Spatiotemporal behavior of predators and prey in an arid environment of Central Asia.

The mechanisms of interactions between apex and smaller carnivores may range from competition to facilitation. Conversely, interactions between predators and prey are mainly driven by the prey reducing the likelihood of encounters with predators. In this study, we investigated (1) the spatiotemporal interactions between an apex (the snow leopard) and a meso-predator (the red fox), and (2) the temporal interactions between the snow leopard and its potential prey (Siberian ibex, argali, Asian wild ass, Tolai hare) through camera trapping in the Mongolian Great Gobi-A. The probability of occurrence for the red fox was higher in the presence of the snow leopard than in its absence. Moreover, the red fox activity pattern matched that of the snow leopard, with both species mostly active at sunset. This positive spatiotemporal interaction suggests that the presence of the snow leopard may be beneficial for the red fox in terms of scavenging opportunities. However, other explanations may also be possible. Amongst prey, the Siberian ibex and the argali were mainly active during the day, whereas the Asian wild ass and the Tolai hare were more nocturnal. These findings suggest that potential prey (especially the Siberian ibex and the argali) may shape their behavior to decrease the opportunity for encounters with the snow leopard. Our results have revealed complex interactions between apex and smaller predators and between apex predator and its potential prey.

Animal conflicts escalate in a warmer world.

The potential for climate change to affect animal behaviour is widely recognized, yet its possible consequences on aggressiveness are still unclear. If warming and drought limit the availability of food resources, climate change may elicit an increase of intraspecific conflicts stemming from resource competition. By measuring aggressivity indices in a group-living, herbivorous mammal (the Apennine chamois Rupicapra pyrenaica ornata) in two sites differing in habitat quality, and coupling them with estimates of plant productivity, we investigated whether harsh climatic conditions accumulated during the growing season influenced agonistic contests at feeding via vegetation-mediated effects, and their interaction with the site-specific habitat quality. We focused on females, which exhibit intra-group contest competition to access nutritious food patches. Accounting for confounding variables, we found that (1) the aggression rate between foraging individuals increased with the warming accumulated over previous weeks; (2) the probability to deliver more aggressive behaviour patterns toward contestants increased with decreasing rainfall recorded in previous weeks; (3) the effects of cumulative warming and drought on aggressivity indices occurred at time windows spanning 15-30 days, matching those found on vegetation productivity; (4) the effects of unfavourable climatic conditions via vegetation growth on aggressivity were independent of the site-specific habitat quality. Simulations conducted on our model species predict a ~50 % increase in aggression rate following the warming projected over the next 60 years. Where primary productivity will be impacted by warming and drought, our findings suggest that the anticipated climate change scenarios may trigger bottom-up consequences on intraspecific animal conflicts. This study opens the doors for a better understanding of the multifactorial origin of aggression in group-living foragers, emphasising how the escalation of agonistic contests could emerge as a novel response of animal societies to ongoing global warming.

The scale-dependent effectiveness of wildlife management: A case study on British deer.

Impacts of herbivory by wild ungulates represent a significant issue world-wide. To be effective, management of populations and impacts needs to be coordinated above the site scale, yet little research has investigated the appropriate spatial scale over which management should be integrated to be fully effective. In consideration of reduction of impacts in deciduous or mixed woodland habitats, we tested scale-specific management effectiveness in a lowland area of UK where moderate- to high-density populations of four deer species were the target of deliberate control programmes, and nonhuman predators were absent. We modelled the annual impact recorded between 2009 and 2015 in 98 woodlands as a function of cumulative culls of deer taken since the commencement of management. Analysis was repeated at different spatial scales by increasing the circular area around each focal woodland, from 2.5 km-radius up to 100 km-radius. Our findings suggest for the first time the geographical scale over which deer management needs to be coordinated for optimum effectiveness in decreasing their impact on woodland across relatively homogenous landscapes. For small bodied and relatively sedentary species (roe deer Capreolus capreolus; Reeves' muntjac Muntiacus reevesi), reductions in impacts within woodlands can be achieved by culling at the immediately local level, but some modest increase in effectiveness (probably relating to reductions in the degree of source-sink movement) may be expected with an increase in spatial scale of culling to around 30-70 km-radius. For larger-bodied, herding species with more extensive home-ranges (fallow deer Dama dama; red deer Cervus elaphus) management for reduction of woodland impacts was only really effective when coordinated above the single woodland-scale, with marked increases shown again up to a scale of 100 km-radius. Whilst future studies for different landscape types are still needed, our work emphasises that the spatial scale at which control plans are conducted can determine the effectiveness of wildlife management, possibly providing an advance on how to manage wildlife populations more effectively.

On the Behavioural Biology of the Mainland Serow: A Comparative Study.

Comparative behavioural studies help reconstruct the phylogeny of closely related species. In that respect, the serows Capricornis spp. occupy an important position as they have been assumed to be the closest forms to the ancestors of Caprinae. In spite of that, information on the behavioural repertoire of the mainland serow Capricornis sumatraensis is exceedingly poor. In this paper, we report data on the activity rhythms and social behaviour of rutting mainland serows in captivity (Central Thailand, January 1986; January-February 1987). Activity was bimodal with peaks in mid-afternoon and late night. Resting and ruminating peaked at noon and twilight. Four patterns of marking behaviour were observed out of a total of 1900 events. Males and females were found to use different marking sites and frequencies. A total of 33 social behaviour patterns were observed: 18 patterns concerned agonistic behaviour, whereas 15 patterns were relevant to courtship behaviour. A comparison across Caprinae species with unritualised piercing weapons (i.e., Capricornis, Naemorhedus, Rupicapra, Budorcas, and Hemitragus) has shown that inter-sexual direct forms of aggressive behaviour are used significantly more often than indirect ones, but for chamois, confirming Rupicapra spp. as the most advanced genus among them in terms of an early ritualisation of weapons. Conversely, horns of the goral Nemorhaedus spp. and the serow lie on the same plane of the frontal bones, thus making possible the usage of a dominance display through frontal pushing.

Temporal variation in foraging activity and grouping patterns in a mountain-dwelling herbivore: Environmental and endogenous drivers.

In temperate ecosystems, seasonality influences animal behaviour. Food availability, weather, photoperiod and endogenous factors relevant to the biological cycle of individuals have been shown as major drivers of temporal changes in activity rhythms and group size/structure of herbivorous species. We evaluated how diurnal female foraging activity and grouping patterns of a mountain herbivore, the Apennine chamois Rupicapra pyrenaica ornata, varied during a decreasing gradient of pasture availability along the summer-autumn progression (July-October), a crucial period for the life cycle of mountain ungulates. Females increased diurnal foraging activity, possibly because of constrains elicited by variation in environmental factors. Size of mixed groups did not vary, in contrast with the hypothesis that groups should be smaller when pasture availability is lower. Proportion of females in groups increased, possibly suggesting that they concentrated on patchily distributed nutritious forbs. Occurrence of yearlings in groups decreased, which may have depended on dispersal of chamois in this age class. Presence of kids in groups did not show variation through summer-autumn, suggesting a close mother-juvenile relationship even at the end of weaning and/or, possibly, low summer mortality. Both endogenous and environmental factors contribute to shape variation in foraging activity and grouping behaviour in mountain-dwelling herbivores.

Joint effects of weather and interspecific competition on foraging behavior and survival of a mountain herbivore.

Weather variations have the potential to influence species interactions, although effects on competitive interactions between species are poorly known. Both weather and competition can influence foraging behavior and survival of herbivores during nursing/weaning, a critical period in the herbivore life cycle. We evaluated the joint effects of weather and competition with red deer Cervus elaphus on the foraging behavior of adult female Apennine chamois Rupicapra pyrenaica ornata in summer, and on winter survival of chamois kids. High temperature and low rainfall during the growing season of vegetation had negative effects on bite rate. Effects of weather were greater in forb patches, including cold-adapted, nutritious plants of key importance to chamois, than in graminoid ones. Our results confirm previous indications of a negative effect of competition on bite rate of female chamois and on kid survival. Furthermore, harsh weather conditions and competition with deer had additive, negative roles on foraging behavior and survival of chamois. Growing temperatures are expected to influence distribution, growth, and/or nutritional quality of plants; competition would reduce pasture quality and food availability through resource depletion. Both factors would limit food/energy intake rates during summer, reducing survival of the youngest cohorts in winter. We suggest that interspecific competition can be an important additive factor to the effects of weather changes on behavior and demography.

Woodland features determining home range size of roe deer.

Use of ecotones by ungulates may be mediated by their movements between main feeding areas and woodland, where they locate their shelter. The roe deer Capreolus capreolus has been termed as a woodland species, although we suggest that it did not evolve as a forest ungulate, but depending on forest glades. Roe deer feed on a wide range of vegetal species, although their diet is mainly dominated by woody plants. Our study was carried out in a fragmented area covered with small forest patches of Mediterranean "macchia" scrubwood, interspersed in an agricultural matrix. Aim of our study has been to test how ranging movements of roe deer are influenced by landscape heterogeneity and to evaluate which features of woodland affect home range size. Radio-locations of 22 female and 12 male adult roe deer, monitored for three years, were used to assess home range size. A linear mixed model was fitted to investigate variation in home range size according to eleven spatial parameters estimated to describe home range size and composition. Throughout the year, no significant difference was found between home range sizes of males (median: 16.70ha, Q1-Q3: 13.20-31.60ha) and females (median: 23.52ha, Q1-Q3: 13.30-44.00ha: lme: F=0.9; P=0.35). Habitat density, edge density, percentage of woodland within home range and woodland structure determined home range size. Home ranges with few habitat types and a small amount of wood were large, while roe deer occupied small home ranges when habitat density was high and when a high proportion of wood was concentrated in a single large patch. Woodland covered a mean±SE of 36.2±17.9% in each home range. In conclusion, roe deer seem to be particularly well adapted to live in human transformed, peripheral habitats, e.g. farmlands, as long as a minimum quantity of woodland is included within their HR.

Predation may counteract climatic change as a driving force for movements of mountain ungulates.

Temperature variations are expected to influence altitudinal movements of mountain herbivores and, in turn, those of their predators, but relevant information is scarce. We evaluated monthly relationships between temperature and altitude used by a large mountain-dwelling herbivore, the Himalayan tahr Hemitragus jemlahicus, and its main predator, the snow leopard Panthera uncia, in an area of central Himalaya for five consecutive years (2006-2010). In contrast to expectations, there was no significant direct relationship between altitude of tahr sightings and temperature. The mean altitude of tahr sightings decreased by c. 200m throughout our study. As expected, snow leopard movements tracked those of tahr, although the core area of the snow leopard did not move downwards. Tahr remained the staple of the snow leopard diet: we suggest that the former did not move upwards in reaction to higher temperature to avoid encounters with the latter. Avoidance of competition with the larger common leopard Panthera pardus at lower altitudes could explain why snow leopards did not shift their core area downwards. Apparently, interspecific interactions (predation; competition) influenced movements of Himalayan tahr and snow leopards more than climatic variations.

Nocturnal activity of a "diurnal" species, the northern chamois, in a predator-free Alpine area.

The reduction of predation risk is widely considered a major factor affecting the nocturnal activity of mammals. Furthermore, on precipitous mountain terrain, moving in very poor light conditions should be avoided by animals with no special eyesight adaptation to darkness. The Northern chamois Rupicapra rupicapra has been for long considered as a diurnal species, with occasional nocturnal movements. For the first time, we have quantified the nocturnal activity of 21 radiotagged female chamois from the Italian Eastern Alps (Paneveggio-Pale di San Martino Natural Park), continuously monitored for two years from sunset to sunrise, with 24h tracking sessions carried out for six months. Large predators were not present in the study site. Despite their mainly diurnal activity pattern, peaks of nocturnal movements were detected throughout the year. The least proportion of active night fixes occurred in January and in July, while the most were in April and in October. The greater nocturnal activity in the warm months compared to cold periods, was probably due to frozen snow cover reducing nocturnal movements. Movements were mainly concentrated in bright moonlight nights, possibly because of the absence of large predators, but more likely because of increased visibility. Changes in activity levels throughout the year may also reflect changes in energy requirements of Northern chamois.

Ectoparasite load in the crested porcupine Hystrix cristata Linnaeus, 1758 in Central Italy.

The crested porcupine Hystrix cristata is a large body-sized rodent, occurring in Europe only in the Italian Peninsula, where it may have been introduced in early Medieval times. Its parasite fauna is currently poorly known and limited to few anecdotal observations. We have analyzed the ectoparasite load of 165 crested porcupines from Tuscany and Latium (Central Italy). Both captured and road-killed individuals were checked for fleas and ticks. Overall, only 39 porcupines were infested by four species of ticks and five of fleas. Abundance of ectoparasites was higher in areas with higher habitat richness, with respect to densely wooded areas. The most frequent species was the flea Pulex irritans (25%), whose prevalence peaked in winter probably because of optimal abiotic conditions in the porcupine's den. The remaining species of both hard ticks (Rhipicephalus bursa, Pholeoixodes hexagonus, and Ixodes ventalloi) and fleas (Paraceras melis, Ctenocephalides canis, Dasypsyllus gallinulae, and Hystrichopsylla talpae), all with prevalence lower than 5%, could be due to den sharing with other vertebrates, mainly carnivores such as, e.g., red foxes and badgers. The second most prevalent species was the generalist tick Ixodes ricinus (21%). An adult male-biased parasitism for ticks has been detected, suggesting a possible role of testosterone related immune-depressive effect. The low richness in dominant ectoparasite species, built up by locally acquired generalist taxa, provides support to the allochthonous origin of this rodent in Italy.

Patterns of spatial overlap in a monogamous large rodent, the crested porcupine.

The crested porcupine Hystrix cristata is a large rodent which pairs for life. We studied the space use of 17 female and 9 male radio-tracked porcupines in an evergreen coastal woodland ("macchia", EW) and in an agricultural estate (AE), with special reference to the use of cultivations. Home range sizes of male porcupines (4.72-323.40) ranged around 114ha (median) during the warm period (April-September) and 162ha during the cold one (October-March). Home ranges of females (2.48-323.40) were c. 91ha during the warm period and c. 143ha during the cold one. Habitat composition and selection changed from the cold to the warm months, with porcupines being present in agricultural areas especially in the latter. Home range overlap between members of the same pair varied from 57% to 97% (median, 75%). Habitat selection was analyzed at the second (within study area) and at the third (within home range) order of selection. Within study areas, porcupines avoided cultivations and selected habitats with dense vegetation, providing cover and food. Within home ranges, in the warm period, porcupines selected agricultural areas in EW, where this habitat represented a minor portion of the study site. In that season, the Mediterranean "macchia" is a poor source of food, forcing porcupines to travel long distances to reach feeding sites. No significant difference of habitat selection within home ranges was detected between members of the same pair. Cultivations may play a key-role for porcupine survival, especially in poor habitats, as they provide abundant food resources in the warm period.

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.

Habitat richness affects home range size in a monogamous large rodent.

In monogamous species, after pair formation, the main reason for ranging movements is not searching for a mate, but for other important resources e.g. food. We monitored a total of 20 radio-tagged adult, paired crested porcupines in four areas of different habitat richness. No sexual size dimorphism was assessed. Body mass and habitat richness showed collinearity. For both sexes, home range size was correlated to habitat richness, with a significant inverse exponential regression. Opposite to natural foragers, living in poor habitats, crop foragers had smaller home ranges, with their dens significantly closer to cultivations. Both availability of food resources and den sites are key variables to determine home range size.

Rutting behaviour of territorial and nonterritorial male chamois: is there a home advantage?

Males using alternative male mating tactics (AMTs) may express their mating effort in a variety of ways. In polygynous species with limited sexual dimorphism, differences in male aggressiveness may affect mating opportunities. We recorded the behaviour of 8 territorial and 7 nonterritorial male Alpine chamois Rupicapra rupicapra, a nearly monomorphic ungulate, during the 2011 rut in the Gran Paradiso National Park (Italy), to analyse differences in mating effort and mating opportunities between AMTs. The chamois showed a rich behavioural repertoire (31 behavioural patterns), with a prevalence of indirect aggression. Territorial males had higher frequency of aggressive and courtship behaviour than nonterritorial males over the early rut. Later, nonterritorials increased their mating effort, possibly because of reduced competition with dominant males. Territorial males monopolised all observed mating events. Our results support the hypothesis that chamois may assert dominance through intense aggressiveness rather than through horn size or body mass as found in other polygynous ungulates. Most important, differences in mating effort mediated by AMTs resulted in different mating opportunities; these benefits, however, are traded off against greater costs, due to higher levels of hormone metabolites and parasitism. Data on AMTs flexibility, lifetime reproductive success and survival are needed to clarify the mechanisms underlying the evolution and maintenance of AMTs within chamois populations.

Effects of sampling regime on the mean and variance of home range size estimates.

1. Although the home range is a fundamental ecological concept, there is considerable debate over how it is best measured. There is a substantial literature concerning the precision and accuracy of all commonly used home range estimation methods; however, there has been considerably less work concerning how estimates vary with sampling regime, and how this affects statistical inferences. 2. We propose a new procedure, based on a variance components analysis using generalized mixed effects models to examine how estimates vary with sampling regime. 3. To demonstrate the method we analyse data from one study of 32 individually marked roe deer and another study of 21 individually marked kestrels. We subsampled these data to simulate increasingly less intense sampling regimes, and compared the performance of two kernel density estimation (KDE) methods, of the minimum convex polygon (MCP) and of the bivariate ellipse methods. 4. Variation between individuals and study areas contributed most to the total variance in home range size. Contrary to recent concerns over reliability, both KDE methods were remarkably efficient, robust and unbiased: 10 fixes per month, if collected over a standardized number of days, were sufficient for accurate estimates of home range size. However, the commonly used 95% isopleth should be avoided; we recommend using isopleths between 90 and 50%. 5. Using the same number of fixes does not guarantee unbiased home range estimates: statistical inferences differ with the number of days sampled, even if using KDE methods. 6. The MCP method was highly inefficient and results were subject to considerable and unpredictable biases. The bivariate ellipse was not the most reliable method at low sample sizes. 7. We conclude that effort should be directed at marking more individuals monitored over long periods at the expense of the sampling rate per individual. Statistical results are reliable only if the whole sampling regime is standardized. We derive practical guidelines for field studies and data analysis.

Chemical immobilization of crested porcupines with tiletamine HCl and zolazepam HCl (Zoletil) under field conditions.

The combination of tiletamine HCl and zolazepam HCl has been used on many species of wild mammals. Short induction time, low dosage, satisfactory safety margins, relatively constant immobilization time, and smooth recovery are benefits reported. This combination (Zoletil 100) was used during a study on behavioural ecology of the crested porcupine (Hystrix cristata) in a Mediterranean coastal area (Maremma Regional Park, Tuscany, Italy). We used this mixture 42 times on 31 individuals. Mean adult dose was (+/- SE) 7.24 +/- 0.37 mg/kg (74.0 +/- 3.0 mg/individual). Average adult induction time was 5.3 min (+/- 1.1) and average adult immobilization time was 22.6 min (+/- 6.0). One adult male porcupine died after chemical restraints. The use of tiletamine-zolazepam seems adequate for chemical immobilization of crested porcupines under field conditions, mainly because of its short induction time, small volume to be injected and wide safety margin.