A comparison of four different methods to estimate population size of Alpine marmot (Marmota marmota)

Obtaining reliable information on animal abundance in mountainous landscapes is challenging. Highly heterogeneous habitats tend to reduce detection probabilities, and the three-dimensional, rugged nature of the terrain poses severe limits to the fulfilment of a number of assumptions underlying several statistical methods. In this study, we aimed to compare the performance of 4 different methods to estimate population size of Alpine marmot (Marmota marmota), a highly social semifossorial rodent widely distributed on the European Alps. Between May and August 2015, in a study area within the Stelvio National Park (Italy) we conducted 8 sessions of capture-mark-recapture, 6 sessions of mark-resight from vantage points, 8 sessions of line distance sampling along 4 transects, and 2 sessions using double-observer methods from vantage points. The minimum number of animals alive, obtained during the mark-resight surveys, was n=54 individuals. Capture-mark-recapture models estimated a population size of n=56 individuals [95% CI (45,87)]; similar, but more precise estimates were obtained with the mark-resight approach {Bowden’s estimator: n=62 [95% CI (54,71)]; Poisson log-normal estimator: n=62 [95% CI (55,69)]}. Line distance sampling and double-observer methods were severely biased low {Line distance sampling: n=24 individuals [95% CI (19,31)]; Independent double-observer: n=24 [95% CI (22, 35)]; Dependent double-observer: n=15 [95% CI (15,20)]}. Our results suggest that the probabilistic approach based on marked individuals yielded fairly robust estimates of population size. The underestimates obtained using distance sampling and double-observer methods were likely due to the violation of some underlying assumptions. While the topography of the mountainous landscape makes it difficult to randomize the sampling scheme, the semifossorial behaviour of the target species is likely to lower the detection probabilities and violate the assumption of perfect detection on the transect

Face markings in Northern chamois: cues of dominance?

AbstractBlack and white face markings in Artiodactyls are thought to be broadly associated with intraspecific communication. Evidence-based studies on the fine-scale mechanisms of signaling, however, are scant. The occurrence of black and white face markings is one of the most distinctive features of the Northern chamoisRupicapra rupicapra. Although their function is unknown, they might possibly signal fighting abilities: if so, the intensity of face color contrast should positively correlate with traits involved in aggressive contests, i.e. body mass and horn size. We collected data on face marking 'blackness' as a proxy of color contrast, dressed body mass and horn length on 103 chamois of either sex harvested during the hunting season of 2019 in the central Italian Alps. To investigate the relationship between color contrast and sex-specific body mass and horn length, we fitted an ordinal multilevel regression model in a Bayesian framework. Contrast score positively associated with increasing body mass in both sexes, but this relationship was much stronger in females than in males. Contrast score positively related with horn length in males but not in females. Our results offer some first insights into the potential correlates of chamois facial mask, suggesting that face markings might provide age-specific cues of mass-based dominance in females, while their role in males appears more uncertain. Behavioral studies on marked or hunted individuals in different environmental settings are necessary to confirm these findings, and provide further understandings of face color patterns in chamois

Linking alternative reproductive tactics and habitat selection in Northern chamois

In polygynous ungulates, males may achieve fertilization through the use of alternative reproductive tactics (ARTs), discrete phenotypic variations evolved to maximize fitness. ARTs are often associated with different male spatial strategies during the rut, from territoriality to female-following. Although variation in space use patterns of rutting male ungulates is known to be largely affected by the spatial distribution of females, information on the year-round habitat selection of alternative reproductive types is scant. Here, we investigate the seasonal variation in habitat choice of a large mammal with ARTs (territoriality and nonterritoriality), the Northern chamois Rupicapra rupicapra. Global Positioning System (GPS) data on 28 adult males were collected between February 2010 and December 2013 in the Gran Paradiso National Park (Italy) and used to fit resource selection functions to explore the ART-specific use of key topographic features, such as elevation, aspect, and slope, and vegetation phenology expressed as NDVI values. Territorial and nonterritorial chamois profoundly differed in their habitat selection not only during the rutting season. Compared to nonterritorial males, territorial males used lower elevations in summer and autumn, preferred southern slopes in spring and summer, and used steeper areas in summer but not in winter. We found no difference in seasonal selection of NDVI values between males adopting ARTs. Our results suggest that territorial males tend to occupy warmer, lower-food-quality habitats in late spring and summer, whereas nonterritorial males are free to follow and exploit vegetation phenology and more favorable temperatures. Different patterns of habitat selection may reflect different trade-offs between the optimization of energy balances throughout the year and the increase of mating opportunities during the rut in males adopting alternative reproductive tactics

Active surveillance of paratuberculosis in Alpine-dwelling red deer (Cervus elaphus)

Paratuberculosis (Johne’s disease) is a globally widespread infectious disease affecting domestic and wild ruminants, caused by Mycobacterium avium subsp. paratuberculosis (MAP). The bacterium is excreted in the feces and is characterized by high environmental resistance. The new Animal Health Law (Regulation EU 2016/429) on transmissible animal diseases, recently in force throughout the European Union, includes paratuberculosis within the diseases requiring surveillance in the EU, listing some domestic and wild Bovidae, Cervidae, and Camelidae as potential reservoirs. Taking advantage of a culling activity conducted in the Stelvio National Park (Italy), this study investigated MAP infection status of red deer (Cervus elaphus) between 2018 and 2022, and evaluated the probability of being MAP-positive with respect to individual and sampling-level variables. A total of 390 subjects were examined macroscopically and tested for MAP, using different diagnostic tools: IS900 qPCR, culture, histopathology, and serology. Twenty-three of them were found positive for MAP by at least one test, with an overall prevalence of 5.9% (95% CI 4.0–8.7), that, respectively, ranged from 12.4% in the first culling season to 2.0 and 2.1% in the 2019–2020 and 2021–2022 culling seasons. Quantitative PCR assay on ileocecal valve and mesenteric lymph nodes detected the highest number of MAP positive animals. The results of the study showed the increased probability of being MAP-positive with increasing age and that red deer with lower body mass values were more likely to be infected with MAP. Overall, the absence of signs of clinical paratuberculosis and gross lesions together with the low level of shedding witness early phases of the disease among the positive red deer and support an improvement of the paratuberculosis status of this population, as shown by the decreased prevalence of the disease over the years

Male-biased gastrointestinal parasitism in a nearly monomorphic mountain ungulate

Background: Pyrenean chamois (Rupicapra pyrenaica pyrenaica) is a nearly monomorphic mountain ungulate with an unbiased sex-specific overwinter adult survival. Few differences in gastrointestinal parasitism have been reported by coprology as yet. This study aims to assess diversity, prevalence, intensity of infection and aggregation of gastrointestinal nematodes in male and female adult chamois. We expect no differences in the parasite infection rates between sexes. Findings: Gastrointestinal tracts of 28 harvested Pyrenean chamois in the Catalan Pyrenees (autumn 2012 and 2013) were necropsied and sexual differences in the diversity and structure of parasite community, prevalence, intensity of infection, and richness were investigated. We found 25 helminth species belonging to 13 different genera. Conclusions: Contrary to our expectations, male chamois showed different parasite communities, higher prevalence, intensity of infection and richness than females. Such sexual differences were clear irrespective of age of individuals. Hence, male chamois must cope with a more diverse and abundant parasite community than females, without apparent biological cost. Further research will be required to confirm this hypothesis

The use of block counts, mark-resight and distance sampling to estimate population size of a mountain-dwelling ungulate

Population size estimates represent indispensable tools for many research programs and for conservation or management issues. Mountain ungulates in open areas are often surveyed through ground counts that normally underestimate population size. While the use of sample counts is desirable, few studies have compared different probabilistic approaches to estimate population size in this taxon. We compared the size estimates of a male population of Alpine chamois using mark-resight and line transect sampling methods, while block counts were used to obtain the minimum number of males alive in the study area. Surveys were conducted within the Gran Paradiso National Park (Italy), in August–September 2013, using block counts along purposely selected trails and vantage points, mark-resight over 5 consecutive resightings from vantage points and trails, and line transect sampling along 12 transects repeated 8 times. Block counts yielded a minimum number of males alive in the population of n = 72 individuals. This value was greater than the upper bound of the 95 % confidence interval achieved using line transect sampling {n = 54, CV = 14 % [95 % CI (40, 71)]} while mark-resight yielded a more realistic result of n = 93 individuals {CV = 18 % [95 % CI (63, 137)]}. Our results suggest that line transect sampling performed poorly in the Alpine environment, leading to underestimates of population size, likely due to violations of some assumptions imposed by the rugged nature of the terrain. The mark-resight yielded lower precision, possibly due to the limited number of marked individuals and resighting occasions, but it provided robustness and accurate estimates as marks were evenly distributed among animals

Disentangling demographic effects of red deer on chamois population dynamics

Abstract 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

Corlatti_et_al_Parasites & ARTs chamois_Supporting Information_2

Parasite data and individual / environmental covariates

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. nonterritorial 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 oldermales in agonistic contests. Social status was an important determinant of FCMexcretion: 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

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