Hugh S. Robinson

Neonatal mortality of elk driven by climate, predator phenology and predator community composition

1. Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2. Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3. Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4. In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5. Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.

Food availability and foraging near human developments by black bears

Abstract Understanding the relationship between foraging ecology and the presence of human-dominated landscapes is important, particularly for American black bears (Ursus americanus), which sometimes move between wildlands and urban areas to forage. The food-related factors influencing this movement have not been explored, but can be important for understanding the benefits and costs to black bear foraging behavior and the fundamental origins of bear conflicts. We tested whether the scarcity of wildland foods or the availability of urban foods can explain when black bears forage near houses, examined the extent to which male bears use urban areas in comparison to females, and identified the most important food items influencing bear movement into urban areas. We monitored 16 collared black bears in and around Missoula, Montana, during 2009 and 2010, while quantifying the rate of change in green vegetation and the availability of 5 native berry-producing species outside the urban area, the rate of change in green vegetation, and the availability of apples and garbage inside the urban area. We used parametric time-to-event models in which an event was a bear location collected within 100 m of a house. We also visited feeding sites located near houses and quantified food items bears had eaten. The probability of a bear being located near a house was 1.6 times higher for males, and increased during apple season and the urban green-up. Fruit trees accounted for most of the forage items at urban feeding sites (49%), whereas wildland foods composed <10%. Black bears foraged on human foods near houses even when wildland foods were available, suggesting that the absence of wildland foods may not influence the probability of bears foraging near houses. Additionally, other attractants, in this case fruit trees, appear to be more important than the availability of garbage in influencing when bears forage near houses.

The Role of Translocation in Recovery of Woodland Caribou Populations

Maintenance of viable populations of many endangered species will require conservation action in perpetuity. Efforts to conserve these species are more likely to be successful if their reliance on conservation actions is assessed at the population level. Woodland caribou (Rangifer tarandus caribou) were extirpated recently from Banff National Park, Canada, and translocations of caribou to Banff and neighboring Jasper National Park are being considered. We used population viability analysis to assess the relative need for and benefits from translocation of individuals among caribou populations. We measured stochastic growth rates and the probability of quasi extinction of four populations of woodland caribou with and without translocation. We used two vital rates in our analysis: mean adult female survival and mean number of calves per breeding-age female as estimates of mean fecundity. We isolated process variance for each vital rate. Our results suggested the Tonquin caribou population in Jasper is likely to remain viable without translocation, but that translocation is probably insufficient to prevent eventual extirpation of the two other populations in Jasper. Simulated reintroductions of caribou into Banff resulted in a 53-98% probability of >8 females remaining after 20 years, which suggests translocation may be an effective recovery tool for some caribou populations.

Structural habitat predicts functional dispersal habitat of a large carnivore: how leopards change spots

Natal dispersal promotes inter-population linkage, and is key to spatial distribution of populations. Degradation of suitable landscape structures beyond the specific threshold of an individuals ability to disperse can therefore lead to disruption of functional landscape connectivity and impact metapopulation function. Because it ignores behavioral responses of individuals, structural connectivity is easier to assess than functional connectivity and is often used as a surrogate for landscape connectivity modeling. However using structural resource selection models as surrogate for modeling functional connectivity through dispersal could be erroneous. We tested how well a second-order resource selection function (RSF) models (structural connectivity), based on GPS telemetry data from resident adult leopard (Panthera pardus L.), could predict subadult habitat use during dispersal (functional connectivity). We created eight non-exclusive subsets of the subadult data based on differing definitions of dispersal to assess the predictive ability of our adult-based RSF model extrapolated over a broader landscape. Dispersing leopards used habitats in accordance with adult selection patterns, regardless of the definition of dispersal considered. We demonstrate that, for a wide-ranging apex carnivore, functional connectivity through natal dispersal corresponds to structural connectivity as modeled by a second-order RSF. Mapping of the adult-based habitat classes provides direct visualization of the potential linkages between populations, without the need to model paths between a priori starting and destination points. The use of such landscape scale RSFs may provide insight into predicting suitable dispersal habitat peninsulas in human-dominated landscapes where mitigation of human-wildlife conflict should be focused. We recommend the use of second-order RSFs for landscape conservation planning and propose a similar approach to the conservation of other wide-ranging large carnivore species where landscape-scale resource selection data already exist.

Integrating resource selection into spatial capture‐recapture models for large carnivores

Wildlife managers need reliable methods to estimate large carnivore densities and population trends; yet large carnivores are elusive, difficult to detect, and occur at low densities making traditional approaches intractable. Recent advances in spatial capture-recapture (SCR) models have provided new approaches for monitoring trends in wildlife abundance and these methods are particularly applicable to large carnivores. We applied SCR models in a Bayesian framework to estimate mountain lion densities in the Bitterroot Mountains of west central Montana. We incorporate an existing resource selection function (RSF) as a density covariate to account for heterogeneity in habitat use across the study area and include data collected from harvested lions. We identify individuals through DNA samples collected by (1) biopsy darting mountain lions detected in systematic surveys of the study area, (2) opportunistically collecting hair and scat samples, and (3) sampling all harvested mountain lions. We included 80 DNA...

Mortality of Amur tigers: The more things change, the more they stay the same

Poaching as well as loss of habitat and prey are identified as causes of tiger population declines. Although some studies have examined habitat requirements and prey availability, few studies have quantified cause-specific mortality of tigers. We used cumulative incidence functions (CIFs) to quantify cause-specific mortality rates of tigers, expanding and refining earlier studies to assess the potential impact of a newly emerging disease. To quantify changes in tiger mortality over time, we re-examined data first collected by Goodrich et al. (; study period 1: 1992-2004) as well as new telemetry data collected since January 2005 (study period 2: 2005-2012) using a total of 57 tigers (27 males and 30 females) monitored for an average of 747 days (range 26-4718 days). Across the entire study period (1992 to 2012) we found an estimated average annual survival rate of 0.75 for all tigers combined. Poaching was the primary cause of mortality during both study periods, followed by suspected poaching, distemper and natural/unknown causes. Since 2005, poaching mortality has remained relatively constant and, if combined with suspected poaching, may account for a loss of 17-19% of the population each year. Canine distemper virus (CDV) may be an additive form of mortality to the population, currently accounting for an additional 5%. Despite this relatively new source of mortality, poaching remains the main threat to Amur tiger survival and, therefore, population growth.

Cats, connectivity and conservation: incorporating data sets and integrating scales for wildlife management

Understanding resource selection and quantifying habitat connectivity are fundamental to conservation planning for both land-use and species management plans. However, datasets available to management authorities for resource selection and connectivity analyses are often highly limited and fragmentary. As a result, measuring connectivity is challenging, and often poorly integrated within conservation planning and wildlife management. To exacerbate the challenge, scale-dependent resource use makes inference across scales problematic, resource use is often modelled in areas where the species is not present, and connectivity is typically measured using a source-to-sink approach, erroneously assuming animals possess predefined destinations. Here, we used a large carnivore, the leopard Panthera pardus, to characterise resource use and landscape connectivity across a vast, biodiverse region of southern Africa. Using a range of datasets to counter data deficiencies inherent in carnivore management, we overcame methodological limitations by employing occupancy modelling and resource selection functions across three orders of selection, and estimated landscape-scale habitat connectivity – independent of a priori source and sink locations – using circuit theory. We evaluated whether occupancy modelling on its own was capable of accurately informing habitat connectivity, and identified conservation priorities necessary for applied management. We detected markedly different scale-dependent relationships across all selection orders. Our multi-data, multi-scale approach accurately predicted resource use across multiple scales and demonstrates how management authorities can more suitably utilise fragmentary datasets. We further developed an unbiased landscape-scale depiction of habitat connectivity, and identified key linkages in need of targeted management. We did not find support for the use of occupancy modelling as a proxy for landscape-scale habitat connectivity and further caution its use within a management context. Synthesis and applications. Maintaining habitat connectivity remains a fundamental component of wildlife management and conservation, yet data to inform these biological and ecological processes are often scarce. We present a robust approach that incorporates multi-scale fragmentary datasets (e.g. mortality data, permit data, sightings data), routinely collected by management authorities, to inform wildlife management and land-use planning. We recommend that management authorities employ a multi-data, multi-scale connectivity approach—as we present here—to identify management units at risk of low connectivity.

Determinants of Distribution Patterns and Management Needs in a Critically Endangered Lion Panthera leo Population

The lion Panthera leo is Critically Endangered in West Africa and is known to occupy only four protected areas within the region. The largest population persists in the trans-boundary W-Arly-Pendjari (WAP) ecosystem, in the border region of Benin, Burkina Faso, and Niger. WAP harbors an estimated 350 individuals, or 90% of West Africa’s lions. We modeled lion occupancy across WAP using systematic, vehicle-based spoor counts to assess how landscape variables related to biotic factors, management and human impact influence lion distribution across WAP. We surveyed 1110 km of roads across WAP in 2012, obtaining 79 lion detections in 32 of our 167 15 x 15 km sampling units (naive occupancy = 0.41). Overall occupancy (Ψ) was 0.71 (95% SE = 0.56-0.83) when accounting for imperfect detection (p = 0.22, 95% SE = 0.18-0.27). The best predictors of lion occupancy were numbers of permanent protected area staff and mean monthly dry season precipitation. Model-averaged estimates suggest greatest lion occupancy in the Arly and Pendjari management blocks, with lowest occupancy in the tri-national W National Park. Our results suggest that lions in WAP are equally limited by management and biotic factors, and demonstrate how unevenly distributed protection effort limits the distribution of an apex predator across a protected landscape. We strongly recommend increased funding and better protection to increase lion occupancy in WAP, most urgently in the W National Park.

Estimating abundance and density of Amur tigers along the Sino–Russian border

As an apex predator the Amur tiger (Panthera tigris altaica) could play a pivotal role in maintaining the integrity of forest ecosystems in Northeast Asia. Due to habitat loss and harvest over the past century, tigers rapidly declined in China and are now restricted to the Russian Far East and bordering habitat in nearby China. To facilitate restoration of the tiger in its historical range, reliable estimates of population size are essential to assess effectiveness of conservation interventions. Here we used camera trap data collected in Hunchun National Nature Reserve from April to June 2013 and 2014 to estimate tiger density and abundance using both maximum likelihood and Bayesian spatially explicit capture-recapture (SECR) methods. A minimum of 8 individuals were detected in both sample periods and the documentation of marking behavior and reproduction suggests the presence of a resident population. Using Bayesian SECR modeling within the 11 400 km(2) state space, density estimates were 0.33 and 0.40 individuals/100 km(2) in 2013 and 2014, respectively, corresponding to an estimated abundance of 38 and 45 animals for this transboundary Sino-Russian population. In a maximum likelihood framework, we estimated densities of 0.30 and 0.24 individuals/100 km(2) corresponding to abundances of 34 and 27, in 2013 and 2014, respectively. These density estimates are comparable to other published estimates for resident Amur tiger populations in the Russian Far East. This study reveals promising signs of tiger recovery in Northeast China, and demonstrates the importance of connectivity between the Russian and Chinese populations for recovering tigers in Northeast China.

Leopard distribution and abundance is unaffected by interference competition with lions

Lay SummaryWe tested the population-level effects of a dominant competitor—lions—on a subordinate—leopards. Although lions were a common cause of leopard mortality, they did not suppress or displace leopards. Population growth was similar between the 2 species and leopards did not avoid lions. Coexistence was likely possible because lions and leopards targeted different sized prey. Widespread poaching of large ungulates may increase levels of competition between lions and leopards by forcing them to target the same prey.