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This is the first study on spatial behaviour of brown hares Lepus europaeus Pallas, 1778 based on radio-telemetry in a natural system, which we contrast with data from agricultural systems. Radio tracking took place in a Dutch salt marsh over a 10-month period, with intensive tracking sessions during April/May and December/January. Six hares could be followed in both periods and in total 1224 fixes were collected. Average home range size was calculated as 28.7 ± 8.5 ha when using Adaptive Kernell method (Mimimum Convex Polygon: 27.3 ± 9.0 ha) on 90% of all fixes. Such values are in the lower end of the range of those obtained for agricultural systems. Home range size did not differ between sexes, day and night, or across seasons. However, the size of the core range (50% of fixes) was twice as large in May compared to the winter period, and thus inversely related to food availability. Unlike in agricultural systems, use of space by hares did not change over the course of the season. This probably reflects the patchy nature of the natural habitat which provides food and shelter throughout the year in a confined area.
We have studied fluctuating asymmetry (FA), as indicator of developmental stability, and between-individual variation, as surrogate of developmental canalization (DC), in long bones (humerus, ulna, radius, femur, tibia) of 72 wild-living adult-sized brown haresLepus europaeus Pallas, 1778 with variable individual heterozygosity (H).H was calculated from 13 polymorphic allozyme loci. According to the “over-dominance hypothesis”, we expected increased developmental stability and canalization at higherH-levels. But at the individual level we did not find any significant correlation between overall FA (FAI) andH. Also, standard deviations (SD) of mean length (over both body sides) of bones did not differ between individuals from two intentionally created groups of hares, namely one with high and one with lowH. FA-indices and variances of FA-indices of bone lengths did not differ significantly when compared between two intentionally created groups of hares with high and low SD of bone lengths, respectively. These latter findings suggest that developmental stability and DC are two separate or partly separate mechanisms of developmental homeostasis in the studied appendicular skeleton, and thatH has no traceable effect on develop-mental homeostasis. If there is still such an effect, it should be clearly smaller than a possibly combined effect of (presently uncontrolled) environmental stressors.
In 1987 - 1990 an investigation was made on the spatial structure of brown hare Lepus europaeus Pallas, 1778 populations using the transect routes method. The in­vestigation was carried out in two different parts of Poland that differed in the charac­ter of agriculture: 1 - northeastern with large fields of state farms, and II - east- central with small fields of individual farmers. The results indicate that independently of the agrarian structure, the distribution of hares in crops corresponds to their per­centage share in the region, with exception of rape fields, which show a tendency to be avoided. There was also a significant relationship between the spatial distribution of the hare and the habitat diversity - highly variegated fields are more consistently inhabited by hares. Besides, hares show a distinct tendency to prefer areas close to the field margins. Positively oblique distance distributions between the individuals and the margins of field crops are particularly strongly expressed in the region of a highly monotonous crop species. It may be expected that a change in type of farming by passing to the large-fields and continuous cropping is one of the factors that make it difficult for the hare to utilize the space within the habitat.
We describe the results of our research on population dynamics among brown hares reared in enclosures and then released into suitable natural habitat. Radio-tracking was used to follow the fate of 60 released brown hares over a 4-year period, extending between November 2005 and November 2009. The survival rate among these animals after 12 months was estimated to be 37 %, with 22 tagged individuals surviving beyond 1 year post-release. The highest (40 %) level of mortality characterised the first month after release, while a second period of enhanced mortality coincided with the breeding season (altogether accounting for a 20 % mortality rate). There was no significant relationship between body mass and mortality rate in the first month following release. A natural cause of death was predation by mammals, which accounted for some 31 % of all losses. Remaining causes were poaching (13 %), hits by vehicles (7 %) and unidentified causes (9 %). However, in at least 40 % of cases, it was not possible to determine the date when a released animal died, to say nothing of the cause of death.
The relationships between the density of brown hare Lepus europaeus Pallas, 1778 and landscape structure were analysed on the basis of data collected for 11 study areas between 1991 and 1994, and hunting reports from all the 49 provinces of Poland covering the period 1981-1995. The average autumn density of hares in study areas (8-28 ind/km2, estimated by strip census) decreased with the number of forest edges and increased with the number of permanent cover areas per km of transects. In the provinces of the country, the hare abundance index (hunting bag, ranging from 0.07 to 5.18 ind/km2) decreased with both the proportion of large fields and forests in all the years. The negative effects of large fields and forests on the hare abundance index increased in the study period.
A total of 193 brown hares, collected from 7 sampling sites in Poland during 1986 - 1990 were examined for genetic variability and differentiation at 39 presumptive isozyme loci by means of horizontal starch gel electrophoresis. Values of polymorphism (mean P = 0.180, SD 0.039) and average heterozygosity (mean H = 0.047, SD 0.006) were similar to those detected in previous studies on the population genetics of the brown hare. Relative (Cst = 0.041) and absolute (mean D/Nei, 1978/ = 0.0012, SD 0.0013) genetic differentiation among populations were very low, which fits well to the high number of migrant individuals per generation (Nm = 12.7), estimated using the private allele method of Slalkin (1985). Average heterozygosity was examined for associations with geographical distribution, the year of culling, population density, age, sex, body weight and health status, whereby a better survival of heterozygous femals could be detected. According Lo our results, the present decline of the brown hare is noL due to genetic depiction. However, once population sizes drop below a critical threshold, a pronounced inbreeding depression can be expected.
Testicular activity in brown hare Lepus europaeus Pallas, 1778 was studied during the annual cycle. Testicular mass, spermatozoa and testosterone were estimated. Percentages of haploid, diploid and tetraploid cells were monitored using DNA flow cytometry and the proportions of somatic and spermatogenetic cells were determined after selective labelling of somatic cells. Testis mass was high from January to July and declined thereafter to the nadir in September. Testis growth was reactivated signifi­cantly in December. Changes in testis mass corresponded with the spermatogenic efficacy (spermatozoa/g testis). High spermatogenic activity was characterized by intensive meiotic transformation of spermatocytes to spermatids, high percentages of haploid cells and low proportions of cells in the G2/M phase of mitosis. Proportions of haploid cells declined rapidly during the testis involution. Spermatogenesis was newly activated from November. The proportions as well as the ratios of spermatogenic and somatic cells differed significantly between the periods of testis involution and recru­descence, Testosterone level showed a pronounced increase in autumn preceding the intensification of spermatogenesis; the lowest concentration was found during prominent testis involution in August. The results suggest that the regulation of seasonal testicular activity is characterized by co-ordinated shifts in the relationships between mitosis, meiosis and testosterone production.
Allozymic variability of 157 brown hares Lepus europaeus (Pallas, 1778) from eight sampling localities in Bulgaria was studied by horizontal starch gel electrophoresis of 50 putative structural gene loci. The present data were compared to an adjusted data set of brown hares from central Europe (Austria), published earlier, in order to test the hypothesis of increased genetic diversity in brown hares from the southeastern European zoogeographical crossroads region, a possible refuge for brown hares during the last glaciation period. No new polymorphic locus was detected, and only one new allele occurred at a low frequency in the Bulgarian hares. Nevertheless, two estimators of genetic diversity (average expected heterozygosity, Shannon-Weaver information index) where slightly higher in Bulgarian hares than in central European hares. In contrast, some alleles found earlier in various parts of central Europe did not occur in the Bulgarian samples. The latter finding suggests that gene pools of central European brown hares might have received gene flow from other parts of Europe during their postglacial (re-(colonization process. This accords to an alternative hypothesis of postglacial colonization of central Europe from both southeastern and southwestern refuges.
Mandibles of 258 brown hares Lepus europaeus Pallas, 1778 from Poland were controlled for abnormal expression of the premolars. In four individuals a reciprocal transposition of the right and left third lower premolars was found. In addition, all the affected premolars were rotated. The rotation showed bilateral symmetry in each of the four specimens. A genetical basis is suggested for this type of tooth anomaly.
Habitat management should be an important part of the brown hare (Lepus europaeus) conservation, but the habitat requirements of this species are not fully recognised. The aim of our research was to estimate these requirements by analysing the effect of various agricultural landscape structure features on the distribution of hares in five agricultural areas in Germany and Poland. The local density of hares was assessed in the spring and autumn of 2006 by using the method of spotlight–strip counts on 9–15 subareas in each research region. The structure of agricultural landscape has been described for each subarea: the share of grain, other crops and grasses as well as the density of crop edges and uncultivated places with wild vegetation. The density of hares was considerably higher in Germany than in Poland (18.8–48.4 vs. 4.1–9.5 indiv./km2). The hare density was positively correlated with non-grain crops in an area, with crop edges in two areas and with wild vegetation without trees in two areas, and negatively correlated with grassfields in two areas. The occurrence of wild vegetation without trees affected the hare density only in the study areas, where this habitat was relatively rare (<3 km/km2). It was suggested that proper projects aimed at habitat management for brown hares should be elastic, i.e. the projects should be modified depending on the structure of local landscapes. Moreover, the protection and creation of structures with wild vegetation among cropland seem to be considerable methods of brown hare or generally wildlife conservation; therefore, such measures should be an important part of agro-environmental packages.
Control of red fox Vulpes vulpes populations is a fundamental game management tool used by hunters interested in increasing prey populations. In Italy, a popular method to control fox populations is hunting with small hound packs. The effects of this hunting technique on non-target species such as the brown hare Lepus europaeus, are unknown. In this study, we analysed for the first time the effects of fox hunting with hound packs on brown hares tagged with VHF collars. Our results showed that hunting with four trained hounds did not significantly modify the spatial behaviour of the brown hare.
Two contrasting hypotheses on the relationship between dental character variability and biochemical-genetic diversity: (1) "influence of developmental homeostasis" and (2) "genetic-phenetic variation correlation" were tested in brown hare Lepus europaeus Pallas, 1778 populations. Interindividual variability (IV) and fluctuating asymmetry (FA) of 12 non-metrical characters of third lower premolars (P3) as well as allozymic heterozygosity (H) at 13 polymorphic loci was examined in 385 individuals from 19 geographical sampling units (GSU) in Austria. Juveniles and adults were discriminated according to dry eye lens weights. Sex was determined from internal reproductive organs. IV was calculated as the mean standard deviation of the 11 tooth characters in each GSU. GSU-specific FA was calculated as the mean FA of individuals (FAin), where FAin was the percentage of characters found asymmetric in individuals of a GSU. While IV did not show any significant relationship with H at the population level, FA of adults was significantly positively correlated (rs = +0.650, p < 0.05) with H. In juveniles a trend (rs = +0.399, ns) for such a correlation was apparent too. This finding corresponds to the "genetic-phenetic variation correlation hypothesis". Variability of both character systems is high in populations with high genomic variability, because both character systems concordantly portray gene pool diversity. Both IV and FA was significantly lower in juveniles than in adults. Since no ontogenic changes in P3 characters were found, this age-specific difference appears to result from selection against juveniles with low P3 variability (i.e. low genomic diversity). However, H was not lower in juveniles as compared to adults.
A total of 469 brown hares Lepus europaeus Pallas, 1778 from 20 sampling sites in Austria were examined for genetic diversity within and among populations by means of horizontal starch gel electrophoresis. Fourteen out of 54 presumptive structural loci were polymorphic, one of which was excluded from further population genetic analyses due to the occurrence of a null-allele. The mean proportion of polymorphic loci (P) was 15.3% (SD 2.2%), and mean expected average heterozygosity (He) was 4.6% (SD 0.5%). Both relative (Fst = 5.4%) and absolute (mean Nei's 1978 D = 0.0016, SD 0.0016) genetic differentiation among populations were low, suggesting a generally high level of migration. Cluster analysis revealed some separation of brown hare populations in western and northern Austria from those in the east and in the south. In 131 individuals, mtDNA was digested with a battery of 16 restriction endonucleases. Besides the standard type I which occurred exclusively in most of the populations, five additional haplotypes, each of them deviating from type I by one base pair substitution, were detected. Together with rare alleles at allozyme loci, the distribution of variant haplotypes corroborated the spatial pattern obtained by allozyme distances and suggested considerable immigration of brown hares from the adjacent countries in the east and south. Twenty non-metric skull traits were scored in 443 individuals. Character variants were dichotomized (0/1) and the respective frequencies were used to calculate C. A. B. Smith's 'mean measure of divergence' (MMD) among five population groups. Morphological differentiation was in accordance with the major population genetic pattern as revealed by molecular techniques. MtDNA variation (nucleon diversity, nucleotide diversity) and morphological variation (mean of SD in single characters) within populations were not significantly associated with one another, and did not show a relationship with indices of genetic variation obtained by allozyme analysis. These findings suggest that variability in only one of these characters cannot be considered representative for overall gene pool diversity within populations.
The content of iron, zinc, copper, manganese, lead, and cadmium was determined in the kidney, liver, and abdominal muscle of the brown hare (Lepus europaeus), a species known for its bioindicative potential in environmental quality assessment. The animals assayed were divided into immature (in their first year of life; n=25) and adult (n=39). The hares were acquired in an area situated far away from major cities and direct impacts of industrial and transport pollution. The zinc content found in the assayed hare's muscles (25.9 μg/g w.w.) and that of manganese in the kidneys, liver, and muscles (2, 2.51 i 0.85 μg/g w.w., respectively) tended to be higher than those reported from the brown hare elsewhere in Europe. The content of cadmium (particularly in the kidneys) and lead proved substantial and close to those typical of the hare inhabiting industry-affected areas. Analysis of the data pooled for all the individuals showed Zn and Cd contents to increase with age in the kidneys, the liver Cd content increasing with age as well; on the other hand, the age-Cu content was negative. In addition, a number of significant correlations between the metals themselves were revealed, particularly with respect to Zn-Cd correlation in kidneys (rs=0.47), Fe-Cd, Zn-Cu, Mn-Zn, Mn-Cu, and Mn-Pb in the liver (rs of 0.42, 0.86, 0.72, 0.79, and 0.42, respectively), and Zn-Cu in the muscle (rs=0.56). The kidney cadmium content was higher by 81% in the adult than in the immature hare, the adult hare muscle copper content being lower by 15.5%.
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