Genetic nature of eastern wolves: Past, present and future

Eastern North American wolves have long been recognized as morphologically distinct from both coyotes and gray wolves. This has led to questions regarding their origins and taxonomic status. Eastern wolves are mainly viewed as: (1) a smaller subspecies of gray wolf (Canis lupus lycaon), potentially the result of historical hybridization between gray wolves (C. lupus) and red wolves (C. rufus), (2) a hybrid, the result of gray wolf (C. lupus) and coyote (C. latrans) interbreeding, or (3) a distinct species, C. lycaon, closely related to the red wolf (C. rufus). Although debate persists, recent molecular studies suggest that the eastern wolf is not a gray wolf subspecies, nor the result of gray wolf/coyote hybridization. Eastern wolves were more likely a distinct species, C. lycaon, prior to the eastward spread of coyotes in the late 1800s. However, contemporary interbreeding exits between C. lycaon to both C. lupus and C. latrans over much of its present range complicating its present taxonomic characterization. While hybridization may be reducing the taxonomic distinctiveness of C. lycaon, it should not necessarily be viewed as negative influence. Hybridization may be enhancing the adaptive potential of eastern wolves, allowing them to more effectively exploit available resources in rapidly changing environments.     

From a Howling Wilderness to Howling Safaris: Science,Policy and Red Wolves in the American South

This article details the complex natural and cultural history of red wolf (Canis rufus) restoration in the American South. The decisions and methods utilized in the red wolf’s recovery after 1960 were unprecedented and creative but not geographically limited. The federal red wolf recovery experiment highlights the debate over what constitutes a species in a dynamic world, and the practical challenges and unexpected results in endangered species management in peopled landscapes. This wildlife restoration story illustrates not only the “hands-on” management role humans played, and continue to play, but also reveals cultural assumptions about what constitutes a “wild” wolf and about the necessity of wilderness. The red wolf recovery project provides constructive lessons for future species restoration involving flora and fauna on public and private land, and demonstrates human and animal engagement in the making of nature and culture.     

Wolf Depredation on Domestic Animals in the Polish Carpathian Mountains

Abstract As wolves (Canis lupus) recover in Poland, their depredation on domestic animals is increasing, as have conflicts between wolves and farmers. From 1998 to 2004, I investigated spatial and temporal patterns of 591 verified incidents of wolf depredation in the eastern part of the Polish Carpathian Mountains. The wolf population I surveyed covered an estimated range of 4,993 km². Depredation occurred over 1,595 km² of that area. Sheep accounted for 84.8% of domestic animals killed by wolves. Depredation on sheep and number of sheep farms attacked by wolves increased between 1998 and 2004 (r² = 0.61, P = 0.04 and r² = 0.89, P = 0.02, respectively). The number of wolf attacks on sheep farms in a given year were negatively correlated to red deer (Cervus elaphus) population numbers (R² = 0.69, P = 0.02). The amount of depredation caused by each of the 4 monitored packs was best explained by farm density in their territories (R² = 0.59, P = 0.004). Number of attacks recorded on farms was positively correlated to distance from the farm to the pack's den and rendezvous sites (R² = 0.16, P = 0.04). Of depredation recorded in the 4 pack's territories I surveyed, 77% occurred in 4 farms with no or inadequate protection. I concluded that wolf depredation in the studied area is opportunistic. Wolf predation intensity is a function of decreasing abundance of red deer, the density of sheep farms, and proximity of farms to the summer activity centers of wolf packs, and it is facilitated by poor husbandry practices. These results can aid in preventing wolf depredation and provide a foundation for a wolf management plan.     

Late-Pleistocene and early Holocene history of the canid fauna of Europe (Canidae)

In the sub-fossil assemblages of Europe the red fox is clearly the most frequent carnivorous mammalian species with a total of 1553 records. In depositions from the Weichselian Glacial the red fox Vulpes vulpes is, a typical representative of the Holocene fauna, already recorded in 100 assemblages. The Iberian peninsula, Italian peninsula and Balkans were theorised as glacial refugia. Well-founded facts give reason to believe that V. vulpes was also distributed in the Carpathian refuge. Later on, the Crimean peninsula would also appear to be a possible glacial refuge of the red fox.In the last warmer complex of interstadials during the Pleni-Glacial (Hengelo-Denekamp, 38,000–25,000 BC) the red fox was distributed in central Europe. Its distribution during this epoch extended at least in part to southern England. The earliest well-dated records of V. vulpes in central Europe after the Maximum Glaciation lie between 14,000 and 13,500 BC. Already during the early Late-Glacial (13,500 BC) the red fox appeared in typical glacial faunal communities. A separation to glacial refugia was only possible for 10,000 years.During the last warmer Pleni-Glacial complex of interstadials (38,000–25,000 BC) in central Europe a sympatric distribution of the arctic fox (Alopex lagopus) and the red fox probably existed. During the Last Glacial Maximum (22,000–18,000 BC) the arctic fox was exclusively distributed in central Europe, outside of the refuges. The combined distribution of A. lagopus and V. vulpes during the Late-Glacial (15,000–9500 BC) in central Europe, with the probable exception of the Allerød, is precisely documented by sub-fossil assemblages.In the Pleni-Glacial the wolf Canis lupus was distributed in geographic regions that served as glacial refugia of more warm-climate adapted species. Concerning the wolf no drastic decrease of the distribution is assumed. The Holocene presence of C. lupus is probably not caused by recolonisation.

Zusammenfassung

Die Entwicklung der Canidenfauna Europas im Spätpleistozän und frühen HolozänIn den subfossilen Ablagerungen ist Vulpes vulpes in Europa mit 1553 Nachweisen das mit Abstand am häufigsten nachgewiesene Raubsäugetier. Allein in den Ablagerungen des Weichselglazials konnte der Rotfuchs bereits in 100 Fundkomplexen nachgewiesen werden. Als Glazialrefugium des Rotfuchses wird die Iberische Halbinsel sicher identifiziert. Eine Verbreitung der Art während des Kältemaximums wird außerdem auf der Apenninen-Halbinsel sowie der Balkan- Halbinsel vermutet. Außerdem liegen fundierte Fakten für die Annahme vor, dass V. vulpes im Karpatenrefugium verbreitet war. Auf der Halbinsel Krim scheint ein Glazialrefugium für des Fuchses möglich.Während des letzten wärmeren Interstadial-Komplexes im Hochglazial (Hengelo-Denekamp, ca. 38.000–25.000 v. Chr.) war Mitteleuropa vom Rotfuchs besiedelt. Die nördliche Arealgrenze der Art war innerhalb dieses Klimaabschnitts mindestens zeitweise bis nach Südengland ausgedehnt. Die frühesten, zeitlich relativ gut abgesicherten Nachweise von V. vulpes nach der Weichsel-Maximalvereisung in Mitteleuropa liegen etwa zwischen 14.000–13.500 v. Chr. Schon im frühen Spätglazial (ca. 13.500 v. Chr.) kam der Rotfuchs im nördlichen Mitteleuropa in typischen glazialen Faunengemeinschaften vor. Eine Disjunktion des Areals während der letzten Vereisung kann für höchstens 10.000 Jahre stattgefunden haben. Es wird angenommen, dass während des Hengelo-Deenekamp Interstadials ein sympatrisches Vorkommen von Eisfuchs Alopex lagopus und Rotfuchs V. vulpes in Mitteleuropa existierte. Nur zur Zeit des absoluten Kältemaximums (ca. 22.000–18.000 v. Chr.) war außerhalb der Refugialgebiete in Mitteleuropa ausschließlich A. lagopus verbreitet. Das gemeinsame Vorkommen (von A. lagopus und V. vulpes) während des gesamten Spätglazials in Mitteleuropa, wahrscheinlich mit Ausnahme des Allerød-Interstadials, ist präzise belegt.Während des Hochglazials war Canis lupus auch in geografischen Regionen verbreitet, die für an wärmeres Klima gebundene Tierarten den Charakter von Glazialrefugien hatten. Beim Wolf C. lupus kann keine extreme Arealverringerung während der Weichseleiszeit angenommen werden. Das holozäne Vorkommen von C. lupus in Mitteleuropa dürfte daher nicht generell auf eine Rekolonisation zurückzuführen sein.     

An efficient method for screening faecal DNA genotypes and detecting new individuals and hybrids in the red wolf (<Emphasis Type="Italic">Canis rufus</Emphasis>) experimental population area

Previously, sequencing of mitochondrial DNA (mtDNA) from non-invasively collected faecal material (scat) has been used to help manage hybridization in the wild red wolf (Canis rufus) population. This method is limited by the maternal inheritance of mtDNA and the inability to obtain individual identification. Here, we optimize the use of nuclear DNA microsatellite markers on red wolf scat DNA to distinguish between individuals and detect hybrids. We develop a data filtering method in which scat genotypes are compared to known blood genotypes to reduce the number of PCR amplifications needed. We apply our data filtering method and the more conservative maximum likelihood ratio method (MLR) of Miller et al. (2002 Genetics 160:357–366) to a scat dataset previously screened for hybrids by sequencing of mtDNA. Using seven microsatellite loci, we obtained genotypes for 105 scats, which were matched to 17 individuals. The PCR amplification success rate was 50% and genotyping error rates ranged from 6.6% to 52.1% per locus. Our data filtering method produced comparable results to the MLR method, and decreased the time and cost of analysis by 25%. Analysis of this dataset using our data filtering method verified that no hybrid individuals were present in the Alligator River National Wildlife Refuge, North Carolina in 2000. Our results demonstrate that nuclear DNA microsatellite analysis of red wolf scats provides an efficient and accurate approach to screen for new individuals and hybrids.     

Evaluating the ability of Bayesian clustering methods to detect hybridization and introgression using an empirical red wolf data set

Bayesian clustering methods have emerged as a popular tool for assessing hybridization using genetic markers. Simulation studies have shown these methods perform well under certain conditions; however, these methods have not been evaluated using empirical data sets with individuals of known ancestry. We evaluated the performance of two clustering programs, baps and structure , with genetic data from a reintroduced red wolf (Canis rufus) population in North Carolina, USA. Red wolves hybridize with coyotes (C. latrans), and a single hybridization event resulted in introgression of coyote genes into the red wolf population. A detailed pedigree has been reconstructed for the wild red wolf population that includes individuals of 50–100% red wolf ancestry, providing an ideal case study for evaluating the ability of these methods to estimate admixture. Using 17 microsatellite loci, we tested the programs using different training set compositions and varying numbers of loci. structure was more likely than baps to detect an admixed genotype and correctly estimate an individual's true ancestry composition. However, structure was more likely to misclassify a pure individual as a hybrid. Both programs were outperformed by a maximum‐likelihood‐based test designed specifically for this system, which never misclassified a hybrid (50–75% red wolf) as a red wolf or vice versa. Training set composition and the number of loci both had an impact on accuracy but their relative importance varied depending on the program. Our findings demonstrate the importance of evaluating methods used for detecting admixture in the context of endangered species management.     

Recolonizing gray wolves increase parasite infection risk in their prey

The recent recolonization of Central Europe by the European gray wolf (Canis lupus) provides an opportunity to study the dynamics of parasite transmission for cases when a definitive host returns after a phase of local extinction. We investigated whether a newly established wolf population increased the prevalence of those parasites in ungulate intermediate hosts representing wolf prey, whether some parasite species are particularly well adapted to wolves, and the potential basis for such adaptations. We recorded Sarcocystis species richness in wolves and Sarcocystis prevalence in ungulates harvested in study sites with and without permanent wolf presence in Germany using microscopy and DNA metabarcoding. Sarcocystis prevalence in red deer (Cervus elaphus) was significantly higher in wolf areas (79.7%) than in control areas (26.3%) but not in roe deer (Capreolus capreolus) (97.2% vs. 90.4%) or wild boar (Sus scrofa) (82.8% vs. 64.9%). Of 11 Sarcocystis species, Sarcocystis taeniata and Sarcocystis grueneri occurred more often in wolves than expected from the Sarcocystis infection patterns of ungulate prey. Both Sarcocystis species showed a higher increase in prevalence in ungulates in wolf areas than other Sarcocystis species, suggesting that they are particularly well adapted to wolves, and are examples of “wolf specialists”. Sarcocystis species richness in wolves was significantly higher in pups than in adults. “Wolf specialists” persisted during wolf maturation. The results of this study demonstrate that (1) predator–prey interactions influence parasite prevalence, if both predator and prey are part of the parasite life cycle, (2) mesopredators do not necessarily replace the apex predator in parasite transmission dynamics for particular parasites of which the apex predator is the definitive host, even if meso‐ and apex predators were from the same taxonomic family (here: Canidae, e.g., red foxes Vulpes vulpes), and (3) age‐dependent immune maturation contributes to the control of protozoan infection in wolves.     

Y‐chromosome evidence supports widespread signatures of three‐species Canis hybridization in eastern North America

There has been considerable discussion on the origin of the red wolf and eastern wolf and their evolution independent of the gray wolf. We analyzed mitochondrial DNA (mtDNA) and a Y‐chromosome intron sequence in combination with Y‐chromosome microsatellites from wolves and coyotes within the range of extensive wolf–coyote hybridization, that is, eastern North America. The detection of divergent Y‐chromosome haplotypes in the historic range of the eastern wolf is concordant with earlier mtDNA findings, and the absence of these haplotypes in western coyotes supports the existence of the North American evolved eastern wolf (Canis lycaon). Having haplotypes observed exclusively in eastern North America as a result of insufficient sampling in the historic range of the coyote or that these lineages subsequently went extinct in western geographies is unlikely given that eastern‐specific mtDNA and Y‐chromosome haplotypes represent lineages divergent from those observed in extant western coyotes. By combining Y‐chromosome and mtDNA distributional patterns, we identified hybrid genomes of eastern wolf, coyote, gray wolf, and potentially dog origin in Canis populations of central and eastern North America. The natural contemporary eastern Canis populations represent an important example of widespread introgression resulting in hybrid genomes across the original C. lycaon range that appears to be facilitated by the eastern wolf acting as a conduit for hybridization. Applying conventional taxonomic nomenclature and species‐based conservation initiatives, particularly in human‐modified landscapes, may be counterproductive to the effective management of these hybrids and fails to consider their evolutionary potential.     

Legal protection of wolves in Poland: implications for the status of the wolf population

Legal protection of wolves (Canis lupus) in Poland was implemented in 1998 after 23 years of management as a game species. Wolves occurring in Poland were interconnected with larger populations in the Carpathian Mountains and Belarus, Baltic States and Russia, stable in numbers, and were not considered endangered before the change in legal status affording protection from hunting. Parties calling for wolf protection wanted to stop killing of wolves because of their symbolic nature, but did not have particular management goals to achieve. The government did not accompany the change in legal status by management plan, and therefore, the ban on wolf hunting was weakly enforced. A wolf distribution monitoring demonstrated that wolf range had not expanded 9 years after the hunting ban was implemented, and no increase in wolf numbers was observed. This failure to recover may be explained by: (1) a significant (up to 35%) reduction in the wolves’ prey base 6 years before wolf hunting was stopped, (2) weak enforcement of the protection law, resulting in lack of poaching control of wolves, (3) probable increasing fragmentation and isolation of wolf habitat caused by rapid economic growth in Poland. Inconsistent application of current management policy toward wolves resulted in weak enforcement of regulations and promoted negative attitudes toward the species. To improve the status of wolves in Poland, I recommend a flexible wolf management planning framework that involves and addresses attitudes of hunters and sheep herders, includes a framework to promote strong law enforcement, and consistent, fair compensation for livestock killed by wolves.     

Camera-trap studies of maned wolf density in the Cerrado and the Pantanal of Brazil

The maned wolf (Chrysocyon brachyurus) is threatened by large-scale habitat loss, in particular due to conversion to agricultural land. This is the first published study on maned wolf density and the first test of individual identification from camera-trap photographs. We present results from two Brazilian regions: the Cerrado and the Pantanal. Using capture–recapture analysis of camera-trap data, we estimated densities per 100 square kilometers of 3.64 ± 0.77 individuals at the Cerrado site and 1.56 ± 0.77 individuals at the Pantanal site. Parallel radio-telemetry studies at the Pantanal site showed that maned wolves occupied home ranges of 39–58 km² (mean = 50.3 ± 7.67 km²). Our study in the Cerrado took place in a private farm with a mixture of agricultural land and native habitats, representative of the majority of the present-day Cerrado. Whereas many other mammalian species have suffered in the region, our results show that the maned wolf may cope better with this highly fragmented landscape than one might have feared. Finally, the paper briefly compares maned wolf density with density of puma (Puma concolor) in the Pantanal site.     

Biogeography and conservation biology of Florida's Geolycosa wolf spiders: threatened spiders in endangered ecosystems

The state of Florida has a complex biogeography which is the product of past sea level changes and associated ocean currents creating numerous inland sand ridge systems which are home to a unique biota adapted to the seasonally dry conditions found on these sandy uplands. Among these xeric-adapted taxa is a suite of wolf spider species in the genus Geolycosa. Geolycosa wolf spiders are habitat specialists and poor dispersers, and many taxa have ranges restricted to specific ridge systems. We found that the six Geolycosa species typical of Florida's scrubs and sandhills (G. escambiensis Wallace, G. ornatipes (Bryant), G. micanopy Wallace, G. patellonigra Wallace, G. hubbelli Wallace and G. xera McCrone) have current distributions that correlate with the historical biogeography of the state. There are species typical of the Lake Wales Ridge and associated ridges (G. xera and G. hubbelli), species found on the more coastal ridges (G. patellonigra and G. micanopy) and species whose ranges are divided by the Apalachicola River drainage (G. escambiensis and G. ornatipes). We also found that Geolycosa wolf spiders tend to occur in species pairs which use different microhabitats: those which burrow in areas covered in leaf litter, and those which use only barren sites. These latter species require habitat management plans that include burns, that maintain the habitat in an open condition.     

Size‐assortative choice and mate availability influences hybridization between red wolves (Canis rufus) and coyotes (Canis latrans)

Anthropogenic hybridization of historically isolated taxa has become a primary conservation challenge for many imperiled species. Indeed, hybridization between red wolves (Canis rufus) and coyotes (Canis latrans) poses a significant challenge to red wolf recovery. We considered seven hypotheses to assess factors influencing hybridization between red wolves and coyotes via pair‐bonding between the two species. Because long‐term monogamy and defense of all‐purpose territories are core characteristics of both species, mate choice has long‐term consequences. Therefore, red wolves may choose similar‐sized mates to acquire partners that behave similarly to themselves in the use of space and diet. We observed multiple factors influencing breeding pair formation by red wolves and found that most wolves paired with similar‐sized conspecifics and wolves that formed congeneric pairs with nonwolves (coyotes and hybrids) were mostly female wolves, the smaller of the two sexes. Additionally, we observed that lower red wolf abundance relative to nonwolves and the absence of helpers increased the probability that wolves consorted with nonwolves. However, successful pairings between red wolves and nonwolves were associated with wolves that maintained small home ranges. Behaviors associated with territoriality are energetically demanding and behaviors (e.g., aggressive interactions, foraging, and space use) involved in maintaining territories are influenced by body size. Consequently, we propose the hypothesis that size disparities between consorting red wolves and coyotes influence positive assortative mating and may represent a reproductive barrier between the two species. We offer that it may be possible to maintain wild populations of red wolves in the presence of coyotes if management strategies increase red wolf abundance on the landscape by mitigating key threats, such as human‐caused mortality and hybridization with coyotes. Increasing red wolf abundance would likely restore selection pressures that increase mean body and home‐range sizes of red wolves and decrease hybridization rates via reduced occurrence of congeneric pairs.     

The distribution of <Emphasis Type="Italic">Echinococcus granulosus</Emphasis> in moose: evidence for parasite-induced vulnerability to predation by wolves?

The role of parasites in influencing the trophic dynamics of hosts is becoming increasingly recognized in the ecological literature. Echinococcus granulosus is a tapeworm that relies on the predator-prey relationship between the definitive host (wolf, Canis lupus) and the intermediate host, (moose, Alces alces) to complete its life cycle. Heavy infection by E. granulosus may predispose moose to increased risk of predation by wolves. Theory predicts that parasite-induced vulnerability to predation will reduce the degree of aggregation of parasites in a host population. We tested for different levels of aggregation of E. granulosus in moose in areas of low, moderate, and high levels of wolf predation using Greens coefficient of dispersion. Parasite aggregation was lower in an area with high predation rate, thus we hypothesize that heavy infection by E. granulosus predisposes moose to predation by wolves. This increase in predation rate due to parasite infection may influence the role of wolves in regulating moose populations. We discuss alternative explanations for the negative correlation between predation rate and parasite aggregation.An erratum to this article can be found at     

Maned Wolf Density in a Central Brazilian Grassland Reserve

ABSTRACT The maned wolf (Chrysocyon brachyurus) inhabits the savannah-like Cerrado of Brazil. Although 80% of this biome has already been affected by human activity, little is known about maned wolf abundance. Using mark—recapture models, we obtained the first density estimate from central Brazil, the core of maned wolf distribution. With 5.19 individuals/100 km², even large reserves support only small maned wolf populations. Therefore, long-term conservation of the maned wolf depends on land management outside of reserves.     

Spatial interactions between grey wolves and Eurasian lynx in Białowieża Primeval Forest, Poland

Various species of large predators are reported to influence each other through interference or exploitation competition that may affect demography and survival of the subordinate species. We analyzed spatial relationships between grey wolf (Canis lupus) and Eurasian lynx (Lynx lynx) in Białowieża Primeval Forest (BPF, eastern Poland) to determine how they partitioned the space. The wolves (n = 8) and lynx (n = 14) were radio-tracked in 1991–1999. Three wolves and seven lynx were radio-tracked simultaneously in 1994–1996. Territories of wolf packs and home ranges of lynx overlapped considerably (76% of wolf territories and 50% of lynx home ranges, on average). In three cases, their core areas were also overlapping. Wolf-lynx dyads with overlapping home ranges were simultaneously located at distances from 0 to 28 km from each other. We found neither avoidance nor attraction between wolves and lynx occupying the same areas. We concluded that in BPF, the two large predators coexist due to specialization on different preferred prey and heterogeneous habitat.     

Genetic and morphometric analysis of sixteenth century <Emphasis Type="Italic">Canis</Emphasis> skull fragments: implications for historic eastern and gray wolf distribution in North America

Resolving the taxonomy and historic ranges of species are essential to recovery plans for species at risk and conservation programs that aim to restore extirpated populations. In eastern North America, planning for wolf population restoration is complicated by the disputed historic distributions of two wolf species: the Old World-evolved gray wolf (Canis lupus) and the New World-evolved eastern wolf (C. lycaon). We used genetic and morphometric data from 4- to 500-year-old Canis samples excavated in London, Ontario, Canada to help clarify the historic range of these two wolf species in the eastern temperate forests of North America. We isolated DNA and sequenced the mitochondrial control region and found that none of the samples were of gray wolf origin. Two of the DNA sequences corresponded to those found in present day coyotes (C. latrans), but morphometric comparisons show an eastern wolf, not coyote, origin. The remaining two sequences matched ancient domestic dog haplotypes. These results suggest that the New World-evolved eastern wolf, not the gray wolf, occupied this region prior to the arrival of European settlers, although eastern-gray wolf hybrids cannot be ruled out. Furthermore, our data support the idea of a shared common ancestry between eastern wolves and western coyotes, and that the distribution of gray wolves at this time probably did not include the eastern temperate forests of North America.     

Species-Specificity of Chemical Signals: Silk Source Affects Discrimination in a Wolf Spider (Araneae: Lycosidae)

Female spiders deposit chemical cues that elicit male courtship behavior with silk. These cues are often assumed to be species-specific although male spiders may court in response to chemical cues of closely-related species. We used behavioral assays to test the extent of species discrimination of female chemical cues by male Schizocosa ocreata, a wolf spider (Lycosidae). Discrimination, expressed as relative courtship intensity of males, varied significantly with phylogenetic distance. Males did not discriminate between female cues of conspecifics and a sibling species, S. rovneri. Courtship response was intermediate for another species within the ocreata clade and not different from control for spiders outside the clade. These findings support the sibling species status of S. ocreata and S. rovneri, and also suggest the composition of female chemical signals is conserved across closely related wolf spider species.     

Analysis of <Emphasis Type="Italic">Canis</Emphasis> mitochondrial DNA demonstrates high concordance between the control region and ATPase genes

Background  

Phylogenetic studies of wild Canis species have relied heavily on the mitochondrial DNA control region (mtDNA CR) to infer species relationships and evolutionary lineages. Previous analyses of the CR provided evidence for a North American evolved eastern wolf (C. lycaon), that is more closely related to red wolves (C. rufus) and coyotes (C. latrans) than grey wolves (C. lupus). Eastern wolf origins, however, continue to be questioned. Therefore, we analyzed mtDNA from 89 wolves and coyotes across North America and Eurasia at 347 base pairs (bp) of the CR and 1067 bp that included the ATPase6 and ATPase8 genes. Phylogenies and divergence estimates were used to clarify the evolutionary history of eastern wolves, and regional comparisons of nonsynonomous to synonomous substitutions (dN/dS) at the ATPase6 and ATPase8 genes were used to elucidate the potential role of selection in shaping mtDNA geographic distribution.     

Prey selection and dietary response by wolves in a high-density multi-species ungulate community

Studies on predation by the wolf (Canis lupus) have often reported contradictory results about the role of prey density and vulnerability on wolf prey use. We investigated dietary response and prey selection by wolves in a high-density and multi-species ungulate community, analysing scats collected over a period of 11 years in the Casentinesi Forests, Italy. The second most abundant species, wild boar (Sus scrofa), was found to be the main wolf prey, and we did not observe any dietary response of wolves to variations in the density of either primary or secondary prey species. Selection patterns were uniform throughout the study period. Wolves strongly selected for wild boar piglets, while roe deer (Capreolus capreolus) fawns and adults, red deer (Cervus elaphus) adults and fallow deer (Dama dama) adults were avoided. Wolf preference for wild boar was inversely density dependent. Within each species, juveniles were preferred to adults. Medium-sized, young individuals of both wild boar and roe deer were optimal prey, although with different selection patterns related to the different anti-predator strategies adopted by each prey species. The results of this study suggest that in productive ecosystems with high density and high renewal rates of prey, selection patterns by wolves are determined by prey vulnerability, which is connected to prey age and body size. The different patterns of wild boar versus cervids use by wolf across Europe seems to be related to their relative abundances, while the strong selection of wild boar in Italian Apennines with respect to the more frequent avoidance in central-eastern Europe is better explained by higher piglet productivity and smaller body size of adults boar in Mediterranean temperate forests.     

Large‐scale patterns of seed removal by small mammals differ between areas of low‐ versus high‐wolf occupancy

Because most tree species recruit from seeds, seed predation by small‐mammal granivores may be important for determining plant distribution and regeneration in forests. Despite the importance of seed predation, large‐scale patterns of small‐mammal granivory are often highly variable and thus difficult to predict. We hypothesize distributions of apex predators can create large‐scale variation in the distribution and abundance of mesopredators that consume small mammals, creating predictable areas of high and low granivory. For example, because gray wolf (Canis lupus) territories are characterized by relatively less use by coyotes (C. latrans) and greater use by foxes (Vulpes vulpes, Urocyon cinereoargentus) that consume a greater proportion of small mammals, wolf territories may be areas of reduced small‐mammal granivory. Using large‐scale, multiyear field trials at 22 sites with high‐ and low‐wolf occupancy in northern Wisconsin, we evaluated whether removal of seeds of four tree species was lower in wolf territories. Consistent with the hypothesized consequences of wolf occupancy, seed removal of three species was more than 25% lower in high‐wolf‐occupancy areas across 2 years and small‐mammal abundance was more than 40% lower in high‐wolf areas during one of two study years. These significant results, in conjunction with evidence of seed consumption in situ and the absence of significant habitat differences between high‐ and low‐wolf areas, suggest that top‐down effects of wolves on small‐mammal granivory and seed survival may occur. Understanding how interactions among carnivores create spatial patterns in interactions among lower trophic levels may allow for more accurate predictions of large‐scale patterns in seed survival and forest composition.