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Various monitoring methods have been developed for large carnivores, but not all are practical or sufficiently accurate for long-term monitoring over large spatial scales. From 2009 to 2010, we used a predictive habitat model to locate gray wolf rendezvous sites in 4 study areas in Idaho, USA and conducted noninvasive genetic sampling (NGS) of scat and hair found at the sites. We evaluated species and individual identification PCR success rates across the study areas, and estimated population size with a single-session population estimator using 2 different recapture-coding methods. We then compared NGS population estimates to estimates generated concurrently from telemetry data. We collected 1,937 scat and 166 hair samples and identified 193 unique individuals over 2 years. For fecal DNA samples, species identification success rates were consistently high (>92%) across areas. Individual identification success rates ranged from 78% to 80% in the drier study areas and dropped to 50% in the wettest study area. The degree of agreement between NGS- and telemetry-derived population estimates varied by recapture-coding method with considerable variability in 95% confidence intervals. Population estimates derived from NGS methods were most influenced by the average number of detections per individual. We demonstrate how changes in field effort and recapture-coding method can affect population estimates in a widely used single-session population estimation model. Our study highlights the need to further develop reliable population estimation tools for single-session NGS data, especially those with large differences in capture frequencies among individuals stemming from severe capture heterogeneity (i.e., overdispersion).

Traditional methods of monitoring gray wolves (Canis lupus) are expensive and invasive and require extensive efforts to capture individual animals. Noninvasive genetic sampling (NGS) is an alternative method that can provide data to answer management questions and complement already-existing methods. In a 2-year study, we tested this approach for Idaho gray wolves in areas of known high and low wolf density. To focus sampling efforts across a large study area and increase our chances of detecting reproductive packs, we visited 964 areas with landscape characteristics similar to known wolf rendezvous sites. We collected scat or hair samples from 20% of sites and identified 122 wolves, using 8–9 microsatellite loci. We used the minimum count of wolves to accurately detect known differences in wolf density. Maximum likelihood and Bayesian single-session population estimators performed similarly and accurately estimated the population size, compared with a radiotelemetry population estimate, in both years, and an average of 1.7 captures per individual were necessary for achieving accurate population estimates. Subsampling scenarios revealed that both scat and hair samples were important for achieving accurate population estimates, but visiting 75% and 50% of the sites still gave reasonable estimates and reduced costs. Our research provides managers with an efficient and accurate method for monitoring high-density and low-density wolf populations in remote areas.

Many animals, including gray wolves (Canis lupus), live in social groups. Genetic techniques can help reveal the structure and composition of social groups, providing valuable information about group and population dynamics. We evaluated the effectiveness of using noninvasive genetic sampling (NGS) of fecal and hair samples at wolf rendezvous sites combined with spatial and genetic assignment criteria for assigning individuals to packs, detecting dispersers and lone wolves, determining the number of packs in an area, and obtaining group metrics. We applied this approach in 4 study areas covering 13,182 km² in Idaho, USA while concurrently monitoring wolves using telemetry techniques. We assigned pack affiliation to 78-97% of individuals across study areas and identified 12 potential dispersers. We detected a successful gene flow event by reconstructing a breeding male's genotype and tracing it back to a pack of origin using genetic assignment techniques. Average pack size was consistent between our NGS- and telemetry-based counts (x̄ = 10 for both), and both methods detected similar age composition within groups (31% pups and 69% adults for NGS and 33% pups and 67% adults for telemetry). Our NGS approach has the advantage of providing pack metrics including sex ratio, inferred breeders, and intra-pack relatedness that telemetry and observational techniques alone cannot. This NGS field sampling strategy combined with our pack assignment method was successful and provides an approach for characterizing functional social groups in the absence of previously acquired NGS, telemetry, or other observational data that may not be available when sampling new areas. © 2016 The Wildlife Society.

Studying the ecology and behavior of pack animals often requires that most, or all, of the pack members are sampled. A unique opportunity to sample all gray wolf (Canis lupus) pack members arises during the summer months when reproductive packs localize in rendezvous sites. We collected 155–296 scat and hair samples from each of 5 wolf rendezvous sites in central Idaho to evaluate intrapack relationships and determine the efficacy of noninvasive genetic sampling (NGS) for estimating pack size and family relationships. We detected 65 wolves (5–20 wolves per pack) with NGS, and the pack counts from NGS were the same or higher for adults and the same or slightly lower for pups compared with the counts from observation and telemetry. The wolves in each pack were closely related to one another, and all packs included at least 2 years of offspring from the current breeding pair. Three of the packs had additional breeding adults present. In 1 pack pups were produced by a parent–offspring pair and a pair of their inbred full siblings, indicating multiple cases of inbreeding. This targeted NGS approach shows great promise for studying pack size and wolf social structure without the use of radiotelemetry or direct observations.

Connectivity is important for population persistence and can reduce the potential for inbreeding depression. Connectivity between populations can also facilitate disease transmission; respiratory diseases are one of the most important factors affecting populations of bighorn sheep (Ovis canadensis). The mechanisms of connectivity in populations of bighorn sheep likely have implications for spread of disease, but the behaviors leading to connectivity between bighorn sheep groups are not well understood. From 2007–2012, we radio-collared and monitored 56 bighorn sheep in the Salmon River canyon in central Idaho. We used cluster analysis to define social groups of bighorn sheep and then estimated connectivity between these groups using a multi-state mark-recapture model. Social groups of bighorn sheep were spatially segregated and linearly distributed along the Salmon River canyon. Monthly probabilities of movement between adjacent male and female groups ranged from 0.08 (±0.004 SE) to 0.76 (±0.068) for males and 0.05 (±0.132) to 0.24 (±0.034) for females. Movements of males were extensive and probabilities of movement were considerably higher during the rut. Probabilities of movement for females were typically smaller than those of males and did not change seasonally. Whereas adjacent groups of bighorn sheep along the Salmon River canyon were well connected, connectivity between groups north and south of the Salmon River was limited. The novel application of a multi-state model to a population of bighorn sheep allowed us to estimate the probability of movement between adjacent social groups and approximate the level of connectivity across the population. Our results suggest high movement rates of males during the rut are the most likely to result in transmission of pathogens among both male and female groups. Potential for disease spread among female groups was smaller but non-trivial. Land managers can plan grazing of domestic sheep for spring and summer months when males are relatively inactive. Removal or quarantine of social groups may reduce probability of disease transmission in populations of bighorn sheep consisting of linearly distributed social groups.

We used rendezvous site locations of wolf (Canis lupus) packs recorded during 1996–2006 to build a predictive model of gray wolf rendezvous site habitat in Idaho, USA. Variables in our best model included green leaf biomass (Normalized Difference Vegetation Index), surface roughness, and profile curvature, indicating that wolves consistently used wet meadow complexes for rendezvous sites. We then used this predictive model to stratify habitat and guide survey efforts designed to document wolf pack distribution and fecundity in 4 study areas in Idaho. We detected all 15 wolf packs (32 wolf pack-yr) and 20 out of 27 (74%) litters of pups by surveying <11% of the total study area. In addition, we were able to obtain detailed observations on wolf packs (e.g., hair and scat samples) once we located their rendezvous sites. Given an expected decrease in the ability of managers to maintain radiocollar contact with all of the wolf packs in the northern Rocky Mountains, rendezvous sites predicted by our model can be the starting point and foundation for targeted sampling and future wolf population monitoring surveys.

The behavioral patterns and large territories of large carnivores make them challenging to monitor. Occupancy modeling provides a framework for monitoring population dynamics and distribution of territorial carnivores. We combined data from hunter surveys, howling and sign surveys conducted at predicted wolf rendezvous sites, and locations of radiocollared wolves to model occupancy and estimate the number of gray wolf (Canis lupus) packs and individuals in Idaho during 2009 and 2010. We explicitly accounted for potential misidentification of occupied cells (i.e., false positives) using an extension of the multi-state occupancy framework. We found agreement between model predictions and distribution and estimates of number of wolf packs and individual wolves reported by Idaho Department of Fish and Game and Nez Perce Tribe from intensive radiotelemetry-based monitoring. Estimates of individual wolves from occupancy models that excluded data from radiocollared wolves were within an average of 12.0% (SD = 6.0) of existing statewide minimum counts. Models using only hunter survey data generally estimated the lowest abundance, whereas models using all data generally provided the highest estimates of abundance, although only marginally higher. Precision across approaches ranged from 14% to 28% of mean estimates and models that used all data streams generally provided the most precise estimates. We demonstrated that an occupancy model based on different survey methods can yield estimates of the number and distribution of wolf packs and individual wolf abundance with reasonable measures of precision. Assumptions of the approach including that average territory size is known, average pack size is known, and territories do not overlap, must be evaluated periodically using independent field data to ensure occupancy estimates remain reliable. Use of multiple survey methods helps to ensure that occupancy estimates are robust to weaknesses or changes in any 1 survey method. Occupancy modeling may be useful for standardizing estimates across large landscapes, even if survey methods differ across regions, allowing for inferences about broad-scale population dynamics of wolves.

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.

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.

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.