Two Different Epidemiological Scenarios of Border Disease in the Populations of Pyrenean chamois (Rupicapra p. pyrenaica) after the First Disease Outbreaks

Since 2001 several outbreaks of a new disease associated with Border disease virus (BDV) infection have caused important declines in Pyrenean chamois (Rupicapra pyrenaica) populations in the Pyrenees. The goal of this study was to analyze the post-outbreak BDV epidemiology in the first two areas affected by disease with the aim to establish if the infection has become endemic. We also investigated if BDV infected wild and domestic ruminants sharing habitat with chamois. Unexpectedly, we found different epidemiological scenarios in each population. Since the disease outbreaks, some chamois populations recuperated quickly, while others did not recover as expected. In chamois from the first areas, prevalence was high (73.47%) and constant throughout the whole study period and did not differ between chamois born before and after the BDV outbreak; in all, BDV was detected by RT-PCR in six chamois. In the other areas, prevalence was lower (52.79%) and decreased during the study period; as well, prevalence was significantly lower in chamois born after the disease outbreak. No BDV were detected in this population. A comparative virus neutralisation test performed with four BDV strains and one Bovine viral diarrhoea virus (BVDV) strain showed that all the chamois had BDV-specific antibodies. Pestivirus antibodies were detected in all the rest of analyzed species, with low prevalence values in wild ruminants and moderate values in domestic ruminants. No viruses were detected in these species. These results confirm the hypothesis that outbreaks of BDV infection only affect the Pyrenean chamois, although other wild ruminants can occasionally be infected. In conclusion, two different scenarios have appeared since the first border disease outbreaks in Pyrenean chamois: on the one hand frequent BDV circulation with possible negative impact on population dynamics in some areas and on the other, lack of virus circulation and quick recovery of the chamois population.     

Predicting seasonal and spatial variations in diet quality of Pyrenean chamois (Rupicapra pyrenaica pyrenaica) using near infrared reflectance spectroscopy

Monitoring nutritional status of wildlife is key to understanding the population response to changes in food availability. Several direct and indirect methods have been proposed for such purposes, but faecal nitrogen (FN) is by far the most commonly used indirect non-invasive indicator of free-ranging ruminant diet quality. In this work, two methods were used to estimate the FN content in 291 pellet samples of Pyrenean chamois (Rupicapra pyrenaica pyrenaica) collected during a 2-year period (May 2009 to May 2011) in two chamois populations from the Catalan Pyrenees, Spain: the Dumas LECO analyzer and near-infrared reflectance spectroscopy (NIRS). In order to optimize further FN estimations we performed three different NIRS calibrations: (1) general, across year, season and populations; (2) population-specific; and (3) season-specific, i.e., different periods of vegetation growth. FN ranged from 1.66 % to 2.85 % (dry matter basis) and was higher in the main vegetation growing season than in winter. Concerning NIRS calibration, our study confirmed that FN of Pyrenean chamois can be accurately determined by NIRS, since the general calibration was accurate enough to predict FN. It is concluded that NIRS represents a non-destructive, emission-free and low-cost analytical technique which may reduce the use of conventional laborious methods for estimating FN in long-term wildlife monitoring.     

A negative association between horn length and survival in a weakly dimorphic ungulate

While all models of sexual selection assume that the development and expression of enlarged secondary sexual traits are costly, males with larger ornaments or weapons generally show greater survival or longevity. These studies have mostly been performed in species with high sexual size dimorphism, subject to intense sexual selection. Here, we examined the relationships between horn growth and several survival metrics in the weakly dimorphic Pyrenean chamois (Rupicapra pyrenaica). In this unhunted population living at high density, males and females were able to grow long horns without any apparent costs in terms of longevity. However, we found a negative relationship between horn growth and survival during prime age in males. This association reduces the potential evolutionary consequences of trophy hunting in male chamois. We also found that females with long horns tended to have lower survival at old ages. Our results illustrate the contrasting conclusions that may be drawn when different survival metrics are used in analyses. The ability to detect trade‐off between the expression of male secondary sexual traits and survival may depend more on environmental conditions experienced by the population than on the strength of sexual selection.     

Vigilance behavior of Pyrenean chamois Rupicapra pyrenaica pyrenaica: Effect of sex and position in the herd

The Pyrenean chamois Rupicapra pyrenaica pyrenaica is a mountain-dwelling ungulate with an extensive presence in open areas. Optimal group size results from the trade off between advantages (a reduction in the risk of predation) and disadvantages (competition between members of the herd) of group living. In addition, advantages and disadvantages of group living may vary depending on the position of each individual within the herd. Our objective was to study the effect of central vs. Peripheral position in the herd on feeding and vigilance behavior in male and female Pyrenean chamois and to ascertain if a group size effect existed. We used focal animal sampling and recorded social interactions when a focal animal was involved. With males, vigilance rate was higher in the central part of the group than at the periphery, probably due to a higher density of animals in the central part of the herd and a higher probability of being disturbed by conspecifics. With females, vigilance rate did not differ according to position in the herd. Females spent more time feeding than males, and males showed a higher frequency of the vigilance behavior than females. We did not observe a clear relationship between group size and vigilance behavior. The differences in vigilance behavior might be due to social interactions.     

Diversity and evolution of the Mhc-DRB1 gene in the two endemic Iberian subspecies of Pyrenean chamois, Rupicapra pyrenaica

Major histocompatibility complex class II locus DRB variation was investigated by single-strand conformation polymorphism analysis and sequence analysis in the two subspecies of Pyrenean chamois (Rupicapra pyrenaica) endemic to the Iberian Peninsula. Low levels of genetic variation were detected in both subspecies, with seven different alleles in R. p. pyrenaica and only three in the R. p. parva. After applying the rarefaction method to cope with the differences in sample size, the low allele number of parva was highlighted. The low allelic repertoire of the R. p. parva subspecies is most likely the result of bottlenecks caused by hunting pressure and recent parasitic infections by sarcoptic mange. A phylogenetic analysis of both Pyrenean chamois and DRB alleles from 10 different caprinid species revealed that the chamois alleles form two monophyletic groups. In comparison with other Caprinae DRB sequences, the Rupicapra alleles displayed a species-specific clustering that reflects a large temporal divergence of the chamois from other caprinids, as well as a possible difference in the selective environment for these species.     

Dynamics of an Infectious Keratoconjunctivitis Outbreak by Mycoplasma conjunctivae on Pyrenean Chamois Rupicapra p. pyrenaica

Between 2006 and 2008, an outbreak of Infectious Keratoconjunctivitis (IKC) affected Pyrenean chamois Rupicapra p. pyrenaica, an endemic subspecies of mountain ungulate that lives in the Pyrenees. The study focused on 14 mountain massifs (180,000 ha) where the species’ population is stable. Cases of IKC were detected in ten of the massifs and, in five of them, mortality was substantial. The outbreak spread quickly from the first location detected, with two peaks in mortality that affected one (2007) and three (2008) massifs. In the latter, the peak was seasonal (spring to autumn) and, in the former, the outbreak persisted through winter. To identify the outbreak’s aetiology, we examined 105 Pyrenean chamois clinically affected with IKC. TaqMan rt-PCR identified Mycoplasma conjunctivae in 93 (88.5%) of the chamois. Another rt-PCR detected Chlamydophila spp. in 14 of chamois, and 12 of those had mixed infections with mycoplasmas. In the period 2000–2007, the chamois population increased slightly (λ 1.026) but decreased significantly during the IKC outbreak (λ 0.8, 2007–2008; λ 0.85, 2008–2009) before increasing significantly after the outbreak (λ 1.1, 2009–2010). Sex-biased mortality shifted the adult sex ratio toward males (from 0.6 to 0.7 males per female) and reduced productivity slightly. Hunting was practically banned in the massifs where chamois experienced significant mortality and allowed again after the outbreak ended. Long-term monitoring of wild populations provides a basis for understanding the impacts of disease outbreaks and improves management decisions, particularly when species are subject to extractive exploitation.     

Molecular identification of a new pestivirus associated with increased mortality in the Pyrenean Chamois (Rupicapra pyrenaica pyrenaica) in Spain

Pestivirus infection was identified in 16 of 17 chamois during an outbreak of a previously unreported disease in Pyrenean chamois (Rupicapra pyrenaica pyrenaica) in northeastern Spain in 2001-02. By analysis of the 5' noncoding regions of the virus, we assigned it to the border disease virus cluster with pairwise similarity values ranging from 82.1% to 88.1%. It will be important to investigate the association of this pestivirus with disease in Pyrenean chamois.     

Concentration of fecal cortisol metabolites in chamois in relation to tourist pressure in Tatra National Park (South Poland)

In protected areas, outdoor recreation, and nature-based tourism can act as potential stressors for wildlife. Given the growing demand for nature recreation, the consequences of high tourist visitation on wildlife need to be assessed in order to avoid incompatibilities between public use and species protection goals. The Tatra National Park (Poland), in the Carpathian Mountains, is a unique alpine ecosystem visited by three million tourists per year. It hosts the only native population of an endemic subspecies of chamois (Rupicapra rupicapra tatrica). We investigated the effects of tourist disturbance, the number of visitors, and the season on the concentration of fecal cortisol metabolites (FCM) in Tatra chamois in 2009. FCM levels of chamois were significantly higher and showed higher variation at high tourist disturbance (mean?±?SD, 46.2?±?31.53 ng/g, n?=?56) than at low disturbance sites (mean?±?SD, 17.2?±?8.05 ng/g, n?=?38). Stress levels increased with the number of visitors and therefore showed a peak in summer, coinciding with the highest number of visitation to the national park. A large portion of chamois habitat in Tatra National Park is within the area of influence of the touristic trail network. The temporal or permanent creation of areas free of disturbance in protected areas should be considered, especially in the periods of high tourist visitation. This study highlights the need to monitor the effects of tourist activities on wildlife and to implement new policies in the management of protected areas.     

The influence of environmental factors on the dynamics of the size and spatial structure of the chamois (Rupicapra rupicapra caucasica) population on the Caucasian Reserve

The size and structure of the chamois (Rupicapra rupicapra caucasica) population as well as the influence of anthropogenic factors on it were studied on the Caucasian Reserve (Western Caucasus). The negative effect of the snow cover, the thickness of which differed from the usual level, is shown. The effect of winters with thick snow cover is confirmed by the relation between these factors and the proportion of yearlings in the population one year after the winter. An increased death rate of animals in winters with a deep snow cover is observed in the case when their density exceeds the optimal one. Wolf predation does not affect the chamois population on the reserve. The number of wolves increases with an increase in the chamois population density to 15–20 ind./1000 ha. The adverse impact of humans on the chamois population is manifested in the regions with motor roads. The chamois, as compared to the deer, suffers from poachers to a lesser degree because it inhabits inaccessible areas and its trophy value is low.     

Seasonal Changes in Fecal Cortisol Metabolites in Pyrenean Chamois

Abstract: We studied seasonal changes in fecal Cortisol metabolites (FCM), which have been widely used as indicators of stress, in a population of Pyrenean chamois (Rupicapra pyrenaica pyrenaica) in the Cadí Range of northeastern Spain. We collected fecal samples from 2001 to 2003 in 3 particular locations with different altitudes and male or female presence, and we analyzed them for FCM and fecal nitrogen as an indicator of diet quality. We observed a clear seasonal pattern, with the highest FCM in winter, and we obtained correlations between FCM and monthly mean minimum temperatures and fecal nitrogen. We observed no effects of tourism presence, trophy hunting, or rut season on FCM. Analysis of cortisol metabolites in feces can be a good measure of winter stress in Pyrenean chamois.     

Effects of livestock and non-native mouflon on use of high-elevation pastures by Alpine chamois

Interspecific interference among livestock, native and non-native large herbivores is a key management and conservation issue, and little is known about its dynamics and implications. We investigated whether native Alpine chamois (Rupicapra r. rupicapra) modify their spatial distribution when non-native mouflon Ovis orientalis musimon and livestock (domestic sheep Ovis aries, and domestic goats Capra hircus) inhabit the same areas in the Alpine meadows of Italian Eastern Alps. We walked 5 hiking trails (5.0 ± 0.1 km) at dawn, twice a month during summers 2007 and 2008. During these surveys, we located each group of free-ranging ungulates (chamois, mouflon, and livestock). We also estimated the quality of meadows, finding that forage availability increased linearly as the distance from rocks (i.e., chamois refuges) increased. We predicted that the linear distance between chamois groups and rocks would depend on the co-presence of livestock and mouflon groups. Our results showed that chamois were more likely to be observed in areas with low food availability, but safer (i.e., closer to the rocks), whenever the nearest livestock group was larger and closer, especially if the shepherd's dog was present. Avoidance of the best feeding patches by the wild species is presumably due to spatial interference (e.g. visual and acoustic disturbance) and/or predation risk perception caused by the presence of shepherd's dogs. Similarly, the larger was the nearest group of mouflon, the closer to rocks was located the chamois group. Interestingly, mouflon group vicinity induced chamois to move closer to rocks only if mouflon rams were within the group. This suggests that physical displacement of the smaller species (i.e. chamois) is likely to occur to avoid direct disturbance of larger mouflon rams. Our study clearly showed how a native herbivorous species adjusts its spatial distribution and decreases the likelihood of using areas with higher food availability when livestock and/or a non-native species co-occur.     

Recombination and the origin of sequence diversity in the DRB MHC class II locus in chamois (Rupicapra spp.)

We examined the evolutionary processes contributing to genetic diversity at the major histocompatibility complex (MHC) class II DRB locus in chamois (Rupicapra spp., subfamily Caprinae). We characterised the pattern of intragenic recombination (or homologous gene conversion) and quantified the amount of recombination in the genealogical history of the two chamois species, Pyrenean chamois (Rupicapra pyrenaica) and Alpine chamois (Rupicapra rupicapra). We found evidence for intragenic recombination, and the estimated amount of population recombination suggests that recombination has been a significant process in generating DRB allelic diversity in the genealogical history of the genus Rupicapra. Moreover, positive selection appears to act on the same peptide-binding residues in both analysed chamois species, but not in identical intensity. Recombination coupled with positive selection drives the rapid evolution at the peptide-binding sites in the MHC class II DRB gene. Many chamois MHC class II DRB alleles are thus much younger than previously assumed.     

Increasing Himalayan tahr and decreasing chamois densities in the eastern Southern Alps, New Zealand: evidence for interspecific competition

There is anecdotal evidence that increasing densities of Himalayan tahr (Hemitragus jemlahicus) are associated with declining densities of chamois (Rupicapra rupicapra) in the Southern Alps, New Zealand. To examine this phenomenon, densities of tahr and chamois were measured at 53 sites within their sympatric range in the eastern Southern Alps during 1978–1979. In sites where only one species was present, tahr density was significantly higher than chamois density (P=0.032), probably reflecting species differences in sociality. Chamois density was higher in catchments without tahr than in those with tahr (P=0.012). Similarly, tahr density was significantly higher at sites without chamois than at those with chamois (P=0.033). Sites with both species present (termed sympatric) were significantly larger than those with only chamois or tahr (P<0.001). Following the prohibition of aerial hunting of tahr in 1983, 16 of the 17 sites where tahr and chamois were sympatric during 1978–1979 were recounted during 1991–1996. There was a 6-fold increase in the mean density of tahr between the two counts (P=0.001), whereas chamois density had declined significantly (P=0.006). Chamois persisted at only three sites, two of which had the highest chamois densities in 1978–1979. This is evidence that increasing densities of tahr exclude chamois from all but the `best' habitats. We conclude that intensive aerial hunting of tahr during 1967–1983 reduced tahr densities such that chamois could co-exist with tahr. Received: 20 March 1997 / Accepted: 8 September 1997     

Detection of a newly described pestivirus of Pyrenean chamois (Rupicapra pyrenaica pyrenaica) in France

A pestivirus was detected and characterized in chamois (Rupicapra pyrenaica pyrenaica) originating from the French part of the Pyrenees. Phylogenetic analysis of the pestivirus was done on the basis of a fragment from the 5' noncoding region including 22 published nucleotide sequences of different pestivirus strains. Our strain was grouped within the clade of border disease viruses (BDV). However, it had an intermediate position between clade BDV and classical swine fever viruses representing a basal position to BDV strains of domestic sheep. Our strain was grouped as a sister unit to a novel pestivirus (Chamois-1) recently described from chamois in Spain. Therefore, we postulate that this virus occurs in the entire population of Pyrenean chamois. On the basis of the phylogenetic grouping of this isolate, a postulated cross-species transmission of pestivirus from domestic sheep to chamois via shared pastures seems to be unlikely.     

Dung avoidance as a possible mechanism in competition between wild and domestic ungulates: two experiments with chamois <Emphasis Type="Italic">Rupicapra rupicapra</Emphasis>

Wild ungulates often adjust spatial behaviour where they coexist with livestock. In European mountains, chamois Rupicapra rupicapra or R. pyrenaica commonly avoid alpine pastures used by domestic sheep. Mechanisms leading to competitive losing out of the wild species are not well understood, but mostly, resource competition is inferred and sometimes demonstrated. We hypothesised that chamois need to minimise the risk of intestinal parasite uptake and therefore would avoid pastures contaminated with sheep dung. We tested this in two experiments by contaminating feeding patches of wild and captive chamois with (parasite-free) sheep faeces. Wild chamois did not avoid grazing sites or plots on which sheep dung had been placed at a (low) density representing the commonly encountered situation in the Swiss Alps. Captive chamois strongly reduced browsing time on small trees given for food when the surroundings of the trees were sprayed with a watery faecal solution. We concluded that the odour signalled a potential high risk of parasite infection to the captive chamois, whereas the density of sheep dung on the grazing site of wild chamois was not high enough to be perceived as a risk. The need to minimise endoparasite uptake from faeces may therefore play a role in driving spatial behaviour of wild ungulates and could result in competitive imbalance between wild and domestic ungulates.     

Mineral Levels in Pyrenean Chamois (Rupicapra pyrenaica)

The effects of mineral deficiencies are often sub-clinical, and the importance of mineral status is often underestimated in wildlife populations. To our knowledge, this is the first study that gives reference intervals of hepatic minerals for Pyrenean chamois (Rupicapra pyrenaica). We determined macro and trace mineral concentrations in liver samples from 100 animals (44 healthy and 56 sick) collected in the Catalan Pyrenees (NE Spain) from 1995 to 2008. After wet digestion, we determined Na, K, Ca, P, Mg, S, and Fe concentrations by inductively coupled plasma optical emission spectrometry (ICP-OES) and Cu, Zn, and Mo concentrations by inductively coupled plasma mass spectrometry (ICP-MS). We observed low hepatic concentrations of Cu in a considerable percentage of chamois, without evidences that these low concentrations increased their susceptibility to infectious diseases. The group of sick chamois had very similar percentage of animals (10/56) with low concentration of Cu (<20 ppm DW) than the group of healthy chamois (9/44). On the other hand, we observed that infectious diseases increased significantly the hepatic concentrations of Na, Ca, Mg, Fe and Zn, very likely, as a consequence of processes associated with the acute phase inflammatory response. The obtained values of liver mineral levels and their sources of variation, such as sex, age and disease, mostly fall within the range of those described for other ruminants, but possible deficiencies and differences between individuals and populations require further study.     

Preliminary results on snow surveys of Pyrenean grey partridge (<Emphasis Type="Italic">Perdix perdix hispaniensis</Emphasis>) in the Cantabrian Mountains

Peripheral populations tend to be smaller and more prone to extinction than those in core areas. Grey partridge Perdix perdix in its southern-most edge occupies mountain habitats, which is unlike any habitat in its northern European range. This makes these mountain ecotypes especially susceptible to range contractions and population decrease is likely the result of the current global change process. In Iberia, the so-called Pyrenean grey partridge subspecies P. p. hispaniensis is considered to be declining but little is known about its real trends and numbers. We locally studied the abundance of the Pyrenean grey partridges in winter by direct observations in the Cantabrian Mountains (NW Spain), while driving on a paved road. In addition, we related observed partridge abundances to different context variables to find the ones better helping to detect this species. The paved road-based censuses were successful, and we detected the presence and abundance of the Pyrenean grey partridge. Individuals and groups were more easily detected from December to February while couples were observed at the end of the winter (i.e. February–March). Wherever open paved roads exist, we strongly recommend using direct observations from them during winter as an adequate tool for Pyrenean grey partridge monitoring. This easy and cost-effective method affordable to managers and conservationists can contribute to a better understanding of the European mountain’s changing ecosystems and help us understanding the population trends of this vulnerable subspecies.     

Population genetics of chamois in the contact zone between the Alps and the Dinaric Mountains: uncovering the role of habitat fragmentation and past management

The chamois is a habitat specialist ungulate occupying “continental archipelagos” of fragmented rocky habitats which are frequently restricted to high altitudes. It is not clear whether forest habitats separating such population fragments act as barriers to gene flow. We studied the genetic makeup of the chamois in a topographically diverse landscape at the contact zone of two mountain ranges in Slovenia. Based on sequences of mitochondrial DNA, all Slovenian populations belong to a Northern chamois (Rupicapra r. rupicapra) subspecies. The range of chamois in Slovenia encompasses three different regions, each with unique topography, habitat connectivity and abundance of chamois: the Alps, the Dinaric Mts., and the Pohorje Mts. The habitat of the chamois is extensive and more or less continuous in the Alps, but suboptimal and fragmented in the remaining regions. In agreement with neutral genetic theory, large Northern chamois populations tended to have higher allelic richness and observed heterozygosity. Spatial clustering bears the differentiation into four geographically associated clusters within Slovenia and also revealed a strong substructure within all mountain ranges with suboptimal chamois habitat. Surprisingly, some small Dinaric populations have stayed genetically isolated in restricted habitat patches, even if they are geographically very close to each other. The four clusters, each having a unique demographic history, should be regarded as independent units for management purposes.     

Social Interactions in Isard Kids Rupicapra pyrenaica pyrenaica (Artiodactyla: Bovidae): Possible Influence on Development of Social Bonding

Assuming that social experiences at an early age may be a determinant factor in the process of formation of social network and attachments, we studied the social interactions of isard, or Pyrenean chamois, kids (Rupicapra pyrenaica pyrenaica) during their first year, to understand “how” sociality arises in isards rather than “why”. Interactions with the mother were affiliative, and approaches of kid to mother or vice versa were generally followed by another “positive” interaction. Interactions between kids were also frequent and playful. Interactions with both mother and peers provided positive reinforcement which may strengthen bonds. However, early interactions of kids with yearlings and other adults provided negative experience, no attachment thus developing from them. Social affinities of adults as well as the social and spatial organization of groups may be understood when ontogenetical history and immediate causality are highlighted.     

Effects of an exceptionally snowy winter on chamois survival

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.