Lake Superior's summer cooling of shorelines and adjacent inland forests: Implications for refugia of boreal forests and disjunct arctic-alpine plants.

Climate refugia can serve as remnant habitat for cold-adapted species and delay forest transitions. The world's largest freshwater lake by surface area, Lake Superior, serves as a model system for understanding cooling-mediated refugia effects, as its cool summer water temperatures have maintained disjunct populations of arctic-alpine plants on its shoreline since deglaciation. It is known to affect local inland climates by providing a summer cooling effect; however, the inland temperature gradient and spatial patterns of cooling have not been well quantified. Here, we describe the extent, degree, and patterns of temperature buffering and examine drivers of buffering and disjunct plant occurrence for Lake Superior's north shore over a 3-year period at distances of 10, 100 m, 1, 10, and 100 km inland. We analyzed temperature data by year, month, summer maximum (July), and growing degree days (GDD0) for each site. Average summertime cooling at shore sites (10 m) was ~5°C cooler than reference sites (100 km inland), with a maximum difference of -19.2°C. The magnitude of cooling varied geographically, with sites further west and southeast showing little to no cooling effect, while the exposed north-central shore showed the highest degree of buffering (5.8°C cooler) and had a shorter growing season than reference sites. Finally, north-central shorelines had fewer days above 16°C, a threshold above which disjunct plants are unlikely to grow. These sites also showed the highest proportion of disjunct arctic-alpine species, reflecting the highest buffering from inland sites. On north-central shores, sites up to 10 km inland had less than 10 days per year warmer than 20°C, a threshold identified for boreal forest transition. An understanding of the extent of lake-mediated cooling on adjacent forests can better inform the risk to disjunct species, inland forests, and vegetation transition models on Lake Superior's north shore.

Mule deer fawn recruitment dynamics in an energy disturbed landscape.

Wildlife population dynamics are modulated by abiotic and biotic factors, typically climate, resource availability, density-dependent effects, and predator-prey interactions. Understanding whether and how human-caused disturbances shape these ecological processes is helpful for the conservation and management of wildlife and their habitats within increasingly human-dominated landscapes. However, many jurisdictions lack either long-term longitudinal data on wildlife populations or measures of the interplay between human-mediated disturbance, climate, and predator density. Here, we use a 50-year time series (1962-2012) on mule deer (Odocoileus hemionus) demographics, seasonal weather, predator density, and oil and gas development patterns from the North Dakota Badlands, USA, to investigate long-term effects of landscape-level disturbance on mule deer fawn fall recruitment, which has declined precipitously over the last number of decades. Mule deer fawn fall recruitment in this study represents the number of fawns per female (fawn:female ratio) that survive through the summer to October. We used this fawn recruitment index to evaluate the composite effects of interannual extreme weather conditions, energy development, and predator density. We found that density-dependent effects and harsh seasonal weather were the main drivers of fawn fall recruitment in the North Dakota Badlands. These effects were further shaped by the interaction between harsh seasonal weather and predator density (i.e., lower fawn fall recruitment when harsh weather was combined with higher predator density). Additionally, we found that fawn fall recruitment was modulated by interactions between seasonal weather and energy development (i.e., lower fawn fall recruitment when harsh weather was combined with higher density of active oil and gas wells). Interestingly, we found that the combined effect of predator density and energy development was not interactive but rather additive. Our analysis demonstrates how energy development may modulate fluctuations in mule deer fawn fall recruitment concurrent with biotic (density-dependency, habitat, predation, woody vegetation encroachment) and abiotic (harsh seasonal weather) drivers. Density-dependent patterns emerge, presumably due to limited quality habitat, being the primary factor influencing fall fawn recruitment in mule deer. Secondarily, stochastic weather events periodically cause dramatic declines in recruitment. And finally, the additive effects of human disturbance and predation can induce fluctuations in fawn fall recruitment. Here we make the case for using long-term datasets for setting long-term wildlife management goals that decision makers and the public can understand and support.

Topographic and vegetation drivers of thermal heterogeneity along the boreal-grassland transition zone in western Canada: Implications for climate change refugia.

Climate change refugia are areas that are relatively buffered from contemporary climate change and may be important safe havens for wildlife and plants under anthropogenic climate change. Topographic variation is an important driver of thermal heterogeneity, but it is limited in relatively flat landscapes, such as the boreal plain and prairie regions of western Canada. Topographic variation within this region is mostly restricted to river valleys and hill systems, and their effects on local climates are not well documented. We sought to quantify thermal heterogeneity as a function of topography and vegetation cover within major valleys and hill systems across the boreal-grassland transition zone. Using iButton data loggers, we monitored local temperature at four hills and 12 river valley systems that comprised a wide range of habitats and ecosystems in Alberta, Canada (N = 240), between 2014 and 2020. We then modeled monthly temperature by season as a function of topography and different vegetation cover types using general linear mixed effect models. Summer maximum temperatures (T max) varied nearly 6°C across the elevation gradient sampled. Local summer mean (T mean) and maximum (T max) temperatures on steep, north-facing slopes (i.e., low levels of potential solar radiation) were up to 0.70°C and 2.90°C cooler than highly exposed areas, respectively. T max in incised valleys was between 0.26 and 0.28°C cooler than other landforms, whereas areas with greater terrain roughness experienced maximum temperatures that were up to 1.62°C cooler. We also found that forest cover buffered temperatures locally, with coniferous and mixedwood forests decreasing summer T mean from 0.23 to 0.72°C and increasing winter T min by up to 2°C, relative to non-forested areas. Spatial predictions of temperatures from iButton data loggers were similar to a gridded climate product (ClimateNA), but the difference between them increased with potential solar radiation, vegetation cover, and terrain roughness. Species that can track their climate niche may be able to compensate for regional climate warming through local migrations to cooler microsites. Topographic and vegetation characteristics that are related to cooler local climates should be considered in the evaluation of future climate change impacts and to identify potential refugia from climate change.

Assessing the nutritional consequences of switching foraging behavior in wood bison.

Diet is one of the most common traits used to organize species of animals into niches. For ruminant herbivores, the breadth and uniqueness of their dietary niche are placed on a spectrum from browsers that consume woody (i.e., browse) and herbaceous (i.e., forbs) plants, to grazers with graminoid-rich diets. However, seasonal changes in plant availability and quality can lead to switching of their dietary niche, even within species. In this study, we examined whether a population of wood bison (Bison bison athabascae) in northeast Alberta, Canada, seasonally switched their foraging behavior, and if so, whether this was associated with changes in nutrient acquisition. We hypothesized that bison should switch foraging behaviors from grazing in the winter when standing, dead graminoids are the only foliar plants readily available to browsing during spring and summer as nutritious and digestible foliar parts of browse and forbs become available. If bison are switching foraging strategy to maximize protein consumption, then there should be a corresponding shift in the nutritional niche. Alternatively, if bison are eating different plants, but consuming similar amounts of nutrients, then bison are switching their dietary niche to maintain a particular nutrient composition. We found wood bison were grazers in the winter and spring, but switch to a browsing during summer. However, only winter nutrient consumption of consumed plants differed significantly among seasons. Between spring and summer, bison maintained a specific nutritional composition in their diet despite compositional differences in the consumed plants. Our evidence suggests that bison are selecting plants to maintain a target macronutrient composition. We posit that herbivore's can and will switch their dietary niche to maintain a target nutrient composition.

A functional perspective on the analysis of land use and land cover data in ecology.

Assessments of large-scale changes in habitat are a priority for management and conservation. Traditional approaches use land use and land cover data (LULC) that focus mostly on "structural" properties of landscapes, rather than "functional" properties related to specific ecological processes. Here, we contend that designing functional analyses of LULC can provide important and complementary information to traditional, structural analyses. We substantiate this perspective with an example of functional changes in habitat due to industrial anthropogenic footprints in Alberta's boreal forest, where there has been little overall forest loss (~ 6% structural change), but high levels of functional change (up to 93% functional change) for species' habitat, biodiversity, and wildfire ignition. We discuss the methods needed to achieve functional LULC analyses, when they are most appropriate to add to structural assessments, and conclude by providing recommendations for analyses of LULC in a future of increasingly high-resolution, dynamic remote sensing data.

Gene flow and climate-associated genetic variation in a vagile habitat specialist.

Previous work in landscape genetics suggests that geographic isolation is of greater importance to genetic divergence than variation in environmental conditions. This is intuitive when configurations of suitable habitat are a dominant factor limiting dispersal and gene flow, but has not been thoroughly examined for habitat specialists with strong dispersal capability. Here, we evaluate the effects of geographic and environmental isolation on genetic divergence for a vagile invertebrate with high habitat specificity and a discrete dispersal life stage: Dod's Old World swallowtail butterfly, Papilio machaon dodi. In Canada, P. m. dodi are generally restricted to eroding habitat along major river valleys where their larval host plant occurs. A series of causal and linear mixed effects models indicate that divergence of genome-wide single nucleotide polymorphisms is best explained by a combination of environmental isolation (variation in summer temperatures) and geographic isolation (Euclidean distance). Interestingly, least-cost path and circuit distances through a resistance surface parameterized as the inverse of habitat suitability were not supported. This suggests that, although habitat associations of many butterflies are specific due to reproductive requirements, habitat suitability and landscape permeability are not equivalent concepts due to considerable adult vagility. We infer that divergent selection related to variation in summer temperatures has produced two genetic clusters within P. m. dodi, differing in voltinism and diapause propensity. Within the next century, temperatures are predicted to rise by amounts greater than the present-day difference between regions of the genetic clusters, potentially affecting the persistence of the northern cluster under continued climate change.

The ecology of human-carnivore coexistence.

With a shrinking supply of wilderness and growing recognition that top predators can have a profound influence on ecosystems, the persistence of large carnivores in human-dominated landscapes has emerged as one of the greatest conservation challenges of our time. Carnivores fascinate society, yet these animals pose threats to people living near them, resulting in high rates of carnivore death near human settlements. We used 41 y of demographic data for more than 2,500 brown bears-one of the world's most widely distributed and conflict-prone carnivores-to understand the behavioral and demographic mechanisms promoting carnivore coexistence in human-dominated landscapes. Bear mortality was high and unsustainable near people, but a human-induced shift to nocturnality facilitated lower risks of bear mortality and rates of conflict with people. Despite these behavioral shifts, projected population growth rates for bears in human-dominated areas revealed a source-sink dynamic. Despite some female bears successfully reproducing in the sink areas, bear persistence was reliant on a supply of immigrants from areas with minimal human influence (i.e., wilderness). Such mechanisms of coexistence reveal a striking paradox: Connectivity to wilderness areas supplies bears that likely will die from people, but these bears are essential to avert local extirpation. These insights suggest carnivores contribute to human-carnivore coexistence through behavioral and demographic mechanisms, and that connected wilderness is critical to sustain coexistence landscapes.

Perceptual Range, Targeting Ability, and Visual Habitat Detection by Greater Fritillary Butterflies Speyeria cybele (Lepidoptera: Nymphalidae) and Speyeria atlantis.

Butterflies are widely invoked as model organisms in studies of metapopulation and dispersal processes. Integral to such investigations are understandings of perceptual range; the maximum distance at which organisms are able to detect patches of suitable habitat. To infer perceptual range, researchers have released butterflies at varying distances from habitat patches and observed their subsequent flight behaviors. It is often assumed that butterflies rely on visual senses for habitat detection; however, this assumption has not been explicitly investigated. Here, we assess the extent and sensory determinants of perceptual range for the great spangled fritillary (Speyeria cybele (Fabricius, 1775)) and Atlantis fritillary (Speyeria atlantis (W.H. Edwards, 1862)). This was achieved by experimentally releasing butterflies over open water at various distances from a lake island, representing an isolated habitat patch in a dichotomous habitat-matrix landscape. To infer whether butterflies rely on vision for habitat detection, we exposed a subset of butterflies to a series of intense light flashes before release to induce flash blindness (bleaching of photoreceptive rhodopsins) without affecting olfaction. Flashed individuals were 30.1 times less likely to successfully navigate to the target island after release, suggesting butterflies rely primarily on visual senses to navigate fragmented landscapes. For unflashed butterflies, the likelihood of successful navigation decreased by a factor of 2.1 for every 10 m increase in release distance. However, no specific distance threshold for perceptual range was observed. We therefore suggest that perceptual range is best viewed as a continuum of probabilities (targeting ability), reflecting the likelihood of habitat detection across a range of distances.

Large carnivore habitat suitability modelling for Romania and associated predictions for protected areas.

Habitat characteristics associated with species occurrences represent important baseline information for wildlife management and conservation, but have rarely been assessed for countries recently joining the EU. We used footprint tracking data and landscape characteristics in Romania to investigate the occurrence of brown bear (Ursus arctos), gray wolf (Canis lupus) and Eurasian lynx (Lynx lynx) and to compare model predictions between Natura 2000 and national-level protected areas (gap analysis). Wolves were more likely to occur where rugged terrain was present. Increasing proportion of forest was positively associated with occurrence of all large carnivores, but forest type (broadleaf, mixed, or conifer) generally varied with carnivore species. Areas where cultivated lands were extensive had little suitable habitat for lynx, whereas bear occurrence probability decreased with increasing proportion of built areas. Pastures were positively associated with wolf and lynx occurrence. Brown bears occurred primarily where national roads with high traffic volumes were at low density, while bears and lynx occurred at medium-high densities of communal roads that had lower traffic volumes. Based on predictions of carnivore distributions, natural areas protected in national parks were most suitable for carnivores, nature parks were less suitable, whereas EU-legislated Natura 2000 sites had the lowest probability of carnivore presence. Our spatially explicit carnivore habitat suitability predictions can be used by managers to amend borders of existing sites, delineate new protected areas, and establish corridors for ecological connectivity. To assist recovery and recolonization, management could also focus on habitat predicted to be suitable but where carnivores were not tracked.

Examining forest resilience to changing fire frequency in a fire-prone region of boreal forest.

Future changes in climate are widely anticipated to increase fire frequency, particularly in boreal forests where extreme warming is expected to occur. Feedbacks between vegetation and fire may modify the direct effects of warming on fire activity and shape ecological responses to changing fire frequency. We investigate these interactions using extensive field data from the Boreal Shield of Saskatchewan, Canada, a region where >40% of the forest has burned in the past 30 years. We use geospatial and field data to assess the resistance and resilience of eight common vegetation states to frequent fire by quantifying the occurrence of short-interval fires and their effect on recovery to a similar vegetation state. These empirical relationships are combined with data from published literature to parameterize a spatially explicit, state-and-transition simulation model of fire and forest succession. We use this model to ask if and how: (a) feedbacks between vegetation and wildfire may modify fire activity on the landscape, and (b) more frequent fire may affect landscape forest composition and age structure. Both field and GIS data suggest the probability of fire is low in the initial decades after fire, supporting the hypothesis that fuel accumulation may exert a negative feedback on fire frequency. Field observations of pre- and postfire composition indicate that switches in forest state are more likely in conifer stands that burn at a young age, supporting the hypothesis that resilience is lower in immature stands. Stands dominated by deciduous trees or jack pine were generally resilient to fire, while mixed conifer and well-drained spruce forests were less resilient. However, simulation modeling suggests increased fire activity may result in large changes in forest age structure and composition, despite the feedbacks between vegetation-fire likely to occur with increased fire activity.

Trophic interactions among vertebrate guilds and plants shape global patterns in species diversity.

Trophic interactions play critical roles in structuring biotic communities. Understanding variation in trophic interactions among systems provides important insights into biodiversity maintenance and conservation. However, the relative importance of bottom-up versus top-down trophic processes for broad-scale patterns in biodiversity is poorly understood. Here, we used global datasets on species richness of vascular plants, mammals and breeding birds to evaluate the role of trophic interactions in shaping large-scale diversity patterns. Specifically, we used non-recursive structural equation models to test for top-down and bottom-up forcing of global species diversity patterns among plants and trophic guilds of mammals and birds (carnivores, invertivores and herbivores), while accounting for extrinsic environmental drivers. The results show that trophic linkages emerged as being more important to explaining species richness than extrinsic environmental drivers. In particular, there were strong, positive top-down interactions between mammal herbivores and plants, and moderate to strong bottom-up and/or top-down interactions between herbivores/invertivores and carnivores. Estimated trophic interactions for separate biogeographical regions were consistent with global patterns. Our findings demonstrate that, after accounting for environmental drivers, large-scale species richness patterns in plant and vertebrate taxa consistently support trophic interactions playing a major role in shaping global patterns in biodiversity. Furthermore, these results suggest that top-down forces often play strong complementary roles relative to bottom-up drivers in structuring biodiversity patterns across trophic levels. These findings underscore the importance of integrating trophic forcing mechanisms into studies of biodiversity patterns.

Combining aggregated and dispersed tree retention harvesting for conservation of vascular plant communities.

Retention harvesting (also called tree retention or structural retention), in which live mature trees are selectively retained within harvested stands at different retention levels and in different patterns (aggregated to dispersed), is increasingly being used to mitigate the negative impacts of forest harvesting on biodiversity. However, the effectiveness of combining different patterns of retention harvesting for conservation and recovery of understory vascular plants in the long term is largely unknown. To address this gap, we compared understory vascular plant diversity, abundance, and composition between aggregated retention and five levels of surrounding dispersed retention (0% [clearcut], 10%, 20%, 50%, 75%) 15 yr postharvest. We also investigated the influence of dispersed retention on the ability of embedded retention patches to support plant communities characteristic of unharvested forests, and whether it varies by patch size of aggregated retention (0.20 ha or 0.46 ha) and position within patches (edge or interior). Species richness, diversity, and cover were higher in the dispersed retention than in the patch retention as the harvested areas favored early-seral plant species. Graminoid cover was greater at the edges than in the interior of large patches. Retention patches as small as 0.2 ha more effectively supported shade-tolerant (forest interior) plant communities when they were surrounded by higher levels of dispersed retention (as compared to patches retained within clearcuts). Overall, the combined use of both aggregated and dispersed retention within a given cutblock benefits both late- and early-seral plant species and thus could effectively conserve understory plant assemblages in harvested landscapes. Sustainable forest management should therefore consider using a range of retention patch sizes combined with varying levels of surrounding dispersed retention in harvest designs to achieve objectives for plant conservation.

Using spatial mark-recapture for conservation monitoring of grizzly bear populations in Alberta.

One of the challenges in conservation is determining patterns and responses in population density and distribution as it relates to habitat and changes in anthropogenic activities. We applied spatially explicit capture recapture (SECR) methods, combined with density surface modelling from five grizzly bear (Ursus arctos) management areas (BMAs) in Alberta, Canada, to assess SECR methods and to explore factors influencing bear distribution. Here we used models of grizzly bear habitat and mortality risk to test local density associations using density surface modelling. Results demonstrated BMA-specific factors influenced density, as well as the effects of habitat and topography on detections and movements of bears. Estimates from SECR were similar to those from closed population models and telemetry data, but with similar or higher levels of precision. Habitat was most associated with areas of higher bear density in the north, whereas mortality risk was most associated (negatively) with density of bears in the south. Comparisons of the distribution of mortality risk and habitat revealed differences by BMA that in turn influenced local abundance of bears. Combining SECR methods with density surface modelling increases the resolution of mark-recapture methods by directly inferring the effect of spatial factors on regulating local densities of animals.

Narrow anthropogenic corridors direct the movement of a generalist boreal butterfly.

Ecological and anthropogenic corridors are becoming more common worldwide, but little is known about how corridor size (width) affects species' movements, and thus their effects. Here we investigated whether 4- and 8-m wide anthropogenic corridors (seismic lines) cleared for petroleum (oil sands) exploration in boreal forests in Alberta, Canada, act on altering the behaviour of a habitat generalist butterfly, the Arctic fritillary (Boloria chariclea). Specifically, we captured 539 Arctic fritillaries and released them in seismic line corridor or control sites with no structural directionality (i.e. forests and clearings), and recorded both their initial direction (along the seismic line or not) and persistence in directional movements. Arctic fritillaries moved inside these lines twice as often as they left them, and maintained their initial direction more often, regardless of line size and independently of forest structure or sex of individuals. Thus, anthropogenic corridors as narrow as 4 m can affect insect movements. Given the vast area of boreal forests disturbed from seismic assessments, investigating if the effects of these dense, localized lines affect population dynamics and species interactions would provide important insights to managing this ecosystem and identifying restoration actions.

Accelerated seed dispersal along linear disturbances in the Canadian oil sands region.

Habitat fragmentation is typically seen as inhibiting movement via erosion in connectivity, although some patterns of early-phase disturbance, such as narrow linear disturbances in otherwise undisturbed forests, may actually facilitate the dispersal of certain species. Such features are common in Alberta's oil sands region as legacies from seismic hydrocarbon exploration used to map oil reserves. Many of the ecological implications of these disturbances are unknown. Here, we investigate the effect of these forest dissections by experimentally testing dispersal patterns along seismic lines compared with adjacent forests using two proxy materials for wind-dispersed seeds, Typha latifolia seed and goose down feathers. We found that wind speeds were up to seven times higher and 95th percentile seed dispersal distances nearly four times farther on seismic lines compared with undisturbed forests and the corresponding effect of these features on seed dispersal distances can be substantial, potentially facilitating future changes in composition and ecological processes in boreal forests. This raises important considerations for native and invasive species, particularly in the context of climate change and the associated importance of seed movement and migration.

Functional macronutritional generalism in a large omnivore, the brown bear.

We combine a recently developed framework for describing dietary generalism with compositional data analysis to examine patterns of omnivory in a large widely distributed mammal. Using the brown bear (Ursus arctos) as a model species, we collected and analyzed data from the literature to estimate the proportions of macronutrients (protein, carbohydrate, and lipid) in the diets of bear populations. Across their range, bears consumed a diversity of foods that resulted in annual population diets that varied in macronutrient proportions, suggesting a wide fundamental macronutrient niche. The variance matrix of pairwise macronutrient log-ratios indicated that the most variable macronutrient among diets was carbohydrate, while protein and lipid were more proportional or codependent (i.e., relatively more constant log-ratios). Populations that consumed anthropogenic foods, such agricultural crops and supplementary feed (e.g., corn), had a higher geometric mean proportion of carbohydrate, and lower proportion of protein, in annual diets. Seasonally, mean diets were lower in protein and higher in carbohydrate, during autumn compared to spring. Populations with anthropogenic subsidies, however, had higher mean proportions of carbohydrate and lower protein, across seasons compared to populations with natural diets. Proportions of macronutrients similar to those selected in experiments by captive brown bears, and which optimized primarily fat mass gain, were observed among hyperphagic prehibernation autumn diets. However, the majority of these were from populations consuming anthropogenic foods, while diets of natural populations were more variable and typically higher in protein. Some anthropogenic diets were close to the proportions selected by captive bears during summer. Our results suggest that omnivory in brown bears is a functional adaptation enabling them to occupy a diverse range of habitats and tolerate variation in the nutritional composition and availability of food resources. Furthermore, we show that populations consuming human-sourced foods have different dietary macronutrient proportions relative to populations with natural diets.

Decoupling habitat fragmentation from habitat loss: butterfly species mobility obscures fragmentation effects in a naturally fragmented landscape of lake islands.

Since the publication of the theory of island biogeography, ecologists have postulated that fragmentation of continuous habitat presents a prominent threat to species diversity. However, negative fragmentation effects may be artifacts; the result of species diversity declining with habitat loss, and habitat loss correlating positively with degree of fragmentation. In this study, we used butterfly assemblages on islands of Lake of the Woods, Ontario, Canada to decouple habitat fragmentation from habitat loss and test two competing hypotheses: (1) the island effect hypothesis, which predicts that decreasing fragment size and increasing fragment isolation reduces species diversity beyond the effects of habitat loss, and (2) the habitat amount hypothesis, which negates fragmentation effects and predicts that only total habitat area determines the diversity of species persisting on fragmented landscapes. Using eight independent size classes of islands (ranging from 0.1 to 8.0 ha) that varied in number of islands while holding total area constant, species diversity comparisons, species accumulation curves, and species-area relationship extrapolations demonstrated that smaller insular habitats contained at least as many butterfly species as continuous habitat. However, when highly mobile species occurring on islands without their larval food plants were excluded from analyses, island effects on potentially reproducing species became apparent. Similarily, generalized linear models suggested that effects of island isolation and vascular plant richness on insular butterfly richness were confounded by species of high mobility. We conclude that inter-fragment movements of highly mobile species may obscure important fragmentation effects on potentially reproducing populations, questioning support for the habitat amount hypothesis.

Species traits modify the species-area relationship in ground-beetle (Coleoptera: Carabidae) assemblages on islands in a boreal lake.

Life-history traits influence colonization, persistence, and extinction of species on islands and are important aspects of theories predicting the geographical distribution and evolution of species. We used data collected from a large freshwater lake (1,413 km2) in central Canada to test the effects of island area and isolation on species richness and abundance of carabid beetles as a function of body size, wing length, and breeding season. A total of 10,018 individual beetles from 37 species were collected during the frost-free period of 2013 using transects of pitfall traps on 30 forested islands ranging in area from 0.2 to 980.7 ha. Life-history traits improved the predictive ability and significantly modified the shape of species-area and abundance-area curves. Abundance and richness of small-bodied (< 13.9 mm), macropterous (winged), and spring-breeding species decreased with island area and increased with isolation. In contrast, richness and abundance of larger-bodied (> 14.0 mm) and flightless species increased with area, but not isolation. Body size of female Carabus taedatus Fabricius, the largest-bodied species, was positively related to island area, while body size on the adjacent mainland was most similar to that on smaller islands. Overall, species with large body size and low dispersal ability, as indicated by flightlessness, were most sensitive to reductions in area. We suggest that large-bodied, flightless species are rare on small islands because habitat is less suitable for them and immigration rates are lower because they depend on freshwater drift for dispersal to islands.

Density-dependent signaling: An alternative hypothesis on the function of chemical signaling in a non-territorial solitary carnivore.

Brown bears are known to use rubbing behavior as a means of chemical communication, but the function of this signaling is unclear. One hypothesis that has gained support is that male bears rub to communicate dominance to other males. We tested the communication of dominance hypothesis in a low-density brown bear population in southeast British Columbia. We contrasted rubbing rates for male and female bears during and after the breeding season using ten years of DNA-mark-recapture data for 643 individuals. Here we demonstrate that male brown bears rub 60% more during the breeding than the non-breeding season, while female rubbing had no seasonal trends. Per capita rub rates by males were, on average, 2.7 times higher than females. Our results suggest that the function of rubbing in the Rocky Mountains may not only be to communicate dominance, but also to self-advertise for mate attraction. We propose that the role of chemical communication in this species may be density-dependent, where the need to self-advertise for mating is inversely related to population density and communicating for dominance increases with population density. We suggest that future endeavors to elucidate the function of rubbing should sample the behavior across a range of population densities using camera trap and genotypic data.

Scale-dependent complementarity of climatic velocity and environmental diversity for identifying priority areas for conservation under climate change.

As most regions of the earth transition to altered climatic conditions, new methods are needed to identify refugia and other areas whose conservation would facilitate persistence of biodiversity under climate change. We compared several common approaches to conservation planning focused on climate resilience over a broad range of ecological settings across North America and evaluated how commonalities in the priority areas identified by different methods varied with regional context and spatial scale. Our results indicate that priority areas based on different environmental diversity metrics differed substantially from each other and from priorities based on spatiotemporal metrics such as climatic velocity. Refugia identified by diversity or velocity metrics were not strongly associated with the current protected area system, suggesting the need for additional conservation measures including protection of refugia. Despite the inherent uncertainties in predicting future climate, we found that variation among climatic velocities derived from different general circulation models and emissions pathways was less than the variation among the suite of environmental diversity metrics. To address uncertainty created by this variation, planners can combine priorities identified by alternative metrics at a single resolution and downweight areas of high variation between metrics. Alternately, coarse-resolution velocity metrics can be combined with fine-resolution diversity metrics in order to leverage the respective strengths of the two groups of metrics as tools for identification of potential macro- and microrefugia that in combination maximize both transient and long-term resilience to climate change. Planners should compare and integrate approaches that span a range of model complexity and spatial scale to match the range of ecological and physical processes influencing persistence of biodiversity and identify a conservation network resilient to threats operating at multiple scales.