John Odden

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. Many populations of brown bears, lynx, grey wolves, and wolverines persist successfully outside protected areas in Europe. Success for Europes large carnivores? Despite pessimistic forecasts, Europes large carnivores are making a comeback. Chapron et al. report that sustainable populations of brown bear, Eurasian lynx, gray wolf, and wolverine persist in one-third of mainland Europe. Moreover, many individuals and populations are surviving and increasing outside protected areas set aside for wildlife conservation. Coexistence alongside humans has become possible, argue the authors, because of improved public opinion and protective legislation. Science, this issue p. 1517

Translocation of carnivores as a method for managing problem animals: a review

Translocation of individual carnivores has been a standard management tool for decades in North America and southern Africa in response to livestock depredation and other conflict behaviours. As carnivore populations across Europe begin to increase it is expected that management problems will also increase. Before translocation becomes established as a management tool in Europe its success needs to be reviewed. In general, there has been very little follow-up of translocated animals. Almost no data exist on the subsequent levels of damage after translocation. Large carnivores have shown a consistent ability to return to the site of capture over distances of up to 400 km. Even those individuals that do not succeed in returning home roam over very large distances, best measured in units of hundreds of kilometres. Very few individuals remain at the release sites. Survival of translocated animals has occasionally been shown to be poor, often as a result of the large movements. In general, there needs to be a large area (hundreds or thousands of square kilometres) without conflict potential where the individuals can be released for the strategy to work. When such areas are not available, management efforts should concentrate on reducing conflict potential, or, where this is not practical, lethal control.

Risk of capture-related mortality in large free-ranging mammals: experiences from Scandinavia

Abstract Chemical capture and anaesthesia of free-ranging mammals will always involve some risk of mortality even in healthy animals. Deaths may be directly or indirectly attributable to the anaesthetic event itself (e.g. drug overdose, drowning during induction and dart trauma) or may be caused by secondary effects from the capture (e.g. stress, myopathy, trauma or instrumentation with radio-transmitters). In long-term research projects on five major wildlife species in Scandinavia, the capture-related mortality rates (number of captures) were: moose Alces alces 0.7% (N = 2,816), brown bears Ursus arctos 0.9% (N = 1,079), wolverines Gulo gulo 2.8% (N = 461), Eurasian lynx Lynx lynx 3.9% (N = 380), and gray wolves Canis lupus 3.4% (N = 89). We suggest that wildlife professionals should strive for a zero mortality rate but adopt the standard that a mortality rate of > 2% probably should not be accepted in any large mammalian species. This can be achieved by: 1) using an experienced professional capture team, 2) developing and following a capture protocol specific to each species, and 3) requiring that a mortality assessment be undertaken after any capture-related death. This assessment should re-evaluate the capture protocol, including how changes in anaesthetics and methodological approaches could have prevented the mortality.

Biomedical Protocols for Free-ranging Brown Bears, Gray Wolves, Wolverines and Lynx Editors

PREFACE Compilation of this document was initiated by the Norwegian Directorate for Nature Management in order to establish recommended protocols for capture, chemical immobilization, anesthesia and radiotagging of free-ranging brown bears (Ursus arctos), gray wolves (Canis lupus), wolverines (Gulo gulo) and Eurasian lynx (Lynx lynx). In addition, procedures to ensure proper sampling of biological materials for management, research and banking purposes have been included. The current protocols are based on nearly 3,000 captures of free-ranging brown bears, wolves, wolverines and lynx carried out from 1984 through 2012 in Scandinavia. Some of the results have been published as peer reviewed papers, conference presentations, theses, and reports. However, a large amount of data are still on file and will be published in the future. In addition, comprehensive reviews of the global literature on brown bears, wolves, wolverines and lynx have been carried out in order to include pertinent information from other sources. These protocols have been approved by all ongoing research projects on brown bears, wolves, wolverines and lynx in Scandinavia. We thank the contributors for their cooperative efforts. We also thank the Norwegian Directorate for Nature Management for their support.

Diet of Eurasian lynx, Lynx lynx, in the boreal forest of southeastern Norway: the relative importance of livestock and hares at low roe deer density

The year-round food habits of lynx were studied using radio-telemetry and snow-tracking in the boreal forest of southeastern Norway. The main objectives of the study were to clarify the importance of domestic sheep and small prey species in the diet of lynx in an area with a very low density of roe deer. During the period 1995–1999, we found 193 scats and 358 kills made by lynx. Our results indicate that roe deer were the most common prey species (contributing to 83 and 34% of the biomass consumed in winter and summer, respectively), although a wide range of other species were also found, including mountain hares, tetranoids, red foxes, domestic sheep, wild reindeer, and even moose. Most of the diet was obtained by predation, although we did document several cases of scavenging. Roe deer were more important in the diet in winter than in summer, perhaps because they were easier to locate in winter as they clustered around feeding sites. In summer, domestic sheep and small prey increased in importance. Despite the very low density of roe deer in this study area, lynx seemed to still specialise on them, although domestic sheep did constitute a significant amount to their diet, especially for males and yearlings. However, the contribution of sheep to summer diet was far from that expected if their relative density was considered.

Climate, season, and social status modulate the functional response of an efficient stalking predator: the Eurasian lynx

1. Predation plays a major role in shaping the structure and dynamics of ecological communities, and the functional response of a predator is of crucial importance to the dynamics of any predator-prey system by linking the trophic levels. For large mammals, there is a dearth of field studies documenting functional responses, and observations at low prey density are particularly scarce. Furthermore, there is a lack of understanding about how variables such as season, social status and climate modulate the functional response curves. 2. We analysed kill rate data collected over a 10-year period based on radio-marked lynx (Lynx lynx) mainly preying on roe deer (Capreolus capreolus) along a steep prey density gradient in south-eastern Norway. 3. The asymptotic kill rate was reached at a very low prey density for both solitary individuals and family groups (i.e. females with their dependent kittens), indicative of an efficient predator. This highlights the importance of understanding the interplay between predator and prey at low prey densities. 4. A purely prey-dependent functional response was a poor descriptor of the data, as the curve was strongly modulated by season and differences between lynx of different social status. In addition, there was a clear effect of abiotic climatic factors (indexed by the North Atlantic Oscillation) on observed kill rates in the more snow-rich portion of our study area. 5. Our analysis suggests that simple functional response curves might be poor descriptors of predator consumption rates in complex natural system, and that auxiliary factors are likely to induce complexity into any predator-prey systems that would not be captured by simple deterministic approaches.

Lynx depredation on domestic sheep in Norway

We studied depredation rates on free-ranging domestic sheep (Ovis aries) by Eurasian lynx (Lynx lynx) in 2 areas in Norway to test whether selected individuals or a demographic group within a lynx population kill a disproportionate share of livestock. During 6 grazing seasons from 1994 to 1999, we monitored 34 radiocollared lynx during 641 tracking periods (either all night or during a full 24-hr period). Sixty-three domestic sheep and 3 domestic goats were killed by radiocollared lynx during these periods. All of the radiocollared lynx had access to free-ranging domestic sheep within their annual home ranges. Male lynx killed sheep more frequently than females and were responsible for 12 out of 13 cases of surplus killing. After adjusting for number of nights monitored, sheep-killing rates among males differed slightly, whereas the differences between individual females were correlated with the number of sheep available. There were no indications of any special problem individuals, but rather a problem sex. Therefore, it seems to be intrinsic in their behavior that males are likely to kill domestic animals more frequently than females. The lack of specific problem individuals in this study might depend on the livestock-herding technique used in Norway, where sheep generally are free-ranging and unattended throughout a lynxs home range. Selective removal of depredating males requires that the gap in the social mosaic either remains unfilled or is filled by an individual that causes less damage to livestock. The effects of such removal on sheep depredation are likely to be only temporary since all adult and yearling male lynx appear to kill sheep.

Habitat differentiation within the large-carnivore community of Norway's multiple-use landscapes

The re-establishment of large carnivores in Norway has led to increased conflicts and the adoption of regional zoning for these predators. When planning the future distribution of large carnivores, it is important to consider details of their potential habitat tolerances and strength of inter-specific differentiation. We studied differentiation in habitat and kill sites within the large-carnivore community of south-eastern Norway. We compared habitat selection of the brown bear Ursus arctos L., Eurasian lynx Lynx lynx L., wolf Canis lupus L. and wolverine Gulo gulo L., based on radio-tracking data. Differences in kill site locations were explored using locations of documented predator-killed sheep Ovis aries L. We modelled each species’ selection for, and differentiation in, habitat and kill sites on a landscape scale using resource selection functions and multinomial logistic regression. Based on projected probability of occurrence maps, we estimated continuous patches of habitat within the study area. Although bears, lynx, wolves and wolverines had overlapping distributions, we found a clear differentiation for all four species in both habitat and kill sites. The presence of bears, wolves and lynx was generally associated with rugged, forested areas at lower elevations, whereas wolverines selected rugged terrain at higher elevations. Some degree of sympatry was possible in over 40% of the study area, although only 1·5% could hold all four large carnivores together. Synthesis and applications. A geographically differentiated management policy has been adopted in Norway, aimed at conserving viable populations of large carnivores while minimizing the potential for conflicts. Sympatry of all four carnivores will be most successful if regional zones are established of adequate size spanning an elevational gradient. High prey densities, low carnivore densities, low dietary overlap and scavenging opportunities have most probably led to reduced competitive exclusion. Although regional sympatry enhances the conservation of an intact guild of large carnivores, it may well increase conflict levels and resistance to carnivore conservation locally.

Estimating total lynx Lynx lynx population size from censuses of family groups

Counts of reproductive units, i.e. family groups, constitute the main monitoring index for lynx Lynx lynx populations in Scandinavia. However, for some purposes it is necessary to extrapolate from the number of family groups to obtain an estimate of total population size. Using data on survival and reproduction from radio-marked lynx from three Scandinavian study areas, we simulated the lynx population structure in February. The average proportions of family groups out of all independent individuals, i.e. adults and yearlings, in these simulations were 21% ± 2.1 (SD), 22% ± 3.6 and 27% ±3.1 for the data sets from northern Sweden (Sarek), southeastern Norway (Hedmark) and south-central Sweden (Bergslagen), respectively, and the overall mean for all three study areas was 23% ± 3.8. This translated into extrapolation factors of 6.14 ± 0.44,6.24 ± 0.73 and 5.48 ± 0.40 for the three study areas, respectively, leading to an overall mean for all three study areas of 5.95 ± 0.64. We conclude, that it is possible to extrapolate from the number of family groups to obtain an estimate of total lynx population size with a statistical measure of uncertainty.

Predicting the potential demographic impact of predators on their prey: a comparative analysis of two carnivore–ungulate systems in Scandinavia

1. Understanding the role of predation in shaping the dynamics of animal communities is a fundamental issue in ecological research. Nevertheless, the complex nature of predator–prey interactions often prevents researchers from modelling them explicitly. 2. By using periodic Leslie–Usher matrices and a simulation approach together with parameters obtained from long-term field projects, we reconstructed the underlying mechanisms of predator–prey demographic interactions and compared the dynamics of the roe deer–red fox–Eurasian lynx–human harvest system with those of the moose–brown bear–gray wolf–human harvest system in the boreal forest ecosystem of the southern Scandinavian Peninsula. 3. The functional relationship of both roe deer and moose λ to changes in predation rates from the four predators was remarkably different. Lynx had the strongest impact among the four predators, whereas predation rates by wolves, red foxes, or brown bears generated minor variations in prey population λ. Elasticity values of lynx, wolf, fox and bear predation rates were −0·157, −0·056, −0·031 and −0·006, respectively, but varied with both predator and prey densities. 4. Differences in predation impact were only partially related to differences in kill or predation rates, but were rather a result of different distribution of predation events among prey age classes. Therefore, the age composition of killed individuals emerged as the main underlying factor determining the overall per capita impact of predation. 5. Our results confirm the complex nature of predator–prey interactions in large terrestrial mammals, by showing that different carnivores preying on the same prey species can exert a dramatically different demographic impact, even in the same ecological context, as a direct consequence of their predation patterns. Similar applications of this analytical framework in other geographical and ecological contexts are needed, but a more general evaluation of the subject is also required, aimed to assess, on a broader systematic and ecological range, what specific traits of a carnivore are most related to its potential impact on prey species.