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Although weather-induced mass mortalities of wild ungulates have been reported, no study has quantified how these episodes may affect the survival of prime-aged adults. Long-term studies of marked ungulates have instead consistently found very weak or no effects of weather on the survival of this age class, particularly for females. We report on the effects of the exceptionally snowy winter of 2008–2009 on three populations of chamois in the western Alps: two in Italy, one in France. In the Alpi Marittime Natural Park in Italy, mortality of prime-aged females (aged 2–9 years) was 43%, about five times higher than reported by previous studies of chamois. Just across the continental divide in the adjacent Mercantour National Park (PNM) in France, however, prime-aged female mortality was only 6%. Senescent females suffered very high mortality in both populations (100% and 56%). In the Gran Paradiso National Park in Italy and in the Alpi Marittime Natural Park, adult male mortality rate was respectively of 81% and 44%, whereas in the PNM, it was only 10%. A recent reduction in population density in the French population, or lower absolute snowfall than in Italy, may explain the difference in survival. Survival of males and prime-aged females can be affected by exceptional weather events, possibly in combination with high population density. Adult chamois of both sexes appeared to show elevated mortality in response to harsh winter conditions. Our results underline the importance of considering sex and age classes in evaluating the impacts of population density in wild ungulates.
The herd size of Cantabrian chamois Rupicapra pyrenaica parva (Cabrera, 1910) varied seasonally in relation to escape terrain and food availability in our study area (Asturias, north of Spain). The median group size of females without kids was 1 (mean ± SD = 1.62 ± 1.00), females with kids was 4 (5.59 ± 5.42), males was 1 (1.73 ± 1.78), and mixed group size was 7 (8.91 ± 7.91). The female-kid group size depended more on escape terrain availability than on food quality. Throughout the early weeks of the life of kids, the mothers remained in difficult access areas (cliffs and steep slopes), and showed a weak tendency to aggregate. These areas provided a wide visual range and hiding places for offspring and their use may be an anti-predation strategy. When the kids were able to run quickly, the mothers used subalpine meadows. These areas were very open and exposed kids to predation and human disturbance, however the forage has high nutritive value, and may compensate for the cost of breeding and suckling by the mothers. Aggregation may be selected as an anti-predation strategy in subalpine meadows, allowing a reduction in time spent vigilant by each individual in the group, and increased time available for other activities. The largest male groups were located in pastures with abundant but poor quality forage. Our results suggest that group size in Cantabrian chamois is very flexible, changing seasonally and in response to resource availability.
Sexual dimorphism in Cantabrian chamois Rupicapra pyrenaica parua (Cabrera, 1911) horn length was recorded annually for the first five years of life. The horns of males were larger than those of females, though horn growth rate in the first two years (ie horn length in the first two years divided by horn length in the first five years) was greater in females than males. Sexual dimorphism in horn length of Cantabrian chamois adults was found to be the highest of genus according to available data. Males and females with faster horn growth in the first two years of life reduced their horn growth rate in the third and fourth year. Chamois' horns from the Eastern Cantabrian Mountains were smaller than those from Western Cantabrian Mountains, although at five years of age these differences were not evident. Total horn length and jaw length showed significant positive correlation (females: r^ = 0.75, males: rp2 = 0.54, p < 0.05 in both cases). This relationship suggests that horn growth dynamics may be used as an estimation of body-size dynamics. Horn growth / jaw length correlation was higher in females than in males. Horn growth was also positively correlated with annual precipitation in the previous year, but not with precipitation in the current year. Winter horn growth was observed in the younger age classes. Post-winter horn growth was recorded in 55% of the animals in March. The use of this information to age Cantabrian chamois is described.
Restriction fragment length polymorphisms (RFLPs) of mitochondrial (mt) DNA were used for investigating genetic differentiation in chamois (genus Rupicapra). Digestion of the mtDNAs of 58 individuals from 6 populations with a battery of 16 six-base cutting (restriction endonucleases yielded a total of 67 restriction sites. Based on the presence and absence of these restriction sites a total of 8 haplotypes could be defined. Six of them served for assessing genetic diversity within and among 4 local populations of R. rupicapra rupicapra. Estimates of nucleotide divergence among those haplotypes ranged from 0.05% to 0.25%. One chamois from the High Tatra (subspecies R. r. tatrica) was examined and showed the standard haplotype found in R. r. rupicapra. MtDNA in chamois from Catalunya, belonging to R. pyrenaica pyrenaica, was polymorphic for two haplotypes not found in any population of R. rupicapra. Mean nucleotide divergence among haplotypes found in R. rupicapra and R. pyrenaica was 0.56% (SD = 0.16%). Based on this value, an estimated divergence time of about 280 000 years suggests that the mtDNA lineages of R. rupicapra and R. pyrenaica separated prior to the Riss glacial in the later Pleistocene.
The relationships between availability and quality of food, diet composition and habitat use of female and male chamois herds Rupicapra pyrenaica parva (Cabrera, 1911), were investigated in the Cantabrian Mountains (Spain). The two vegetation types studied were grass-forb and shrub. Grass-forb contained higher crude protein and lower fibre than shrub throughout year (crude protein: 17.2% vs 8.6%, p = 0,018; acid detergent fibre: 27.0% vs 35.8%, p = 0.018). The diet of female and male herds showed more grass-forb than shrub (p = 0.012), however, a discrepancy between site selection and bite selection was found. Generally male herds were found in shrub areas although they mainly chose grass-forb in their diet. Female herds had a higher percentage of grass-forb in the diet than male herds throughout the year (81.6% vs 65.6%). Diet composition pattern was similar in both sexes, with a peak of grass-forb in spring and autumn, and a minimum in summer. Both sexes selecLed grass-forb throughout the year, but its use was increased when its quality was high, inde­pendently of quantity available. Females showed higher grazing activity than males throughout year (53% vs 37%, p < 0.0001). Male herds showed less grazing activity during the rut (October-November) in comparison with the rest of the year (25% vs 43%, p < 0.0001). Some hypotheses to explain the segregation of the herds of both sexes are presented and discussed.
Examination of 599 faecal samples taken over the years 1984-1993 from Alpine chamois in the Slovak Paradise National Park revealed the presence of lung nematodes Muellerius capillaris, Muellerius tenuispiculatus, Neostrongylus linearis and Dictyocaulus viviparus. Their total prevalence in chamois herds was 57.7%. Muellerius spp. were predominant (56.5%), N. linearis occurred in about half less amount (29.2%) and D. viviparus was prevalent in 2.5%. The prevalence of lung nematodes in chamois herds fluctuated with years from 41.4 to 69.4% and with seasons of the year from 33.3 to 83.3%. Mean larval count per gram faeces in individual years ranged between 73.7 ± 60.1 and 148.5 ± 58.0. Chamois herds were significantly less infected in summer seasons (P<0.05) than in other seasons, except for the summer of 1986, 1988, 1989 and winter of 1989. A necropsy confirmed the presence of adult stages of all three nematodes in the lung parenchyma, but in bronchi and trachea no adults of D. viviparus were observed. Mean larval count per gram lung tissue was high (152 ± 20.9 to 270 ± 19). The wide diversity of snail species in the territory of the Slovak Paradise, their great ability to serve as intermediate hosts of lung nematodes in various biotopes create appropriate conditions for the development of lung nematodes and consequently for the infection of chamois in the Slovak Paradise reserve.
Reproductive parameters, kidney fat index and grazing activity of both sexes of Cantabrian chamois Rupicapra pyrenaica parua (Cabrera, 1910) in relation to age, season and year are presented and compared with those of other chamois subspecies and cervids. Females younger than three years old had not ovulated. Ovulated ovaries were significantly heavier than non-ovulated ovaries in March-April, and these dif­ferences remained until the following rutting season. All pregnant females carried a single foetus. A significant decrease in the percentage of parous females was found in 3-11 years age class (94%) compared with > 11 years (50%). There were no differences in the kidney fat index (KFI) between age classes for either of the sexes throughout the study period. Females showed a significant higher KFI in August­-October 1993 than in August-October 1992, coinciding with a significantly greater abundance of grass in 1993. Throughout the year KFI in females showed less fluc­tuations than in mates. Males began the rutting season with a significant KFI three times higher than did females. However, by the spring, the KFI of the mates had fallen to half of that of the females. In March-April pregnant females showed a significantly higher KFI than non-pregnant or late-conception adult females. Feeding activity of females and sward height throughout the year were inversely related. Females grazed more actively than males in the summer, rut and spring, except in winter where no differences in feeding activity between sexes were found. The dif­ferences in the KFI and feeding activity are discussed in relation to the parental investment and the sexual dimorphism-body size theory.
Measures of horn growth of Bulgarian chamois Rupicapra rupicapra balcanica Bolkay, 1925 were compared between males and females. The hypothesis that rapid early growth of horn is followed by slower subsequent growth was tested through the regression of the horn increment in the first two years against the third, fourth and fifth year respectively in the same individual. Sexual dimorphism in annual growth increment was significant up to the third year, males having higher values than females. Negative correlation coefficients were found when regressing the third, fourth and fifth individual annual increment on the second one in males, thus showing that individuals which had horns which grew fastest in the first two years tended to have slower horn growth in subsequent years.
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