Genetic Structure and Migration in Native and Reintroduced Rocky Mountain Wolf Populations

Gray wolf (Canis lupus) recovery in the Rocky Mountains of the U.S. is proceeding by both natural recolonization and managed reintroduction. We used DNA microsatellite analysis of wolves transplanted from Canada to two reintroduction sites in the U.S. to study population structure in native and reintroduced wolf populations. Gene flow due to migration between regions in Canada is substantial, and all three recovery populations in the U.S. had high genetic variation. The reintroduced founders were moderately genetically divergent from the naturally colonizing U.S. population. These findings corroborate that the reintroduction more than meets generally accepted genetic guidelines. Maintaining this variation, however, will depend on ample reproduction in the first few generations. In the long term genetic variation will best be retained if migration occurs among the recolonizing and the two transplanted populations. Evidence from field observation and genetic studies shows extensive dispersal by wolves, and we conclude that exchange among these groups due to natural dispersal is likely if public tolerance and legal protection are adequate outside lands designated for wolf recovery.     

Use of linkage mapping and centrality analysis across habitat gradients to conserve connectivity of gray wolf populations in western North America

Centrality metrics evaluate paths between all possible pairwise combinations of sites on a landscape to rank the contribution of each site to facilitating ecological flows across the network of sites. Computational advances now allow application of centrality metrics to landscapes represented as continuous gradients of habitat quality. This avoids the binary classification of landscapes into patch and matrix required by patch-based graph analyses of connectivity. It also avoids the focus on delineating paths between individual pairs of core areas characteristic of most corridor- or linkage-mapping methods of connectivity analysis. Conservation of regional habitat connectivity has the potential to facilitate recovery of the gray wolf (Canis lupus), a species currently recolonizing portions of its historic range in the western United States. We applied 3 contrasting linkage-mapping methods (shortest path, current flow, and minimum-cost-maximum-flow) to spatial data representing wolf habitat to analyze connectivity between wolf populations in central Idaho and Yellowstone National Park (Wyoming). We then applied 3 analogous betweenness centrality metrics to analyze connectivity of wolf habitat throughout the northwestern United States and southwestern Canada to determine where it might be possible to facilitate range expansion and interpopulation dispersal. We developed software to facilitate application of centrality metrics. Shortest-path betweenness centrality identified a minimal network of linkages analogous to those identified by least-cost-path corridor mapping. Current flow and minimum-cost-maximum-flow betweenness centrality identified diffuse networks that included alternative linkages, which will allow greater flexibility in planning. Minimum-cost-maximum-flow betweenness centrality, by integrating both land cost and habitat capacity, allows connectivity to be considered within planning processes that seek to maximize species protection at minimum cost. Centrality analysis is relevant to conservation and landscape genetics at a range of spatial extents, but it may be most broadly applicable within single- and multispecies planning efforts to conserve regional habitat connectivity.     

Mitochondrial DNA Variability in Italian and East European Wolves: Detecting the Consequences of Small Population Size and Hybridization

Abstract: The Italian wolf ( Canis lupus ) population has declined continuously over the last few centuries and become isolated as a result of the extermination of other populations in central Europe and the Alps during the nineteenth century. In the 1970s, approximately 100 wolves survived in 10 isolated areas in the central and southern Italian Apennines. Loss of genetic variability, as suggested by preliminary studies of mitochondrial DNA (mtDNA) sequences, hybridization with feral dogs, and the illegal release of captive, non-native wolves are considered potential threats to the viability of the Italian wolf population. We sequenced 546 base pairs of the mtDNA control region in a comprehensive set of Italian wolves and compared them to those of dogs and other wolf populations from Europe and the Near East. Our data confirm the absence of mtDNA variability in Italian wolves: all 101 individuals sampled across their distribution in Italy had the same, unique haplotype, whereas seven haplotypes were found in only 26 wolves from an outbred population in Bulgaria. Most haplotypes were specific either to wolves or dogs, but some east European wolves shared haplotypes with dogs, indicative of hybridization. In contrast, neither hybridization with dogs nor introgression of non-native wolves was detected in the Italian population. These findings exclude the introgression of dog genes via matings between male wolves and female dogs, the most likely direction of hybridization. The observed mtDNA monomorphism is the possible outcome of random drift in the declining and isolated Italian wolf population, which probably existed at low effective population size during the last 100–150 years. Low effective population size and the continued loss of genetic variability might be a major threat to the long-term viability of Italian wolves. A controlled demographic increase, leading to recolonization of the historical wolf range in Italy, should be enforced.     

Molecular Genetics of Pre-1940 Red Wolves

Conservation of the endangered red wolf (Canis rufus) has become a controversial issue because its genetic and morphological composition has been altered by hybridization with coyotes (C. latrans) and possibly gray wolves (C. lupus) making its evolutionary origins difficult to ascertain. The evolutionary hypothesis based on morphological data is that the red wolf had an Early Pleistocene origin and was the predecessor of both modern coyotes and gray wolves. After 1940 red wolves hybridized with coyotes as the species vanished from the wild. In contrast to this ancient origin-recent introgression hypothesis, molecular data are more consistent with an origin through hybridization between gray wolves and coyotes. Interspecific hybridization may have occurred repeatedly over time prior to European settlement in the southcentral United States or may have been induced recently by anthropogenic changes. We review recent molecular evidence and present new results from the analysis of mitochondrial and nuclear DNA markers in pre-1940 populations of red wolves. Our results are inconsistent with an ancient origin of the red wolf and support the hybridization model. We discuss possible hybridization scenarios and reasons for the red wolf reintroduction program to be concerned with the effects of genetic introgression from coyotes.     

Mitochondrial DNA Variability of the Gray Wolf: Genetic Consequences of Population Decline and Habitat Fragmentation

The gray wolf is a large, highly mobile predator whose original geographic range included most of the Northern Hemisphere. High rates of genetic exchange probably characterized even distantly-separated populations in the past, but recent population declines and habitat fragmentation have isolated previously contiguous populations, especially in the Old World. We examine mitochondrial DNA (mtDNA) variability among twenty-six populations of wolves from throughout their geographic range. We find eighteen mtDNA genotypes in gray wolves, seven of these are derived from hybridization with coyotes, four are confined to the New World, six are confined to the Old World and one is shared by both areas. Genetic differentiation among wolf populations is significant but small in magnitude. In the Old World, most localities have a single unique genotype, whereas in the New World several genotypes occur at most localities and three of the five genotypes are nearly ubiquitous. The pattern of genetic differentiation in the gray wolf contrasts with that of another large, highly vagile canid, the coyote, in which genetic differentiation among populations is not significant even among widely separated localities. We suggest that the difference between these two species reflects the rapid, recent increase in coyote numbers and expansion of their geographic range, and the coincident decline in gray wolf populations. Apparent genetic differences among extant wolf populations may be a recent phenomenon reflecting population declines and habitat fragmentation rather than a long history of genetic isolation.     

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

Summary 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) 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.USA Offprint requests to: N. Lehman at the present address     

Longitudinal Analysis of Attitudes Toward Wolves

Understanding individual attitudes and how these predict overt opposition to predator conservation or direct, covert action against predators will help to recover and maintain them. Studies of attitudes toward wild animals rely primarily on samples of individuals at a single time point. We examined longitudinal change in individuals’ attitudes toward gray wolves (Canis lupus). In the contiguous United States, amidst persistent controversy and opposition, abundances of gray wolves are at their highest in 60 years. We used mailed surveys to sample 1892 residents of Wisconsin in 2001 or 2004 and then resampled 656 of these individuals who resided in wolf range in 2009. Our study spanned a period of policy shifts and increasing wolf abundance. Over time, the 656 respondents increased agreement with statements reflecting fear of wolves, the belief that wolves compete with hunters for deer (Odocoileus virginianus), and inclination to poach a wolf. Endorsement of lethal control of wolves by the state and public hunting of wolves also increased. Neither the time span over which respondents reported exposure to wolves locally nor self‐reported losses of domestic animals to wolves correlated with changes in attitude. We predict future increases in legal and illegal killing of wolves that may reduce their abundance in Wisconsin unless interventions are implemented to improve attitudes and behavior toward wolves. To assess whether interventions change attitudes, longitudinal studies like ours are needed. Análisis Longitudinal de las Actitudes Hacia Lobos     

Willow on Yellowstone's northern range: evidence for a trophic cascade?

Reintroduction of wolves (Canis lupus) to Yellowstone National Park in 1995-1996 has been argued to promote a trophic cascade by altering elk (Cervus elaphus) density, habitat-selection patterns, and behavior that, in turn, could lead to changes within the plant communities used by elk. We sampled two species of willow (Salix boothii and S. geyeriana) on the northern winter range to determine whether (1) there was quantitative evidence of increased willow growth following wolf reintroduction, (2) browsing by elk affected willow growth, and (3) any increase in growth observed was greater than that expected by climatic and hydrological factors alone, thereby indicating a trophic cascade caused by wolves. Using stem sectioning techniques to quantify historical growth patterns we found an approximately twofold increase in stem growth-ring area following wolf reintroduction for both species of willow. This increase could not be explained by climate and hydrological factors alone; the presence of wolves on the landscape was a significant predictor of stem growth above and beyond these abiotic factors. Growth-ring area was positively correlated with the previous year's ring area and negatively correlated with the percentage of twigs browsed from the stem during the winter preceding growth, indicating that elk browse impeded stem growth. Our results are consistent with the hypothesis of a behaviorally mediated trophic cascade on Yellowstone's northern winter range following wolf reintroduction. We suggest that the community-altering effects of wolf restoration are an endorsement of ecological-process management in Yellowstone National Park.     

Recolonizing wolves and mesopredator suppression of coyotes: impacts on pronghorn population dynamics.

Food web theory predicts that the loss of large carnivores may contribute to elevated predation rates and, hence, declining prey populations, through the process of mesopredator release. However, opportunities to test predictions of the mesopredator release hypothesis are rare, and the extent to which changes in predation rates influence prey population dynamics may not be clear due to a lack of demographic information on the prey population of interest. We utilized spatial and seasonal heterogeneity in wolf distribution and abundance to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus) throughout much of the United States, contributes to high rates of neonatal mortality in ungulates. To test this hypothesis, we contrasted causes of mortality and survival rates of pronghorn (Antilocapra americana) neonates captured at wolf-free and wolf-abundant sites in western Wyoming, USA, between 2002 and 2004. We then used these data to parameterize stochastic population models to heuristically assess the impact of wolves on pronghorn population dynamics due to changes in neonatal survival. Coyote predation was the primary cause of mortality at all sites, but mortality due to coyotes was 34% lower in areas utilized by wolves (P < 0.001). Based on simulation modeling, the realized population growth rate was 0.92 based on fawn survival in the absence of wolves, and 1.06 at sites utilized by wolves. Thus, wolf restoration is predicted to shift the trajectory of the pronghorn population from a declining to an increasing trend. Our results suggest that reintroductions of large carnivores may influence biodiversity through effects on prey populations mediated by mesopredator suppression. In addition, our approach, which combines empirical data on the population of interest with information from other data sources, demonstrates the utility of using simulation modeling to more fully evaluate ecological theories by moving beyond estimating changes in vital rates to analyses of population-level impacts.     

Assessment of prey vulnerability through analysis of wolf movements and kill sites.

Within predator-prey systems behavior can heavily influence spatial dynamics, and accordingly, the theoretical study of how spatial dynamics relate to stability within these systems has a rich history. However, our understanding of these behaviors in large mammalian systems is poorly developed. To address the relationship between predator selection patterns, prey density, and prey vulnerability, we quantified selection patterns for two fine-scale behaviors of a recovering wolf (Canis lupus) population in Yellowstone National Park, Wyoming, USA. Wolf spatial data were collected between November and May from 1998-1999 until 2001-2002. Over four winters, 244 aerial locations, 522 ground-based telemetry locations, 1287 km of movement data from snow tracking, and the locations of 279 wolf kill sites were recorded. There was evidence that elk (Cervus elaphus) and bison (Bison bison) densities had a weak effect on the sites where wolves traveled and made kills. Wolf movements showed a strong selection for geothermal areas, meadows, and areas near various types of habitat edges. Proximity to edge and habitat class also had a strong influence on the locations where elk were most vulnerable to predation. There was little evidence that wolf kill sites differed from the places where wolves traveled, indicating that elk vulnerability influenced where wolves selected to travel. Our results indicate that elk are more vulnerable to wolves under certain conditions and that wolves are capable of selecting for these conditions. As such, vulnerability plays a central role in predator-prey behavioral games and can potentially impact the systems to which they relate.     

Developing Metapopulation Connectivity Criteria from Genetic and Habitat Data to Recover the Endangered Mexican Wolf

Restoring connectivity between fragmented populations is an important tool for alleviating genetic threats to endangered species. Yet recovery plans typically lack quantitative criteria for ensuring such population connectivity. We demonstrate how models that integrate habitat, genetic, and demographic data can be used to develop connectivity criteria for the endangered Mexican wolf (Canis lupus baileyi), which is currently being restored to the wild from a captive population descended from 7 founders. We used population viability analysis that incorporated pedigree data to evaluate the relation between connectivity and persistence for a restored Mexican wolf metapopulation of 3 populations of equal size. Decreasing dispersal rates greatly increased extinction risk for small populations (<150–200), especially as dispersal rates dropped below 0.5 genetically effective migrants per generation. We compared observed migration rates in the Northern Rocky Mountains (NRM) wolf metapopulation to 2 habitat‐based effective distance metrics, least‐cost and resistance distance. We then used effective distance between potential primary core populations in a restored Mexican wolf metapopulation to evaluate potential dispersal rates. Although potential connectivity was lower in the Mexican wolf versus the NRM wolf metapopulation, a connectivity rate of >0.5 genetically effective migrants per generation may be achievable via natural dispersal under current landscape conditions. When sufficient data are available, these methods allow planners to move beyond general aspirational connectivity goals or rules of thumb to develop objective and measurable connectivity criteria that more effectively support species recovery. The shift from simple connectivity rules of thumb to species‐specific analyses parallels the previous shift from general minimum‐viable‐population thresholds to detailed viability modeling in endangered species recovery planning. Desarrollo de Criterios de Conectividad Metapoblacional a Partir de Datos Genéticos y de Hábitat para Recuperar al Lobo Mexicano en Peligro de Extinción     

Human Culture and Large Carnivore Conservation in North America

We examined and compared human attitudes toward wolves ( Canis lupus ), grizzly bears ( Ursus arctos horribilis ), and mountain lions ( Puma concolor ) in North America, with an emphasis on the Rocky Mountains of the United States and Canada. Primary research, literature review, and secondary data were included in the analysis. Wolves were historically persecuted by Euro-American settlers, but they have been the focus of a significant attitudinal transformation during the latter half of this century. Many now view the wolf in positive and protective ways. Considerable variation remains, however, in contemporary attitudes. Attitudes toward mountain lions have been highly ambivalent, never assuming the prominence or clarity of views on wolves. The behavior and physiology of the species, along with human psychological factors, are important reasons for the difference in human perceptions of wolves and mountain lions. We also examined attitudes toward grizzly bears. Many indigenous cultures revered grizzly bears, although European settlers were interested in exploiting and eliminating them. Today, perceptions of grizzly bears range from positive to negative. Our recommendations include targeting key groups with education programs, building support through the use of spokespeople within the target groups, integrating human and ecological concerns, and designing species-specific education initiatives in some cases.     

Dynamics of Hybridization and Introgression in Red Wolves and Coyotes

Abstract:  Hybridization and introgression are significant causes of endangerment in many taxa and are considered the greatest biological threats to the reintroduced population of red wolves ( Canis rufus ) in North Carolina (U.S.A.). Little is known, however, about these processes in red wolves and coyotes ( C. latrans ). We used individual-based simulations to examine the process of hybridization and introgression between these species. Under the range of circumstances we considered, red wolves in colonizing and established populations were quickly extirpated, persisted near the carrying capacity, or had intermediate outcomes. Sensitivity analyses suggested that the probabilities of quasi extinction and persistence of red wolves near the carrying capacity were most affected by the strength of two reproductive barriers: red wolf challenges and assortative mating between red wolves and coyotes. Because model parameters for these barriers may be difficult to estimate, we also sought to identify other predictors of red wolf population fate. The proportion of pure red wolves in the population was a strong predictor of the future probabilities of red wolf quasi extinction and persistence. Finally, we examined whether sterilization can be effective in minimizing introgression while allowing the reintroduced red wolf population to grow. Our results suggest sterilization can be an effective short-term strategy to reduce the likelihood of extirpation in colonizing populations of red wolves. Whether red wolf numbers are increased by sterilization depends on the level of sterilization effort and the acting reproductive barriers. Our results provide an outline of the conditions likely required for successful reestablishment and long-term maintenance of populations of wild red wolves in the presence of coyotes. Our modeling approach may prove generally useful in providing insight into situations involving complex species interactions when data are few.     

Paying for Tolerance: Rural Citizens' Attitudes toward Wolf Depredation and Compensation

Abstract:   As wolf ( Canis lupus ) populations recover in Wisconsin (U.S.A.), their depredations on livestock, pets, and hunting dogs have increased. We used a mail-back survey to assess the tolerance of 535 rural citizens of wolves and their preferences regarding the management of "problem" wolves. Specifically, we tested whether people who had lost domestic animals to wolves or other predators were less tolerant of wolves than neighboring residents who had not and whether compensation payments improved tolerance of wolves. We assessed tolerance via proxy measures related to an individual's preferred wolf population size for Wisconsin and the likelihood she or he would shoot a wolf. We also measured individuals' approval of lethal control and other wolf-management tactics under five conflict scenarios. Multivariate analysis revealed that the strongest predictor of tolerance was social group. Bear ( Ursus americanus ) hunters were concerned about losing valuable hounds to wolves and were more likely to approve of lethal control and reducing the wolf population than were livestock producers, who were more concerned than general residents. To a lesser degree, education level, experience of loss, and gender were also significant. Livestock producers and bear hunters who had been compensated for their losses to wolves were not more tolerant than their counterparts who alleged a loss but received no compensation. Yet all respondents approved of compensation payments as a management strategy. Our results indicate that deep-rooted social identity and occupation are more powerful predictors of tolerance of wolves than individual encounters with these large carnivores.     

Hybridization between Wolves and Dogs

Concern has been expressed that European populations of gray wolves ( Canis lupus ) have extensively hybridized with domestic dogs ( C. familiaris ). We reviewed and analyzed surveys of mitochondrial and biparentally inherited genetic markers in dogs and wild populations of wolf-like canids. Although dog-wolf hybrids have been observed in the wild, significant introgression of dog markers into wild wolf populations has not yet occurred. Our investigation suggests that hybridization may not be an important conservation concern even in small, endangered wolf populations near human settlements. The behavioral and physiological differences between domestic dogs and gray wolves may be sufficiently great such that mating is unlikely and hybrid offspring rarely survive to reproduce in the wild.     

Effects of Social Structure and Prey Dynamics on Extinction Risk in Gray Wolves

Extinction models based on diffusion theory generally fail to incorporate two important aspects of population biology—social structure and prey dynamics. We include these aspects in an individual-based extinction model for small, isolated populations of the gray wolf (Canis lupus). Our model predicts mean times to extinction significantly longer than those predicted by more general (diffusion) models. According to our model, an isolated population of 50 wolves has a 95% chance of surviving just 9 years and only a 30% chance of surviving beyond 100 years. Reflecting the influence of social structure, a wolf population initially comprising 50 individuals is expected to persist only a few years longer, on average (71 years), than is a population initially comprising just a single reproductive pair (62 years). In contrast, substantially greater average prey abundance leads to dramatically longer expected persistence times. Autocorrelated prey dynamics result in a more complex distribution of extinction times than predicted by many extinction models. We contend that demographic stochasticity may pose the greatest threat to small, isolated wolf populations, although environmental stochasticity and genetic effects may compound this threat. Our work highlights the importance of considering social structure and resource dynamics in the development of population viability analyses.     

Hunting Effects on Favourable Conservation Status of Highly Inbred Swedish Wolves

The wolf (Canis lupus) is classified as endangered in Sweden by the Swedish Species Information Centre, which is the official authority for threat classification. The present population, which was founded in the early 1980s, descends from 5 individuals. It is isolated and highly inbred, and on average individuals are more related than siblings. Hunts have been used by Swedish authorities during 2010 and 2011 to reduce the population size to its upper tolerable level of 210 wolves. European Union (EU) biodiversity legislation requires all member states to promote a concept called “favourable conservation status” (FCS) for a series of species including the wolf. Swedish national policy stipulates maintenance of viable populations with sufficient levels of genetic variation of all naturally occurring species. Hunting to reduce wolf numbers in Sweden is currently not in line with national and EU policy agreements and will make genetically based FCS criteria less achievable for this species. We suggest that to reach FCS for the wolf in Sweden the following criteria need to be met: (1) a well‐connected, large, subdivided wolf population over Scandinavia, Finland, and the Russian Karelia‐Kola region should be reestablished, (2) genetically effective size (N_e) of this population is in the minimum range of N_e = 500–1000, (3) Sweden harbors a part of this total population that substantially contributes to the total N_e and that is large enough to not be classified as threatened genetically or according to IUCN criteria, and (4) average inbreeding levels in the Swedish population are <0.1. Efectos de la Cacería sobre el Estatus de Conservación Favorable de Lobos Suecos con Endogamia Alta     

Biological, Conservation, and Ethical Implications of Exploiting and Controlling Wolves

The widespread claim that wolf populations can withstand 25–50% or greater annual reductions without major biological consequences is based primarily on the observation that populations often maintain their size from year to year as harvest or control continues or recover within a few years afterward. This emphasis on numerical status overlooks the likelihood of major, lingering impacts on the size, number, stability, and persistence of family-group social units, on reproductive, hunting, and territorial behavior, on the role of learning and related traditions, on within- and between-group patterns of genetic variation, and on overall mortality rates. The tendency of biologists and agencies in northern North America to promote wolf harvests that are four to eight times greater than ungulate harvests, in accord with the wolf versus ungulate difference in reproductive rates but contradictory to a broad array of differences in social organization and related behavior, is reason enough to question the logic of this prevailing management view. True sustained-yield management requires more emphasis on qualitative biological features to determine the extent to which wolves and other species with evolutionary histories as predators rather than as prey should be harvested. Most recent government-sponsored wolf control programs and proposals, including sterilization, relocation, and "redirected" killing, have been based on questionable claims about ungulate or livestock problems and have not adequately considered potential biological costs (especially to the target wolf populations), benefits, or management alternatives. The high sentience of wolves justifies overlapping biological-ethical concerns about such programs and especially about the heavy, indiscriminate, deceptively reported public hunting and trapping of wolves that is currently permitted throughout most of Alaska (U.S.A.)—including in national parks—and elsewhere.     

Inbreeding Depression in a Captive Wolf (Canis lupus) Population

Abstract. Uncertainty currently exists regarding the extent to which mammalian carnivores suffer from inbreeding depression. In particular, it has been proposed that wolves and species with a similar social structure are adapted to close inbreeding. Empirical data, however, are scarce. This paper provides strong evidence against the contention that natural populations of wolves are resistant to inbreeding depression. We analyzed studbook data of a captive wolf population bred in Scandinavian zoos and found negative effects of inbreeding expressed as reductions in juvenile weight, reproduction, and longevity. The occurrence of an apparently bereditary form of blindness is also associated with inbreeding. Different effects of inbreeding can be attributed to genes originating from different founder pairs, thus indicating that alleles that are deleterious in the homozygous state are fairly common in natural wolf populations.     

Evaluating Wolf Translocation as a Nonlethal Method to Reduce Livestock Conflicts in the Northwestern United States

Abstract:  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.