Lepus Timidus Research Articles

Effect of landscape type, elevation, vegetation period, and taxonomic plant identification level on diet preferences of Alpine mountain hares (Lepus timidus varronis)

Diet is important for understanding the ecology and evolution of populations. When examining animals’ diets, the lowest taxonomic rank (i.e., species level) is generally used. However, it is questionable whether dietary description varies with respect to the plant taxonomy or with the extent of plant species diversity in landscapes. We studied the diet of the Alpine mountain hare (Lepus timidus varronis) by analyzing the botanical composition of fecal pellets and comparing the results with the availability of food plants. Overall, Alpine mountain hares selected ligneous plants, whereas grasses were neutrally consumed or avoided. Plant identification at the genus and family level revealed the most information, whereas little additional information was gleaned by identifying food plants at the species level. In low-diversity spruce forests and at higher elevations, hares exhibited greater diet breadth compared to high-diversity vegetative mosaics or at lower elevations, even if they had access to the same types of plants. This is likely due to the hares’ need to diversify their diet when plant quantity is low. Greater diet breadth and selection of a larger number of different plant types at the beginning of the plant vegetation growth period suggested that more food plants suitable for Alpine mountain hares were available during this time compared to the end of the vegetation period.

Distribution of mountain hares Lepus timidus in Scotland in 2016/2017 and changes relative to earlier surveys in 1995/1996 and 2006/2007

Mountain hares Lepus timidus, incorporating the subspecies L. t. varronis, L. t. hibernicus and L. t. scoticus, have declined in range throughout continental Europe where they face pressures from climate-change, competition and land-management. In Scotland, the absence of a national monitoring scheme, including mandatory reporting of hunting records, means that producing robust estimates of mountain hare population trends is difficult. We repeated questionnaire surveys conducted in 1995/1996 and 2006/2007 to assess the 2016/2017 distribution and hunting records of mountain hares in Scotland and describe regional changes in their distribution over a 20-year period in relation to management for red grouse Lagopus lagopus scotica shooting. Comparisons of areas covered in all surveys indicated no net change in the area of Scotland occupied by mountain hares, but within that we found changes in range between regions and sites of differing grouse management intensity. Between 1995/1996 and 2016/2017, range contractions in southern Scotland contrasted with no changes in north-–east Scotland. In north-west Scotland range expanded by 61% in areas practicing driven grouse shooting, declined by 57% in areas practicing walked-up grouse shooting and remained low and stable in areas which did not shoot grouse. A total of 33 582 mountain hares were killed in 2016/2017 representing a 71% and 48% increase from 1995/1996 and 2006/2007 respectively. However, the average kill density in 2016/2017 (12.4 ± 3.3 hares km–2) was comparable to 2006/2007 (10.8 ± 3.0 km–2) and we found no relationship between kill density and contractions in range. Despite increases in numbers of mountain hares killed over the last 20 years, it appears that range contraction may be attributed to factors other than culling, such as changes in habitat and management. Disentangling these factors should be the focus of future research.

Quantification of sex-related diet composition by free-ranging mountain hares ( Lepus timidus )

Quantification of sex-related diet composition by free-ranging mountain hares ( Lepus timidus )

Weak Habitat Segregation between Male and Female Mountain Hares (Lepus timidus)

Sexual habitat segregation affects animal distribution and can lead to different life-histories across sexes. We investigated sex-related habitat segregation in the mountain hare (Lepus timidus) during the early-breeding and post-reproductive periods at the macro- (home range) and microscale (pellet location) by using pellet data sets from a non-invasive genetic population monitoring in the Swiss Alps. The data sets comprise six years (2014–2019) of sampling and include 119 individuals (70 males, 49 females). At the macroscale, the sex-related habitat segregation was weak in both periods but higher in the early-breeding period as compared with that in the post-reproductive period. Home ranges of females contained a higher proportion of forest stands in the early-breeding period. At the microscale, the sex-related habitat segregation for habitat characteristics was low in both periods. We conclude that habitat segregation between male and female mountain hares is weak during the early-breeding and post-reproductive periods.

Cases of melanism in mountain hare (Lepus timidus) in Yakutia

Cases of melanism in mountain hare (Lepus timidus) in Yakutia

Lagovirus europeus GI.2 (rabbit hemorrhagic disease virus 2) infection in captive mountain hares (Lepus timidus) in Germany

BackgroundRabbit hemorrhagic disease virus (RHDV, Lagovirus europeus GI.1) induces a contagious and highly lethal hemorrhagic disease in rabbits. In 2010 a new genotype of lagovirus (GI.2), emerged in Europe, infecting wild and domestic population of rabbits and hares.Case presentationWe describe the infection with a GI.2 strain, “Bremerhaven-17”, in captive mountain hares (Lepus timidus) in a zoo facility in Germany. Postmortem examination revealed RHD-like lesions including necrotizing hepatitis. RT-qPCR and AG-ELISA confirmed presence of GI.2. Recombination and phylogenetic analysis grouped the identified strain with other GI.2 strains, sharing nucleotide identity of 91–99%.ConclusionOur findings confirm that mountain hares are susceptible to GI.2 infection, due to a past recombination event facilitating virus spillover from sympatric rabbits.

Identifying the accidental‐natural mortality of leporids in the archaeological record: insights from a taphonomical analysis of a pitfall without evidence of human presence

ABSTRACTLeporids (Lagomorpha, Mammalia), especially the mountain hare (Lepus timidus) and the European rabbit (Oryctolagus cuniculus), have been among the most abundant small game in western Europe since the Pleistocene. These animals are preferred prey for more than 40 predators, which increases the chances of finding their skeletal remains at archaeological or natural sites, and in particular karstic formations alternately occupied by Palaeolithic human societies and other predators. Moreover, specific eco‐ethological characteristics for both species also offer the possibility to produce a substantial quantity of their bones accumulated by natural mortality without predation, making it more difficult to identify their origin in the fossil record. Despite this fact, the taphonomic signature of accidental‐natural accumulations of leporids in karsts, such as pitfalls, has never been properly characterised. In order to address these issues, we carried out a detailed taphonomic study of leporid remains in a pitfall without any evidence of human activity, namely at Coulet des Roches (Monieux, Vaucluse, south‐eastern France). At this site, leporids are the most abundant species throughout the Last Glacial Maximum sedimentary sequence. The bone accumulation was analysed in order to determine the most relevant criteria to identify a natural accumulation that results from the accidental fall of individuals in the pitfall. The relative proportions of each species are consistent with their current ethology, and their mortality profiles are compatible with the structure of a living natural population. The completeness rate of the different skeletal portions for both species is relatively high, bones are less fragmented and, unlike what might be expected, very few anatomical connections were preserved. The observed anatomical representation, breakage, and bone surface modification patterns are discussed and compared with available data concerning accumulations with predation. Taken together, the identified criteria contribute valuable help in the recognition of accidental mortality in the fossil record, as well as to evaluate the proportion of leporid bones accumulated naturally and those accumulated by predation, human or not, in a deposit.

The complete mitochondrial genome sequence of Lepus tolai in Xinjiang

Lepus tolai has a wide geographic distribution in the world, and there are some taxonomic confusions due to lack of studies. The complete mitochondrial genome of L. tolai in Xinjiang is 17,047 bp long with 13 protein-coding genes (PCGs), 22 tRNAs, two rRNAs, and a control region. The overall base composition is 31.81% of A, 29.40% of T, 25.67%of C, and 13.12% of G, showing AT-rich. Phylogenetic analysis results showed that L. tolai is closely related to Lepus timidus, and distantly related to Lepus earoupenses and Lepus capensis. The new mtDNA data will supplement the mitochondrial gene pool of Lepus and helpful for validating taxonomic status of L. tolai on mtDNA level.

First record of Lepus timidus (Linnaeus, 1758) from Novaya Zemlya, Russian Arctic (Lagomorpha, Leporidae)

Mountain hare (Lepus timidus) is a widespread species that lives throughout the continental tundra zone of Eurasia, as well as over the greater part of the northern forest zone. For a long time, the presence of the mountain hare in the fauna of the Novaya Zemlya Archipelago was assumed. However, documented evidence from the archipelago was lacking. In 2017, two fragments of the lower jaw of a single specimen were found in the Bezymyannaya Bay area on Yuzhny Island, Novaya Zemlya, Russia. This record confirms the presence of the species on the archipelago, while its status remains unclear. The jaw fragments from Novaya Zemlya contained dry meat remnants, indicating that this individual died relatively recently (i.e., no more than 10 years ago). It is likely that the species periodically visits the archipelago, without forming a viable population there.

Welfare Implications for Hares, Lepus timidus hibernicus, Taken from the Wild for Licensed Hare Coursing in Ireland.

Simple SummaryHare coursing is a widespread and controversial activity in the Republic of Ireland in which the speed and agility of greyhounds are measured against that of a hare. Every year, several thousands of hares are taken from the wild by coursing clubs under licenses issued by the National Parks and Wildlife Service. While the majority are returned to the wild at the end of the coursing season, no research has been done on the impact of hare coursing on the welfare of individual hares. Despite greyhounds in licensed coursing events being muzzled since 1993, hares may be pinned to the ground by the dogs and killed or so severely injured that they have to be euthanized by a veterinary practitioner. In addition to animal welfare concerns, the recent emergence in Ireland of rabbit haemorrhagic disease (RHD2) has led to calls for licensed hare coursing to be prohibited on animal welfare, disease control, conservation and ethical grounds. In this paper, publicly available information provided by licensed hare coursing clubs on the number of hares captured, used for coursing events, killed or injured and the percentage returned to the wild, over four coursing seasons is reviewed. A reported 19,402 hares were taken from the wild over the four coursing seasons. Whilst 19,080 hares (98%) were returned to the wild, 75 were killed by greyhounds or had to be euthanized due to their injuries. Policy makers should take animal welfare, disease control, conservation, ethics and public opinion into account and fund independent research where gaps in knowledge are identified.Hare coursing is legal in the Republic of Ireland under licenses issued to coursing clubs but is illegal in other jurisdictions in the British Isles including Northern Ireland. Supporters of coursing maintain that coursing contributes to the conservation of the hare whilst opponents claim that coursing is cruel and the welfare of the hares is compromised. However, while the contribution of coursing to conservation has been considered, the impact of coursing on hare welfare has not been investigated. This paper reviews publicly available information from licensed hare coursing clubs over four coursing seasons, including the number of hares taken from the wild, numbers coursed, and numbers pinned to the ground by dogs, killed or injured during coursing events. In total, 19,402 hares were taken from the wild—98% of which were subsequently released back to the wild at the end of the coursing season. Almost 600 hares were pinned by greyhounds during coursing events and 75 were either killed or had to be euthanized as a result of their injuries. While the number of hares killed or injured is relatively small compared to the number caught, the welfare of all captured hares will have been compromised and has not been investigated. Policy makers must fill this knowledge gap or take a precautionary approach and further regulate or indeed prohibit the capture of hares which are otherwise fully protected.

Transcriptomic regulation of seasonal coat color change in hares.

Color molts from summer brown to winter white coats have evolved in several species to maintain camouflage year‐round in environments with seasonal snow. Despite the eco‐evolutionary relevance of this key phenological adaptation, its molecular regulation has only recently begun to be addressed. Here, we analyze skin transcription changes during the autumn molt of the mountain hare (Lepus timidus) and integrate the results with an established model of gene regulation across the spring molt of the closely related snowshoe hare (L. americanus). We quantified differences in gene expression among three stages of molt progression—“brown” (early molt), “intermediate,” and “white” (late molt). We found 632 differentially expressed genes, with a major pulse of expression early in the molt, followed by a milder one in late molt. The functional makeup of differentially expressed genes anchored the sampled molt stages to the developmental timeline of the hair growth cycle, associating anagen to early molt and the transition to catagen to late molt. The progression of color change was characterized by differential expression of genes involved in pigmentation, circadian, and behavioral regulation. We found significant overlap between differentially expressed genes across the seasonal molts of mountain and snowshoe hares, particularly at molt onset, suggesting conservatism of gene regulation across species and seasons. However, some discrepancies suggest seasonal differences in melanocyte differentiation and the integration of nutritional cues. Our established regulatory model of seasonal coat color molt provides an important mechanistic context to study the functional architecture and evolution of this crucial seasonal adaptation.

When to count? Indices of population size for mountain hares of the north-west Lammermuir Hills, Scotland

Surveys with binoculars at two different times of day and in various months of the year, along with camera trap studies of diel activity patterns, were used to inform an appropriate period to count mountain hares ( Lepus timidus) on managed red grouse (Lagopus lagopus ) moorland of the Lammermuir Hills, south-east Scotland. Factors affecting the numbers of hares counted were time relative to sunrise, the presence of winter coat colour and reproductive behaviour in spring. Counts of hares in March and April starting one hour before dawn were used as an index of population size of the mountain hare over three years of observations on three hill-tops, with densities of 23-33 hares km-2. The number of hares seen was stable or rose slightly over the three years despite a partial cull on one of the hills. In support of suitability of the timing of surveys used, camera trap studies revealed that the period around dawn in March and April was associated with high levels of hare activity.

Distribution of the Mountain Hare (Lepus timidus ainu) during the Snow Season in the Central Tokachi District, Hokkaido

Distribution of the Mountain Hare (<i>Lepus timidus ainu</i>) during the Snow Season in the Central Tokachi District, Hokkaido

An Annotated Draft Genome of the Mountain Hare (Lepus timidus).

Hares (genus Lepus) provide clear examples of repeated and often massive introgressive hybridization and striking local adaptations. Genomic studies on this group have so far relied on comparisons to the European rabbit (Oryctolagus cuniculus) reference genome. Here, we report the first de novo draft reference genome for a hare species, the mountain hare (Lepus timidus), and evaluate the efficacy of whole-genome re-sequencing analyses using the new reference versus using the rabbit reference genome. The genome was assembled using the ALLPATHS-LG protocol with a combination of overlapping pair and mate-pair Illumina sequencing (77x coverage). The assembly contained 32,294 scaffolds with a total length of 2.7 Gb and a scaffold N50 of 3.4 Mb. Re-scaffolding based on the rabbit reference reduced the total number of scaffolds to 4,205 with a scaffold N50 of 194 Mb. A correspondence was found between 22 of these hare scaffolds and the rabbit chromosomes, based on gene content and direct alignment. We annotated 24,578 protein coding genes by combining ab-initio predictions, homology search, and transcriptome data, of which 683 were solely derived from hare-specific transcriptome data. The hare reference genome is therefore a new resource to discover and investigate hare-specific variation. Similar estimates of heterozygosity and inferred demographic history profiles were obtained when mapping hare whole-genome re-sequencing data to the new hare draft genome or to alternative references based on the rabbit genome. Our results validate previous reference-based strategies and suggest that the chromosome-scale hare draft genome should enable chromosome-wide analyses and genome scans on hares.

Using by‐catch data from wildlife surveys to quantify climatic parameters and timing of phenology for plants and animals using camera traps

AbstractGaining a better understanding of global environmental change is an important challenge for conserving biodiversity. Shifts in phenology are an important consequence of environmental change. Measuring phenology of different taxa simultaneously at the same spatial and temporal scale is necessary to study the effects of changes in phenology on ecosystems. Camera traps that take both time‐lapse as well as motion‐triggered images are increasingly used to study wildlife populations. The by‐catch data of these networks of camera traps provide a potential alternative for measuring several climatic and phenological variables. Here, we tested this ability of camera traps, and quantified climatic variables as well as the timing of changes in plant and animal phenology. We obtained data from 193 camera‐unit deployments during a year of camera trapping on a peninsula in northern Sweden aimed at studying wildlife. We estimated daily temperature at noon and snow cover using recordings provided by cameras. Estimates of snow cover were accurate, but temperature estimates were higher compared with a local weather station. Furthermore, we were able to identify the timing of leaf emergence and senescence for birches (Betula sp.) and the presence of bilberry berries (Vaccinium myrtillus), as important food sources for herbivores. These were linked to the timing of the growth of antlers and the presence of new‐born young for three ungulate species as well as the presence of migratory Eurasian cranes (Grus grus). We also identified the timing of spring and autumn moulting of mountain hares (Lepus timidus) in relation to snow cover. In this novel study, we show the potential of (by‐catch) data from camera traps to study phenology across a broad range of taxa, suggesting that a global network of camera traps has great potential to simultaneously track wildlife populations and the phenology of interactions between animals and plants.

Home Ranges of Semi-Urban Brown Hares (Lepus europaeus) and Mountain Hares (Lepus timidus) at Northern Latitudes

Many species shift their distribution northwards due to changing climate, which may bring previously isolated species into contact. In some cases, this can lead to competition and crossbreeding among congeneric species. Due to milder winters, several hare species (Lepus spp.) have shifted poleward or to higher elevations and currently occur in sympatry. There are limited data on the spatial ecology of northern brown hares (L. europaeus), and on possible joint space use with mountain hares (L. timidus), although species are known to frequently crossbreed. We estimated home ranges of 12 brown and 12 mountain hares in eastern Finland using GPS telemetry. The total home-range median areas were 111 ha for brown hares and 105 ha for mountain hares. The median core areas of these species were 5 and 3 ha, respectively. Individual home ranges and core areas of the studied species overlapped. Interspecific interaction seems to be frequent during the breeding season, which also enables crossbreeding.

Vocalisation in the mountain hare: calls of a mostly silent species

While the occurrence and function of vocalisation by pikas has been studied (genus Ochotona), little is known about vocalisation by hares (genus Lepus). In this study, we investigate the vocalisation activity of mountain hare (Lepus timidus) over the course of a year to better understand its occurrence and function. We found that mountain hares have a low vocalisation rate of one vocalisation every 33 night hours. We hypothesise that this low rate is the result of a strategy to minimise detection by predators. However, the 20 recorded vocal events correspond with intraspecific communication events such as warning others about danger and defending resources. This case study identifies further research directions to help strengthen our understanding of communication between solitary hares.

Introgression drives repeated evolution of winter coat color polymorphism in hares

Changing from summer-brown to winter-white pelage or plumage is a crucial adaptation to seasonal snow in more than 20 mammal and bird species. Many of these species maintain nonwhite winter morphs, locally adapted to less snowy conditions, which may have evolved independently. Mountain hares (Lepus timidus) from Fennoscandia were introduced into the Faroe Islands in 1855. While they were initially winter-white, within ∼65 y all Faroese hares became winter-gray, a morph that occurs in the source population at low frequency. The documented population history makes this a valuable model for understanding the genetic basis and evolution of the seasonal trait polymorphism. Through whole-genome scans of differentiation and single-nucleotide polymorphism (SNP) genotyping, we associated winter coat color polymorphism to the genomic region of the pigmentation gene Agouti, previously linked to introgression-driven winter coat color variation in the snowshoe hare (Lepus americanus). Lower Agouti expression in the skin of winter-gray individuals during the autumn molt suggests that regulatory changes may underlie the color polymorphism. Variation in the associated genomic region shows signatures of a selective sweep in the Faroese population, suggesting that positive selection drove the fixation of the variant after the introduction. Whole-genome analyses of several hare species revealed that the winter-gray variant originated through introgression from a noncolor changing species, in keeping with the history of ancient hybridization between the species. Our findings show the recurrent role of introgression in generating winter coat color variation by repeatedly recruiting the regulatory region of Agouti to modulate seasonal coat color change.

Petrography and provenance of floor sediments from the Loutra Almopias Cave (Pella, Macedonia, Greece)

Thirty seven floor sediment samples of Upper Pleistocene age from the Loutra Almopias Cave were collected from different beds and stratigraphic columns on the basis of their induration grade, grain distribution, and paleontological findings. Channel facies make up the bulk of the clastic sediments found in the cave passages. Slackwater facies compose the final layer of all the stratigraphic sections of the examined cave. The floor sediments are mineralogically immature, since they contain many ferromagnesian minerals, feldspars (especially plagioclase) and quartz. The extensive presence of silicate minerals means that the phyllites, gneisses, schists, ophiolitic rocks and the clastic Mariam Formation of the Almopia Zone are the main detrital load source, along with the flysch of the neighboring Pelagonian Zone. The presence of sand-sized grains, pebbles and cobbles of dolomitic or calcitic composition also designates the carbonate rocks of the Almopia and Pelagonian Zones as primary sources. The dolomite and calcite content of secondary chemical origin in the cave sediments is very limited. Kutnohorite, isomorphous with dolomite, was found for first time in a Greek cave. The provenance of the sediments is mixed; they are composed mainly of the weathering materials of the Alpine metamorphic basement and the carbonate rocks outcropping adjacent to the cave. The sediments were transported and deposited inside the cave, after rapid weathering and erosion of the surrounding rocks, under a tectonically active regime. The mineralogical variation in the stratigraphic columns demonstrates variations in the clastic load, due to the different weathering intensity periods. Most of the sediments are fluvial deposits, and one is considered a glacial deposit. Fossils of Lepus timidus (mountain hare) found within the cave represent the southernmost record of this species in Europe. Its presence signifies a cool phase at the end of the last glacial period before the onset of the warm Holocene Epoch.

The complete mitochondrial genome sequence of Yarkand hare (Lepus yarkandensis)

We first reported the mitochondrial genome of Lepus yarkandensis. The mitogenome of L. yarkandensis contains 17,011 base pairs. The overall base composition of complete mitogenome is 28.13% A, 27.67% T, 22.02% C, and 22.17% G, with 44.20% of the GC content. All genes exhibit the typical mitochondrial gene arrangement and transcribing directions. Phylogenetic analysis of 9 Lepus species was performed based on the sequence of cytochrome b gene using the Maximum Likelihood method in MEGA 7.0. The results suggested that L. yarkandensis is closely related to Lepus timidus. The results are helpful to future studies on molecular evolution, population genetics, and wildlife protection of L. yarkandensis.