Pasture Quality Affects Juvenile Survival through Reduced Maternal Care in a Mountain‐Dwelling Ungulate

Lactation exerts heavy energetic and physiological costs to mothers, whilst determining early growth and survival of offspring. To mountain ungulates, access to high‐quality forage during nursing and weaning is crucial for reproductive success. We have evaluated the effects of pasture quality on suckling behaviour and winter survival of Apennine chamois Rupicapra pyrenaica ornata kids, across three areas. Areas A‐B (‘poor’ areas) were characterised by a reduced availability of nutritious forage, thus a lower diet quality for female chamois and kids; Area C (a ‘rich’ area) included a much greater availability of nutritious forage. In poor areas, pasture quality has been reduced by climatic and plant composition changes, as well as the presence of a herbivore competitor (red deer Cervus elaphus). In poor areas, we recorded a significantly (1) lower suckling success of chamois kids (number of suckling bouts/number of suck attempts); (2) lower frequency of suckling bouts (n. suckling bouts/kid/h); and (3) lower suckling intensity (suck duration/kid/h) in respect to the rich area. Conversely, frequencies of suckling rejections and those of suckling attempts (n. events/kid/h) were the lowest in the rich area. Winter survival of chamois kids was c. 2 times greater in the rich area (45%) than in poor areas (20–26%). In the poor areas, resource scarcity induced adult female chamois to decrease maternal cares and favour their own maintenance, ultimately affecting population dynamics through kid winter mortality.     

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.     

Contrasting Effects of Climate Change on Alpine Chamois

Global climate change can affect animal ecology in numerous ways, but researchers usually emphasize undesirable consequences. Temperature increases, for instance, can induce direct physiological costs and indirect effects via mismatches in resource needs and availability. Species living in mountainous regions, however, could experience beneficial effects because winters might become less severe. We examined the potentially opposing effects of climate change during spring, summer, and winter on recruitment in Alpine chamois (Rupicapra rupicapra). We examined initial recruitment (i.e., the ratio of kids to adult females) and net recruitment (i.e., the ratio of yearlings to adult females) of Alpine chamois through the use of linear mixed effects models and data from block count censuses performed across a 1,500-km² study area in the Italian Alps during summer from 2001 to 2015. Initial recruitment was relatively resistant to the effects of climate change, declining slightly over the study period. We suggest that the effects of increased forage availability and lower snow cover in winter may benefit the reproductive output of adult females, compensating for any negative effects of trophic mismatch and higher temperatures during summer. By contrast, net recruitment strongly declined throughout the study period, consistent with the slight decline of initial recruitment and the negative effects of increasing summer temperatures on the survival of kids during their first winter. These negative effects seemed to outweigh positive effects of climate change, even in a species strongly challenged by winter conditions. These findings provide important information for hunted populations; setting more appropriate hunting bags for yearling chamois should be considered. The ecological plasticity of the chamois, which also inhabits low altitudes, may allow a possible evolutionary escape for the species. © 2020 The Wildlife Society.     

The role of geological substrate for horn growth in ungulates: a case study on Alpine chamois

Ecological factors such as environmental and climatic conditions affect the growth of ornaments in ungulates. Studies about their influence can reveal key information on individual life histories and detect the events that can have important evolutionary consequences. Among the ecological factors commonly thought to play a role in such ecological processes, researches have so far neglected the possible influence of the geological feature which may either promote or restrain the early investment in horn growth. This study takes into consideration a broad range of ecological factors that are expected to affect horn growth and analyses how the substrate (calcareous versus siliceous) could modify the horn size in 1,685 yearling Alpine chamois (Rupicapra rupicapra) which were legally shot during 5 consecutive years in Central-Eastern Italian Alps. Interestingly, we found shorter horns in chamois shot on siliceous than on calcareous substrate, irrespective of the sex, showing how the substrate (and related ecological conditions) may affect horn growth. The substrate interacted with two ecological factors in shaping the horn size, i.e. aspect and snow cover. Differences between horn lengths recorded for chamois shot on these two substrates were less pronounced where South-facing slopes prevailed. During an average winter (about 100 days with more than 10 cm of snow on the ground) chamois horns were longer in individuals using calcareous areas, although the increase of number of days with more than 10 cm of snow on the ground had a stronger impact in reducing horn length in calcareous environment. We demonstrated that substrate, and related ecological conditions, influence horn growth. In ungulate species where ornament size is related to reproductive success, environmental factors, such as geological feature, can thus cause evolutionary consequences in the expression of ornaments and individual fitness.     

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.     

Heterogeneity in Primary Productivity Influences Competitive Interactions between Red Deer and Alpine Chamois

Habitat heterogeneity can promote coexistence between herbivores of different body size limited to different extents by resource quantity and quality. Red deer (Cervus elaphus) are known as superior competitors to smaller species with similar diets. We compared competitive interactions and habitat use between red deer and Alpine chamois (Rupicapra rupicapra) in two adjacent valleys in a strictly protected area in the Central Alps. Red deer density was higher in the valley with higher primary productivity. Only here was horn growth in kid and yearling chamois (as a measure for body condition) negatively correlated with red deer population size, suggesting interspecific competition, and chamois selected meadows with steeper slopes and lower productivity than available on average. Conversely, red deer selected meadows of high productivity, particularly in the poorer area. As these were located mainly at lower elevations, this led to strong altitudinal segregation between the two species here. Local differences in interspecific competition thus coincided with differences in habitat preference and–segregation between areas. This suggests that spatial habitat and resource heterogeneity at the scale of adjacent valleys can provide competition refuges for competitively inferior mountain ungulates which differ from their superior competitor in their metabolic requirements.     

Disentangling demographic effects of red deer on chamois population dynamics

Investigating the impact of ecological factors on sex‐ and age‐specific vital rates is essential to understand animal population dynamics and detect the potential for interactions between sympatric species. We used block count data and autoregressive linear models to investigate variation in birth rate, kid survival, female survival, and male survival in a population of Alpine chamois Rupicapra rupicapra rupicapra monitored over 27 years within the Stelvio National Park, Central Italian Alps, as function of climatic variables, density dependence, and interspecific competition with red deer Cervus elaphus. We also used path analysis to assess the indirect effect of deer abundance on chamois growth rate mediated by each demographic parameter. Based on previous findings, we predicted that birth rate at [t] would negatively relate to red deer abundance at year [t − 1]; survival rates between [t] and [t + 1] would negatively relate to red deer abundance at year [t − 1] and to the interactive effect of winter precipitation at [t + 1] and chamois density at [t]. Our results showed that birth rate was positively related to spring–summer precipitation in the previous year, but this effect was hampered by increasing red deer abundance. Kid and female survival rates were negatively related to the combined effect of chamois abundance and winter precipitation. Male and female survival rates were negatively related to lagged red deer abundance. The path analysis supported a negative indirect effect of red deer abundance on chamois growth rate mediated by birth rate and female survival. Our results suggest that chamois population dynamics was largely explained by the synergistic effect of density dependence and winter harshness, as well as by interspecific competition with red deer, whose effects were seemingly stronger on the kid–female segment of the population.     

Food searching in griffon vultures

Observations are presented on food searching in griffon vultures. To locate food directly, griffon vultures do not use a sense of smell but rely on vision. However, most birds locate carcasses indirectly—by watching the activities of neighbouring birds. The method of food searching is described and the number of birds which arrive at a carcass is shown to depend on the amount of food that is available. Birds do not hold feeding ranges, but travel widely during food searching, one bird travelling 180 km in 6 days. The density of searching birds varies greatly according to the density of ungulates in an area, birds being most numerous over high ungulate concentrations. The altitude at which birds search also varies, birds flying at higher altitudes over areas of low ungulate density and at lower altitudes over ungulate concentrations. These variations in searching density and height are shown to affect the efficiency of food searching. Carcasses are located rapidly and the food consumed quickly in high ungulate density areas, while in low ungulate density conditions carcasses can take a long time to be located. It is concluded that griffon vultures are more likely to find food and are better able to compete with mammalian competitors by searching over migratory ungulate herds than over areas containing resident ungulate species. Their adaptations for gliding flight enable these birds to follow these migratory ungulates throughout the year.     

Birthdate, mass and survival in mountain goat kids: effects of maternal characteristics and forage quality

In temperate environments, early-born ungulates may enjoy a longer growth period before winter, and so attain a higher body mass and an increased probability of survival compared to late-born ones. We assessed the effects of maternal characteristics, forage quality and population density on kid birthdate, mass and survival in a population of marked mountain goats (Oreamnos americanus) in Alberta. The duration and timing of the birth season were similar in all years. Births were highly synchronised: 80% of kids were born within 2 weeks of the first birth. Maternal age, maternal social rank and density did not affect kid birthdate or mass. Previous breeding experience was not related to kid birthdate, but kids born to pluriparous mothers were heavier during summer than kids born to primiparous mothers. Male and female kids had similar mass and accumulated mass linearly during summer. Early-born kids were heavier than late-born kids. Faecal crude protein (FCP) in late spring and maternal mass were positively related to kid mass. Survival to weaning appeared higher for males (90%) than for females (78%), but survival to 1 year was 65% for both sexes. FCP in late spring, density, birthdate and mass did not affect kid survival to weaning in either sex. Survival to 1 year increased with FCP in late spring for females, but not for males. Survival to 1 year was independent of birthdate for both sexes, but heavy females survived better than light ones. Multiple logistic regression revealed a positive effect of mass on survival to 1 year when the sexes were pooled. Our results suggest that mountain goats are constrained to give birth in a short birth season synchronised with forage productivity.     

Rewilding traditional grazing areas affects scavenger assemblages and carcass consumption patterns

The abandonment of traditional livestock farming systems in Mediterranean countries is triggering a large-scale habitat transformation, which, in general, consists of the replacement of open grazing areas by woodlands through non-managed regeneration. As a consequence, wild ungulates are occupying rapidly the empty niche left by domestic ungulates. Both types of ungulates represent the main trophic resource for large vertebrate scavengers. However, a comparison of how vertebrate scavengers consume ungulate carcasses in different habitats with different ungulate species composition is lacking. This knowledge is essential to forecast the possible consequences of the current farmland abandonment on scavenger species. Here, we compared the scavenging patterns of 24 wild and 24 domestic ungulate carcasses in a mountainous region of southern Spain monitored through camera trapping. Our results show that carcasses of domestic ungulates, which concentrate in large numbers in open pasturelands, were detected and consumed earlier than those of wild ungulate carcasses, which frequently occur in much lower densities at more heterogenous habitats such as shrublands and forest. Richness and abundance of scavengers were also higher at domestic ungulate carcasses in open habitats. Vultures, mainly griffons (Gyps fulvus), consumed most of the carcasses, although mammalian facultative scavengers, mainly wild boar (Sus scrofa) and red fox (Vulpes vulpes), also contributed importantly to the consumption of wild ungulate carcasses in areas with higher vegetation cover. Our findings evidence that the abandonment of traditional grazing may entail consequences for the scavenger community, which should be considered by ecologists and wildlife managers.     

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.     

Physical size,condition, and demography of chamois (Rupicapra rupicapra L.) in the Avoca River region,Canterbury, New Zealand

Abstract

The physical and demographic characteristics of chamois in the Avoca region are evaluated from 306 animals shot and autopsied between 1975 and 1978. These data are compared with published and unpublished information for chamois populations in Westland and Canterbury. Avoca chamois were large-framed, but weighed less than Westland chamois. The weight difference suggests better habitat condition and food resources for Westland animals, but the large skeletal size of Avoca chamois is unexplained.

High rates of juvenile mortality were caused by acute bacterial-pneumonia infections (Pasteurella). These deaths and other losses by natural causes were offset by the good breeding success of adult females so that stable population numbers were maintained.     

Reflections of ecological differences? Stress responses of sympatric Alpine chamois and red deer to weather,forage quality,and human disturbance

Depending on the habitats they live in, temperate ungulates have adapted to different degrees to seasonally changing forage and weather conditions, and to specific escape strategies from predators. Alpine chamois, a mountain ungulate, and red deer, originally adapted to open plains, would therefore be expected to differ in their physiological responses to potential stressors. Based on 742 chamois and 1557 red deer fecal samples collected year‐round every 2 weeks for 4 years at the same locations within a strictly protected area in the Swiss Alps, we analyzed glucocorticoid metabolite (FGM) concentrations for both species. Results from linear mixed effects models revealed no physiological stress response to changing visitor numbers, but instead to drought conditions for both species during summer. In winter, FGM concentrations increased with increasing snow height in both species, but this response was modulated by temperature in red deer. Chamois showed a stronger stress response to increasing snow height during November and December than between January and March, while FGM concentrations increased with decreasing temperature throughout winter. An increase in FGM concentrations with decreasing forage digestibility during winter was found only for red deer. The results are thus partly in contradiction to expectations based on feeding type and adaptations to different habitats between the two species. The lack of a response to forage digestibility in chamois may reflect either better adaptation to difficult feeding conditions in subalpine forests, or, by contrast, strong constraints imposed by forage quality. The similar responses of both species to weather conditions in winter suggest that climatic factors at the elevations examined here are sufficiently harsh to be limiting to temperate ungulates regardless of their specific adaptations to this environment.     

Comparative evaluation of effort, capture and handling effects of drive nets to capture roe deer (Capreolus capreolus), Southern chamois (Rupicapra pyrenaica) and Spanish ibex (Capra pyrenaica)

The objective of this study is to assess the usefulness of drive nets to capture roe deer (Capreolus capreolus), Southern chamois (Rupicapra pyrenaica) and Spanish ibex (Capra pyrenaica), comparing the results obtained with other capture methods and amongst the three species. Sixty-five drive net capture operations using beaters were conducted from January 1998 to September 2004. A total of 161 wild ungulates (31 roe deer, 95 Southern chamois and 35 Spanish ibexes) were captured. The average number of animals captured per operation was 1.07 for roe deer, 3.96 for Southern chamois and 2.92 for Spanish ibex. The average number of person–days per captured animal was 21.5, 7.1 and 10.6 for roe deer, Southern chamois and Spanish ibex, respectively. Specificity was 100% for Southern chamois and Spanish ibex (only the target species captured) and 77.5% for roe deer. Risk for the animals (mortality plus injuries) was 3.23% for roe deer, 5.27% for Southern chamois and 0% for Spanish ibex, whereas injuries to the operators occurred with 3.1% of the handled animals. Sex ratio was skewed towards females in roe deer, towards males in Southern chamois and balanced in Southern chamois. Drive nets showed good performance, although many operators were required. Safety for the animals and specificity were higher than traditionally attributed to this capture method. It is concluded that drive nets are an efficient and safe method to capture many ungulate species.     

Effect of vitamin E supplementation on development of reproductive organs in Boer goat

The aim of this study was to evaluate the supplementation of Vitamin E in diet on development of reproductive organs in Boer goat. Twenty-four healthy, Boer male kids of similar body weight (BW) were selected at 3 months of age from the kid flock. The Boer kids were randomly divided into four groups and each group was supplemented with Vitamin E at 0, 80, 320 or 880 IU kid(-1)d(-1) for 5 months. Three Boer goats in each group were slaughtered at the age of 8 months and measured for testis and epididymises and a ductus deferens biopsy by H&E stained slides was run. The results showed that supplementing 80 and 320 IU kid(-1)d(-1) Vitamin E increased weights of the epididymis and the numeric density of convoluted seminiferous tubules. Diameters of convoluted seminiferous tubules and epididymis ductules and numeric density of spermatogenic cells tended to be larger in 80 IU kid(-1)d(-1) compared to the control group (P<0.05). Weight of ductus deferens and thickness of epididymis wall were increased significantly with 320 IU kid(-1)d(-1) (P<0.01). However, no significant effects were observed on thickness of germinal epithelium, numeric density of leydig cells, testis weight or indexes of ductus deferens (P>0.05). In conclusion, supplementing 80 and 320 IU kid(-1)d(-1) Vitamin E in diets significantly simulates the development of reproductive organs in Boer goats. No significantly improved effects were found from supplementing 880 IU kid(-1)d(-1).     

Differential effects of undernutrition during pregnancy on the behaviour of does and their kids at parturition and on the establishment of mutual recognition

We investigated whether undernutrition during the second half of pregnancy impaired the behaviour of does and their kids at parturition and early mutual recognition. Twenty-two control and 22 underfed mixed-breed, multiparous dairy goats were used, together with their respective kids (control, n = 31: nine singles, 16 twins and six triplets; underfed, n = 32: 11 singles, 18 twins and three triplets). Undernutrition involved limiting protein and energy intake at 70% of the nutritional requirements for maintenance and foetal growth from day 70 of pregnancy until birth. The behaviour of mothers and their two first-born kids was observed for 90 min from the birth of each kid. Maternal olfactory recognition of the kid was assessed at 4 h post partum by testing selective nursing behaviour. Non-olfactory recognition was assessed at 8 h in a two-choice test excluding olfactory cues. In kids, preference for the mother was assessed in a two-choice test at either 12 or 24 h post partum. Bodyweight of does and kids were lower in the underfed group up to 2 weeks post partum. At parturition, licking, maternal bleating frequency and latency to nursing did not differ between nutritional groups. Control kids were faster than underfed kids to stand, search for and reach the udder, but underfed kids bleated more and tended to spend more time at the udder. Both control and underfed does accepted their own kid and rejected the alien in the selectivity test at 4 h. In contrast, at 8 h post partum, only control goats showed a significant preference for their own kid in the non-olfactory recognition test. Both control and underfed kids showed a preference for their own mother at 12 and 24 h and undernutrition during pregnancy had little influence on the performance of kids. However, 12 h-old underfed kids tended to be less active than control kids and visited their own mothers less than control kids. There were no significant correlations between the behaviour of the mother or of the kid at parturition and their performance in the discrimination tests. Overall, undernutrition in the second half of pregnancy appears to be more detrimental for the behaviour of the mother than for the kid. Furthermore, it has more impact on the establishment of maternal non-olfactory recognition than on maternal care at parturition or the establishment of maternal selectivity.     

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.     

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.     

Gastro-intestinal nematode infections in four Alpine chamois herds: influence of host density on helminth egg output

A three years survey on gastro-intestinal nematode egg output was carried out in four Alpine chamois herds (Rupicapra rupicapra rupicapra) from Belluno province (Italy). Data were analysed in order to detect the influence of cattle presence and chamois density on egg output, and to evaluate the stability of host-parasite relationship. As expected, variations related to the season were present, with the major peak of egg output observed in the summer. Results highlighted an unexpected inverse influence of host density on egg counts. The results are in relation to host-parasite relationship stability. Parasites seem to be able to self-regulate in the absence of parasite-induced host mortality. No effect due to cattle presence was detected.