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.
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 L 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 5 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 ...
Summary1. The importance of individuals to the dynamics of populations may depend on reproductive status, especially for species with complex social structure. Loss of reproductive individuals in socially complex species could disproportionately affect population dynamics by destabilizing social structure and reducing population growth. Alternatively, compensatory mechanisms such as rapid replacement of breeders may result in little disruption. The impact of breeder loss on the population dynamics of social species remains poorly understood. 2. We evaluated the effect of breeder loss on social stability, recruitment and population growth of grey wolves (Canis lupus) in Denali National Park and Preserve, Alaska using a 26-year dataset of 387 radiocollared wolves. Harvest of breeding wolves is a highly contentious conservation and management issue worldwide, with unknown population-level consequences. 3. Breeder loss preceded 77% of cases (n = 53) of pack dissolution from 1986 to 2012. Packs were more likely to dissolve if a female or both breeders were lost and pack size was small. Harvest of breeders increased the probability of pack dissolution, likely because the timing of harvest coincided with the breeding season of wolves. Rates of denning and successful recruitment were uniformly high for packs that did not experience breeder loss; however, packs that lost breeders exhibited lower denning and recruitment rates. Breeder mortality and pack dissolution had no significant effects on immediate or longer term population dynamics. 4. Our results indicate the importance of breeding individuals is context dependent. The impact of breeder loss on social group persistence, reproduction and population growth may be greatest when average group sizes are small and mortality occurs during the breeding season. This study highlights the importance of reproductive individuals in maintaining group cohesion in social species, but at the population level socially complex species may be resilient to disruption and harvest through strong compensatory mechanisms.
Gray wolf (Canis lupus) populations have persisted and expanded in northwest Montana since 1986, while reintroduction efforts in Idaho and Yellowstone have further bolstered the regional population. However, rigorous analysis of either the availability of wolf habitat in the entire region, or the specific habitat requirements of local wolves, has yet to be conducted. We examined wolf‐habitat relationships in the northern Rocky Mountains of the U.S. by relating landscape/habitat features found within wolf pack home ranges (n = 56) to those found in adjacent non‐occupied areas (n = 56). Logistic regression revealed that increased forest cover, lower human population density, higher elk density, and lower sheep density were the primary factors related to wolf occupation. Similar factors promoted wolf pack persistence. Further, our analysis indicated that relatively large tracts of suitable habitat remain unoccupied in the Rocky Mountains, suggesting that wolf populations likely will continue to increase in the region. Analysis of the habitat linkage between the 3 main wolf recovery areas indicates that populations in central Idaho and northwest Montana have higher connectivity than either of the 2 recovery areas to the Greater Yellowstone recovery area. Thus, for the northern Rocky Mountains to function as a metapopulation for wolves, it will be necessary that dispersal corridors to the Yellowstone ecosystem be established and conserved.
DNA 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) shortrange 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.
Successful nonlethal management of livestock predation is important for conserving rare or endangered carnivores. In the northwestern United States, wolves (Canis lupus) have been translocated away from livestock to mitigate conflicts while promoting wolf restoration. We assessed predation on livestock, pack establishment, survival, and homing behavior of 88 translocated wolves with radiotelemetry to determine the effectiveness of translocation in our region and consider how it may be improved. More than one-quarter of translocated wolves preyed on livestock after release. Most translocated wolves (67%) never established or joined a pack, although eight new packs resulted from translocations. Translocated wolves had lower annual survival (0.60) than other radio-collared wolves (0.73), with government removal the primary source of mortality. In northwestern Montana, where most wolves have settled in human-populated areas with livestock, survival of translocated wolves was lowest (0.41) and more wolves proportionally failed to establish packs (83%) after release. Annual survival of translocated wolves was highest in central Idaho (0.71) and more wolves proportionally established packs (44%) there than in the other two recovery areas. Translocated wolves showed a strong homing tendency; most of those that failed to home still showed directional movement toward capture sites. Wolves that successfully returned to capture sites were more likely to be adults, hard (immediately) rather than soft (temporarily held in enclosure) released, and translocated shorter distances than other wolves that did not return home. Success of translocations varied and was most affected by the area in which wolves were released. We suggest managers translocating wolves or other large carnivores consider soft releasing individuals (in family groups, if social) when feasible because this may decrease homing behavior and increase release-site fidelity.
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