Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale.

Understanding the distribution of biodiversity across the Earth is one of the most challenging questions in biology. Much research has been directed at explaining the species latitudinal pattern showing that communities are richer in tropical areas; however, despite decades of research, a general consensus has not yet emerged. In addition, global biodiversity patterns are being rapidly altered by human activities. Here, we aim to describe large-scale patterns of species richness and diversity in terrestrial vertebrate scavenger (carrion-consuming) assemblages, which provide key ecosystem functions and services. We used a worldwide dataset comprising 43 sites, where vertebrate scavenger assemblages were identified using 2,485 carcasses monitored between 1991 and 2018. First, we evaluated how scavenger richness (number of species) and diversity (Shannon diversity index) varied among seasons (cold vs. warm, wet vs. dry). Then, we studied the potential effects of human impact and a set of macroecological variables related to climatic conditions on the scavenger assemblages. Vertebrate scavenger richness ranged from species-poor to species rich assemblages (4-30 species). Both scavenger richness and diversity also showed some seasonal variation. However, in general, climatic variables did not drive latitudinal patterns, as scavenger richness and diversity were not affected by temperature or rainfall. Rainfall seasonality slightly increased the number of species in the community, but its effect was weak. Instead, the human impact index included in our study was the main predictor of scavenger richness. Scavenger assemblages in highly human-impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. Our results highlight the effect of human impact at a global scale. As species-rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human-dominated landscapes in the Anthropocene.

Does despotic behavior or food search explain the occurrence of problem brown bears in Europe?

Bears foraging near human developments are often presumed to be responding to food shortage, but this explanation ignores social factors, in particular despotism in bears. We analyzed the age distribution and body condition index (BCI) of shot brown bears in relation to densities of bears and people, and whether the shot bears were killed by managers (i.e., problem bears; n = 149), in self-defense (n = 51), or were hunter-killed nonproblem bears (n = 1,896) during 1990-2010. We compared patterns between areas with (Slovenia) and without supplemental feeding (Sweden) of bears relative to 2 hypotheses. The food-search/food-competition hypothesis predicts that problem bears should have a higher BCI (e.g., exploiting easily accessible and/or nutritious human-derived foods) or lower BCI (e.g., because of food shortage) than nonproblem bears, that BCI and human density should have a positive correlation, and problem bear occurrence and seasonal mean BCI of nonproblem bears should have a negative correlation (i.e., more problem bears during years of low food availability). Food competition among bears additionally predicts an inverse relationship between BCI and bear density. The safety-search/naivety hypothesis (i.e., avoiding other bears or lack of human experience) predicts no relationship between BCI and human density, provided no dietary differences due to spatiotemporal habitat use among bears, no relationship between problem bear occurrence and seasonal mean BCI of nonproblem bears, and does not necessarily predict a difference between BCI for problem/nonproblem bears. If food competition or predation avoidance explained bear occurrence near settlements, we predicted younger problem than nonproblem bears and a negative correlation between age and human density. However, if only food search explained bear occurrence near settlements, we predicted no relation between age and problem or nonproblem bear status, or between age and human density. We found no difference in BCI or its variability between problem and nonproblem bears, no relation between BCI and human density, and no correlation between numbers of problem bears shot and seasonal mean BCI for either country. The peak of shot problem bears occurred from April to June in Slovenia and in June in Sweden (i.e., during the mating period when most intraspecific predation occurs and before fall hyperphagia). Problem bears were younger than nonproblem bears, and both problem and nonproblem bears were younger in areas of higher human density. These age differences, in combination with similarities in BCI between problem and nonproblem bears and lack of correlation between BCI and human density, suggested safety-search and naïve dispersal to be the primary mechanisms responsible for bear occurrence near settlements. Younger bears are less competitive, more vulnerable to intraspecific predation, and lack human experience, compared to adults. Body condition was inversely related to the bear density index in Sweden, whereas we found no correlation in Slovenia, suggesting that supplemental feeding may have reduced food competition, in combination with high bear harvest rates. Bears shot in self-defense were older and their BCI did not differ from that of nonproblem bears. Reasons other than food shortage apparently explained why most bears were involved in encounters with people or viewed as problematic near settlements in our study.

Impact of Long-Term Floods on Spatial Dynamics of Myrmica scabrinodis, a Host Ant of a Highly Threatened Scarce Large Blue (Phengaris teleius).

Extensively used wet meadows with high species diversity are under threat in Europe by anthropogenic pressure. The increasing frequency of prolonged flooding is emerging as an additional threat to this fragile environment. In our study, we investigated how prolonged flooding affects the spatial distribution and temporal dynamics (through mortality and recolonization process) of the host ant species Myrmica scabrinodis, which is essential for the survival of the endangered Scarce large blue (Phengaris teleius). The study was conducted in the flood-prone Ljubljansko barje plain situated on the southern edge of the species' global range. Prolonged flooding in the study area, possibly affecting the past and current distribution of the host ant M. scabrinodis, was recorded in 2010, 2013, and 2017. In 2020, we set 160 ant traps to estimate the distribution of host ants in a system of meadows covering the entire gradient of flood history. Results indicate that M. scabrinodis survives the flooding for up to three days, starting to disappear if flooding persists longer. After the flooding recedes, ants gradually recolonize empty habitats from the surrounding upland refugia. Our spatial analyses predict that the average recolonization speed was about 29 m per year and that in a year, ants compensate for the mortality effects of 1.8 days of flooding by recolonization in a year. These results show that flooding should be considered as an additional (in some areas, a major) threat to the endangered P. teleius through its deleterious effects on the host ant species.

Monitoring the effective population size of a brown bear (Ursus arctos) population using new single-sample approaches.

The effective population size (N(e) ) could be the ideal parameter for monitoring populations of conservation concern as it conveniently summarizes both the evolutionary potential of the population and its sensitivity to genetic stochasticity. However, tracing its change through time is difficult in natural populations. We applied four new methods for estimating N(e) from a single sample of genotypes to trace temporal change in N(e) for bears in the Northern Dinaric Mountains. We genotyped 510 bears using 20 microsatellite loci and determined their age. The samples were organized into cohorts with regard to the year when the animals were born and yearly samples with age categories for every year when they were alive. We used the Estimator by Parentage Assignment (EPA) to directly estimate both N(e) and generation interval for each yearly sample. For cohorts, we estimated the effective number of breeders (N(b) ) using linkage disequilibrium, sibship assignment and approximate Bayesian computation methods and extrapolated these estimates to N(e) using the generation interval. The N(e) estimate by EPA is 276 (183-350 95% CI), meeting the inbreeding-avoidance criterion of N(e) > 50 but short of the long-term minimum viable population goal of N(e) > 500. The results obtained by the other methods are highly consistent with this result, and all indicate a rapid increase in N(e) probably in the late 1990s and early 2000s. The new single-sample approaches to the estimation of N(e) provide efficient means for including N(e) in monitoring frameworks and will be of great importance for future management and conservation.

Passive Disease Surveillance of Alpine Chamois (Rupicapra r. rupicapra) in Slovenia between 2000 and 2020.

In this paper, we provide an overview of the causes of death of Alpine chamois (Rupicapra r. rupicapra) diagnosed in the national passive health surveillance of chamois in Slovenia. From 2000 to 2020, 284 free-ranging chamois provided by hunters were necropsied at the Veterinary Faculty, University of Ljubljana, Slovenia. Depending on the results of complete necropsy, histopathological, bacteriological, parasitological, and virological examinations, a descriptive data analysis was performed. The most common causes of death in chamois were infectious diseases (82.2%), followed by non-infectious diseases (11.8%). Of all the causes of death, parasitic infections accounted for 70.3%, trauma for 9.7%, and bacterial infections for 9.3% of all cases. Less common diseases were viral infections, neoplasms, winter starvation, and metabolic disorders.

Twenty Years of Passive Disease Surveillance of Roe Deer (Capreolus capreolus) in Slovenia.

In this paper, we provide an overview of the causes of death of roe deer (Capreolus capreolus) diagnosed within the national passive health surveillance of roe deer in Slovenia. From 2000 to 2019, postmortem examinations of 510 free-ranging roe deer provided by hunters were conducted at the Veterinary Faculty, Slovenia. A comprehensive necropsy was performed. According to the results of the necropsy, the samples were subjected to microscopic, histopathological, bacteriological, parasitological, or virological examination. The most frequent causes of death in roe deer were infectious diseases (67%), followed by noninfectious diseases (28%). Of all deaths, parasitic infections represented 48%, bacterial infections 14.8%, trauma 12.5%, and metabolic disorders 9.8%. Less frequent causes were diseases like neoplasia and mycotic infections, winter starvation, hernias, and lightning strike. This study covered an estimated 1% of the total disease-related mortality of roe deer in Slovenia. Comparisons of sex/age structure indicated that hunters did not provide random samples (e.g., young males were disproportionately represented). Therefore, such monitoring does not ensure an unbiased assessment of the significance of the individual disease for the mortality of the population; however, it can provide credible evidence of whether or not a particular disease is present in a population. We show that no identified disease in roe deer in Slovenia can be considered a significant health threat to roe deer, other wildlife species, or humans.

Exposure of Free-Ranging Wild Animals to Zoonotic Leptospira interrogans Sensu Stricto in Slovenia.

A total of 249 serum samples from 13 wild animal species namely fallow deer (Dama dama, n = 1), roe deer (Capreolus capreolus, n = 80), red deer (Cervus elaphus, n = 22), chamois (Rupicapra rupicapra, n = 21), mouflon (Ovis musimon, n = 4), brown hare (Lepus europaeus, n = 2), nutria (Myocastor coypus, n = 1), red fox (Vulpes vulpes, n = 97), stone marten (Martes foina, n = 12), European badger (Meles meles, n = 2), golden jackal (Canis aureus, n = 2) Eurasian lynx (Lynx lynx, n = 2) and grey wolf (Canis lupus, n = 3) were analysed for the presence of antibodies against Leptospira interrogans sensu stricto. Serum samples were examined via the microscopic agglutination test for the presence of specific antibodies against Leptospira serovars Icterohaemorrhagiae, Bratislava, Pomona, Grippotyphosa, Hardjo, Sejroe, Australis, Autumnalis, Canicola, Saxkoebing and Tarassovi. Antibodies to at least one of the pathogenic serovars were detected in 77 (30.9%; CI = 25-37%) sera. The proportion of positive samples varied intraspecifically and was the biggest in large carnivores (lynx, wolf and jackal; 86%), followed by mezzo predators: stone marten (67%) and red fox (34%), and large herbivores: red deer (32%), roe deer (25%), alpine chamois (10%) and mouflon (0%). Out of the 77 positive samples, 42 samples (53.8%) had positive titres against a single serovar, while 35 (45.4%) samples had positive titres against two or more serovars. The most frequently detected antibodies were those against the serovar Icterohaemorrhagiae. The present study confirmed the presence of multiple pathogenic serovars in wildlife throughout Slovenia. It can be concluded that wild animals are reservoirs for at least some of the leptospiral serovars and are a potential source of leptospirosis for other wild and domestic animals, as well as for humans.

Functional traits driving species role in the structure of terrestrial vertebrate scavenger networks.

Species assemblages often have a non-random nested organization, which in vertebrate scavenger (carrion-consuming) assemblages is thought to be driven by facilitation in competitive environments. However, not all scavenger species play the same role in maintaining assemblage structure, as some species are obligate scavengers (i.e., vultures) and others are facultative, scavenging opportunistically. We used a database with 177 vertebrate scavenger species from 53 assemblages in 22 countries across five continents to identify which functional traits of scavenger species are key to maintaining the scavenging network structure. We used network analyses to relate ten traits hypothesized to affect assemblage structure with the "role" of each species in the scavenging assemblage in which it appeared. We characterized the role of a species in terms of both the proportion of monitored carcasses on which that species scavenged, or scavenging breadth (i.e., the species "normalized degree"), and the role of that species in the nested structure of the assemblage (i.e., the species "paired nested degree"), therefore identifying possible facilitative interactions among species. We found that species with high olfactory acuity, social foragers, and obligate scavengers had the widest scavenging breadth. We also found that social foragers had a large paired nested degree in scavenger assemblages, probably because their presence is easier to detect by other species to signal carcass occurrence. Our study highlights differences in the functional roles of scavenger species and can be used to identify key species for targeted conservation to maintain the ecological function of scavenger assemblages.

Barbary sheep tissues as bioindicators of radionuclide and stabile element contamination in Croatia: exposure assessment for consumers.

Muscle, liver and kidney of 21 Barbary sheep (Ammotragus lervia) from Mosor Mountain, Croatia, were sampled to quantify the activity of caesium and potassium radionuclides and five toxic and ten essential stabile elements in order to establish reference values for this species and to evaluate the potential of Barbary sheep tissues to reflect environmental pollution. We also assessed seasonal diet (botanical composition and dry matter content) of Barbary sheep based on analyses of a rumen content of culled animals. None of the 19 plant species (mostly grasses) identified as part of the Barbary sheep diet is known as a stabile element or radionuclide hyperaccumulator. Measured levels reflected low environmental pollution with arsenic, cadmium, mercury and lead, with levels generally less than those reported for wild herbivorous ungulates. Methodological differences (detection limit of elements in muscle) were shown to hamper interpretation and comparison of the Toxic Contamination Index (TCI) values with those published for other species. There was no homeostasis disturbance of trace elements in Barbary sheep, either due to inadequate intake via food or as an adverse effect due to a high toxic metal(loid) burden. Consumption of the muscle and liver of wild Barbary sheep can be considered safe for the health of adult consumers regarding toxic metal(loid)s and radioactive caesium, though the liver should be avoided as a food item in vulnerable population groups due to the possible adverse effects of cadmium and lead. Otherwise, muscle and liver are a rich source of copper, iron, selenium and zinc for consumers and, as such, can benefit the overall dietary intake of essential elements.

Age-related effects of body mass on fertility and litter size in roe deer.

We analysed effects of females' body mass and age on reproductive capacity of European roe deer (Capreolus capreolus) in a large sample set of 1312 females (305 yearlings and 1007 adults), hunted throughout Slovenia, central Europe, in the period 2013-2015. Body mass positively affected probability of ovulation and potential litter size (number of corpora lutea), although its effect was more pronounced in yearlings than in adults. Between age groups, we found clear differences in responses of both reproductive parameters to body mass which influences primarily reproductive performance of younger, and in particular, lighter individuals: at the same body mass yearlings would at average have smaller litters than adults, and at lower body mass also young to middle-aged adults would have smaller litters than old ones. In addition, while yearlings have to reach a critical threshold body mass to attain reproductive maturity, adult females are fertile (produce ova) even at low body mass. However, at higher body mass also younger individuals shift their efforts into the reproduction, and after reaching an age-specific threshold the body mass does not have any further effects on the reproductive output of roe deer females. Increased reproductive capacity at more advanced age, combined with declining body mass suggests that old does allocate more of their resources in reproduction than in body condition.

Recovery of large carnivores in Europe's modern human-dominated landscapes.

The conservation of large carnivores is a formidable challenge for biodiversity conservation. Using a data set on the past and current status of brown bears (Ursus arctos), Eurasian lynx (Lynx lynx), gray wolves (Canis lupus), and wolverines (Gulo gulo) in European countries, we show that roughly one-third of mainland Europe hosts at least one large carnivore species, with stable or increasing abundance in most cases in 21st-century records. The reasons for this overall conservation success include protective legislation, supportive public opinion, and a variety of practices making coexistence between large carnivores and people possible. The European situation reveals that large carnivores and people can share the same landscape.