Habitat and climatic associations of climate-sensitive species along a southern range boundary.

Climate change and habitat loss are recognized as important drivers of shifts in wildlife species' geographic distributions. While often considered independently, there is considerable overlap between these drivers, and understanding how they contribute to range shifts can predict future species assemblages and inform effective management. Our objective was to evaluate the impacts of habitat, climatic, and anthropogenic effects on the distributions of climate-sensitive vertebrates along a southern range boundary in Northern Michigan, USA. We combined multiple sources of occurrence data, including harvest and citizen-science data, then used hierarchical Bayesian spatial models to determine habitat and climatic associations for four climate-sensitive vertebrate species (American marten [Martes americana], snowshoe hare [Lepus americanus], ruffed grouse [Bonasa umbellus] and moose [Alces alces]). We used total basal area of at-risk forest types to represent habitat, and temperature and winter habitat indices to represent climate. Marten associated with upland spruce-fir and lowland riparian forest types, hares with lowland conifer and aspen-birch, grouse with lowland riparian hardwoods, and moose with upland spruce-fir. Species differed in climatic drivers with hares positively associated with cooler annual temperatures, moose with cooler summer temperatures and grouse with colder winter temperatures. Contrary to expectations, temperature variables outperformed winter habitat indices. Model performance varied greatly among species, as did predicted distributions along the southern edge of the Northwoods region. As multiple species were associated with lowland riparian and upland spruce-fir habitats, these results provide potential for efficient prioritization of habitat management. Both direct and indirect effects from climate change are likely to impact the distribution of climate-sensitive species in the future and the use of multiple data types and sources in the modelling of species distributions can result in more accurate predictions resulting in improved management at policy-relevant scales.

Short-term responses of small mammal diversity to varying stand-scale patterns of retention tree patches.

Retention forestry is a common practice for biodiversity conservation in forests managed for wood production. Retention forestry often leaves unharvested patches of trees that vary in size and spatial pattern but experiments evaluating the effects of different retention patch configurations at a constant level of retention are lacking for many regions and taxonomic groups. We implemented an experimental study in clearcut conifer stands with retention across the U.S. Pacific Northwest region. The study consisted of five stand-level (11-55 ha) experimental treatments each replicated 10 times within a randomized complete block design, resulting in 50 treated stands. Retained tree density was comparable across treatments but size, number, and location (upland or riparian) of patches within stands varied among the five treatments. Within experimental treatments, we measured small mammal (<1kg) species and functional trait (i.e., body size, diet, activity stratum) richness in retention patches, surrounding harvested portions of stands, and nearby unharvested stands. We evaluated species and functional trait richness by treatment using generalized linear mixed-effects models and species-specific responses to retention placement using a community occupancy model. We obtained repeat captures of 21 species of small mammals but found limited evidence of a treatment effect on species richness, and no differences in functional trait richness. Species richness was highest where all retained trees were aggregated into one patch placed adjacent to a forested riparian buffer (mean = 6.6 species, 95% CI = 5.7-7.5), and lowest in the treatment containing one retention patch in the upland portion of a harvested stand (mean = 4.7 species, 95% CI = 3.8-5.6). Furthermore, estimates of species richness within retention patches of harvested stands (i.e., not considering species in harvested areas) did not differ among treatments, indicating that the slightly elevated species richness in riparian-associated retention results from 1-2 species in these patches that do not occur in adjacent harvested portions of each treated stand. Patch occupancy of several species was higher in riparian patches than harvested portions of the treated stands, and fewer species had higher occupancy in upland patches compared to harvested portions of treated stands. Our results indicated that at retention densities currently required in Oregon and Washington, the location of retention patches had a small influence on stand-scale measures of small mammal diversity, but local increases in species richness may be obtained by retaining trees adjacent to riparian buffers.

Use of environmental DNA (eDNA) in streams to detect feral swine (Sus scrofa).

Invasive feral swine can damage ecosystems, disrupt plant and animal populations, and transmit diseases. Monitoring of feral swine populations requires expensive and labor-intensive techniques such as aerial surveys, field surveys for sign, trail cameras, and verifying landowner reports. Environmental DNA (eDNA) provides an alternative method for locating feral swine. To aid in detection of this harmful invasive species, a novel assay was developed incorporating molecular methods. From August 2017 to April 2018, water samples and stream data were collected along 400 m transects in two different stream types where swine DNA was artificially introduced to investigate potential factors affecting detection. A generalized linear model (family binomial) was used to characterize environmental conditions affecting swine DNA detection; detection was the dependent variable and stream measurements included stream type, distance downstream, water temperature, velocity, turbidity, discharge, and pH as independent variables. Parameters from the generalized linear model were deemed significant if 95% confidence intervals did not overlap 0. Detection probability for swine DNA negatively related to water temperature (ß =  - 0.21, 95% CI [-0.35 to -0.09]), with the highest detection probability (0.80) at 0 °C and lowest detection probability (0.05) at 17.9 °C water temperature. Results indicate that sampling for swine eDNA in free-flowing stream systems should occur at lower water temperatures to maximize detection probability. This study provides a foundation for further development of field and sampling techniques for utilizing eDNA as a viable alternative to monitoring a terrestrial invasive species in northern regions of the United States.

Multispecies hierarchical modeling reveals variable responses of African carnivores to management alternatives.

Carnivore communities face unprecedented threats from humans. Yet, management regimes have variable effects on carnivores, where species may persist or decline in response to direct or indirect changes to the ecosystem. Using a hierarchical multispecies modeling approach, we examined the effects of alternative management regimes (i.e., active vs. passive enforcement of regulations) on carnivore abundances and group sizes at both species and community levels in the Masai Mara National Reserve, Kenya. Alternative management regimes have created a dichotomy in ecosystem conditions within the Reserve, where active enforcement of regulations maintains low levels of human disturbance in the Mara Triangle and passive enforcement of regulations in the Talek region permits multiple forms of human disturbance. Our results demonstrate that these alternative management regimes have variable effects on 11 observed carnivore species. As predicted, some species, such as African lions and bat-eared foxes, have higher population densities in the Mara Triangle, where regulations are actively enforced. Yet, other species, including black-backed jackals and spotted hyenas, have higher population densities in the Talek region where enforcement is passive. Multiple underlying mechanisms, including behavioral plasticity and competitive release, are likely causing higher black-backed jackals and spotted hyena densities in the disturbed Talek region. Our multispecies modeling framework reveals that carnivores do not react to management regimes uniformly, shaping carnivore communities by differentially producing winning and losing species. Some carnivore species require active enforcement of regulations for effective conservation, while others more readily adapt (and in some instances thrive in response) to lax management enforcement and resulting anthropogenic disturbance. Yet, high levels of human disturbance appear to be negatively affecting the majority of carnivores, with potential consequences that may permeate throughout the rest of the ecosystem. Community approaches to monitoring carnivores should be adopted as single species monitoring may overlook important intra-community variability.

Improving inferences from short-term ecological studies with Bayesian hierarchical modeling: white-headed woodpeckers in managed forests.

Pilot studies are often used to design short-term research projects and long-term ecological monitoring programs, but data are sometimes discarded when they do not match the eventual survey design. Bayesian hierarchical modeling provides a convenient framework for integrating multiple data sources while explicitly separating sample variation into observation and ecological state processes. Such an approach can better estimate state uncertainty and improve inferences from short-term studies in dynamic systems. We used a dynamic multistate occupancy model to estimate the probabilities of occurrence and nesting for white-headed woodpeckers Picoides albolarvatus in recent harvest units within managed forests of northern California, USA. Our objectives were to examine how occupancy states and state transitions were related to forest management practices, and how the probabilities changed over time. Using Gibbs variable selection, we made inferences using multiple model structures and generated model-averaged estimates. Probabilities of white-headed woodpecker occurrence and nesting were high in 2009 and 2010, and the probability that nesting persisted at a site was positively related to the snag density in harvest units. Prior-year nesting resulted in higher probabilities of subsequent occurrence and nesting. We demonstrate the benefit of forest management practices that increase the density of retained snags in harvest units for providing white-headed woodpecker nesting habitat. While including an additional year of data from our pilot study did not drastically alter management recommendations, it changed the interpretation of the mechanism behind the observed dynamics. Bayesian hierarchical modeling has the potential to maximize the utility of studies based on small sample sizes while fully accounting for measurement error and both estimation and model uncertainty, thereby improving the ability of observational data to inform conservation and management strategies.

Comparing population patterns to processes: abundance and survival of a forest salamander following habitat degradation.

Habitat degradation resulting from anthropogenic activities poses immediate and prolonged threats to biodiversity, particularly among declining amphibians. Many studies infer amphibian response to habitat degradation by correlating patterns in species occupancy or abundance with environmental effects, often without regard to the demographic processes underlying these patterns. We evaluated how retention of vertical green trees (CANOPY) and coarse woody debris (CWD) influenced terrestrial salamander abundance and apparent survival in recently clearcut forests. Estimated abundance of unmarked salamanders was positively related to CANOPY (ß Canopy  = 0.21 (0.02-1.19; 95% CI), but not CWD (ß CWD  = 0.11 (-0.13-0.35) within 3,600 m2 sites, whereas estimated abundance of unmarked salamanders was not related to CANOPY (ß Canopy  = -0.01 (-0.21-0.18) or CWD (ß CWD  = -0.02 (-0.23-0.19) for 9 m2 enclosures. In contrast, apparent survival of marked salamanders within our enclosures over 1 month was positively influenced by both CANOPY and CWD retention (ß Canopy  = 0.73 (0.27-1.19; 95% CI) and ß CWD  = 1.01 (0.53-1.50). Our results indicate that environmental correlates to abundance are scale dependent reflecting habitat selection processes and organism movements after a habitat disturbance event. Our study also provides a cautionary example of how scientific inference is conditional on the response variable(s), and scale(s) of measure chosen by the investigator, which can have important implications for species conservation and management. Our research highlights the need for joint evaluation of population state variables, such as abundance, and population-level process, such as survival, when assessing anthropogenic impacts on forest biodiversity.

Where wolves kill moose: the influence of prey life history dynamics on the landscape ecology of predation.

The landscape ecology of predation is well studied and known to be influenced by habitat heterogeneity. Little attention has been given to how the influence of habitat heterogeneity on the landscape ecology of predation might be modulated by life history dynamics of prey in mammalian systems. We demonstrate how life history dynamics of moose (Alces alces) contribute to landscape patterns in predation by wolves (Canis lupus) in Isle Royale National Park, Lake Superior, USA. We use pattern analysis and kernel density estimates of moose kill sites to demonstrate that moose in senescent condition and moose in prime condition tend to be wolf-killed in different regions of Isle Royale in winter. Predation on senescent moose was clustered in one kill zone in the northeast portion of the island, whereas predation on prime moose was clustered in 13 separate kill zones distributed throughout the full extent of the island. Moreover, the probability of kill occurrence for senescent moose, in comparison to prime moose, increased in high elevation habitat with patches of dense coniferous trees. These differences can be attributed, at least in part, to senescent moose being more vulnerable to predation and making different risk-sensitive habitat decisions than prime moose. Landscape patterns emerging from prey life history dynamics and habitat heterogeneity have been observed in the predation ecology of fish and insects, but this is the first mammalian system for which such observations have been made.

The influence of winter severity, predation and senescence on moose habitat use.

Habitat use is widely known to be influenced by abiotic and biotic factors, such as climate, population density, foraging opportunity and predation risk. The influence of the life-history state of an individual organism on habitat use is less well understood, especially for terrestrial mammals. There is good reason to expect that life-history state would affect habitat use. For example, organisms exhibiting poor condition associated with senescence have an increased vulnerability to predation and that vulnerability is known to alter habitat use strategies. We assessed the influence of life-history stage on habitat use for 732 moose (Alces alces) killed by wolves (Canis lupus) over a 50-year period in Isle Royale National Park, an island ecosystem in Lake Superior, USA. We developed regression models to assess how location of death was associated with a moose's life-history stage (prime-aged or senescent), presence or absence of senescent-associated pathology (osteoarthritis and jaw necrosis), and annual variation in winter severity, moose density and ratio of moose to wolves, which is an index of predation risk. Compared to senescent moose, prime-aged moose tend to make greater use of habitat farther from the shoreline of Isle Royale. That result is ecologically relevant because shoreline habitat on Isle Royale tends to provide better foraging opportunities for moose but is also associated with increased predation risk. During severe winters prime-aged moose tend to make greater use of habitat that is closer to shore in relation to senescent-aged moose. Furthermore, moose of both age classes were more likely to die in riskier, shoreline habitat during years when predation risk was lower in the preceding year. Our results highlight a complicated connection between life history, age-structured population dynamics and habitat-related behaviour. Our analysis also illustrates why intraspecific competition should not be the presumed mechanism underlying density-dependent habitat use, if predation risk is related to density, as it is expected to be in many systems.

Poaching risks in community-based natural resource management.

Poaching can disrupt wildlife-management efforts in community-based natural resource management systems. Monitoring, estimating, and acquiring data on poaching is difficult. We used local-stakeholder knowledge and poaching records to rank and map the risk of poaching incidents in 2 areas where natural resources are managed by community members in Caprivi, Namibia. We mapped local stakeholder perceptions of the risk of poaching, risk of wildlife damage to livelihoods, and wildlife distribution and compared these maps with spatially explicit records of poaching events. Recorded poaching events and stakeholder perceptions of where poaching occurred were not spatially correlated. However, the locations of documented poaching events were spatially correlated with areas that stakeholders perceived wildlife as a threat to their livelihoods. This result suggests poaching occurred in response to wildlife damage occurred. Local stakeholders thought that wildlife populations were at high risk of being poached and that poaching occurred where there was abundant wildlife. These findings suggest stakeholders were concerned about wildlife resources in their community and indicate a need for integrated and continued monitoring of poaching activities and further interventions at the wildlife-agricultural interface. Involving stakeholders in the assessment of poaching risks promotes their participation in local conservation efforts, a central tenet of community-based management. We considered stakeholders poaching informants, rather than suspects, and our technique was spatially explicit. Different strategies to reduce poaching are likely needed in different areas. For example, interventions that reduce human-wildlife conflict may be required in residential areas, and increased and targeted patrolling may be required in more remote areas. Stakeholder-generated maps of human-wildlife interactions may be a valuable enforcement and intervention support tool.

Modeling protected species habitat and assigning risk to inform regulatory decisions.

In the United States, environmental regulatory agencies are required to use "best available" scientific information when making decisions on a variety of issues. However, agencies are often hindered by coarse or incomplete data, particularly as it pertains to threatened and endangered species protection. Stakeholders often agree that more resolute and integrated processes for decision-making are desirable. We demonstrate a process that uses species occurrence data for a federally endangered insect (Karner blue butterfly), a readily available habitat modeling tool, and spatially explicit information about an important Michigan commodity (tart cherries). This case study has characteristics of many protected species regulatory decisions in that species occurrence data were sparse and unequally distributed; regulatory decisions (on pesticide use) were required with potentially significant impacts on a viable agricultural industry; and stakeholder relations were diverse, misinformed, and, in some situations, unjustly contentious. Results from our process include a large-scale, empirically derived habitat suitability map for the focal species and a risk ranking of tart cherry orchards with risk based on the likelihood that pesticide applications will influence the focal protected species. Although the majority (77%) of pesticide-influence zones overlapped Karner blue butterfly habitat, risk scores associated with each orchard were low. Through our process we demonstrated that spatially explicit models can help stakeholders visualize and quantify potential protected species effects. In addition, model outputs can serve to guide field activities (e.g., species surveys and implementation of pesticide buffer zones) that help minimize future effects.