodland cover. Edge species showed different responses: they were influenced by the diversity of structural features in woods, and were a more-dominant component of the avifauna in isolated woods in open fenland environments of the region. Significant relationships between coarse regional gradients (25 km² units) and bird assemblages in small woods (0.5-30 ha) suggest that population and community processes in the avifauna operate across a broader scale than local patch neighbourhoods. They also highlight the importance of adopting a landscape or regional perspective on potential changes to land-use in rural environments, and on the conservation management of small reserves.     

Habitat fragmentation in marine landscapes: relative effects of habitat cover and configuration on juvenile crab survival in California and North Carolina seagrass beds

Processes that dictate faunal abundance (e.g. predation) may be linked primarily to the configuration of habitat (e.g. patch size, patch isolation, proximity to edge) or simply to the amount of habitat (i.e. habitat cover) in marine and terrestrial systems. I asked whether juvenile crab survival in marine seagrass habitats is linked to habitat cover or to habitat configuration, and whether relationships between crab survival and habitat features were similar in two widely separated seagrass landscapes (Bodega Harbor, California and Back Sound, North Carolina, USA). I also evaluated the independent effects of seagrass structural complexity (shoot density and shoot biomass per unit area) and habitat fragmentation on crab survival. Juvenile red rock crabs (Cancer productus: 1+year class) were tethered in Bodega Harbor and juvenile blue crabs (Callinectes sapidus: 0+year class) were tethered in Back Sound. Seagrass cover in both landscapes ranged from <10 to 100% over 10×10 m areas. Crab survival was correlated with elements of landscape structure and structural complexity in both landscapes, but relationships between crab survival and specific habitat features differed between the two landscapes. In California, juvenile red rock crab survival was negatively correlated with seagrass cover and was positively correlated with seagrass shoot density, and was marginally (P=0.06) negatively correlated with seagrass shoot biomass. In North Carolina, juvenile blue crab survival was positively correlated with log₁₀ patch area and was negatively correlated with seagrass shoot biomass. The results indicate that (1) both seagrass cover and configuration may influence crab survival; (2) seagrass structural complexity influences crab survival independently of landscape structure; (3) the relative influence of cover, configuration and structural complexity on survival likely differ among seagrass landscapes; and (4) increasing structural complexity (e.g. shoot biomass) may not always lead to increased faunal survival. The results correspond to those of recent studies in marine and terrestrial landscapes showing widely variable faunal responses to landscape structure among species, geographic location and through time, and imply that seagrass restoration and conservation strategies should incorporate species-specific responses to habitat structure at multiple scales.     

Effects of habitat loss and fragmentation on amphibians: A review and prospectus

Habitat loss and fragmentation are among the largest threats to amphibian populations. However, most studies have not provided clear insights into their population-level implications. There is a critical need to investigate the mechanisms that underlie patterns of distribution and abundance. In order to understand the population- and species-level implications of habitat loss and fragmentation, it is necessary to move from site-specific inferences to assessments of how the influences of multiple factors interact across extensive landscapes to influence population size and population connectivity. The goal of this paper is to summarize the state of knowledge, identify information gaps and suggest research approaches to provide reliable knowledge and effective conservation of amphibians in landscapes experiencing habitat loss and fragmentation. Reliable inferences require attention to species-specific ecological characteristics and their interactions with environmental conditions at a range of spatial scales. Habitat connectivity appears to play a key role in regional viability of amphibian populations. In amphibians, population connectivity is predominantly effected through juvenile dispersal. The preponderance of evidence suggests that the short-term impact of habitat loss and fragmentation increases with dispersal ability. However, species with limited dispersal abilities are likely to be equally imperiled by habitat loss and fragmentation over longer time periods. Rigorous understanding of the effects of habitat loss and fragmentation on amphibians will require species-specific, multi-scale, mechanistic investigations, and will be benefit from integrating large empirical field studies with molecular genetics and simulation modeling. Molecular genetic methods are particularly suited to quantifying the influences of habitat structure across large spatial extents on gene flow and population connectivity. Conservation strategies would benefit by moving from generalizations to species and process specific recommendations and by moving from site-specific actions to implementing conservation plans at multiple scales across broad landscapes.     

Small mammals in agricultural landscapes of Prince Edward Island (Canada): Effects of habitat characteristics at three different spatial scales

We examined the influence of habitat characteristics at the microhabitat, macrohabitat, and landscape spatial scales on small mammals occurring in 12 forest patches within four agricultural landscapes of Prince Edward Island (Canada). Landscape features were important determinants of small mammal variables at all levels, but especially at the community level, whereas microhabitat characteristics tended to influence small mammals at the population level. Macrohabitat characteristics had only minor effects on small mammals occurring in our study sites. Species richness was most strongly influenced by patch area, reaching a threshold at forest patches of roughly 8-10 ha. The proportions of both forest and hedgerow cover within 400 m from the study site were also significant determinants of small mammals species diversity, possibly reflecting their ability to perceive suitable habitats, forage in areas outside the forest patches, and/or disperse in agricultural landscapes. At least one small mammal species (Napaeozapus insignis) benefitted from the presence of agricultural fields at distances up to 1000 m. Tamias striatus benefitted from the presence of hedgerow cover within 400 m from forest patches, possibly allowing them to move between forest patches. Clearly, the maintenance of forest patches of 8-10 ha and of forest cover within 400 m from them is fundamental for the conservation of small mammals inhabiting agricultural landscapes on the Island. Conservation strategies should also consider the establishment of more effective regulations to prevent and/or reduce hedgerow removal on Prince Edward Island.     

Predation of artificial nests in a fragmented landscape in the tropical region of Los Tuxtlas, Mexico

Predation rates of artificial nests were investigated in a fragmented landscape in the lowlands of Los Tuxtlas in southern Mexico. Hen and plasticine eggs were used to assess predation pressure in four habitats: the interior of forest fragments, the forest-pasture edge, corridors of residual forest vegetation and linear strips of live fences across pastures. Three sites per habitat were used in three experimental trials. Hen and plasticine ground nests with three eggs each were alternated every 50 m along transects at each site. Predation rates on each type of nest were monitored for 9 days. Survey of potential avian and mammalian potential nest predators were conducted at each site prior to the experimental trails. Readings of amount of light illuminating the ground were taken by each nest at each site to assess exposure of nests. In general, average predation rates were significantly higher for both hen and plasticine nests in the forest-pasture edge and in the corridors than in the interior of the forest fragments. While birds and mammals were the principal predators on hen eggs in the forests, mammals were responsible for the majority (?70%) of eggs damaged at the other habitats. Surveys of potential nest predators showed that avian and mammalian potential nest predators were significantly more common at the forest-pasture edges and at the other habitats than in the forests. Readings of light reaching the ground suggest that concealment of nests by the vegetation may play an important role in predation risk. Our results are consistent with reports from other Neotropical rainforests indicating an increase of artificial nest predation pressures from forest interior to open habitats. Restoration of forest fragments, allowing the vegetation to grow along the forest-pasture edge and the planting of arboreal crops at the forest-pasture edges may be measures that could increase cover and nest protection.     

Effect of adjacent agricultural habitat on the distribution of passerines in natural grasslands

Habitat heterogeneity has often been acknowledged as a key factor determining which species occur in a landscape, but we still know little on the nature and relative importance of the processes controlling species distribution and abundance within complex landscape mosaics. We used a framework defined by the compensation, supplementation and complementation models to study the distribution of four species of passerines typical of steppe habitats and identify the degree to which landscape context accounts for their distribution in the steppe-agriculture mosaic of the Crau, southern France. Steppe habitat was the best predictor of species distribution in all cases and that densities were highest in steppe. One of these species, the Calandra lark, is strictly tied to the natural steppe habitats independently of its context. For another species, the Tawny pipit, we document a significant positive effect of the presence, near steppe habitat, of extensive pastoral habitats, such as fallows and grazed crops. Finally, the abundance of the two remaining species, Skylark and Short-toed lark, was negatively affected by the presence of non-herbaceous habitats in the vicinity of steppe habitat. This suggests that the quality of the steppe habitat, as perceived by these species is conditional to the nature of adjacent habitats. Our results suggest that the management of the context in which steppe habitat is imbedded will significantly affect the distribution and abundance of this group of passerines and determine both habitat quality and conservation value at a landscape scale.     

Effects of habitat degradation on the abundance, richness and diversity of raptors across Neotropical biomes

Population growth and human development result in biodiversity loss and biological homogenization not only in developed countries, but increasingly in the less developed countries as well. In those countries, where urbanization and agricultural intensification occur at a faster rate than in developed countries, habitat degradation appears to be the leading cause of wildlife loss. During the breeding seasons of 2002–2005 we conducted road surveys across five biomes of Argentina to detect variations in raptor community attributes as potential indicators of broad scale habitat degradation. Abundance of individuals, richness and diversity of species were calculated to assess the effects of habitat transformation and patch size on these community attributes. Raptor communities strongly varied in relation to habitat transformations, with lower abundance of individuals, richness and diversity of species in more transformed landscapes. Small patches of natural vegetation and locations in which natural and cultivated lands where interspersed showed lower richness and diversity of raptors than large patches. Fragmentation was the main cause of reductions in abundance of individuals. Although the relative contribution of our two estimates of habitat degradation to abundance, richness and diversity of raptors varied among biomes, these community attributes proved useful as predictors of habitat degradation. This was especially true in habitats where raptor communities are more complex although overall patterns remained constant across biomes, from forests to deserts. Taking into account current trends of habitat transformation (drastic increments in monocultures, urban areas, and habitat patchiness), the conservation of raptor communities in these biomes could be seriously compromised. In terms of species-specific responses of raptors to habitat degradation, a rapid process of homogenization can be expected, resulting in only a few winner species within a general scenario of losers.     

Effects of fragmentation on artificial nest predation in a tropical forest in Kenya

Nest predation has been established as a leading source of reproductive failure in songbirds. Numerous studies, conducted primarily in the Northern Hemisphere, have linked high rates of nest predation with fragmentation and habitat edges. To date, virtually no such studies have been conducted in Tropical Africa. Here we investigate the effects of disturbance level and edge on depredation of artificial avian nests in six fragments of Kakamega Forest, Kenya. We found that nest losses were higher at the edge than in the interior as often reported in previous studies. Surprisingly, nest predation was highest in the least disturbed fragments. One possible explanation for these findings is that top predators are missing from Kakamega Forest, allowing nest predator populations to increase in all fragments, while in the highly disturbed fragments, domestic dogs might be surrogates for top predators. Alternatively, human activity may itself lead to a decline in mesopredator populations.     

Partitioning of molecular variation at local spatial scales in the vulnerable neotropical freshwater turtle, Hydromedusa maximiliani (Testudines, Chelidae): implications for the conservation of aquatic organisms in natural hierarchical systems

Hydromedusa maximiliani is a vulnerable freshwater turtle endemic to mountainous regions of the Atlantic rainforest in southeastern Brazil. Random amplified polymorphic DNA (RAPD) were surveyed with the purpose of assessing the genetic structure and determining the partitioning of molecular variation in H. maximiliani across the natural spatial hierarchical scale of its habitat. The goal of the study was to integrate ecological data with estimates of molecular genetics diversity to develop strategies for the conservation of this freshwater turtle. Specimens were sampled from rivers and streams across three drainages. Nine of the 80 primers used generated 27 scoreable bands of which 10 (37%) were polymorphic and produced 16 RAPD phenotypes. Significant heterogeneity was found in the distribution of RAPD molecular phenotypes across the three drainages. Analysis of molecular variance for molecular phenotypes showed that the heterogeneity had a spatial structure since a significant amount (22%) of the total variance was attributable to variation among rivers and streams. Since the genetic variation of this turtle seems to be structured according to the natural hierarchical system of rivers and streams within drainages, it is suggested that local populations should be considered as separate management units.     

Forest fragment size and microhabitat effects on palm seed predation

The establishment of plant species depends crucially on where the seeds are deposited. However, since most studies have been conducted in continuous forests, not much is known about the effects of forest fragmentation on the maintenance of abiotic and biotic characteristics in microhabitats and their effects on seed survival. In this study, we evaluated the effects of forest fragmentation on the predation upon the seeds of the palm Syagrus romanzoffiana in three microhabitats (interior forest, forest edge and gaps) in eight fragments of semi-deciduous Atlantic forest ranging in size from 9.5 ha to 33,845 ha in southeastern Brazil. Specifically, we examined the influence of the microhabitat structure, fauna and fragment size on the pattern of seed predation. Fragments <100 ha showed similar abiotic and biotic characteristics to those of the forest edge, with no seed predation in these areas. Forest fragments 230-380 ha in size did not present “safe sites” for S. romanzoffiana seed survival and showed high seed predation intensity in all microhabitats evaluated. In fragments larger than 1000 ha, the seed predation was lower, with abiotic and biotic differences among gaps, interior forests and forest edges. In these fragments, the survival of S. romanzoffiana seeds was related to squirrel abundance and interior forest maintenance. Based on these results, we concluded that there are no safe sites for S. romanzoffiana seed establishment in medium- and small-sized fragments as result of the biotic and abiotic pressure, respectively. We suggest that on these forest fragments, management plans are needed for the establishment of S. romanzoffiana, such as interior forest improvement and development in small-sized sites in order to minimize the edge effects, and on medium-sized fragments, we suggest post-dispersal seed protection in order to avoid seed predation by vertebrates. Our findings also stress the importance of assessing the influence of forest fragmentation on angiosperm reproductive biology as part of the effective planning for the management of fragmented areas.     

Patch and landscape characteristics associated with the distribution of woodland amphibians in an agricultural fragmented landscape: an information-theoretic approach

In the Midwestern United States, agricultural landscapes with scattered patches of fragmented forest are common. To investigate the relationship between amphibian distributions and wetland, woodlot, and landscape characteristics, we studied the pond-breeding amphibians within a 15,450-ha plot in rural north-central Ohio. We surveyed 25 woodlots and one area of continuous riparian forest for amphibians, and each surveyed woodland contained at least one temporary wetland. We used Akaike's Information Criterion (AIC) to evaluate the effectiveness of 13 a priori models in predicting total amphibian species richness, anuran richness, caudate richness, and the presence of individual species in woodlots. We identified 13 species of amphibians within the study plot, and every woodlot contained at least one amphibian species. The most important variable in predicting total amphibian and anuran species richness was hydroperiod. For caudates, woodlot edge-to-area ratio, hydroperiod, pH, and ammonia were important characteristics in predicting species richness. Woodlots within agricultural landscapes are important refuges for amphibians.     

Influence of resource quality on the composition of soil decomposer community in fragmented and continuous habitat

The aim of this field experiment was to explore the combined effects of two factors potentially affecting the local composition of soil decomposer community: resource quality and habitat fragmentation. We created humus (habitat) patches with three different resource quality: (1) pure homogenised humus; (2) humus enriched with needle litter; and (3) humus enriched with needle and leaf litter. These patches were embedded either in a mineral soil matrix, thus representing fragmented habitat, or in natural forest soil, representing continuous (non-fragmented) habitat. The development of faunal (colonisations/extinctions of soil animal populations) and microbial communities in the patches was followed for 12 months. Our results partly supported the hypothesized strong influence of resource quality on the structure of local soil food webs: the abundances of practically all groups of soil fauna, together with biomass of fungi, were higher in the litter-enriched patches than in the pure humus patches. The manifestation and magnitude of the responses of fauna were, however, strongly affected by complex interactions between the characteristics (especially colonisation capacity) of the faunal group in question, habitat quality and time of sampling. In microarthropods and nematodes, the effect of resource quality cascaded up to the predatory level, rendering further support to the existence of strong bottom-up control in soil food webs. Contrary to our expectations, species richness of the communities was not unanimously affected by resource quality. Habitat fragmentation affected the communities only through different number and identity of patch-colonising species in the fragmented and continuous habitat: fragmentation induced no extinctions of species during the experiment at any resource quality level. Consequently, the results indicate that resource quality is more important factor than habitat fragmentation in determining the local structure of communities in soils. On the other hand, colonisation capacities of soil organisms appear to set limits to the exploitation of local resources.     

Effects of landscape characteristics on amphibian distribution in a forest-dominated landscape

We investigated the influence of landscape and wetland characteristics on pond-breeding amphibian assemblages in south-central New Hampshire, a relatively low populated and heavily forested region of the northeastern United States. This allowed us to better understand landscape influences in less disturbed areas, and to determine critical landscape disturbance thresholds, above which amphibians are negatively impacted. We sampled 61 wetlands for larval amphibians in 1998 and 1999 to examine the influence of forest cover and road density (at seven buffer distances between 100 and 2000 m) and wetland characteristics on larval amphibian assemblages. Assemblages were influenced primarily by forest cover and wetland hydroperiod. Species richness was most strongly influenced by the proportion of forest cover within 1000 m of the wetland. Several species were also influenced by forest cover, but were differentially influenced by buffer widths. Our study suggests that, at least in the northeast US, wetlands with <40% forest cover within a 1000 m radius may have depauperate larval amphibian assemblages, and forest cover above 60% within a 1000 m radius is likely to ensure species rich and abundant larval amphibian assemblages. Given the above, current federal and state regulations that focus amphibian protection efforts on narrow terrestrial buffers surrounding wetlands are likely to be inadequate.