Håkan Sand

Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant.

The fragmentation of populations is an increasingly important problem in the conservation of endangered species. Under these conditions, rare migration events may have important effects for the rescue of small and inbred populations. However, the relevance of such migration events to genetically depauperate natural populations is not supported by empirical data. We show here that the genetic diversity of the severely bottlenecked and geographically isolated Scandinavian population of grey wolves (Canis lupus), founded by only two individuals, was recovered by the arrival of a single immigrant. Before the arrival of this immigrant, for several generations the population comprised only a single breeding pack, necessarily involving matings between close relatives and resulting in a subsequent decline in individual heterozygosity. With the arrival of just a single immigrant, there is evidence of increased heterozygosity, significant outbreeding (inbreeding avoidance), a rapid spread of new alleles and exponential population growth. Our results imply that even rare interpopulation migration can lead to the rescue and recovery of isolated and endangered natural populations.

Severe inbreeding depression in a wild wolf Canis lupus population

The difficulty of obtaining pedigrees for wild populations has hampered the possibility of demonstrating inbreeding depression in nature. In a small, naturally restored, wild population of grey wolves in Scandinavia, founded in 1983, we constructed a pedigree for 24 of the 28 breeding pairs established in the period 1983–2002. Ancestry for the breeding animals was determined through a combination of field data (snow tracking and radio telemetry) and DNA microsatellite analysis. The population was founded by only three individuals. The inbreeding coefficient F varied between 0.00 and 0.41 for wolves born during the study period. The number of surviving pups per litter during their first winter after birth was strongly correlated with inbreeding coefficients of pups (R2=0.39, p<0.001). This inbreeding depression was recalculated to match standard estimates of lethal equivalents (2B), corresponding to 6.04 (2.58–9.48, 95% CI) litter-size-reducing equivalents in this wolf population.

Life history patterns in female moose (Alces alces): the relationship between age, body size, fecundity and environmental conditions

I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (≥2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.

Home-Range Size in Relation to Age and Sex in Moose

During 1983–1989, 49 moose ( Alces alces ) were monitored in central Sweden to estimate annual and seasonal home ranges. Males had larger annual home ranges (25.9 km2 ± 3.3 SE ) than did females (13.7 km2 ± 2.2 SE, P 0.05), except females without young ( P < 0.05). Only in autumn did females with young have different ( P < 0.01) average home-range size compared with females without young; those with young were, on average, twice as large as those without young (5.8 km2 versus 2.2 km2, respectively). We suggest that variation in home-range size may have two main causes. First, body size, and thus nutritional demands, varies between sexes throughout the year; second, the strong relationship between age and home-range size among males is likely caused by social activities associated with rut, and, in particular with age-related dominance.

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.

Selection for Heterozygosity Gives Hope to a Wild Population of Inbred Wolves

Recent analyses have questioned the usefulness of heterozygosity estimates as measures of the inbreeding coefficient (f), a finding that may have dramatic consequences for the management of endangered populations. We confirm that f and heterozygosity is poorly correlated in a wild and highly inbred wolf population. Yet, our data show that for each level of f, it was the most heterozygous wolves that established themselves as breeders, a selection process that seems to have decelerated the loss of heterozygosity in the population despite a steady increase of f. The markers contributing to the positive relationship between heterozygosity and breeding success were found to be located on different chromosomes, but there was a substantial amount of linkage disequilibrium in the population, indicating that the markers are reflecting heterozygosity over relatively wide genomic regions. Following our results we recommend that management programs of endangered populations include estimates of both f and heterozygosity, as they may contribute with complementary information about population viability.

Effects of hunting group size, snow depth and age on the success of wolves hunting moose

To study factors important to the success of wolves, Canis lupus, hunting moose, Alces alces, we analysed data from more than 4000 km of snow tracking of wolves during 1998e2003 in Scandinavia. We used two methods to estimate hunting success for 17 wolf territories from 185 observations of wolf attacks on moose. On average, hunting success was estimated at 45 and 64% for the two methods, respectively. We used a smaller data set (N ¼ 142) to examine the effect of age of breeding wolves, hunting group size, snow depth and moose density on hunting success. Multiple logistic regression showed that age of breeding males was the only variable significantly related to hunting success, with maximum hunting success at 4.5e5.5þ years of age. We also studied prey selection of radiocollared adult wolves over successive winters in two wolf packs that lost one of the breeding wolves. Whereas the surviving adult female switched to prey on roe deer, Capreolus capreolus, the surviving adult male continued mainly to select moose. Our results suggest that the positive effect of male age on hunting success reflects both increased experience of attacking prey and possibly the greater size of adult male wolves (25e30%) compared to adult female wolves.

Cross-continental differences in patterns of predation: will naive moose in Scandinavia ever learn?

Predation has been recognized as a major selective force in the evolution of behavioural characteristics of mammals. As a consequence of local predator extinction, prey may lose knowledge about natural predators but usually express behavioural adjustments after return of predators. Human harvest may replace natural predation but prey selection may differ from that of natural predators leading to a change in the behavioural response of prey. We show that hunting success (HS) of re-colonizing wolves (Canis lupus) on moose (Alces alces) in Scandinavia was higher than reported in North America, where moose have been continuously exposed to wolves and grizzly bears. We found no evidence that moose expressed behavioural adjustments that lowered the HS of wolves in territories that had been occupied by wolves for up to 21 years. Moose behaviour towards wolves and humans typically differs in Scandinavia compared to North America. We explain the differences found to be caused by variation in predation pressure by large carnivores and the rate, and mode, of human harvest during the twentieth century.

Building a mechanistic understanding of predation with GPS-based movement data

Quantifying kill rates and sources of variation in kill rates remains an important challenge in linking predators to their prey. We address current approaches to using global positioning system (GPS)-based movement data for quantifying key predation components of large carnivores. We review approaches to identify kill sites from GPS movement data as a means to estimate kill rates and address advantages of using GPS-based data over past approaches. Despite considerable progress, modelling the probability that a cluster of GPS points is a kill site is no substitute for field visits, but can guide our field efforts. Once kill sites are identified, time spent at a kill site (handling time) and time between kills (killing time) can be determined. We show how statistical models can be used to investigate the influence of factors such as animal characteristics (e.g. age, sex, group size) and landscape features on either handling time or killing efficiency. If we know the prey densities along paths to a kill, we can quantify the ‘attack success’ parameter in functional response models directly. Problems remain in incorporating the behavioural complexity derived from GPS movement paths into functional response models, particularly in multi-prey systems, but we believe that exploring the details of GPS movement data has put us on the right path.

Body Mass Dynamics of Moose Calves in Relation to Winter Severity

We measured winter and spring loss of body mass among moose (Alces alces) calves in a population in southcentral Sweden (Grimso). The sample consisted of 522 calves of both sexes killed during October-May from 1973 to 1988. Males were consistently heavier than females (P<0.05). Body masses of moose calves were significantly associated with both winter severity (P<0.05) and the date of kill (P<0.001). Mean loss of body mass was highest in hard winters (13% from Nov to Feb-Mar). Conversely, average net gain in body mass from autumn (Nov) to spring (Apr-May) (20%) was highest in mild winters. Snow depth, but not temperature and precipitation, was significantly associated with loss of body masses during winter (P<0.02). During winter, small calves lost not only a larger proportion of their body mass, but also more kilograms than large calves