Sénescence, sélection sexuelle et dynamique de population du bouquetin des Alpes (Capra Ibex)

Le principal objectif de cette thèse est d’étudier les variations intra et inter individuelle pour deux traits phénotypiques reliés à la valeur sélective, et d’analyser la dynamique d’une population sauvage de bouquetin des Alpes (Capra ibex). Le premier chapitre analyse la relation de deux traits phénotypiques reliés à la valeur sélective (le gain de masse corporelle, et l’intensité d’infection parasitaire gastro-intestinal) et d’un caractère sexuel secondaire (la croissance des cornes) avec l’âge, des facteurs environnementaux, la survie, et avec l’hétérozygotie, ceci au sein d’un large échantillon de bouquetins des Alpes mâles en libertés et marqués individuellement. À travers la croissance corporelle et les comptages fécaux d’oeufs de nématodes gastro-intestinaux, il a été mis en évidence un phénomène de sénescence et de coût pour la reproduction. En analysant un large échantillon de squelettes de bouquetins mâles trouvés morts dans l’hiver pour cause de famine, nous avons trouvé que le début de la sénescence était caractérisé par la longueur de segments de croissance annuels âges spécifiques des cornes, mais pas par leur asymétrie. Enfin, des corrélations entre l’hétérozygotie et la valeur sélective ont été mises en évidence pour la croissance des cornes, mais pas pour la masse corporelle et l’intensité d’infection parasitaire. Le deuxième chapitre traite de l’importance relative de la densité dépendance et de la variabilité stochastique du climat sur la dynamique d’une population d’ongulé de montagne, à travers l’analyse de série temporelle de 45 ans de recensements de bouquetins des Alpes collectés dans le Parc National du Grand Paradis en Italie. Pendant les 28 premières années de l’étude, le nombre total de bouquetins a varié entre 2600 et 4000 sans tendance visuelle apparente. Au cours de cette période, il est apparu que des oscillations avec une périodicité de 3 à 8 ans étaient présentes. Dès 1982, les comptages de bouquetin ont augmentés régulièrement pour atteindre un pic de 5000 en 1993 et ensuite décroître. Nous avons montré que la taille de population du bouquetin était limitée à la fois par la densité dépendance et par l’épaisseur de neige. Un modèle basé sur ces deux facteurs et ajusté pour les 19 premières années de données, réussi avec succès à prédire l’augmentation et le déclin subséquent de la taille totale de la population pour les 20 dernières années de l’étude. Cette thèse conclue avec un troisième chapitre présentant deux techniques innovatrices utiles pour l’étude de populations d’ongulés en liberté.Abstract: The main objectives of this thesis are to investigate within and between-individual variation in fitness related phenotypic traits and to analyse the population dynamics of a wild population of Alpine ibex (Capra ibex). The first chapter analyzes the relationship of two fitness-related phenotypic traits (body mass gain and the intensity of gastrointestinal parasite infection) and one secondary sexual trait (horn growth) with age, environmental factors, survival and heterozygosity in a large sample of individually tagged free-ranging Alpine ibex males. Evidence for senescence and costs of reproduction in body growth and fecal counts of gastrointestinal nematode eggs was found. Analysing a large sample of male ibex skulls, found dead in winter from starvation, we found that the onset of senescence was signaled by the length of age specific yearly horn growth segments, but not by their asymmetry. Finally evidence for heterozygosity-fitness correlations was found for horn growth, but not for body mass and the intensity of parasite infection. The second chapter investigates the relative importance of density dependence and of stochastic climatic variability in the population dynamics of a mountain ungulate, analysing a 45 year long time series of Alpine ibex censuses collected in the Gran Paradiso National Park, Italy During the first 28 years of the study, the total number of ibex ranged from about 2600 to about 4000 with no visually apparent trend. During this period oscillations with periodicities of about 3 and 8 years appeared to be present. From 1982 onwards, ibex counts increased steadily and peaked at almost 5000 in 1993, decreasing afterwards. We show that the ibex population size was limited by both density dependence and deep snow. A model based on these factors fit to the first 19 years of data was successful in forecasting the increase and subsequent decline in total population size over the final 20 years of the study. The thesis concludes with a third chapter presenting two innovative techniques useful in the study of free-ranging populations of ungulates

Predicting the potential distribution of the Endangered huemul deer Hippocamelus bisulcus in North Patagonia

Habitat loss is one of the main threats to wildlife, particularly large mammals. Estimating the potential distribution of threatened species to guide surveys and conservation is crucial, primarily because such species tend to exist in small fragmented populations. The Endangered huemul deer Hippocamelus bisulcus is endemic to the southern Andes of Chile and Argentina. Although the species occurs in the Valdivian Ecoregion, a hotspot for biodiversity, we have no information on its occupancy and potential distribution in this region. We built and compared species distribution models for huemul using the maximum entropy approach, using 258 presence records and sets of bioclimatic and geographical variables as predictors, with the objective of assessing the potential distribution of the species in the Valdivian Ecoregion. Annual temperature range and summer precipitation were the predictive variables with the greatest influence in the best-fitting model. Approximately 12,360 km2 of the study area was identified as suitable habitat for the huemul, of which 30% is included in the national protected area systems of Chile and Argentina. The map of potential distribution produced by our model will facilitate prioritization of future survey efforts in other remote and unexplored areas in which huemul have not been recorded since the 1980s, but where there is a high probability of their occurrence

Flight Initiation Distance and Starting Distance: Biological Effect or Mathematical Artefact?

In many studies, flight initiation distance (FID, the distance at which a prey starts to flee at the approach of a walker) is positively related to starting distance (SD, the distance at which the walker begins to approach) and alert distance (AD, the distance at which the focal individual becomes alert to the threat). In spite of the fundamental differences between SD, a covariate that may not have any biological effect, and AD, a measure related to the behaviour of the animal, it is common to use SD as a proxy for AD when AD is hard to measure (e.g. in species that do not exhibit distinguishable alert postures). However, the relationship between SD and AD or FID may not have any biological reasons, but may instead simply result from a mathematical artefact because of the constraints SD = AD = FID. Under such constrains, the homoscedasticity assumption is violated, and thus, the classical null hypothesis of linear regression (slope = 0) is invalid. In this study, we first show that using SD as a proxy for AD can strongly affect the results on FID. Using data from FID tests on alpine marmots (Marmota marmota), a linear mixed model with AD as a covariate, suggested that the interaction between previous activity and AD had an effect on FID, while this effect was not detected when SD replaced AD as the covariate in the analysis. We then propose that the actual statistical test of the relationship between SD, AD and FID should be based on a null hypothesis that incorporates the constraint SD = AD = FID = 0 and generate 95% CI of simulated slopes obtained from random values under this constraint. This null hypothesis can be rejected if the observed slope of the relationship between two of these variables is outside the 95% CI. We demonstrated that, for alpine marmots, the observed slope of the relationship between AD and SD was within the 95% CI of the simulated slopes. The absence of a statistically significant biological effect in the relationship between SD and AD raises important questions on the outcome of relationship between SD and FID. In Alpine marmot flight, decision should be studied separating the effect of SD on AD and the effect of AD on FID

Early onset of vegetation growth vs. rapid green-up : impacts on juvenile mountain ungulates

Seasonal patterns of climate and vegetation growth are expected to be altered by global warming. In alpine environments, the reproduction of birds and mammals is tightly linked to seasonality; therefore such alterations may have strong repercussions on recruitment. We used the normalized difference vegetation index (NDVI), a satellite-based measurement that correlates strongly with aboveground net primary productivity, to explore how annual variations in the timing of vegetation onset and in the rate of change in primary production during green-up affected juvenile growth and survival of bighorn sheep (Ovis canadensis), Alpine ibex (Capra ibex), and mountain goats (Oreamnos americanus) in four different populations in two continents. We indexed timing of onset of vegetation growth by the integrated NDVI (INDVI) in May. The rate of change in primary production during green-up (early May to early July) was estimated as (1) the maximal slope between any two successive bimonthly NDVI values during this period and (2) the slope in NDVI between early May and early July. The maximal slope in NDVI was negatively correlated with lamb growth and survival in both populations of bighorn sheep, growth of mountain goat kids, and survival of Alpine ibex kids, but not with survival of mountain goat kids. There was no effect of INDVI in May and of the slope in NDVI between early May and early July on juvenile growth and survival for any species. Although rapid changes in NDVI during the green-up period could translate into higher plant productivity, they may also lead to a shorter period of availability of high-quality forage over a large spatial scale, decreasing the opportunity for mountain ungulates to exploit high-quality forage. Our results suggest that attempts to forecast how warmer winters and springs will affect animal population dynamics and life histories in alpine environments should consider factors influencing the rate of changes in primary production during green-up and the timing of vegetation onset

Involvement of recreational anglers in the eradication of alien brook trout from high altitude lakes

Stocking programmes for recreational angling are primarily responsible for the spread and ecological impact of introduced sh in high-altitude, originally shless lakes. In 2013, the Gran Paradiso National Park started an eradication campaign of brook trout by intensive gill-netting. Local anglers were invited to attend two angling sessions to start the eradication before gill-netting in an experimental lake, as part of an education action devoted to these critical stakeholders. The angling sessions turned out to be a valuable help for the eradication campaign and the aim of this study is to report on the outcomes of these angling sessions. Angling techniques were highly size-selective, removing a substantial part of the adult population and of the sh biomass, but their contribution to the eradication of small sh (<15cm) was irrelevant. Therefore, angling cannot completely eradicate age-structured populations. However, there is scope to use angling sessions as a support for eradication campaigns and as an emergency measure for recent sh introduc- tions. Similar actions should be considered whenever a sh eradication programme is planned. These ndings, however, do not imply a general endorsement for angling within protected areas

Technical validation and a comparison of two methods to quantify individual levels of glucocorticoids in Alpine marmot hair

Quantification of cortisol concentration in hair has become a promising conservation tool for non-invasive monitoring of “stress” in wild populations, yet this method needs to be carefully validated for each species. The goals of the study were: • Immunologically validate two methods (study 1 and 2 respectively) to extract and quantify cortisol in the hair of wild Alpine marmots. • Compare the amount of cortisol extracted from hair samples using two methods i.e. cut into fine pieces (study 1) and hair samples pulverized using a ball mill (study 2). • Determine the extent to which methods in study 2 could provide individual specific hair cortisol (HC) measures when samples were taken from the same body location. Within and between individual variations in HC levels were examined from multiple hair samples from 14 subjects in study 2. We evaluated if inter-individual variations in HC levels could be explained by sex and age. At least twice the amount of cortisol was obtained per g/hair when samples were pulverized in a ball mill prior to extraction compared to when cut into pieces. Our methods demonstrated intra-individual consistency in HC at a given time point: inter-individual variation in HC was three times larger than within individual variance. Sex and age did not impact HC levels

Behavioural Indicators of Intra- and Inter-Specific Competition: Sheep Co-Grazing with Guanaco in the Patagonian Steppe

In extensive livestock production, high densities may inhibit regulation processes, main- taining high levels of intraspecific competition over time. During competition, individuals typically modify their behaviours, particularly feeding and bite rates, which can therefore be used as indicators of competition. Over eight consecutive seasons, we investigated if variation in herd density, food availability, and the presence of a potential competitor, the guanaco (Lama guanicoe), was related with behavioural changes in domestic sheep in Chilean Patagonia. Focal sampling, instantaneous scan sampling, measures of bite and movement rates were used to quantify behavioural changes in domestic sheep. We found that food availability increased time spent feeding, while herd density was associated with an increase in vigilant behaviour and a decrease in bite rate, but only when food availability was low. Guanaco presence appeared to have no impact on sheep behaviour. Our results suggest that the observed behavioural changes in domestic sheep are more likely due to intraspecific competition rather than interspecific competition. Consideration of intraspecific competition where guanaco and sheep co-graze on pastures could allow management strategies to focus on herd density, according to rangeland carrying capacity

Social network analysis of small social groups: Application of a hurdle GLMM approach in the Alpine marmot ( Marmota marmota )

From Wiley via Jisc Publications RouterHistory: received 2020-09-21, rev-recd 2021-03-09, accepted 2021-03-10, pub-electronic 2021-03-24, pub-print 2021-06Article version: VoRPublication status: PublishedAbstract: Social network analysis (SNA) has recently emerged as a fundamental tool to study animal behavior. While many studies have analyzed the relationship between environmental factors and behavior across large, complex animal populations, few have focused on species living in small groups due to limitations of the statistical methods currently employed. Some of the difficulties are often in comparing social structure across different sized groups and accounting for zero‐inflation generated by analyzing small social units. Here, we use a case study to highlight how Generalized Linear Mixed Models (GLMMs) and hurdle models can overcome the issues inherent to study of social network metrics of groups that are small and variable in size. We applied this approach to study aggressive behavior in the Alpine marmot (Marmota marmota) using an eight‐year long dataset of behavioral interactions across 17 small family groups (7.4 ± 3.3 individuals). We analyzed the effect of individual and group‐level factors on aggression, including predictors frequently inferred in species with larger groups, as the closely related yellow‐bellied marmot (Marmota flaviventris). Our approach included the use of hurdle GLMMs to analyze the zero‐inflated metrics that are typical of aggressive networks of small social groups. Additionally, our results confirmed previously reported effects of dominance and social status on aggression levels, thus supporting the efficacy of our approach. We found differences between males and females in terms of levels of aggression and on the roles occupied by each in agonistic networks that were not predicted in a socially monogamous species. Finally, we provide some perspectives on social network analysis as applied to small social groups to inform subsequent studies

Monitoring wildlife population trends with sample counts: a case study on the Alpine ibex (Capra ibex)

Monitoring population dynamics is of fundamental importance in conservation but assessing trends in abundance can be costly, especially in large and rough areas. Obtaining trend estimations from counts performed in only a portion of the total area (sample counts) can be a cost‐effective method to improve the monitoring and conservation of species difficult to count. We tested the effectiveness of sample counts in monitoring population trends of wild animals, using as a model population the Alpine ibex Capra ibex in the Gran Paradiso National Park (Italy), both with computer simulations and using historical count data collected over the last 65 years. Despite sample counts failed to correctly estimate the true population abundance, sampling half of the target area could reliably monitor the trend of the target population. In case of strong changes in abundance, an even lower proportion of the total area could be sufficient to identify the direction of the population trend. However, when there is a high yearly trend variability, the required number of samples increases and even counting in the entire area can be ineffective to detect population trends. The effect of other parameters, such as which portion of the area is sampled and detectability, was lower, but these should be tested case by case. Sample counts could therefore constitute a viable alternative to assess population trends, allowing for important, cost‐effective improvements in the monitoring of wild animals of conservation interest

Marginal habitats provide unexpected survival benefits to the alpine marmot

From Wiley via Jisc Publications RouterHistory: received 2021-04-15, rev-recd 2021-09-30, accepted 2021-10-27, pub-print 2022-01, pub-electronic 2022-01-06Article version: VoRPublication status: PublishedFunder: University of Turin; Id: http://dx.doi.org/10.13039/501100006692Abstract: Age‐specific survival trajectories can vary significantly among wild populations. Identifying the environmental conditions associated with such variability is of primary importance to understand the dynamics of free‐ranging populations. In this study, we investigated survival variations among alpine marmot (Marmota marmota) families living in areas with opposite environmental characteristics: the typical habitat of the species (alpine meadow) and a marginal area bordering the forest. We used data collected during an 11‐year study in the Gran Paradiso National Park (Italy) and performed a Bayesian survival trajectory analysis on marked individuals. Furthermore, we investigated, at a territorial level, the relationships among demographic parameters and habitat variables by using a path analysis approach. Contrary to our expectations, for most of the marmot's lifespan, survival rate was higher in the marginal site closer to the forest and with lower visibility than in the alpine meadow site. Path analysis indicated that the number of families living close to each other negatively affected the stability of the dominant couple, which in turn affected both juvenile survival and reproduction. Given the lower number of neighboring families which inhabited the marginal site and the potentially different predation pressure by the most effective predator in the area (Aquila chrysaetos), our results suggest that species adapted to live in open habitats may benefit from living in a marginal habitat. This study highlights the importance of habitats bordering the forest in the conservation of alpine marmots