Prevalence of tick-borne pathogens in ticks collected from the wild mountain ungulates mouflon and chamois in 4 regions of France.

Ticks are major vectors of various pathogens of health importance, such as bacteria, viruses and parasites. The problems associated with ticks and vector-borne pathogens are increasing in mountain areas, particularly in connection with global climate change. We collected ticks (n = 2,081) from chamois and mouflon in 4 mountainous areas of France. We identified 6 tick species: Ixodes ricinus, Rhipicephalus bursa, Rh. sanguineus s.l., Haemaphysalis sulcata, H. punctata and Dermacentor marginatus. We observed a strong variation in tick species composition among the study sites, linked in particular to the climate of the sites. We then analysed 791 ticks for DNA of vector-borne pathogens: Babesia/Theileria spp., Borrelia burgdorferi s.l., Anaplasma phagocytophilum, A. marginale, A. ovis, and Rickettsia of the spotted fever group (SFG). Theileria ovis was detected only in Corsica in Rh. bursa. Babesia venatorum (2 sites), Borrelia burgdorferi s.l. (B. afzelii and B. garinii; 2 sites) and Anaplasma phagocytophilum (3 sites) were detected in I. ricinus. Anaplasma ovis was detected at one site in I. ricinus and Rh. sanguineus s.l. SFG Rickettsia were detected at all the study sites: R. monacensis and R. helvetica in I. ricinus at the 3 sites where this tick is present; R. massiliae in Rh. sanguineus s.l. (1 site); and R. hoogstraalii and Candidatus R. barbariae in Rh. bursa in Corsica. These results show that there is a risk of tick-borne diseases for humans and domestic and wild animals frequenting these mountain areas.

Title: Prévalence d'agents pathogènes vectorisés chez des tiques collectées chez des ongulés sauvages (mouflons, chamois) dans 4 zones montagneuses en France. Abstract: Les tiques sont des vecteurs majeurs de différents agents pathogènes d'importance sanitaire, tels que des bactéries, des virus et des parasites. Les problématiques liées aux tiques et aux pathogènes vectorisés augmentent en zones de montagne, en lien notamment avec le réchauffement climatique. Nous avons collecté des tiques (n = 2 081) sur des chamois et des mouflons dans 4 zones montagneuses en France. Six espèces ont été identifiées : Ixodes ricinus, Rhipicephalus bursa, Rh. sanguineus s.l., Haemaphysalis sulcata, H. punctata et Dermacentor marginatus. Nous avons observé une forte variation de la composition en espèces de tiques entre les sites d'étude, en lien notamment avec le climat des sites. Nous avons ensuite recherché les ADN d'agents pathogènes vectorisés sur 791 tiques : Babesia/Theileria spp, Borrelia burgdorferi s.l., Anaplasma phagocytophilum, A. marginale, A. ovis, et de Rickettsia du groupe des fièvres boutonneuses (SFG). Theileria ovis a été détecté uniquement en Corse chez Rh. bursa. Babesia venatorum (2 sites), Borrelia burgdorferi s.l. (B. afzelii and B. garinii; 2 sites) et Anaplasma phagocytophilum (3 sites) ont été détectés chez I. ricinus. Anaplasma ovis a été détecté dans un site chez I. ricinus et Rh. sanguineus s.l.. Les Rickettsia SFG ont été détectées dans tous les sites d'étude : Rickettsia monacensis et R. helvetica chez I. ricinus dans les 3 sites où cette tique est présente; R. massiliae chez Rh. sanguineus s.l. (1 site); et R. hoogstraalii et Candidatus R. barbariae chez Rh. bursa en Corse. Ces résultats montrent un risque de transmission de maladies par les tiques pour les personnes et les animaux domestiques et sauvages fréquentant ces zones de montagne.

Interacting lethal and nonlethal human activities shape complex risk tolerance behaviors in a mountain herbivore.

Animals perceive human activities as risky and generally respond with fear-induced proactive behaviors to buffer the circadian patterns of lethal and nonlethal disturbances, such as diel migrations (DMs) between risky places during safe nighttime and safer places during risky daytime. However, such responses potentially incur costs through movement or reduced foraging time, so individuals should adjust their tolerance when human activities are harmless, through habituation. Yet this is a challenging cognitive task when lethal and nonlethal risks co-occur, forming complex landscapes of fear. The consequences of this human-induced complexity have, however, rarely been assessed. We studied the individual DM dynamics of chamois (Rupicapra rupicapra rupicapra), 89 GPS-tracked individual-years, from/to trails in the French Alps in areas with co-occurring lethal (hunting) and nonlethal (hiking and skiing) disturbances, with different intensities across seasons. We developed a conceptual framework relying on the risk-disturbance hypothesis and habituation to predict tolerance adjustments of chamois under various disturbance contexts and across contrasted seasonal periods. Based on spatial and statistical analyses combining periodograms and multinomial logistic models, we found that DM in relation to distance to a trail was a consistent response by chamois (~85% of individuals) to avoid human disturbance during daytime, especially during the hiking and hunting periods. Such behavior revealed a low tolerance of most chamois to human activities, although there was considerable interindividual heterogeneity in DM. Interestingly, there was an increased tolerance among the most disturbed diel migrants, potentially through habituation, with chamois performing shorter DMs in areas highly disturbed by hikers. Crucially, chamois that were most human-habituated during the hiking period remained more tolerant in the subsequent harvesting period, which could increase their risk of being harvested. In contrast, individuals less tolerant to hiking performed longer DMs when hunting risk increased, and compared to hiking, hunting exacerbated the threshold distance to trails triggering DMs. No carryover effect of hunting beyond the hunting period was observed. In conclusion, complex human-induced landscapes of fear with co-occurring disturbances by nature-based tourism and hunting may shape unexpected patterns of tolerance to human activities, whereby animal tolerance could become potentially deleterious for individual survival.

On this side of the fence: Functional responses to linear landscape features shape the home range of large herbivores.

Understanding the consequences of global change for animal movement is a major issue for conservation and management. In particular, habitat fragmentation generates increased densities of linear landscape features that can impede movements. While the influence of these features on animal movements has been intensively investigated, they may also play a key role at broader spatial scales (e.g. the home range scale) as resources, cover from predators/humans, corridors/barriers or landmarks. How space use respond to varying densities of linear features has been mostly overlooked in large herbivores, in contrast to studies done on predators. Focusing on large herbivores should provide additional insights to understand how animals solve the trade-off between energy acquisition and mortality risk. Here, we investigated the role of anthropogenic (roads and tracks) and natural (ridges, valley bottoms and forest edges) linear features on home range features in five large herbivores. We analysed an extensive GPS monitoring database of 710 individuals across nine populations, ranging from mountain areas mostly divided by natural features to lowlands that were highly fragmented by anthropogenic features. Nearly all of the linear features studied were found at the home range periphery, suggesting that large herbivores primarily use them as landmarks to delimit their home range. In contrast, for mountain species, ridges often occurred in the core range, probably related to their functional role in terms of resources and refuge. When the density of linear features was high, they no longer occurred predominantly at the home range periphery, but instead were found across much of the home range. We suggest that, in highly fragmented landscapes, large herbivores are constrained by the costs of memorising the spatial location of key features, and by the requirement for a minimum area to satisfy their vital needs. These patterns were mostly consistent in both males and females and across species, suggesting that linear features have a preponderant influence on how large herbivores perceive and use the landscape.

Positive effect of spring advance on the diet quality of an alpine herbivore.

Changes in vegetation phenology related to global warming are having alarming effects on the life history traits of many herbivore species. Such changes are particularly critical in alpine ecosystems, where strong climate limitations on plant growth make seasonal synchronization imperative for the growth, reproduction and survival of herbivores. However, despite the pivotal role of resource-use strategies on the performances of such species, few studies have explicitly assessed the mechanistic impact of climate change on their diets. We aimed to fill this gap by studying the effect of spring onset on the dietary composition and quality of a medium-size alpine herbivore while considering density-dependent processes and age- and sex-specific differences in foraging behavior. Using an exceptional, long-term (24 years) direct individual-based dietary monitoring of a Pyrenean chamois population (Rupicapra pyrenaica pyrenaica), we showed that ongoing earlier onsets of spring are leading to an earlier access to high-quality forage and therefore a higher diet quality at a fixed date, without apparent changes in diet composition. We also showed that at high densities, intraspecific competition reduced diet quality by driving animals to feed more on woody plants and less on nutritious forbs and graminoids. By assessing the mechanistic effects of global warming on the dietary patterns of species at the center of trophic networks, this study is an essential step for predictive models aiming at understanding the ongoing ecosystem consequences of the global climatic crisis.

Near Infrared Reflectance Spectroscopy Analysis to Predict Diet Composition of a Mountain Ungulate Species.

The diet composition of ungulates is important to understand not only their impact on vegetation, but also to understand the consequences of natural and human-driven environmental changes on the foraging behavior of these mammals. In this work, we evaluated the use of near infrared reflectance spectroscopy analysis (NIRS), a quick, economic and non-destructive method, to assess the diet composition of the Pyrenean chamois Rupicapra pyrenaica pyrenaica. Fecal samples (n = 192) were collected from two chamois populations in the French and Spanish Pyrenees. Diet composition was initially assessed by fecal cuticle microhistological analysis (CMA) and categorized into four functional groups, namely: woody, herbaceous, graminoid and Fabaceae plants. Regressions of modified partial least squares and several combinations of scattering correction and derivative treatments were tested. The results showed that models based on the second derivative processing obtained the higher determination coefficient for woody, herbaceous and graminoid plants (R2CAL, coefficient of determination in calibration, ranged from 0.86 to 0.91). The Fabaceae group, however, was predicted with lower accuracy (R2CAL = 0.71). Even though an agreement between NIRS and CMA methods was confirmed by a Bland-Altman analysis, confidence limits of agreement differed by up to 25%. Our results support the viability of fecal NIRS analysis to study spatial and temporal variations of the Pyrenean chamois' diets in summer and winter when differences in the consumption of woody and annual plants are the greatest. This new use for the NIRS technique would be useful to assess the consequences of global change on the feeding behavior of this mountain ungulate and also in other ungulate counterparts.

Reproductive females and young mouflon (Ovis gmelini musimon × Ovis sp.) in poor body condition are the main spreaders of gastrointestinal parasites.

Several individual, environmental and parasitic factors can influence the impacts of parasites on host's fitness and on host's ability to transmit these parasites to new hosts. Identifying these factors and the individuals who play a greater role in parasite transmission is of main concern for the development of parasite control strategies. In the present study, we aimed to describe the diversity of gastrointestinal parasites and to identify the individual factors influencing the faecal spreading of parasites in a free-ranging population of Mediterranean mouflon. From the analysis of 433 faecal samples, we found Eimeria spp. and gastrointestinal strongyles (GIS) were the most common parasites (>94%). The faecal oocyst counts of Eimeria spp. were the highest during the first years of life. It was 1.6 times higher in females than in males and 2.5 times higher in individuals in poor than in good body condition. Similarly, the faecal egg count of GIS was higher in females and decreased with age, but only in males. Finally, reproductive females had GIS faecal egg count values 2.6 times higher than non-reproductive females. Management strategies of parasites should thus primarily focus on reproductive females and young individuals in poor body condition as they represent the main contamination source of the environment.

Variation in the ontogenetic allometry of horn length in bovids along a body mass continuum.

Allometric relationships describe the proportional covariation between morphological, physiological, or life-history traits and the size of the organisms. Evolutionary allometries estimated among species are expected to result from species differences in ontogenetic allometry, but it remains uncertain whether ontogenetic allometric parameters and particularly the ontogenetic slope can evolve. In bovids, the nonlinear evolutionary allometry between horn length and body mass in males suggests systematic changes in ontogenetic allometry with increasing species body mass. To test this hypothesis, we estimated ontogenetic allometry between horn length and body mass in males and females of 19 bovid species ranging from ca. 5 to 700 kg. Ontogenetic allometry changed systematically with species body mass from steep ontogenetic allometries over a short period of horn growth in small species to shallow allometry with the growth period of horns matching the period of body mass increase in the largest species. Intermediate species displayed steep allometry over long period of horn growth. Females tended to display shallower ontogenetic allometry with longer horn growth compared to males, but these differences were weak and highly variable. These findings show that ontogenetic allometric slope evolved across species possibly as a response to size-related changes in the selection pressures acting on horn length and body mass.

Does host socio-spatial behavior lead to a fine-scale spatial genetic structure in its associated parasites?

Gastro-intestinal nematodes, especially Haemonchus contortus, are widespread pathogenic parasites of small ruminants. Studying their spatial genetic structure is as important as studying host genetic structure to fully understand host-parasite interactions and transmission patterns. For parasites having a simple life cycle (e.g., monoxenous parasites), gene flow and spatial genetic structure are expected to strongly rely on the socio-spatial behavior of their hosts. Based on five microsatellite loci, we tested this hypothesis for H. contortus sampled in a wild Mediterranean mouflon population (Ovis gmelini musimon × Ovis sp.) in which species- and environment-related characteristics have been found to generate socio-spatial units. We nevertheless found that their parasites had no spatial genetic structure, suggesting that mouflon behavior was not enough to limit parasite dispersal in this study area and/or that other ecological and biological factors were involved in this process, for example other hosts, the parasite life cycle, or the study area history.

TITLE: Le comportement socio-spatial de l'hôte conduit-il à une structure génétique à fine échelle de ses parasites ? ABSTRACT: Les nématodes gastro-intestinaux, et plus particulièrement Haemonchus contortus, sont cosmopolites et pathogènes chez les petits ruminants. Étudier leur structure génétique spatiale est aussi important que d'étudier celle des hôtes pour pleinement comprendre les interactions hôtes-parasites et les processus de transmission. Pour les parasites ayant des cycles de vie simples (par exemple, les parasites monoxènes), on s'attend à ce que les flux de gènes et la structure génétique spatiale dépendent fortement du comportement socio-spatial de leurs hôtes. En utilisant cinq loci microsatellites, nous avons testé cette hypothèse pour des H. contortus échantillonnés dans une population sauvage de mouflons méditerranéens (Ovis gmelini musimon × Ovis sp.) dans laquelle les caractéristiques de l'espèce et de l'environnement génèrent des unités socio-spatiales. Nous avons néanmoins mis en évidence que leurs parasites ne présentent pas de structure génétique spatiale, ce qui suggère que le comportement des mouflons ne restreint pas la dispersion des parasites dans cette aire d'étude et/ou que d'autres facteurs biologiques et écologiques tels que d'autres hôtes, le cycle de vie du parasite, ou l'histoire de l'aire d'étude jouent un rôle dans ce processus.

Detection of tick-borne pathogens in questing Ixodes ricinus in the French Pyrenees and first identification of Rickettsia monacensis in France.

Ticks are important vectors of several human and animal pathogens. In this study, we estimated the prevalence of important tick-borne infections in questing ticks from an area in Southwestern France (Hautes-Pyrénées) inhabited by Pyrenean chamois (Rupicapra pyrenaica pyrenaica) experiencing high tick burden. We examined adult and nymph ticks collected by the flag dragging method from 8 to 15 sites in the Pic de Bazès during the years 2009, 2011, 2013 and 2015. PCR assays were conducted on selected ticks for the detection of Borrelia burgdorferi s.l., Babesia spp., Rickettsia spp., spotted fever group (SFG) Rickettsia and Anaplasma phagocytophilum. Randomly selected positive samples were submitted for sequence analysis. A total of 1971 questing ticks were collected including 95 males, 101 females and 1775 nymphs. All collected ticks were identified as Ixodes ricinus. Among them, 696 ticks were selected for pathogen detection and overall prevalence was 8.4% for B. burgdorferi s.l.; 0.4% for Babesia spp.; 6.1% for A. phagocytophilum; 17.6% for Rickettsia spp.; and 8.1% for SFG Rickettsia. Among the sequenced pathogens, we detected in this population of ticks the presence of Babesia sp. EU1 and Rickettsia helvetica, as well as Rickettsia monacensis for the first time in France. The detection of these pathogens in the Pic de Bazès highlights the potential infection risks for visitors to this area and the Pyrenean chamois population.

TITLE: Détection d'agents pathogènes présents chez les tiques errantes, Ixodes ricinus, dans les Pyrénées françaises et première identification de Rickettsia monacensis en France. ABSTRACT: Les tiques sont des vecteurs importants de plusieurs maladies animales et humaines. Dans cette étude, nous avons estimé la prévalence de maladies vectorisées chez les tiques errantes dans une région du sud-ouest de la France (Hautes-Pyrénées) occupée par des chamois des Pyrénées (Rupicapra pyrenaica pyrenaica) fortement infestés par les tiques. Nous avons utilisé des tiques adultes et des larves collectées par la technique du drapeau sur 8­15 sites du Pic de Bazès au cours des années 2009, 2011, 2013 et 2015. Des analyses PCR ont été menées sur des tiques sélectionnées pour la détection de Borrelia burgdorferi s.l., Babesia spp., Rickettsia spp., Rickettsia du groupe de la fièvre pourprée (GFP) et Anaplasma phagocytophilum. Des échantillons positifs sélectionnés au hasard ont été soumis à une analyse de séquences. Un total de 1971 tiques a été collecté dont 95 mâles, 101 femelles et 1775 larves. Toutes les tiques recueillies ont été identifiées comme étant Ixodes ricinus. Parmi celles-ci, 696 tiques ont été sélectionnées pour la détection des agents pathogènes et la prévalence globale était de 8,4 % pour B. burgdorferi s.l., 0,4 % pour Babesia spp., 6,1 % pour A. phagocytophilum, 17,6 % pour Rickettsia spp. et 8,1 % pour Rickettsia GFP. Parmi les agents pathogènes séquencés, nous avons détecté dans cette population de tiques la présence de Babesia sp. EU1, Rickettsia helvetica et, pour la première fois en France, Rickettsia monacensis. La détection de ces agents pathogènes dans le Pic de Bazès souligne les risques potentiels d'infection pour les visiteurs de cette région et pour la population de chamois des Pyrénées.

Both candidate gene and neutral genetic diversity correlate with parasite resistance in female Mediterranean mouflon.

BACKGROUND: Parasite infections can have substantial impacts on population dynamics and are accordingly a key challenge for wild population management. Here we studied genetic mechanisms driving parasite resistance in a large herbivore through a comprehensive approach combining measurements of neutral (16 microsatellites) and adaptive (MHC DRB1 exon 2) genetic diversity and two types of gastrointestinal parasites (nematodes and coccidia). RESULTS: While accounting for other extrinsic and intrinsic predictors known to impact parasite load, we show that both neutral genetic diversity and DRB1 are associated with resistance to gastrointestinal nematodes. Intermediate levels of multi-locus heterozygosity maximized nematodes resistance, suggesting that both in- and outbreeding depression might occur in the population. DRB1 heterozygosity and specific alleles effects were detected, suggesting the occurrence of heterozygote advantage, rare-allele effects and/or fluctuating selection. On the contrary, no association was detected between genetic diversity and resistance to coccidia, indicating that different parasite classes are impacted by different genetic drivers. CONCLUSIONS: This study provides important insights for large herbivores and wild sheep pathogen management, and in particular suggests that factors likely to impact genetic diversity and allelic frequencies, including global changes, are also expected to impact parasite resistance.

Making sense of ultrahigh-resolution movement data: A new algorithm for inferring sites of interest.

Decomposing the life track of an animal into behavioral segments is a fundamental challenge for movement ecology. The proliferation of high-resolution data, often collected many times per second, offers much opportunity for understanding animal movement. However, the sheer size of modern data sets means there is an increasing need for rapid, novel computational techniques to make sense of these data. Most existing methods were designed with smaller data sets in mind and can thus be prohibitively slow. Here, we introduce a method for segmenting high-resolution movement trajectories into sites of interest and transitions between these sites. This builds on a previous algorithm of Benhamou and Riotte-Lambert (2012). Adapting it for use with high-resolution data. The data's resolution removed the need to interpolate between successive locations, allowing us to increase the algorithm's speed by approximately two orders of magnitude with essentially no drop in accuracy. Furthermore, we incorporate a color scheme for testing the level of confidence in the algorithm's inference (high = green, medium = amber, low = red). We demonstrate the speed and accuracy of our algorithm with application to both simulated and real data (Alpine cattle at 1 Hz resolution). On simulated data, our algorithm correctly identified the sites of interest for 99% of "high confidence" paths. For the cattle data, the algorithm identified the two known sites of interest: a watering hole and a milking station. It also identified several other sites which can be related to hypothesized environmental drivers (e.g., food). Our algorithm gives an efficient method for turning a long, high-resolution movement path into a schematic representation of broadscale decisions, allowing a direct link to existing point-to-point analysis techniques such as optimal foraging theory. It is encoded into an R package called SitesInterest, so should serve as a valuable tool for making sense of these increasingly large data streams.

Seasonal diet composition of Pyrenean chamois is mainly shaped by primary production waves.

In alpine habitats, the seasonally marked climatic conditions generate seasonal and spatial differences in forage availability for herbivores. Vegetation availability and quality during the growing season are known to drive life history traits of mountain ungulates. However, little effort has been made to understand the association between plant phenology and changes in the foraging strategies of these mountain dwellers. Furthermore, this link can be affected by the seasonal presence of livestock in the same meadows. The objective of this work was to study the seasonal changes in diet composition of Pyrenean chamois (Rupicapra p. pyrenaica) and its relationship to primary production trends in a Mediterranean alpine environment. Moreover, diet composition in two populations with contrasting livestock pressure was compared in order to study the effect of sheep flocks on the feeding behaviour of chamois. From 2009 to 2012, monthly diet composition was estimated by cuticle microhistological analysis of chamois faeces collected in the eastern Pyrenees. The primary production cycle was assessed by remote sensing, using the Normalized Difference Vegetation Index. Additionally, the diet of sheep sharing seasonally the subalpine and alpine meadows with chamois was analysed. Diet selection of chamois and sheep and their overlap was also assessed. Our results show an intra-annual variation in the diet composition of Pyrenean chamois and demonstrate a strong relationship between plant consumption dynamics and phenology in alpine areas. In addition, Calluna vulgaris, Cytisus spp. and Festuca spp., as well as forbs in the summer, are found to be key forage species for Pyrenean chamois. Furthermore, this study couldn't detect differences between both chamois populations despite the presence of sheep flocks in only one area. However, the detection of a shift in the diet of chamois in both areas after the arrival of high densities of multi-specific livestock suggest a general livestock effect. In conclusion, Pyrenean chamois are well adapted to the variations in the seasonal availability of plants in alpine habitats but could be disturbed by the seasonal presence of livestock. Due to the key plants in their diet, we suggest that population management programmes should focus on the preservation of mixed grasslands composed of patches of shrubs and herbs. The effects of climate change and shrub expansion should be studied as they may potentially affect chamois population dynamics through changes in habitat composition and temporal shifts in forage availability.

Demographic stochasticity drives epidemiological patterns in wildlife with implications for diseases and population management.

Infectious diseases raise many concerns for wildlife and new insights must be gained to manage infected populations. Wild ungulates provide opportunities to gain such insights as they host many pathogens. Using modelling and data collected from an intensively monitored population of Pyrenean chamois, we investigated the role of stochastic processes in governing epidemiological patterns of pestivirus spread in both protected and hunted populations. We showed that demographic stochasticity led to three epidemiological outcomes: early infection fade-out, epidemic outbreaks with population collapse, either followed by virus extinction or by endemic situations. Without re-introduction, the virus faded out in >50% of replications within 4 years and did not persist >20 years. Test-and-cull of infected animals and vaccination had limited effects relative to the efforts devoted, especially in hunted populations in which only quota reduction somewhat improve population recovery. Success of these strategies also relied on the maintenance of a high level of surveillance of hunter-harvested animals. Our findings suggested that, while surveillance and maintenance of population levels at intermediate densities to avoid large epidemics are useful at any time, a 'do nothing' approach during epidemics could be the 'least bad' management strategy in populations of ungulates species facing pestivirus infection.

Circadian periodicity in space use by ungulates of temperate regions: How much, when and why?

When they visit and revisit specific areas, animals may reveal what they need from their home range and how they acquire information. The temporal dimension of such movement recursions, that is, periodicity, is however rarely studied, yet potentially bears a species, population or individual-specific signature. A recent method allows estimating the contribution of periodic patterns to the variance in a movement path. We applied it to 709 individuals from five ungulate species, looking for species signatures in the form of seasonal variation in the intensity of circadian patterns. Circadian patterns were commonplace in the movement tracks, but the amount of variance they explained was highly variable among individuals. It increased in intensity during spring and summer, when key resources were spatially segregated, and decreased during winter, when food availability was more uniformly low. Other periodicity-inducing mechanisms supported by our comparison of species- and sex-specific patterns involve young antipredator behaviour, territoriality and behavioural thermoregulation. Model-based continuous-time movement metrics represent a new avenue for researchers interested in finding individual-, population- or species-specific signatures in heterogeneous movement databases featuring various study designs and sampling resolutions. However, we observed large amounts of individual variation, so comparative analyses should ideally use both GPS and animal-borne loggers to augment the discriminatory power and be based on large samples. We briefly outline potential uses of the intensity of circadian patterns as a metric for the study of animal personality and community ecology.

Self-clearance of Pestivirus in a Pyrenean Chamois ( Rupicapra pyrenaica) Population.

Understanding the dynamics of host-pathogen interaction is key to the management of epidemics. A pestivirus belonging to the border disease virus group 4 emerged around 2001 in Pyrenean chamois ( Rupicapra pyrenaica) in Spain and France. The virus had significant demographic impact in some populations, but it was less harmful and more endemic in other places. The determinants of these local variations are still unclear. Here, we documented empirical evidence of self-clearance of the virus in a chamois population in France. This population has regularly been counted, and chamois were trapped and harvested each year, providing unique demographic and epidemiologic surveys of the population since 1984 and 1994, respectively. The virus was detected using direct (PCR) and indirect (antibody) testing. We showed that virus transmission declined in 2011-12 and likely ceased in 2013, leading to a decline in antibody prevalence since 2014. Self-clearance may be due to limited exchanges with other populations, decrease in population size after an epizootic, and herd immunity. The age structure of captured animals shifted to younger age classes after virus self-clearance, suggesting a return to a colonizing population structure. The possible consequences of virus re-entry are discussed. This observation suggests that pestivirus dynamics occurs at the scale of the metapopulation of Pyrenean chamois. Local self-clearance and re-emergence may help explain the variation of virus dynamics at the local scale.

Introduction history overrides social factors in explaining genetic structure of females in Mediterranean mouflon.

Fine-scale spatial genetic structure of populations results from social and spatial behaviors of individuals such as sex-biased dispersal and philopatry. However, the demographic history of a given population can override such socio-spatial factors in shaping genetic variability when bottlenecks or founder events occurred in the population. Here, we investigated whether socio-spatial organization determines the fine-scale genetic structure for both sexes in a Mediterranean mouflon (Ovis gmelini musimon × Ovis sp.) population in southern France 60 years after its introduction. Based on multilocus genotypes at 16 loci of microsatellite DNA (n = 230 individuals), we identified three genetic groups for females and two for males, and concurrently defined the same number of socio-spatial units using both GPS-collared individuals (n = 121) and visual resightings of marked individuals (n = 378). The socio-spatial and genetic structures did not match, indicating that the former was not the main driver of the latter for both sexes. Beyond this structural mismatch, we found significant, yet low, genetic differentiation among female socio-spatial groups, and no genetic differentiation in males, with this suggesting female philopatry and male-biased gene flow, respectively. Despite spatial disconnection, females from the north of the study area were genetically closer to females from the south, as indicated by the spatial analysis of the genetic variability, and this pattern was in accordance with the common genetic origin of their founders. To conclude, more than 14 generations later, genetic signatures of first introduction are not only still detectable among females, but they also represent the main factor shaping their present-time genetic structure.

From gestation to weaning: Combining robust design and multi-event models unveils cost of lactation in a large herbivore.

The cost of current reproduction on survival or future reproduction is one of the most studied trade-offs governing resource distribution between fitness components. Results have often been clouded, however, by the existence of individual heterogeneity, with high-quality individuals able to allocate energy to several functions simultaneously, at no apparent cost. Surprisingly, it has also rarely been assessed within a breeding season by breaking down the various reproductive efforts of females from gestation to weaning, even though resource availability and energy requirements vary greatly. We filled this gap by using an intensively monitored population of Pyrenean chamois and by expanding a new methodological approach integrating robust design in a multi-event framework. We distinguished females that gave birth or not, and among reproducing females whether they lost their kid or successfully raised it until weaning. We estimated spring and summer juvenile survival, investigated whether gestation, lactation or weaning incurred costs on the next reproductive occasion, and assessed how individual heterogeneity influenced the detection of such costs. Contrary to expectations if trade-offs occur, we found a positive relationship between gestation and adult survival suggesting that non-breeding females are in poor condition. Costs of reproduction were expressed through negative relationships between lactation and both subsequent breeding probability and spring juvenile survival. Such costs could be detected only once individual heterogeneity (assessed as two groups contrasting good vs. poor breeders) and time variations in juvenile survival were accounted for. Early lactation decreased the probability of future reproduction, providing quantitative evidence of the fitness cost of this period recognized as the most energetically demanding in female mammals and critical for neonatal survival. The new approach employed made it possible to estimate two components of kid survival that are often considered practically unavailable in free-ranging populations, and also revealed that reproductive costs appeared only when contrasting the different stages of reproductive effort. From an evolutionary perspective, our findings stressed the importance of the temporal resolution at which reproductive cost is studied, and also provided insights on the reproductive period during which internal and external factors would be expected to have the greatest fitness impact.

Predicting herbivore faecal nitrogen using a multispecies near-infrared reflectance spectroscopy calibration.

Optimal management of free-ranging herbivores requires the accurate assessment of an animal's nutritional status. For this purpose 'near-infrared reflectance spectroscopy' (NIRS) is very useful, especially when nutritional assessment is done through faecal indicators such as faecal nitrogen (FN). In order to perform an NIRS calibration, the default protocol recommends starting by generating an initial equation based on at least 50-75 samples from the given species. Although this protocol optimises prediction accuracy, it limits the use of NIRS with rare or endangered species where sample sizes are often small. To overcome this limitation we tested a single NIRS equation (i.e., multispecies calibration) to predict FN in herbivores. Firstly, we used five herbivore species with highly contrasting digestive physiologies to build monospecies and multispecies calibrations, namely horse, sheep, Pyrenean chamois, red deer and European rabbit. Secondly, the equation accuracy was evaluated by two procedures using: (1) an external validation with samples from the same species, which were not used in the calibration process; and (2) samples from different ungulate species, specifically Alpine ibex, domestic goat, European mouflon, roe deer and cattle. The multispecies equation was highly accurate in terms of the coefficient of determination for calibration R2 = 0.98, standard error of validation SECV = 0.10, standard error of external validation SEP = 0.12, ratio of performance to deviation RPD = 5.3, and range error of prediction RER = 28.4. The accuracy of the multispecies equation to predict other herbivore species was also satisfactory (R2 > 0.86, SEP < 0.27, RPD > 2.6, and RER > 8.1). Lastly, the agreement between multi- and monospecies calibrations was also confirmed by the Bland-Altman method. In conclusion, our single multispecies equation can be used as a reliable, cost-effective, easy and powerful analytical method to assess FN in a wide range of herbivore species.

The physiological cost of male-biased parasitism in a nearly monomorphic mammal.

BACKGROUND: Even though male-biased parasitism is common in mammals, little effort has been made to evaluate whether higher parasitic burden in males results in an extra biological cost, and thus a decrease in fitness. Body condition impairment and the augmentation of oxidative stress can be used as indicators of the cost of parasite infections. Here, we examined relationships between gastrointestinal and respiratory helminths, body condition and oxidative stress markers (glutathione peroxidase, paraoxonase-1) in 28 Pyrenean chamois (Rupicapra p. pyrenaica) sampled in autumn. RESULTS: Only male chamois showed a reduction in body condition and higher oxidative stress due to parasite infection, likely because of the extremely high parasite burdens observed in males. CONCLUSIONS: This study made evident a disparity in the physiological cost of multiple parasitism between sexes in a wild mammal, mainly due to parasitic richness. Because of the similar life expectancy in male and female chamois, we suggest that males may have developed natural mechanisms to compensate for higher parasite loads during the rut.

Combining familiarity and landscape features helps break down the barriers between movements and home ranges in a non-territorial large herbivore.

Recent advances in animal ecology have enabled identification of certain mechanisms that lead to the emergence of territories and home ranges from movements considered as unbounded. Among them, memory and familiarity have been identified as key parameters in cognitive maps driving animal navigation, but have been only recently used in empirical analyses of animal movements. At the same time, the influence of landscape features on movements of numerous species and on space division in territorial animals has been highlighted. Despite their potential as exocentric information in cognitive maps and as boundaries for home ranges, few studies have investigated their role in the design of home ranges of non-territorial species. Using step selection analyses, we assessed the relative contribution of habitat characteristics, familiarity preferences and linear landscape features in movement step selection of 60 GPS-collared Mediterranean mouflon Ovis gmelini musimon × Ovis sp. monitored in southern France. Then, we evaluated the influence of these movement-impeding landscape features on the design of home ranges by testing for a non-random distribution of these behavioural barriers within sections of space differentially used by mouflon. We reveal that familiarity and landscape features are key determinants of movements, relegating to a lower level certain habitat constraints (e.g. food/cover trade-off) that we had previously identified as important for this species. Mouflon generally avoid crossing both anthropogenic (i.e. roads, tracks and hiking trails) and natural landscape features (i.e. ridges, talwegs and forest edges) while moving in the opposite direction, preferentially toward familiar areas. These specific behaviours largely depend on the relative position of each movement step regarding distance to the landscape features or level of familiarity in the surroundings. We also revealed cascading consequences on the design of home ranges in which most landscape features were excluded from cores and relegated to the peripheral areas. These results provide crucial information on landscape connectivity in a context of marked habitat fragmentation. They also call for more research on the role of landscape features in the emergence of home ranges in non-territorial species using recent methodological developments bridging the gap between movements and space use patterns.