Seeking temporal refugia to heat stress: increasing nocturnal activity despite predation risk.

Flexibility in activity timing may enable organisms to quickly adapt to environmental changes. Under global warming, diurnally adapted endotherms may achieve a better energy balance by shifting their activity towards cooler nocturnal hours. However, this shift may expose animals to new or increased environmental challenges (e.g. increased predation risk, reduced foraging efficiency). We analysed a large dataset of activity data from 47 ibex (Capra ibex) in two protected areas, characterized by varying levels of predation risk (presence versus absence of the wolf-Canis lupus). We found that ibex increased nocturnal activity following warmer days and during brighter nights. Despite the considerable sexual dimorphism typical of this species and the consequent different predation-risk perception, males and females demonstrated consistent responses to heat in both predator-present and predator-absent areas. This supports the hypothesis that shifting activity towards nighttime may be a common strategy adopted by diurnal endotherms in response to global warming. As nowadays different pressures are pushing mammals towards nocturnality, our findings emphasize the urgent need to integrate knowledge of temporal behavioural modifications into management and conservation planning.

Weather-dependent changes in habitat use by Alpine chamois.

BACKGROUND: Alterations in weather patterns due to climate change are accelerated in alpine environments, but mountains also provide a wide range of niches and potential refuge areas. In order to identify future critical habitat for mountain ungulates for effective protection, it is important to understand their spatial responses to changing weather conditions without movement constraints by human disturbance. METHODS: Using integrated step selection functions, we investigated fine-scale changes in seasonal habitat use in response to weather and time of day for 55 GPS-collared adult Alpine chamois in summer and 42 individuals in winter in a strictly protected area. RESULTS: Chamois reacted to increasing precipitation and wind speeds primarily by moving to lower elevations in summer and winter. However, reactions to high summer temperatures predominantly involved preferences for increasing tree cover density and northerly slopes. Snow depth had little effect on habitat choice, and southerly slopes were preferred in winter regardless of temperature. At night, chamois moved to steeper slopes and lower elevations than during daytime in both seasons, and to more open areas in summer. Steeper slopes were also preferred with increasing tree cover density. CONCLUSIONS: Chamois employ adaptive fine-scale adjustments in their habitat choice consistent with respect to efficient thermoregulation and protection from both weather extremes and predation risk in summer and winter. Movement responses to climate change are therefore expected to be far more complex than simple altitudinal changes in distribution. Particularly the role of forest cover must not be underestimated, as it appears to provide important thermal refuge habitat from high summer temperatures.

Green-up selection by red deer in heterogeneous, human-dominated landscapes of Central Europe.

The forage maturation hypothesis (FMH) assumes that herbivores cope with the trade-off between digestibility and biomass in forage by selecting vegetation at intermediate growth. The green wave hypothesis (GWH) extends the FMH to suggest how spatiotemporal heterogeneity in plant quality shapes migratory movements of herbivores. Growing empirical support for these hypotheses mainly comes from studies in vast landscapes with large-scale habitat heterogeneity. It is unclear, however, to what extent ungulates surf green waves in human-altered landscapes with small-scale heterogeneity in terms of land use and topography. We used plant phenological proxies derived from Sentinel 2 satellite data to analyze the habitat selection of 93 collared red deer (Cervus elaphus) in montane and alpine habitats. Using a step selection analysis, we investigated how plant phenology, that is, the instantaneous rate of green-up (IRG) and normalized difference vegetation index (NDVI), and a set of variables describing topography and human presence influenced red deer resource selection in open habitats. We learned that red deer selected areas with high biomass at green-up and avoided habitats with possible exposure to human activity. Additionally, landscape structure and topography strongly influenced spatial behavior of red deer. We further compared cumulative access to high-quality forage across migrant strategies and found migrants gained better access than residents. Many migratory individuals surfed the green wave, and their surfing behavior, however, became less pronounced with decreasing distance to settlements. Within the constraints of topography and human land use, red deer track spring green-up on a fine spatiotemporal scale and follow the green wave across landscapes in migration movements. Thus, they benefit from high-quality forage even in human-dominated landscapes with small-scale heterogeneity and vegetation emerging in a heterogenic, dynamic mosaic.

Multi-event capture-recapture analysis in Alpine chamois reveals contrasting responses to interspecific competition, within and between populations.

Understanding components of interspecific competition has long been a major goal in ecological studies. Classical models of competition typically consider equal responses of all individuals to the density of competitors, however responses may differ both among individuals from the same population, and between populations. Based on individual long-term monitoring of two chamois populations in sympatry with red deer, we built a multi-event capture-recapture model to assess how vital rates of the smaller chamois are affected by competition from the larger red deer. In both populations, mortality and breeding probabilities of female chamois depend on age and in most cases, breeding status the preceding year. Successful breeders always performed better the next year, indicating that some females are of high quality. In one population where there was high spatial overlap between the two species, the survival of old female chamois that were successful breeders the preceding year (high-quality) was negatively related to an index of red deer population size suggesting that they tend to skip reproduction instead of jeopardizing their own survival when the number of competitors increases. The breeding probability of young breeders (ages 2 and 3) was similarly affected by red deer population size. In contrast, in the second site with low spatial overlap between the two species, the vital rates of female chamois were not related to red deer population size. We provide evidence for population-specific responses to interspecific competition and more generally, for context-, age- and state-dependent effects of interspecific competition. Our results also suggest that the classical assumption of equal responses of all individuals to interspecific competition should be relaxed, and emphasize the need to move towards more mechanistic approaches to better understand how natural populations respond to changes in their environment.

Ungulate management in European national parks: Why a more integrated European policy is needed.

1. Primary objectives of national parks usually include both, the protection of natural processes and species conservation. When these objectives conflict, as occurs because of the cascading effects of large mammals (i.e., ungulates and large carnivores) on lower trophic levels, park managers have to decide upon the appropriate management while considering various local circumstances. 2. To analyse if ungulate management strategies are in accordance with the objectives defined for protected areas, we assessed the current status of ungulate management across European national parks using the naturalness concept and identified the variables that influence the management. 3. We collected data on ungulate management from 209 European national parks in 29 countries by means of a large-scale questionnaire survey. Ungulate management in the parks was compared by creating two naturalness scores. The first score reflects ungulate and large carnivore species compositions, and the second evaluates human intervention on ungulate populations. We then tested whether the two naturalness score categories are influenced by the management objectives, park size, years since establishment, percentage of government-owned land, and human impact on the environment (human influence index) using two generalized additive mixed models. 4. In 67.9% of the national parks, wildlife is regulated by culling (40.2%) or hunting (10.5%) or both (17.2%). Artificial feeding occurred in 81.3% of the national parks and only 28.5% of the national parks had a non-intervention zone covering at least 75% of the area. Furthermore, ungulate management differed greatly among the different countries, likely because of differences in hunting traditions and cultural and political backgrounds. Ungulate management was also influenced by park size, human impact on the landscape, and national park objectives, but after removing these variables from the full model the reduced models only showed a small change in the deviance explained. In areas with higher anthropogenic pressure, wildlife diversity tended to be lower and a higher number of domesticated species tended to be present. Human intervention (culling and artificial feeding) was lower in smaller national parks and when park objectives followed those set by the International Union for the Conservation of Nature (IUCN). 5. Our study shows that many European national parks do not fulfil the aims of protected area management as set by IUCN guidelines. In contrast to the USA and Canada, Europe currently has no common ungulate management policy within national parks. This lack of a common policy together with differences in species composition, hunting traditions, and cultural or political context has led to differences in ungulate management among European countries. To fulfil the aims and objectives of national parks and to develop ungulate management strategies further, we highlight the importance of creating a more integrated European ungulate management policy to meet the aims of national parks.

Does selection on horn length of males and females differ in protected and hunted populations of a weakly dimorphic ungulate?

Weaponry in ungulates may be costly to grow and maintain, and different selective pressures in males and females may lead to sex-biased natural survival. Sexual differences in the relationship between weapon growth and survival may increase under anthropogenic selection through culling, for example because of trophy hunting. Selection on weaponry growth under different scenarios has been largely investigated in males of highly dimorphic ungulates, for which survival costs (either natural or hunting related) are thought to be greatest. Little is known, however, about the survival costs of weaponry in males and females of weakly dimorphic species. We collected information on horn length and age at death/shooting of 407 chamois Rupicapra rupicapra in a protected population and in two hunted populations with different hunting regimes, to explore sexual differences in the selection on early horn growth under contrasting selective pressures. We also investigated the variation of horn growth and body mass in yearling males (n = 688) and females (n = 539) culled in one of the hunted populations over 14 years. The relationship between horn growth and survival showed remarkable sexual differences under different evolutionary scenarios. Within the protected population, under natural selection, we found no significant trade-off in either males or females. Under anthropogenic pressure, selection on early horn growth of culled individuals showed diametrically opposed sex-biased patterns, depending on the culling regime and hunters' preferences. Despite the selective bias between males and females in one of the hunted populations, we did not detect significant sex-specific differences in the long-term pattern of early growth. The relationship between early horn growth and natural survival in either sex might suggest stabilizing selection on horn size in chamois. Selection through culling can be strongly sex-biased also in weakly dimorphic species, depending on hunters' preferences and hunting regulations, and long-term data are needed to reveal potential undesirable evolutionary consequences.

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.

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.

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.

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.

Detection of Mycoplasma conjunctivae in the eyes of healthy, free-ranging Alpine ibex: possible involvement of Alpine ibex as carriers for the main causing agent of infectious keratoconjunctivitis in wild Caprinae.

Mycoplasma conjunctivae is considered the major cause of infectious keratoconjunctivitis (IKC) in Alpine ibex (Capra i. ibex) and chamois (Rupicapra r. rupicapra). While it is known that domestic sheep can act as healthy carriers for M. conjunctivae, this question has not been addressed in wild ungulates so far. In this study, bacteriological investigations and field observations were performed to assess whether free-ranging Alpine ibex can be healthy carriers of M. conjunctivae. Among 136 ibex without clinical signs of IKC, M. conjunctivae was identified 26 times (19.1%) by TaqMan PCR. To assess the potential pathogenicity of M. conjunctivae strains isolated from asymptomatic eyes, strains from three healthy ibex and from 15 IKC-ibex and IKC-chamois were analysed genetically by DNA sequence analysis of the variable part of the lppS gene. No significant differences were observed between strains from asymptomatic and clinically affected animals, reflecting the assumption that healthy ibex may act as carriers for M. conjunctivae strains that may be pathogenic for other individuals. Our results further indicate that development of IKC is associated with M. conjunctivae load in the eyes. In addition, a questionnaire survey revealed that IKC is generally less common in ibex than chamois and that infection in wild ungulates is not necessarily linked to the presence of sheep. These data support the hypothesis that apparently healthy ibex may be important in the epizootiology of IKC and indicate that host predilection may play a role in IKC development.

Predicting fluctuations of reintroduced ibex populations: the importance of density dependence, environmental stochasticity and uncertain population estimates.

1. Development of population projections requires estimates of observation error, parameters characterizing expected dynamics such as the specific population growth rate and the form of density regulation, the influence of stochastic factors on population dynamics, and quantification of the uncertainty in the parameter estimates. 2. Here we construct a Population Prediction Interval (PPI) based on Bayesian state space modelling of future population growth of 28 reintroduced ibex populations in Switzerland that have been censused for up to 68 years. Our aim is to examine whether the interpopulation variation in the precision of the population projections is related to differences in the parameters characterizing the expected dynamics, in the effects of environmental stochasticity, in the magnitude of uncertainty in the population parameters, or in the observation error. 3. The error in the population censuses was small. The median coefficient of variation in the estimates across populations was 5.1%. 4. Significant density regulation was present in 53.6% of the populations, but was in general weak. 5. The width of the PPI calculated for a period of 5 years showed large variation among populations, and was explained by differences in the impact of environmental stochasticity on population dynamics. 6. In spite of the high accuracy in population estimates, the uncertainty in the parameter estimates was still large. This uncertainty affected the precision in the population predictions, but it decreased with increasing length of study period, mainly due to higher precision in the estimates of the environmental variance in the longer time-series. 7. These analyses reveal that predictions of future population fluctuations of weakly density-regulated populations such as the ibex often become uncertain. Credible population predictions require that this uncertainty is properly quantified.

Seroprevalence of anaplasmosis among cattle in Switzerland in 1998 and 2003: no evidence of an emerging disease.

Anaplasma marginale infection in Europe has been limited to the Mediterranean and eastern countries, to Austria and to very sporadic cases in Switzerland. There are no reports of its occurrence in the countries north of Switzerland. A severe outbreak of anaplasmosis in August 2002 in a cattle farm in the canton Grisons, Switzerland, north of the Alps, with more than 300 cattle that had to be culled, came unexpected and gave reason to hypothesize presence of an increased yet undetected prevalence of A. marginale in Switzerland. Randomly selected bovine serum samples collected in 1998 and 2003 were tested using a competitive inhibitory ELISA (cELISA) to test the hypothesis. Our validation of the diagnostic sensitivity and specificity of this test, done in the outbreak herd, yielded 99.2 and 83.3%, respectively, probably underestimating the true specificity. The true seroprevalence of anaplasmosis in Swiss cattle determined by cELISA was likely to be zero with upper 95% confidence limits of 2.49% in the canton Grisons and 1.17% in the rest of Switzerland, respectively, in 1998. For 2003, these estimates were even lower. There was no significant difference in apparent prevalences between 1998 and 2003. In search of a possible reservoir, three chamoises out of 46 free ranging wild ruminants from the Swiss National Park, Grisons, tested positive in the cELISA. This reaction is in accordance with A. marginale or a cross reacting agent such as Anaplasma ovis. From our results we conclude that the hypothesis of an increased prevalence of anaplasmosis in cattle in Switzerland must be rejected.