Diet selection by European hares (Lepus europaeus) in the alpine zone of the Snowy Mountains, Australia

European hares (Lepus europaeus) are grazers and open grassland specialists that are replaced in mountain areas of their natural range in the northern hemisphere by browsing/intermediate feeding mountain hares (Lepus timidus), but in their introduced range in the southern hemisphere, occupy the alpine zone. We used micro-histological identification of plant fragments and germination of seeds in faecal pellets of L. europaeus from the Snowy Mountains, Australia, to determine diet. We asked whether diet shifted and/or diet breadth expanded in response to seasonally reduced food availability, particularly during winter. If so, did the constraints of food availability in the alpine zone lead to the diet mirroring that of L. timidus in its native alpine habitat. The diet of L. europaeus was dominated by grasses, herbs and shrubs. The main diet items in summer were grasses (70 %) and herbs (28 %). Grasses declined in the diet between summer and autumn when herbs increased to co-dominance, with a further change after establishment of the winter snowpack to a greater preponderance of shrubs (43 % compared with a maximum of 3 % in snow-free months). L. europaeus selected a wider range of plants in winter (59 species compared with 39 in summer) and diet was significantly more variable in winter than in autumn or summer (and in autumn than summer). We concluded that the persistence of L. europaeus in alpine areas of the southern hemisphere is testament to their ability to expand their dietary breadth to occupy mountain climatic zones normally occupied by L. timidus.     

Intensity of livestock grazing in relation to habitat use by brown hares (Lepus europaeus)

There is very limited information concerning livestock (sheep and goats) and brown hare Lepus europaeus interaction when both coexist. The effect of the intensity of livestock grazing on seasonal habitat use by hares, in a typical Mediterranean rangeland, was evaluated using the pellet-count method. Lightly grazed pastures were less preferred by hares compared with moderately grazed ones, whereas ungrazed pastures were used less intensively than grazed ones. Because livestock grazing reduces the quantity of standing biomass proportionally to its grazing intensity, forage resource was not the driving force for pasture selection. The increased use of moderately grazed pastures by hares in relation to lightly and ungrazed ones, where vegetation was more abundant, could be attributed to their reduced herbage height and density. This behaviour is probably a tactic that hares follow for predator avoidance, because they are more likely to detect visually approaching predators when feeding in a biotope with a limited herbaceous layer. The conclusion of this research is that livestock and brown hare coexistence may be compatible and beneficial rather than competitive when stocking rates do not exceed grazing capacity, leading to the conclusion that proper livestock grazing and hare population management can be feasible in practice.     

Biochemical genetic variability in brown hares (Lepus europaeus) from Greece

Allozyme variability of 91 brown hares (Lepus europaeus) from seven regions in Greece was compared to existing data of Bulgarian populations to test the hypothesis of the occurrence of specific alleles in Greece, likely stemming from an isolated Late Pleistocene refugial population in the southern Balkans. This hypothesis is particularly suggested by some subfossil Late Pleistocene hare remains in Greece and the reported high mtDNA diversity in Greek hares. Allozymic diversity could be higher in Greek hares than in hares from neighboring regions as a result of the accumulation of variants in a long-lasting Pleistocene refugium. Conversely, Greek hares could exhibit reduced genetic diversity because of long-lasting low effective population sizes during the Late Glacial Maximum and a lower chance of postglacial gene flow from other populations into this rather marginal part in the southern Balkans. Horizontal starch gel electrophoresis of proteins from 35~loci revealed three alleles (Es-1 ^–162, Pep-2 ¹¹⁴, Mpi ⁸⁸) at low frequencies, which were not found in Bulgarian or any other brown hare population. In contrast, some alleles from the populations from Bulgaria and other regions of Europe were absent in the Greek samples. Population genetic statistics indicated only a slight tendency of increased gene pool diversity in Greek hares, little substructuring in Greek and Bulgarian populations, respectively, as well as an only slightly lower level of gene flow between the two neighboring regions, as compared to the gene flow within each region. The results conform to the hypothesis of a Late Pleistocene refugial population in the southern Balkans, with some few specific nuclear gene pool characteristics, but little effect on the overall genetic differentiation between Greek and Bulgarian hares.     

Excretion of corticosteroids in urine and faeces of hares (Lepus europaeus)

Increased production of glucocorticoids by the adrenal cortex is found in mammals under stress. As cortisol itself is absent in the faeces, an enzyme immunoassay (11-oxoaetiocholanolone) measuring 11,17-dioxoandrostanes has already been established to measure faecal cortisol metabolites in ruminants for non-invasive monitoring of adrenocortical activity. The aim of this study was to establish route and delay of excretion of glucocorticoids in hares and to determine whether a cortisol-, corticosterone- or this new enzyme immunoassay is best suited to detect faecal glucocorticoid metabolites. In the first experiment radioactive-labelled glucocorticoids (¹⁴C-cortisol and ³H-corticosterone) were administered intravenously to two groups of three hares in metabolic cages. All voided urine and faecal samples were collected for 4 days. Metabolites of both steroids were found predominantly in the urine (91 ± 4%). Peak concentrations were observed in the first urinary sample following infusion (13 ± 6 h) and in the faeces with a delay of about 1 day (23 ± 7 h). Most of the radioactivity was not extractable with diethylether, indicating that the metabolites excreted in urine and faeces are mainly conjugated or polar unconjugated ones. This was confirmed by reverse-phase high-performance liquid chromatography separations of the metabolites, which also revealed marked differences concerning the metabolism of the two glucocorticoids injected. Compared with the cortisol and the corticosterone enzyme immunoassay, only the group-specific enzyme immunoassay for 11,17-dioxoandrostanes detected high quantities of immunoreactive metabolites. In a second experiment hares (n=20) were stressed by rousing them three times (5 min, 10 min and another 5 min) with a 20-min break in-between. Faecal samples were collected 2 days before until 4 days after stress and analysed using the 11-oxoaetiocholanolone enzyme immunoassay. After stress significantly (P < 0.001) increased 11,17-dioxoandrostane concentrations were found. Based on these results, measuring 11,17-dioxoandrostanes in faeces enables non-invasive monitoring of disturbances in hares and thus provides a tool for field investigations elucidating the role of stress in hare populations. Accepted: 24 November 1999     

Sexing European rabbits (Oryctolagus cuniculus), European brown hares (Lepus europaeus) and mountain hares (Lepus timidus) with ZFX and ZFY loci

We describe a new method for the sex determination of tissue originating from Oryctolagus cuniculus (European rabbit), Lepus europaeus (European brown hare) and Lepus timidus (mountain hare) based on PCR-RFLP analysis of point mutations that differentiate the ZFX and ZFY gene sequences. Among several applications, this PCR-RFLP method could be used to investigate gender ratio and evaluate the population dynamics of these species using samples collected when sex cannot be identified.     

Taxon 20 (Fam. Pasteurellaceae) infections in European brown hares (Lepus europaeus).

Hemolytic bacteria, phenotypically related to organisms previously identified as Pasteurella haemolytica and tentatively named Taxon 20, were isolated from cases of purulent bronchopneumonia and from conjunctivitis in European brown hares (Lepus europaeus). The bronchopneumonia, sometimes accompanied by lesions in other organs, occurred without other concomitant disease. The conjunctivitis was found mainly in animals suffering from the European brown hare syndrome.     

Evidence of autumn reproduction in female European hares (Lepus europaeus) from southern Europe

Information on reproductive biology of the European hare (Lepus europaeus) in different environmental and landscape conditions comprises part of fundamental knowledge regarding species’ adaptive responses as well as many aspects of its biology. Most of the studies conducted on European hare reproduction are confined to midlatitude and northern populations, whereas no data exist on the indigenous southern populations. Here, we present information on reproductive characteristics of European hares inhabiting Mediterranean ecosystems on the island of Crete, Greece for two successive hunting seasons. Although the annual reproductive cycle of the species is well known, with an autumn sexual inactivity, the duration of this period is subjected to fluctuations in different years and for different areas. According to our data, hare populations of Crete present an autumn–early winter reproductive activity with high proportions of pregnant females observed in all the months of the study. Furthermore, the estimated mean litter size (1.54 SE ± 0.07) while signed to the lowest values ever observed for European hares is similar to values obtained in continuous breeding species of the same genus, Lepus granatensis, Lepus corsicanus, Lepus (capensis) mediterraneus, and Lepus capensis also inhabiting warm climates. In conclusion, our results suggest that Cretan European hare populations exhibit a reproductively active period during autumn–early winter where proportions of pregnant females and litter size give a strong indication of a continuous reproduction throughout the year.     

High incidence of missing posterior upper molars in hares (Lepus europaeus) in New Zealand

Abstract

Of 140 skulls, 11 (7.9%) had one or both M³s reduced or missing. Males and females were affected, in both the North Island and the South Island.     

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.     

Annual patterns in reproduction and perirenal fat of hares (Lepus europaeus) in sub-alpine Canterbury, New Zealand

The aim of this paper was to describe how the age, sex and reproductive state of hares affected the amount of fat found around their left kidney. A total of 2208 hares ( Lepus europaeus ) was shot and autopsied over 13 years from sub-alpine grasslands in Canterbury, South Island, New Zealand. Adult females may be pregnant from July to mid-March. Adult males' testis weights were highest from July to January, and some fertile males may have been present in all months but April. The amount of perirenal fat in adults increased in autumn, peaked in early winter, and decreased thereafter to low levels through spring and summer. Autumn fat deposition in males was not clearly related to reproductive state in individuals, although it coincided with seasonal testicular development. Autumn fat deposition in females may have been caused in part by the cessation of lactation, but, like that in males, the increase was also part of an endogenous pattern. Fat utilization in winter was related to breeding. In males, utilization coincided with reproductive activity and the amount of perirenal fat remained consistently low throughout the breeding season. In females, net fat utilization was associated with lactation but this was balanced to some extent by fat deposition induced by pregnancy. Females simultaneously lactating and pregnant were in finer balance between utilization and deposition. Other factors, such as the number of embryos and the stage of gestation, altered the fat balance in these hares. Fat was also deposited as a 'yolk-sac' in embryos.     

Pituitary and ovarian responses to luteinizing-hormone-releasing hormone during pregnancy and after parturition in brown hares (Lepus europaeus)

Female hares were given an i.v. injection of 5 micrograms luteinizing-hormone-releasing hormone (LHRH) between Days 7 and 19 (n = 21), 20 and 33 (n = 17) and 34 and 41 (n = 17) of pregnancy, and in the 3 days after parturition (n = 16). Whatever the stage of pregnancy, the LHRH injection induced a release of luteinizing hormone (LH) and follicle-stimulating hormone (FSH) and an acute secretion of progesterone; these hormonal responses increased significantly during pregnancy, to reach values similar to those observed in nonpregnant, nonpseudopregnant females during the breeding season in the 3 days after parturition. However, the release of LH remained monophasic in pregnant and post-partum females, in contrast to the unmated females during the reproductive season, in which there was a biphasic profile. The proportion of ovulating females after LHRH treatment was approximately 60% at the beginning and end of pregnancy; and, after parturition, fell to 23% between Days 20 and 33. After Day 33, the pituitary response to LHRH was significantly higher in ovulating than in nonovulating females. At the beginning of pregnancy, 67% of females aborted after LHRH injection; after Day 20, the incidence of abortion decreased significantly and was 0% from Day 34. The amplitude and duration of progesterone secretion by the new corpora lutea resulting from ovulation after LHRH injection were similar to those of corpora lutea induced in nonpregnant females during the breeding season.(ABSTRACT TRUNCATED AT 250 WORDS)     

Influence of management regime and population history on genetic diversity and population structure of brown hares (Lepus europaeus) in an Italian province

In many areas, the management of overexploited populations of brown hare (Lepus europaeus) is based on annual restocking. While in some cases exotic hares are introduced, in some others hares are captured locally within protected areas and subsequently released into hunting grounds. We evaluated the genetic effects of this management regime in an Italian province where the brown hare population has recovered in the last few decades, by sequencing the hypervariable domain 1 of the mitochondrial control region and by genotyping eight autosomal microsatellites in hares sampled in both hunting and non-hunting areas. Both nuclear (H _e?=?0.68 and H _o?=?0.65) and mitochondrial variability (h?=?0.853 and π?=?0.012) were in line with other European populations. When comparing our data with mitochondrial sequences retrieved from GenBank, out of the 21 detected haplotypes, 14 were private to our study area. While 4.6 % of the individuals were found to carry haplotypes attributable to past introductions, 41.5 % grouped within a well-supported lineage, previously identified with a presumed native Italian taxon, L. e. meridiei. Despite the detectable geographic partitioning of mitochondrial haplotypes across the province, no genetic structure resulted from microsatellites analysis, indicating that no reproductive barriers exist among hares carrying different mitochondrial lineages. In conclusion, the local management seems to have contributed to the recovery of the species and to a full admixture of nuclear genes in the province. However, neither the extensive translocations nor the possible introductions of exotic heads seem to have completely undermined the local mitochondrial lineages.     

Epizootiologic and ecologic investigations of European brown hares (Lepus europaeus) in selected populations from Schleswig-Holstein, Germany

From 1997-99 European brown hare (Lepus europaeus) population densities were estimated by spotlight surveys within different areas in Schleswig-Holstein, Germany. These areas showed a wide variation in local hare population densities. In addition, red fox (Vulpes vulpes) densities were estimated in 1997 by surveys of fox dens and litters. Sera of 321 hares (shot between 1998-2000) from four study areas were examined for antibodies against European brown hare syndrome virus (EBHSV) by enzyme linked immunosorbent assay (ELISA), Yersinia spp. (n = 299) and Francisella tularensis (n = 299) by western blotting, Brucella spp. by Rose Bengal test, and Toxoplasma gondii by Sabin-Feldman test (n = 318). Tissue samples comprising lung, liver, spleen, kidney, heart, and adrenal glands were collected for histopathology. Liver (n = 201) and spleen (n = 201) samples were processed for the detection of T. gondii-antigen in tissue sections and 321 liver and spleen samples were investigated for EBHSV-antigen by ELISA. Furthermore, 116 hares were examined macro- and microscopically for lungworms. Significant negative correlations between hare and fox densities were found in spring and autumn 1997. Antibodies against EBHSV were detected in 92 of 321 (29%), against Yersinia spp. in 163 of 299 (55%), and against T. gondii in 147 of 318 (46%) hares. We evaluated the potential influence of origin and hunting season on exposure rates of hares using logistic regression analysis. A strong association between hare densities and exposure rates was observed for various agents. One hundred and eight of 201 (57%) hares were positive for T. gondii-antigen. All sera were negative for antibodies against Brucella spp. and F. tularensis and all lung samples were negative for lungworms. In conclusion, variation in red fox densities may have an impact on the hare populations examined and the infectious diseases we studied seem to play a subordinate role in the dynamics of European brown hare populations from Schleswig-Holstein.     

Purifying selection shaping the evolution of the Toll-like receptor 2 TIR domain in brown hares (Lepus europaeus) from Europe and the Middle East

Molecular Biology Reports - Toll-like receptors (TLRs) are transmembrane proteins of the innate immune system, composed of the ectodomain involved in pathogen recognition and the intracellular...     

Differences in body mass, health status and genetic variation between insular and mainland brown hares (Lepus europaeus) in Sweden

Introduced populations can be affected by random processes such as genetic drift, deterministic processes given by the local environmental conditions and anthropogenic factors such as hunting and management. Geographically constrained populations are particularly exposed to these processes, and altogether, these factors may result in rapid differentiation from the ancestral populations. The introduced European brown hare (Lepus europaeus) population on the island Ven is isolated from the Swedish and Danish mainland. Undocumented observations suggest that the hares on the island have been increasingly diseased in recent years and also decreased in body size. To test the substance of these observations, as well as the potential for inbreeding depression in this geographically constrained population, a total of 321 hares from Ven and three reference populations on the Swedish mainland were analysed with respect to body mass, general health status and genetic variation. The results confirm that the hares on Ven have lower body mass than hares on the mainland, but there are no indications of health deficits. We argue that the difference in body mass primarily is an island effect of stress and/or nutritional shortage, possibly induced by high population density, anthropogenic selection regimes and absence of mammalian meso-predators. Further, the genetic data indicate that the insular population is substructured, and subadults from these two subpopulations differ in body mass. This apparent substructuring could be due to chance effects, but may also be related to assortative mating or presence of sustained populations with different ancestry.     

Scale-dependent habitat selection by wintering geese: implications for landscape management

The management of Arctic migrant geese is complex, because they frequently use landscapes under intensive human use, and are conflict species in multiple respects. Some populations are of high conservation concern, but they also cause agricultural damage, are quarry for hunters, and may be particularly sensitive to infrastructure developments. In Bulgarian Dobrudzha, large wintering populations of greater white-fronted geese Anser albifrons and red-breasted geese Branta ruficollis feed in agricultural land, and cause management dilemmas. We developed linear models to investigate fine- and meso-scale foraging habitat selection of geese foraging on winter wheat in the area, and used these models to make suggestions for zoning landscape use in order to reduce conflict and conserve geese. Habitat selection was scale-dependent. Geese selected fields that were near to major roosts and had low proximity to roads and tree-lines, which may be a proxy for hunting disturbance. We found some evidence for selection of wheat fields with high nutritional quality. Within fields, geese strongly avoided features which cause landscape ‘clutter’: power-lines, tree-lines and wind-turbines, but primarily over distances of less than a few hundred metres. Optimal management might involve encouraging goose populations to feed in areas close to roosts, by means of agri-environmental measures and creation of hunting-free refuges. This would allow efficient use of agri-environment funds, might reduce conflict with farmers, and would mean that infrastructure development—notably wind farms—could be sited at greater distance from roosts with relatively minor impact on foraging habitat availability.     

Evolutionary relationships among hares from North Africa (Lepus sp. or Lepus spp.), cape hares (L. capensis) from South Africa, and brown hares (L. europaeus), as inferred from mtDNA PCR-RFLP and allozyme data

Systematics and taxonomy of hares of the genus Lepus (Lagomorpha) are under contentious debate, and phylogenetic relationships among many taxa are not well understood. Here we study genetic differentiation and evolutionary relationships among North African hares, currently considered subspecies of Lepus capensis , cape hares ( L. capensis ) from the Cape province in South Africa, and brown hares ( L. europeaus ) from Europe and Anatolia, using maternally (mtDNA) and biparentally (allozymes) inherited markers. A polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) analysis of a c. 1.8 kb long segment of the mitochondrial control region using eight hexanucleotide-recognizing restriction endonucleases yielded 28 haplotypes, and horizontal starch gel electrophoresis of proteins encoded by 25 structural gene loci revealed 52 alleles at 18 polymorphic loci. Diverse phylogenetic analyses (neighbor joining dendrogram, median joining network, multidimensional scaling of pairwise distances, AMOVA, F -statistics, hierarchical F -statistics) of genetic variants revealed marked substructuring of mtDNA into three phylogeographic groups, namely an African, a central European, and an Anatolian, but a somewhat less pronounced overall differentiation of the nuclear genome, despite a relatively high number of population-specific (private) alleles. However, all our results are not incongruent with Petter's (1959: Mammalia 23 , 41; 1961: Z. f. Säugetierkunde 26 , 30; 1972 : Société Des Sciences Naturelles et Physiques du Maroc 52 , 122) hypothesis that North African hares generally belong to L. capensis and that brown hares should be included in this species as well.