Body Size Variation of Mammals in a Fragmented, Temperate Rainforest

Abstract:  Body size is perhaps the most important trait of an organism, affecting all of its physiological and ecological processes and, therefore, fundamentally influencing its ability to survive and reproduce in different environments, including those that have been modified by human activities. We tested the hypothesis that anthropogenic transformation of old-growth forest landscapes can result in significant intraspecific changes in body size of resident biotas. We collected data on five species of nonvolant mammals (common deer mouse [Peromyscus maniculatus ], northwestern deer mouse [P. keeni] , southern red-backed vole [Clethrionomys gapperi ], montane shrew [Sorex monticolus] , and Trowbridge's shrew [S. trowbridgii] ) to test whether body size (mass and length) of these species varied across types of land cover (macrohabitats) and along elevational gradients of the fragmented, temperate rainforest of Olympic National Forest (Washington, U.S.A.). We measured 2168 and 1134 individuals for body mass and body length, respectively. Three species ( P. keeni , S. monticolus , and S. trowbridgii ) exhibited significantly different body size among macrohabitats: individuals from fragments were smaller than those in old-growth corridors and those in more extensive stands of old-growth forest. Body size of P. keeni was significantly correlated with elevation along corridors, peaking near the medial reaches of the corridors. The effects of anthropogenic transformations of this landscape of old-growth, temperate rainforest, although not universal among the five species, were significant and rapid—developing in just a few decades following tree harvests. Thus, anthropogenic fragmentation may influence not only the diversity, species composition, and densities of local biotas, but also one of the most fundamental and defining characteristics of native species—their body size.     

Habitat Split as a Cause of Local Population Declines of Amphibians with Aquatic Larvae

Abstract:  Most amphibian species have biphasic life histories and undergo an ontogenetic shift from aquatic to terrestrial habitats. In deforested landscapes, streams and forest fragments are frequently disjunct, jeopardizing the life cycle of forest-associated amphibians with aquatic larvae. We tested the impact of habitat split—defined as human-induced disconnection between habitats used by different life-history stages of a species—on four forest-associated amphibian species in a severely fragmented landscape of the Brazilian Atlantic Forest. We surveyed amphibians in forest fragments with and without streams (referred to as wet and dry fragments, respectively), including the adjacent grass-field matrix. Our comparison of capture rates in dry fragments and nearby streams in the matrix allowed us to evaluate the number of individuals that engaged in high-risk migrations through nonforested habitats. Adult amphibians moved from dry fragments to matrix streams at the beginning of the rainy season, reproduced, and returned at the end of the breeding period. Juveniles of the year moved to dry fragments along with adults. These risky reproductive migrations through nonforested habitats that expose individuals to dehydration, predation, and other hazards may cause population declines in dry fragments. Indeed, capture rates were significantly lower in dry fragments compared with wet fragments. Declining amphibians would strongly benefit from investments in the conservation and restoration of riparian vegetation and corridors linking breeding and nonbreeding areas.     

Fragments as Islands: a Synthesis of Faunal Responses to Habitat Patchiness

Abstract:  Scientific interest in the impact of habitat fragmentation on biodiversity is increasing, but our understanding of fragmentation is clouded by a lack of appreciation for fundamental similarities and differences across studies representing a wide range of taxa and landscape types. In an effort to synthesize data describing ecological responses of animals to fragmentation across two classes of independent variables (taxonomic group and landscape), we sampled 148 studies of five major faunal groups from the primary literature and analyzed data on 13 variables extracted from those studies. We focused our analyses on three classes of dependent variables (effects of area and isolation on species richness, z values, and nestedness and species composition). Area ranged over more orders of magnitude than isolation and tended to explain more variation in species richness than isolation. There were few matrix or taxon effects on the patterns we investigated, although we did find that sky islands tended to manifest isolation effects on both species richness and nestedness more frequently than other patch types. Sky islands may offer insight into the future of habitat patches fragmented by contemporary habitat loss, and because they show a stronger effect of isolation than other patch types, we suggest that isolation will play an increasing role in the biology of habitat fragments. We use multiple lines of evidence to suggest that our understanding of the role of isolation on community assembly in fragmented landscapes is inadequate. Finally, our observation that consistent taxonomic differences in community patterns were minimal suggests that conservation actions intended to mitigate the negative effects of extinction may have far-reaching effects across taxonomic groups.     

Conservation Value of Remnant Riparian Forest Corridors of Varying Quality for Amazonian Birds and Mammals

Abstract:  Forest corridors are often considered the main instrument with which to offset the effects of habitat loss and fragmentation. Brazilian forestry legislation requires that all riparian zones on private landholdings be maintained as permanent reserves and sets fixed minimum widths of riparian forest buffers to be retained alongside rivers and perennial streams. We investigated the effects of corridor width and degradation status of 37 riparian forest sites (including 24 corridors connected to large source-forest patches, 8 unconnected forest corridors, and 5 control riparian zones embedded within continuous forest patches) on bird and mammal species richness in a hyper-fragmented forest landscape surrounding Alta Floresta, Mato Grosso, Brazil. We used point-count and track-sampling methodology, coupled with an intensive forest-quality assessment that combined satellite imagery and ground truthed data. Vertebrate use of corridors was highly species-specific, but broad trends emerged depending on species life histories and their sensitivity to disturbance. Narrow and/or highly disturbed riparian corridors retained only a depauperate vertebrate assemblage that was typical of deforested habitats, whereas wide, well-preserved corridors retained a nearly complete species assemblage. Restriction of livestock movement along riparian buffers and their exclusion from key areas alongside deforested streams would permit corridor regeneration and facilitate restoration of connectivity .     

Predicting the Deleterious Effects of Mutation Load in Fragmented Populations

Abstract:  Human-induced habitat fragmentation constitutes a major threat to biodiversity. Both genetic and demographic factors combine to drive small and isolated populations into extinction vortices. Nevertheless, the deleterious effects of inbreeding and drift load may depend on population structure, migration patterns, and mating systems and are difficult to predict in the absence of crossing experiments. We performed stochastic individual-based simulations aimed at predicting the effects of deleterious mutations on population fitness (offspring viability and median time to extinction) under a variety of settings (landscape configurations, migration models, and mating systems) on the basis of easy-to-collect demographic and genetic information. Pooling all simulations, a large part (70%) of variance in offspring viability was explained by a combination of genetic structure ( F_ST ) and within-deme heterozygosity ( H_S ). A similar part of variance in median time to extinction was explained by a combination of local population size ( N ) and heterozygosity ( H_S ). In both cases the predictive power increased above 80% when information on mating systems was available. These results provide robust predictive models to evaluate the viability prospects of fragmented populations.     

Efficacy of Land-Cover Models in Predicting Isolation of Marbled Salamander Populations in a Fragmented Landscape

Abstract:  Amphibians worldwide are facing rapid declines due to habitat loss and fragmentation, disease, and other causes. Where habitat alteration is implicated, there is a need for spatially explicit conservation plans. Models built with geographic information systems (GIS) are frequently used to inform such planning. We explored the potential for using GIS models of functional landscape connectivity as a reliable proxy for genetically derived measures of population isolation. We used genetic assignment tests to characterize isolation of marbled salamander populations and evaluated whether the relative amount of modified habitat around breeding ponds was a reliable indicator of population isolation. Using a resampling analysis, we determined whether certain land-cover variables consistently described population isolation. We randomly drew half the data for model building and tested the performance of the best models on the other half 100 times. Deciduous forest was consistently associated with lower levels of population isolation, whereas salamander populations in regions of agriculture and anthropogenic development were more isolated. Models that included these variables and pond size explained 65–70% of variation in genetically inferred isolation across sites. The resampling analysis confirmed that these habitat variables were consistently good predictors of isolation. Used judiciously, simple GIS models with key land-cover variables can be used to estimate population isolation if field sampling and genetic analysis are not possible.     

Appreciating Ecological Complexity: Habitat Contours as a Conceptual Landscape Model

Abstract:  Organisms respond to their surroundings at multiple spatial scales, and different organisms respond differently to the same environment. Existing landscape models, such as the "fragmentation model" (or patch-matrix-corridor model) and the "variegation model," can be limited in their ability to explain complex patterns for different species and across multiple scales. An alternative approach is to conceptualize landscapes as overlaid species-specific habitat contour maps. Key characteristics of this approach are that different species may respond differently to the same environmental conditions and at different spatial scales. Although similar approaches are being used in ecological modeling, there is much room for habitat contours as a useful conceptual tool. By providing an alternative view of landscapes, a contour model may stimulate more field investigations stratified on the basis of ecological variables other than human-defined patches and patch boundaries. A conceptual model of habitat contours may also help to communicate ecological complexity to land managers. Finally, by incorporating additional ecological complexity, a conceptual model based on habitat contours may help to bridge the perceived gap between pattern and process in landscape ecology. Habitat contours do not preclude the use of existing landscape models and should be seen as a complementary approach most suited to heterogeneous human-modified landscapes.     

Spillover of Agriculturally Subsidized Predators as a Potential Threat to Native Insect Herbivores in Fragmented Landscapes

Abstract:  Habitat loss and fragmentation can have strong negative impacts on populations of some native species. Spillover of generalist natural enemies from the surrounding landscape matrix is one mechanism potentially generating such effects, yet this has been rarely studied in insects. We examined the influence of habitat conversion to agriculture on the abundance and potential effects of predatory coccinellid beetles on native insect herbivores within 12 grassland remnants in central Nebraska (U.S.A.). Results of sweep sampling revealed that coccinellids were three to six times more abundant at native grassland sites embedded within cropland-dominated landscapes compared with control sites in grassland-dominated landscapes over the 3 years of the study. Exclusion experiments further demonstrated that predation intensity was strongly related to coccinellid abundances across sites and that coccinellids can dramatically reduce densities of a native aphid herbivore. In contrast to studies of specialized insect parasitoids, which have generally found reduced enemy pressure in fragmented landscapes, our results suggest that native herbivores may in some cases experience increased consumer pressure in landscapes with increasing habitat loss because of spillover of generalist predators from surrounding cropland habitats.     

Fish Responses to Experimental Fragmentation of Seagrass Habitat

Abstract: Understanding the consequences of habitat fragmentation has come mostly from comparisons of patchy and continuous habitats. Because fragmentation is a process, it is most accurately studied by actively fragmenting large patches into multiple smaller patches. We fragmented artificial seagrass habitats and evaluated the impacts of fragmentation on fish abundance and species richness over time (1 day, 1 week, 1 month). Fish assemblages were compared among 4 treatments: control (single, continuous 9‐m² patches); fragmented (single, continuous 9‐m² patches fragmented to 4 discrete 1‐m² patches); prefragmented/patchy (4 discrete 1‐m² patches with the same arrangement as fragmented); and disturbance control (fragmented then immediately restored to continuous 9‐m² patches). Patchy seagrass had lower species richness than actively fragmented seagrass (up to 39% fewer species after 1 week), but species richness in fragmented treatments was similar to controls. Total fish abundance did not vary among treatments and therefore was unaffected by fragmentation, patchiness, or disturbance caused during fragmentation. Patterns in species richness and abundance were consistent 1 day, 1 week, and 1 month after fragmentation. The expected decrease in fish abundance from reduced total seagrass area in fragmented and patchy seagrass appeared to be offset by greater fish density per unit area of seagrass. If fish prefer to live at edges, then the effects of seagrass habitat loss on fish abundance may have been offset by the increase (25%) in seagrass perimeter in fragmented and patchy treatments. Possibly there is some threshold of seagrass patch connectivity below which fish abundances cannot be maintained. The immediate responses of fish to experimental habitat fragmentation provided insights beyond those possible from comparisons of continuous and historically patchy habitat.     

Demographic Limitations of the Ability of Habitat Restoration to Rescue Declining Populations

Abstract:  Habitat restoration is often recommended in conservation without first evaluating whether populations are in fact habitat limited and thus whether declining populations can be stabilized or recovered through habitat restoration. We used a spatially structured demographic model coupled with a dynamic neutral landscape model to evaluate whether habitat restoration could rescue populations of several generic migratory songbirds that differed in their sensitivity to habitat fragmentation (i.e., severity of edge effects on nesting success). Simulating a best-case scenario, landscapes were instantly restored to 100% habitat before, at, or after habitat loss exceeded the species' vulnerability threshold. The vulnerability threshold is a measure of extinction risk, in which the change in population growth rate ( δλ ) scaled to the rate of habitat loss ( δh ) falls below −1% ( δλ/δh ≤ −0.01). Habitat restoration was most effective for species with low-to-moderate edge sensitivities and in landscapes that had not previously experienced extensive fragmentation. To stabilize populations of species that were highly edge sensitive or any species in heavily fragmented landscapes, restoration needed to be initiated long before the vulnerability threshold was reached. In practice, habitat restoration is generally not initiated until a population is at risk of extinction, but our model results demonstrate that some populations cannot be recovered at this point through habitat restoration alone. At this stage, habitat loss and fragmentation have seriously eroded the species' demographic potential such that halting population declines is limited more by demographic factors than the amount of available habitat. Evidence that populations decline in response to habitat loss is thus not sufficient to conclude that habitat restoration will be sufficient to rescue declining populations.     

Value of Small Patches in the Conservation of Plant-Species Diversity in Highly Fragmented Rainforest

Abstract:  We evaluated the importance of small (<5 ha) forest patches for the conservation of regional plant diversity in the tropical rainforest of Los Tuxtlas, Mexico. We analyzed the density of plant species (number of species per 0.1 ha) in 45 forest patches of different sizes (1–700 ha) in 3 landscapes with different deforestation levels (4, 11, and 24% forest cover). Most of the 364 species sampled (360 species, 99%) were native to the region, and only 4 (1%) were human-introduced species. Species density in the smallest patches was high and variable; the highest (84 species) and lowest (23 species) number of species were recorded in patches of up to 1.8 ha. Despite the small size of these patches, they contained diverse communities of native plants, including endangered and economically important species. The relationship between species density and area was significantly different among the landscapes, with a significant positive slope only in the landscape with the highest deforestation level. This indicates that species density in a patch of a given size may vary among landscapes that have different deforestation levels. Therefore, the conservation value of a patch depends on the total forest cover remaining in the landscape. Our findings revealed, however, that a great portion of regional plant diversity was located in very small forest patches (<5 ha), most of the species were restricted to only a few patches (41% of the species sampled were distributed in only 1–2 patches, and almost 70% were distributed in 5 patches) and each landscape conserved a unique plant assemblage. The conservation and restoration of small patches is therefore necessary to effectively preserve the plant diversity of this strongly deforested and unique Neotropical region.     

Relationships between Human Disturbance and Wildlife Land Use in Urban Habitat Fragments

Abstract: Habitat remnants in urbanized areas typically conserve biodiversity and serve the recreation and urban open‐space needs of human populations. Nevertheless, these goals can be in conflict if human activity negatively affects wildlife. Hence, when considering habitat remnants as conservation refuges it is crucial to understand how human activities and land uses affect wildlife use of those and adjacent areas. We used tracking data (animal tracks and den or bed sites) on 10 animal species and information on human activity and environmental factors associated with anthropogenic disturbance in 12 habitat fragments across San Diego County, California, to examine the relationships among habitat fragment characteristics, human activity, and wildlife presence. There were no significant correlations of species presence and abundance with percent plant cover for all species or with different land‐use intensities for all species, except the opossum (Didelphis virginiana), which preferred areas with intensive development. Woodrats (Neotoma spp.) and cougars (Puma concolor) were associated significantly and positively and significantly and negatively, respectively, with the presence and prominence of utilities. Woodrats were also negatively associated with the presence of horses. Raccoons (Procyon lotor) and coyotes (Canis latrans) were associated significantly and negatively and significantly and positively, respectively, with plant bulk and permanence. Cougars and gray foxes (Urocyon cinereoargenteus) were negatively associated with the presence of roads. Roadrunners (Geococcyx californianus) were positively associated with litter. The only species that had no significant correlations with any of the environmental variables were black‐tailed jackrabbits (Lepus californicus) and mule deer (Odocoileus hemionus). Bobcat tracks were observed more often than gray foxes in the study area and bobcats correlated significantly only with water availability, contrasting with results from other studies. Our results appear to indicate that maintenance of habitat fragments in urban areas is of conservation benefit to some animal species, despite human activity and disturbance, as long as the fragments are large.     

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 .     

Connectivity in an Agricultural Landscape as Reflected by Interpond Movements of a Freshwater Turtle

Abstract:  Connectivity is a measure of how landscape features facilitate movement and thus is an important factor in species persistence in a fragmented landscape. The scarcity of empirical studies that directly quantify species movement and determine subsequent effects on population density have, however, limited the utility of connectivity measures in conservation planning. We undertook a 4-year study to calculate connectivity based on observed movement rates and movement probabilities for five age-sex classes of painted turtles ( Chrysemys picta ) inhabiting a pond complex in an agricultural landscape in northern Virginia (U.S.A.). We determined which variables influenced connectivity and the relationship between connectivity and subpopulation density. Interpatch distance and quality of habitat patches influenced connectivity but characteristics of the intervening matrix did not. Adult female turtles were more influenced by the habitat quality of recipient ponds than other age-sex classes. The importance of connectivity on spatial population dynamics was most apparent during a drought. Population density and connectivity were low for one pond in a wet year but dramatically increased as other ponds dried. Connectivity is an important component of species persistence in a heterogeneous landscape and is strongly dependent on the movement behavior of the species. Connectivity may reflect active selection or avoidance of particular habitat patches. The influence of habitat quality on connectivity has often been ignored, but our findings highlight its importance. Conservation planners seeking to incorporate connectivity measures into reserve design should not ignore behavior in favor of purely structural estimates of connectivity.     

Modeling the Effects of Anthropogenic Habitat Change on Savanna Snake Invasions into African Rainforest

Abstract:  We used a species-distribution modeling approach, ground-based climate data sets, and newly available remote-sensing data on vegetation from the MODIS and Quick Scatterometer sensors to investigate the combined effects of human-caused habitat alterations and climate on potential invasions of rainforest by 3 savanna snake species in Cameroon, Central Africa: the night adder (Causus maculatus) , olympic lined snake (Dromophis lineatus) , and African house snake (Lamprophis fuliginosus) . Models with contemporary climate variables and localities from native savanna habitats showed that the current climate in undisturbed rainforest was unsuitable for any of the snake species due to high precipitation. Limited availability of thermally suitable nest sites and mismatches between important life-history events and prey availability are a likely explanation for the predicted exclusion from undisturbed rainforest. Models with only MODIS-derived vegetation variables and savanna localities predicted invasion in disturbed areas within the rainforest zone, which suggests that human removal of forest cover creates suitable microhabitats that facilitate invasions into rainforest. Models with a combination of contemporary climate, MODIS- and Quick Scatterometer-derived vegetation variables, and forest and savanna localities predicted extensive invasion into rainforest caused by rainforest loss. In contrast, a projection of the present-day species-climate envelope on future climate suggested a reduction in invasion potential within the rainforest zone as a consequence of predicted increases in precipitation. These results emphasize that the combined responses of deforestation and climate change will likely be complex in tropical rainforest systems.