Lepus timidus, Linnaeus, 1758
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publication ID |
https://doi.org/ 10.5281/zenodo.6625539 |
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DOI |
https://doi.org/10.5281/zenodo.6625488 |
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persistent identifier |
https://treatment.plazi.org/id/03822308-B761-FFD1-FAFC-F9FFFC16F4AB |
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Carolina |
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scientific name |
Lepus timidus |
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59. View On
Mountain Hare
French: Liévre variable / German: Schneehase / Spanish: Liebre de montana
Other common names: Blue Hare, Snow Hare, Eurasian Arctic Hare, Varying Hare, White Hare; Irish Hare (hibernicus), Alpine Mountain Hare (varronis)
Taxonomy. Lepus timidus Linnaeus, 1758 View in CoL ,
“in Europa.”
Formerly, the three arctic species, L. timidus , L. arcticus , and L. othus , were included in L. timidus based on morphological characteristics that, relative to their distributions, form a circumpolar “ring species.” This is also supported by genetic analysis of mtDNA, although evidence based only on mtDNA should be treated cautiously. There is also the view that two species exist: L. timidus in the Old World and L. arcticus in Greenland, northern Canada, Alaska, and the Chukchi Peninsula, Russia. Other lagomorph taxonomists consider that L. arcticus is conspecific with L. timidus and distinct from L. othus . Until conclusive evidenceis available, the three species are considered to be distinct with L. timidus in the Old World, L. othus in Alaska, and L. arcticus in northern Canada and Greenland. Nevertheless, the border between L. timidus and L. othusis not clear and might be either in the Bering Strait or in the Kolyma region, Russia. Recent molecular phylogenetic study suggests that the break occursin the Bering Strait, but more studies are required for a conclusive demarcation because the study was only based on mtDNA. Six populations of L. timidus are geographically separated and morphologically distinct and therefore can be treated as subspecies (abe, ainu, hibernicus, orii, scoticus, and varronis). Because division into subspecies is mostly based on pelage characteristics, it must be considered preliminary. Three studies analyzing geographic variation of metric characters of the skull of L. timidus demonstrated clinal variations. This doubts the advisability of identifying subspecies in L. timidus . Origin of subspecies of L. timidus in Europe from a panmictic population during the Weichsel period is supported by genetic assessments. Hybridization between L. timidus and L. europaeus and between L. timidus and L. tolai are recorded in contact zones. No interbreeding has been noted between scoticus and hibernicus on the island of Mull where both subspecies were introduced 60 years ago. Sixteen subspecies recognized.
Subspecies and Distribution.
L. t. timidus Linnaeus, 1758 — Scandinavia S to 59° N, N Russia (S to 57° N to 58° N) to E of Ural Mts, Estonia, perhaps Poland, and Altai and Beita Mts in N Xinjiang (China); from Sweden extending in hybrid zones with the subspecies sylvaticus to S coast of Norway and W Latvia.
L. t. abet Kuroda, 1938 — Kurile Is (Russia).
L. t. ainu Barrett-Hamilton, 1900 — Hokkaido (Japan).
L. t. begitschevi Koljuschev, 1936 — Taimyr Peninsula (Russia).
L. t. gichiganus J. A. Allen, 1903 — C Yakutia, Kamchatka and Okhotsk Sea coast (Russia).
L. t. hibernicus Bell, 1837 — Ireland.
L. t. kolymensis Ognev, 1922 — NE Siberia (Russia).
L. t. kozhevnikovi Ognev, 1929 — C Russia from 57° to 58°N, whereit intergrades with the nomonotypical timidus S to 53° N. The W boundary is unclear.
L. t. lugubris Kastschenko, 1899 — Siberian Altai Mts (Russia).
L. t. mordeni Goodwin, 1933 — Russian Far East (Ussuri Valley and lower and middle Amur River) and Heilongjiang (China).
L. t. ori Kuroda, 1928 — Sakhalin (Russia).
L. t. scoticus Hilzheimer, 1906 — Scotland.
L. t. sibiricorumJohansen, 1923 — W & SW Siberia (Russia), N Kazakhstan, and Tacheng Mts in N Xinjiang (China).
L. t. sylvaticus Nilsson, 1831 —S Sweden, extending in hybrid zones with nominotypical timidus to S coast of Norway and W Latvia.
L. t. transbaikalicus Ognev, 1929 — Trans-Baikal region in E Siberia (Russia) and E Inner Mongolia (= Nei Mongol, China).
L. t. varronis Miller, 1901 — at high altitudes in the Alps, C Europe. View Figure
The Mountain Hare inhabits the Palearctic from Scandinavia to Russian Far East, except E Chukchi Peninsula, S to Baltic States, E Poland, N Ukranie, and E to Tian Shan Mts, Altai Mts, N Mongolia, NW & NE China (N Xinjiang, N Inner Mongolia, Heilongjiang), and Sikhote-Alin Mts. There are isolated populations in Scotland, Ireland, the Alps, Sakhalin (Russia), Hokkaido (Japan), and the Kurile Is. The nominotypical subspecies has been introduced into Faroe Is (Denmark), Scottish Is, and parts of England. Introduced populations on Spitsbergen I failed to persist.
Descriptive notes. Head-body 510-550 mm, tail 59-65 mm, ear 90-100 mm, hindfoot 159-165 mm; weight 2.4-3.4 kg (depending on subspecies). The smallest subspecies is the “Alpine Mountain Hare” (varronis), and the largest subspecies is the nominate timidus . Male Mountain Hares tend to be smaller than females. In general, northern populations are larger. The Mountain Hare is completely white in winter, and its head and body are brown in summer. Summer fur differs among populations and can be pale gray or dusky brown in mountainous areas to rich rustbrown on plains. Exceptionally, fur of the Irish Hare is white on rump, flanks, and legs, with its upper back and head remaining brown in winter. Ears are short and narrow, with 30mm black tips on exterior surfaces and 5mm on interior surfaces. Relatively large head has prominent eyes, with yellowish irises in adults but dark brown in young. Hindfeet have hairy soles and widely spreading toes. Molt occurs three times per year: white to brown in spring; brown to brown in late summer; and brown to white in autumn. Molt is initiated by daylength, and its duration is influenced by temperature. Mountain Hares at high elevations and latitudes remain white longer, relative to average ambient temperature and duration of snow cover.
Habitat. Wide range of biomes from extreme tundra with permafrost in northern Russia to rich agricultural areas in Ireland. Mountain Hares prefer to live in forests of pine, birch, and juniper. In Scotland and Ireland, heather moors and bog land are favored, and in Russia, copses in the middle of open steppe and reed belts around lakes are selected. In the Alps, Mountain Hares prefer highly structured forests and thickets at forest edges. In spring, abundant herbaceous food in meadows is important to Mountain Hares. When a habitat offers food and shelter against predators, Mountain Hares use it for rest and activity. Forms are usually a depression they make. In snow, Mountain Hares burrow to make forms and enter rock crevices or marmot (Marmota) burrows for shelter or escape. They sometimes dig permanent burrows 1-2 m long in the ground. Mountain Hares in Ireland and the Alps do not burrow, but in Russia, complex burrows up to 7 m long have been reported. On agricultural land, Mountain Hares tends to be replaced by European Hares ( L. europaeus ) in Europe and Cape Hares ( L. capensis ) further east.
Food and Feeding. Mountain Hares are grazers in summer and browsers in winter. Their diets vary with habitat. Plant species used as food in forest habitats include leaves and twigs of Salix and Populus (both Salicaceae ), Sorbus (Rosaceae) , Betula (Betulaceae) , Juniperus (Cupressaceae) , and Vaccinium (Ericaceae) ; on Scottish moorlands and Swedish islands, Calluna (Ericaceae) is eaten. Different grass species and clovers are eaten when available. With snow cover, Mountain Hares feed on twigs, bark, moss, and lichens. They seldom drink water but may eat snow. Re-ingestion ofsoft pellets takes place at 09:00-16:00 h and increases dietary digestibility up to 25%; these pellets contain three times as much protein as regular pellets. Mountain Hares feed with their back to the wind and can clear snow from vegetation with their forepaws. When snow is hard, they move to lower ground to feed.
Breeding. Gestation of the Mountain Hare lasts 46-55 days. Postpartum estrus with copulations a few hours after parturition is common. Superfetation does occur. Females have predominantly 2-3 litters/year, with litter sizes of 1-3 young, resulting in 5-9 offspring/female during a reproductive season. Weather has an important influence on numberof litters produced during a reproductive season.
The second litter is on average larger than the first one. Start of breeding varies with temperature, and gestating females have been recorded from January/February until August/September. Juvenile females do not breed in the same year of their birth. Differences in reproduction exist among populations or subspecies.
For example, subspecies timidus in northern Norway normally has only 1-2 litters/ year, but in south-western Sweden,it regularly has three litters. Another difference is the length of the reproductive period with the Irish Hare having a longer one than scoticus. At birth, young are fully furred, their eyes are open, and they start suckling immediately. Adult sex ratio is 1:1. Overall juvenile survival from birth to the following spring averages 20%. In January-July, male Mountain Hares often follow 2-20 m behind a female for hours. Males use scent to track females. During courtship, several males chase a female, but males do not guard females. Females dominate males and strike with their forepaws with ears laid backward if the male approaches too closely. Copulation can involve five or more males and a female, and males can be involved in long fights.
Activity patterns. Mountain Hares are nocturnal throughout the year, with a prolonged resting phase during the day. Activity patterns change seasonally with circadian rhythms. Mountains Hares rest by day in a form, with ears back and eyes half closed, but they only sleep for a few minutes at a time. In summer when daylight increases and in winter when food is scarce, more Mountain Hares are active during the day. Activity is highest during the first two hours after sunset.
Movements, Home range and Social organization. Mountain Hares move an average of 1400 m/mnight, but they can travel up to 2400 m/night. Home rangesizes vary depending on climate. Home ranges are ¢.200 ha in boreal coniferous forests of Scandinavia. In moorland and pastures of Scotland and Ireland, respectively, and in the Alps, home ranges are 5-435 ha. Home ranges at night are larger than during the day, and they overlap ¢.35%. Densities vary from 1 ind/km? in Russia to 400 ind/km? on the Swedish Island of Vedholmen. Massive migrations of groups of 100-500, but up to 1500-2000 hares have been recorded in years of high numbers in northern Russia (Barents and Kara seas, Yamal Peninsula, Yakutia). Mountain Hares often move, rest, and feed together in groups of 2-5 individuals. Groups of 20-100 individuals have been recorded feeding in the same place.
Status and Conservation. Classified as Least Concern on The IUCN Red List. The Mountain Hare is widespread, populations appear to be stable across much ofits distribution, and it occurs in many protected areas. Nevertheless, historical game bag records in Northern Ireland indicate that there has been a substantial decline in abundance. Moreover, historical and recent records indicate that distribution of Mountain Haresis decreasing in southern Sweden. This decline started simultaneously with introduction of European Hares and continues while they expand northward. Furthermore, hunting statistics from Grisons, Switzerland, indicate a possible shift of both species to higher elevations. From 1990 to 2010, European Hares and Mountain Hares annually ascended more than 6 m and 3 m, respectively. Results of the study showed that European Hares advanced faster to higher elevations than Mountain Hares. A possible threat to the Irish Hare might be an introduced and expanding population of the European Hare in Ireland. The two examples from Sweden and Switzerland suggest that European Hares benefit from global warming and expand their distribution northward and to higher elevations and, thereby, might displace the Mountain Hares from their current distribution. Competitive exclusion and hybridization and introgression might explain how the European Hare is involved in the continuous decline of the Mountain Hare. Comparison of distributions of European Hares and Mountain Hares in Europe reveals that wherever European Hares occur, Mountain Hares are restricted to higher elevations or deep forests. It seems that European Hares and Mountain Hares are ecologically equivalent and strongly compete with one another. Studies of food preferences show that Mountain Haresin Ireland eat more grasses and herbs than Mountain Hares that live in sympatry with European Hares. Consequently,it has been argued that competitive exclusion by the European Hare restricts Mountain Hare distribution. Besides, in sympatric areas the forms of Mountain Hares are located in denser forest and further away from open fields than in allopatric areas. European Hares also expand their distribution and use of various habitats when densities of Mountain Hare are low. All these studies show that European Hares have the potential to expand into Mountain Hare territory and,in sympatry, Mountain Hares disappear from optimal European Hare habitat. European Hares and Mountain Hares hybridize with each other. Offspring are morphological intermediate and fertile. Normally, a male European Hare will mate with a female Mountain Hare. Reciprocal hybridization is also possible, but very rare due to typical mating behavior of European Hares and Mountain Hares in natural surroundings. Each time a Mountain Hare female hybridizes with a European Hare male, the local Mountain Hare population loses a species-specific litter. Consequently, observed decline in distribution and densities of Mountain Hares may be a direct consequence of unidirectional hybridization. This phenomenon is described as extinction by hybridization in the literature.
After hybridization, mtDNA of Mountain Hares can be detected in European Hares if the hybrids backcross with true European Hares, which is often the case. Therefore, introgressed individuals can be found in both species. In areas where European Hares live currently in sympatry with Mountain Hares, the proportion of European Hares with Mountain Hare mtDNA is higher than in areas where both species lived in sympatry in the past. It seems that European Hares with Mountain Hare mtDNA have a reduced fitness compared with European Hares with species-specific mtDNA. This means that Mountain Hare mtDNA is incorporated into European Hares in sympatric areas but gradually disappears if European Hares remain allopatric. After the introduction of European Hares in Sweden, European Hares and Mountain Hares occurred together for a period of time before Mountain Hares disappeared and only European Hares remained. Hybrids were only observed a couple of years before they disappeared. Accordingly, hybridization, introgression, and the eventual disappearance of Mountain Hare mtDNA might be regarded as refined competitive exclusion in which traces of Mountain Hares are not only ecologically but also genetically erased from the European Hare. In addition to interactions with the European Hare, other causes for the Mountain Hare decline are under consideration, such as diseases (e.g. European Brown Hare Syndrome and tularemia), deforestation and agricultural expansion, predation, and negative interactions with other herbivores. Increasing tourism in the Alps has a negative impact on Mountain Hares and other biota. Nevertheless, none of these factors alone can explain declines of Mountain Hares. Research is needed to identify causes of regional decline.
Current conservation status of the endemic Irish Hare and the Alpine Mountain Hare require specific investigations.
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