The Sustainability of Wolverine Trapping Mortality in Southern Canada

Range declines, habitat connectivity, and trapping have created conservation concern for wolverines throughout their range in North America. Previous researchers used population models and observed estimates of survival and reproduction to infer that current trapping rates limit population growth, except perhaps in the far north where trapping rates are lower. Assessing the sustainability of trapping requires demographic and abundance data that are expensive to acquire and are therefore usually only achievable for small populations, which makes generalization risky. We surveyed wolverines over a large area of southern British Columbia and Alberta, Canada, used spatial capture-recapture models to estimate density, and calculated trapping kill rates using provincial fur harvest data. Wolverine density averaged 2 wolverines/1,000 km² and was positively related to spring snow cover and negatively related to road density. Observed annual trapping mortality was >8.4%/year. This level of mortality is unlikely to be sustainable except in rare cases where movement rates are high among sub-populations and sizable un-trapped refuges exist. Our results suggest wolverine trapping is not sustainable because our study area was fragmented by human and natural barriers and few large refuges existed. We recommend future wolverine trapping mortality be reduced by ≥50% throughout southern British Columbia and Alberta to promote population recovery. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Assessing population structure and gene flow in Montana wolverines (Gulo gulo) using assignment-based approaches

In North America, wolverines once occupied a continuous range from Alaska southward to New Mexico. In the lower 48 states, small remnant populations remain only in the northwestern United States. Among these remnant populations, the Montana population has the highest probability of long-term persistence given its size and proximity to healthy populations in Canada. In this study, we evaluate population genetic structure and gene flow among Montana wolverines using 10 polymorphic microsatellite loci. Bayesian and frequency-based assignment tests revealed significant population substructure and provide support for at least three subpopulations in Montana. FST values between subpopulations ranged from 0.08 to 0.10 and provide evidence for male-biased dispersal. The high degree of population substructure and low levels of gene flow contrast results from wolverine population genetic studies in less fragmented landscapes of Alaska and Canada. This study provides additional support for the hypothesis that large carnivore populations of Montana are becoming increasingly fragmented due to human development and disturbance.     

Genetic structure of North American wolverine (Gulo gulo) populations

Wolverines (Gulo gulo) are found in low densities throughout their circumpolar distribution. They are also potentially susceptible to human-caused population fragmentation (development, recreation and fur harvesting). The combination of these factors has contributed to this species being listed as having either vulnerable or endangered status across much of its current range. The effects of inherently low densities and anthropogenic pressures on the genetic structure and variation of wolverine populations are, as yet, unknown. In this study, 461 individuals were typed at 12 microsatellite loci to investigate the population genetic structure of wolverines from north-western Alaska to eastern Manitoba. Levels of gene flow and population differentiation among the sampled regions were estimated via a genotype assignment test, pairwise F(ST), and two genetic distance measures. Our results suggest that wolverine populations from southernmost regions, in which anthropogenic factors are strongest, revealed more genetic structuring than did northern populations. Furthermore, these results suggest that reductions in this species' range may have led to population fragmentation in the extreme reaches of its southern distribution. The continued reduction of suitable habitat for this species may lead to more populations becoming isolated remnants of a larger distribution of northern wolverines, as documented in other North American carnivore species.     

Nonlinear responses of wolverine populations to declining winter snowpack

Understanding the population-level impacts of climate change is critical for effectively managing ecosystems. Predators are important components of many systems because they provide top−down control of community structure. Ecological theory suggests that these species could be particularly susceptible to climate change because they generally occur at low densities and have resource-limited populations. Yet, our understanding of climate-change impacts on predators is hindered by the difficulty in assessing complex, nonlinear dynamics over the large spatial scales necessary to depict a species’ general response to abiotic forcing. Here we use fur-return data to characterize population dynamics of a snow-adapted carnivore, the wolverine, across most of its North American range. Using novel modeling techniques, we simultaneously measured the impact of winter snowpack on wolverine dynamics across critical thresholds in snowpack depth and two domains of population growth. Winter snowpack declined from 1970 to 2004 in nearly the entire region studied, concordant with increases in Northern Hemisphere temperature anomalies. Fur returns have declined in many areas; our models show that snowpack has strong, nonlinear effects on wolverine population dynamics. Importantly, wolverine harvests dropped the fastest in areas where snowpack declined most rapidly and also where snowpack had the greatest effect on population dynamics. Moreover, declining snow cover appears to drive trends in wolverine population synchrony, with important implications for overall persistence. These results illustrate the vulnerability and complex responses of predator populations to climate change. We also suggest that declining snowpack may be an important and hitherto little-analyzed mechanism through which climate change alters high-latitude ecosystems.     

The Abundance and Distribution of Wolverines in British Columbia,Canada

ABSTRACT The abundance and distribution of carnivores and their habitat are key information needed for status assessment, conservation planning, population management, and assessment of the effects of human development on their habitat and populations. We developed a habitat quality rating system, using existing wolverine (Gulo gulo) distribution, wolverine food, ecosystem mapping, and human development data. We used this and empirically derived estimates of wolverine density to predict wolverine distribution and abundance at a provincial scale. Density estimates for wolverines in high-quality habitat averaged 6.2 wolverines/1,000 km² (95% CI = 4.2–9.5). We predicted mean densities ranging from 0.3/1,000 km² in rare-quality habitat to 4.1/1,000 km² in moderate-quality habitat. Our predicted population estimate for wolverines in British Columbia was 3,530 (95% CI = 2,700-4,760). We predicted highest densities of wolverines in interior mountainous regions, moderate densities in interior plateau and boreal forest regions, and low densities in mainland coastal regions and drier interior plateaus. We predicted that wolverines would be rare on Vancouver Island, along the outer mainland coast, and in the dry interior forests, and absent from the Queen Charlotte Islands, interior grassland environments, and areas of intensive urban development.     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

Testing Pleistocene refugia theory: phylogeographical analysis of Desmognathus wrighti,a high-elevation salamander in the southern Appalachians

During the colder climates of the Pleistocene, the ranges of high-elevation species in unglaciated areas may have expanded, leading to increased gene flow among previously isolated populations. The phylogeography of the pygmy salamander, Desmognathus wrighti, an endemic species restricted to the highest mountain peaks of the southern Appalachians, was examined to test the hypothesis that the range of D. wrighti expanded along with other codistributed taxa during the Pleistocene. Analyses of genetic variation at 14 allozymic loci and of the 12S rRNA gene in the mtDNA genome was conducted on individuals sampled from 14 population isolates throughout the range of D. wrighti. In contrast to the genetic patterns of many other high-elevation animals and plants, genetic distances derived from both molecular markers showed significant isolation by distance and genetic structuring of populations, suggesting long-term isolation of populations. Phylogeographical analyses revealed four genetically distinct population clusters that probably remained fragmented during the Pleistocene, although there was also evidence supporting recent gene flow among some population groups. Support for isolation by distance is rare among high-elevation species in unglaciated areas of North and Middle America, although not uncommon among Plethodontid Salamanders, and this pattern suggests that populations of D. wrighti did not expand entirely into suitable habitat during the Pleistocene. We propose that intrinsic barriers to dispersal, such as species interactions with other southern Appalachian plethodontid salamanders, persisted during the Pleistocene to maintain the fragmented distribution of D. wrighti and allow for significant genetic divergence of populations by restricting gene flow.     

Genetic structure among remnant populations of a migratory passerine,the Northern Wheatear Oenanthe oenanthe

Continuous animal populations often become fragmented due to anthropogenic habitat alterations. These small, fragmented populations are fragile due to demographic and genetic factors, whereas immigration can enhance their long‐term viability. Previously, we showed that high philopatry affected the local dynamics of three small and remnant subpopulations of Northern Wheatears in The Netherlands. Here, we show that these three populations together with an additional larger population in the European lowlands are highly genetically differentiated based on 22 microsatellite markers. In contrast, we found no evidence for differentiation using two mitochondrial DNA markers. An IMa2 analysis indicates that gene flow has occurred regularly among our sampled populations. As immigration of colour‐ringed birds among our sampled populations is rare at best, our results suggest that the populations have recently become isolated from one another. Low dispersal rates in highly mobile birds may occur when suitable habitat becomes highly fragmented, and will accentuate stochastic demographic processes and inbreeding, both reducing population viability. As dispersal rates are low among populations of Northern Wheatears in The Netherlands, there is only a small probability of recolonization of habitat patches where populations have become locally extinct.     

Natal habitat effects drive density—dependent scaling of dispersal decisions

The dispersal behavior of a species is critical for the stability and persistence of its populations across a landscape. How population density affects dispersal decisions is important for predicting these dynamics, as the form of density‐dependent dispersal influences the stability and persistence of populations. Natal habitat experience often has strong impacts on individual dispersal behavior as well, but its influence on density‐dependent dispersal behaviors remains unexplored. Here we address this conceptual gap in two experiments separately examining habitat selection and emigration from recently colonized patches for two species of flour beetle Tribolium sp. We found that interactions between the quality of habitat experienced during natal development and current habitat for dispersal capable adults can strongly affect the form of density dependence, including reversing the direction of nonlinearities (accelerating to decelerating), or even negating the influence of population density for individual dispersal decisions. Across heterogeneous landscapes, where individuals from different populations may experience different natal habitats, this altering of density‐dependent relationships is predicted by theory to fundamentally influence regional population dynamics. Our results indicate that species which occur across heterogeneous environments, such as during conservation reintroductions, or as invasive species spread, have much potential for natal experience to interact with density dependence and influence local and regional population dynamics.     

Distribution and Broadscale Habitat Relations of the Wolverine in the Contiguous United States

ABSTRACT Conservation of the wolverine (Gulo gulo) at the southern extent of its North American range requires reliable understandings of past and present distribution patterns and broad-scale habitat relations. We compiled 820 verifiable and documented records of wolverine occurrence (specimens, DNA detections, photos, and accounts of wolverines being killed or captured) in the contiguous United States from museums, the literature, and institutional archives. We spatially referenced 729 records with areal precision ≤1 township (93.2 km²) and temporal precision ≤10 years. Historical records (1827–1960) were located primarily in the western mountains and Great Lakes region. However, our data suggest that the historical distribution of wolverines in the Cascade Range and Sierra Nevada was disjunct, contradicting previous interpretations. Our results indicate that wolverine range in the contiguous United States had contracted substantially by the mid-1900s. Current records (1995–2005) are limited to north-central Washington, northern and central Idaho, western Montana, and northwestern Wyoming. We investigated potential relations between wolverines and alpine vegetation, cold temperatures, and spring snow cover by comparing the distribution of historical wolverine records with Kuchler's potential natural vegetation types, Holdridge's climatic life zones, and EASE snow-cover maps during the latter portion of the wolverine denning period (15 Apr-14 May). In the western mountains, historical wolverine records generally occurred in or near alpine vegetation and climatic conditions, especially at the limits of their distribution in the Cascade Range, Sierra Nevada, and southern Rocky Mountains. However, the only habitat layer that fully accounted for historical distribution patterns was spring snow cover. Causal factors for the extirpation of wolverines from the southern portions of their range in the contiguous United States are unknown, but are likely related to high levels of human-caused mortality and low to nonexistent immigration rates.     

HOT SPOTS OF GENETIC DIVERSITY DESCENDED FROM MULTIPLE PLEISTOCENE REFUGIA IN AN ALPINE UNGULATE

Species that inhabit naturally fragmented environments are expected to be spatially structured and exhibit reduced genetic diversity at the periphery of their range. Patterns of differentiation may also reflect historical processes such as recolonization from glacial refugia. We examined the relative importance of these factors in shaping the spatial patterns of genetic differentiation across the range of an alpine specialist, the North American mountain goat (Oreamnos americanus). Contrary to fossil evidence that suggests a single southern refugium, we detected evidence for additional refugia in northern British Columbia and the Alaskan coast using both mitochondrial and microsatellite DNA. A core area of elevated genetic diversity characterized both regions, and molecular dating suggested a recent Pleistocene split was followed by demographic expansion. Across their range, mountain goats were highly genetically structured and displayed the expected pattern of declining diversity toward the periphery. Gene flow was high within contiguous mountain ranges, but cross‐assignments paradoxically suggest that long‐distance contemporary dispersal movements are not uncommon. These results improve our understanding of how historical vicariance and contemporary fragmentation influence population differentiation, and have implications for conserving the adaptive potential of alpine populations and habitat.     

The role of Allee effects in gypsy moth, <Emphasis Type="Italic">Lymantria dispar</Emphasis> (L.), invasions

Allee effects have been applied historically in efforts to understand the low-density population dynamics of rare and endangered species. Many biological invasions likewise experience the phenomenon of decreasing population growth rates at low population densities because most founding populations of introduced nonnative species occur at low densities. In range expansion of established species, the initial colonizers of habitat beyond the organism’s current range are usually at low density, and thus could be subject to Allee dynamics. There has been consistent empirical and theoretical evidence demonstrating, and in some cases quantifying, the role of Allee dynamics in the gypsy moth, Lymantria dispar (L.), invasion of North America. In this review, we examine the potential causes of the Allee effect in the gypsy moth and highlight the importance of mate-finding failure as a primary mechanism behind an Allee effect, while the degree to which generalist predators induce an Allee effect remains unclear. We then explore the role of Allee effects in the establishment and spread dynamics of the gypsy moth system, which conceptually could serve as a model system for understanding how Allee effects manifest themselves in the dynamics of biological invasions.     

Interior Least Tern (Sternula antillarum) breeding distribution and ecology: implications for population‐level studies and the evaluation of alternative management strategies on large,regulated rivers

Interior Least Terns (Sternula antillarum) (ILT) are colonial, fish‐eating birds that breed within active channels of large sand bed rivers of the Great Plains and in the Lower Mississippi Valley. Multipurpose dams, irrigation structures, and engineered navigation systems have been present on these rivers for many decades. Despite severe alteration of channels and flow regimes, regulation era floods have remained effective at maintaining bare sandbar nesting habitat on many river segments and ILT populations have been stable or expanding since they were listed as endangered in 1985. We used ILT breeding colony locations from 2002 to 2012 and dispersal information to identify 16 populations and 48 subpopulations. More than 90% of ILT and >83% of river km with suitable nesting habitat occur within the two largest populations. However, replicate populations remain throughout the entire historical, geophysical, and ecological range of ILT. Rapid colonization of anthropogenic habitats in areas that were not historically occupied suggests metapopulation dynamics. The highest likelihood of demographic connectivity among ILT populations occurs across the Southern Plains and the Lower Mississippi River, which may be demographically connected with Least Tern populations on the Gulf Coast. Paired ecological and bird population models are needed to test whether previously articulated threats limit ILT population growth and to determine if management intervention is necessary and where. Given current knowledge, the largest sources of model uncertainty will be: (1) uncertainty in relationships between high flow events and subsequent sandbar characteristics and (2) uncertainty regarding the frequency of dispersal among population subunits. We recommend research strategies to reduce these uncertainties.     

Inferring Geographic Isolation of Wolverines in California Using Historical DNA

ABSTRACT Delineating a species' geographic range using the spatial distribution of museum specimens or even contemporary detection-non-detection data can be difficult. This is particularly true at the periphery of a species range where species' distributions are often disjunct. Wolverines (Gulo gulo) are wide-ranging mammals with discontinuous and potentially isolated populations at the periphery of their range. One potentially disjunct population occurred in the Sierra Nevada Mountains, California, USA, and appears to have been extirpated by the 1930s. Many early 20th century naturalists believed that this population was connected to other populations occurring in the Cascade Range of northern California, Oregon, and Washington, USA, but a recent analysis of historical records suggests that California wolverines were isolated from other populations in North America. We used DNA extracted from museum specimens to examine whether California wolverines were isolated. Both nuclear and mitochondrial DNA data indicate that California wolverines were genetically distinct from extant populations, suggesting long-term isolation. We identified 2 new control region (mitochondrial DNA) haplotypes located only within California. We used these data and referenced sequences from the Rocky Mountains, USA, to make inferences regarding potential wolverine translocations into California. In addition, we used these genetic data to make inferences about wolverine conservation throughout western North America.     

Population genetic structure of a common host predicts the spread of white‐nose syndrome,an emerging infectious disease in bats

Landscape complexity influences patterns of animal dispersal, which in turn may affect both gene flow and the spread of pathogens. White‐nose syndrome (WNS) is an introduced fungal disease that has spread rapidly throughout eastern North America, causing massive mortality in bat populations. We tested for a relationship between the population genetic structure of the most common host, the little brown myotis (Myotis lucifugus), and the geographic spread of WNS to date by evaluating logistic regression models of WNS risk among hibernating colonies in eastern North America. We hypothesized that risk of WNS to susceptible host colonies should increase with both geographic proximity and genetic similarity, reflecting historical connectivity, to infected colonies. Consistent with this hypothesis, inclusion of genetic distance between infected and susceptible colonies significantly improved models of disease spread, capturing heterogeneity in the spatial expansion of WNS despite low levels of genetic differentiation among eastern populations. Expanding our genetic analysis to the continental range of little brown myotis reveals strongly contrasting patterns of population structure between eastern and western North America. Genetic structure increases markedly moving westward into the northern Great Plains, beyond the current distribution of WNS. In western North America, genetic differentiation of geographically proximate populations often exceeds levels observed across the entire eastern region, suggesting infrequent and/or locally restricted dispersal, and thus relatively limited opportunities for pathogen introduction in western North America. Taken together, our analyses suggest a possibly slower future rate of spread of the WNS pathogen, at least as mediated by little brown myotis.     

Climate and density shape population dynamics of a marine top predator

Long-term studies have documented that climate fluctuations affect the dynamics of populations, but the relative influence of stochastic and density-dependent processes is still poorly understood and debated. Most studies have been conducted on terrestrial systems, and lacking are studies on marine systems explicitly integrating the fact that most populations live in seasonal environments and respond to regular or systematic environmental changes. We separated winter from summer mortality in a seabird population, the blue petrel Halobaena caerulea, in the southern Indian Ocean where the El Niño/Southern Oscillation effects occur with a 3-4-year lag. Seventy per cent of the mortality occurred in winter and was linked to climatic factors, being lower during anomalous warm events. The strength of density dependence was affected by climate, with population crashes occurring when poor conditions occurred at high densities. We found that an exceptionally long-lasting warming caused a ca. 40% decline of the population, suggesting that chronic climate change will strongly affect this top predator. These findings demonstrate that populations in marine systems are particularly susceptible to climate variation through complex interactions between seasonal mortality and density-dependent effects.     

Genetic variation and population structure in Scandinavian wolverine (Gulo gulo) populations

Wolverine (Gulo gulo) numbers in Scandinavia were significantly reduced during the early part of the century as a result of predator removal programmes and hunting. Protective legislation in both Sweden and Norway in the 1960s and 1970s has now resulted in increased wolverine densities in Scandinavia. We report here the development of 15 polymorphic microsatellite markers in wolverine and their use to examine the population sub-structure and genetic variability in free-ranging Scandinavian wolverine populations as well as in a sample of individuals collected before 1970. Significant subdivision between extant populations was discovered, in particular for the small and isolated population of southern Norway, which represents a recent recolonization. Overall genetic variability was found to be lower than previously reported for other mustelids, with only two to five alleles per locus and observed heterozygosities (H(O)) ranging from 0.269 to 0.376 across the examined populations, being lowest in southern Norway. Analysis of the mitochondrial DNA control region revealed no variation throughout the surveyed populations. As the historical sample did not show higher levels of genetic variability, our results are consistent with a reduction in the genetic variation in Scandinavian wolverines that pre-dates the demographic bottleneck observed during the last century. The observed subdivision between populations calls for management caution when issuing harvest quotas, especially for the geographically isolated south Norwegian population.     

Which plant traits predict species loss in calcareous grasslands with extinction debt?

Aim Habitat loss and degradation pose a major threat to biodiversity, which can result in the extinction of habitat characteristic species. However, many species exhibit a delayed response to environmental changes because of the slow intrinsic dynamics of populations, resulting in extinction debt. We assess directly the changes in habitat characteristic species composition by comparing historical (1923) and current inventories in highly fragmented grasslands. We aim to characterize the species that constitute extinction debt in European calcareous grasslands. Location Europe, Estonia, 59–60° N, 24–25° E. Methods We related eleven life‐history traits and selected habitat preferences to local extinctions of populations in grasslands where extinction debt has been largely paid. Traits were chosen to describe species dispersal and persistence abilities and were quantified from databases. Results The studied grasslands have lost 90% of their area and 30% of their characteristic plant populations in 90 years. Species more prone to local population extinction were characterized by shorter life span, self‐pollination, a lack of clonal growth, fewer seeds per shoot, lower average height, lower soil nitrogen preference and higher requirements for light, indicating a limited ability to tolerate the range of changes in biotic and abiotic conditions of the sites. Locally extinct populations were also characterized by wind‐dispersed seeds, lower seed weight and lower terminal velocity of seeds, suggesting that species strategies for long‐distance dispersal are not favoured in highly fragmented landscapes. Thus, both increased habitat isolation and decreased habitat quality are important in determining local population extinction. Main conclusions Populations more prone to local extinction were characterized by a number of life‐history traits, demonstrating a greater extinction risk for species with poorer abilities for local persistence and competition. Our results can be applied to less degraded grasslands where the extinction debt is not yet paid to determine those species most susceptible to future extinction.     

Social information and emigration: lessons from immigrants

‘Should I stay or should I go?’ is a fundamental question facing any candidate for emigration, as emigrating without outside information has major costs. Most studies on this topic have concentrated on risk‐reducing strategies (e.g. exploration) developed after leaving the natal habitat. The idea that information might be acquired before leaving has not been investigated. Immigrants carrying information about their origins could provide such information to potential emigrants in their initial habitat. We manipulated the density of common lizard (Lacerta vivipara) populations, to investigate whether immigrants originating from these populations transmitted such information to the population they joined. Emigration of the residents of this new population clearly depended on the origin of the immigrant. Immigrants are therefore a source of information, in this case about surrounding population densities, and may have a major effect on dispersal and species persistence in a fragmented habitat.     

Variation in fine‐scale genetic structure and local dispersal patterns between peripheral populations of a South American passerine bird

The distribution of suitable habitat influences natal and breeding dispersal at small spatial scales, resulting in strong microgeographic genetic structure. Although environmental variation can promote interpopulation differences in dispersal behavior and local spatial patterns, the effects of distinct ecological conditions on within‐species variation in dispersal strategies and in fine‐scale genetic structure remain poorly understood. We studied local dispersal and fine‐scale genetic structure in the thorn‐tailed rayadito (Aphrastura spinicauda), a South American bird that breeds along a wide latitudinal gradient. We combine capture‐mark‐recapture data from eight breeding seasons and molecular genetics to compare two peripheral populations with contrasting environments in Chile: Navarino Island, a continuous and low density habitat, and Fray Jorge National Park, a fragmented, densely populated and more stressful environment. Natal dispersal showed no sex bias in Navarino but was female‐biased in the more dense population in Fray Jorge. In the latter, male movements were restricted, and some birds seemed to skip breeding in their first year, suggesting habitat saturation. Breeding dispersal was limited in both populations, with males being more philopatric than females. Spatial genetic autocorrelation analyzes using 13 polymorphic microsatellite loci confirmed the observed dispersal patterns: a fine‐scale genetic structure was only detectable for males in Fray Jorge for distances up to 450 m. Furthermore, two‐dimensional autocorrelation analyzes and estimates of genetic relatedness indicated that related males tended to be spatially clustered in this population. Our study shows evidence for context‐dependent variation in natal dispersal and corresponding local genetic structure in peripheral populations of this bird. It seems likely that the costs of dispersal are higher in the fragmented and higher density environment in Fray Jorge, particularly for males. The observed differences in microgeographic genetic structure for rayaditos might reflect the genetic consequences of population‐specific responses to contrasting environmental pressures near the range limits of its distribution.