Daniel H. Thornton

The influence of landscape, patch, and within-patch factors on species presence and abundance: a review of focal patch studies

Understanding the influence of large and small-scale heterogeneity on species distribution and abundance is one of the major foci of landscape ecology research in fragmented environments. Although a large number of studies have addressed this issue individually, little effort has been made to synthesize the vast amount of literature published in the last decade. We reviewed 122 focal patch studies on 954 species published between 1998 and 2009 to determine the probability of species responding significantly to landscape, patch, and within-patch variables. We assessed the influence of taxonomic, life history, and methodological variables on probability of response to these 3 levels. Species in diverse taxa responded at high rates to factors at all three levels, suggesting that a multi-level approach is often necessary for understanding species response in patchy systems. Mammals responded at particularly high rates to landscape variables and therefore may benefit more than other taxa from landscape-level conservation efforts in fragmented environments. The probability of detecting a species response to landscape context, patch, and within-patch factors was influenced by a variety of methodological aspects of the studies such as type of landscape metric used, type of response variable, and sample size. Study design issues rarely are discussed by authors as reasons why a particular study did not find an effect of a variable, but should be given more consideration in future studies.

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.

ECOLOGICAL SEPARATION WITHIN NEWLY SYMPATRIC POPULATIONS OF COYOTES AND BOBCATS IN SOUTH-CENTRAL FLORIDA

Abstract The coyote (Canis latrans) has recently expanded its geographic range into Florida, and the impacts of this range expansion on Florida ecosystems are likely to be complex. An area of particular concern is the effect on native carnivores. From May 2001 to May 2002, we investigated the ecological relationships between the coyote and bobcat (Lynx rufus) in south-central Florida to determine how they partition space, habitat, time, and food. Ecological separation was facilitated by dietary differences. Coyotes preyed primarily upon large ungulates and consumed substantial quantities of fruit, whereas bobcats primarily consumed rodents and lagomorphs. Coyotes and bobcats displayed similar habitat selection and activity patterns, and their high interspecific overlap in home ranges indicated a lack of large-scale spatial segregation. However, at the finer scale of core areas, patterns of spatial segregation were present. The lack of evidence for negative interactions at our study site suggests that non-overlapping core areas reduces agonistic encounters between the 2 species.

Reconsidering the Specialist-Generalist Paradigm in Niche Breadth Dynamics: Resource Gradient Selection by Canada Lynx and Bobcat

The long-standing view in ecology is that disparity in overall resource selection is the basis for identifying niche breadth patterns, with species having narrow selection being classified “specialists” and those with broader selection being “generalists”. The standard model of niche breadth characterizes generalists and specialists as having comparable levels of overall total resource exploitation, with specialists exploiting resources at a higher level of performance over a narrower range of conditions. This view has gone largely unchallenged. An alternate model predicts total resource use being lower for the specialized species with both peaking at a comparable level of performance over a particular resource gradient. To reconcile the niche breadth paradigm we contrasted both models by developing range-wide species distribution models for Canada lynx, Lynx canadensis, and bobcat, Lynx rufus. Using a suite of environmental factors to define each species’ niche, we determined that Canada lynx demonstrated higher total performance over a restricted set of variables, specifically those related to snow and altitude, while bobcat had higher total performance across most variables. Unlike predictions generated by the standard model, bobcat level of exploitation was not compromised by the trade-off with peak performance, and Canada lynx were not restricted to exploiting a narrower range of conditions. Instead, the emergent pattern was that specialist species have a higher total resource utilization and peak performance value within a smaller number of resources or environmental axes than generalists. Our results also indicate that relative differences in niche breadth are strongly dependent on the variable under consideration, implying that the appropriate model describing niche breadth dynamics between specialists and generalists may be more complex than either the traditional heuristic or our modified version. Our results demonstrate a need to re-evaluate traditional, but largely untested, assumptions regarding resource utilization in species with broad and narrow niches.

Passive sampling effects and landscape location alter associations between species traits and response to fragmentation

As tropical reserves become smaller and more isolated, the ability of species to utilize fragmented landscapes will be a key determinant of species survival. Although several ecological and life history traits commonly are associated with vulnerability to fragmentation, the combination of traits that are most highly influential and the effectiveness of those traits in predicting vulnerability across distinct landscapes, remains poorly understood. We studied use of forest fragments by 25 mid- and large-sized neotropical mammals in Guatemala to determine how seven species traits influence vulnerability to fragmentation. We measured vulnerability in two ways: one measure that did not remove passive sampling effects (proportion of fragments occupied), and one that did (difference in occupancy rates within continuous and fragmented sites). We also examined the influence of species traits on patch occupancy rates of the same set of mammals on two landscapes in Mexico. When not accounting for passive sampling effects, body size, home range size, and vulnerability to hunting influenced how species responded to fragmentation. However, after controlling for passive sampling effects, only vulnerability to hunting strongly influenced sensitivity to fragmentation. Species that were heavily hunted were much less common in forest patches than in continuous forest sites of the same sampling size. The cross-landscape comparison revealed both similarities and differences in the species traits that influenced patch occupancy patterns on each landscape. Given the ubiquity of hunting in tropical environments, our findings indicate that management efforts in fragmented landscapes that do not account for hunting pressure may be ineffective in conserving heavily hunted tropical species. Our study also indicates that species traits may be useful in predicting relative patch occupancy rates and/or vulnerability to fragmentation across distinct landscapes, but that caution must be used as certain traits can become more or less influential on different landscapes, even when considering the same set of species.

Evidence for large-scale effects of competition: niche displacement in Canada lynx and bobcat.

Determining the patterns, causes and consequences of character displacement is central to our understanding of competition in ecological communities. However, the majority of competition research has occurred over small spatial extents or focused on fine-scale differences in morphology or behaviour. The effects of competition on broad-scale distribution and niche characteristics of species remain poorly understood but critically important. Using range-wide species distribution models, we evaluated whether Canada lynx (Lynx canadensis) or bobcat (Lynx rufus) were displaced in regions of sympatry. Consistent with our prediction, we found that lynx niches were less similar to those of bobcat in areas of sympatry versus allopatry, with a stronger reliance on snow cover driving lynx niche divergence in the sympatric zone. By contrast, bobcat increased niche breadth in zones of sympatry, and bobcat niches were equally similar to those of lynx in zones of sympatry and allopatry. These findings suggest that competitively disadvantaged species avoid competition at large scales by restricting their niche to highly suitable conditions, while superior competitors expand the diversity of environments used. Our results indicate that competition can manifest within climatic niche space across species’ ranges, highlighting the importance of biotic interactions occurring at large spatial scales on niche dynamics.

Predicting shifts in parasite distribution with climate change: a multitrophic level approach

Climate change likely will lead to increasingly favourable environmental conditions for many parasites. However, predictions regarding parasitisms impacts often fail to account for the likely variability in host distribution and how this may alter parasite occurrence. Here, we investigate potential distributional shifts in the meningeal worm, Parelaphostrongylosis tenuis, a protostrongylid nematode commonly found in white-tailed deer in North America, whose life cycle also involves a free-living stage and a gastropod intermediate host. We modelled the distribution of the hosts and free-living larva as a complete assemblage to assess whether a complex trophic system will lead to an overall increase in parasite distribution with climate change, or whether divergent environmental niches may promote an ecological mismatch. Using an ensemble approach to climate modelling under two different carbon emission scenarios, we show that whereas the overall trend is for an increase in niche breadth for each species, mismatches arise in habitat suitability of the free-living larva vs. the definitive and intermediate hosts. By incorporating these projected mismatches into a combined model, we project a shift in parasite distribution accounting for all steps in the transmission cycle, and identify that overall habitat suitability of the parasite will decline in the Great Plains and southeastern USA, but will increase in the Boreal Forest ecoregion, particularly in Alberta. These results have important implications for wildlife conservation and management due to the known pathogenicity of parelaphostrongylosis to alternate hosts including moose, caribou and elk. Our results suggest that disease risk forecasts which fail to consider biotic interactions may be overly simplistic, and that accounting for each of the parasites life stages is key to refining predicted responses to climate change.

Assessing the umbrella value of a range-wide conservation network for Jaguars (Panthera onca)

Umbrella species are employed as conservation short-cuts for the design of reserves or reserve networks. However, empirical data on the effectiveness of umbrellas is equivocal, which has prevented more widespread application of this conservation strategy. We perform a novel, large-scale evaluation of umbrella species by assessing the potential umbrella value of a jaguar (Panthera onca) conservation network (consisting of viable populations and corridors) that extends from Mexico to Argentina. Using species richness, habitat quality, and fragmentation indices of ~1500 co-occurring mammal species, we show that jaguar populations and corridors overlap a substantial amount and percentage of high-quality habitat for co-occurring mammals and that the jaguar network performs better than random networks in protecting high-quality, interior habitat. Significantly, the effectiveness of the jaguar network as an umbrella would not have been noticeable had we focused on species richness as our sole metric of umbrella utility. Substantial inter-order variability existed, indicating the need for complementary conservation strategies for certain groups of mammals. We offer several reasons for the positive result we document, including the large spatial scale of our analysis and our focus on multiple metrics of umbrella effectiveness. Taken together, our results demonstrate that a regional, single-species conservation strategy can serve as an effective umbrella for the larger community and should help conserve viable populations and connectivity for a suite of co-occurring mammals. Current and future range-wide planning exercises for other large predators may therefore have important umbrella benefits.

The subtle role of climate change on population genetic structure in Canada lynx

Anthropogenically driven climatic change is expected to reshape global patterns of species distribution and abundance. Given recent links between genetic variation and environmental patterns, climate change may similarly impact genetic population structure, but we lack information on the spatial and mechanistic underpinnings of genetic-climate associations. Here, we show that current genetic variability of Canada lynx (Lynx canadensis) is strongly correlated with a winter climate gradient (i.e. increasing snow depth and winter precipitation from west-to-east) across the Pacific-North American (PNO) to North Atlantic Oscillation (NAO) climatic systems. This relationship was stronger than isolation by distance and not explained by landscape variables or changes in abundance. Thus, these patterns suggest that individuals restricted dispersal across the climate boundary, likely in the absence of changes in habitat quality. We propose habitat imprinting on snow conditions as one possible explanation for this unusual phenomenon. Coupling historical climate data with future projections, we also found increasingly diverging snow conditions between the two climate systems. Based on genetic simulations using projected climate data (2041-2070), we predicted that this divergence could lead to a threefold increase in genetic differentiation, potentially leading to isolated east-west populations of lynx in North America. Our results imply that subtle genetic structure can be governed by current climate and that substantive genetic differentiation and related ecological divergence may arise from changing climate patterns.

Response of large galliforms and tinamous (Cracidae, Phasianidae, Tinamidae) to habitat loss and fragmentation in northern Guatemala

The potential conservation value of fragmented or countryside landscapes in the tropics is being increasingly recognized. However, the degree to which fragmented landscapes can support species and the key patch and landscape features that promote population persistence remain poorly understood for elusive species such as ground-dwelling birds. We examined the presence/absence of seven species of galliforms and tinamous in 50 forest patches of 2.9–445 ha in northern Guatemala using camera traps and audiovisual surveying. After accounting for differences in detectability among species we found great variation in patterns of vulnerability of these species to habitat loss and fragmentation, with the three largest species being the most vulnerable. Distribution patterns of species among patches was influenced more strongly by measures of landscape context, such as the amount and configuration of habitat in the surrounding landscape, than within-patch variation in vegetation structure or disturbance. Our results indicate that large-bodied game birds may be particularly sensitive to habitat loss and fragmentation and emphasize that management efforts for these species need to go beyond consideration of local, within-patch factors to consider the impact of processes in the surrounding landscape. Our findings also demonstrate the utility of camera traps as a methodology for surveying large terrestrial bird species in fragmented landscapes.