Different effects of alpine woody plant expansion on domestic and wild ungulates.

Changes in land-use and climate affect the distribution and diversity of plant and animal species at different spatiotemporal scales. The extent to which species-specific phenotypic plasticity and biotic interactions mediate organismal adaptation to changing environments, however, remains poorly understood. Woody plant expansion is threatening the extent of alpine grasslands worldwide, and evaluating and predicting its effects on herbivores is of crucial importance. Here, we explore the impact of shrubification on the feeding efficiency of Pyrenean chamois (Rupicapra p. pyrenaica), as well as on the three most abundant coexisting domestic ungulate species: cattle, sheep and horses. We use observational diet composition from May to October and model different scenarios of vegetation availability where shrubland and woodland proliferate at the expense of grassland. We then predicted if the four ungulate species could efficiently utilize their food landscapes with their current dietary specificities measuring their niche breath in each scenario. We observed that the wild counterpart, due to a higher trophic plasticity, is less disturbed by shrubification compared to livestock, which rely primarily on herbaceous plants and will be affected 3.6 times more. Our results suggest that mixed feeders, such as chamois, could benefit from fallow landscapes, and that mountain farmers are at a growing economic risk worldwide due to changing land-use practices and climate conditions.

From gestation to weaning: Combining robust design and multi-event models unveils cost of lactation in a large herbivore.

The cost of current reproduction on survival or future reproduction is one of the most studied trade-offs governing resource distribution between fitness components. Results have often been clouded, however, by the existence of individual heterogeneity, with high-quality individuals able to allocate energy to several functions simultaneously, at no apparent cost. Surprisingly, it has also rarely been assessed within a breeding season by breaking down the various reproductive efforts of females from gestation to weaning, even though resource availability and energy requirements vary greatly. We filled this gap by using an intensively monitored population of Pyrenean chamois and by expanding a new methodological approach integrating robust design in a multi-event framework. We distinguished females that gave birth or not, and among reproducing females whether they lost their kid or successfully raised it until weaning. We estimated spring and summer juvenile survival, investigated whether gestation, lactation or weaning incurred costs on the next reproductive occasion, and assessed how individual heterogeneity influenced the detection of such costs. Contrary to expectations if trade-offs occur, we found a positive relationship between gestation and adult survival suggesting that non-breeding females are in poor condition. Costs of reproduction were expressed through negative relationships between lactation and both subsequent breeding probability and spring juvenile survival. Such costs could be detected only once individual heterogeneity (assessed as two groups contrasting good vs. poor breeders) and time variations in juvenile survival were accounted for. Early lactation decreased the probability of future reproduction, providing quantitative evidence of the fitness cost of this period recognized as the most energetically demanding in female mammals and critical for neonatal survival. The new approach employed made it possible to estimate two components of kid survival that are often considered practically unavailable in free-ranging populations, and also revealed that reproductive costs appeared only when contrasting the different stages of reproductive effort. From an evolutionary perspective, our findings stressed the importance of the temporal resolution at which reproductive cost is studied, and also provided insights on the reproductive period during which internal and external factors would be expected to have the greatest fitness impact.

Onset of autumn shapes the timing of birth in Pyrenean chamois more than onset of spring.

In seasonal environments, birth dates are a central component for a species' life history, with potential long-term fitness consequences. Yet our understanding of selective pressures of environmental changes on birth dates is limited in wild mammals due to the difficulty of data collection. In a context of rapid climate change, the question of a possible mismatch between plant phenology and birth phenology also remains unanswered for most species. We assessed whether and how the timing of birth in a mountain mammal (isard, also named Pyrenean chamois, Rupicapra pyrenaica pyrenaica) tracked changes in plant growing season, accounting for maternal traits, individual heterogeneity and population density. We not only focused on spring conditions but also assessed to what extent onset of autumn can be a driver of phenological biological events and compared the magnitude of the response to the magnitude of the environmental changes. We relied on a 22-year study based on intensively monitored marked individuals of known age. Births were highly synchronized (80% of kids born within 25 days) and highly repeatable (84%; between-female variation of 9.6 days, within-female variation of 4.2 days). Individual phenotypic plasticity allows females to respond rapidly to interannual changes in plant phenology but did not prevent the existence of a mismatch: a 10-day advance in the autumn or spring plant phenology led to 3.9 and 1.3 days advance in birth dates, respectively. Our findings suggest that plant phenology may act as a cue to induce important stages of the reproductive cycle (e.g. conception and gestation length), subsequently affecting parturition dates, and stressed the importance of focusing on long-term changes during spring for which females may show much lower adaptive potential than during autumn. These results also question the extent to which individual plasticity along with high heterogeneity among individuals will allow species to cope with demographic consequences of climate changes.

Tall Grass Invasion After Grassland Abandonment Influences the Availability of Palatable Plants for Wild Herbivores: Insight into the Conservation of the Apennine Chamois Rupicapra pyrenaica ornata.

Invasion of the tall grass Brachypodium genuense was observed in an area of the central Apennines (Italy) where the population size of Apennine chamois (Rupicapra pyrenaica ornata) was in strong decline. Since this dominant tall grass threatens biodiversity and forage quality, our hypothesis was that B. genuense abundance influenced that of palatable species for the chamois, depending on their functional traits and distribution patterns. Our sampling design used plots of 10 × 10 m and 1 × 1 m to investigate the plant community level and fine-scale interactions. We analyzed data using correlation, generalized linear models, and redundancy analyses. We found that B. genuense can reach high abundance values on the deepest soils. Its high cover value influences plant community composition by competitive exclusion of subordinate species and suppression of functional features because of temporal or spatial niche overlap. This leads to low cover of palatable species at a fine scale, and to poor pasture quality for chamois at a wider scale. Therefore, we postulated that B. genuense invasion, enhanced by long-term grazing cessation, may reduce the availability of palatable plants for Apennine chamois, especially below the potential timberline (1900-2000 m a.s.l.). The high abundance of B. genuense may amplify the effect of other negative factors, such as competition with red deer (Cervus elaphus) and climate change, in restricting the suitable habitat of the Apennine chamois to the higher sectors of the central Apennines. Thus, we suggested that B. genuense spread should be monitored carefully and plans to control its invasion should be implemented.

A novel epidemiological model to better understand and predict the observed seasonal spread of Pestivirus in Pyrenean chamois populations.

Seasonal variations in individual contacts give rise to a complex interplay between host demography and pathogen transmission. This is particularly true for wild populations, which highly depend on their natural habitat. These seasonal cycles induce variations in pathogen transmission. The seasonality of these biological processes should therefore be considered to better represent and predict pathogen spread. In this study, we sought to better understand how the seasonality of both the demography and social contacts of a mountain ungulate population impacts the spread of a pestivirus within, and the dynamics of, this population. We propose a mathematical model to represent this complex biological system. The pestivirus can be transmitted both horizontally through direct contact and vertically in utero. Vertical transmission leads to abortion or to the birth of persistently infected animals with a short life expectancy. Horizontal transmission involves a complex dynamics because of seasonal variations in contact among sexes and age classes. We performed a sensitivity analysis that identified transmission rates and disease-related mortality as key parameters. We then used data from a long-term demographic and epidemiological survey of the studied population to estimate these mostly unknown epidemiological parameters. Our model adequately represents the system dynamics, observations and model predictions showing similar seasonal patterns. We show that the virus has a significant impact on population dynamics, and that persistently infected animals play a major role in the epidemic dynamics. Modeling the seasonal dynamics allowed us to obtain realistic prediction and to identify key parameters of transmission.

Senescence in breeding success of female Alpine chamois (Rupicapra rupicapra): the role of female quality and age.

Although numerous studies have reported reproductive senescence in wild animal populations, we still know very little on inter-individual differences in rates of ageing and on the factors accounting for these differences. To investigate age-related variation in breeding success in a natural population of Alpine chamois (Rupicapra rupicapra), we used 15 years of data obtained by monitoring individual ear-tagged females. Analyses at the population level confirmed the occurrence of a decline in female breeding success, which is most noticeable from 9 years of age onward. Using an age-reverse approach, we showed that senescence in female breeding success occurs at the same age, since only very old individuals (older than 16 years) showed a decline in breeding success in the years preceding death. Interestingly, we also found evidence that 'success comes from success', as females that gave birth in a given year were more likely to procreate again in the following year. Moreover, results showed that discrepancies between successful and unsuccessful individuals tend to become more relevant in the oldest age classes. There was no evidence of a terminal allocation. These results emphasize the importance of age-dependent effects and individual quality in shaping reproductive senescence in a wild long-lived ungulate.

Age-specific survival and annual variation in survival of female chamois differ between populations.

In many species, population dynamics are shaped by age-structured demographic parameters, such as survival, which can cause age-specific sensitivity to environmental conditions. Accordingly, we can expect populations with different age-specific survival to be differently affected by environmental variation. However, this hypothesis is rarely tested at the intra-specific level. Using capture-mark-recapture models, we quantified age-specific survival and the extent of annual variations in survival of females of alpine chamois in two sites. In one population, survival was very high (>0.94; Bauges, France) until the onset of senescence at approximately 7 years old, whereas the two other populations (Swiss National Park, SNP) had a later onset (12 years old) and a lower rate of senescence. Senescence patterns are therefore not fixed within species. Annual variation in survival was higher in the Bauges (SD = 0.26) compared to the SNP populations (SD = 0.20). Also, in each population, the age classes with the lowest survival also experienced the largest temporal variation, in accordance with inter-specific comparisons showing a greater impact of environmental variation on these age classes. The large difference between the populations in age-specific survival and variation suggests that environmental variation and climate change will affect these populations differently.

Upscaling the niche variation hypothesis from the intra- to the inter-specific level.

The "niche variation hypothesis" (NVH) predicts that populations with wider niches should display higher among-individual variability. This prediction originally stated at the intra-specific level may be extended to the inter-specific level: individuals of generalist species may differ to a greater extent than individuals of a specialist species. We tested the NVH at intra- and inter-specific levels based on a large diet database of three large herbivore feces collected in the field and analyzed using DNA metabarcoding. The three herbivores (roe deer Capreolus capreolus, chamois Rupicapra rupicapra and mouflon Ovis musimon) are highly contrasted in terms of sociality (solitary to highly gregarious) and diet. The NVH at the intraspecific level was tested by relating, for the same population, diet breadth and inter-individual variation across the four seasons. Compared to null models, our data supported the NVH both at the intra- and inter-specific levels. Inter-individual variation of the diet of solitary species was not larger than in social species, although social individuals feed together and could therefore have more similar diets. Hence, the NVH better explained diet breadth than other factors such as sociality. The expansion of the population niche of the three species was driven by resource availability, and achieved by an increase in inter-individual variation, and the level of inter-individual variability was larger in the generalist species (mouflon) than in the specialist one (roe deer). This mechanism at the base of the NVH appears at play at different levels of biological organization, from populations to communities.

Physiological response to etho-ecological stressors in male Alpine chamois: timescale matters!

From a life history perspective, glucocorticoids secreted by the neuroendocrine system, integrating different sources of stress through an adaptive feedback mechanism, may have important consequences on individual fitness. Although stress responses have been the object of several investigations, few studies have explored the role of proximate mechanisms responsible for the potential trade-offs between physiological stress and life history traits integrating social and environmental stressors. In 2011 and 2012, we collected data on faecal cortisol metabolites (FCM) in a marked male population of Alpine chamois, within the Gran Paradiso National Park (Italy). Using a model selection approach we analysed the effect of potential etho-ecological stressors such as age, social status (territorial vs. non-territorial males), minimum temperature, snow depth and precipitation on FCM variation. To correctly interpret environmentally and socially induced stress responses, we conducted model selections over multiple temporal scales defined a priori: year, cold months, spring, warm months, mating season. Over the year, FCM levels showed a negative relationship with minimum temperature, but altogether, climatic stressors had negligible effects on glucocorticoid secretion, possibly owing to good adaptations of chamois to severe weather conditions. Age was negatively related to FCM during the rut, possibly due to greater experience of older males in agonistic contests. Social status was an important determinant of FCM excretion: while both the 'stress of subordination' and the 'stress of domination' hypotheses received some support in spring and during the mating season, respectively, previous data suggest that only the latter may have detrimental fitness consequences on male chamois.

Intraseasonal variation in reproductive effort: young males finish last.

Age-dependent reproductive timing has been observed in females of a number of species; older females often breed earlier in the season and experience higher reproductive success as a result. However, to date, evidence for within-season variation in reproductive effort (RE) for males has been relatively weak. Males are expected to time RE in light of intraseasonal variations in the availability of receptive females and competition with other males. Young males, which are typically smaller and less experienced, might benefit from breeding later in the season, when male-male competition is less intense. Using a long-term data set of Alpine chamois Rupicapra rupicapra, we sought to evaluate the hypothesis that younger males allocate highest RE late in the breeding season, at a time when older male RE has decreased substantially. Our results support this hypothesis, which suggests that intraseasonal variation in RE may be an adaptive life-history trait for males as well as females.

Effects of early horn growth on reproduction and hunting mortality in female chamois.

1. Environmental conditions during early development can affect the growth patterns of vertebrates, influencing future survival and reproduction. In long-lived mammals, females that experience poor environmental conditions early in life may delay primiparity. In female bovids, annual horn growth increments may provide a record of age-specific reproduction and body growth. Horn length, however, may also be a criterion used by hunters in selecting animals to harvest, possibly leading to artificial selection. 2. We studied three populations of chamois (Rupicapra rupicapra) in the western Alps to explore the relationships between female horn length and early growth, age of primiparity and age-specific reproduction. We also compared the risk of harvest to reproductive status and horn length. 3. Early horn growth was positively correlated with body mass in pre-reproductive females and with reproduction in very young and senescent adults. Females with strong early horn growth attained primiparity at an earlier age than those with weak early growth. Horn length did not affect hunter selection, but we found a strong hunter preference for nonlactating females. 4. Our research highlights the persistent effects of early development on reproductive performance in mammals. Moderate sport harvests are unlikely to affect the evolution of phenotypic traits and reproductive strategies in female chamois. A policy of penalizing hunters that harvest lactating females, however, may increase the harvest of 2-year-old females, which have high reproductive potential.

Analyzing long-term impacts of ungulate herbivory on forest-recruitment dynamics at community and species level contrasting tree densities versus maximum heights.

Herbivores are constitutive elements of most terrestrial ecosystems. Understanding effects of herbivory on ecosystem dynamics is thus a major, albeit challenging task in community ecology. Effects of mammals on plant communities are typically explored by comparing plant densities or diversity in exclosure experiments. This might over-estimate long-term herbivore effects at community levels as early life stage mortality is driven by a multitude of factors. Addressing these challenges, we established a set of 100 pairs of ungulate exclosures and unfenced control plots (25 m2) in mixed montane forests in the Alps in 1989 covering a forest area of 90 km2. Investigations ran until 2013. Analogous to the gap-maker-gap-filler approach, dynamically recording the height of the largest trees per tree species in paired plots with and without exclosures might allow for assessing herbivore impacts on those individuals with a high probability of attaining reproductive stages. We thus tested if recording maximum heights of regenerating trees would better reflect effects of ungulate herbivory on long-term dynamics of tree regeneration than recording of stem density, and if species dominance patterns would shift over time. For quantifying the effects of ungulate herbivory simultaneously at community and species level we used principle response curves (PRC). PRCs yielded traceable results both at community and species level. Trajectories of maximum heights yielded significant results contrary to trajectories of total stem density. Response patterns of tree species were not uniform over time: e.g., both Norway spruce and European larch switched in their response to fencing. Fencing explained about 3% of the variance of maximum tree heights after nine years but increased to about 10% after 24 years thus confirming the importance of long-term surveys. Maximum height dynamics of tree species, addressed in our study, can thus reflect local dominance of tree species via asymmetric plant competition. Such effects, both within and among forest patches, can accrue over time shaping forest structure and composition.

Analysing the isotopic life history of the alpine ungulates Capra ibex and Rupicapra rupicapra rupicapra through their horns.

The horn of ungulate grazers offers a valuable isotopic record of their diet and environment. However, there have been no reports of the spatio-temporal variation of the isotopic composition of horns. We investigated patterns of carbon (delta(13)C) and nitrogen (delta(15)N) isotopic composition along and perpendicular to the horn axis in Capra ibex and Rupicapra rupicapra rupicapra to assess the effects of animal age, within-year (seasonal) and inter-annual variation, natural contamination and sampling position on horn isotope composition. Horns of male C. ibex (n = 23) and R. r. rupicapra (n = 1) were sampled longitudinally on the front (only R. r. rupicapra) and back side and on the surface and sub-surface. The sides of the R. r. rupicapra horn did not differ in delta(13)C. In both species, the horn surface had a 0.15 per thousand lower delta(13)C and a higher carbon-to-nitrogen (C/N) ratio than the sub-surface. Washing the horn with water and organic solvents removed material that caused these differences. With age, the delta(15)N of C. ibex horns increased (+0.1 per thousand year(-1)), C/N ratio increased, and (13)C discrimination relative to atmospheric CO(2) ((13)Delta) increased slightly (+0.03 per thousand year(-1)). Geostatistical analysis of one C. ibex horn revealed systematic patterns of inter-annual and seasonal (13)C changes, but (15)N changed only seasonally. The work demonstrates that isotopic signals in horns are influenced by natural contamination (delta(13)C), age effects ((13)Delta and delta(15)N), and seasonal (delta(13)C and delta(15)N) and inter-annual variation (delta(13)C). The methods presented allow us to distinguish between these effects and thus allow the use of horns as isotopic archives of the ecology of these species and their habitat.

Seasonal diet composition of Pyrenean chamois is mainly shaped by primary production waves.

In alpine habitats, the seasonally marked climatic conditions generate seasonal and spatial differences in forage availability for herbivores. Vegetation availability and quality during the growing season are known to drive life history traits of mountain ungulates. However, little effort has been made to understand the association between plant phenology and changes in the foraging strategies of these mountain dwellers. Furthermore, this link can be affected by the seasonal presence of livestock in the same meadows. The objective of this work was to study the seasonal changes in diet composition of Pyrenean chamois (Rupicapra p. pyrenaica) and its relationship to primary production trends in a Mediterranean alpine environment. Moreover, diet composition in two populations with contrasting livestock pressure was compared in order to study the effect of sheep flocks on the feeding behaviour of chamois. From 2009 to 2012, monthly diet composition was estimated by cuticle microhistological analysis of chamois faeces collected in the eastern Pyrenees. The primary production cycle was assessed by remote sensing, using the Normalized Difference Vegetation Index. Additionally, the diet of sheep sharing seasonally the subalpine and alpine meadows with chamois was analysed. Diet selection of chamois and sheep and their overlap was also assessed. Our results show an intra-annual variation in the diet composition of Pyrenean chamois and demonstrate a strong relationship between plant consumption dynamics and phenology in alpine areas. In addition, Calluna vulgaris, Cytisus spp. and Festuca spp., as well as forbs in the summer, are found to be key forage species for Pyrenean chamois. Furthermore, this study couldn't detect differences between both chamois populations despite the presence of sheep flocks in only one area. However, the detection of a shift in the diet of chamois in both areas after the arrival of high densities of multi-specific livestock suggest a general livestock effect. In conclusion, Pyrenean chamois are well adapted to the variations in the seasonal availability of plants in alpine habitats but could be disturbed by the seasonal presence of livestock. Due to the key plants in their diet, we suggest that population management programmes should focus on the preservation of mixed grasslands composed of patches of shrubs and herbs. The effects of climate change and shrub expansion should be studied as they may potentially affect chamois population dynamics through changes in habitat composition and temporal shifts in forage availability.

The factorial decomposition of the Mahalanobis distances in habitat selection studies.

The Mahalanobis distances have been introduced in habitat selection studies for the estimation of environmental suitability maps (ESMs). The pixels of raster maps of a given area correspond to points in the multidimensional space defined by the mapped environmental variables (ecological space). The Mahalanobis distances measure the distances in this space between these points and the mean of the ecological niche (i.e., the hypothesized optimum for the species) regarding the structure of the niche. The map of these distances over the area of interest is an estimated ESM. Several authors recently noted that the use of a single optimum for the niche of a species may lead to biased predictions of animal occurrence. They proposed to use instead a minimum set of basic habitat requirements, found by partitioning the Mahalanobis distances into a restricted set of biologically meaningful axes. However, the statistical approach they proposed does not take into account the environmental conditions on the area where the niche was sampled (i.e., the environmental availability), and we show that including this availability is necessary. We used their approach as a basis to develop a new exploratory tool, the Mahalanobis distance factor analysis (MADIFA), which performs an additive partitioning of the Mahalanobis distances taking into account this availability. The basic habitat requirements of a species can be derived from the axes of the MADIFA. This method can also be used to compute ESMs using only this small number of basic requirements, therefore including only the biologically relevant information. We also prove that the MADIFA is complementary to the commonly used ecological-niche factor analysis (ENFA). We used the MADIFA method to analyze the niche of the chamois Rupicapra rupicapra in a mountainous area. This method adds to the existing set of tools for the description of the niche.

Semen evaluation in the chamois of Abruzzi (Rupicapra pyrenaica ornata).

The chamois of Abruzzi (Rupicapra pyrenaica ornata) has been classified as endangered by the World Union for Conservation. The objective of this study was to analyze seasonal differences in the characteristics of various male reproductive organs and in semen quality. The study was conduced during 2004 in the reserve of Lama dei Peligni (Italy) on three chamois males aged between 2 and 5 years. Males were captured during March-May months and October-December months. Various testicular and scrotal measurements were taken and semen was collected using an electroejaculator. Sperm motility pattern was evaluated using computer assisted sperm analyzer, membrane integrity using differential staining and morphology with phase contrast microscopy. Testicular size, sperm motility membrane integrity and the percent of morphological normal spermatozoa were greater during October-December. The authors concluded that sperm characteristics are strongly influenced by season and that semen collected in this period (breeding season) has suitable quality for manipulation and long-term preservation.

Effects of acepromazine on the stress response in Southern chamois (Rupicapra pyrenaica) captured by means of drive-nets.

This study was conducted to assess the stress response of Southern chamois (Rupicapra pyrenaica) to capture and physical restraint and the effects of acepromazine (a short-acting neuroleptic) on this response. Forty free-ranging Southern chamois were captured, injected intramuscularly with acepromazine (19 animals, randomly selected) or saline (the other 21 animals), and physically restrained for 3 h. Heart rate and body temperature were monitored with telemetric devices, and blood samples were obtained at capture and every hour thereafter to determine hematologic and serum biochemical parameters. The lower heart-rate variability, temperature, erythrocyte count, hemoglobin concentration, packed cell volume (PCV), and serum creatine kinase activity in the animals treated with acepromazine indicated that this agent reduced the adverse effects of stress. According to the differences in heart rate, erythrocyte count, hemoglobin concentration, PCV, lymphocyte count, and serum concentrations of glucose, creatinine, chloride, and potassium, alpha-adrenergic stimulation by catecholamines seemed to be stronger in females, whereas the adrenal-cortex reaction seemed to be stronger in males. The differences in erythrocyte parameters, temperature, serum creatine kinase activity, and serum concentrations of potassium and chloride indicated that acepromazine's beneficial effects were greater in females.

Steroid hormone related male biased parasitism in chamois, Rupicapra rupicapra rupicapra.

Parasites are linked with their host in a trophic interaction with implications for both hosts and parasites. Interaction stretches from the host's immune response to the structuring of communities and the evolution of biodiversity. As in many species sex determines life history strategy, response to parasites may be sex-specific. Males of vertebrate species tend to exhibit higher rates of parasites than females. Sex-associated hormones may influence immunocompetence and are hypothesised to lead to this bias. In a field study, we tested the prediction of male biased parasitism (MBP) in free ranging chamois (Rupicapra rupicapra rupicapra), which are infested intensely by gastrointestinal and lung helminths. We further investigated sex differences in faecal androgen (testosterone and epiandrosterone), cortisol and oestrogen metabolites using enzyme immunoassays (EIA) to evaluate the impact of these hormones on sex dependent parasite susceptibility. Non-invasive methods were used and the study was conducted throughout a year to detect seasonal patterns. Hormone levels and parasite counts varied significantly throughout the year. Male chamois had a higher output of gastrointestinal eggs and lungworm larvae when compared to females. The hypothesis of MBP originating in sex related hormone levels was confirmed for the elevated output of lungworm larvae, but not for the gastrointestinal nematodes. The faecal output of lungworm larvae was significantly correlated with androgen and cortisol metabolite levels. Our study shows that sex differences in steroid levels play an important role to explain MBP, although they alone cannot fully explain the phenomenon.

Transport stress in Southern chamois (Rupicapra pyrenaica) and its modulation by acepromazine.

Chamois have been translocated for more than 100 years but their stress response to transport remains to be elucidated. In this study, 21 free-ranging Southern chamois were captured, physically restrained and then transported. The animals were randomly injected intramuscularly with acepromazine (nine adult males, one adult female and one yearling male) or saline (five adult males, two adult females and three yearling males). Heart rate and body temperature were monitored with telemetric devices, and blood samples were obtained at capture, immediately before and immediately after transport to determine haematological and serum biochemical parameters. Heart rate and blood parameters (erythrocytes, haemoglobin, packed cell volume, leukocytes, monocytes, band neutrophils, lactate, muscular enzymes and creatinine) indicated that transport was more stressful than previous physical restraint. Acepromazine reduced the adverse consequences of transport stress, as demonstrated by heart rate, body temperature, cortisol, creatinine, muscular enzymes, urea, sodium and potassium.

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.