Thomas J. Meier

A study of the genetic relationships within and among wolf packs using DNA fingerprinting and mitochondrial DNA

SummaryDNA fingerprinting and mitochondrial DNA analyses have not been used in combination to study relatedness in natural populations. We present an approach that involves defining the mean fingerprint similarities among individuals thought to be unrelated because they have different mtDNA genotypes. Two classes of related individuals are identified by their distance in standard errors above this mean value. The number of standard errors is determined by analysis of the association between fingerprint similarity and relatedness in a population with a known genealogy. We apply this approach to gray wolf packs from Minnesota, Alaska, and the Northwest Territories. Our results show that: (1) wolf packs consist primarily of individuals that are closely related genetically, but some packs contain unrelated, non-reproducing individuals; (2) dispersal among packs within the same area is common; and (3) short-range dispersal appears more common for female than male wolves. The first two of these genetically-based observations are consistent with behavioral data on pack structure and dispersal in wolves, while the apparent sex bias in dispersal was not expected.

Survival of Colonizing Wolves in the Northern Rocky Mountains of the United States, 1982-2004

Abstract After roughly a 60-year absence, wolves (Canis lupus) immigrated (1979) and were reintroduced (1995–1996) into the northern Rocky Mountains (NRM), USA, where wolves are protected under the Endangered Species Act. The wolf recovery goal is to restore an equitably distributed metapopulation of ≥30 breeding pairs and 300 wolves in Montana, Idaho, and Wyoming, while minimizing damage to livestock; ultimately, the objective is to establish state-managed conservation programs for wolf populations in NRM. Previously, wolves were eradicated from the NRM because of excessive human killing. We used Andersen–Gill hazard models to assess biological, habitat, and anthropogenic factors contributing to current wolf mortality risk and whether federal protection was adequate to provide acceptably low hazards. We radiocollared 711 wolves in Idaho, Montana, and Wyoming (e.g., NRM region of the United States) from 1982 to 2004 and recorded 363 mortalities. Overall, annual survival rate of wolves in the recovery areas was 0.750 (95% CI  =  0.728–0.772), which is generally considered adequate for wolf population sustainability and thereby allowed the NRM wolf population to increase. Contrary to our prediction, wolf mortality risk was higher in the northwest Montana (NWMT) recovery area, likely due to less abundant public land being secure wolf habitat compared to other recovery areas. In contrast, lower hazards in the Greater Yellowstone Area (GYA) and central Idaho (CID) likely were due to larger core areas that offered stronger wolf protection. We also found that wolves collared for damage management purposes (targeted sample) had substantially lower survival than those collared for monitoring purposes (representative sample) because most mortality was due to human factors (e.g., illegal take, control). This difference in survival underscores the importance of human-caused mortality in this recovering NRM population. Other factors contributing to increased mortality risk were pup and yearling age class, or dispersing status, which was related to younger age cohorts. When we included habitat variables in our analysis, we found that wolves having abundant agricultural and private land as well as livestock in their territory had higher mortality risk. Wolf survival was higher in areas with increased wolf density, implying that secure core habitat, particularly in GYA and CID, is important for wolf protection. We failed to detect changes in wolf hazards according to either gender or season. Maintaining wolves in NWMT will require greater attention to human harvest, conflict resolution, and illegal mortality than in either CID or GYA; however, if human access increases in the future in either of the latter 2 areas hazards to wolves also may increase. Indeed, because overall suitable habitat is more fragmented and the NRM has higher human access than many places where wolves roam freely and are subject to harvest (e.g., Canada and AK), monitoring of wolf vital rates, along with concomitant conservation and management strategies directed at wolves, their habitat, and humans, will be important for ensuring long-term viability of wolves in the region.

The Effects of Breeder Loss on Wolves

Abstract Managers of recovering wolf (Canis lupus) populations require knowledge regarding the potential impacts caused by the loss of territorial, breeding wolves when devising plans that aim to balance population goals with human concerns. Although ecologists have studied wolves extensively, we lack an understanding of this phenomenon as published records are sparse. Therefore, we pooled data (n = 134 cases) on 148 territorial breeding wolves (75 M and 73 F) from our research and published accounts to assess the impacts of breeder loss on wolf pup survival, reproduction, and territorial social groups. In 58 of 71 cases (84%), ≥1 pup survived, and the number or sex of remaining breeders (including multiple breeders) did not influence pup survival. Pups survived more frequently in groups of ≥6 wolves (90%) compared with smaller groups (68%). Auxiliary nonbreeders benefited pup survival, with pups surviving in 92% of cases where auxiliaries were present and 64% where they were absent. Logistic regression analysis indicated that the number of adult-sized wolves remaining after breeder loss, along with pup age, had the greatest influence on pup survival. Territorial wolves reproduced the following season in 47% of cases, and a greater proportion reproduced where one breeder had to be replaced (56%) versus cases where both breeders had to be replaced (9%). Group size was greater for wolves that reproduced the following season compared with those that did not reproduce. Large recolonizing (>75 wolves) and saturated wolf populations had similar times to breeder replacement and next reproduction, which was about half that for small recolonizing (≤75 wolves) populations. We found inverse relationships between recolonizing population size and time to breeder replacement (r = −0.37) and time to next reproduction (r = −0.36). Time to breeder replacement correlated strongly with time to next reproduction (r = 0.97). Wolf social groups dissolved and abandoned their territories subsequent to breeder loss in 38% of cases. Where groups dissolved, wolves reestablished territories in 53% of cases, and neighboring wolves usurped territories in an additional 21% of cases. Fewer groups dissolved where breeders remained (26%) versus cases where breeders were absent (85%). Group size after breeder loss was smaller where groups dissolved versus cases where groups did not dissolve. To minimize negative impacts, we recommend that managers of recolonizing wolf populations limit lethal control to solitary individuals or territorial pairs where possible, because selective removal of pack members can be difficult. When reproductive packs are to be managed, we recommend that managers only remove wolves from reproductive packs when pups are ≥6 months old and packs contain ≥6 members (including ≥3 ad-sized wolves). Ideally, such packs should be close to neighboring packs and occur within larger (≥75 wolves) recolonizing populations.

People and Wildlife: Managing wolf–human conflict in the northwestern United States

INTRODUCTION The grey wolf ( Canis lupus ) is the most widely distributed large carnivore in the northern hemisphere (Nowak 1995) and has a reputation for killing livestock and competing with human hunters for wild ungulates (Young 1944; Fritts et al . 2003). Wolves rarely threaten human safety, but many people still fear them. In the western USA, widespread extirpation of ungulates by colonizing settlers, wolf depredation on livestock and negative public attitudes towards wolves resulted in extirpation of wolf populations by 1930 (Mech 1970; McIntyre 1995). By 1970, mule deer ( Odocoileus hemionus ), white-tailed deer ( O. virginianus ), elk ( Cervus elaphus ), moose ( Alces alces ) and bighorn sheep ( Ovis canadensis ) populations had been restored throughout the western USA while bison ( Bison bison ) were recovered only in Yellowstone National Park. However, grey wolves were still persecuted. In 1974, grey wolves were protected and managed by the US Fish and Wildlife Service under the federal Endangered Species Act of 1973. In 1986, the first recorded den in the western USA in over 50 years was established in Glacier National Park by wolves that naturally dispersed from Canada (Ream et al . 1989). Restoration of wolves in that region emphasized legal protection and building local public tolerance. Wolves from Canada were reintroduced to central Idaho and Yellowstone National Park in 1995 and 1996 to accelerate restoration (Bangs and Fritts 1996; Fritts et al . 1997). The Northern Rocky Mountains wolf population grew from 10 wolves in 1987 to 663 wolves by 2003 (US Fish and Wildlife Service et al . 2003) (Fig. 21.1, Table 21.1).

Kin encounter rate and inbreeding avoidance in canids

Mating with close kin can lead to inbreeding depression through the expression of recessive deleterious alleles and loss of heterozygosity. Mate selection may be affected by kin encounter rate, and inbreeding avoidance may not be uniform but associated with age and social system. Specifically, selection for kin recognition and inbreeding avoidance may be more developed in species that live in family groups or breed cooperatively. To test this hypothesis, we compared kin encounter rate and the proportion of related breeding pairs in noninbred and highly inbred canid populations. The chance of randomly encountering a full sib ranged between 1–8% and 20–22% in noninbred and inbred canid populations, respectively. We show that regardless of encounter rate, outside natal groups mates were selected independent of relatedness. Within natal groups, there was a significant avoidance of mating with a relative. Lack of discrimination against mating with close relatives outside packs suggests that the rate of inbreeding in canids is related to the proximity of close relatives, which could explain the high degree of inbreeding depression observed in some populations. The idea that kin encounter rate and social organization can explain the lack of inbreeding avoidance in some species is intriguing and may have implications for the management of populations at risk.

Estimation of Successful Breeding Pairs for Wolves in the Northern Rocky Mountains, USA

Abstract Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ≥2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.

Temporal validation of an estimator for successful breeding pairs of wolves Canis lupus in the U.S. northern Rocky Mountains

Abstract Model-based predictors derived from historical data are rarely evaluated before they are used to draw inferences. We performed a temporal validation, (i.e. assessed the performance of a predictive model using data collected from the same population after the model was developed) of a statistical predictor for the number of successful breeding pairs of wolves Canis lupus in the northern Rocky Mountains (NRM). We predicted the number of successful breeding pairs, β, in Idaho, Montana and Wyoming based on the distribution of pack sizes observed through monitoring in 2006 and 2007 (β̂), and compared these estimates to the minimum number of successful breeding pairs, βMIN, observed through intensive monitoring. βMIN was consistently included within the 95% confidence intervals of β̂ for all states in both years (except for Idaho in 2007), generally following the pattern β̂L (lower 95% prediction interval for β̂) < β̂MIN < β̂. This evaluation of β̂ estimates for 2006 and 2007 suggest it will be a robust model-based method for predicting successful breeding pairs of NRM wolves in the future, provided influences other than those modeled in β̂ (e.g. disease outbreak, severe winter) do not have a strong effect on wolf populations. Managers can use β̂ models with added confidence as part of their post-delisting monitoring of wolves in NRM.