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Abstract Background Animal personality has emerged as a key concept in behavioral ecology. While many studies have demonstrated the influence of personality traits on behavioral patterns, its quantification, especially in wild animal populations, remains a challenge. Only a few studies have established a link between personality and recurring movements within home ranges, although these small-scale movements are of key importance for identifying ecological interactions and forming individual niches. In this regard, differences in space use among individuals might reflect different exploration styles between behavioral types along the shy-bold continuum. Methods We assessed among-individual differences in behavior in the European hare (Lepus europaeus), a characteristic mammalian herbivore in agricultural landscapes using a standardized box emergence test for captive and wild hares. We determined an individuals’ degree of boldness by measuring the latencies of behavioral responses in repeated emergence tests in captivity. During capture events of wild hares, we conducted a single emergence test and recorded behavioral responses proven to be stable over time in captive hares. Applying repeated novel environment tests in a near-natural enclosure, we further quantified aspects of exploration and activity in captive hares. Finally, we investigated whether and how this among-individual behavioral variation is related to general activity and space use in a wild hare population. Wild and captive hares were treated similarly and GPS-collared with internal accelerometers prior to release to the wild or the outdoor enclosure, respectively. General activity was quantified as overall dynamic body acceleration (ODBA) obtained from accelerometers. Finally, we tested whether boldness explained variation in (i) ODBA in both settings and (ii) variation in home ranges and core areas across different time scales of GPS-collared hares in a wild population. Results We found three behavioral responses to be consistent over time in captive hares. ODBA was positively related to boldness (i.e., short latencies to make first contact with the new environment) in both captive and wild hares. Space use in wild hares also varied with boldness, with shy individuals having smaller core areas and larger home ranges than bold conspecifics (yet in some of the parameter space, this association was just marginally significant). Conclusions Against our prediction, shy individuals occupied relatively large home ranges but with small core areas. We suggest that this space use pattern is due to them avoiding risky, and energy-demanding competition for valuable resources. Carefully validated, activity measurements (ODBA) from accelerometers provide a valuable tool to quantify aspects of animal personality along the shy-bold continuum remotely. Without directly observing—and possibly disturbing—focal individuals, this approach allows measuring variability in animal personality, especially in species that are difficult to assess with experiments. Considering that accelerometers are often already built into GPS units, we recommend activating them at least during the initial days of tracking to estimate individual variation in general activity and, if possible, match them with a simple novelty experiment. Furthermore, information on individual behavioral types will help to facilitate mechanistic understanding of processes that drive spatial and ecological dynamics in heterogeneous landscapes.

Abstract Although many plants are dispersed by wind and seeds can travel long distances across unsuitable matrix areas, a large proportion relies on co‐evolved zoochorous seed dispersal to connect populations in isolated habitat islands. Particularly in agricultural landscapes, where remaining habitat patches are often very small and highly isolated, mobile linkers as zoochorous seed dispersers are critical for the population dynamics of numerous plant species. However, knowledge about the quali‐ or quantification of such mobile link processes, especially in agricultural landscapes, is still limited. In a controlled feeding experiment, we recorded the seed intake and germination success after complete digestion by the European brown hare (Lepus europaeus) and explored its mobile link potential as an endozoochoric seed disperser. Utilizing a suite of common, rare, and potentially invasive plant species, we disentangled the effects of seed morphological traits on germination success while controlling for phylogenetic relatedness. Further, we measured the landscape connectivity via hares in two contrasting agricultural landscapes (simple: few natural and semi‐natural structures, large fields; complex: high amount of natural and semi‐natural structures, small fields) using GPS‐based movement data. With 34,710 seeds of 44 plant species fed, one of 200 seeds (0.51%) with seedlings of 33 species germinated from feces. Germination after complete digestion was positively related to denser seeds with comparatively small surface area and a relatively slender and elongated shape, suggesting that, for hares, the most critical seed characteristics for successful endozoochorous seed dispersal minimize exposure of the seed to the stomach and the associated digestive system. Furthermore, we could show that a hare's retention time is long enough to interconnect different habitats, especially grasslands and fields. Thus, besides other seed dispersal mechanisms, this most likely allows hares to act as effective mobile linkers contributing to ecosystem stability in times of agricultural intensification, not only in complex but also in simple landscapes.

Abstract Numerous studies have examined whether the primary and/or secondary sex ratio in mammals, including humans, deviates from an equilibrium of 1:1. Although effect size in the sex ratio variation is expected to be low, a large sample size allows the identification of even small deviations from parity. In this study, we investigated whether the sex ratio of roe deer (Capreolus capreolus) offspring at birth approaches parity, using a large data set from roe deer offspring tagged in Baden‐Württemberg (Germany, 1972–2019, N = 12,437). In addition, a systematic re‐analysis of available data on the secondary sex ratios of roe deer was conducted to test whether our finding withstood the accumulation of further data. The null hypothesis that the sex ratio of roe deer (prenatal sex ratio and sex ratio at birth) approaches parity was rejected. Moreover, the secondary sex ratio of roe deer offspring deviated from the male‐biased mean for relatively cold or warm weather conditions during autumn and winter. Our study provides strong evidence for a male‐biased sex ratio in a large herbivore and weak evidence for variations in the secondary sex ratio owing to environmental conditions. The pattern is highly relevant in the context of climate change and its impact on the population dynamics of large herbivores.

Abstract Some carnivores are known to survive well in urban habitats, yet the underlying behavioral tactics are poorly understood. One likely explanation for the success in urban habitats might be that carnivores are generalist consumers. However, urban populations of carnivores could as well consist of specialist feeders. Here, we compared the isotopic specialization of red foxes in urban and rural environments, using both a population and an individual level perspective. We measured stable isotope ratios in increments of red fox whiskers and potential food sources. Our results reveal that red foxes have a broad isotopic dietary niche and a large variation in resource use. Despite this large variation, we found significant differences between the variance of the urban and rural population for δ13C as well as δ15N values, suggesting a habitat‐specific foraging behavior. Although urban regions are more heterogeneous regarding land cover (based on the Shannon index) than rural regions, the dietary range of urban foxes was smaller compared with that of rural conspecifics. Moreover, the higher δ13C values and lower δ15N values of urban foxes suggest a relatively high input of anthropogenic food sources. The diet of most individuals remained largely constant over a longer period. The low intraindividual variability of urban and rural red foxes suggests a relatively constant proportion of food items consumed by individuals. Urban and rural foxes utilized a small proportion of the potentially available isotopic dietary niche as indicated by the low within‐individual variation compared to the between‐individual variation. We conclude that generalist fox populations consist of individual food specialists in urban and rural populations at least over those periods covered by our study.

Abstract Mammalian species express a broad variety in the shape of their extrahepatic biliary tract. While a gall bladder is present in many species, others are lacking this organ. Evolutionary driving forces for these differences have not been determined yet, and organ-specific pathologies might present potentially influencing factors. We reviewed the literature regarding extrahepatic biliary tract pathologies reported in mammalian species of zoo animals and wildlife. Specific pathologies have been reported in the taxonomic orders Proboscidea, Chiroptera, Carnivora, Artiodactyla, Primates, Rodentia, and Lagomorpha with variable frequencies of etiological categories. While metabolic disorders with cholelith formation have been reported mainly in captive populations, parasitological infestation was found particularly in free-ranging animals. Based on the available data, we hypothesize Proboscidea, Primates, and Rodentia species to be prone to cholelithiasis. Species belonging to the Artiodactyla seem to be more susceptible to parasitological infestation while in representatives of the Carnivora infectious, metabolic, neoplastic, and parasitological disorders have been reported nearly equally. Extending our knowledge on extrahepatic biliary tract pathologies in exotic pets and wildlife will support the work of veterinary practitioners as well as scientists in evolutionary biology, making further research in this area strongly recommendable.

Spatial and temporal variability in the availability of food resources will lead to variation in a species' diet, which can then influence patterns of space use, sociality, and life history characteristics. Despite such potential impacts, little information is available about dietary variability for some species with large geographical ranges. Here we quantify the diet and nutritional content of plants consumed by western lowland gorillas (Gorilla gorilla gorilla) in Loango National Park, Gabon over a 2.6 year period and make comparisons with two study sites located 800 km away. The major foods consumed by the Loango gorillas differed greatly from the other two study sites, but gorillas at all three locations spent a similar proportion of feeding time consuming herbaceous vegetation and tree leaves (~ 50%) and fruit (35%). The Loango gorillas spent approximately 10% of feeding time eating nuts, which were not consumed at the other two study sites. Gorillas at those sites spent about 5% of feeding time eating insects, which were not consumed by Loango gorillas. Even though the species composition of the diet differed among the three sites, the nutritional composition of the major food items differed very little, suggesting that western gorillas consume foods of similar nutritional values to meet their dietary needs. This study shows the flexibility of diet of a species with a wide geographic distribution, which has implications for understanding variation in life history characteristics and can be useful for conservation management plans.

Abstract In temperate habitats, the vegetation onset in spring affects the life cycle activities of large herbivores. For one species of large herbivore, roe deer (Capreolus capreolus), parturition is tied to peak resource availability. However, recent long‐term studies of the parturition date of roe deer found only limited support for its earlier occurrence with an earlier onset of spring. In this study, data from a long‐term project (1973–2019) in southern Germany (Baden‐Württemberg; elevation 101–1036 m; 16,130 tagged roe deer fawns) were used to investigate whether the advance in the roe deer parturition date reflected different environmental conditions and was thus explained by temporal and spatial factors. Our results showed a consistent advance toward an earlier parturition date (between 0.16 and 0.33 d/yr), especially for elevations above 750 m, that coincided with an advance in plant phenology. Up to 37% of the variance of the detrended time series of the mean annual parturition was accounted for by the detrended time series of the flowering date of forsythia. Our results suggest a plasticity that allows roe deer to cope with climate‐induced changes along environmental gradients.

Abstract Background Expansion of brain tissue and development of advanced cognitive skills are characteristic traits of human evolution. Their emergence has been causally linked to the intake of nutrients that promote brain development and iodine is considered a critical resource. Rich sources of iodine exist in coastal areas and evolutionary scenarios associate the progressive development of brain size and cognitive skills to such landscapes. This raises the question of how early hominins living in continental areas could have met their iodine requirements. One way to explore this question is to use information from hominoid primates as a proxy for the nutritional ecology of early hominins. Bonobos are particularly interesting in this context as they are restricted to the central part of the Congo basin, an area considered to be iodine deficient based on human standards. Methods Pooled samples of fruit, terrestrial and aquatic herbs were used to assess mineral content with an inductively coupled plasma optical emission spectrometer. Iodine content was measured with the catalytic technique of Sandell-Kolthoff and two separate inductively coupled plasma mass spectrometry methods. Results Nutritional analyses revealed that the mineral content of aquatic herbs is higher than in other plant foods. Moreover, two species of aquatic herbs consumed by bonobos contain iodine concentrations that are almost equivalent to marine algae. Conclusions These data challenge the general notion that the Congo basin is iodine deficient and demonstrate that its lowland forest offers natural sources of iodine in concentrations high enough to prevent iodine deficiency in hominoids and humans.

Frugivorous primates have developed several strategies to deal with wild fruit scarcity, such as modifying their activity budget or enlarging their diet. Agricultural expansion threatens primate habitats and populations (e.g., disease transmission, agrochemical exposure), but it also increases crop feeding opportunities. We aimed at understanding whether maize presence close to the natural habitat of chimpanzees, a threatened species, would lead to significant behavioral modifications. We monitored 20 chimpanzees over 37 months in Kibale National Park, Uganda, with maize gardens at the forest edge. Based on focal nest-to-nest data, we analyzed their diet, activity budget, and energy balance depending on wild fruit and maize availability. We found that the Sebitoli area is a highly nutritive habitat for chimpanzees, with large and caloric wild fruits available all year long. The chimpanzees opportunistically consume maize and exploit it by resting less during maize season. However, no significant variation was found in daily paths and energy expenditures according to maize availability. No behavioral or energy modification was observed regarding wild resources either. Despite the availability of nutritious domestic resources, chimpanzees still exploit wild fruits and do not limit their movements. Thus, their contribution to seed dispersal and forest regeneration in this area is not affected.

In urban areas with a high level of human disturbance, wildlife has to adjust its behavior to deal with the so called “landscape of fear.” This can be studied in risk perception during movement in relation to specific habitat types, whereby individuals trade-off between foraging and disturbance. Due to its high behavioral plasticity and increasing occurrence in urban environments, wild boar (Sus scrofa) is an excellent model organism to study adjustment to urbanization. With the help of GPS tracking, space use of 11 wild boar from Berlin's metropolitan region was analyzed: we aimed at understanding how animals adjust space use to deal with the landscape of fear in urban areas compared to rural areas. We compared use vs. availability with help of generalized linear mixed models. First, we studied landscape types selected by rural vs. urban wild boar, second, we analyzed distances of wild boar locations to each of the landscape types. Finally, we mapped the resulting habitat selection probability to predict hotspots of human-wildlife conflicts. A higher tolerance to disturbance in urban wild boar was shown by a one third shorter flight distance and by an increased re-use of areas close to the trap. Urban wild boar had a strong preference for natural landscapes such as swamp areas, green areas and deciduous forests, and areas with high primary productivity, as indicated by high NDVI (normalized difference vegetation index) values. The areas selected by urban wild boar were often located closely to roads and houses. The spatial distribution maps show that a large area of Berlin would be suitable for urban wild boar but not their rural conspecifics, with the most likely reason being a different perception of anthropogenic disturbance. Wild boar therefore showed considerable behavioral plasticity suitable to adjust to human-dominated environments in a potentially evolutionarily adaptive manner.

Most wildlife species are urban avoiders, but some became urban utilizers and dwellers successfully living in cities. Often, they are assumed to be attracted into urban areas by easily accessible and highly energetic anthropogenic food sources. We macroscopically analysed stomachs of 247 wild boar (Sus scrofa, hereafter WB) from urban areas of Berlin and from the surrounding rural areas. From the stomach contents we determined as predictors of food quality modulus of fineness (MOF,), percentage of acid insoluble ash (AIA) and macronutrients such as amount of energy and percentage of protein, fat, fibre and starch. We run linear mixed models to test: (1) differences in the proportion of landscape variables, (2) differences of nutrients consumed in urban vs. rural WB and (3) the impact of landscape variables on gathered nutrients. We found only few cases of anthropogenic food in the qualitative macroscopic analysis. We categorized the WB into five stomach content categories but found no significant difference in the frequency of those categories between urban and rural WB. The amount of energy was higher in stomachs of urban WB than in rural WB. The analysis of landscape variables revealed that the energy of urban WB increased with increasing percentage of sealing, while an increased human density resulted in poor food quality for urban and rural WB. Although the percentage of protein decreased in areas with a high percentage of coniferous forests, the food quality increased. High percentage of grassland decreased the percentage of consumed fat and starch and increased the percentage of fibre, while a high percentage of agricultural areas increased the percentage of consumed starch. Anthropogenic food such as garbage might serve as fallback food when access to natural resources is limited. We infer that urban WB forage abundant, natural resources in urban areas. Urban WB might use anthropogenic resources (e.g. garbage) if those are easier to exploit and more abundant than natural resources. This study shows that access to natural resources still is mandatory and drives the amount of protein, starch, fat or fibre in wild boar stomachs in urban as well as rural environments.

The daily energy requirements of animals are determined by a combination of physical and physiological factors, but food availability may challenge the capacity to meet nutritional needs. Western gorillas (Gorilla gorilla) are an interesting model for investigating this topic because they are folivore-frugivores that adjust their diet and activities to seasonal variation in fruit availability. Observations of one habituated group of western gorillas in Bai-Hokou, Central African Republic (December 2004-December 2005) were used to examine seasonal variation in diet quality and nutritional intake. We tested if during the high fruit season the food consumed by western gorillas was higher in quality (higher in energy, sugar, fat but lower in fibre and antifeedants) than during the low fruit season. Food consumed during the high fruit season was higher in digestible energy, but not any other macronutrients. Second, we investigated whether the gorillas increased their daily intake of carbohydrates, metabolizable energy (KCal/g OM), or other nutrients during the high fruit season. Intake of dry matter, fibers, fat, protein and the majority of minerals and phenols decreased with increased frugivory and there was some indication of seasonal variation in intake of energy (KCal/g OM), tannins, protein/fiber ratio, and iron. Intake of non-structural carbohydrates and sugars was not influenced by fruit availability. Gorillas are probably able to extract large quantities of energy via fermentation since they rely on proteinaceous leaves during the low fruit season. Macronutrients and micronutrients, but not digestible energy, may be limited for them during times of low fruit availability because they are hind-gut fermenters. We discuss the advantages of seasonal frugivores having large dietary breath and flexibility, significant characteristics to consider in the conservation strategies of endangered species.