The vocal repertoires of the bush dog (Speothos venaticus), crab-eating fox (Cerdocyon thous), and maned wolf (Chrysocyon brachyurus)

The vocalizations of the bush dog (Speothos venaticus), crab-eating fox (Cerdocyon thous), and maned wolf (Chrysocyon brachyurus) were studied in a captive setting. The three species' repertoires were similar and included whines, long-distance calls, and growls. The whines and growls varied in amplitude, duration, and repetition rate; these variations and the frequencies of occurrence probably were influenced by both social organization and habitat characteristics. The bush dog, which lives in family groups and inhabits rain forests, had an elaborate whine-scream continuum that at close distances conveyed a wide range of moods. On the other hand, the maned wolf, which is solitary and frequents open pampas, used amplitude variations of the growl to signal hostile intentions over short, medium, and long distances.The long-distance vocalizations of the three species were the least variable in structure; the bush dog and crab-eating fox used the call to promote contact with separated pairmates or family members. Based on recordings during development, these two calls appeared to arise from the repetitive whine, a variant of the basic whine syllable. The maned wolf also had a contact-promoting howl, apparently an extended whine of high amplitude, but it was rarely recorded during the present study. More prominent was the maned wolf roar-bark, which functioned to space individuals over large distances, and appeared to be a growl of high amplitude.     

Detection of the maned wolf,a cryptic and vulnerable species,through environmental DNA studies

The maned wolf (Chrysocyon brachyurus) is a vulnerable and ubiquitous species endemic to South America. Their populations are declining due to habitat loss as a consequence of agricultural expansion and exposure to anthropic threats such as road-kills, illegal hunting and diseases. In Argentina, there is a lack of information about its accurate distribution because its populations are sparse and the specimens are very difficult to observe due to its cryptic and lonely behavior, low population density, and type of habitat. In this article, we describe the results of the application of the environmental DNA methodology to carry out indirect monitoring of maned wolf specimens. The data obtained show that this technique is effective in detecting specimens of this mammal, which was verified through direct monitoring. This information represents an adjustment of the methods that can be applied in large-scale monitoring to study the structure and distribution of the maned wolf populations, as well as to obtain genetic data about this species, to design optimal conservation plans to preserve it in Argentina.     

Predicting occurrence of wolf territories in Scandinavia

We used logistic regression to compare a set of habitat features inside known Scandinavian wolf Canis lupus territories with the same habitat features in areas outside known territories, but still close enough to be available for wolf colonization. In addition, we analysed changes in habitat variables over time within wolf territories. Wolf territories had lower densities of roads, built-up areas and open land than areas outside wolf territories, but there was no difference in the density of the wolves' main prey, elk Alces alces . The logistic regression model classified 79% of Scandinavia outside the reindeer husbandry area as suitable wolf habitat, that is with a probability of wolf occurrence >0.5. The proportion of built-up areas within the wolf territory decreased as the 'borders' of the wolf territory changed over time. Our model had a reasonably high predictive power, with correct classification in 90% (18 of 20) of the observed wolf territories in the study area. Polygons, randomly distributed outside the observed wolf territories, were correctly classified as not being occupied by wolves in 85% of the cases (17 of 20). This allows a more effective use of resources to, for example, prevent wolf depredation on livestock and dogs.     

Traditional Versus Multivariate Methods for Identifying Jaguar,Puma, and Large Canid Tracks

ABSTRACT The jaguar (Panthera onca) and puma (Puma concolor) are the largest felids of the American Continent and live in sympatry along most of their distribution. Their tracks are frequently used for research and management purposes, but tracks are difficult to distinguish from each other and can be confused with those of big canids. We used tracks from pumas, jaguars, large dogs, and maned wolves (Chrysocyon brachyurus) to evaluate traditional qualitative and quantitative identification methods and to elaborate multivariate methods to differentiate big canids versus big felids and puma versus jaguar tracks (n = 167 tracks from 18 zoos). We tested accuracy of qualitative classification through an identification exercise with field-experienced volunteers. Qualitative methods were useful but there was high variability in accuracy of track identification. Most of the traditional quantitative methods showed an elevated percentage of misclassified tracks (≥20%). We used stepwise discriminant function analysis to develop 3 discriminant models: 1 for big canid versus big felid track identification and 2 alternative models for jaguar versus puma track differentiation using 1) best discriminant variables, and 2) size-independent variables. These models had high classification performance, with <10% of error in the validation procedures. We used simpler discriminant models in the elaboration of identification keys to facilitate track classification process. We developed an accurate method for track identification, capable of distinguishing between big felids (puma and jaguar) and large canids (dog and maned wolf) tracks and between jaguar and puma tracks. Application of our method will allow a more reliable use of tracks in puma and jaguar research and it will help managers using tracks as indicators of these felids' presence for conservation or management purposes.     

Investigating the distribution of prairie dogs in an urban landscape

Habitat fragmentation is a prevalent threat to biological diversity, and urbanization is a primary agent of fragmentation and a leading cause of species endangerment. Landscape biogeographic and local habitat characteristics can be important determinants of the distribution of species in habitat patches in urban landscapes. However, the specifics of which characteristics are most critical to maintaining biological diversity are not fully known for prairie ecosystems, especially in fragmented urban habitat. This study focuses on black-tailed prairie dogs along an urban gradient in Denver, CO. Prairie dogs have declined precipitously throughout the region and are an essential part of the prairie ecosystem, making them excellent study subjects. We identified a series of habitat fragments along a gradient of urbanization in the fully urbanized areas and south suburbs of Denver, CO, both containing and not containing prairie dogs. Local characteristics, including fragment slope and vegetative cover, and landscape characteristics, including fragment size, age and connectivity, were measured on each fragment. We used likelihood-based methods to explore which variables most accurately predicted prairie dog occurrence within our study area. Multiple factors influenced the distribution of prairie dogs in urban settings, with colony connectivity the strongest predictor of occupancy. Large and recently isolated fragments near other prairie dog colonies, flat areas and those with high graminoid cover were most likely to support prairie dog populations. Our study provides the first attempt to model prairie dog occurrence in highly fragmented urban habitat and has important implications for the management and conservation of prairie dogs.     

Genetic variability in maned wolf based on heterologous short-tandem repeat markers from domestic dog

The maned wolf (Chrysocyon brachyurus) is the largest South American canid. Habitat loss and fragmentation, due to agricultural expansion and predatory hunting, are the main threats to this species. It is included in the official list of threatened wildlife species in Brazil, and is also protected by IUCN and CITES. Highly variable genetic markers such as microsatellites have the potential to resolve genetic relationships at all levels of the population structure (among individuals, demes or metapopulations) and also to identify the evolutionary unit for strategies for the conservation of the species. Tests were carried out to verify whether a class of highly polymorphic tetranucleotide repeats described for the domestic dog effectively amplifies DNA in the maned wolf. All five loci studied were amplified; however, one of these, was shown to be monomorphic in 69 maned wolf samples. The average allele number and estimated heterozygosity per polymorphic locus were 4.3 and 67%, respectively. The genetic variability found for this species, which is considered threatened with extinction, showed similar results when compared to studies of other canids.     

Bucking the Trend in Wolf-Dog Hybridization: First Evidence from Europe of Hybridization between Female Dogs and Male Wolves

Studies on hybridization have proved critical for understanding key evolutionary processes such as speciation and adaptation. However, from the perspective of conservation, hybridization poses a concern, as it can threaten the integrity and fitness of many wild species, including canids. As a result of habitat fragmentation and extensive hunting pressure, gray wolf (Canis lupus) populations have declined dramatically in Europe and elsewhere during recent centuries. Small and fragmented populations have persisted, but often only in the presence of large numbers of dogs, which increase the potential for hybridization and introgression to deleteriously affect wolf populations. Here, we demonstrate hybridization between wolf and dog populations in Estonia and Latvia, and the role of both genders in the hybridization process, using combined analysis of maternal, paternal and biparental genetic markers. Eight animals exhibiting unusual external characteristics for wolves - six from Estonia and two from Latvia - proved to be wolf-dog hybrids. However, one of the hybridization events was extraordinary. Previous field observations and genetic studies have indicated that mating between wolves and dogs is sexually asymmetrical, occurring predominantly between female wolves and male dogs. While this was also the case among the Estonian hybrids, our data revealed the existence of dog mitochondrial genomes in the Latvian hybrids and, together with Y chromosome and autosomal microsatellite data, thus provided the first evidence from Europe of mating between male wolves and female dogs. We discuss patterns of sexual asymmetry in wolf-dog hybridization.     

Resource selection and its implications for wide-ranging mammals of the brazilian cerrado

Conserving animals beyond protected areas is critical because even the largest reserves may be too small to maintain viable populations for many wide-ranging species. Identification of landscape features that will promote persistence of a diverse array of species is a high priority, particularly, for protected areas that reside in regions of otherwise extensive habitat loss. This is the case for Emas National Park, a small but important protected area located in the Brazilian Cerrado, the world's most biologically diverse savanna. Emas Park is a large-mammal global conservation priority area but is too small to protect wide-ranging mammals for the long-term and conserving these populations will depend on the landscape surrounding the park. We employed novel, noninvasive methods to determine the relative importance of resources found within the park, as well as identify landscape features that promote persistence of wide-ranging mammals outside reserve borders. We used scat detection dogs to survey for five large mammals of conservation concern: giant armadillo (Priodontes maximus), giant anteater (Myrmecophaga tridactyla), maned wolf (Chrysocyon brachyurus), jaguar (Panthera onca), and puma (Puma concolor). We estimated resource selection probability functions for each species from 1,572 scat locations and 434 giant armadillo burrow locations. Results indicate that giant armadillos and jaguars are highly selective of natural habitats, which makes both species sensitive to landscape change from agricultural development. Due to the high amount of such development outside of the Emas Park boundary, the park provides rare resource conditions that are particularly important for these two species. We also reveal that both woodland and forest vegetation remnants enable use of the agricultural landscape as a whole for maned wolves, pumas, and giant anteaters. We identify those features and their landscape compositions that should be prioritized for conservation, arguing that a multi-faceted approach is required to protect these species.     

Prey availability and diet of maned wolf in Serra da Canastra National Park,southeastern Brazil

Analysis of 400 scats of the maned wolfChrysocyon brachyurus (Illiger, 1815), collected in Serra da Canastra National Park from November 1998 to January 2000 yielded 1688 occurrences of 69 identified food items, 49.5% of which were represented by plant items and 50.5% by animal items. According to the frequency of occurrence analysis, the most important food item categories were wolf’s fruit,Parinari obtusifolia (Chrysobalanaceae), small mammals and birds. When the estimated biomass consumption is considered, the animal items became more important (56.8%) than plant items (43.2%). Small mammals were consumed more in the dry season, while miscellaneous fruits and arthropods were consumed mostly in the wet season. Functional response was found only for the category miscellaneous fruits; however, seasonality analysis revealed that wolf’s fruit and small mammals were consumed according to their availability in the area. Prey selection was observed for most of the small mammal species, emphasizing a high level of consumption ofNecromys lasiurus, Monodelphis sp. andGracilinanus sp. The finding that most of the diet was composed of fruits and animal items from open areas of the savannah-like Cerrado led us to suggest that the maned wolf spends most of its time preying in this habitat, indicating a need to establish conservation policies for the Cerrado.     

The Distributional Ecology of the Maned Sloth: Environmental Influences on Its Distribution and Gaps in Knowledge

The maned sloth Bradypus torquatus (Pilosa, Bradypodidae) is endemic to a small area in the Atlantic Forest of coastal Brazil. It has been listed as a threatened species because of its restricted geographic range, habitat loss and fragmentation, and declining populations. The major objectives of this study were to estimate its potential geographic distribution, the climatic conditions across its distributional range, and to identify suitable areas and potential species strongholds. We developed a model of habitat suitability for the maned sloth using two methods, Maxent and Mahalanobis Distance, based on 42 occurrence points. We evaluated environmental variable importance and the predictive ability of the generated distribution models. Our results suggest that the species distribution could be strongly influenced by environmental factors, mainly temperature seasonality. The modeled distribution of the maned sloth included known areas of occurrence in the Atlantic Forest (Sergipe, Bahia, Espírito Santo, and Rio de Janeiro), but did not match the observed distributional gaps in northern Rio de Janeiro, northern Espírito Santo or southern Bahia. Rather, the model showed that these areas are climatically suitable for the maned sloth, and thus suggests that factors other than climate might be responsible for the absence of species. Suitable areas for maned sloth were located mainly in the mountainous region of central Rio de Janeiro throughout Espírito Santo and to the coastal region of southern Bahia. We indicate 17 stronghold areas and recommended survey areas for the maned sloth. In addition, we highlight specific areas for conservation, including the current network protected areas. Our results can be applied for novel surveys and discovery of unknown populations, and help the selection of priority areas for management and conservation planning, especially of rare and relatively cryptic species directed associated with forested habitats.     

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.     

Transit time and digestibility of two experimental diets in the maned wolf (Chrysocyon brachyurus) and domestic dog (Canis lupus)

Recent research involving nutritional management of clinical disease secondary to cystinuria in captive maned wolves (Chrysocyon brachyurus) in the United States resulted in the production of a commercially manufactured maintenance diet and subsequently, a further modified experimental diet. These two diets differed only in their protein sources and sodium concentrations. The purpose of this study was to further investigate these diets by determining their digestibility and transit time in maned wolves, and also to compare these results to the digestibility and transit time of the diets in domestic dogs. The experiment consisted of two diets (maned wolf commercial maintenance diet and experimental diet) and two species (maned wolf and domestic dog [beagle]) arranged as a 2 × 2 factorial design. For the transit time study, titanium dioxide (TiO₂, 5 g/kg diet) was administered as a pulse dose and total collection of feces was carried out for 50 hr. There was no effect of diet or species on initial Ti recovery time (IRT), the amount of Ti recovered at IRT, or the time to reach 50% of Ti excretion in the feces. Maned wolves had lower total percent recovery (P<0.01) of Ti than dogs and shorter time to last recovery of Ti in the feces (P<0.05), however, marker excretion in the maned wolves may not have been complete. For the digestibility study, in which chromic oxide was used as a marker, fecal samples were collected on 2 consecutive days immediately after 12 days of being fed the diets and then analyzed for dry matter, energy, protein, and minerals. Crude protein, calcium, phosphorus, and zinc apparent digestibilities were unaffected by species or diet (P>0.05). Apparent digestibility of dry matter (P<0.05) and energy (P<0.01) were higher in the experimental diet and in the dogs. Both dogs and maned wolves fed the maintenance diet exhibited a greater apparent retention of copper, iron, and magnesium (P<0.01). Dogs had a higher apparent retention of magnesium and sodium than the maned wolves. Zoo Biol 0:1–13, 2006. © 2006 Wiley‐Liss, Inc.     

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.     

Symptomatic Raccoon Dogs and Sarcoptic Mange Along an Urban Gradient

We quantitatively evaluated the effects of landscape factors on the distribution of symptomatic raccoon dogs with sarcoptic mange along an urban gradient. We used 246 camera traps (182 traps from April 2005 to December 2006; 64 traps from September 2009 to October 2010) to record the occurrence of asymptomatic and symptomatic raccoon dogs at 21 survey sites along an urban–rural gradient in the Tama Hills area of Tokyo. Each occurrence was explained in terms of the surrounding forest, agricultural, and grassland areas and additional factors (i.e., seasonal variations and survey methods) at various spatial scales using a generalized additive mixed model (GAMM). In our analysis, a 1000-m radius was identified as the important spatial scale for asymptomatic and symptomatic raccoon dog occurrence. The peak of the predicted occurrence probability of asymptomatic raccoon dogs appeared in the intermediate forest landscape as opposed to non-forest and forest landscapes. However, a high occurrence probability of symptomatic raccoon dogs was detected in non-forest and intermediate forest landscapes (i.e., urban and suburban) as opposed to a forest landscape, presumably because of animals occurring at much higher densities in more urbanized areas. Therefore, our results suggest that human-modified landscapes play an important role in the high occurrence of sarcoptic mange in raccoon dogs.     

Diet of the maned wolf, Chrysocyon brachyurus, in central Brazil

Analysis of 105 scats of the maned wolf ( Chrysocyon brachyurus ) collected in central Brazil yielded 304 occurrences of food items of which fruits of Solanum lycocarpum , rodents, and birds accounted for 61.5%. By analysis of minimum number of individual animals, rodents and birds were 75% of a total of 156 prey. On the other hand, fruits of S. lycocarpum and armadillos ( Dasypus spp.) were the bulk of the total estimated biomass consumption, yielding 63.7% in a total of 73.5 kg. Rodents were mostly captured during the dry season, while miscellaneous fruits were consumed mostly in the wet season. The consumption of S. lycocarpum fruits and armadillos was aseasonal. Small rodents were taken in about the ranks of abundance in the study area, but S. lycocarpum fruits were actively searched by maned wolves, for its occurrence is limited to secondary savanna. Prey and fruits typical of savanna ('cerrado') and grassland ('campo') were the bulk of species, occurrences, prey numbers, and biomass consumed. These findings reinforce the importance of conservation of cerrado and campo in central Brazil. Despite maned wolf being a vulnerable species, its feeding habits are opportunistic, and the main reasons for its decline are likely to be habitat destruction and human disturbance.     

Polymorphism of transferrin and esterase in Alaskan wolves: evidence of close molecular homology with the dog

By the use of isoelectric focusing in polyacrylamide gels serum samples from 146 Alaskan wolves were studied with regard to transferrin (Tf) and esterase (ArE) polymorphism, comparing the phenotypic band patterns with those of selected Norwegian dogs. The study revealed Tf and ArE polymorphisms in the wolf with phenotypic band patterns being indistinguishable from the corresponding ones in dogs. This suggests the occurrence of the same two common Tf alleles in the wolf as in the dog. In the ArE system the results are consistent with the occurrence of three alleles which also occur in dogs whereas a fourth allele, so far not seen in dogs, is seen in Alaskan wolves.     

Bear-Baiting May Exacerbate Wolf-Hunting Dog Conflict

Background

The influence of policy on the incidence of human-wildlife conflict can be complex and not entirely anticipated. Policies for managing bear hunter success and depredation on hunting dogs by wolves represent an important case because with increasing wolves, depredations are expected to increase. This case is challenging because compensation for wolf depredation on hunting dogs as compared to livestock is less common and more likely to be opposed. Therefore, actions that minimize the likelihood of such conflicts are a conservation need.

Methodology/Principal Findings

We used data from two US states with similar wolf populations but markedly different wolf/hunting dog depredation patterns to examine the influence of bear hunting regulations, bear hunter to wolf ratios, hunter method, and hunter effort on wolf depredation trends. Results indicated that the ratio of bear hunting permits sold per wolf, and hunter method are important factors affecting wolf depredation trends in the Upper Great Lakes region, but strong differences exist between Michigan and Wisconsin related in part to the timing and duration of bear-baiting (i.e., free feeding). The probability that a wolf depredated a bear-hunting dog increases with the duration of bear-baiting, resulting in a relative risk of depredation 2.12–7.22× greater in Wisconsin than Michigan. The net effect of compensation for hunting dog depredation in Wisconsin may also contribute to the difference between states.

Conclusions/Significance

These results identified a potential tradeoff between bear hunting success and wolf/bear-hunting dog conflict. These results indicate that management options to minimize conflict exist, such as adjusting baiting regulations. If reducing depredations is an important goal, this analysis indicates that actions aside from (or in addition to) reducing wolf abundance might achieve that goal. This study also stresses the need to better understand the relationship among baiting practices, the effect of compensation on hunter behavior, and depredation occurrence.     

Effects of habitat fragmentation and black-tailed prairie dogs on urban avian diversity

Urbanization and habitat fragmentation have the potential to influence bird communities. In addition, these phenomena, as well as ongoing lethal control measures, have also greatly reduced the range of the black-tailed prairie dog (Cynomys ludovicianus) since the beginning of the 20th century. Although prairie dogs are highly interactive species that can influence avian communities, few studies have investigated whether these interactions persist in urban settings. Our goal was to investigate the relative impacts of habitat fragmentation and prairie dogs on bird communities within an urban matrix. We performed bird surveys on 20 habitat fragments (10 colonized by prairie dogs, 10 uncolonized by prairie dogs) distributed throughout the Denver metropolitan area, and calculated Shannon–Weiner diversity and richness of all birds and native species, as well as total counts of grassland birds and raptors. Diversity, richness, and counts of many species increased with increasing fragment connectivity, and decreased on fragments isolated for longer periods of time. Avian diversity and richness did not differ between fragments with and without prairie dogs, suggesting that this element of the ecological role of prairie dogs is not fully retained in urban habitat. Future studies of the role of prairie dogs as keystone species in urban systems should include other taxa as well as consider the influence of the urban matrix surrounding prairie dog habitat. Our results emphasize that conservation of urban avian diversity should focus on landscape connectivity as well as local habitat features.     

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.     

Food composition and food niche overlap of three kinds of canidae

The diets of the wolf Canis lupus, red fox Vulpes vulpes and raccoon dog Nyctereutes ussurienusis Matschie were studied in the grassland of eastern Inner Mongolia from August to October in 2007 and from March to July in 2008. Feces were collected from dens and latrines. The frequency of occurrence and the dry weight of the remains of each food item in the feces were calculated. Livestock (50.65% frequency of occurrence) constituted the bulk of the wolf diet, while small rodents (76.31% frequency of occurrence) were most frequently eaten by the foxes. Small rodents (22.69% frequency of occurrence) were important to the raccoon dogs but they also frequently consumed birds (39.81% frequency of occurrence) and insects (26.39% frequency of occurrence). The wolf was the most omnivorous and the red fox was the least omnivorous of these three kinds of canidae according to the diversity index. The food niches of the wolf and the red fox overlapped more than that of the raccoon dog and other species. These three kinds of canidae shared many resources and consequently competition occurred to them. However, their diets also differed to some extent which helped them to avoid competition. Furthermore, raccoon dogs were dormant in winter when food was scarce, which may be the reason why all these species could coexist in a rather unproductive grassland.