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Alpine marmots are thought to be herbivores but to be selective regarding food choice with focus on plant material rich in proteins and fatty acids. Carnivorism has been occasionally observed in Al...

Studying natal dispersal in natural populations using capture-recapture data is challenging as an unknown proportion of individuals leaves the study area when dispersing and are never recaptured. Most dispersal (and survival) estimates from capture-recapture studies are thus biased and only reflect what happens within the study area, not the population. Here, we elaborate on recent methodological advances to build a spatially-explicit multi-state capture-recapture model to study natal dispersal in a territorial mammal while accounting for imperfect detection and movement in and out of the study area. We validate our model using a simulation study where we compare it to a non-spatial multi-state capture-recapture model. We then apply it to a long-term individual-based dataset on Alpine marmot (Marmota marmota). Our model was able to accurately estimate natal dispersal and survival probabilities, as well as mean dispersal distance for a large range of dispersal patterns. By contrast, the non-spatial multi-state estimates underestimated both survival and natal dispersal even for short dispersal distances relative to the study area size. We discuss the application of our approach to other species and monitoring set-ups. We estimated higher inheritance probabilities of female Alpine marmots, which suggests higher levels of philopatry, although the probability to become dominant after dispersal did not differ between sexes. Nonetheless, the lower survival of young adult males suggests higher costs of dispersal for males. We further discuss the implications of our findings in light of the life-history of the species.

Author(s): Viviano, Andrea; Mori, Emiliano | Abstract: The distribution of the Alpine marmot released in the Northern Apennines has been largely unstudied. In this note, we summarise the current distribution and the altitude range of the Alpine marmot in the Apennine ridge, 80 years after their first releases. We searched for marmot occurrence on the Apennines (i) on citizen-science platforms and (ii) through a webmail on Sciuridae distribution in Italy. We collected 80 marmot records validated by photos and by field investigations. We showed that Alpine marmots are present on over 70,000 ha in the Apennines, between Emilia Romagna and Tuscany. Most occurrences were recorded between 1600 and 1700 m a.s.l., in lines with other works on this species. Although the introduction of the Alpine marmot in the Apennines appears to have been successful, further molecular and ecological data are needed to assess origins and potential environmental impacts (e.g. on soil stability) of these established populations. This work may represent a description of the current status of this species, to be compared with future monitoring. In turn, updating the distribution of the Alpine marmot in the Apennines in the next years may be useful to assess potential distribution shift towards higher altitudes as a response to local climatic change.n

This is the peer reviewed version of the following article: Panaccio, M., Ferrari, C., Bassano, B., Stanley, C.R. & von Hardenberg, A. (2021). Social Network Analysis of small social groups: application of a hurdle GLMM approach in the Alpine marmot (Marmota marmota). Ethology, which has been published in final form at [Link to final article using the DOI]. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving

The importance of vectors and vector-borne diseases (VBDs) is increasing on a global scale. Many vectors and pathogens benefit from global warming and can spread to novel habitats where they were formerly not present, including higher altitudes. Various vector-borne pathogens (VBPs), such as Anaplasma phagocytophilum, have been reported in, for instance, red foxes and wild ungulates in the Western Austrian Alps. However, these animals are known to migrate to lower regions in the winter season, and therefore, it is of interest to investigate if VBPs are also present in mammals faithful to their higher altitude alpine habitat all year round. Blood parasites and other VBPs, namely. Trypanosomatidae, piroplasms, Hepatozoon spp., filarioid helminths, Anaplasmataceae, and Rickettisa spp., were thus analysed with PCR in 148 alpine marmots (Marmota marmota). None of the marmots’ blood samples was positive for these VBPs, indicating a low abundance or absence of competent vectors in the alpine region. Alpine marmots seem to be naïve for VBPs (at least in our study area). An overview of VBD agents in other marmot species is given.

We sequenced the genome of the North American groundhog, Marmota monax, also known as the woodchuck. Our sequencing strategy included a combination of short, high-quality Illumina reads plus long reads generated by both Pacific Biosciences and Oxford Nanopore instruments. Assembly of the combined data produced a genome of 2.74 Gbp in total length, with an N50 contig size of 1,094,236 bp. To annotate the genome, we mapped the genes from another M. monax genome and from the closely related Alpine marmot, Marmota marmota, onto our assembly, resulting in 20,559 annotated protein-coding genes and 28,135 transcripts. The genome assembly and annotation are available in GenBank under BioProject PRJNA587092.

Geographic variation in acoustic signals has been investigated for five decades to better understand the evolution of communication. When receivers are able to discriminate among signals and to react accordingly, geographic variation can have major impacts on the ability of conspecifics to communicate. Surprisingly, geographic variation in alarm calls and its consequences for the communication process have been so far neglected despite their crucial role on individual survival. Working with four wild populations of Alpine marmots (Marmota marmota), we found differences in the acoustic structure of their alarm calls. These differences cannot be explained by geographic or genetic distances but more likely by other mechanisms including random processes. Moreover, playback experiments provided evidence that receivers discriminate between alarm calls from their own versus other populations, with responses at lower intensity when the alarm calls played back originated from their own population. Research on the mechanistic causes of geographic variation and on the relationship between alarm call variation, familiarity, and intelligibility of signal and behavioral responses is now required to better understand how predation pressure, and more widely natural selection, could drive the evolution of communication. Dialects (i.e., geographic variation) can have major impacts on the ability of conspecifics to communicate. Surprisingly, dialects in alarm calls have been neglected despite their crucial role on survival of individuals. Alpine marmots have dialects in alarm calls and discriminate their own dialects from others, being more frightened by alarm calls from another population than by those from their own. Confronted with an unknown dialect, marmots may adopt a self-preserving strategy and choose to run away before assessing the danger.

Development of winter outdoor leisure activities in areas of high natural value is a key issue in the sustainable use of mountain environments. Ski touring, an emerging outdoor activity in the Tatra Mountains, is believed to affect protected mammal species such as the Tatra chamois (Rupicapra rupicapra tatrica), alpine marmot (Marmota marmota latirostris), red deer (Cervus elaphus), European roe deer (Capreolus capreolus), brown bear (Ursus arctos), and red fox (Vulpes vulpes). To assess the impact of ski touring on these species, various studies are underway in Tatra National Park. The aim of this study was to investigate the interactions between ski tourers and large mammals in the park using a social science approach. A total of 274 skiers completed an online questionnaire about their encounters with large mammals during their ski tours in Tatra National Park and their perceptions of animals' reactions to their presence. Just over half of the respondents reported encounters with large mammals—most often with chamois, followed by deer (red or roe), foxes, and marmots. Only 6% reported encounters with brown bears. The most commonly reported animal reactions were vigilance and indifference (no reaction). Flight or aggression occurred less often—in about 22% of encounters with all species and 12% of encounters with chamois. An online survey on human–wildlife interactions can provide a cost- and labor-efficient complement to field research such as direct observation, GPS tracking, and physiological testing.

To reproduce, animals have to form pairs and large variations in the degree of mate switching are observed. Extrinsic and intrinsic factors can constrain individual's mate switching. Among intrinsic factors, genes involved in pair-bonding, such as Avpr-1a, receive increasing attention. The length of microsatellites present in the regulatory region of Avpr-1a determines the neural densities and distributions of the vasopressin receptors known to impact pair-bonding behaviours. For the first time, we investigated whether and how the genetic makeup at Avpr-1a, an intrinsic factor, and the social context, an extrinsic factor, experienced by wild Alpine marmot (Marmota marmota) females affect the proportion of extra-pair young. This proportion was positively correlated with the length of their Avpr-1a regulatory region but only when the social constraints were relaxed, that is when mature male subordinates were present. When ignoring the interactive effect between the length of their Avpr-1a regulatory region and the social constraints, the genetic makeup at Avpr-1a was not associated with the proportion of extra-pair young. Under natural conditions, the genetic regulation of pair-bonding could be hidden by extrinsic factors constraining mate choice.

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Habitat Suitability Models (HSMs) are central tools in physical and environmental planning because they are able to predict potential distribution of species. In this study two spatial methods were compared to define the HSM for Marmota marmota L. in the Adamello Brenta Nature Park in the Italian central southern part of the Alps. The first model was based on Weighted Linear Combination (WLC), a nonparametric overlay procedure, the second one was a parametric method based on logistic regression algorithm. Data collected in two sample areas were used to build the models, while data recorded along linear transects evenly distributed throughout the study area were used for testing the model performance. The models were validated calculating the Area Under the Curve (AUC) of the Relative Operating Characteristics (ROC). The two models had similar performances, but the accuracy of the WLC model was slightly higher than the logistic model (AUC of 0.833 instead of 0.821). The habitat preferences of Marmota marmota L. were also investigated with Jacobs index and chi-square test and compared with other studies in different environments. The results demonstrated that the relative importance of environmental factors for Marmota marmota L. changes depending on local conditions.

Summary Some species responded successfully to prehistoric changes in climate [1, 2], while others failed to adapt and became extinct [3]. The factors that determine successful climate adaptation remain poorly understood. We constructed a reference genome and studied physiological adaptations in the Alpine marmot (Marmota marmota), a large ground-dwelling squirrel exquisitely adapted to the “ice-age” climate of the Pleistocene steppe [4, 5]. Since the disappearance of this habitat, the rodent persists in large numbers in the high-altitude Alpine meadow [6, 7]. Genome and metabolome showed evidence of adaptation consistent with cold climate, affecting white adipose tissue. Conversely, however, we found that the Alpine marmot has levels of genetic variation that are among the lowest for mammals, such that deleterious mutations are less effectively purged. Our data rule out typical explanations for low diversity, such as high levels of consanguineous mating, or a very recent bottleneck. Instead, ancient demographic reconstruction revealed that genetic diversity was lost during the climate shifts of the Pleistocene and has not recovered, despite the current high population size. We attribute this slow recovery to the marmot’s adaptive life history. The case of the Alpine marmot reveals a complicated relationship between climatic changes, genetic diversity, and conservation status. It shows that species of extremely low genetic diversity can be very successful and persist over thousands of years, but also that climate-adapted life history can trap a species in a persistent state of low genetic diversity., Highlights • The Alpine marmot is among the least genomically diverse animal species • Its diversity was lost during consecutive ice-age climate-related events • An extreme lifestyle hampered the subsequent recovery of genetic variation • Alpine marmots show why large populations can coexist with very low genetic variation, Despite being highly abundant and well adapted, Gossmann et al. report that the Alpine marmot is among the least genetically diverse animal species. The low diversity is found to be the consequence of consecutive, climate-related events, including long-term extreme niche adaptation, that also greatly retarded the recovery of its genetic diversity.

In the context of global change, an increasing challenge is to understand the interaction between weather variables and life histories. Species-specific life histories should condition the way climate influences population dynamics, particularly those that are associated with environmental constraints, such as lifestyles like hibernation and sociality. However, the influence of lifestyle in the response of organisms to climate change remains poorly understood. Based on a 23-year longitudinal study on Alpine marmots, we investigated how their lifestyle, characterized by a long hibernation and a high degree of sociality, interacts with the ongoing climate change to shape temporal variation in age-specific survival. As generally reported in other hibernating species, we expected survival of Alpine marmots to be affected by the continuous lengthening of the growing season of plants more than by changes in winter conditions. We found, however, that Alpine marmots displayed lower juvenile survival over time. Colder winters associated with a thinner snow layer lowered juvenile survival, which in turn was associated with a decrease in the relative number of helpers in groups the following years, and therefore lowered the chances of over-winter survival of juveniles born in the most recent years. Our results provide evidence that constraints on life-history traits associated with hibernation and sociality caused juvenile survival to decrease over time, which might prevent Alpine marmots coping successfully with climate change.

Maintaining dominance status had long been considered to be less stressful than subordination. However, no consistency in stress levels of dominant and subordinate individuals has been demonstrated. Tactics used to achieve and maintain dominance could be determinant. In cooperatively breeding species, conflicts between dominants and subordinates are expected since dominant individuals tend to monopolize reproduction while subordinates seldom reproduce. Reproductive skew models predict that subordinates’ reproductive opportunities are either allotted or subject to competition with dominants. In the former case, no policing of subordinates by dominants is expected. In the latter, dominant should exert a control over the subordinates possibly leading to higher stress levels in dominants than in subordinates, which could be further elevated as the number of potential competitors in the group increases. In the present study, we aimed to test these hypotheses by assessing individual’s stress level using the neutrophils to lymphocytes ratio (N:L) in a wild cooperatively breeding rodent, the Alpine marmot (Marmota marmota). We found that dominants exhibit higher N:L ratio than subordinates and that dominants’ N:L ratio increases with the number of unrelated same-sex subordinates in the group. We conclude that controlling unrelated subordinates is stressful for dominants, as expected under tug-of-war models. These stress patterns reveal conflicting relationships between dominants and subordinates over the reproduction and social status acquisition. This study highlights the influence of the nature, strength, and direction of conflicts on stress levels. In cooperatively breeding species, reproductive skew models predict that subordinates’ reproductive opportunities are either allotted or subject to competition with dominants and, thus, can modulate the relative stress level between dominants and subordinates. In the first case, no policing of subordinates by dominants is expected, while in the second, dominant should exert a control over the subordinates which should lead to higher stress level in dominants than in subordinates, and particularly when subordinates are unrelated to the dominants. In Alpine marmots, we found that dominants exhibit higher stress level than subordinates and dominants’ stress level increased with the number of unrelated same-sex subordinates. These patterns are in agreement with the predictions of the tug-of war models of reproductive skew and indicate that controlling subordinates is costly for dominants.

The Alpine Marmot (Marmota marmota) is a rodent remnant of the ‘ice-age’ climate of the Pleistocene steppe, and since the disappearance of this habitat, persists in the high altitude Alpine meadow. Sequencing its genome, we reveal that the long-term cold-climate adaptation has altered its metabolism, in particular biosynthesis and storage of fatty acids. Paradoxically, despite successful adaptation and a large population size, we detected levels of genetic variation that are among the lowest for mammals, and that purifying selection is ineffective. Reconstructing its demographic past revealed that the two apparently contradictory observations are, in fact, intimately interlinked. The climate-adaptive life history prevented genetic diversity to recover from Pleistocene perturbations. The case of the Alpine marmot demonstrates that populations of extremely low genetic diversity can be very successful and persist over thousands of years, but that a climate-adapted life history can deprive a species from the possibility to recover genetic diversity.

The function of Alpine marmot subspecies burrows and factors influencing their distribution were investigated at 17 sites in the Tatra Mountains in Slovakia, Europe. Topographic variables and habitat types expected to influence the location of the Tatra marmot burrows were observed and examined. In the various locations of the studied home ranges (summer, winter and grazing areas, marmot trails) and outside the family territories, we recorded 36 vegetation samples including several reliefs. The collected data were analyzed using multivariate analysis. The summer areas of the marmot home ranges in the Tatra Mountains are often located within the alpine zone in communities of the Braun-Blanquet alliance Juncion trifidi (siliceous short stem grasslands) and in chionophilous communities on stable scree slopes of the alliance Festucion picturatae (tall stem grasslands). Marmots usually avoid habitats that have the lowest trophic benefits and the most extreme sites with low-stems or mat-grass communiti...

In social mammals, territory size and shape vary according to the number and strength of neighbour individuals competing for resources. Two main theories have been proposed to explain this variability: the Group Augmentation (GA) and the realized Resource Holding Potential (rRHP) hypotheses. The first states that the outcome of the interactions among groups depends on the total number of individuals in the group while the second states that only the number of animals directly involved in intergroup competition determines this outcome. We collected data on space use of individually tagged Alpine marmots (Marmota marmota), a cooperative breeding species that overlaps part of its territory with neighbouring groups. In accordance with the rRHP hypothesis, we found that groups having higher proportion of helpers, rather than higher total number of individuals, had lower percentage of the territory overlapping with neighbouring groups and a larger area available for individual exclusive use.

Alpine marmots Marmota marmota occupy a narrow altitudinal niche within high elevation alpine environments. For animals living at such high elevations where resources are limited, parasitism represents a potential major cost in life history. Using occupancy models, we tested if marmots living at higher elevation have a reduced risk of being infected with gastro-intestinal helminths, possibly compensating the lower availability of resources (shorter feeding season, longer snow cover and lower temperature) than marmots inhabiting lower elevations. Detection probability of eggs and oncospheres of two gastro-intestinal helminthic parasites, Ascaris laevis and Ctenotaenia marmotae, sampled in marmot feces, was used as a proxy of parasite abundance. As predicted, the models showed a negative relationship between elevation and parasite detectability (i.e. abundance) for both species, while there appeared to be a negative effect of solar radiance only for C. marmotae. Site-occupancy models are used here for the first time to model the constrains of gastrointestinal parasitism on a wild species and the relationship existing between endoparasites and environmental factors in a population of free-living animals. The results of this study suggest the future use of site-occupancy models as a viable tool to account for parasite imperfect detection in eco-parasitological studies, and give useful insights to further investigate the hypothesis of the contribution of parasite infection in constraining the altitudinal niche of Alpine marmots.

Coping styles represent alternative response patterns in reaction to a stressor. The coping style model provides a set of predictions about correlations between behavioural and neurophysiological reactions to a stressful situation. According to this model, high levels of activity should be correlated with high levels of aggressiveness at the behavioural level, and to high sympathetic reactivity, low parasympathetic reactivity (higher heart rate levels) and low hypothalamus–pituitary–adrenocortical (HPA) axis reactivity (low production of glucocorticoids in response to a stressor). More recent versions of the model, however, are challenging this view and consider the possibility of independent axes of coping style and stress reactivity. The coping style model has mainly been tested on artificially selected or inbred lines in laboratory settings. Such a situation restricts its generalization to a larger number of species and there is a need for studies testing it in the wild under more natural situations. Here, we test the predictions of the coping style model in a wild alpine marmot, Marmota marmota, population. We show that several behavioural (i.e. exploration in an open field, impulsivity and docility) and neurophysiological traits (i.e. heart rate, breathing rate and cortisol production) assumed to represent individual differences in coping style were significantly repeatable over 2–3 years. Not all the correlations between traits predicted by the coping style model were found in marmots, which supports the more recent two-axes model. Furthermore, most correlations were observed at the between-individual level, and the within-individual correlations (i.e. phenotypic plasticity) were weaker. Overall, our results support the prediction of the coping style model, but highlight the fact that the association between traits found in artificial conditions may be weaker in a more natural setting.

Reintroductions inherently involve a small number of founders leading reintroduced populations to be prone to genetic drift and, consequently, to inbreeding depression. Assessing the origins as the genetic diversity and structure of reintroduced populations compared to native populations are thus crucial to foresee their future. Here, we aim to clarify the origins of the Alpine marmots reintroduced in the Pyrenees and to evaluate the genetic consequences of this reintroduction after almost 30 years without monitoring. We search for the origins and compare the genetic structure and the genetic variability of three reintroduced Pyrenean and eight native Alpine populations using pairwise genetic distances, Bayesian clustering method and multivariate analyses. Our results reveal that the Alpine marmots reintroduced in the Pyrenees originated both from the Northern and the Southern Alps, and that, despite these multiple origins, none of the current Pyrenean marmots are admixed. The reintroduction led to a strong genetic differentiation and to a decrease in genetic diversity. This pattern likely results from the small number of founders and the low dispersal capacities of Alpine marmots and thus, highlight the necessity to consider both genetic characteristics and natural history when reintroducing a species.

We investigated age-specific variation in survival of dominant individuals in a long-lived and monogamous mammal, the alpine marmot ( Marmota marmota ), from a large dataset collected during a 24-year intensive monitoring of a free-ranging population. We found evidence of actuarial senescence in dominant individuals for both sexes. Survivorship was constant with age until dominant marmots were between 6 and 8 years of age and declined markedly thereafter. No between sex differences occurred in the intensity of actuarial senescence, which might be related to the weak intensity of sexual selection in this socially monogamous mammal. More investigations are needed to know whether cooperative breeding, hibernation, and monogamy, which are key features of the alpine marmot life history, could have shaped the patterns of actuarial senescence we report.

Sociality should evolve when the fitness benefits of group living outweigh the costs. Theoretical models predict an optimal group size maximizing individual fitness. However, beyond the number of individuals present in a group, the characteristics of these individuals, like their sex, are likely to affect the fitness payoffs of group living. Using 20 years of individually based data on a social mammal, the Alpine marmot (Marmota marmota), we tested for the occurrence of an optimal group size and composition, and for sex-specific effects of group characteristics on fitness. Based on lifetime data of 52 males and 39 females, our findings support the existence of an optimal group size maximizing male fitness and an optimal group composition maximizing fitness of males and females. Additionally, although group characteristics (i.e., size, composition and instability) affecting male and female fitness differed, fitness depended strongly on the number of same-sex subordinates within the social group in the two sexes. By comparing multiple measures of social group characteristics and of fitness in both sexes, we highlighted the sex-specific determinants of fitness in the two sexes and revealed the crucial role of intrasexual competition in shaping social group composition.

We warmly thank all students and Earthwatch volunteers involved in catching the marmots and the authorities of the Vanoise National Park for granting us permission to work in the Grande Sassière Nature Reserve. We thank M. Harrington for English editing and Dr. E. Rajon for helpful discussions on the manuscript. This work was supported by the "Agence Nationale de la Recherche" (ANR, project ANR-08-BLAN-0214-03, ANR-13-JSV7-0005), the "Centre National de la Recherche Scientifique" (CNRS), the "FR41 BioEnvironnement et Santé de l'Université de Lyon" and the "Earthwatch Institute", the scholarship for postgraduate studies "Obra Social Fundació La Caixa" and Vetagro-Sup. Sexual selection through female mate choice for genetic characteristics has been suggested to be an important evolutionary force maintaining genetic variation in animal populations. However, the genetic targets of female mate choice are not clearly identified and whether female mate choice is based on neutral genetic characteristics or on particular functional loci remains an open question. Here, we investigated the genetic targets of female mate choice in Alpine marmots (Marmota marmota), a socially monogamous mammal where extra-pair paternity (EPP) occurs. We used 16 microsatellites to describe neutral genetic characteristics and two MHC loci belonging to MHC class I and II as functional genetic characteristics. Our results reveal that (1) neutral and MHC genetic characteristics convey different information in this species, (2) social pairs show a higher MHC class II dissimilarity than expected under random mate choice, and (3) the occurrence of EPP increases when social pairs present a high neutral genetic similarity or dissimilarity but also when they present low MHC class II dissimilarity. Thus, female mate choice is based on both neutral and MHC genetic characteristics, and the genetic characteristics targeted seem to be context dependent (i.e., the genes involved in social mate choice and genetic mate choice differ). We emphasize the need for empirical studies of mate choice in the wild using both neutral and MHC genetic characteristics because whether neutral and functional genetic characteristics convey similar information is not universal.

Ranging behaviour is one of the most important aspects of the life history of many species. Spatial distributions of individuals in the wild is linked to factors such as foraging, mating, population density, availability of resources and competition. Accurate data on the spatial location of individuals over time is often difficult to collect. Here, we propose a new simple, non-invasive and economic method for collecting accurate spatial data usable for many different species of free-ranging animals. Our instrument for collecting animal locations consists of three elements: a laser range finder, a laser tilt sensor and a protractor. This instrument can obtain three-dimensional parameters of the space from a fixed point allowing the user to collect geographical locations of the animals and, in general, of any point of interest. The device we tested showed a very low average error among (1.76 6 0.643 m) and within (1.79 6 0.058 m) observers, and the locations we obtained were all within the 95% probability of the tolerance intervals for the 20 positions which we measured repeatedly with a Global Position System for each of 10 different test locations. We tested a range of different distances to the target points (from 20 to 222 m), and we propose formulas to calculate precision of the instrument inside this interval. Precision of estimated locations was between 0.32 to 3.55 m from the real location and it was slightly related to distance of the target point (r ¼ 0.38, P ¼ 0.054). As an example of its practical application, we present data on the use of the instrument within the framework of a study on a population of free-ranging individually tagged alpine marmots Marmota marmota.

The major histocompatibility complex (MHC) genes code for proteins that play a critical role in the immune system response. The MHC genes are among the most polymorphic genes in vertebrates, presumably due to balancing selection. The two MHC classes appear to differ in the rate of evolution, but the reasons for this variation are not well understood. Here, we investigate the level of polymorphism and the evolution of sequences that code for the peptide-binding regions of MHC class I and class II DRB genes in the Alpine marmot (Marmota marmota). We found evidence for four expressed MHC class I loci and two expressed MHC class II loci. MHC genes in marmots were characterized by low polymorphism, as one to eight alleles per putative locus were detected in 38 individuals from three French Alps populations. The generally limited degree of polymorphism, which was more pronounced in class I genes, is likely due to bottleneck the populations undergone. Additionally, gene duplication within each class might have compensated for the loss of polymorphism at particular loci. The two gene classes showed different patterns of evolution. The most polymorphic of the putative loci, Mama-DRB1, showed clear evidence of historical positive selection for amino acid replacements. However, no signal of positive selection was evident in the MHC class I genes. These contrasting patterns of sequence evolution may reflect differences in selection pressures acting on class I and class II genes.

The introduction of non-native species can pose environmental and economic risks, but under some conditions, introductions can serve conservation or recreational objectives. To minimize risks, introductions should be conducted following the International Union for Conservation of Nature's guidelines and should include an initial assessment and a follow-up. In 1948, to reduce the predation pressure on Pyrenean chamois Rupicapra pyrenaica pyrenaica by golden eagles Aquila chrysaetos, the alpine marmot Marmota marmota was introduced to the Pyrenees in Western Europe. In successive introductions, about 500 marmots were released, but the fate of the released animals and their impacts on the environment remain largely unstudied. The aim of this study was to assess the success of the introduction of the alpine marmot into the Pyrenees, 60 years after the initial release, and the potential impacts of this species on Pyrenean ecosystems. We reviewed what is known about the marmot populations introduced to the Pyrenees and other populations within their native range in the Alps, particularly in terms of population structure and dynamics, habitat use and potential environmental impacts. The alpine marmot is widely distributed and, apparently, well established in the Pyrenees. Population structure and demographic parameters are similar within and outside the historical distribution range of the species, and habitat suitability is one of the main reasons for the species' success in the Pyrenees. Few researchers have investigated the impacts of alpine marmots in the Pyrenees; thus, those impacts have to be inferred from those observed in the species' native range or in other species of marmot. Introduced alpine marmots are likely to impact on Pyrenean grasslands through grazing and burrowing, have the potential to alter Pyrenean food webs and could act as vectors of parasites and disease. Although the introduction of the alpine marmot in the Pyrenees appears to have been successful, more needs to be known about the effects of the established populations on the environment before informed management actions can be taken in the Pyrenees.

Archaeozoological investigations carried out at Grotta del Clusantin in the Italian north-eastern Alps suggest that this late Upper Palaeolithic site hosted small groups of marmot hunters. Taxonomic data describe an alpine grassland environment with arboreal and shrubby vegetation in proximity. The bone assemblage is extremely fragmented as a result of anthropic activities and post-depositional agents. The presence of percussion cones, butchering cutmarks, burnt bones, and the almost total absence of carnivore traces and animal lairs, suggests that humans are responsible for faunal accumulation. Experimental approaches on marmot specimens supported archaeological evidence, demonstrating elements of systematics and/or explaining anatomical absences. From field-evidence, lithic, and functional data, it has been inferred that this site was a specialized camp, used for exploiting and consuming several rodent carcasses, in the context of the first human occupations of the highlands at the onset of the LateGlacial interstadial.

SUMMARY Variation in how individuals cope behaviourally and physiologically with stressors is widespread and can have a significant impact on life-history traits and fitness. Individual coping styles are characterised by differential behavioural and adrenocortical reactivity to various challenges. As stress hormones can affect the production of reactive chemical species and the antioxidant status, individuals with different coping styles may differ also in oxidative status. Field studies on wild mammalian populations are few in number and none so far has simultaneously tested the relationship between coping style, adrenocortical reactivity and oxidative status in the same individuals. We measured individual variation in coping styles along a proactive-reactive continuum together with variation in baseline and stress-induced plasma oxidative damage, plasma non-enzymatic antioxidant capacity and cortisol in wild alpine marmots, Marmota marmota. Confirmatory path analysis revealed that different coping styles are accompanied by different baseline and stress-induced plasma oxidative statuses. Our findings also highlight the potential role of cortisol as a mediator of such differences.

In litter‐bearing mammals, adult phenotype is influenced during prenatal life by the sex of neighbouring foetuses. This phenomenon, found so far only in laboratory studies, may have ecological importance in nature. We present the first evidence that litter sex ratio has consequences for lifetime reproductive success in mammals. Female Alpine marmots born in a male‐biased litter, i.e. more likely to be located in utero next to male foetuses, are more likely to be dominant later in life – a prerequisite for reproduction in this species. We found no evidence for lower reproductive output of these females, a common cost of prenatal masculinization.

The behavioural ecology of the Alpine marmot Marmota marmota (Linne, 1758) was studied Jun.—Sep. 1990 in the Vanoise National Park (French Alps). We describe the socio-spatial structure of a high-altitude population, to give additional information on the formerly unclear marmot social organization. The social unit was the family group, with a common home range between 0.9 and 2.8 ha; a slight overlapping occurred (9 to 12.5%). Each home range had a central area of main burrows, where the hibernaculum was located, and peripheral areas. Space utilization and distribution of activities during the season were analysed. Some activities took place exclusively in the centre area while peripheral parts were used for foraging. This latter increased in Aug. and decreased in Sep., marmots tending to remain more and more at the centre area as the hibernation period approached.

We studied the degree to which alpine marmot (Marmota marmota L.) alarm calls function as communication about specific external stimuli. Alpine marmots emit variable alarm calls when they encounter humans, dogs, and several species of aerial predators. The first part of the study involved observations and manipulations designed to document contextual variation in alarm calls. Alarm calls varied along several acoustic parameters, but only along one that we examined, the number of notes per call, was significantly correlated with the type of external stimulus. Marmots were more likely to emit single-note alarm calls as their first or only call in response to an aerial stimulus, and multiple-note alarm calls when first calling to a terrestrial stimulus. This relationship was not without exceptions; there was considerable variation in the number of notes they emitted to both aerial and terrestrial stimuli, and a single stimulus type — humans — elicited a wide range of acoustic responses. The second part of the study involved playing back three types of alarm calls to marmots and observing their responses. Marmots did not have overtly different responses to the three types of played-back alarm calls. Our results are consistent with the hypotheses that: 1. Alarm calls do not refer to specific external stimuli; 2. Alarm calls function to communicate the degree of risk a caller experiences; and 3. Alarm calls require additional contextual cues to be properly interpreted by conspecifics.

Scent-deposition was investigated in Marmota marmota L. at Gran Paradiso National Park. 448 h of focal and scan sampling were performed on 22 marmots in four subsequent years. Results show that: (1) 97.4 % of the scent-deposition is done within 25 m of the edge of the territories or of the main burrows (X2 = 100.4, df 9, p < 0.001); (2) being a reproducing adult is the most prominent factor in scent-deposition (X2 = 32.4, df = 18, p < 0.002). The role of factors affecting scent-deposition in territorial defence and social communication is discussed.

Hibernators display severe changes in brain structure during deep torpor, including alterations in synaptic constitution. To address a possible effect on long-term memory, we examined learning behavior and memory of the hibernator Marmota marmota. In two operant conditioning tasks, the marmots learned to jump on two boxes or to walk through a tube. The animals were trained during their active season. Performance improved during the training phase and remained stable in a last test, four weeks before entrance into hibernation. When retested after six months of hibernation, skills were found to be unimpaired (box: before hibernation: 258.2+/-17.7 s, after hibernation: 275.0+/-19.8 s; tube: before hibernation: 158.4+/-9.0 s, after hibernation: 137.7+/-6.3 s). Contrary to these findings, memory seemed to be less fixed during the active season, since changes in test procedure resulted in impaired test performance. Besides the operant conditioning, we investigated the animals' habituation to a novel environment by repeated open field exposure. In the first run, animals showed exploratory behavior and thus a high locomotor activity was observed (63.6+/-10.7 crossed squares). Upon a second exposure, all animals immediately moved into one corner and locomotion ceased (7.2+/-1.9 crossed squares). This habituation was not altered even after hibernation (6.1+/-1.1 crossed squares). We thus conclude that long-term memory is unaffected by hibernation in Alpine marmots.

The project of marmot reintroduction in the Dolomiti Bellunesi National Park started in 2006. In May 2006 and May 2007, 80 ear-marked (by sex and family) marmots were released in two separate areas (20 ind./ year/area). In summer 2008 the populations produced 36 and 39 individuals, respectively; the animals born in the areas exceeded the number of released animals that survived. Only one pair bred successfully in the year of release; 87.5% of the surviving pairs bred the following year, including the pair breeding the previous year. We recorded the reproduction of a single female, and the acceptance of the litter by the male who later joined her. Mortality of the animals released in 2006 was 39% before the first winter and increased to 53.6% after the second summer. The first summer mortality of animals released in 2007 was higher, probably because of greater predation by golden eagles. The population density increased in late summer 2008 to 13.5 and 22.2 family units/100 ha, respectively in the two areas...

Rock ptarmigan is a vulnerable species in the southern Pyrenees, with less than 300 pairs and two unconnected populations. Alpine marmot was introduced in the northern Pyrenees between 1955 and 1988, but they rapidly colonised the southern slopes, with an actual estimated population of around 10,000 individuals. Both species are mainly herbivores, develop their activities on the ground and have their offspring at the same time, so they may compete at various levels. We studied the possible interactions between the two species based on field observations, bibliographic data and Geographical Information System (GIS) analysis. We found that alpine marmot has colonised all habitats occupied by rock ptarmigan in the southern Pyrenees, but their plant diet is absolutely differentiated. We also analysed the composition of golden eagle nests and confirmed that this predator predates the two species. From field observations in areas where the two species have coexisted for more than 10 years, we observed no behavi...

A small population of Alpine marmots was introduced near the top of the Mezenc Massif (France) in 1980 and reinforced 7 times until 1991. An annual monitoring was performed since 1988. It took 12 years to the population size to reach the number of marmots released. Afterwards the population doubled every 7 years, despite several population crashes (in 1997 and 2001). The currently censused 492 marmots were distributed into 86 sites on an area of about 200 km2 in 2007. A Multiple Correspondence Analysis of demographic and environmental parameters of each site showed that marmots preferred, these last years, anthropogenic sites (meadows, dry stone walls, elevation under 1200 m) where reproduction rate was highest. Thus, marmots are now competing with farmers for space. In the same time, tourism increased strongly, especially tourism related to marmot spotting. The interactions between the presence of marmots in this agricultural environment and human activities are described, and their consequences are cons...

Predation risk can strongly affect behaviour of preys, and natural selection should have favoured the evolution of behavioural plasticity in the strategies used to avoid predation. The level of predation risk can be affected by environmental characteristics and we expect individuals to adjust vigilance behaviour to the level of risk. In this study we investigate if Alpine marmots (Marmota marmota) living in an area delimited by the forest and with scarce visibility (Closed Site) differ in their vigilance behaviour from marmots inhabiting an open meadow further away from the forest edge and with greater visibility (Open Site). The rate of vigilance and the time spent vigilant in the Closed Site were higher than in the Open Site, while the mean duration of each vigilance bout did not differ between the two sites. Our results suggest that Alpine marmots adjust vigilance behaviour according to the environmental characteristics of their surroundings while there appears to exist an optimal duration of scan bout...

Alpine marmots were introduced in the French Pyrenees between 1948 and 1988. The exact number of re-introduced individuals is unknown, but it oscillated around 400. The likely preference of marmots for the southern sunny slopes rapidly facilitated their expansion to the southern Pyrenees, where the lack of both natural predators and of important interspecific competitors also likely facilitated an important expansion of this species. There is only one attempt to broadly calculate the population of marmots in the southern Pyrennes, estimating a population of around 10,000 individuals. However, there exist no reliable data to calculate the potential distribution of this new colonizing species in the southern Pyrenees, and a map of the potential distribution of the species is necessary to see whether alpine marmots can potentially establish in sites where it might be necessary to manage its populations for various reasons. We developed a map of potential distribution based on census carried out in summer of ...

As revealed by allozyme studies, the genetic variation of the Alpine marmot (Marmota m. marmota) has been reduced by a species-wide bottleneck at the end of the last glaciation. Therefore the more variable microsatellite loci were used as a genetic marker system to investigate variablility and differentiation of four autochthonous and four allochthonous populations founded by the release of small numbers of individuals during the last 150 years. The microsatellite loci detected by the DNA-probe (ATCC)4 were found to be polymorphic in all populations, but the amount of variation was lower than in comparable mammalian species. In spite of founder effects the variation in the allochthonous populations was not significanlty reduced compared to the autochthonous populations. The autochthonous populations from Austria and from the eastern part of Switzerland were genetically similar, only the population from western Switzerland was clearly differentiated from the others. In the allochthonous populations similarities in the microsatellite patterns reveal genetic affinities to putative autochthonous source populations of the founder individuals.

We measured body temperatures in two large hibernating mammals, the eutherian alpine marmot (Marmota marmota) and the egg-laying echidna (Tachyglossus aculeatus) from unrestrained animals in their natural environment. In both species hibernation is broken every 13 days on average by rewarming to euthermic temperatures. We found that the time course of a rewarming could be closely fitted with a sigmoid curve, allowing calculation of peak rewarming rate and corresponding body temperature. Maximum rewarming rates were twice as high in marmots as in echidnas (12.1±1.3 °C h−1, n=10 cf. 6.2±1.2 °C h−1, n=10). Peak rewarming rates were positively correlated with body temperature in echidnas, but negatively correlated in marmots.

Microsatellite variation was studied in 11 populations of the Alpine marmot along a west–east transect through the present distribution range. The samples represent five autochthonous and six introduced populations. Eleven loci were analysed in nine populations and six loci in the two populations from France. In the populations from the Western Alps, there is no indication for reduced variability as has been assumed in previous studies. However, a decrease of variation in the autochthonous populations was observed from the west to the east. The introduced populations showed a heterogeneous pattern reflecting the geographic origin of the released individuals. The population from the Spanish Pyrenees harbours a high level of variation and is genetically closest to the French populations. In Austria, three of the introduced populations have low variation and are closely related to the autochthonous populations from the western part of Austria. In contrast, two introduced populations from the central part of Austria are highly variable and resemble the populations from France. At least for one of these populations an early introduction of founder individuals from the Western Alps has been documented. Zusammenfassung Die Mikrosatellitenvariation wurde in 11 Populationen des Alpenmurmeltieres entlang eines west-ost Gradienten durch das gegenwartige Verbreitungsgebiet untersucht. Die Stichproben reprasentieren funf autochthone und sechs eingeburgerte Populationen. Elf Loci wurden in neun Populationen analysiert und sechs Loci in den beiden Populationen aus Frankreich. In den Populationen aus den Westalpen ergab sich im Gegensatz zu fruheren Untersuchungen kein Hinweis auf eine Reduktion der Variabilitat. Innerhalb der autochthonen Populationen ist jedoch eine Abnahme der Variation von Westen nach Osten festzustellen. Die eingeburgerten Populationen zeigten ein uneinheitliches Muster, welches den geografischen Ursprung der ausgesetzten Individuen widerspiegelt. Die Population aus den spanischen Pyrenaen besitzt eine hohe Variation und ist genetisch den franzosischen Populationen am ahnlichsten. In Osterreich zeigen drei der eingeburgerten Populationen niedrige Variation und sind den autochthonen Populationen aus Westosterreich am nachsten verwandt. Hingegen sind die beiden eingeburgerten Populationen aus Zentralosterreich hoch variabel und den Populationen aus Frankreich ahnlich. Zumindest fur eine dieser beiden Populationen ist eine fruhe Einburgerung von Grunderindividuen aus den Westalpen dokumentiert.

In social species, the hierarchical status of an individual has important consequences for its fitness. While many studies have focused on individual condition to explain access to dominance, very few have investigated the influence of the social environment, especially during early life. Yet it is known that environmental conditions early in life may influence several traits at adulthood. Here, we examine the influence of early social environment on accession to dominance by investigating the influence of litter size and sex composition on survival and the probability of ascending to dominance later in life using a 20-year dataset from a wild population of Alpine marmots (Marmota marmota). Although litter size had no effect on the fate of individuals, litter sex composition affected male juvenile survival and both male and female probabilities of reaching dominant status when adult. Male juveniles incur lower survival when the number of male juveniles in the litter increases, and individuals of both sexes from male-biased litters are more likely to become dominant than individuals from female-biased litters. However, the absolute number of sisters in the litter, rather than the sex ratio, seems to be an important predictor of the probability of acquiring dominant status: pups having more sisters are less likely to become dominant. Several potential mechanisms to explain these results are discussed.

Extra-pair paternity (EPP) can be influenced by both social setting and female mate choice. If evidence suggests that females try to obtain extra-pair copulations (EPCs) in order to gain genetic benefits when mated to a homozygous and/or to a related male, females may not be able to choose freely among extra-pair mates (EPMs) as the social mate may constrain female access to EPMs. In this study, we investigated, first, how EPP depended on social setting and specifically on the number of subordinate males in the family group in a highly social and monogamous mammal, the alpine marmot. Second, we investigated how EPP depended on female mate choice for genetic benefits measured as male mate-heterozygosity and within-pair relatedness. Our results reveal, first, that EPP depended on the social setting, increasing with the number of subordinate males. Second, EPPs were related to relatedness between mates. Third, EPMs were found to be more heterozygous than within-pair males. Thus, social setting may constrain female choice by limiting opportunities for EPC. However, after accounting for social confounding factors, female choice for genetic benefits may be a mechanism driving EPP in monogamous species.

Two hundred and six samples of alpine marmot (Mamota marmota) hair (148 from adults and 58 from young subjects), 102 soil samples from the entrances to the burrows of the above individuals and 20 control specimens (obtained from adjoining areas away from the burrow systems where the rodents are not usually present) were examined from May 1994 to September 1997. Seventy-five isolates belonging to six species of dermatophytes were found in 69 of the 206 hair samples examined (33.5%). Two species were zoophilic, Microsporum canis (7.8%) and Trichophyton mentagrophytes (11.2.%), and four geophilic, Microsporum cookei (2%), M. gypseum (5.8%), Trichophyton ajelloi (3.9%) and T. terrestre (5.8%). The prevalence of each species in the hair samples did not change significantly according to year, season (chi-squared test [limit significance: P 0.05] both in year and in season comparison) or age/sex (adult versus juvenile: P = 0.1; male versus female: P=0.8) of the marmot. Twen...

The consequence of helping behavior on breeders fitness is still controversial. We used multivariate analyses to investigate for the effects of male and female subordinates on breeders’ components of fitness in the Alpine marmot, Marmota marmota. We found that male and female subordinates, respectively, increased and decreased juvenile survival during winter. Thus, we give evidence that male subordinates should be considered as helpers, and that helpers provided breeders with immediate reproductive success gains, whereas subordinates females were costly. Helpers had no positive effects on female body condition, on persistence (future survival) of dominants, and on future reproduction (occurrence and size of a litter). Helpers thus did not provide breeders with delayed fitness benefits, and therfore, the load-lightening hypothesis was not supported. On the contrary, helpers had delayed fitness cost for dominant males and, consequently, for dominant females. Immediate benefits counterbalanced by delayed costs suggested an optimal number of helpers in the family group both from male and female perspectives. An optimality model well predicted the observed mean number of helpers in Alpine marmot family groups. Optimal numbers of helpers were slightly different for males and females, suggesting a potential conflict of interest between dominants. We finally discuss the possible mechanisms of helping that may explain the observed pattern in the Alpine marmot. Key words: cooperative breeding, loadlightening hypothesis, Marmota marmota, mixed models, optimality. [Behav Ecol 15:916–924 (2004)]

Circannual changes in serum parameters and body mass were studied in free-ranging Marmota marmota that had been shot in Switzerland (Grisons) for management reasons in May, July, and September of 1995, 1996, and 1997; and in April 1996. Markers of lipid (triglycerides, cholesterol), protein (total protein, urea-nitrogen) and mineral (calcium, inorganic phosphate) metabolism were evaluated in 111 haemolysis-free serum samples; the effects of sex, age, reproductive status, season and year were tested. Mean body mass was higher in adult males than in adult females in July and September, and serum cholesterol concentrations were lower in adult males in May. Pregnant females had lower concentrations of total protein than non-pregnant females, and triglyceride concentrations were negatively correlated with the number of uterine ampullae. Inorganic phosphate decreased and total protein increased with age. In adults, triglycerides, cholesterol and urea-nitrogen increased mainly from May to July and decreased between September and May. Total protein, calcium and phosphate did not change throughout the year. This suggests that, during the active season, ingested lipids and amino acids were utilised for metabolic needs or lipid storage, while in winter, lipids were catabolised and protein was spared. Quantitative differences between years, observed for triglycerides and total protein during the active season, were probably due to different climatic conditions in each year.

In cooperatively breeding vertebrate species, a clear theoretical prediction about the direction of sex ratio adjustment can be made: mothers should bias the sex ratio of their offspring towards the helping sex when helpers are absent. A consistent trend in the direction predicted by theory exists in cooperative birds, but theory is still poorly tested in cooperative mammals. Here, multivariate analyses are applied to a long-term data set to test this prediction in two ways in the alpine marmot: (1) across females in a population and (2) in individual females across multiple years. It was shown that in the alpine marmot offspring sex ratio was biased towards the helping sex (males) when helpers were absent, whereas helped mothers produced unbiased sex ratio. Unhelped mothers did not adjust the litter size but produced more sons and fewer daughters than helped mothers. These results support the theoretical prediction and explain well the male bias observed among juvenile alpine marmots at the population level. The occurrence of possible sex ratio manipulations in cooperatively breeding vertebrates is also discussed. Copyright 2004.

An alpine marmot's home range is a structured portion of soil, vegetation and rocky outcrops. The dynamics of home range and resource use were investigated in Marmota marmota L. at Gran Paradiso National Park, south-west Alps, Italy. Data were collected during 8 years from 73 individually known marmots, occupying three adjacent locations, in 264 hr of scanning and 551 hr of focal animal sampling. Three different locations did not differ in the total dimension of the home range but they differed in the total dimension of pasture. The home range inherited by immigrant monogamous couples supplanting the unrelated previous groups did not contract, and the dimension were not dictated by immediate feeding necessities, thus suggesting that in some way alpine marmots may envisage future needs. The main burrow systems and hibernacula were inherited by the new couples as well, while the spotting/resting points and other burrows varied across years and groups. The capability of the alpine marmots to maintain stable ...