Estimating spatially variable and density-dependent survival using open-population spatial capture-recapture models.

Open-population spatial capture-recapture (OPSCR) models use the spatial information contained in individual detections collected over multiple consecutive occasions to estimate not only occasion-specific density, but also demographic parameters. OPSCR models can also estimate spatial variation in vital rates, but such models are neither widely used nor thoroughly tested. We developed a Bayesian OPSCR model that not only accounts for spatial variation in survival using spatial covariates but also estimates local density-dependent effects on survival within a unified framework. Using simulations, we show that OPSCR models provide sound inferences on the effect of spatial covariates on survival, including multiple competing sources of mortality, each with potentially different spatial determinants. Estimation of local density-dependent survival was possible but required more data due to the greater complexity of the model. Not accounting for spatial heterogeneity in survival led to up to 10% positive bias in abundance estimates. We provide an empirical demonstration of the model by estimating the effect of country and density on cause-specific mortality of female wolverines (Gulo gulo) in central Sweden and Norway. The ability to make population-level inferences on spatial variation in survival is an essential step toward a fully spatially explicit OPSCR model capable of disentangling the role of multiple spatial drivers of population dynamics.

GPS collars have an apparent positive effect on the survival of a large carnivore.

Are instrumented animals representative of the population, given the potential bias caused by selective sampling and the influence of capture, handling and wearing bio-loggers? The answer is elusive owing to the challenges of obtaining comparable data from individuals with and without bio-loggers. Using non-invasive genetic data of a large carnivore, the wolverine (Gulo gulo) in Scandinavia, and an open-population spatial capture-recapture model, we found a 16 (credible interval: 4-30) percentage points lower mortality probability for GPS-collared individuals compared with individuals without GPS collars. While the risk of dying from legal culling was comparable for collared and non-collared wolverines, the former experienced lower probability of mortality due to causes other than legal culling. The aforementioned effect was pronounced despite a potentially lower age-and therefore likely higher natural mortality-of collared individuals. Reports of positive effects of bio-loggers on the survival of individuals are uncommon and we argue that GPS collars could shield animals from poaching. Our results highlight the challenges of drawing population-level inferences for populations subjected to poaching when using data from instrumented individuals.

Estimating and forecasting spatial population dynamics of apex predators using transnational genetic monitoring.

The ongoing recovery of terrestrial large carnivores in North America and Europe is accompanied by intense controversy. On the one hand, reestablishment of large carnivores entails a recovery of their most important ecological role, predation. On the other hand, societies are struggling to relearn how to live with apex predators that kill livestock, compete for game species, and occasionally injure or kill people. Those responsible for managing these species and mitigating conflict often lack fundamental information due to a long-standing challenge in ecology: How do we draw robust population-level inferences for elusive animals spread over immense areas? Here we showcase the application of an effective tool for spatially explicit tracking and forecasting of wildlife population dynamics at scales that are relevant to management and conservation. We analyzed the world's largest dataset on carnivores comprising more than 35,000 noninvasively obtained DNA samples from over 6,000 individual brown bears (Ursus arctos), gray wolves (Canis lupus), and wolverines (Gulo gulo). Our analyses took into account that not all individuals are detected and, even if detected, their fates are not always known. We show unequivocal quantitative evidence of large carnivore recovery in northern Europe, juxtaposed with the finding that humans are the single-most important factor driving the dynamics of these apex predators. We present maps and forecasts of the spatiotemporal dynamics of large carnivore populations, transcending national boundaries and management regimes.

Harvest models of small populations of a large carnivore using Bayesian forecasting.

Harvesting large carnivores can be a management tool for meeting politically set goals for their desired abundance. However, harvesting carnivores creates its own set of conflicts in both society and among conservation professionals, where one consequence is a need to demonstrate that management is sustainable, evidence-based, and guided by science. Furthermore, because large carnivores often also have high degrees of legal protection, harvest quotas have to be carefully justified and constantly adjusted to avoid damaging their conservation status. We developed a Bayesian state-space model to support adaptive management of Eurasian lynx harvesting in Scandinavia. The model uses data from the annual monitoring of lynx abundance and results from long-term field research on lynx biology, which has provided detailed estimates of key demographic parameters. We used the model to predict the probability that the forecasted population size will be below or above the management objectives when subjected to different harvest quotas. The model presented here informs decision makers about the policy risks of alternative harvest levels. Earlier versions of the model have been available for wildlife managers in both Sweden and Norway to guide lynx harvest quotas and the model predictions showed good agreement with observations. We combined monitoring data with data on vital rates and were able to estimate unobserved additional mortality rates, which are most probably due to poaching. In both countries, the past quota setting strategy suggests that there has been a de facto threshold strategy with increasing proportion, which means that there is no harvest below a certain population size, but above this threshold there is an increasing proportion of the population harvested as the population size increases. The annual assessment of the monitoring results, the use of forecasting models, and a threshold harvest approach to quota setting will all reduce the risk of lynx population sizes moving outside the desired goals. The approach we illustrate could be adapted to other populations of mammals worldwide.

A local evaluation of the individual state-space to scale up Bayesian spatial capture-recapture.

Spatial capture-recapture models (SCR) are used to estimate animal density and to investigate a range of problems in spatial ecology that cannot be addressed with traditional nonspatial methods. Bayesian approaches in particular offer tremendous flexibility for SCR modeling. Increasingly, SCR data are being collected over very large spatial extents making analysis computational intensive, sometimes prohibitively so. To mitigate the computational burden of large-scale SCR models, we developed an improved formulation of the Bayesian SCR model that uses local evaluation of the individual state-space (LESS). Based on prior knowledge about a species' home range size, we created square evaluation windows that restrict the spatial domain in which an individual's detection probability (detector window) and activity center location (AC window) are estimated. We used simulations and empirical data analyses to assess the performance and bias of SCR with LESS. LESS produced unbiased estimates of SCR parameters when the AC window width was ≥5σ (σ: the scale parameter of the half-normal detection function), and when the detector window extended beyond the edge of the AC window by 2σ. Importantly, LESS considerably decreased the computation time needed for fitting SCR models. In our simulations, LESS increased the computation speed of SCR models up to 57-fold. We demonstrate the power of this new approach by mapping the density of an elusive large carnivore-the wolverine (Gulo gulo)-with an unprecedented resolution and across the species' entire range in Norway (> 200,000 km2). Our approach helps overcome a major computational obstacle to population and landscape-level SCR analyses. The LESS implementation in a Bayesian framework makes the customization and fitting of SCR accessible for practitioners working at scales that are relevant for conservation and management.

Noninvasive genetic sampling reveals intrasex territoriality in wolverines.

Due to its conspicuous manifestations and its capacity to shape the configuration and dynamics of wild populations, territorial behavior has long intrigued ecologists. Territoriality and other animal interactions in situ have traditionally been studied via direct observations and telemetry. Here, we explore whether noninvasive genetic sampling, which is increasingly supplementing traditional field methods in ecological research, can reveal territorial behavior in an elusive carnivore, the wolverine (Gulo gulo). Using the locations of genotyped wolverine scat samples collected annually over a period of 12 years in central Norway, we test three predictions: (1) male home ranges constructed from noninvasive genetic sampling data are larger than those of females, (2) individuals avoid areas used by other conspecifics of the same sex (intrasexual territoriality), and (3) avoidance of same-sex territories diminishes or disappears after the territory owner's death. Each of these predictions is substantiated by our results: sex-specific differences in home range size and intrasexual territoriality in wolverine are patently reflected in the spatial and temporal configuration of noninvasively collected genetic samples. Our study confirms that wildlife monitoring programs can utilize the spatial information in noninvasive genetic sampling data to detect and quantify home ranges and social organization.

Recovery of large carnivores in Europe's modern human-dominated landscapes.

The conservation of large carnivores is a formidable challenge for biodiversity conservation. Using a data set on the past and current status of brown bears (Ursus arctos), Eurasian lynx (Lynx lynx), gray wolves (Canis lupus), and wolverines (Gulo gulo) in European countries, we show that roughly one-third of mainland Europe hosts at least one large carnivore species, with stable or increasing abundance in most cases in 21st-century records. The reasons for this overall conservation success include protective legislation, supportive public opinion, and a variety of practices making coexistence between large carnivores and people possible. The European situation reveals that large carnivores and people can share the same landscape.

The role of predation and food limitation on claims for compensation, reindeer demography and population dynamics.

A major challenge in biodiversity conservation is to facilitate viable populations of large apex predators in ecosystems where they were recently driven to ecological extinction due to resource conflict with humans.Monetary compensation for losses of livestock due to predation is currently a key instrument to encourage human-carnivore coexistence. However, a lack of quantitative estimates of livestock losses due to predation leads to disagreement over the practice of compensation payments. This disagreement sustains the human-carnivore conflict.The level of depredation on year-round, free-ranging, semi-domestic reindeer by large carnivores in Fennoscandia has been widely debated over several decades. In Norway, the reindeer herders claim that lynx and wolverine cause losses of tens of thousands of animals annually and cause negative population growth in herds. Conversely, previous research has suggested that monetary predator compensation can result in positive population growth in the husbandry, with cascading negative effects of high grazer densities on the biodiversity in tundra ecosystems.We utilized a long-term, large-scale data set to estimate the relative importance of lynx and wolverine predation and density-dependent and climatic food limitation on claims for losses, recruitment and population growth rates in Norwegian reindeer husbandry.Claims of losses increased with increasing predator densities, but with no detectable effect on population growth rates. Density-dependent and climatic effects on claims of losses, recruitment and population growth rates were much stronger than the effects of variation in lynx and wolverine densities.Synthesis and applications. Our analysis provides a quantitative basis for predator compensation and estimation of the costs of reintroducing lynx and wolverine in areas with free-ranging semi-domestic reindeer. We outline a potential path for conflict management which involves adaptive monitoring programmes, open access to data, herder involvement and development of management strategy evaluation (MSE) models to disentangle complex responses including multiple stakeholders and individual harvester decisions.

The risks of learning: confounding detection and demographic trend when using count-based indices for population monitoring.

Theory recognizes that a treatment of the detection process is required to avoid producing biased estimates of population rate of change. Still, one of three monitoring programmes on animal or plant populations is focused on simply counting individuals or other fixed visible structures, such as natal dens, nests, tree cavities. This type of monitoring design poses concerns about the possibility to respect the assumption of constant detection, as the information acquired in a given year about the spatial distribution of reproductive sites can provide a higher chance to detect the species in subsequent years. We developed an individual-based simulation model, which evaluates how the accumulation of knowledge about the spatial distribution of a population process can affect the accuracy of population growth rate estimates, when using simple count-based indices. Then, we assessed the relative importance of each parameter in affecting monitoring performance. We also present the case of wolverines (Gulo gulo) in southern Scandinavia as an example of a monitoring system with an intrinsic tendency to accumulate knowledge and increase detectability. When the occupation of a nest or den is temporally autocorrelated, the monitoring system is prone to increase its knowledge with time. This happens also when there is no intensification in monitoring effort and no change in the monitoring conditions. Such accumulated knowledge is likely to increase detection probability with time and can produce severe bias in the estimation of the rate and direction of population change over time. We recommend that a systematic sampling of the population process under study and an explicit treatment of the underlying detection process should be implemented whenever economic and logistical constraints permit, as failure to include detection probability in the estimation of population growth rate can lead to serious bias and severe consequences for management and conservation.

Implementation uncertainty when using recreational hunting to manage carnivores.

1. Wildlife managers often rely on resource users, such as recreational or commercial hunters, to achieve management goals. The use of hunters to control wildlife populations is especially common for predators and ungulates, but managers cannot assume that hunters will always fill annual quotas set by the authorities. It has been advocated that resource management models should account for uncertainty in how harvest rules are realized, requiring that this implementation uncertainty be estimated.2. We used a survival analysis framework and long-term harvest data from large carnivore management systems in three countries (Estonia, Latvia and Norway) involving four species (brown bear, grey wolf, Eurasian lynx and wolverine) to estimate the performance of hunters with respect to harvest goals set by managers.3. Variation in hunter quota-filling performance was substantial, ranging from 40% for wolverine in Norway to nearly 100% for lynx in Latvia. Seasonal and regional variation was also high within country-species pairs. We detected a positive relationship between the instantaneous potential to fill a quota slot and the relative availability of the target species for both wolverine and lynx in Norway.4. Survivor curves and hazards - with survival time measured as the time from the start of a season until a quota slot is filled - can indicate the extent to which managers can influence harvest through adjustments of season duration and quota limits.5.Synthesis and applications. We investigated seven systems where authorities use recreational hunting to manage large carnivore populations. The variation and magnitude of deviation from harvest goals was substantial, underlining the need to incorporate implementation uncertainty into resource management models and decisions-making. We illustrate how survival analysis can be used by managers to estimate the performance of resource users with respect to achieving harvest goals set by managers. The findings in this study come at an opportune time given the growing popularity of management strategy evaluation (MSE) models in fisheries and a push towards incorporating MSE into terrestrial harvest management.

Limited gene flow among brown bear populations in far Northern Europe? Genetic analysis of the east-west border population in the Pasvik Valley.

Noninvasively collected genetic data can be used to analyse large-scale connectivity patterns among populations of large predators without disturbing them, which may contribute to unravel the species' roles in natural ecosystems and their requirements for long-term survival. The demographic history of brown bears (Ursus arctos) in Northern Europe indicates several extinction and recolonization events, but little is known about present gene flow between populations of the east and west. We used 12 validated microsatellite markers to analyse 1580 hair and faecal samples collected during six consecutive years (2005-2010) in the Pasvik Valley at 70°N on the border of Norway, Finland and Russia. Our results showed an overall high correlation between the annual estimates of population size (N(c) ), density (D), effective size (N(e) ) and N(e) /N(c) ratio. Furthermore, we observed a genetic heterogeneity of ∼0.8 and high N(e) /N(c) ratios of ∼0.6, which suggests gene flow from the east. Thus, we expanded the population genetic study to include Karelia (Russia, Finland), Västerbotten (Sweden) and Troms (Norway) (477 individuals in total) and detected four distinct genetic clusters with low migration rates among the regions. More specifically, we found that differentiation was relatively low from the Pasvik Valley towards the south and east, whereas, in contrast, moderately high pairwise F(ST) values (0.91-0.12) were detected between the east and the west. Our results indicate ongoing limits to gene flow towards the west, and the existence of barriers to migration between eastern and western brown bear populations in Northern Europe.

Is hunting mortality additive or compensatory to natural mortality? Effects of experimental harvest on the survival and cause-specific mortality of willow ptarmigan.

1. The effects of harvest on the annual and seasonal survival of willow ptarmigan Lagopus lagopus L. were tested in a large-scale harvest experiment. Management units were randomly assigned to one of three experimental treatments: 0%, 15% or 30% harvest. Seasonal quotas were based on the experimental treatment and estimates of bird density before the hunting season. Survival rates and hazard functions for radio-marked ptarmigan were then estimated under the competing risks of harvest and natural mortality. 2. The partially compensatory mortality hypothesis was supported: annual survival of ptarmigan was 0·54 ± 0·08 SE under 0% harvest, 0·47 ± 0·06 under 15% harvest, and was reduced to 0·30 ± 0·05 under 30% harvest. Harvest mortality increased linearly from 0·08 ± 0·05, 0·27 ± 0·05 and 0·42 ± 0·06 from 0% to 30% harvest, whereas natural mortality was 0·38 ± 0·08, 0·25 ± 0·05 and 0·28 ± 0·06 under the same treatments. 3. Realized risk of harvest mortality was 0·08-0·12 points higher than our set harvest treatments of 0-30% because birds were exposed to risk if they moved out of protected areas. The superadditive hypothesis was supported because birds in the 30% harvest treatment had higher natural mortality during winter after the hunting season. 4. Natural mortality was mainly because of raptor predation, with two seasonal peaks in fall and spring. Natural and harvest mortality coincided during early autumn with little potential for compensation during winter months. Peak risk of harvest mortality was 5× higher than natural mortality. Low natural mortality during winter suggests that most late season harvest would be additive mortality. 5. Environmental correlates of natural mortality of ptarmigan included seasonal changes in snow cover, onset of juvenile dispersal, and periods of territorial activity. Natural mortality of ptarmigan was highest during autumn movements and nesting by gyrfalcons Falco rusticolus L. Mortality was low when gyrfalcons had departed for coastal wintering sites, and during summer when ptarmigan were attending nests and broods. 6. Our experimental results have important implications for harvest management of upland gamebirds. Seasonal quotas based on proportional harvest were effective and should be set at ≤ 15% of August populations for regional management plans. Under threshold harvest of a reproductive surplus, 15% harvest would be sustainable at productivity rates ≥ 2·5 young per pair. Impacts of winter harvest could be minimized by closing the hunting season in early November or by reducing late season quotas.

The cost of maturing early in a solitary carnivore.

Central to the theory of life history evolution is the existence of trade-offs between different traits, such as the trade-off between early maturity and an extended period of body growth. Based on analysis of the reproductive tracts of harvested Eurasian lynx (Lynx lynx) females in Norway, we find that females that mature early are generally heavier than those that postpone maturation. A higher proportion of 1.5-year-old females showed signs of ovulation in areas with high prey density, where they were also heavier. Further, we show that female Eurasian lynx that mature early have the same number of placental scars (an index of breeding investment and litter size) as older females, suggesting that they have a relatively high investment in their first litter. This induces a cost in terms of body weight development, as those females that had matured at the age of 1.5 years were substantially lighter by the age of 2.5 years than those that postponed breeding. This effect tended to be more pronounced in areas with low prey density. We discuss to what extent this might affect their future fitness prospects, and suggest that such costs of maturing early in terms of body weight development might be high in terrestrial large carnivores due to a prolonged period of postnatal care.

Sustainably harvesting a large carnivore? Development of Eurasian lynx populations in Norway during 160 years of shifting policy.

The management of large carnivores in multi-use landscapes is always controversial, and managers need to balance a wide range of competing interests. Hunter harvest is often used to limit population size and distribution but is proving to be both controversial and technically challenging. Eurasian lynx (Lynx lynx) are currently managed as a game species in Norway. We describe an adaptive management approach where quota setting is based on an annual census and chart the population development through the period 1996-2008, as management has become significantly more sophisticated and better informed by the increased availability of scientific data. During this period the population has been through a period of high quotas and population decline caused by fragmented management authority and over optimistic estimates of lynx reproduction, followed by a period of recovery due to quota reductions. The modern management regime is placed in the context of shifting policy during the last 160 years, during which management goals have moved from extermination stimulated by bounties, through a short phase of protection, and now to quota-regulated harvest. Much management authority has also been delegated from central to local levels. We conclude that adaptive management has the potential to keep the population within some bounded limits, although there will inevitably be fluctuation.

Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves.

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.