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.     

Extremes in the resource consumption process: A mathematical model of the irruption of the moose population on Isle Royale. Part 1—Ecological background and model development

The development of a mathematical model of the moose-browse interaction on Isle Royale, Michigan, from the time of the establishment of the moose population in the early 1900s until the time of the arrival of wolves on the island about the mid-twentieth century is described. A brief history of the observations that were made during that period of the dynamics of the moose population and its resulting impact upon the vegetation is given. The simplifying assumptions that were made, the processes that were judged important enough to be represented in the model, the state variables that were selected and the input variables that were considered are delineated. A rationale is developed for the mathematical form of each of the terms used in the state equations. The literature that was utilized and the methodologies that were employed to obtain estimates of the parameters of the equations and the values of the initial conditions of the state variables are described.     

Genetic Variation for Morphological and Allozyme Variation in Relation to Population Size in Clarkia dudleyana, an Endemic Annual

Abstract: The maintenance of genetic variation within populations is expected to allow species to respond to evolutionary challenges such as selection and environmental stress. Larger populations are generally expected to maintain larger amounts of genetic variation. Although several studies have found a positive relationship between population size and levels of genetic variation for molecular markers such as allozymes, few comparisons have been made between molecular measures of variation and genetic variation that is likely to be ecologically important. Most ecologically important traits require quantitative genetic analyses. I examined the relationship between levels of genetic variation and population size for both allozymes and morphological traits in a California endemic annual plant, Clarkia dudleyana . Levels of genetic variation for allozymes did not show a significant positive relationship with population size. The level of genetic variance for all of the 18 morphological traits exhibited no significant relationship with population size. Further, allozyme heterozygosities were not related to levels of quantitative genetic variation. These results indicate that levels of allozyme variability do not predict levels of genetic variation for morphological traits in C. dudleyana , suggesting that molecular measures of variation, in general, differ from quantitative genetic measures. These results imply that conservation genetic studies should generally focus on aspects other than measuring levels of genetic variation found within populations.     

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     

Genetic Variation of Naturally Colonizing Wolves in the Central Rocky Mountains

Recovery of gray wolf ( Canis lupus ) populations in North America depends on minimizing human-caused mortality and enhancing migration from stable source populations to suitable habitat unoccupied by wolves. We used a combination of field observation and DNA microsatellite genotyping to examine natural wolf colonization of Glacier National Park, Montana, and surrounding lands. We found high genetic variation in the colonizing population, showing that these packs were founded by multiple, unrelated wolves from Canada. High dispersal rates, long dispersal distances, and lack of a founding population bottleneck indicate that wolves in the United States and Canada should be viewed and managed as a single population. Restoration in the United States by artificial transplants from Alberta to Yellowstone National Park and central Idaho began in 1995. The transplanted wolves will likely aid demographic recovery, but permanently retaining the high genetic variation of wolves in the United States will require assuring gene flow throughout the central Rocky Mountains.     

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.     

Genetic Variation and the Immune Response in Natural Populations of Pocket Gophers

It is thought that genetic variation can affect the persistence of a population through its influence on disease susceptibility. We assessed genome-wide genetic variation, variation at a locus involved in the immune system, and acceptance or rejection of skin grafts in three natural populations of the pocket gopher ( Thomomys bottae ). Multilocus DNA fingerprints confirmed previous allozyme data, revealing high levels of variation among Hastings Reserve pocket gophers and almost complete within-population identity for individuals from the two Patricks Point populations (Patricks J and Patricks F), although Patricks J animals were dissimilar to animals from Patricks F despite their proximity. Individuals from the high-variation population consistently rejected within-population reciprocal skin grafts, whereas Patricks J and Patricks F individuals accepted within-population grafts. Patricks J and Patricks F individuals were found to be immunocompetent, however, as revealed by the ability of all individuals to reject between-population grafts, including those that previously accepted within-population grafts. A DNA heteroduplex analysis was then used to directly characterize variability at DQα, a locus of the immune system's major histocompatability complex. Both populations low in genetic variation were fixed for unique DQα alleles, whereas observed heterozygosity in the Hastings population was 0.43, ascribable to at least three unique alleles. These data are in accord with previous cheetah skin-graft results and confirm that skin grafts can be used to assess genetic similarity. We suggest that although many animal populations can persist with extremely low levels of genetic variation in the wild, such populations may be at a greater risk of extinction from particular pathogens because of their genetic uniformity.     

Revisiting the classics: considering nonconsumptive effects in textbook examples of predator-prey interactions

Predator effects on prey dynamics are conventionally studied by measuring changes in prey abundance attributed to consumption by predators. We revisit four classic examples of predator-prey systems often cited in textbooks and incorporate subsequent studies of nonconsumptive effects of predators (NCE), defined as changes in prey traits (e.g., behavior, growth, development) measured on an ecological time scale. Our review revealed that NCE were integral to explaining lynx-hare population dynamics in boreal forests, cascading effects of top predators in Wisconsin lakes, and cascading effects of killer whales and sea otters on kelp forests in nearshore marine habitats. The relative roles of consumption and NCE of wolves on moose and consequent indirect effects on plant communities of Isle Royale depended on climate oscillations. Nonconsumptive effects have not been explicitly tested to explain the link between planktonic alewives and the size structure of the zooplankton, nor have they been invoked to attribute keystone predator status in intertidal communities or elsewhere. We argue that both consumption and intimidation contribute to the total effects of keystone predators, and that characteristics of keystone consumers may differ from those of predators having predominantly NCE. Nonconsumptive effects are often considered as an afterthought to explain observations inconsistent with consumption-based theory. Consequently, NCE with the same sign as consumptive effects may be overlooked, even though they can affect the magnitude, rate, or scale of a prey response to predation and can have important management or conservation implications. Nonconsumptive effects may underlie other classic paradigms in ecology, such as delayed density dependence and predator-mediated prey coexistence. Revisiting classic studies enriches our understanding of predator-prey dynamics and provides compelling rationale for ramping up efforts to consider how NCE affect traditional predator-prey models based on consumption, and to compare the relative magnitude of consumptive and NCE of predators.     

The Pending Extinction of the Uncompahgre Fritillary Butterfly

Previous mark-recapture studies and population size estimates indicated that in the 1980s populations of the endangered Uncompahgre fritillary ( Boloria acrocnema ) declined precipitously, apparently leading to extirpation at its type locality. This locality and a nearby second site, both high in the San Juan Mountains of southwestern Colorado, constitute the total known range of B. acrocnema . A survey of 20 presumptive allozyme loci from one B. acrocnema population and seven populations of the closely related B. improba , sampled along the Rocky Mountain cordillera from the Yukon Territory, Canada, to the Wind River Range of Wyoming, revealed that B. acrocnema populations are less variable genetically than are B. improba populations in the northern part of that species' North American range. Genetic variability was also assayed at 18 presumptive allozyme loci in populations of Boloria titania, which were sampled from five of the same locations for comparison with the B. improba-B. acrocnema clade. Low estimates of heterozygosity indicate that genetic and demographic factors preceded by historic human disturbance are causing the decline of B. acrocnema . In addition, persistent drought conditions in southwestern Colorado throughout the 1980s probably contributed to the population declines. Conservation recommendations include strict enforcement of the ban against collecting pursuant to the Endangered Species Act, careful monitoring of known B. acrocnema sites, and possible translocations. Extreme measures beyond these will be costly and may be inappropriate for this relict species, which has an unknown capacity to persist in the face of local weather fluctuations and predicted regional warming trends.     

Population History, Genetic Variability, and Horn Growth in Bighorn Sheep

Bighorn sheep ( Ovis canadensis ) are restricted in distribution and numbers relative to presettlement conditions. Some populations have alledgedly suffered losses of fitness resulting from small, insular populations and a breeding system that reduces effective population size. Large horns in rams, which confer breeding superiority, are absent from some populations, and this absence may result in part from loss of genetic variability. We investigated the relationship among allozyme variability, population history, and horn growth in bighorn sheep from the Rocky Mountains. Heterozygosity was higher for bighorn sheep than has been reported for Dall sheep ( O. dalli ). Heterozygosity and allelic variability were marginally related to effective population size for the proceeding 15 years. Horn growth was significantly higher in more heterozygous than in less heterozygous rams for years 6, 7, and 8 of life. By the end of year 8, more heterozygous rams had 13% higher horn volumes than less heterozygous rams. Most hunting of bighorn sheep involves selective removal of large-horned rams, which we hypothesize may reduce genetic variability of these populations and contribute to losses in fitness.     

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.     

Genetic population structure in the black-spot sea bream (<Emphasis Type="Italic">Pagellus bogaraveo</Emphasis> Brünnich, 1768) from the NE Atlantic

The depletion of shallow-water fish stocks through overexploitation has led to increasing fishing pressure on deep-sea species. Poor knowledge of the biology of commercially valuable deep-water fish has led to the serial depletion of stocks of several species across the world. Data regarding the genetic structure of deep-sea fish populations is important in determining the impact of overfishing on the overall genetic variability of species and can be used to estimate the likelihood of recolonisation of damaged populations through immigration of individuals from distant localities. Here the genetic structure of the commercially fished deep-water species the blackspot sea bream, Pagellus bogaraveo is investigated in the northeastern Atlantic using partial DNA sequencing of mitochondrial cytochrome b (cyt-b) and D-loop regions and genotyping of microsatellite loci. An absence of variation in cyt-b and low genetic variation in D-loop sequences potentially indicate that P. bogaraveo may have undergone a severe bottleneck in the past. Similar bottlenecks have been detected in other Atlantic species of fish and have possibly originated from the last glaciation. P. bogaraveo may have been particularly vulnerable to the effects of low temperature and a fall in sea level because stages of its life history occur in shallow water and coastal sites. However, there are other explanations of low genetic variability in populations of P. bogaraveo, such as a low population size and the impacts of fishing on population structure. Analysis of population structure using both D-loop and microsatellite analysis indicates low to moderate, but significant, genetic differentiation between populations at a regional level. This study supports studies on other deep-sea fish species that indicate that hydrographic or topographic barriers prevent dispersal of adults and/or larvae between populations at regional and oceanographic scales. The implications for the management and conservation of deep-sea fish populations are discussed.Communicated by J.P. Thorpe, Port Erin     

Environmental Surrogates for Predicting and Conserving Adaptive Genetic Variability in Tree Species

Abstract: Adaptive genetic variability within species is an essential component of biodiversity but has been largely ignored in studies aimed at assessing and predicting biodiversity of the forest environment. We used factorial regression and structuring models to test easily measured surrogates, such as ecological attributes, as predictors of adaptive genetic variation between populations of a tree species ( Eucalyptus delegatensis ). Adaptive variability was defined in terms of variation in average growth performance of 68 populations and of population-by-environment interaction across seven different environments. The best surrogates of genetic variability were measures of solar radiation and temperature range, each predicting more than 50% of the genetic variability within the species. Rock and understory types, when used either alone or in combination with other covariates, also were very efficient in discriminating between populations in groups showing similar adaptation. Significant relationships between particular surrogates and growth patterns of variation were attributed to effects of natural selection that had occurred in the population source locations. We recommend the development of studies focusing on the population level of biodiversity to improve the conservation of forest ecosystems in Australia.     

Genetic Diversity and Tests of the Hybrid Origin of the Endangered Yellow Larkspur

Abstract: Delphinium luteum ( Ranunculaceae), an endangered larkspur, is restricted to two wild populations near Bodega Bay, California. The total number of individuals in these two populations is estimated to be <100. We used allozyme and random amplified polymorphic DNA ( RAPD) markers to (1) assess levels and patterns of genetic diversity in one wild population and two cultivated populations and (2) test the hypothesis that D. luteum is of hybrid origin between D. decorum and D. nudicaule . These data will be used to aid in developing a management plan to conserve the species. The wild population maintains high levels of genetic diversity. Genetic data indicate that both cultivated populations, especially the north Sonoma population, have several allozymes and RAPD markers not found in the wild population and could be used to establish new populations of D. luteum or to enhance the diversity and size of the wild population. The allozyme data did not reveal any fixed differences between D. decorum and D. nudicaule , although allele frequencies of the putative parental populations differed. At these loci, D. luteum resembled D. nudicaule more than D. decorum  . Many unique RAPD markers distinguish each of the three species. The diagnostic markers from populations of D. nudicaule and D. decorum were not additive in the putative hybrid, and these data indicate that D. luteum is not of recent hybrid origin. Conservation of the yellow larkspur should include strategies that use the cultivated populations of D. luteum , but hybridizing D. decorum and D. nudicaule to "recreate" D. luteum is not recommended.     

Darwin's Fox: A Distinct Endangered Species in a Vanishing Habitat

The temperate rain forest of Chiloé Island, Chile, is inhabited by an endemic fox ( Dusicyon fulvipes ) first described by Charles Darwin and now designated Darwin's fox. Despite morphological differences, Darwin's fox has been considered only an insular subspecies of the mainland chilla fox ( D. griseus ). This follows the assumption that the island population, with an estimated population of less than 500, has been separated from the mainland chilla fox for only about 15,000 years and may have received occasional immigrants from the mainland. Consequently, this island population has not been protected as endangered or bred in captivity. Recently, a population of Darwin's fox was discovered on the Chilean mainland 600 km north of Chiloé Island. This population exists in sympatry with chilla and possibly culpeo ( D. culpaeus ) foxes, which suggests that Darwin's fox may be reproductively isolated. To clarify the phylogenetic position of Darwin's fox, we analyzed 344 bp of mitochondrial DNA control-region sequence of the three species of Chilean foxes. Darwin's foxes from the island and mainland populations compose a monophyletic group distinct from the two other Chilean fox species. This indicates that Darwin's fox was probably an early inhabitant of central Chile, and that its present distribution on the mainland may be a relict of a once much wider distribution. Our results highlight the ability of molecular genetic techniques to uncover historical relationships masked by recent events, such as local extinctions. The "rediscovery" of Darwin's fox as a distinct species implies that greater significance should be given to the protection of this species and its unique habitat and to documenting the extent of its mainland distribution.     

High frequency of extra-pair paternity in Swedish pied flycatchers revealed by allozyme electrophoresis and DNA fingerprinting

Summary We used two genetic techniques to study multiple parentage in the pied flycatcher (Ficedula hypoleuca). Allozyme electrophoresis of 131 nestlings from 22 clutches sampled in 1982 and 1987 showed that one nestling had a mismatched allele compared with its putative parents. In one additional nest the devitation from Mendelian inheritance of parental genotypes suggested extra-pair paternity. The calculated probability of detecting multiple paternity from the genetic variation at four loci was 0.115. The estimated population frequency of extra-pair fertilization (EPF) was 13%, based on two mismatches and the probability of detecting multiple paternity. The seven families (n = 38 nestlings) in the 1987 sample were also analyzed by DNA fingerprinting using the M13 and Jeffrey's 33.15 probes. Overall, 24% (n = 9) of the nestlings analyzed were genetically inconsistent with their putative father, with EPF occurring in three (43%) of the seven clutches. One nestling originated by intraspecific brood parasitism. Comparison of the two techniques using the same samples showed that allozyme electrophoresis has a resolution only 11% of that of DNA fingerprinting, close to the calculated probability of detecting multiple paternity (0.115). Both techniques suggest that extra-pair fertilization is relatively common in the Swedish populations investigated compared to the low frequency reported from a Norwegian population. Correspondence to: H.P. Gelter at the present address     

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.     

Genetic Introgression of Endemic Taxa by Non-natives: A Case Study with Leon Springs Pupfish and Sheepshead Minnow

Genetic studies of a pupfish (Cyprinodon bovinus) endemic to a small, spring-fed system in west Texas illustrate the potential for small introductions of non-native species to cause large-scale genetic changes through hybridization and genetic introgression. We performed a genetic survey (allozymes and RFLP analysis of mtDNA) of four samples of C. bovinus representing all wild populations of the species and a captive population maintained since 1976 at Dexter, New Mexico. The results indicate genetic introgression of the entire wild population by sheepshead minnow (C. variegatus), a coastal species with a history of introductions in west Texas. Frequencies of foreign genetic elements averaged across four diagnostic allozyme loci and mtDNA varied from 6.1 to 15.1%. The captive population appears free of foreign genetic material. Comparisons with past studies of C. bovinus indicate the present situation is largely due to a recent introduction of C. variegatus, not to an introduction in the mid-1970s; however, residual effects from the earlier introduction cannot be completely discounted. Genetic analysis indicates that the source of introduced C. variegatus in Diamond Y Draw is the nearest known population, an introduced stock in Lake Balmorhea approximately 90 km away. The results demonstrate the value of maintaining imperiled species in captivity. Captive C. bovinus provide an opportunity to restore the genetic integrity of wild populations.     

Implications from Mitochondrial DNA for Management to Conserve the Eastern Barred Bandicoot (Perameles gunnii)

On mainland Australia, the eastern barred bandicoot, Perameles gunnii , is confined to a relic wild population numbering less than 100 individuals in the city of Hamilton. Animals derived from this population are being bred in captivity in order to promote their recovery. The species also exists in Tasmania, where tittle is known of its conservation and taxonomic status. Mitochondrial DNA variability was compared within and between populations of P. gunnii using restriction fragment length polymorphisms. Genetic variability was found to be high among P. gunnii in Hamilton compared to those in Tasmania (higher diversity index, nucleotide sequence divergence, and greater number of haplotypes), despite the known decline and subdivision of the Hamilton population. Restriction fragment length polymorphisms distinguished animals from the east and the west of Hamilton and from the north and south of Tasmania. Nucleotide sequence divergence was substantial (2.2–2.5%) between Hamilton and Tasmania. Implications are that captive breeding and reintroduction should be designed to genetically represent the structure within Hamilton in order to minimize inbreeding and that the introduction of Tasmanian P. gunnii would not benefit the Hamilton population. It is concluded that mitochondrial DNA markers clearly can provide useful information about the history and current status of endangered marsupial populations, to the benefit of conservation management.     

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.