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In seasonal environments, the timing of reproduction often matches with the peak of food resources. One well-known effect of global warming is an earlier phenology of resources, leading to a possible mismatch between the timing of reproduction for consumers and food peak. However, global warming may also change the dynamics of food resources, such as the intensity and frequency of pulsed mast seeding. How quantitative changes in mast seeding influence the timing of reproduction of seed consumers remains unexplored. Here, we assess how yearly variation in mast seeding influences mating time in wild boar ( Sus scrofa ), a widespread seed consumer species. We took advantage of the intensive monitoring of both female reproduction (1636 females) and acorn production over 6 consecutive years across 15 populations of wild boar in the wild. We found that mating time occurs earlier when acorn production increases in most but not all populations. In two out of 15 populations, heavy females mated earlier than light ones. Our findings demonstrate that mast seeding advances the mating time in some populations, which could perhaps impact how boars respond to climate change.

European wildlife has been subjected to intensifying levels of anthropogenic impact throughout the Holocene, yet the main genetic partitioning of many species is thought to still reflect the late-Pleistocene glacial refugia. We analyzed 26,342 nuclear SNPs of 464 wild boar (Sus scrofa) across the European continent to infer demographic history and reassess the genetic consequences of natural and anthropogenic forces. We found that population fragmentation, inbreeding and recent hybridization with domestic pigs have caused the spatial genetic structure to be heterogeneous at the local scale. Underlying local anthropogenic signatures, we found a deep genetic structure in the form of an arch-shaped cline extending from the Dinaric Alps, via Southeastern Europe and the Baltic states, to Western Europe and, finally, to the genetically diverged Iberian peninsula. These findings indicate that, despite considerable anthropogenic influence, the deeper, natural continental structure is still intact. Regarding the glacial refugia, our findings show a weaker signal than generally assumed, but are nevertheless suggestive of two main recolonization routes, with important roles for Southern France and the Balkans. Our results highlight the importance of applying genomic resources and framing genetic results within a species’ demographic history and geographic distribution for a better understanding of the complex mixture of underlying processes. Nasl. z nasl. zaslona. Opis vira z dne 3. 2. 02023. Skupno št. avtorjev: 21. Bibliografija: str. 142-143.

African Swine Fever (ASF) has been slowly but steadily increasing its endemic range throughout Europe, posing an imminent risk to the pig industry. ASF transmission among wild boar occurs mainly through wild boar population movements, hence wild boar presence and density are important risk factors for introducing, maintaining, and spreading the disease. The understanding of wild boar population dynamics and their role in ASF transmission and persistence remains limited. It is crucial to gain knowledge in this area to improve wildlife management while minimizing the risks for ASF introduction and spread. We adapted an individual-based spatio-temporal stochastic model developed by Halasa et al. (2019) and tailored it to two regions in France. The model assessed yearly hunting activity, the carcass persistence seasonality, and the specific landscape characteristics of the Franco-Belgian border region and the Pyrénées-Atlantiques department. Following the establishment of local population dynamics through preliminary runs of the model, the model was run 100 iterations over 8 years in the two study areas where ASF was randomly seeded after the 2nd year of simulation. For each scenario, the model was initiated with 500 wild boar groups randomly spread across the study areas. Hunting activities were included and excluded to assess the impact on population growth and ASF spread. Results showed an ever-growing wild boar population for all scenarios, which was balanced when hunting activities were included. When introducing ASF, the wild boar populations were dramatically impacted in both areas with a decrease of 63 % of the population at the Franco-Belgian border and 86 % in the Pyrénées-Atlantiques department. Habitat fragmentation and landscape connectivity were highlighted as important factors shaping ASF propagation. The Franco-Belgian border, which had the most fragmented habitat with unsuitable areas for wild boars, was shown to limit wild boar movements, reducing the probability, and spread of ASF across the landscape. The lack of connectivity was reflected in a less effective transmission and lower number of infected groups (406 versus 467). In contrast, the epidemic duration was lengthened in the fragmented habitat compared to the homogenous area (2.6 years vs 1.6 years). This study provided information on defining and implementing control measures in case of an ASF incursion, since delimitation of the area via fences artificially induces landscape fragmentation, which is important for controlling ASF outbreaks.

International audience; In Europe, African swine fever virus (ASFV) is one of the most threatening infectious transboundary diseases of domestic pigs and wild boar. In September 2018, ASF was detected in wild boar in the South of Belgium. France, as a bordering country, is extremely concerned about the ASF situation in Belgium, and an active preparedness is ongoing in the country. One of the questions raised by this situation relates to disturbing activities that may affect wild boar movements and their possible impact on the spread of ASFV. Despite evidence of disturbance related to hunting practices, there is a paucity of information on the impact of forestry and human leisure activities. To assess this impact on wild boar movements, a systematic review was first conducted but very few useful data were obtained. For this reason, an expert elicitation was carried out by the French Agency for Food, Environmental and Occupational Health & Safety in order to deal with this knowledge gap. A total of 30 experts originating from France and adjacent neighbouring countries (Spain, Belgium and Switzerland) were elicited about the relative importance of six factors of spatial disturbance of wild boar (noise, smell, invasion of space, modification of the environment, duration and frequency of the activity). Then, for each factor of disturbance, they were asked about the impact of 16 different commercial forestry and human leisure activities. A global weighted score was estimated in order to capture the variability of a wide range of territorial conditions and the uncertainty of expert elicitation. This estimate permitted ranking all 16 activities and aggregating them in three groups according to their potential for disturbance of wild boar, using a regression tree analysis. The results of this expert elicitation provide a methodological approach that may be useful for French and other European decision makers and stakeholders involved in the crisis management of ASF.

Intraspecific variations in mating systems have been reported in numerous species, especially when they live in varying ecological contexts. This leads to variability between populations with regard to the proportion of females engaging in multiple male mating, which depends on the number of males available. For hunted ungulate species, hunting is known to influence population structure, especially when males are preferentially targeted for trophy hunting. Here, we investigated how variations in hunting pressure and the yearly proportion of heavy males removed have impacted multiple paternity rates in five wild boar (Sus scrofa scrofa) populations located in similar ecological contexts. We found high rates of multiple paternity in all studied populations, confirming the recently reported promiscuous mating system of wild boar. However, variations in hunting pressure and removal of heavy males did not significantly influence multiple paternity rates, contrary to our expectation. Nonetheless, a slight tendency for a decreasing multiple paternity rate with increasing hunting pressure and for increasing multiple paternity rate with increasing removal of heavy males from the population was detected. Based on these results, we discuss an alternative hypothesis on the ecological processes sustaining the influence of hunting regimes on the mating system. Overall, hunting pressure and management rules might be sufficient to disrupt the mating system in any of the populations, so it is important to continue the sampling of wild boar populations at the European scale, especially in populations with little hunting pressure.

International audience; For sustainable management of exploited populations, it is required to have good knowledge on temporal trends in population density to adapt the harvest. In this regard, hunting statistics are often collected routinely by government agencies and associations. These data are used to assess demographic trends through the development of indices, which are in turn used to manage exploited populations in a sustainable way. However, these population indices depend on features of the hunting process (e.g. hunting effort, hunting conditions, probability of catch). In this study, we show how to use hunting logs to assess demographic trends in exploited populations while accounting for the components of the hunting process. In particular, we developed a catch-effort model to study how the hunting effort leads to mortality rate – hunting pressure – within a given habitat type and during a given period. We illustrated the usefulness of this approach using exploited wild boar (Sus scrofa) populations as a case study. We used a large hunting logs dataset to perform our study, with several hundreds of thousands hunting events for more than 10 years in two French departments in France, including information about the number of hunters, of wild boars culled and the date of the hunt. We showed that catchability is a key parameter to assess hunting pressure at a given time and place. This parameter varies both within the hunting season and between habitat types. Once this variation in catchability was accounted for, our catch-effort model allowed us to obtain estimates of relative densities of wild boar populations over the study period at the management unit scale. Thus, catch-effort models are powerful tools to assess population density and to understand the underlying hunting process. Our study offers straightforward and reproducible conceptual framework that can be applied routinely by wildlife managers on exploited populations and practitioners from hunting statistics logs.

International audience; Many animal populations are subject to hunting or fishing in the wild. Detailed knowledge of demographic parameters (e.g. survival, reproduction) and temporal dynamics of such populations is crucial for sustainable management. Despite their relevance for management decisions, structure and size of exploited populations are often not known, and data limited. Recently, joint analysis of different types of demographic data, such as population counts, reproductive data and capture-mark-recapture data, within integrated population models (IPMs) has gained much popularity as it may allow estimating population size and structure, as well as key demographic rates, while fully accounting for uncertainty. IPMs built so far for exploited populations have typically been built as age-structured population models. However, the age of harvested individuals is usually difficult and/or costly to assess and therefore often not available. Here, we introduce an IPM structured by body size classes, which allows making efficient use of data commonly available in exploited populations for which accurate information on age is often missing. The model jointly analyzes size-at-harvest data, capture-mark-recapture-recovery data and reproduction data from necropsies, and we illustrate its applicability in a case study involving heavily hunted wild boar. This species has increased in abundance over the last decades despite intense harvest, and the IPM analysis provides insights into the roles of natural mortality, body growth, maturation schedules and reproductive output in compensating for the loss of individuals to hunting. Early maturation and high reproductive output contributed to wild boar population persistence despite a strong hunting pressure. We thus demonstrate the potential of size-class-structured IPMs as tools to investigate the dynamics of exploited populations with limited information on age, and highlight both the applicability of this framework to other species and its potential for follow-up analyses highly relevant to management.

Weather conditions and population density individuals experience at birth influence their life‐history traits and thereby population dynamics. Early‐life individual growth is a key fitness‐related trait; however, how it is affected by such conditions at birth remains to be explored. Taking advantage of long‐term monitoring of three wild boar (Sus scrofa) populations living in contrasting ecological contexts, we assess how weather conditions (temperature and precipitation) and the number of removed individuals at birth influence early‐life growth rates. We found that the number of individuals removed before the early‐growth period had a positive effect on early‐life growth rate across sites. This might be interpreted as a density‐dependent response involving an increase in food availability per capita that favors faster growth. Alternatively, if the number of removed individuals increases with population density, this result might be attributable to decreasing litter sizes at high density, leading mothers to allocate more resources to individual offspring, which favors higher juvenile growth rates. Early‐life growth rates also increased with springtime temperature and decreasing precipitation. Thus, early‐life growth is expected to increase in response to warmer and drier springs, which should become more frequent in the future under current climate change. We found that conditions at birth explained very little among‐year variation in early‐life growth rates (i.e., weak cohort effects) and that within‐year variation in early‐life growth rates was more likely caused by strong individual differences.

In the last few decades, populations of wild boars have been spreading throughout Europe. This expansion is problematic because of conflicts with human activities and/or sanitary reasons. Thus, the need to develop tools to understand wild boar populations’ increase as well as to manage efficiently their populations appears crucial but challenging. As ancestor of the domestic pig, one of the main issues associated with this population spread is the possible hybridization between the two subspecies which complicates the identification of pure individuals. In this study, we propose a set of 20 single nucleotide polymorphism (SNP) markers, allocated in two 10-plexes, selected in order to identify the two subspecies. The multiplexes were validated on reference populations of wild boars and domestic pigs in order to assess their polymorphism and their discriminative power. The selected markers allowed for an exacerbated differentiation between the wild boars and the domestic pigs compared to other studies and thus allows for an efficient identification of species. The low size of this panel of 20 SNP markers makes it a tool compatible with economic constraints associated with wildlife management compared to tools based on chips.

Wild boar populations have increased dramatically over the last decades throughout Europe and in France in particular. While hunting is considered the most efficient way to control game populations, many local conflicts persist after the hunting period due to remaining high densities of wild boar despite the large number of animals culled every year. Therefore, increasing the efficiency of hunting is a timely issue. Herein, we assessed how hunting effort can be measured, and we determined whether the hunting effort carried out by hunters explains the observed hunting pressure. We measured the characteristics and results of all hunts that occurred in the experimental forest of Châteauvillain-Arc-en-Barrois (Northeastern France), and we modelled the number of animals culled as a function of the hunting effort, measured by the number of beaters, hunters, and dogs, as well as the size of the hunting area. We also accounted for variables suspected to affect the hunting efficiency achieved with a given effort, such as time of day (AM/PM), the month during which hunting occurred. We found that more posted hunters, larger hunted areas, and hunts carried out early in the season, i.e. before February, increased the number of culled animals. Our model can be used by wildlife managers to adjust hunting effort in order to reach the hunting pressure expected to meet management objectives.

Many populations are affected by hunting or fishing. Models designed to assess the sustainability of harvest management require accurate estimates of demographic parameters (e.g. survival, reproduction) hardly estimable with limited data collected on exploited populations. The joint analysis of different data sources with integrated population models (IPM) is an optimal framework to obtain reliable estimates for parameters usually difficult to estimate, while accounting for imperfect detection and observation error. The IPM built so far for exploited populations have integrated count-based surveys and catch-at-age data into ageclass structured population models. But the age of harvested individuals is difficult to assess and often not recorded, and population counts are often not performed on a regular basis, limiting their use for the monitoring of exploited populations.Here, we propose an IPM that makes efficient use of data commonly collected in exploited marine and terrestrial populations of vertebrates. As individual measures of body mass at both capture and death are often collected in fish and terrestrial game species, our model integrates capture-mark-recapture-recovery data and data collected at death into a body mass-structured population model. It allows the observed number of individuals harvested to be compared with the expected number and provides accurate estimates of demographic parameters.We illustrate the usefulness of this IPM using an emblematic game species distributed worldwide, the wild boarSus scrofa, as a case study. For this species that has increased in distribution and abundance over the last decades, the model provides accurate and precise annual estimates of key demographic parameters (survival, reproduction, growth) and of population size while accounting for imperfect detection and observation error.To avoid an overexploitation of declining populations or an under-exploitation of increasing populations, it is crucial to gain a good understanding of the dynamics of exploited populations. When managers or conservationists have limited demographic data, the IPM offers a powerful framework to assess population dynamics. Being highly flexible, the approach is broadly applicable to both terrestrial and marine exploited populations for which measures of body mass are commonly recorded and more generally, to all populations suffering from anthropogenic mortality causes.

Mast seeding in temperate oak populations shapes the dynamics of seed consumers and numerous communities. Mast seeding responds positively to warm spring temperatures and is therefore expected to increase under global warming. We investigated the potential effects of changes in oak mast seeding on wild boar population dynamics, a widespread and abundant consumer species. Using long-term monitoring data, we showed that abundant acorn production enhances the proportion of breeding females. With a body-mass-structured population model and a fixed hunting rate of 0.424, we showed that high acorn production over time would lead to an average wild boar population growth rate of 1.197 whereas non-acorn production would lead to a stable population. Finally, using climate projections and a mechanistic model linking weather data to oak reproduction, we predicted that mast seeding frequency might increase over the next century, which would lead to increase in both wild boar population size and the magnitude of its temporal variation. Our study provides rare evidence that some species could greatly benefit from global warming thanks to higher food availability and therefore highlights the importance of investigating the cascading effects of changing weather conditions on the dynamics of wild animal populations to reliably assess the effects of climate change.

Understanding how some species may be able to evolve quickly enough to deal with anthropogenic pressure is of prime interest in evolutionary biology, conservation, and management. Wild boar (Sus scrofa scrofa) populations keep growing all over Europe despite increasing hunting pressure. In wild boar populations subject to male-selective harvesting, the initially described polygynous mating system may switch to a promiscuous/polyandrous one. Such a change in the mating system, where potentially more males sire a litter at one reproductive event, may be associated with the retention of high genetic diversity and an increase of litter size. We tested these hypotheses by estimating the number of sires per litter based on a six-year long monitoring of a wild boar population subject to particularly high harvesting pressure. Our results show a high and stable genetic diversity and high rates of multiple paternity compared to other populations, thus depicting a promiscuous/polyandrous mating system in this population. We also show that litter size is positively linked to the number of sires, suggesting that multiple paternity increases fecundity. We finally discuss that multiple paternity may be one of the factors allowing rapid evolution of this population by maintaining both genetic and phenotypic diversity.

International audience; Herbivorous ungulates are key species in the functioning of terrestrial ecosystems, and their recent demographic and geographic expansion in some temperate regions is likely to influence ecological processes, particularly if we consider plants and the frequency of mobile links among plant populations. In forests, long-distance seed dispersal essentially relies on animal movements. Owing to their ability to cover daily long distances, large herbivorous ungulates are assumed to be important vectors for long-distance seed dispersal (hundreds to thousands of metres). To quantify patterns of seed dispersal, we analysed the paths of roe deer (Capreolus capreolus), red deer (Cervus elaphus) and wild boar (Sus scrofa) which had been equipped with GPS collars. We used seed gut passage times from a recent companion paper (Picard et al., Ecology and Evolution, 2015;5:2621–2632). We combined the animal movements and seed gut passage times to estimate seed dispersal curves for herbivorous ungulates. On average, the forest ungulates might disperse consumed seeds over distances greater than 2 km, during a 48-h path in the forest. Maximal dispersal distance was longer for red deer (3.5 km) and wild boar (3.2 km) than for roe deer (2.0 km). By combining high excretion rates and long dispersal distance, wild boar appears to be potentially an efficient seed disperser. Seed dispersal by common herbivorous ungulates is likely to play a crucial role in the today's context of rapid environmental changes such as global warming and landscape fragmentation. Differences in the distribution of seed gut passage times and seasonal distances covered underline the relevance of cross comparative approaches and the complementarity of herbivorous ungulates in long-distance seed dispersal.

Hunting, including drive hunts, can be considered as a predation process. Although drive hunts are spatially and temporally well defined, the scale at which the hunting risk triggers anti-predator responses of prey remains poorly documented. The present study aims at 1) characterizing the delayed movement responses of female red deer Cervus elaphus to hunters and 2) understanding the main environmental determinants modulating these post-disturbance responses at the individual level. We hypothesized that red deer alter their movement behaviour for several days after a drive hunt by increasing their speed and exploring more space. Then, we predicted that individuals close to the hunting area would exhibit stronger responses (i.e. higher speed and larger home range size) than individuals located further away. We also expected topographic obstacles and vegetation cover to reduce the magnitude of the post-disturbance responses. We used GPS data from a large dataset collected on 91 female red deer in the National Hunting and Wildlife Reserve of La Petite Pierre (France), between 2004 and 2018. Each year, drive hunts are conducted by about 43 hunters and 6 dogs, once a week from October to February. Our results suggest that, over a 72-h period after a drive hunt, female red deer could show two contrasting strategies: 1) flight-type movements (i.e. higher maximum speed) associated with more space explored due to a temporary avoidance of the disturbed area and 2) lower movements to avoid detection and contacts with hunters. These two delayed responses are situation-specific, depending on risk proximity, landscape features, hunting conditions and individual characteristics. This study highlights that ungulate management using a ‘hunting for fear’ approach may be adjusted considering spatial and temporal scales of the effects of drive hunts on red deer movement patterns.

« Dans le cadre de la planification, les services de la DGAL se préparent, aux différents échelons, à la mise en œuvre des mesures de lutte contre la peste porcine africaine (PPA) en identifiant les ressources humaines et matérielles nécessaires. Pour la question du dépeuplement en élevage, il est prévu un prestataire national pour la réalisation des chantiers de dépeuplement de gros effectifs porcins en élevage, le reste des chantiers étant et devant être piloté et organisé par les directions départementales en charge de la protection des populations DDecPP avec des moyens locaux. Pour autant, selon la dynamique de diffusion de la maladie, ces dispositifs pourraients’avérer insuffisants et une option pourrait être de transférer des animaux issus de cheptels infectés pour une mise à mort à l'abattoir. Par ailleurs, une stratégie de dépeuplement préemptive à l'image de ce qui a été mis en place lors de l’épisode influenza aviaire 2016-2017 pourrait être également mise en place et nécessiterait également une mobilisation importante des moyens de dépeuplement.Il s'agirait donc plus précisément d'évaluer le risque de diffusion de la PPA, à la fois par le transport et par la persistance du virus au sein des sites d’abattage, lors de la mobilisation d'abattoirs pour dépeupler des foyers. Cette même question est posée pour du dépeuplementpréemptif d'animaux a priori non suspects en périphérie de foyer. Ces cas de figure pourraient se présenter en cas de dépassement des capacités d'intervention sur site Des questions complémentaires liées aux opérations de dépeuplement d'animaux infectés sontégalement posées.- Parmi les méthodes d’euthanasie individuelle, un dispositif à tige perforante suivi d'un jonchage ou une injection létale peuvent être utilisés pour les porcs ainsi que le tir à balle pour les sangliers. Ces méthodes qui peuvent entraîner des effusions de sang sont-elles denature à engendrer un risque supplémentaire de diffusion du virus ?- Les intervenants dans un foyer sont équipés de protections individuelles (EPI) et passent par un sas sanitaire pour sortir de l'exploitation en vue de limiter les risques de diffusion de la maladie. La procédure recommandée pour l'entrée et la sortie d'un élevage infecté estjointe ; quel serait le délai de carence à respecter entre l'intervention dans un foyer et l'intervention dans un élevage sain afin de prévenir tout risque de diffusion de la maladie ? Le délai historiquement préconisé est de 72h et demande à être documenté. »

During pregnancy, littermates compete to extract maternal resources from the placenta. Unequal extraction of resources leads to developmental differences among offspring and thus within-litter variation in offspring mass. Because competition among littermates can be stronger among half-sibs, multiple paternity may represent an adaptive strategy allowing females to increase within-litter phenotypic variation among offspring when facing variable environments. Wild boar (Sus scrofa) females produce large litters with diversified offspring in terms of body mass. Additionally, multiple paternities within a litter have been observed in this promiscuous species. One can hypothesize that multiple paternity represents the mechanism by which females increase within-litter phenotypic variation. Combining long-term monitoring data with paternity analyses in a wild boar population, we tested whether the increase in the number of fathers within a litter explained the increase in within-litter variation in offspring mass observed in large litters. We showed that heavy females mated earlier during the rut, produced larger litters with a higher number of fathers and more variable fetus mass than lighter females. Within-litter variation of offspring mass increased with gestation stage and litter size, suggesting differential allocation of maternal resource among offspring “in utero.” However, we found only a weak paternal effect on offspring mass and no direct effect of the number of fathers on the within-litter variation in offspring mass. These results indicate that differential maternal allocation to offspring during pregnancy is unlikely related to paternal identity in this species. This is a pre-copyedited, author-produced version of an article accepted for publication in Behavioral Ecology following peer review. The version of record is available online at: https://doi.org/10.1093/beheco/ary056

We estimated wild boar abundance and density using capture–resight methods in the western part of the Canton of Geneva (Switzerland) in the early summer from 2004 to 2006. Ear-tag numbers and transmitter frequencies enabled us to identify individuals during each of the counting sessions. We used resights generated by self-triggered camera traps as recaptures. Program Noremark provided Minta–Mangel and Bowden’s estimators to assess the size of the marked population. The minimum numbers of wild boars belonging to the unmarked population (juveniles and/or piglets) were added to the respective estimates to assess total population size. Over the 3 years, both estimators showed a stable population with a slight diminishing tendency. We used mean home range size determined by telemetry to assess the sampled areas and densities. Mean wild boar population densities calculated were 10.6 individuals/km2 ± 0.8 standard deviation (SD) and 10.0 ind/km2 ± 0.6 SD with both estimators, respectively, and are among the highest reported from Western Europe. Because of the low proportion of marked animals and, to a lesser extent, of technical failures, our estimates showed poor precision, although they displayed similar population trends compared to the culling bag statistics. Reported densities were consistent with the ecological conditions of the study area.

International audience; Masting, or mast-seeding, defined as a synchronized and highly variable seed production from year-to-year within a population of plants, is one of the most common example of pulsed resources in terrestrial ecosystems. In oaks, the dramatic fluctuations of acorn production impact its reproductive success and regeneration, the dynamics of a large diversity of seed consumers that rely on it, and, by cascade effects, the dynamics of the entire forest community. However, reproductive effort is difficult to quantify and there is therefore an urgent need of a reliable assessment of the dynamic of acorn production based on a low-cost, unbiased, and robust tool. One of the most commonly used method, the “visual on-tree” method, is very easy and quick to carry out, but is biased under high seed production or when branches are difficult to see. We here assessed the robustness of an alternative method, the “ground plot” (GP), based on a unique annual ground survey after peak of acorn fall, which has not been tested so far. We compared this method at tree and site levels (10 forests throughout France) with the costly and time-consuming trap acorn collection (TNR) method (used here as a reference method). We show that results from the GP method closely matched with those obtained using the TNR method, which demonstrates the efficiency and robustness of the GP method at both tree and forest site levels. Despite some limitations in specific environmental contexts we review, this GP method offers a powerful tool to quantify acorn production and should be deployed to understand mechanisms underlying oak masting and/or to assess its ecological or economic consequences.

Across Europe, wild boar numbers increased in the 1960s-1970s but stabilised in the 1980s; recent evidence suggests that the numbers and impact of wild boar has grown steadily since the 1980s. As hunting is the main cause of mortality for this species, we reviewed wild boar hunting bags and hunter population trends in 18 European countries from 1982 to 2012. Hunting statistics and numbers of hunters were used as indicators of animal numbers and hunting pressure. The results confirmed that wild boar increased consistently throughout Europe, while the number of hunters remained relatively stable or declined in most countries. We conclude that recreational hunting is insufficient to limit wild boar population growth and that the relative impact of hunting on wild boar mortality had decreased. Other factors, such as mild winters, reforestation, intensification of crop production, supplementary feeding and compensatory population responses of wild boar to hunting pressure might also explain population growth. As populations continue to grow, more human-wild boar conflicts are expected unless this trend is reversed. New interdisciplinary approaches are urgently required to mitigate human-wild boar conflicts, which are otherwise destined to grow further.

Actuarial senescence is widespread in age-structured populations. In growing populations, the progressive decline of Hamiltonian forces of selection with age leads to decreasing survival. As actuarial senescence is overcompensated by a high fertility, actuarial senescence should be more intense in species with high reproductive effort, a theoretical prediction that has not been yet explicitly tested across species. Wild boar (Sus scrofa) females have an unusual life-history strategy among large mammals by associating both early and high reproductive effort with potentially long lifespan. Therefore, wild boar females should show stronger actuarial senescence than similar-sized related mammals. Moreover, being polygynous and much larger than females, males should display higher senescence rates than females. Using a long-term monitoring (18 years) of a wild boar population, we tested these predictions. We provided clear evidence of actuarial senescence in both sexes. Wild boar females had earlier but not stronger actuarial senescence than similar-sized ungulates. Both sexes displayed similar senescence rates. Our study indicates that the timing of senescence, not the rate, is associated with the magnitude of fertility in ungulates. This demonstrates the importance of including the timing of senescence in addition to its rate to understand variation in senescence patterns in wild populations.

In many parts of Europe, wild boar Sus scrofa population increase, and thus, high densities and dispersal into new areas are accompanied by economic problems. Due to many factors like insufficient hunting strategies as well as underestimation of population densities and reproduction rates, harvest rates seem to be insufficient. Thus, we calculated mortality rates of several wild boar populations from 1998 to 2009, to show the efficiency of hunting within several studies distributed over eight European states. For calculating mortality rates, the daily probability of survival of radio telemetrically observed wild boar was analysed according to Mayfield (Wilson Bull 73:255-261, 1961) and with survival analysis in R for three age classes (0, 1, ≥2 years) and both sexes. The mortality rates of wild boar per annum, especially piglets, were comparably low (about 0.5 for piglets and similar for total population). About three third of all observed animals survived at least until the next period of reproduction. Mortality rates differed between some study areas, the sexes and age classes. The sex ratio of the shot piglets equals the sex ratio of captured piglets; there seems to be no sex-biased hunting in this age class, but in an older age. Shooting was the main cause of death; only very few animals died by natural causes, e.g. diseases. The comparative analysis of all studies reflects a low mortality of wild boar in highly productive populations. Our results certified the findings of several studies that predation, natural mortality, and road mortality have only small impact on wild boar populations, whereas especially, nutrition or hunting are mainly decisive. Assuming net reproduction rates of more than 200 % according to literature data, our results indicate that harvest rates are not sufficient at our study sites. In all our studies, mortality rates and, thus, harvest rates are less than the assumed total net reproduction. Especially, the harvest rate of piglets seems to be insufficient. Thus, the population will increase further. High reproduction has to be counteracted by regulating mainly the reproductive animals. For regulating a population, combined and effective hunting methods have to be conducted to harvest at least the net reproduction. Thus, we recommend higher hunting rates of piglets (80 % of the offspring should be harvested) and of adult females. Intensified hunting of piglets by drive hunts and at an early age as well as intensified single hunt on adult females might help regulating wild boar populations.

Agro-ecosystems can experience elevated human-wildlife conflicts, especially crop damage. While game management often aims at reducing number to mitigate conflicts, there is on-going debate about the role of hunting disturbance in promoting game to range over wider areas, thereby potentially exacerbating conflicts. Herein, we hypothesised that landscape configuration and non-lethal disturbance modulate the response to harvest disturbance. We used an information theoretic approach to test the effects of landscape and anthropogenic variables on wild boar ranging patterns across contrasting harvest regimes. We used 164 seasonal home ranges from 95 wild boar (Sus scrofa) radio-tracked over 6 years in the Geneva Basin where two main harvest regimes coexist (day hunt and night cull). Mean seasonal 95% kernel home range size was 4.01 ± 0.20 km2 (SE) and 50% core range size 0.79 ± 0.04 km2, among the smallest recorded in Europe. Range sizes were larger in the day hunt area than in the night cull area, with no seasonal effect. However, when accounting for landscape variables, we demonstrate that these patterns were likely confounded by the underlying landscape configuration, and that landscape variables remain the primary drivers of wild boar ranging patterns in this human-dominated agro-ecosystem with range size best explained by a model including landscape variables only. Therefore, we recommend accounting for landscape configuration and sources of non-lethal disturbance in the design of harvest strategies when the aim is to limit wide-ranging behaviour of wild boar in order to mitigate conflicts.

Summary 1. Harvest models are often built to explore the sustainability of the dynamics of exploited populations and to help evaluate hunting management scenarios. Age-structured models are commonly used for ungulate population dynamics. However, the age of hunted individuals is usually not recorded, and hunting data often only include body weight and sex limiting the usefulness of traditional models. 2. We propose a new modelling approach that fits data collected by hunters to develop management rules when age is not available. Using wild boar Sus scrofa scrofa as a case study, we built a matrix model structured according to sex and body weight whose output can be directly compared with the observed distribution of hunted individuals among sex and body weight classes. 3. In the face of the current wide scale increase in populations of wild boar, the best feasible option to stop or slow down population growth involves targeting the hunting effort to specific sex and body weight classes. The optimal harvest proportion in the target body weight classes is estimated using sensitivity analyses. 4. The number of individuals shot in each sex and body weight class predicted by our model was closely associated with those recorded in the hunting bag. Increasing the hunting pressure on medium-sized females by 14·6% was the best option to limit growth rate to a target of 0·90. 5. Synthesis and applications. We demonstrate that targeting hunting effort to specific body weight classes could reliably control population growth. Our modelling approach can be applied to any game species where group composition, phenotypic traits or coat colour allows hunters to easily identify sex and body weight classes. This offers a promising tool for applying selective hunting to the management of game species.

Exploitation by humans affects the size and structure of populations. This has evolutionary and demographic consequences that have typically being studied independent of one another. We here applied a framework recently developed applying quantitative tools from population ecology and selection gradient analysis to quantify the selection on a quantitative trait—birth date— through its association with multiple fitness components. From the long-term monitoring (22 years) of a wild boar (Sus scrofa scrofa) population subject to markedly increasing hunting pressure, we found that birth dates have advanced by up to 12 days throughout the study period. During the period of low hunting pressure, there was no detectable selection. However, during the period of high hunting pressure, the selection gradient linking breeding probability in the first year of life to birth date was negative, supporting current life-history theory predicting selection for early births to reproduce within the first year of life with increasing adult mortality.

Summary 1. Demographic tactics within animal populations are shaped by selective pressures. Exploitation exerts additional pressures so that differing demographic tactics might be expected among populations with differences in levels of exploitation. Yet little has been done so far to assess the possible consequences of exploitation on the demographic tactics of mammals, even though such information could influence the choice of effective management strategies. 2. Compared with similar-sized ungulate species, wild boar Sus scrofa has high reproductive capabilities, which complicates population management. Using a perturbation analysis, we investigated how population growth rates (λ) and critical life-history stages differed between two wild boar populations monitored for several years, one of which was heavily harvested and the other lightly harvested. 3. Asymptotic λ was 1·242 in the lightly hunted population and 1·115 in the heavily hunted population, while the ratio between the elasticity of adult survival and juvenile survival was 2·63 and 1·27, respectively. A comparative analysis including 21 other ungulate species showed that the elasticity ratio in the heavily hunted population was the lowest ever observed. 4. Compared with expected generation times of similar-sized ungulates (more than 6 years), wild boar has a fast life-history speed, especially when facing high hunting pressure. This is well illustrated by our results, where generation times were 3·6 years in the lightly hunted population and only 2·3 years in the heavily hunted population. High human-induced mortality combined with non-limiting food resources accounted for the accelerated life history of the hunted population because of earlier reproduction. 5. Synthesis and applications. For wild boar, we show that when a population is facing a high hunting pressure, increasing the mortality in only one age-class (e.g. adults or juveniles) may not allow managers to limit population growth. We suggest that simulations of management strategies based on context-specific demographic models are useful for selecting interventions for population control. This type of approach allows the assessment of population response to exploitation by considering a range of plausible scenarios, improving the chance of selecting appropriate management actions.

Spatial variation of the ‘predation risk’ due to human activities or distribution may increase the sexual difference in habitat selection. Indeed, females with offspring are usually more risk adverse than males. Based on a long-term wild boar study, we analysed the diurnal distribution of female and male wild boar before, during and after the hunting period. Hunting, food and foliation were investigated as factors affecting patterns of forest parcel selection. As expected, dense vegetative covers were selected during resting periods, but wild boar decreased this pattern of habitat selection in response to hunting disturbance. Moreover, the habitat selection of wild boar did not fit with the variation of food availability (presence or absence of mast) and the vegetation cycle. As expected, sows responded more to the hunting disturbance than males, leading to a more pronounced sexual difference during the riskier season. The unexpected decrease of bush use may be explained either by the increased hunting effort in this habitat or by the increased movements between resting sites due to disturbance, leading to a more random habitat selection pattern. The observed difference between sexes could result from a higher response of females with offspring to hunting, leading to an increased frequentation of secondary habitats, whereas males can tolerate more risks and remain hidden in thicket plots. Our results highlight how hunting disturbance can lead game species to change their patterns of refuge habitat selection and may affect the habitat segregation between the sexes.

Understanding how some species may be able to evolve quickly enough to deal with anthropogenic pressure is of prime interest in evolutionary biology, conservation, and management. Wild boar (Sus scrofa scrofa) populations keep growing all over Europe despite increasing hunting pressure. In wild boar populations subject to male-selective harvesting, the initially described polygynous mating system may switch to a promiscuous/polyandrous one. Such a change in the mating system, where potentially more males sire a litter at one reproductive event, may be associated with the retention of high genetic diversity and an increase of litter size. We tested these hypotheses by estimating the number of sires per litter based on a six-year long monitoring of a wild boar population subject to particularly high harvesting pressure. Our results show a high and stable genetic diversity and high rates of multiple paternity compared to other populations, thus depicting a promiscuous/polyandrous mating system in this population. We also show that litter size is positively linked to the number of sires, suggesting that multiple paternity increases fecundity. We finally discuss that multiple paternity may be one of the factors allowing rapid evolution of this population by maintaining both genetic and phenotypic diversity.

Mother–daughter postweaning associations in wild boars (Sus scrofa L., 1758) were investigated using 12 years' data from a wild population in Champagne, France. In the wild boar, a polygynous ungulate species, females (i) can reproduce as soon as they are yearlings and (ii) generally have large litters, in contrast to many other ungulate species. It is generally thought that their social organization is centered around groups of adult females and their offspring, but genealogical relationships in female groups have never been studied. Hence this species is suitable for testing the hypothesis of a matrilineal social organization. We studied the occurrence and strength of mother–daughter associations before and after the first potential breeding of yearling females, using a total of 85 individuals. Seasonal fluctuations in associations were observed, but after weaning, daughters generally remained with the mother. When leaving their natal group, yearling females formed new kin groups with sisters. Two important factors involved in the postweaning associations were adult and yearling reproductive participation and maternal age. The present study constitutes the first clear demonstration that family groups, with overlapping generations of females, represent the typical social organization in a forest wild boar population.

International audience; Most mammalian populations suffer from natural or human-induced disturbances; populations are no longer at the equilibrium (i.e., at stable [st]age distribution) and exhibit transient dynamics. From a literature survey, we studied patterns of transient dynamics for mammalian species spanning a large range of life-history tactics and population growth rates. For each population, we built an age-structured matrix and calculated six metrics of transient dynamics. After controlling for possible confounding effects of the phylogenetic relatedness among species using a phylogenetic principal component analysis and phylogenetic generalized least squares models, we found that short-term demographic responses of mammalian populations to disturbance are shaped by generation time and growth rate. Species with a slow pace of life (i.e., species with a late maturity, a low fecundity, and a long life span) displayed decreases in population size after a disturbance, whereas fast-living species increased in population size. The magnitude of short-term variation in population size increased with asymptotic population growth, being buffered in slow-growing species (i.e., species with a low population growth rate) but large in fast-growing species. By demonstrating direct links between transient dynamics, life history (generation time), and ecology (demographic regime), our comparative analysis of transient dynamics clearly improves our understanding of population dynamics in variable environments and has clear implications for future studies of the interplay between evolutionary and ecological dynamics. As most populations in the wild are not at equilibrium, we recommend that analyses of transient dynamics be performed when studying population dynamics in variable environments.

Across Europe, wild boar numbers increased in the 1960s-1970s but stabilised in the 1980s; recent evidence suggests that the numbers and impact of wild boar has grown steadily since the 1980s. As hunting is the main cause of mortality for this species, we reviewed wild boar hunting bags and hunter population trends in 18 European countries from 1982 to 2012. Hunting statistics and numbers of hunters were used as indicators of animal numbers and hunting pressure. The results confirmed that wild boar increased consistently throughout Europe, while the number of hunters remained relatively stable or declined in most countries. We conclude that recreational hunting is insufficient to limit wild boar population growth and that the relative impact of hunting on wild boar mortality had decreased. Other factors, such as mild winters, reforestation, intensification of crop production, supplementary feeding and compensatory population responses of wild boar to hunting pressure might also explain population growth. As populations continue to grow, more human-wild boar conflicts are expected unless this trend is reversed. New interdisciplinary approaches are urgently required to mitigate human-wild boar conflicts, which are otherwise destined to grow further.

Across Europe wild boar increased in the ‘60s-‘70s but stabilised in the ‘80s ; however, their impact grew steadily. As hunting is the main cause of mortality for wild boar, this review described wild boar and hunter population trends in Europe in the last three decades. Hunting statistics and numbers of hunters in 18 countries were used as indicators of wild boar numbers and hunting pressure in 1982-2012. The results showed that wild boar are increasing consistently throughout Europe whilst the number of hunters are relatively stable or even declining. We concluded that the generalised growth in number wild boar harvested is not related to increasing hunting and that wild boar numbers are indeed growing. As the main cause of wild boar mortality, hunting, is decreasing, more human-wild boar conflicts are expected unless this trend is reversed. The results suggested that recreational hunting is not enough to limit wild boar population growth. Other factors, such as mild winters, reforestation, intensification in crop production, supplementary feeding and compensatory population responses of wild boar to hunting pressure might also explain population growth. New interdisciplinary approaches are urgently required to mitigate human-wild boar conflicts that are otherwise destined to grow further.

Summary In highly variable environments, the optimal reproductive tactics of iteroparous organisms should minimize variance in yearly reproductive success to maximize the long-term average reproductive success. To minimize among-year variation in reproductive success, individuals can either minimize the variance in the number of offspring produced at each reproductive attempt (classical bet-hedging) or maximize the phenotypic diversity of offspring produced within or among reproductive attempts (coin-flipping). From a long-term detailed study of an intensively exploited population facing a highly unpredictable environment, we identify a continuum of reproductive tactics in wild boar females depending on their body mass. At one end, light females adjusted litter size to their body mass and produced highly similar-sized offspring within a litter. These females fitted the hypothesis of individual optimization commonly reported in warm-blooded species, which involves both an optimal mass and an optimal number of offspring for a given mother. At the other end of the continuum, heavy females produced litters of variable size including a mixture of heavy and light offspring within litters. Prolific heavy wild boar females diversify the phenotype of their offspring, providing a first evidence for coin-flipping in a warm-blooded species.

No-take reserves are sometimes implemented for sustainable population harvesting because they offer opportunities for animals to spatially avoid harvesters, whereas harvesters can benefit in return from the reserve spillover. Here, we used the framework of predator-prey spatial games to understand how protected areas shape spatial interactions between harvesters and target species and determine animal mortality. In these spatial games, the "predator" searches for "prey" and matches their habitat use, unless it meets spatial constraints offering the opportunity for prey to avoid the mortality source. However, such prey refuges could attract predators in the surroundings, which questions the potential benefits for prey. We located, in the Geneva Basin (France), hunting dogs and wild boar Sus scrofa L. during hunting seasons with global positioning systems and very-high-frequency collars. We quantified how the proximity of the reserve shaped the matching between both habitat uses using multivariate analyses and linked these patterns to animals' mortality with a Cox regression analysis. Results showed that habitat uses by both protagonists disassociated only when hunters were spatially constrained by the reserve. In response, hunters increased hunting efforts near the reserve boundary, which induced a higher risk exposure for animals settled over the reserve. The mortality of adult wild boar decreased near the reserve as the mismatch between both habitat uses increased. However the opposite pattern was determined for younger individuals that suffered from the high level of hunting close to the reserve. The predator-prey analogy was an accurate prediction of how the protected area modified spatial relationships between harvesters and target species. Prey-searching strategies adopted by hunters around reserves strongly impacted animal mortality and the efficiency of the protected area for this harvested species. Increasing reserve sizes and/or implementing buffer areas with harvesting limitations can dampen this edge effect and helps harvesters to benefit durably from source populations of reserves. Predator-prey spatial games therefore provide a powerful theoretical background for understanding wildlife-harvester spatial interactions and developing substantial application for sustainable harvesting.

Exploitation by humans affects the size and structure of populations. This has evolutionary and demographic consequences that have typically being studied independent of one another. We here applied a framework recently developed applying quantitative tools from population ecology and selection gradient analysis to quantify the selection on a quantitative trait-birth date-through its association with multiple fitness components. From the long-term monitoring (22 years) of a wild boar (Sus scrofa scrofa) population subject to markedly increasing hunting pressure, we found that birth dates have advanced by up to 12 days throughout the study period. During the period of low hunting pressure, there was no detectable selection. However, during the period of high hunting pressure, the selection gradient linking breeding probability in the first year of life to birth date was negative, supporting current life-history theory predicting selection for early births to reproduce within the first year of life with increasing adult mortality.

International audience; Whether different sources of mortality are additive, compensatory, or depensatory is a key question in population biology. A way to test for additivity is to calculate the correlation between cause-specific mortality rates obtained from marked animals. However, existing methods to estimate this correlation raise several methodological issues. One difficulty is the existence of an intrinsic bias in the correlation parameter. Although this bias can be formally expressed, it requires knowledge about natural survival without any competing mortality source, which is difficult to assess in most cases. Another difficulty lies in estimating the true process correlation while properly accounting for sampling variation. Using a Bayesian approach, we developed a state–space model to assess the correlation between two competing sources of mortality. By distinguishing the mortality process from its observation through dead recoveries and live recaptures, we estimated the process correlation. To correct for the intrinsic bias, we incorporated experts’ opinions on natural survival. We illustrated our approach using data on a hunted population of wild boars. Mortalities were not additive and natural mortality increased with hunting mortality likely as a consequence of non-controlled mortality by crippling loss. Our method opens perspectives for wildlife management and for the conservation of endangered species.

Wild boars Sus scrofa have a social organization based on female groups that can include several generations of adults and offspring, and are thus likely matrilineal. However, little is known about the degree of relatedness between animals living in such groups or occupying the same core area of spatial activity. Also, polygynous male mating combined with matrilineal female groups can have strong influences on the genetic structure of populations. We used microsatellite genotyping combined with behavioral data to investigate the fine-scale population genetic structure and the mating system of wild boars in a multi-year study at Châteauvillain-Arc-en-Barrois (France). According to spatial genetic autocorrelation, females in spatial proximity were significantly inter-related. However, we found that numerous males contributed to the next generation, even within the same social group. Based on our genetic data and behavioral observations, wild boars in this population appear to have a low level of polygyny associated with matrilineal female groups, and infrequent multiple paternity. Mortality due to hunting may facilitate the breakup of what historically has been a more predominantly polygynous mating system, and likely accelerates the turnover of adults within the matrilineal groups.

Predators generate a “landscape of fear” within which prey can minimize the risk of predation by selecting low-risk areas. Depending on the spatial structure of this “landscape”, i.e., whether it is coarse- or fine-grained, prey may respond to increased risk by shifting their home ranges or by fine-scale redistributions within these ranges, respectively. We studied how wild boar ( Sus scrofa L., 1758) responded to temporal changes in risk in hunted areas (risky habitat) surrounding a nature reserve (refuge habitat). Animals with home ranges “in contact” with the reserve during the low-risk season were the only ones to shift toward the refuge when the risk increased. These shifts occurred at two temporal scales in response to the increased risk during the daytime and during the hunting season. Whereas animals not influenced by the reserve found food and shelter in forest during the hunting season, shifts to the refuge area were detrimental to the rather scarce forest areas in the reserve. This confirms that spatiotemporal changes in risk are major drivers of animal distribution when predation strongly limits their fitness. Their response is, however, scale-dependent and reflects at the individual level the perceived structure of their “landscape of fear”.

We assessed age-specific natural mortality (i.e., excluding hunting mortality) and hunting mortality of 1,175 male and 1,076 female wild boar (Sus scrofa) from Châteauvillain-Arc en Barrois (eastern France), using a 22-year dataset (1982–2004) and mark–recapture–recovery methods. Overall yearly mortality was >50% for all sex and age-classes. Low survival was mostly due to high hunting mortality; a wild boar had a >40% of chance of being harvested annually, and this risk was as high as 70% for adult males. Natural mortality rates of wild boar were similar for males and females (approx. 0.15). These rates were comparable to rates typical of male ungulates but high for female ungulates. Wild boar survival did not vary across sex and age-classes. Despite high hunting mortality, we did not detect evidence of compensatory mortality. Whereas natural mortality for males was constant over time, female mortality varied annually, independent of fluctuations in mast availability. Female wild boar survival pa...

For species in which reproductive success is more variable in one sex than the other, the Trivers and Willard model (TWM) predicts that females are able to adjust their offspring sex ratio. High-quality mothers should provide greater investment to one sex than the other. Previous tests of the TWM have been inconsistent, and whether the TWM applies to species with several offspring per litter is unclear due to possible trade-offs between size, number, and sex of the offspring. Williams’ model (WM) accounts for confounding effects of these trade-offs on sex ratio variation. Lastly, the ‘‘extrinsic modification hypothesis’’ predicts changes in offspring sex ratio in relation to climatic conditions and population density. Using wild boar as a model, we tested 1) whether the WM fitted observed sex ratio variation and 2) whether sex ratio variations were related to maternal attributes (test of the TWM) and/or to resource availability (test of the extrinsic modification hypothesis). Females adjusted their litter size rather than their litter composition, so that the WM was not supported. Likewise, changes in resource availability did not influence the fetal sex ratio, so that the extrinsic modification hypothesis was not supported. The fetal sex ratio was negatively related to increasing litter size, providing some support for the TWM. Sex ratio was male biased for litter sizes up to 6 and then became female biased in larger litters. Our results provide the first case study showing marked changes in sex ratio in relation to litter size in a large mammal. Key words: litter size, primary sex ratio, sex ratio variation, Sus scrofa scrofa, Trivers and Willard model, Williams model. [Behav Ecol]

To maximize long-term average reproductive success, individuals can diversify the phenotypes of offspring produced within a reproductive event by displaying the ‘coin-flipping’ tactic. Wild boar (Sus scrofa scrofa) females have been reported to adopt this tactic. However, whether the magnitude of developmental plasticity within a litter depends on stochasticity in food resources has not been yet investigated. From long-term monitoring, we found that juvenile females produced similar-sized fetuses within a litter independent of food availability. By contrast, adult females adjusted their relative allocation to littermates to the amount of food resources, by providing a similar allocation to all littermates in years of poor food resources but producing highly diversified offspring phenotypes within a litter in years of abundant food resources. By minimizing sibling rivalry, such a plastic reproductive tactic allows adult wild boar females to maximize the number of littermates for a given breeding event.