Habitat selection and activity patterns in Alpine mountain hare (Lepus timidus varronis)

We investigated activity patterns and habitat use of 34 radio-tracked mountain hares (Lepus timidus) in the Italian Alps. We first showed that hares were nocturnal and that activity patterns changed seasonally in parallel with circadian rhythms. We predicted that day home ranges will include suitable resting (shelter) habitats, and night home ranges will primarily include suitable foraging habitats. A hare's night-range was larger than its day-range. On average, night and day ranges overlapped by 36%, suggesting that selective pressures affecting space use were, at least partly, different at night than day. Dwarf mountain-pine was the most preferred habitat in all seasons and was selected both for active behaviour (night) and resting (day) and hares avoided the most open habitats. Exploring the effects of season, time of day (day vs. night) and site, we found that habitat use by mountain hares did not differ between seasons or between the active (night) and resting (day) period of circadian cycle. Also, we found no effects of differences in landscape structure (habitat patchiness and heterogeneity) on the patterns of habitat selection. Hares always preferred the dense, forested habitats, which seemed to provide food resources as well as shelter from predators throughout the year.     

The 24-hour cycle of the mountain hare Lepus timidus Linnaeus, 1758

The behavioral decisions of wild animals are influenced by the often conflicting needs to both feed efficiently and to avoid predators. Information regarding how lagomorphs such as the mountain hare (Lepus timidus) balance these factors throughout the day-night cycle is largely lacking despite its importance to management.To better understand the type and proportion of various behaviors the mountain hare engages in over the course of a 24-hour period, I monitored the behavior of six mountain hares under controlled conditions from early to mid-winter. The mountain hares spend the majority of their time resting (mean ± standard deviation: 32.4 ± 14.4%) and under canopy (22.4 ± 18.7%), and a smaller proportion of their time feeding (16.2 ± 6.1%). Activity peaks over the course of the 24-hour cycle reflect natural behavior patterns that emphasize the reduction of predation risk. The sunrise phase is characterized by the search for a suitable resting place; the day phase by resting and hiding. The sunset phase is characterized by the search for food, and the night phase by an equal mix of food intake and movement through the home range. These results can help land managers predict how factors such as increasing tourist activity (perceived by the hare as a safety threat) will influence the behavior of free-ranging mountain hares.     

The potential role of habitat on intestinal helminths of mountain hares, Lepus timidus

Over the last century in the uplands of Scotland, the extent of heather moorland which supports high densities of mountain hares Lepus timidus has diminished and has gradually been replaced by large-scale commercial forestry plantations or expanding natural woodlands. The potential impact of such a change in land use on host-parasite interactions was investigated by comparing the intensity and prevalence of infection of hares by parasites in two separate habitats: a large hare-fenced young forestry plantation and the adjacent open moorland. Carcasses were collected in November 1990 from within both habitats and after the woodland had been enclosed for nine months. Age, sex, fatness (kidney fat index) and degree of infection of hares were noted. Two parasites were recorded: the nematode Trichostrongylus retortaeformis and the cestode Mosgovoyia pectinata. Clear differences in the intensity of infection of adults occupying the different habitats had occurred in the nine months since woodland enclosure. Adult mountain hares in the woodland had levels of infections approaching four times that observed in hares occupying the open moorland and although not significant, the prevalence of infection was greater in hosts inhabiting the woodland than the open moorland. It is suggested that the parasite-host relationship differs between the two habitats and as heather-dominated moorland landscapes become more fragmented with the increasing establishment of woodlands, the impact of parasites on the life history strategies of mountain hares needs to be reconsidered.     

Contagious mucocutaneous dermatitis of the mountain hare (Lepus timidus): pathology and cause

Contagious mucocutaneous dermatitis is a frequently encountered disease of mountain hares (Lepidus timidus) in Finland. We describe the histopathologic changes and propose an etiologic cause for this disorder. Fifty-three cases collected during 1982-2000 were examined histologically. Transmission electron microscopy was performed in one case. In fully developed lesions, keratinocytes in epidermis and follicular infundibula were swollen and contained large eosinophilic intracytoplasmic inclusion bodies with marked reticular and ballooning degeneration. In later stages, there was marked necrosis and ulceration with severe pyogranulomatous and suppurative inflammation. At this stage, no viral inclusions were detectable, but secondary Staphylococcus warnerii infection was present in most cases. In late lesions, there was dermal fibrosis with epidermal hyperplasia. No spiral-shaped bacteria suggesting treponematosis were detected at any stage. Ultrastructurally, swollen epidermal and follicle infundibular cells contained round intracytoplasmic inclusion bodies with a myriad of virions typical of poxvirus with a biconcave nucleocapsid core, two lateral bodies, and a clearly discernible outer lipoprotein capsule. The findings suggest that contagious mucocutaneous dermatitis in mountain hares is a viral disease caused by a poxvirus. The disease is often complicated by secondary bacterial infection, most commonly S. warneri.     

Behaviour, population changes and dispersal of mountain hares (Lepus timidus) in Scotland

Mountain hares in north-east Scotland spent the day in forms in long heather and moved downhill in the evening to feed in hill pastures. Here they grazed intensively, often in groups of 4–6 individuals. In June and July adult females, then pregnant or lactating, grazed in daylight on the pastures. Leverets spent less time grazing, and more in play and exploration. They spent the day in cover near the feeding areas, and if disturbed during grazing crouched or went into cover, while adults fled. During 1982 and 1983 males predominated in groups of grazing hares, but in 1984 both sexes were equally represented. These changes in sex ratio were reflected in the increased proportion of females trapped for marking between February and late July during 1982–86. There was a dominance order related to weight among male hares and dominant hares approached more females. There was no firm evidence of mate-guarding. Males approached females regardless of their oestrous state and were usually rebuffed with varying degrees of intensity including striking and chasing. There were no interactions between females. Adults of both sexes chased leverets for short distances but leverets joined groups of feeding adults. Neither leverets nor first-winter hares showed evidence of dispersal. Mountain hares avoided sheep and cattle and there were fewer hares after the arrival of sheep in May.     

Molecular phylogeny of Japanese Leporidae,the Amami rabbit Pentalagus furnessi,the Japanese hare Lepus brachyurus,and the mountain hare Lepus timidus,inferred from mitochondrial DNA sequences

We determined mitochondrial 12S ribosomal RNA (rRNA) and cytochrome b (cyt b) gene sequences in three leporid species of Japan, the Amami rabbit Pentalagus furnessi from the Ryukyu Islands, the Japanese hare Lepus brachyurus from Honshu, and a Japanese form of the mountain hare Lepus timidus ainu from Hokkaido. We compared the sequences with those of other taxa of leporids available in databases. Phylogenetic trees of the 12S rRNA gene sequences indicated that the lineage of P. furnessi diversified during the generic radiation of the leporids at an ancient time, which was estimated to have been the middle Miocene. Cyt-b gene trees revealed that the lineage of L. brachyurus branched off at an early stage in the speciation of Lepus, probably at the beginning of the Pliocene. The cyt b sequences of L. t. ainu were somewhat distinct from those of continental conspecific populations; this lineage divergence is likely to have occurred during the middle or late Pleistocene. The results show that the three regions of the Japanese archipelago, Ryukyu, Honshu-Shikoku-Kyushu, and Hokkaido, now preserve their own leporid taxa, each with a different extent of genetic endemicity. It is possible that the zoogeographic traits of the Japanese leporids are a consequence of the evolutionary dynamics of leporids in East Asia, in that the radiation centers of leporids are likely to have shifted from tropical, through temperate, to arctic zones.     

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.     

Weights and growth rates in the Mountain hare Lepus timidus scoticus

Mountain hares were weighed during live trapping on a study area near Dufftown, Banffshire, from June 1958 to August 1966. Hares shot or killed by other means on the study area and elsewhere were also weighed. Females were heavier than males throughout the year and this difference became apparent in juveniles by August of their year of birth. Male hares lost weight during the breeding season (January to June) but regained it in late summer. Young hares gained weight initially at about 14 g per day, then at about 6 g per day to adult weight. Small juveniles, or those born late in the season, tended to become small adults, large or early juveniles to become large adults. Small hares moulted less completely andjbegan to breed later in the season than large hares. The effects of disease, starvation, severe weather and injury on hare weights were considered. Female weights were greater when the population was small, due to more late pregnancies or more embryos per female. Weight could not be used to distinguish between adult and juvenile hares above 2.1 kg, nor between young and older adults.     

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.     

A preliminary study on the seasonal body temperature rhythms of the captive mountain hare (Lepus timidus)

The mountain hare (Lepus timidus) is a year-round active herbivore adapted to survive the boreal winter. Captive mountain hares (N = 4) were implanted with intraabdominal thermosensitive loggers to record their core body temperature (T_b) for a year and during food deprivation (8–48 h) in summer and winter. The average T_b was 38.7 ± 0.01 °C in summer and 38.3 ± 0.01 °C in winter. The yearly T_b correlated positively with the ambient temperature. The 24-h T_b was the highest from late scotophase to early photophase in summer and winter and the lowest during middle-late photophase in summer or during early-middle scotophase in winter. The range of the 24-h oscillations in T_b increased in three animals in winter. Food deprivation did not induce hypothermia in summer or winter. These preliminary data suggest that the mountain hare can spare a modest amount of energy with the wintertime reduction in T_b.     

Diet of the Iberian hare (Lepus granatensis) in a mountain ecosystem

The diet of the Iberian hare (Lepus granatensis) was studied through microhistological pellet analysis in two areas from a mountain ecosystem in Central Portugal. Fecal pellets were collected monthly in 24 plots spatially distributed throughout the two study areas. For each period, a sample of 15 to 20 pellets was milled and 400 epidermal fragments were identified, by comparison with a reference collection. A wide range of plant species was observed in hare’s diet. Grasses represent the basis of the Iberian hare diet, with frequencies always higher than 50% in both study areas (annual average = 69.98%). Most of the 35 species of grasses assembled for the reference collection (91.43%) were identified in the pellets. Nevertheless, only six of these were consumed in proportions greater than 5%, being Anthoxanthum odoratum, Secale cereale and Agrostis spp. the species ingested in higher frequencies. The rate of grasses consumption reached 80.69% in winter but decreased in summer to around 55%. In this season, a concurrent rise in the ingestion of other plant groups, like herbs and shrubs, and of plant inflorescences was observed. This work provides the first results on the Iberian hare’s diet on mountain ecosystems, and suggests that the Iberian hare diet in a mountain ecosystem is similar to the observed in L. europaeus and L. timidus.     

Growth and hare, Lepus timidus, resistance of white birch, Betula pendula, clones grown in different soil types

Many plant species have the ability to expand laterally through space by clonal growth. Plant pathogens can affect clonal growth characteristics thereby altering the success of host plants within populations and of clonal species within communities. We conducted a greenhouse experiment to determine the effects of the systemic fungal pathogen, Epichloë glyceriae , on clonal growth patterns of its host grass, Glyceria striata . We found that infected and uninfected plants produced similar total biomass and numbers of tillers plus primary stolons per mother ramet. However, biomass allocation to tillering (vegetative growth) vs stolon production (clonal growth) was significantly affected by pathogen infection. Infected plants produced more stolons and clonal growth biomass than uninfected plants while mother ramets of uninfected plants produced more tillers and biomass than infected plants. Stolon architecture of infected and uninfected plants also differed. In two of three populations, infected plants produced stolons with greater biomass and shorter spacer lengths, even though mean stolon lengths were similar for infected and uninfected plants. These results contrast strongly with most other clonal plant-pathogen systems where infected plants are less vigorous and have reduced clonal growth compared to uninfected plants. Greater clonal growth may be an effective mechanism for host genotypes to persist and spread when seed production is prevented, as is the case with castrating pathogens like Epichloë glyceriae .     

The rise and fall of the mountain hare (Lepus timidus) during Pleistocene glaciations: expansion and retreat with hybridization in the Iberian Peninsula

The climatic fluctuations during glaciations have affected differently arctic and temperate species. In the northern hemisphere, cooling periods induced the expansion of many arctic species to the south, while temperate species were forced to retract in southern refugia. Consequently, in some areas the alternation of these species set the conditions for competition and eventually hybridization. Hares in the Iberian Peninsula appear to illustrate this phenomenon. Populations of Iberian hare (Lepus granatensis), brown hare (Lepus europaeus) and broom hare (Lepus castroviejoi) in Northern Iberia harbour mitochondrial haplotypes from the mountain hare (Lepus timidus), a mainly boreal and arctic species presently absent from the peninsula. To understand the history of this past introgression we analysed sequence variation and geographical distribution of mitochondrial control region and cytochrome b haplotypes of L. timidus origin found in 378 specimens of these four species. Among 124 L. timidus from the Northern Palaearctic and the Alps we found substantial nucleotide diversity (2.3%) but little differentiation between populations. Based on the mismatch distribution of the L. timidus sequences, this could result from an expansion at a time of temperature decrease favourable to this arctic species. The nucleotide diversity of L. timidus mtDNA found in Iberian L. granatensis, L. europaeus and L. castroviejoi (183, 70 and 1 specimens, respectively) was of the same order as that in L. timidus over its range (1.9%), suggesting repeated introgression of multiple lineages. The structure of the coalescent of L. granatensis sequences indicates that hybridization with L. timidus was followed by expansion of the introgressed haplotypes, as expected during a replacement with competition, and occurred when temperatures started to rise, favouring the temperate species. Whether a similar scenario explains the introgression into Iberian L. europaeus remains unclear but it is possible that it hybridized with already introgressed L. granatensis.     

Daytime habitat selection by introduced eastern cottontail Sylvilagus floridanus and native european hare Lepus europaeus in Northern Italy

We used radiotelemetry to investigate resting sites habitat selection by introduced eastern cottontail (Sylvilagus floridanus) and native European hare (Lepus europaeus) under sympatric conditions. We tracked 24 hares and 34 cottontails in a protected area of northwestern Italy. Hares were found in different sites every week, while cottontails used the same site for two weeks, and occasionally for longer. It is supposed that this periodic nest switching reduces the risk of predation and parasitism. Hares and cottontails forms were located in different habitats and characterized by dense vegetation cover near the ground. This cover increased from winter to summer in both species, while in autumn it continued to increase in cottontails only, and decreased in hares. Cottontails selected shrubby habitats near the river, and avoided crop fields in all seasons. Hares were more adaptive in their search, using high herbs and shrubs all year round, wheat fields in spring, maize in spring and summer, and stubbles in winter. Arguably, partial niche differentiation is necessary to allow the coexistence of similar species. In our study area, hares and cottontails differentiated in the use of resting sites habitats, presumably so as not to compete in this part of their ecological niche.