Panthera Tigris Altaica Research Articles

Serosurvey of Free-ranging Amur Tigers in the Russian Far East

Wild Amur tigers (Panthera tigris altaica, n=44) from the Russian Far East were tested for antibodies to feline leukemia virus, feline corona virus (FCoV), feline immunodeficiency virus, feline parvovirus (FPV), canine distemper virus (CDV), Toxoplasma gondii, and Bartonella henselae. Antibodies to FCoV, CDV, FPV, and T. gondii were detected in 43, 15, 68, and 42% of tigers, respectively. No differences were detected in antibody prevalence estimates between tigers captured as part of a research program and those captured to mitigate human-tiger conflicts. Domestic dogs (Canis familiaris) were tested as a potential source for CDV; 16% were vaccinated against CDV and 58% of unvaccinated dogs were antibody positive for CDV. A high percentage of tigers were exposed to potential pathogens that could affect the survival of this species. We recommend continued monitoring of wild tigers throughout Asia, development of standardized sampling and postmortem examination procedures, and additional research to better understand potential domestic and wild animal sources for these pathogens.

中国野生东北虎数量监测方法有效性评估

Information on abundance and variation in abundance are bases for the effective wildlife management.Accurate survey methods are necessary to monitor the status and dynamics of endangered population of Amur tiger.The tigers are notoriously hard to monitor because they are secretive,occur at low density,wander widely,and are very rarely observed.Suitable monitoring methods vary in their reliability depending on the monitoring objectives,the spatial and temporal scale,tiger population density,prey density,climate,terrain and other environmental parameters. The accuracy and feasibility of three methods(Information collecting networks,traditional transect survey and tiger-prey biomass relationships) for monitoring Amur tiger population abundance were assessed in the eastern Wanda Mountains,Heilongjiang Province and 8 bordering forest area between the southern Laoye Mountains in Heilongjiang Province and the northern Dalong Mountains in Jilin province during 2002—2011.The results showed that: 1) there were at least 6—9 wild Amur tiger in the eastern Wanda Mountains(1 male,2—3 adult females,2—4 sub-adults and 1 cub),in 2006 by using an information network for tigers;2) there were 22—27 wild tigers in the eastern Wanda Mountains in 2002—2004 based on the prey biomass relationship method,which obviously exaggerated the tiger population size;and 3) 0 tigers in 8 bordering forestry areas between the southern Laoye mountains in Heilongjiang Province and the northern Dalong Mountains in Jilin province,in 2011 by using traditional transect method,which underestimated the true tiger population size. The reasons for exaggeration of the tiger population using the biomass method could be previous losses of tigers from the area due to snares and competition with another carnivore,especially with people for ungulates.The transect method may have underestimated tiger densities in the survey areas because it was based on little prior knowledge of tiger behavior.It could only be usefully applied when tigers exist in at least moderate densities(i.e.,when there is a high probability of encountering tiger tracks along a suite of routes).Although there is inherent potential error and bias,such as the unclear relationships of expert estimates and the true density,and between tiger track densities and actual tiger densities,same as the transect method,the monitoring of tiger populations using information networks provides a record of minimum tiger presence,and may be an appropriate approach when tiger presence is extremely rare,transitory and unstable,such as in northeast China.This approach is economically efficient and should be further improved by established a wider network across the landscape to encompass all potential tiger habitat using better trained monitoring staff. An occupancy survey would be an ideal method to monitor tigers at the landscape scale when tigers are determined to be permanently present in some areas of northeastern China.Camera traps,DNA identification,dog identification,track identification,monitoring methodology based on capture-recapture systems may only work when tiger densities are high enough to provide ample capture-recapture data.They may be appropriate for future use in China.The density overestimates as determined by the large ungulate biomass method indicated that tiger numbers might be stabilized if strict measures were taken to alleviate the factors threating both the tigers and their prey.We should also monitor populations of large ungulates and threat factors to those ungulates while at the same time make efforts to monitor abundance trends in the tiger population itself.

Application of Fuzzy Closeness in Siberian Tiger Skin Texture Recognition

In order to improve the accuracy of tiger skin texture recognition, a skin texture identification method based on fuzzy closeness is put forward in this paper: four skin texture stripes extracted from a tiger texture image are selected as a research object. The image treatment process is: firstly, connected their endpoints to form three quadrilaterals, then regarded the quadrilateral area, bottom length, and left bottom corner features as the characteristics index of the object, the fuzzy closeness degree derived from the characteristics is compared with the standard models, and the final closeness degree is obtained. Experimental results show that this method can improve the accuracy of tiger skin texture recognition, and identify the tiger skin texture effectively to achieve individual identification of tigers finally.

Application of satellite collars to the study of home range and activity of the Amur tiger (Panthera tigris altaica)

The possibility of application of satellite collars to the study of home range and activity of Amur tigers has been analyzed. The possibility of obtaining information about the size and structure of a home range, discerning the home range core areas, seasonal changes in the use of space by tigers, and collecting detailed data on animals’ activity and characteristics of the use of different elements of landscape and terrain has been demonstrated. In contrast to VHF transmitters, satellite collars allow tigers to be tracked even in the cases of very long travels.

Population viability of the Siberian Tiger in a changing landscape: Going, going and gone?

The Amur tiger ( Panthera tigris altaica) is a flagship species of the boreal forest ecosystem in northeastern China and Russia Far East. During the past century, the tiger population has declined sharply from more than 3000 to fewer than 600 individuals, and its habitat has become much smaller and greatly fragmented. Poaching, habitat degradation, habitat loss, and habitat fragmentation have been widely recognized as the primary causes for the observed population decline. Using a population viability analysis tool (RAMAS/GIS), we simulated the effects of poaching, habitat degradation, habitat loss, and habitat fragmentation on the population dynamics and extinction risk of the Amur tiger, and then explored the relative effectiveness of three conservation strategies involving improving habitat quality and establishing movement corridors in China and Russia. A series of controlled simulation experiments were performed based on the current spatial distribution of habitat and field-observed vital rates. Our results showed that the Amur tiger population could be viable for the next 100 years if the current habitat area and quality were well-maintained, with poaching strictly prohibited of the tigers and their main prey species. Poaching and habitat degradation (mainly prey scarcity) had the largest negative impacts on the tiger population persistence. While the effect of habitat loss was also substantial, habitat fragmentation per se had less influence on the long-term fate of the tiger population. However, to sustain the subpopulations in both Russia and China would take much greater conservation efforts. The viability of the Chinese population of tigers would rely heavily on its connectivity with the largest patch on the other side of the border. Improving the habitat quality of small patches only or increasing habitat connectivity through movement corridors alone would not be enough to guarantee the long-term population persistence of the Amur tiger in both Russia and China. The only conservation strategy that allowed for long-term persistence of tigers in both countries required both the improvement of habitat quality and the establishment of a transnational reserve network. Our study provides new insights into the metapopulation dynamics and persistence of the Amur tiger, which should be useful in landscape and conservation planning for protecting the biggest cat species in the world.

Complete sequence determination of hemoglobin from endangered feline species using a combined ESI-MS and X-ray crystallography approach

We present the first complete amino acid sequences of Snow leopard (Uncia uncia) and Amur tiger (Panthera tigris altaica) hemoglobin (Hb) determined using a combined ESI MS and X-ray crystallography approach. Using mass spectrometry alone, we were able to achieve 81%, 90%, 87% and 72% sequence coverage for the α-, and β-globins of Snow leopard Hb and the α-, and β-chain of Amur tiger Hb, respectively. The incomplete mass spectrometry-determined sequences were used to process X-ray diffraction data of Snow leopard and Amur tiger Hb crystals solved to 2.1Å and 1.8Å, respectively. Using tandem mass spectrometry in concert with the electron density maps from X-ray structures, we were able to not only achieve 100% coverage of all amino acid sequences, but to also distinguish between isomeric (leucine and isoleucine) and isobaric (glutamine and lysine) residues. These results demonstrate the synergy between mass spectrometry and X-ray crystallography for the determination of full amino acid sequence coverage of abundant proteins.

The Aumr Tiger’s Individual Identification Based on the Tiger Fur’s Texture Characteristic

In order to identify the Amur tiger individual more accurately and conveniently, a new characteristic of Amur tiger is proposed in this paper. The images of the Amur tiger fur’s texture were first processed by computer through the method of image processing technology. After that it was easier to extract the characteristic from the image. In the Amur tiger’s sideways image, the eigenvalue is the ratio of the specified texture’s area in tiger sideways and standard area. The specified texture’s area is formed with the black stripes whose shape is similar to diamond. Standard area is a triangle area and its vertices are the spine’s highest point, abdomen’s lowest point and caudal vertebra’s central point. Experiments show that the characteristic is a unique characteristic of the Amur tiger individual and it can be used to identify the Amur tiger individual.

Activity of the hypothalamo-pituitary-adrenals axis in the Siberian tiger (Panthera tigris altaica) in captivity and in the wild, and its dynamics throughout the year

A noninvasive evaluation method of hypothalamo-pituitary-adrenals axis (HPA) activity in the Siberian tiger was verified. Comparison of the activity level of HPA in Siberian tigers in the wild and in captivity, and their alterations over the year was carried out. Significant seasonal deviations between activity levels of HPA in tigers in captivity were not found. In the wild, this level was significantly higher, reaching the maximum from November to January, which can be related with an unfavorable influence on tigers in low temperatures and deep snow cover.

A non-invasive identification of hormone metabolites, gonadal event and reproductive status of captive female tigers

Putranto HD (2011) A non-invasive identification of hormone metabolites, gonadal event and reproductive status of captive female tigers. Biodiversitas 12: 131-135. As a non-invasive method, fecal sample provides some advantage for animal and collector. The purpose of the present study were to monitor the reproductive status of female Siberian tigers (Panthera tigris altaica) by assessing changes in fecal during natural ovarian activity and pregnancy and to identify whether progesterone (P4) exists and what kinds of P4 metabolites excreted into the feces. Two female tigers were fed a diet consisting of meat. Drinking water was available ad libitum. Feces were collected ones to twice a week. The fecal contents of P4 and estradiol-17β (E2) were determined by EIA and P4 metabolites were separated by a modified HPLC. The EIA results shown that during its natural ovarian activity the E2 contents showed cyclic changes at the average of 27.0 d interval, however, no distinct cycles were shown in fecal P4 contents of non-pregnant tiger. In contrary, the fecal P4 contents in pregnant tiger increased remarkably after copulation approximately 2- to 6-fold higher than the mean value. The HPLC results indicated that two peaks were primarily detected fraction 63- 64 min (identified metabolites) and fraction 85 min (not identified metabolite) in feces of pregnant tiger. However, P4 detected only small amount in feces. It is possible to assess non-invasively gonadalevents such as luteal or follicular activity or ovulation of Siberian tigers by endocrine monitoring based on fecal P4 and E2 to understand reproductive status.

Comparative cat studies: Are tigers auditory specialists?

Although representatives of the 41 extant cat species inhabit nearly every biome on the planet and may have faced a highly diverse set of selection pressures during their evolution, relatively little effort has been made to compare commonly measured features of peripheral auditory function among species representing the family. Given their extensive geographic range, it is reasonable to suggest that auditory system adaptations may have occurred, leading to functional specialization in a subset of species. In that light, frequency-threshold curves and response latency-level and latency-frequency relationships will be compared in cats of widely varying body mass inhabiting a variety of habitats. The body mass of felids spans a range of more than two orders of magnitude, with small cats like the desert sandcat, Felis margarita, weighing as little as 2 kg and the Amur tiger, Panthera tigris altaica, weighing as much as 300 kg. While most cats studied thus far appear to satisfy the conditions necessary to be labeled auditory generalists, the tiger, and perhaps members of the Panthera genus generally, may be exceptions. We will consider the possibility that big cats are auditory generalists with regard to acoustic sensitivity, but exhibit a peripheral specialization affecting low-frequency neural latencies. [Funding provided by NSF Grant No. 0823417].

Siberian tiger's recent population bottleneck in the Russian Far East revealed by microsatellite markers

Because size reduced, or bottlenecked, populations are more prone to adverse events, the detection of genetic bottleneck signatures in wildlife species is highly relevant for conservation. Here we applied 11 microsatellite markers to the endangered Siberian tiger ( Panthera tigris altaica) using tissue and blood samples of animals from the Primorsky region of the Russian Far East. Excess heterozygosity and mode shift in allele frequencies tests were positive, while the M-ratio test was negative, indicating the likelihood of a contemporary rather than a historical population bottleneck. The recent genetic population bottleneck could be attributed to the well-documented demographic collapse of the Siberian tiger population in the 1940s, when population size hit bottom with 20–30 surviving animals. The mean effective population size N e was 14 Siberian tigers (CI 95: 12–25 animals), and the effective population size/census size ratio ( N e / N ratio) was 0.028. This is the first molecular evidence of a recent Siberian tiger population bottleneck, which is of great interest for further conservation and management plans of the highly endangered largest felid species, while the worryingly low effective population size challenges the optimism for the recovery of the huge Siberian cat.

Canine distemper in Siberian tiger cubs from Zagreb ZOO: case report

Canine distemper is a contagious, potentially lethal disease of mainly domestic and wild canids, but also of many other mammalian species including large felids. In February 2004, two Siberian tiger (Panthera tigris altaica) cubs at the age of six months died at the Zagreb ZOO. The animals were presented for necropsy with two days history of severe digestive disorders, characterized mainly by haemathemesis. Dissections revealed catarrhal to pseudomembranous gastroenteritis (depending on the animal) accompanied with haemorrhagic oedema of the lungs. Necrotic tonsillitis and disseminated depletion of the lymphocytes were the most prominent histological findings in both examined animals, while intranuclear and intracytoplasmatic inclusion bodies were found in the samples of the tongues and intestines. Representative portions of the livers, intestines, tonsils and lymph nodes were submitted for bacteriological and mycological analysis. The presence ofClostridiumspp.,Campylobacter coliandEscherichia coliwas detected in gut samples, coli-like bacteria were found in samples of liver, tonsils and lymph nodes, whileCandidasp. was found in the gut and pharynx samples. Toxicological analysis excluded anticoagulant and organophosphorous intoxication as the cause of death. Immunohistochemical analysis was positive for canine distemper virus. Based on all this, epizootiological, clinical and additional findings, canine distemper was recognized as the cause of the observed condition in these animals.

The Once and Future Tiger

In reeds tinged red by the Central Asian sun, a tiger once roamed. There, in riparian forests that line rivers like the Vakhsh on the border of the former Soviet country Tajikistan, the Caspian tiger (Panthera tigris virgata) prowled. Until its extinction in the mid1900s, the tiger stalked Bukhara deer along tugai, thicketed watercourses that flow through Central Asia’s otherwise vast, arid deserts. The tiger’s route took it across shaded rivers and onto muddy banks. Peering into shoreline bracken, the tiger crawled through a latticework of tangled low shrubs, finally emerging into the willow and poplar forests favored by its deer prey. And into trouble. In the early 20th century, the Russian government instructed its army to exterminate all tigers as part of an agricultural conversion project across Central Asia. Once Caspian tigers were almost gone, farmers moved in, clearing tugai and planting crops such as cotton. The tigers retreated, first from lowland streams to marshes around larger rivers. Finally, they crept into forested mountain ranges. The Caspian tiger’s last stronghold was a reserve in Tajikistan: Tigrovaya Balka, “tiger former river channel.” The name was given to this first of Central Asia’s protected areas, or In 1947, Russia banned hunting of the Caspian tiger’s close relative, the Siberian, or more properly Amur, tiger (Panthera tigris altaica), still found today near the Amur River in the Russian Far East. But the edict came too late for the Caspian tiger, says Carlos Driscoll, a biologist at the US National Cancer Institute’s Laboratory of Genomic Diversity in Frederick, Maryland. The last tiger in Tigrovaya Balka was glimpsed in 1958. Although still a matter of debate, the legendary final wild Caspian tiger may have been killed in February 1970, in Hakkari Province, Turkey. New DNA evidence, however, reveals that we may not have seen the last of this tugai tiger.

Conflicts between Amur (Siberian) tigers and humans in the Russian Far East

In 1999, the Russian Federation created a Tiger Response Team (TRT) to investigate and intervene in human–tiger conflicts. We examined data collected on human-Amur tiger ( Panthera tigris altaica) conflicts from January 2000 through February 2009 to: (1) summarize and characterize human–tiger conflicts in the area, (2) examine causes of human–tiger conflicts, and (3) attempt to evaluate the effectiveness of the TRT. The team investigated 202 conflicts. Both attacks on humans and depredations were greatest in winter ( X 2 = 9, df = 3, P = 0.03 and X 2 = 64, df = 3, P < 0.001, respectively). Tiger depredation on domestic animals was the most common type of conflict reported (57%), followed by tigers near human habitations (22%), miscellaneous conflicts (12%), and attacks on humans (9%). Dogs were killed more commonly than other domestic animals (63% of 254 animals), likely because livestock were well managed. Nineteen attacks on humans were recorded resulting in 11 injuries and 2 deaths. Nearly four tigers per year ( n = 32 tigers) died, were killed, or were removed from the wild, and all but 1 of 20 tigers killed or removed from the wild by the TRT were considered unfit to survive in the wild. Attacks on humans ( n = 19) were most often (77%) by wounded tigers (80% of injuries were human-caused) and commonly provoked (47% of attacks). The effectiveness of interventions focused on reducing depredation on domestic animals was unclear, but data suggested that removal of injured and other unhealthy tigers from the wild by the Tiger Response Team resulted in fewer human deaths. Our recommendations include that the TRT continues to work to reduce conflict by rapidly removing debilitated tigers from the wild, explores different methods and technologies for reducing depredation on domestic animals, and increases their efforts to maintain tigers in the wild through telemetry monitoring, translocation, and rehabilitation of orphaned cubs. Further, standardized data should be collected to evaluate all interventions, with information from evaluation guiding an adaptive management component of their human–tiger conflict mitigation activities. This process should occur across tiger landscapes in Asia to allow rapid assessment of interventions.

Clinical Course of a Malignant Peripheral Nerve Sheath Tumor in a Siberian Tiger (Panthera Tigris Altaica)

A 14-year-old male Siberian tiger (Panthera tigris altaica) was admitted with an ulcerating mass on the right thoracic wall. Radiographic and computed tomographic evaluation indicated 2 isolated cutaneous masses without any signs of metastasis. Histology of a Tru-Cut biopsy revealed an anaplastic sarcoma with giant cells. Both tumors were resected with appropriate normal tissue margins. The size of the defect did not allow primary closure of the wound; therefore, a mesh expansion technique was attempted. Three months later, the tiger had to be euthanized due to extensive metastasis to the lungs. Histomorphological features and immunohistochemical results confirmed the diagnosis of malignant peripheral nerve sheath tumor. In contrast to domestic animal experience, the tumor had spread extensively to the lungs without local reccurrence in a short period of time. Correct diagnosis requires various immunohistochemical evaluations of the tumor tissue.

Morbillivirus Infection in a Wild Siberian Tiger in the Russian Far East

We report the first documented case of morbillivirus infection in a wild, free-ranging Siberian tiger (Panthera tigris altaica). The tigress entered a small village in the Russian Far East in an ambulatory but stuporous state with no apparent recognition or fear of humans. Her condition progressed rapidly with neurological signs, anorexia, and ultimately death. Histologic lesions included vacuolated to malacic white matter in the brain stem, cerebellum, and thalamus, with associated lymphocytic meningoencephalitis. Large, intranuclear, eosinophilic inclusions were within regional astrocytes, and the brain lesions were immunohistochemically positive when stained for canine distemper viral antigen. Hematologic and blood chemistry results were consistent with overwhelming systemic infection and starvation. The animal also was antibody-positive for canine distemper virus, feline panleukopenia, and feline coronavirus.

The acoustic features of the long distance advertisement call produced by Panthera tigris altaica, the Amur (Siberian) tiger.

The long distance advertisement call (LDAC) frequently referred to as a moan, an intense mew or a territorial roar, is commonly exchanged by adult members of Panthera tigris living both in captivity and in the wild. While the functionality of the call is not understood ethologically, its social nature is unambiguous, and isolated individuals living in captivity exchange the call repeatedly over extended periods of time. In this report, we concentrate on LDACs produced by P. tigris altaica. The average duration of the LDAC was just under 2 s. The average fundamental frequency, f0, was 0.155 kHz, ranging from ∼0.113 to 0.176 kHz. Spectrograms were dominated by distinct harmonically related frequency modulated frequency sweeps, although subharmonics were also observed. Overall peak frequency, the frequency exhibiting maximum acoustic power, was centered at ∼0.330 kHz and instantaneous peak frequency frequently shifted from one harmonic to another throughout the call’s time course. The bulk of broadcast energy was contained in a band of frequencies ranging from ∼0.1 to 0.8 kHz at 20 dB down from the peak frequency, and bandwidth expanded to ∼3.7 and 4.9 octaves at the 30 and 40 dB down points, respectively. [Funding provided by NSF Grant 0823417.]

Habitat evaluation of wild Amur tiger ( Panthera tigris altaica) and conservation priority setting in north-eastern China

The Amur Tiger ( Panthera tigris altaica) is one of the world’s most endangered species. Recently, habitat fragmentation, food scarcity and human hunting have drastically reduced the population size and distribution areas of Amur tigers in the wild, leaving them on the verge of extinction. Presently, they are only found in the north-eastern part of China. In this study, we developed a reference framework using methods and technologies of analytic hierarchy process (AHP), remote sensing (RS), geographic information system (GIS), GAP analysis and Natural Break (Jenks) classification to evaluate the habitat and to set the conservation priorities for Amur tigers in eastern areas of Heilongjiang and Jilin Provinces of northeast China. We proposed a Habitat Suitability Index (HSI) incorporating 7 factors covering natural conditions and human disturbance. Based on the HSI values, the suitability was classified into five levels from the most to not suitable. Finally, according to results of GAP analysis, we identified six conservation priorities and designed a conservation landscape incorporating four new nature reserves, enlarging two existing ones, and creating four linkages for Amur tigers in northeast China. The case study showed that the core habitats (the most suitable and highly suitable habitats) identified for Amur tigers covered 35,547 km 2, accounting for approximately 26.71% of the total study area (1,33,093 km 2). However, existing nature reserves protected only (7124 km 2 or) 20.04% of the identified core habitats. Thus, enlargement of current reserves is necessary and urgent for the tiger’s conservation and restoration. Moreover, the establishment of wildlife corridors linking core habitats will provide an efficient reserve network for tiger conservation to maintain the evolutionary potential of Amur tigers facing environmental changes.

Evaluation of nutrient digestibility and fecal characteristics of exotic felids fed horse‐ or beef‐based diets: use of the domestic cat as a model for exotic felids

The objective of this study was to determine the effects of feeding commercially available beef- and horse-based diets on nutrient digestibility and fecal characteristics of large captive exotic felids and domestic cats. Four species of large exotic felids including cheetahs, Malayan tigers, jaguars, and Amur tigers, and domestic cats were utilized in a crossover design. Raw meat diets included a beef-based diet (57% protein; 28% fat) and a horse-based diet (51% protein; 30% fat). All cats were acclimated to the diet for 16 days followed by a 4 day collection period, where total feces, including one fresh sample, were collected. All feces were scored on collection. Intake did not differ due to diet, but fecal output was greater when cats consumed the horse-based diet. Total tract apparent dry matter (DM) digestibility was higher (P<0.05) and organic matter (OM) and crude protein (CP) digestibilities were lower (P<0.05) when cats were fed the beef-based diet compared with the horse-based diet. CP digestibility was similar in domestic cats and cheetahs, and greater (P<0.05) than Amur tigers. Fecal scores were lower and fecal DM was greater (P<0.05) when cats consumed the horse-based diet compared with the beef-based diet. Domestic cats had lower (P<0.05) fecal ammonia concentrations compared with all other species. Fecal ammonia concentrations were lowest (P<0.05) when cats were fed the horse-based diet. Fecal total short-chain fatty acid (SCFA), branched-chain fatty acid (BCFA), and butyrate concentrations were higher (P<0.05) when cats consumed the beef-based diet. Our results suggest that the domestic cat serves as an appropriate model for large exotic felid species, but differences among the species exist. Decreased nutrient digestibility by tigers and jaguars should be considered when developing feeding recommendations for these species based on domestic cat data.

Spatial structure of Amur (Siberian) tigers (Panthera tigris altaica) on Sikhote-Alin Biosphere Zapovednik, Russia

Abstract Spacing patterns of large carnivores can affect demographic parameters of populations that, in turn, influence effective population size. As a result, better understanding of spatial structure can provide insight into effective conservation strategies. We examined home-range size, spacing characteristics, and changes in land tenure of radiocollared Amur tigers (Panthera tigris altaica) on the Sikhote-Alin Biosphere Zapovednik, Russia, from 1992 to 2006. We predicted that both sexes would maintain spatially exclusive home ranges and that subadult female tigers would tend toward philopatry and males would disperse. Home ranges (95% fixed kernel estimates; mean ± SD) of resident females (n = 20 home ranges of 14 females; 390 ± 136 km2) were significantly (P = 0.003) smaller than those of males (n = 6 home ranges of 5 males; 1,385 ± 539 km2). Geometric mean overlap between adjacent females (0.11 ± 0.11 SD) did not differ from that between adjacent males (0.14 ± 0.12). All radiocollared male cub...