Clues to the cause of the Tsushima leopard cat (Prionailurus bengalensis euptilura) decline from isotopic measurements in three species of Carnivora

The estimated population of the Tsushima leopard cat Prionailurus bengalensis euptilura is only 80–110 individuals. However, the cause of the population decline is not clear. We investigated temporal changes in the food habits of the cat and two other species of Carnivora (marten and weasel) inhabiting the Tsushima Islands by measuring δ¹³C and δ¹⁵N values in hair samples. Hair samples of the cat were collected not only from specimens and furs, but also from feces. The gathering of hair from cat feces was most efficient when the feces were collected in the spring. The food habit of male cats seemed to be more diverse and tended to comprise prey of higher trophic levels than the food habits of the females. The δ¹³C and δ¹⁵N measurements suggested that the trophic level of the food sources has been decreasing over the last several decades for the cat and weasel, but not for the marten. Increased consumption of prey from lower trophic levels in the food habit of the cat seems to be related to the decline of the cat population because these phenomena occurred simultaneously.     

Fecal DNA analysis for identifying species and sex of sympatric carnivores: a noninvasive method for conservation on the Tsushima Islands, Japan

Fecal analysis is a useful tool for the investigation of food habits and species identity in mammals. However, it is generally difficult to identify the species based on the morphological features and contents of feces deposited by mammals of similar body size. Therefore we developed noninvasive DNA analysis methods using fecal samples for identification of the species and sex of four small sympatric carnivores living on the Tsushima Islands of Japan: the leopard cat (Felis bengalensis), Japanese marten (Martes melampus), Siberian weasel (Mustela sibirica), and feral cat (Felis catus). Based on DNA sequence data from previous phylogenetic studies, we designed species-specific primers for polymerase chain reaction (PCR) amplification of the partial mitochondrial cytochrome b gene (112-347 bp) to identify the species and primers for the partial SRY gene (135 bp) to determine the sex. Due to the adjustment of PCR conditions, those specific DNA fragments were successfully amplified and then applied for species and sex identification. Nucleotide sequences obtained from the PCR products corresponded with cytochrome b sequences of the carnivore species expected. The protocol developed could be a valuable tool in the management and conservation of the four carnivore species occurring on the Tsushima Islands.     

Predation on rodents in Białowieza primeval forest, Poland

Autumn and winter predation on bank vole Clethrtonomys glareolus and yellow-necked mouse Apodemus flavicollis was studied in 1985/86-88/89 in an 11 2 km² area of the Bialowieza National Park Rodents regularly increased in numbers from spring to autumn and decreased throughout winter Out of 23 species of predators, the most common were tawny owl Strix aluco (43-57 adult ind 10 km⁻²) weasel Mustela ntvahs (17-27 ind), buzzard Buteo buteo (12-16) and pine marten Martes martes (5-8) Voles and mice were the staple food for two specialists the stoat Mustela erminea and the weasel, and two generahsts the tawny owl and the pine marten The generalists exploited different alternative prey when rodents were scarce tawny owl -amphibians, marten - small mammals, and the red fox Vulpes vulpes - ungulate carcasses and hares The depth of snow and abundance of voles were two major factors shaping the contribution of voles to tawny owl and marten diets No such relationships were found for mice and generalist predators The predation by 8 species (tawny owl, buzzard, marten, weasel, stoat, polecat Mustela putortus fox, and raccoon dog Nyctereutes procyonoides) from 1 October to 15 April in 1986/87, 87/88, and 88/89 was estimated to be on average 28-35 voles and 14-17 mice ha Estimates suggested that three species were responsible for 86-95% of the total predation impact tawny owl (56-71%), weasel (11-21%) and marten (10-15%) The predation impact was similar to the annual decrease in rodent numbers from autumn till spring voles 35 (SD 111) lnds ha^-1in autumn and 8 (SD 2 6) lnds ha^-1in spring, and mice 24 (SD 16 3) mds ha⁻¹ in autumn and 3 (SD 2 5) in spring Predation, therefore, was regarded as the main agent of rodent mortality throughout autumn and winter     

Genetic diversity in domestic cats Felis catus of the Tsushima Islands, based on mitochondrial DNA cytochrome b and control region nucleotide sequences

Nucleotide sequences of mitochondrial DNA (mtDNA) of 50 domestic cats (Felis catus) obtained from the Tsushima Islands were determined and the genetic diversity was analyzed. In the cats, six haplotypes of the complete cytochrome b sequences (1,140 base-pairs, bp) and ten haplotypes of the partial control region sequences (350 bp) were identified. Haplotypes obtained from both genes showed existence of at least 11 maternal lineages of domestic cats in Tsushima. Mean values of polymorphic site numbers and sequences differences in the control region were 2.4 times and 1.8 times higher than those in the cytochrome b gene, respectively. Our results support the idea that the evolutionary rate of the control region was faster than that of the cytochrome b as reported in other mammals. Molecular phylogenetic trees showed the similar clustering of haplotypes for both genes. Meanwhile, no individual variations within the Tsushima leopard cat (Felis bengalensis euptilura), which is native to Tsushima, were observed, possibly as a result of genetic drift in the small ancestral population by geographical isolation. In contrast, the diversity of the domestic cat population was higher than that of the leopard cats, because the genetic variability of the former's founders, which were repeatedly brought to Tsushima in the past, still remains. In addition, no sequences of the leopard cat mtDNA were detected in any domestic cats. However, because the possibility that the domestic cat would crossbreed with the leopard cat cannot be denied, genetic monitoring of two species is necessary to biological conservation in Tsushima.     

红外相机技术在北京雾灵山自然保护区兽类资源调查中的应用

为了调查北京市雾灵山自然保护区兽类资源情况,自2013年7月至2014年5月,在雾灵山32个位点处布放红外相机进行系统调查,累积相机工作日3630天,共拍摄到兽类有效照片494张,鉴定出10种动物,包括：狍（Capreolus capreolus）、斑羚（Nemorhaedus goral）、猪獾（Arctonyx collaris）、岩松鼠（Sciurotamias davidianus）、野猪（Sus scrofa）、狗獾（Meles meles）、豹猫（Felis bengalensis）、花鼠（Tamias sibiricus）、草兔（Lepus capensis）和黄鼬（Mustela sibirica）,隶属于4目7科;从分布型来看,这10种兽类主要以古北型为主（占70%）。常拍种有狍、岩松鼠、斑羚和猪獾;较常拍种有野猪、狗獾、豹猫、花鼠、草兔;偶拍种有黄鼬。文中还对不同海拔和不同植被类型动物相对丰富度、累积拍摄物种数与相机工作日关系及兽类G-F指数等进行了探讨。     

Fecal steroid metabolites and reproductive monitoring in a female Tsushima leopard cat (Prionailurus bengalensis euptilurus)

Although the Tsushima leopard cat (Prionailurus bengalensis euptilurus) is one of the most endangered mammals in Japan, its reproductive physiology and endocrinology have been not elucidated. The objective was to establish the non-invasive monitoring of reproductive endocrinology in a female Tsushima leopard cat and to identify the types of fecal reproductive steroid metabolites in this species. Fecal concentrations of estrogen and progestin were determined by enzyme immunoassays, from 60 d before to 60 d after the last copulation, during three pregnancies. Fecal estrogen metabolite concentrations were increased before/around the mating period and after mid-pregnancy. Fecal progestin metabolite concentrations increased after the last copulation and remained high during pregnancy. The gestation period was 65.0 ± 0.6 d (mean ± SD). Fecal extracts were separated by high-performance liquid chromatography for identification of fecal metabolites. Fecal estrogens were identified as estradiol-17β and estrone. Fecal progestins during pregnancy contained 5α-reduced pregnanes: 5α-pregnan-3α-ol-20-one, 5α-pregnan-3β-ol-20-one and 5α-pregnan-3,20-dione, and nonmetabolized progesterone was barely detected in feces. In conclusion, measurement of fecal estrogen and progestin metabolites was effective for noninvasive reproductive monitoring in the Tsushima leopard cat. An immunoassay for fecal estradiol-17β concentrations seemed useful to monitor follicular activity, whereas an immunoassay with high cross reactivity for 5α-reduced pregnanes was useful to monitor ovarian luteal activity and pregnancy.     

Endozoochorous seed dispersal by sympatric mustelids, Martes melampus and Mustela itatsi, in western Tokyo, central Japan

We investigated seed dispersal by two sympatric mustelid species, the Japanese marten (Martes melampus) and Japanese weasel (Mustela itatsi), along an intercity forest path in western Tokyo, central Japan, from Jul 2007 to Jul 2008. We aimed to investigate the effect of food/habitat preference of these mustelids (martens are semi-arboreal frugivores while weasels are terrestrial carnivores) on their seed dispersal characteristics, which determine their efficacy as seed dispersers. In total, we analyzed 478 fecal samples collected from the two mustelids (N_marten = 381, N_weasel = 97). The proportions of feces containing seeds for martens and weasels were 81.4% and 55.7%, respectively. The number of plant species whose seeds were found within the feces were 28 and 17, respectively. Almost all seeds within feces of both mustelids were intact. The number of plant species whose seeds were found within a single fecal sample ranged from one to four, but no significant difference was detected between the two mustelids. However, marten feces contained a significantly greater number of seeds of most plant species as well as total number of seeds than did weasel feces. The numbers of plant species and seeds represented in marten feces varied seasonally, but those represented in weasel feces did not. Our findings suggest the possibility that both mustelids act in some ways as seed dispersers, although martens seem to disperse a greater diversity and total amount of seeds.     

Lure sticks as a method to detect pine martens <Emphasis Type="Italic">Martes martes</Emphasis>

Marten species are usually surveyed by trapping, snow tracking or cameratrapping with baits on trees. While testing the efficiency of a monitoring scheme for wildcats Felis silvestris silvestris in north-western Switzerland, we noticed that martens are attracted by lure sticks scented with valerian. On these sticks, the animals leaved some hairs that allowed us to identify the genus Martes by microscopic analysis. Additionally, the animal can be identified on pictures made by a camera trap posed close to the lure stick. In this paper, we compared the efficiency of different methods to find the most appropriate one in order to survey the pine marten Martes martes (Linnaeus, 1758) in Switzerland. For the method of valerian lure sticks we estimated a detectability of 0.08 per 14 days during the whole year. The detectability raised, when we applied American Hawbaker’s marten lure instead of the valerian tincture. In addition, the detectability was higher during the period April to June (p=0.2) compared to the whole year. If we identified the pine marten on the lure stick with pictures from the camera traps we reached a detectability of 0.21 during the whole year. Using only camera traps with baits on trees we could not take any picture of a pine marten.     

Interspecies Transmission of Feline Immunodeficiency Virus from the Domestic Cat to the Tsushima Cat (Felis bengalensis euptilura) in the Wild

Feline immunodeficiency virus (FIV) was isolated from a wild-caught Tsushima cat (Felis bengalensis euptilura), an endangered Japanese nondomestic subspecies of leopard cat (F. bengalensis). Phylogenetic analysis of the env gene sequences indicated that the FIV from the Tsushima cat belonged to a cluster of subtype D FIVs from domestic cats. FIVs from both the Tsushima cat and the domestic cat showed similar levels of replication and cytopathicity in lymphoid cell lines derived from these two species. The results indicated the occurrence of interspecies transmission of FIV from the domestic cat to the Tsushima cat in the wild.     

Selection of waterbirds by Geoffroy's cat: Effects of prey abundance, size, and distance

In spite of its wide distribution, Geoffroy's cat (Oncifelis geoffroyi) has been recently upgraded from Least Concern IUCN category to Near Threatened because of lack of knowledge. In this study, we examine at the Mar Chiquita Biosphere Reserve (37° 46`S, 57° 27`W; Argentina) their seasonal diet, and their feeding preference in relation to waterbird abundance, size, and distance of waterbird prey before attack. The diet, evaluated by feces analysis, was comprised of 42% waterbirds and 40% small mammals. The relative importance of waterbirds in the diet changed throughout the year associated with seasonal fluctuations in their availability. Predation on waterbirds increased from autumn to summer, while predation on small mammals decreased during this period. Trophic-niche breadth for prey occurrence was higher in autumn and lower in spring. We found that abundance, distance of waterbird prey before attack, and prey size were significant predictors of waterbird consumption by the Geoffroy's cat. This small cat appears to change its kill rate in relation to waterbird abundance changes. The overconsumption of prey close to vegetation and big prey indicates selective predatory behavior by this cat.     

内蒙古赛罕乌拉自然保护区4 种小型食肉目动物的食性构成的初步分析

为了解同域分布动物的食物竞争关系,采用样线法收集粪便样品,依据残遗物鉴别物种的方法,于2007年8 ～12 月对赛罕乌拉国家级自然保护区的赤狐、狗獾、香鼬和豹猫的食物构成进行比较。结果表明:赤狐取食所有15 类食物,并捕食小型鼬类;在赤狐粪样(n =70) 中出现率较大的植物(32.6% )以重量比率计算时其贡献极大下降(0.02% ),而出现率较低的草兔和鸟类重量比率较高(草兔65.4% , 鸟类22.9% ), 相对重要值的计算表示,赤狐的食物构成均一。狗獾(n = 64)以取食昆虫和植物果实为主(出现率29.3% 和57.8% )。香鼬(n =14)集中取食鼠类(出现率73.7% ,相对重量99.8% ),豹猫(n = 11) 则以鸟类居多(出现率27.8% ,相对重量69.0% )。食物生态位的宽度以赤狐最大(0.2),其次为豹猫(0.07)。根据食物生态位的重
叠度分析,赤狐与狗獾(O = 0.99)和豹猫(O = 0.7)以及狗獾与豹猫(O = 0.6)的种间竞争较强, 但在主要食物上出现分离,实现捕食者之间的共存。赤狐对香鼬的捕食可能限制了其种群的发展。     

Food habits of genetically identified pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>) expanding in agricultural lowlands (NW Italy)

We assessed the diet of pine marten (Martes martes) expanding in the heavily human-altered agricultural plain of the River Po, northern Italy. Between February 2008 and November 2009, surveys were carried out twice a week during seasonal sampling sessions of three to four consecutive weeks. To distinguish the faecal samples from those of sympatric carnivores, a polymerase chain reaction-restriction fragment length polymorphism method was applied. The availability of small mammals was assessed by the analysis of 59 barn owl (Tyto alba) pellets. A total of 109 pine marten faeces were analysed. Its diet consisted of fruit, rodents, lagomorphs and birds. Seasonal variation occurred for fruit, which prevailed in summer, and rodents, which were more preyed upon in autumn. In winter, the diet of the pine marten was almost totally based on vertebrates, and lagomorphs were the main source of protein in summer. Use of small mammals differed significantly from their availability, voles, particularly bank vole Myodes glareolus, being preferred to mice (Apodemus sp.) and rats (Rattus sp.). Medium-size mammals formed about 18% of pine marten diet, a value generally reported for high-latitude habitats. In agricultural areas, the pine marten proved to be an opportunistic predator, able to face the reduced availability of small mammals by preying upon medium-size prey and fruit.     

Multi-annual variation in the diet composition and frugivory of the Japanese marten (Martes melampus) in western Tokyo,central Japan

To examine multi-annual variations in the food habits of the Japanese marten (Martes melampus), we analyzed the composition of marten feces in the Bonbori Forest Path in western Tokyo, central Japan, in two time periods a decade apart (1997–1998 and 2007–2008). The staple foods of martens in both periods were fruits/seeds and animal materials (mainly insects and mammals). The martens fed frequently on fruits/seeds and insects throughout the year in both periods, but the consumption of mammals, birds, and arthropods/other animals showed seasonal variations. The composition of fruits/seeds and the frequency of occurrence for each fruit-bearing species differed between the two periods. These results suggest that both the foraging strategy and role of martens as a seed dispersal agent changes yearly, presumably according to multi-annual variation in the availability of prey animals and/or fruits. We emphasize the importance of multi-annual studies both on food habits and to monitor food availability in the temperate region where the food environment changes among seasons, as well as years.     

Carnivores (Mammalia: Carnivora) in South China: a status review with notes on the commercial trade

• 1 Evidence from the literature, interviews, market surveys, general rapid field surveys and camera trapping was reviewed to infer the regional status of 33 recorded carnivores in South China (Guangxi, Guangdong, Hainan, Hong Kong and Macau).
• 2 The carnivore fauna in South China is among the most depleted for any continental area in the world.
• 3 The tiger Panthera tigris, leopard Panthera pardus, grey wolf Canis lupus and binturong Arctictis binturong are probably extirpated.
• 4 The dhole Cuon alpinus, Asiatic golden cat Catopuma temmincki and clouded leopard Neofelis nebulosa are at great risk of regional extirpation, as are the more ecologically adaptable red fox Vulpes vulpes, raccoon dog Nyctereutes procyonoides, Asian badger Meles leucurus, Eurasian otter Lutra lutra, Oriental small‐clawed otter Aonyx cinereus and large Indian civet Viverra zibetha.
• 5 Though more widely reported, the Asiatic black bear Ursus thibetanus, yellow‐throated marten Martes flavigula, hog badger Arctonyx collaris and Siberian weasel Mustela sibirica are at regional risk.
• 6 The status of the Burmese ferret‐badger Melogale personata, stripe‐backed weasel Mustela strigidorsa, smooth‐coated otter Lutrogale perspicillata, large‐spotted civet Viverra megaspila and Owston's palm civet Chrotogale owstoni is uncertain due to inadequate information.
• 7 Five carnivore species – the Chinese ferret‐badger Melogale moschata, masked palm civet Paguma larvata, yellow‐bellied weasel Mustela kathiah, small Indian civet Viverricula indica and leopard cat Prionailurus bengalensis– are relatively secure (though not common) in the region, along with the Asian palm civet Paradoxurus hermaphroditus and the Javan mongoose Herpestes javanicus in some areas, while the spotted linsang Prionodon pardicolor and crab‐eating mongoose Herpestes urva may have been under‐reported.
• 8 The main threats have been habitat destruction and unsustainable exploitation. Only in Hong Kong, where enforcement of protection legislation has been stronger, are the surviving carnivore species easily encountered.
• 9 Improved enforcement and monitoring are essential to retaining the remainder of South China's carnivores. Support could be boosted by conservation education and better research.

     

Marten and fisher responses to fluctuations in prey populations and mast crops in the northern hardwood forest

Many forest tree species produce seed (mast) crops that are consumed by a variety of wildlife species and these pulsed resources may mediate interactions among predator and prey populations. In the northern hardwood forests of New York, we investigated interactions among mast production, prey abundance, and harvests of American martens (Martes americana) and fishers (Martes pennanti) during 1988–2009. Mast production for beech (Fagus grandifolia), sugar maple (Acer saccharum), and mountain ash (Sorbus americana) was synchronous and an alternate-year pattern in production was evident for most of the time series. We documented considerable temporal variation in summer small mammal relative abundance and our numerical response models received substantial support for 5 of the 8 species, indicating lagged responses to autumn mast crops. Trap response of martens to the autumn production of beech mast and mountain ash berries was immediate and numerical responses to the relative abundance of small mammal prey occurred during the preceding summer. The age structure of the marten harvest differed based on the dominant alternate-year pattern of summer prey relative abundance and autumn mast production (χ²₄ = 33.06, P < 0.001). The proportion of juvenile marten in the autumn harvest was 52% and 34% following summers when small mammal relative abundance was high and low, respectively and these differences resulted in a persistent cohort effect that was apparent until age 3.5. Trap response of fishers to the autumn production of beech mast was immediate and numerical responses to the relative abundance of Sciurid prey occurred during the preceding summer. Marten and fisher harvests fluctuated similarly among New York, Maine, and New Brunswick, which may indicate regional synchronization of mast crops and responses of martens and fishers to similar prey dynamics. A better understanding of how food availability influences demographic responses and trapping vulnerability of martens and fishers would aid our ability to manage harvests of these species on a sustained yield basis. © 2011 The Wildlife Society.     

Fasting in the American marten (<Emphasis Type="Italic">Martes americana</Emphasis>): a physiological model of the adaptations of a lean-bodied animal

The American marten (Martes americana) is a boreal forest marten with low body adiposity throughout the year. The aim of this study was to investigate the adaptations of this lean-bodied species to fasting for an ecologically relevant duration (48 h) by exposing eight farm-bred animals to total food deprivation with seven control animals. Selected morphological and hematological parameters, plasma and serum biochemistry, endocrinological variables and liver and white adipose tissue (WAT) enzyme activities were determined. After 48 h without food, the marten were within phase II of fasting with depleted liver and muscle glycogen stores, but with active lipid mobilization indicated by the high lipase activities in several WAT depots. The plasma ghrelin concentrations were higher due to food deprivation, possibly increasing appetite and enhancing foraging behavior. The lower plasma insulin and higher cortisol concentrations could mediate augmented lipolysis and the lower triiodothyronine levels could suppress the metabolic rate. Fasting did not affect the plasma levels of stress-associated catecholamines or variables indicating tissue damage. In general, the adaptations to short-term fasting exhibited some differences compared to the related farm-bred American mink (Mustela vison), an example of which was the better ability of the marten to hydrolyze lipids despite its significantly lower initial fat mass.     

Habitat characteristics of Eurasian pine martens Martes martes in an insular Mediterranean environment

Eurasian pine martens are considered habitat specialists, associated primarily with mature stands of mesic mixed wood forest habitats, and avoid areas without overhead cover The species is found throughout the temperate and boreal regions of the continent but on the Mediterranean island of Minorca, introduced pine martens thrive in a competitor- and predator-free environment I test the prediction that because of evolved prey-capture and predator avoidance strategies Minorcan martens should select habitats most similar to temperate and northern parts of their range Scat index routes were used to quantify pine marten habitat selection Marten did not demonstrate any habitat type preferences although observed use of pine forests and coastal shrublands was slighly greater than expected Marten were indifferent to overhead cover whereas mesic sites and areas of tall high shrub density were favored Small mammal trap indices and preferred prey suggested that martens commonly used non-forested areas My results demonstrated that on Minorca pine martens were habitat generalists In the absence of predators open non-forested habitats were equally important to pine marten as were forested ones     

Consumption of Domestic Cat in Madagascar: Frequency,Purpose, and Health Implications

ABSTRACT

The domestic cat Felis catus has a long history of interaction with humans, and is found throughout the world as a household pet and a feral animal. Despite people's often sentimental association with cats, cat meat is sometimes consumed by them; this practice can have important implications for public health. In Madagascar, a least developed country that has experienced recent political instability, cat consumption is known to occur, but remains poorly understood. To improve our understanding of cat consumption practices in Madagascar we interviewed 512 respondents in five towns. We used semi-structured interviews to: 1) clarify the preference for, and prevalence, correlates, and timing of, cat consumption; 2) describe methods used to procure cats for consumption; 3) identify motives for consuming cat meat; and, 4) evaluate to what extent patterns of cat-meat consumption are influenced by taboos. We found that, although cat was not a preferred source of meat, many (34%) Malagasy respondents had consumed cat meat before, with most (54%) of these indicating such consumption occurred in the last decade. We did not detect a link between consumption of cat meat and recent access to meat (a proxy for food security). Cat meat was almost never purchased, but rather was obtained when the owners consumed their own pet cat, as a gift, or by hunting feral cats. Cat meat was usually consumed in smaller towns following cat–human conflict such as attacks on chickens, but in the large capital city, cat meat was procured primarily from road-killed individuals. These results suggest cat-meat consumption is typically an opportunistic means to obtain inexpensive meat, rather than principally serving as a response to economic hardship. These results further suggest cat handling and consumption may present a potential pathway for transmission of several diseases, including toxoplasmosis, that may warrant heightened public health efforts.     

Spring and summer food habits and habitat use of the Europena pine marten (Martes martes) on the island of Micorca, Spain

The food habits and habitat use of pine marten ( Martes martes L.) on the Balearic Island of Mincorca were studied from March-August 1990. In a 28.2 km² area, a series of hiking trails/forest roads were used to collect pine marten faeces bi-monthly and study te species' differential use of habitats. A total of 28 different food items were identified in 723 faeces. Small mammals were the most important food overall, constituting 34% of the volume During March-April, small mammals were the principal food consumed (63% of volume), followed by birds (19%). In May-June, birds were the main food (40%), then small mammals. Plant material and insects were the most important foods in July-August, both made up 68% of the pine marten diet. The abundance of certain foods and the degree of difficulty in obtaining them appear to be important factors that govern pine marten food selection in Minorca. Habitat use was studied along 12 routes totalling 32.6 km. Pine marten showed preference in March-April for Coastal Shrubland habitat, while during May-June they were found most in the Upland Pine type. During July-August, the Upland Pine and Open Pine habitats were most frequented by pine marten. According to bimonthly food habits data, habitat use seems largely determined by the abundance and availability of their prey items in the available habitat types. Martens may concentrate hunting efforts in areas different from when they are not hunting. The Minorcan pine marten differs from continental populations in its wide use of shrublands, in addition to forests.