Biochemical Genetic Relationships Among Tunisian Hares (Lepus sp.), South African Cape Hares (L. capensis), and European Brown Hares (L. europaeus)

Tunisian hares (n = 45), currently assigned to Lepus capensis, were assayed for allelic variation at 40 allozyme loci, and allele frequencies at 32 loci were directly compared with earlier data of South African cape hares (L. capensis, n = 9) and European brown hares (L. europaeus, n = 244) to reveal genetic relationships among them. European mountain hares (L. timidus, n = 200) were used for outgroup comparison. In the Tunisian hares 27.5% of the loci were polymorphic with 2–4 alleles. Among all alleles at polymorphic loci, 15.1% occurred exclusively in Tunisian hares, 5.7% exclusively in cape hares, and 7.5% exclusively in brown hares at low frequencies. Not a single locus showed alternately fixed alleles between the samples of the L. capensis/L. europaeus complex. Levels of absolute and relative genetic differentiation among the samples of the L. capensis/ L. europaeus complex were low, relative to pairwise comparisons involving mountain hares. Diverse cluster analyses and multidimensional scaling of various pairwise genetic distance matrices concordantly grouped Tunisian hares with brown hares, and South African cape hares clustered only slightly farther apart, whereas mountain hares were distinctly separate. These results suggest regionally distinct phylogenetic units within an overall cohesive gene pool in the L. capensis/ L. europaeus complex, supporting Petter's view that all North African hares belong to L. capensis except for one local population of savanna hares, and that cape hares and brown hares are conspecific.     

Infections with Francisella tularensis biovar palaearctica in hares (Lepus timidus, Lepus europaeus) from Sweden

The occurrence of tularemia was studied in 1,500 hares submitted to the National Veterinary Institute, Uppsala, Sweden for postmortem examination during 1973 through 1985. A total of 109 tularemia cases was recorded based on the fluorescent antibody (FA) test for Francisella tularensis and on the gross and microscopic pathology. Tularemia was diagnosed only in the varying hare (Lepus timidus) and not in the European brown hare (Lepus europaeus). The geographical distribution of the 109 cases indicates that tularemia has not spread in Sweden during the last 45 yr, with the exception of an endemic occurrence of the disease on the island of Stora Karls? in the Baltic sea. The disease was most frequent in the autumn and only a few cases were recorded during winter. Cases were not seen in the spring. The annual prevalence varied, with several cases in 1974 and 1981, but there were no cases in 1976 and 1980. The postmortem findings in hares dying of tularemia in the autumn were characterized by focal coagulative necrosis in liver, spleen and bone marrow, with high numbers of bacteria FA-positive for F. tularensis. In hares dying during winter months, the most characteristic findings were hemorrhagic enteritis and typhlitis, although necrotic lesions could occur in liver, spleen and bone marrow. Diseased hares on the island of Stora Karls? were demonstrated to be infected with ticks, while hares on the mainland of Sweden generally were fed upon by mosquitoes. Twenty-six of the 109 hares with tularemia were examined bacteriologically and F. tularensis biovar palaearctica was isolated from eight. The lung extract antibody test for F. tularensis was performed in 18 of the 109 hares. All were negative. In addition to the field study, an experimental study with F. tularensis biovar palaearctica was performed. Four varying hares and three European brown hares were inoculated. None of the hares died from tularemia, and generalized infection was not demonstrated.     

Comparative in vitro study of caecal microbial activity in brown hares and domestic rabbits which were offered the same diet

The products of the microbial activity in the large intestine are an important source of energy for herbivores. A previous study showed differences in caecal methanogenesis and short-chain fatty acid (SCFA) profiles between brown hares and domestic rabbits. The present study was performed on animals which were offered the same diet to eliminate the impact of diet on the comparative analyses of microbial metabolites. Caecal samples of hares and rabbits were incubated in triplicate, i.e. without any supplementary substrate (control) or with the addition of wheat bran or oat bran. Calculated as percentage of body weight, the stomachs of the rabbits were heavier than those of the hares, but caeca of the hares weighed more than those of the rabbits. The total SCFA concentration in caecal samples was higher in rabbits than in hares, and it increased in both species when the supplementary fermentation substrates were added. In hares, the molar proportion of propionate was higher and that of butyrate was lower compared to rabbits. The addition of substrate decreased acetate and propionate but increased the molar proportion of butyrate. Microbial fermentation resulted in greater gas release in rabbit caecal samples compared to those of hares. Methanogenesis tended to be lower in hares than in rabbits, but high individual variability was observed, especially in hares. Our study stated lower microbial activity in the caeca of brown hares compared to domestic rabbits. The presented results might lead to assumption that differences between fermentation patterns were not caused by diet but resulted from the peculiarities of both species.     

Helminth communities of an introduced hare (Lepus granatensis) and a native hare (Lepus europaeus) in southern France

We investigated the parasite communities of introduced Iberian hares (Lepus granatensis) and native European hares (Lepus europaeus) in southern France, where Iberian hares were introduced locally 20 yr ago as a game animal. Parasite communities of sympatric populations of the two hare species and of allopatric populations of European hares were compared. Iberian hares in France harbored a depauperate community of parasites relative to the population in its native habitat in Spain. European hares in areas of sympatry also were infected by Nematodiroides zembrae, which normally infects Iberian hares on their native range.     

Owl predation on snowshoe hares: consequences of antipredator behaviour

We show evidence of differential predation on snowshoe hares (Lepus americanus) by great horned owls (Bubo virginianus) and ask whether predation mortality is related to antipredator behaviour in prey. We predicted higher predation on (1) young and inexperienced hares, (2) hares in open habitats lacking cover to protect from owl predation, and (3) hares in above average condition assuming that rich food patches are under highest risk of predation. Information on killed hares was obtained at nest sites of owls and by monitoring hares using radio-telemetry. The availability of age classes within the hare population was established from live-trapping and field data on reproduction and survival. Great horned owls preferred juvenile over adult hares. Juveniles were more vulnerable to owl predation before rather than after dispersal, suggesting that displacement or increased mobility were not causes for this increased mortality. Owls killed ratio-collared hares more often in open than in closed forest types, and they avoided or had less hunting success in habitats with dense shrub cover. Also, owls took hares in above average condition, although it is unclear whether samples from early spring are representative for other seasons. In conclusion, these results are consistent with the hypothesis that variation in antipredator behaviours of snowshoe hares leads to differential predation by great horned owls.     

The effect of establishing native woodland on habitat selection and ranging of moorland mountain hares (Lepus timidus), a flexible forager

Mountain hares Lepus timidus L. typify species that occupy a broad geographic range and have flexible foraging and nutritional strategies. Such species may show a range of responses to habitat modification. This study aimed to provide a basis for prediction of the impact of mountain hares on woodland establishment, and of woodland establishment on mountain hare distribution. The selection of and the extent of incorporation of new woodland into the home range of mountain hares was investigated in an area where Scots pine Pinus sylvestris L. woodland was establishing within their usual habitat in Britain, upland heather moorland. Seasonal home, day and night-range sizes of radio-tracked mountain hares were determined using the multinuclear probability polygon technique and analysed using residual maximum likelihood (REML). Habitat selection was analysed using compositional analysis. Three main habitat types were available to hares: heather moorland with trees, heather moorland and grassland-mire. Mean home-range size of mountain hares in summer was 10.3 ha and in winter 9.6 ha. There were no significant seasonal or sex differences in home-range size. Females selected grassland-mire habitat in summer and showed no strong selection for any habitat in winter. Males selected heather moorland in both summer and winter. Heather moorland with trees was not selected preferentially by mountain hares of either sex in summer or winter. The absence of selection for areas of newly establishing-Scots pine woodland suggests that any browsing damage to trees by hares is most likely to be a function of the local abundance of mountain hares, rather than a result of active preference of hares for the modified habitat.     

Borrelia burgdorferi sensu lato detected in skin of Norwegian mountain hares (Lepus timidus) without signs of dissemination

The mountain hare (Lepus timidus) population in southern Norway appears to be in decline. Necropsy and laboratory examinations of 36 hares found dead or diseased during 2007-2009 in Vest- and Aust-Agder counties showed that disease and deaths were attributed to multiple causes, with no specific etiology emerging as a cause for population decline. To investigate whether Borrelia burgdorferi sensu lato (s.l.) infection is associated with mortality in mountain hares, tissues and ticks collected from hares were investigated for infection with the spirochete. Borrelia burgdorferi s.l. DNA was not detected in samples from internal organs, whereas Borrelia afzelii, B. burgdorferi sensu stricto (s.s.), and the not-yet-defined Borrelia sp. SV1 were found in skin samples from hares and in adult and nymphal Ixodes ricinus feeding on hares. Only B. burgdorferi s.s. and Borrelia sp. SV1 were detected in larvae feeding on hares. Our results indicate that disseminated Borrelia infection in hares rarely occurs and, presumably, does not play a central role in the suspected population decline. The results also suggest that the mountain hare to some degree functions as a transmission host for B. burgdorferi s.s. and Borrelia sp. SV1.     

Natural toxoplasma gondii infections in European brown hares and mountain hares in Finland: proportional mortality rate, antibody prevalence, and genetic characterization

In material examined postmortem in Finland from May 2006 to April 2009, acute generalized toxoplasmosis was the immunohistochemically confirmed cause of death in 14 (8.1%) of 173 European brown hares (Lepus europaeus) and four (2.7%) of 148 mountain hares (Lepus timidus). Sera from 116 of the European brown hares and 99 of the mountain hares were screened with a commercial direct agglutination test for Toxoplasma gondii-specific IgG antibodies at a dilution of 1:40. All sera from cases of fatal toxoplasmosis had high titers of antibodies reactive to T. gondii. In contrast, none of 107 European brown hares and four (4%) of 96 mountain hares that died of other causes were antibody-positive. The proportional mortality rates and the T. gondii antibody prevalences among noncases differed significantly between the two host species (P<0.05). Direct genetic characterization of the causative agent was performed on DNA extracted from formalin-fixed, paraffin-embedded tissue of the hares with fatal toxoplasmosis. Based on the results with six microsatellite markers (B18, TUB2, TgM-A, W35, B17, and M33; all six in 15 cases and four in three cases), all the cases were caused by T. gondii genotype II; the size of the PCR product at the seventh marker (M48) varied (213-229 base pairs). The presence of T. gondii genotype II, which is endemic in Europe, is now confirmed in Finnish wildlife: Natural infections with T. gondii parasites belonging to this widespread genotype caused fatal generalized toxoplasmosis in the two species of wild hares.     

Blood chemistry and endoparasites of the mountain hare (Lepus timidus L.) in high and low density populations.

In order to study the effect of high population density on the condition, blood characteristics and helminth parasitism of mountain hares (Lepus timidus), 12 specimens were shot in December 1982 and 12 more in February 1983 on the west coast of central Finland (group 1, dense population). In addition 14 hares were shot in December 1982 about 100 km from group 1 (group 2, dense population). Group 3 consists of 15 hares from stable, rather low density populations shot in southern Finland during three previous winters. The hares in group 1 were the lightest, had the least fat and were the most seriously infected with Protostrongylus pulmonalis and Trichostrongylus retortaeformis, while those in group 2 were the heaviest and had the highest Ca, Mg, alkaline phosphatase and creatinine values. The group 3 hares had the most fat. The group 1 animals shot in February 1983 had higher Ca, Mg, triglyceride and cholesterol values than those shot in December 1982. It seems that high population density combined with a lack of suitable food leads to poor condition and high endoparasite abundances. The differences in Ca and Mg are probably due to diet. The higher creatinine values in group 2 and in the hares with little or no T. retortaeformis infection may be due to the greater muscle mass.     

Performance and mortality of farmed hares

Performance and mortality of hares were evaluated for 2 consecutive years in a large farm in Veneto Region (Northern Italy). On average, fertile reproductive pairs (n=318) gave birth 4.8 times and produced 11.4 live leverets, weaned 8.4 leverets and produced 7.0 growing hares (60 days) every year. Mean mortality was 3.6%, 22.9%, 9.7% and 2.5% in newborn (0 to 2 days of age), suckling (3 to 25 days), growing (26 to 60 days) and sub-adult (61 days until sale) hares, respectively. The main causes of mortality were enteric diseases (75.5%, 75.9% and 12.1% in suckling, growing and sub-adult hares, respectively), followed by respiratory diseases (3.4%, 8.0% and 36.2% in suckling, growing and sub-adult hares, respectively), starvation (11.3% and 8.8% in suckling and growing hares, respectively) and trauma (7.1%, 2.3% and 30.2% in suckling, growing and sub-adult hares, respectively). In reproducing hares, mortality was 24.7% and 15.4% in 2010 and 2011, respectively. Respiratory diseases (34.8%) and ulcerative pododermatitis (18.9%) were the most common pathological changes detected in reproducing hares. Farmed hares seem to be affected by diseases resembling those of rabbits reared under intensive conditions. It seems necessary to improve the husbandry of hares to reach satisfactory technical standards and to preserve their health.     

Overwinter mass loss of snowshoe hares in the Yukon: starvation, stress, adaptation or artefact?

1. Overwinter mass loss can reduce energetic requirements in mammals (Dehnel's phenomenon). Alternatively, mass loss can result from food limitation or high predation risk. 2. We use data from fertilizer, food-supplementation and predator-exclusion experiments in the Yukon during a population cycle from 1986 to 1996 to test the causes of overwinter mass loss by snowshoe hares (Lepus americanus). In all years, some hares on control sites gained mass overwinter. During the increase phase the majority gained mass, but in all other phases the majority lost mass. 3. Snowshoe hares weighing <1000 g in autumn always gained mass overwinter, as did the majority that weighed 1000-1400 g. Hares weighing >1800 g in autumn usually lost mass. 4. Snowshoe hares on the predator-exclosure + food site gained mass overwinter in all years. Hares on the food-supplementation sites lost mass during the decline but gained mass in all other phases. Fertilization had little effect on mass dynamics. 5. Snowshoe hares were more likely to lose mass during winters with low survival rates. Snowshoe hares on the predator-exclosure treatments were more likely to gain mass than were hares on control sites. 6. Overwinter mass loss was correlated with maximum snow depth. At equivalent snow depths, hares on food-supplemented areas lost 98 g (+/- 14.6 SE) less on average than hares on the controls and predator-exclosure treatment. 7. Bone-marrow fat was related to body mass and cause of death. Small hares had the lowest marrow fat. Hares killed by humans had higher marrow fat than those killed by predators; hares that simply died had the lowest marrow fat. Hares on food-supplemented sites had the highest kidney and marrow fat. 8. Overwinter-mass loss for snowshoe hares is explained interactively by winter conditions, food supply, predation risk and autumn mass. Some snowshoe hares lost mass overwinter in all years and on all treatments, suggesting that reducing body mass may facilitate survival, especially in cases where foraging costs are high energetically or increase predation risk.     

Characteristics of natural infections of the stomach worm, Obeliscoides cuniculi (Graybill), in lagomorphs and woodchucks in Canada

Wild lagomorphs and woodchucks collected predominantly in southern Ontario, Canada were examined for subspecies of Obeliscoides cuniculi (Graybill). Obeliscoides cuniculi multistriatus was found in snowshoe hares (Lepus americanus). Obeliscoides cuniculi cuniculi was found in cottontail rabbits (Sylvilagus floridanus), European hares (Lepus capensis) and woodchucks (Marmota monax). Prevalence of Obeliscoides cuniculi multistriatus in snowshoe hares was 100% and mean intensity (and range of intensity) was 760 (9-4, 198) in Lindsay, Ontario in 1980. Mean intensity in hares varied trimonthly. The highest mean intensity of worms occurred in spring when most worms were adult. Transmission occurred mainly in spring. Most worms present in fall (70%) and winter (54%) were fourth stage. Immature fifth-stage and gravid females were present in hares during fall and winter. Prevalence and mean intensity of O. c. cuniculi in cottontails was 15% and 29 (1-118). Prevalence and mean intensity of O. c. cuniculi in woodchucks was 6% and 56 (16-118). European hares were infected with O. c. cuniculi, prevalence was 10% and mean intensity was 60 (36-83). In Ontario woodchucks and European hares were common in areas frequented by cottontail rabbits and probably acquired sporadic infections of O. c. cuniculi from infected cottontails.     

Artificial insemination in the European hare (Lepus europaeus syriacus)

A simple technique for artificial insemination in hares was performed and used routinely in mass breeding of hares in captivity. Sperm was collected directly from the epididymides of males. Ovulation was induced by intravenous injection of human chorionic gonadotropin. Female hares were successfully inseminated throughout the year. Pregnancy occurred in 70% of inseminated females; the range in litter size was 1--3 offspring with a mean of 1.4. It was concluded that artificial insemination greatly simplified the breeding of hares in captivity.     

Hare and vole browsing preferences during winter

The aim with this study was to, under controlled conditions, determine the food preference of mountain haresLepus timidus Linnaeus, 1758 and bank volesClethrionomys glareolus (Schreber, 1780) for a substantial part of the woody plants potentially available for these herbivores during winter. In addition, we compared hare and vole preference patterns. Thirteen woody plant species were simultaneously presented to 9 captive voles and 9 captive hares in preference tests during winter. Consumption by hares from 50 g bundles (one per species) was measured after 3 h, whereas shoot consumption by voles was measured after 12 h. Both hares and voles preferred deciduous species to conifers,Populus tremula, andVaccinium myrtillus being the most preferred species. However, there was considerable variation in palatability among deciduous plant species and only a marginally significant correlation was found between hare and vole preference. One striking differences between hares and voles was that Sorbusaucuparia was the most utilised by voles but the least preferred by hares. In conclusion, deciduous plant species were generally considerably more palatable to hares and voles than conifers, which is consistent with current theories. Nevertheless, the high variation in palatability among deciduous trees and the difference in preference between hares and voles indicate more complex and species-specific patterns with regard to plant— animal interactions. The latter also suggests that hares and voles differ in their dietary adaptations and have different dietary constrains.     

中国更新世兔属化石的厘定

在对我国现生兔属各种骨骼特征研究的基础上,通过形态特征的观察与测量数据的分析,修订了已有的兔属化石材料。将周口店第1,2,6,20,山顶洞和东岭子洞等地点以及山东平邑发现的兔标本归入草兔Lepus capensis。以前被归入Lepus wongi种内的标本,除周口店第13地点及蓝田陈家窝子的应属于Lepus teilhardi外,其余地点的皆为Lepus capensis,因此L.wongi失去了原有的意义,为无效种。周口店第16地点的标本归入华南兔Lepus sinensis,是该种目前惟一的化石记录。辽宁营口金牛山的标本暂时归入东北兔Lepus mandschuricus。我国更新世共发现两个化石种和4个现生种。其中化石种Lepus teilhardi和L.ziboensis出现在早更新世晚期至中更新世早期,现生种L.capensis,L.mandschuricus,L.sinensis和L.comus分别出现在中更新世与晚更新世。初步讨论了兔属的起源,认为兔属可能起源于上新世的Trischizolagus。     

Factors related to the occurrence of hybrids between brown hares Lepus europaeus and mountain hares L. timidus in Sweden

Hybridization occurs among many species, and may have implications for conservation as well as for evolution. Interspecific gene flow between brown hares Lepus europaeus and mountain hares L. timidus has been documented in Sweden and in continental Europe, and has probably to some extent occurred throughout history in sympatric areas. What local factors or ecological relationships that correlate with or trigger hybridization between these species has however been unclear. We studied spatial distribution of hybrids between brown hares and mountain hares in Sweden in relation to characteristics of the sampled localities (hunting grounds). In a sample of 70 brown hares collected from 39 populations in south‐central Sweden during 2003–2005, 11 (16%) showed introgressed mtDNA from mountain hares. Among the brown hares from their northern range, i.e. in general the most recent establishments, the corresponding figure was 75% (9/12). The frequency of samples with hybrid ancestry increased significantly with latitude, altitude and hilliness, and were higher (p<0.1) in recently established populations and/or where the proportion of arable land was low. Several site‐specific parameters were correlated, e.g. latitude as expected to hilliness, and no parameter explained the occurrence of hybrids exclusively. Instead, the appearance of mountain hare mtDNA among brown hares was associated with a conglomerate of parameters reflecting landscapes atypical for the brown hare, e.g. forest dominated and steep areas where the species quite recently was established. We suggest that these abiotic factors mirror the main aspect influencing hybridization frequency, namely the density or relative frequency of the two species. In atypical brown hare landscapes with recent establishment, mountain hares are probably relatively more common. When one species dominate in numbers, or when both species display low densities, increased frequency of hybridization is expected due to low availability of conspecific partners, a phenomenon referred to as Hubbs’ principle.     

European brown hare syndrome in free-ranging European brown and mountain hares from Switzerland.

From 1997 to 2000, complete necropsy and histopathologic investigations were performed on 157 free-ranging European brown hares (Lepus europaeus) found dead throughout Switzerland. Organ samples of all these individuals (157 livers and 107 spleens available) were tested for European brown hare syndrome virus (EBHSV)-antigen by enzyme-linked immunosorbent assay (ELISA) test kit. Furthermore, 60 additional blood samples were tested for antibodies against EBHSV by ELISA. In addition, liver samples of 87 free-ranging mountain hares (Lepus timidus) hunted in 1996 were tested for EBHSV-antigen. In two European brown hares from southern Switzerland lesions suggestive of changes induced by EBHSV were present, and high titers of EBHSV-antigen were detected in both liver and spleen samples of these animals. Based on negative staining electron microscopy investigations of liver and spleen homogenates, we observed calicivirus in one antigen-positive hare. Low EBHSV-antigen titers were found in three additional European brown hares from central and western Switzerland, but EBHS-lesions were absent. Antibodies against EBHSV were not detected in any of the sera of European brown hares, and EBHSV-antigen was not found in the samples of mountain hares. This is the first report of EBHS in European brown hares from Switzerland.     

The occurrence of mountain hare mitochondrial DNA in wild brown hares

If interspecific hybrids are fertile and backcross to either parental species, transmission of mitochondrial DNA over the species barrier can occur. To investigate if such transmission has occurred between the brown hare Lepus europeus Pall and the mountain hare L. timidus L. in Scandinavia, an analysis of genetic variation in mitochondrial DNA from 36 hares, collected from 15 localities, was performed. Sequence divergence of mtDNA between species was estimated at 8 ± 1% (SD). Intraspecific mtDNA sequence divergence varied between 0.09 and 0.38% in brown hares and 0.10 and 1.44% in mountain hares. In six out of 18 brown hares examined, two different haplotypes of mountain hare origin were detected, demonstrating a transmission of mtDNA haplotypes from mountain hares to brown hares. The results indicate that interspecific hybridization between the two species occurs in wild populations.