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Institute
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.
Fire and grazing shape biodiversity in savannah landscapes. In land use management, knowing the effects of fire and grazing on biodiversity are important in order to ensure environmental sustainability. Beetles specifically are indicators of the biodiversity response to fire and grazing. A grazing exclusion and burning experiment in a split-plot design was used in order to investigate the interacting effects of fire and wildlife grazing on biomass, diversity, and species composition of darkling beetles (Coleoptera, Tenebrionidae) over time after fire. Darkling beetle species richness and diversity were responding in a three-way-interaction to fire, grazing, and time after fire, whereby biomass of darkling beetles remained unaffected and species compositional changes were attributed to seasonal changes of time only. Fire on ungrazed plots had a negative effect on species diversity and richness 2 weeks and 6 months post fire, whereas fire on grazed plots had no impact on species diversity and richness. Grazing only lowered species diversity and richness 6 months after fire treatments. Results suggest that grazing overrides the effects of fire and that the similar effects caused by fire and grazing are due to niche and assemblage simplification of the habitat.
To assess the ecological and economic implications of the redistributive land reform in semi-arid Namibia, we investigated to what extent land reform beneficiaries adjust herd size and herd composition according to environmental (rainfall, vegetation) and economic variables (herd size, financial assets, running costs). We performed model-based role-plays with Namibian land reform beneficiaries, simulating 10 years of rangeland management.
Our study revealed that the farmers surveyed mainly manage their herds according to their economic situation (herd size and account balance) but do not take environmental variability (rainfall and vegetation) into account. Further, our results indicate that, due to financial pressure, farmers are not able to apply their desired management strategies, and that owners of small farms face a higher risk of economic failure. However, farmers apply rather conservative and constant stocking rates and will thus, given the current economic limitations, likely not contribute to semi-arid savanna degradation.
We conclude that land reform beneficiaries need support to be able to apply straightforward and efficient management strategies. This could be achieved by facilitating cooperation between small farming businesses and by supporting initial investment in productive cattle herds at the time of redistribution of the land.
Major drivers of savanna shrub encroachment are climatic conditions, CO2 and unsustainable grazing management including fire prevention. Although all drivers affect ecohydrological processes, and given that water is a seasonally scarce resource in savannas, it remains largely unclear how shrub encroachment itself affects hydrological conditions that feed back into water use and community assembly of the remaining plant community. Hence, understanding direct ecohydrological effects of shrubs that may limit the recovery of the perennial herbaceous vegetation in grazed areas and promote the establishment of shrub seedlings facilitates the identification of areas that are most sensitive to further encroachment. In our trait-based approach, we determined relationships among shrub cover, soil and plant trait characteristics sensitive to water limitation in 120 plots along a shrub cover gradient. We focused on two functional response traits indicating immediate drought stress and subsequent water use for drought stress recovery with associated competition for water (midday leaf/xylem water potential and diurnally recovery rate of leaf water potential), and three functional response traits indicating long-term stress adaptation and related resource use strategies (SLA, plant height and seed release height). To understand species assembly and the associated mechanisms of resource use, we calculated community weighted mean traits, intraspecific trait variability as a proxy for the mechanism of coexistence, and mean traits at plant functional type level including 2-year-old Acacia mellifera-saplings. We found a low intraspecific trait variability in drought stress recovery rate and height suggesting that competitive exclusion via active resource acquisition (i.e. water exploitation) played a minor role for community assembly in a shrub encroaching savanna. The dominant community assembly process was passive stress avoidance via resource conservation up to stress tolerance indicated by the high variability in SLA and midday leaf water potential. Correlations of traits with soil moisture suggest a rooting niche differentiation between annual and perennial grasses and that Acacia-shrub saplings within the first 50 cm of soil already escaped the highest drought stress. Interestingly, immediate drought stress for the herbaceous community was lowest on moderately shrub encroached sites and not on grass dominated sites. Since passive stress avoidance accompanied by a distinct stress tolerance in semi-arid savannas is more important than active competition, and assuming that the low drought stress of the herbaceous community at intermediate levels of shrub cover also applies to newly emerging shrub seedlings, these areas are likely to be most sensitive to further encroachment. As such, they should be considered as focal areas for prevention management.
Understanding the drivers underlying disease dynamics is still a major challenge in disease ecology, especially in the case of long-term disease persistence. Even though there is a strong consensus that density-dependent factors play an important role for the spread of diseases, the main drivers are still discussed and, more importantly, might differ between invasion and persistence periods. Here, we analysed long-term outbreak data of classical swine fever, an important disease in both wild boar and livestock, prevalent in the wild boar population from 1993 to 2000 in Mecklenburg-Vorpommern, Germany. We report outbreak characteristics and results from generalized linear mixed models to reveal what factors affected infection risk on both the landscape and the individual level. Spatiotemporal outbreak dynamics showed an initial wave-like spread with high incidence during the invasion period followed by a drop of incidence and an increase in seroprevalence during the persistence period. Velocity of spread increased with time during the first year of outbreak and decreased linearly afterwards, being on average 7.6 km per quarter. Landscape- and individual-level analyses of infection risk indicate contrasting seasonal patterns. During the persistence period, infection risk on the landscape level was highest during autumn and winter seasons, probably related to spatial behaviour such as increased long-distance movements and contacts induced by rutting and escaping movements. In contrast, individual-level infection risk peaked in spring, probably related to the concurrent birth season leading to higher densities, and was significantly higher in piglets than in reproductive animals. Our findings highlight that it is important to investigate both individual- and landscape-level patterns of infection risk to understand long-term persistence of wildlife diseases and to guide respective management actions. Furthermore, we highlight that exploring different temporal aggregation of the data helps to reveal important seasonal patterns, which might be masked otherwise.
Semiarid woodlands and savannas are globally important biomes that provide ecosystem goods and services such as habitat for biota and sinks for carbon, support millions of people that rely primarily on pastoralism, and supply livelihoods for about a third of the global human population. Savannas, however, are prone to degradation by overgrazing, and encroachment by woody plants, reducing their capacity to produce forage that pastoral enterprises depend on. We examined the impacts of livestock grazing and woody encroachment on soil hydrological processes, hypothesizing that heavy grazing by livestock would reduce hydrological function, whereas woody plants would increase hydrological function, therefore, partially offsetting any negative effects of overgrazing by livestock. Understanding the major drivers of soil hydrology in savanna ecosystems is important because water is a critical, yet limited resource in savannas. We found that livestock grazing reduced the early (sorptivity) and late (steady-state infiltration) stages of infiltration under both ponding and tension, and attributed this to a reduction in porosity caused by livestock trampling. Steady-state infiltration and sorptivity under ponding were greater under the canopies of woody shrubs than in open areas, partly compensating for any negative effect of grazing. Structural equation modeling revealed a direct positive effect of shrub height on hydrological functions, and an indirect effect via increases in litter cover. Our results suggest that woody plants can play important roles in driving hydrological function in savannas, counteracting the suppressive effect of livestock overgrazing on infiltration processes. Management strategies in semiarid savannas should aim to reduce trampling by livestock and retain large woody plants in order to maintain hydrological function. (C) 2019 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.
Context Human land use intensified over the last century and simultaneously, extreme weather events have become more frequent. However, little is known about the interplay between habitat structure, direct short-term weather effects and indirect seasonal effects on animal space use and behavior. Objectives We used the European hare (Lepus europaeus) as model to investigate how habitat structure and weather conditions affect habitat selection and home range size, predictors for habitat quality and energetic requirements. Methods Using > 100,000 GPS positions of 60 hares in three areas in Denmark and Germany, we analyzed habitat selection and home range size in response to seasonally changing habitat structure, measured as vegetation height and agricultural field size, and weather. We compared daily and monthly home ranges to disentangle between direct short-term weather effects and indirect seasonal effects of climate. Results Habitat selection and home range size varied seasonally as a response to changing habitat structure, potentially affecting the availability of food and shelter. Overall, habitat structure and seasonality were more important in explaining hare habitat selection and home range size compared to direct weather conditions. Nevertheless, hares adjusted habitat selection and daily home range size in response to temperature, wind speed and humidity, possibly in response to thermal constrains and predation risk. Conclusions For effective conservation, habitat heterogeneity should be increased, e.g. by reducing agricultural field sizes and the implementation of set-asides that provide both forage and shelter, especially during the colder months of the year.
Agricultural land-use practices have intensified over the last decades, leading to population declines of various farmland species, including the European hare (Lepus europaeus). In many European countries, arable fields dominate agricultural landscapes. Compared to pastures, arable land is highly variable, resulting in a large spatial variation of food and cover for wildlife over the course of the year, which potentially affects habitat selection by hares. Here, we investigated within-home-range habitat selection by hares in arable areas in Denmark and Germany to identify habitat requirements for their conservation. We hypothesized that hare habitat selection would depend on local habitat structure, that is, vegetation height, but also on agricultural field size, vegetation type, and proximity to field edges. Active hares generally selected for short vegetation (1-25 cm) and avoided higher vegetation and bare ground, especially when fields were comparatively larger. Vegetation >50 cm potentially restricts hares from entering parts of their home range and does not provide good forage, the latter also being the case on bare ground. The vegetation type was important for habitat selection by inactive hares, with fabaceae, fallow, and maize being selected for, potentially providing both cover and forage. Our results indicate that patches of shorter vegetation could improve the forage quality and habitat accessibility for hares, especially in areas with large monocultures. Thus, policymakers should aim to increase areas with short vegetation throughout the year. Further, permanent set-asides, like fallow and wildflower areas, would provide year-round cover for inactive hares. Finally, the reduction in field sizes would increase the density of field margins, and farming different crop types within small areas could improve the habitat for hares and other farmland species.
Shrub encroachment in semi-arid savannas is induced by interacting effects of climate, fire suppression, and unsustainable livestock farming; it carries a severe risk of land degradation and strongly influences natural communities that provide key ecosystem functions. However, species-specific effects of shrub cover on many animal groups that act as indicators of degradation remain largely unknown. We analysed the consequences of shrub encroachment for ground-dwelling beetles in a semi-arid Namibian savanna rangeland, where beetles and vegetation were recorded along a shrub cover gradient (30%). Focusing on species niche breadths and optima, we identified two crucial shrub cover thresholds (2.9% and 10.0%), corresponding to major changes in the beetle communities with implications for savanna ecosystem functioning. Niche optima of most species were between the first and second thresholds; beyond the second threshold, saprophagous, coprophagous, and rare predatory beetles declined in numbers and diversity. This is problematic because beetles provide important ecosystem functions, such as decomposition and nutrient cycling. However, we also found that certain species were adapted to high shrub cover, thus providing examples of niche differentiation. Despite the predominantly negative effects of heavy shrub encroachment on beetle communities, shrubs in their early life stages apparently provide essential structures, which enhance habitat quality for ground-dwelling beetles. Our results demonstrate that shrub encroachment can have mixed effects on ground-dwelling beetle communities and hence on savanna ecosystem functioning. We, therefore, conclude that rangeland management and restoration should consider the complex trade-offs between species-specific effects and the level of encroachment for sustainable land use.
Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home-range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied block cross-validation to quantify bias in empirical home-range estimates. Area requirements of mammals <10 kg were underestimated by a mean approximately15%, and species weighing approximately100 kg were underestimated by approximately50% on average. Thus, we found area estimation was subject to autocorrelation-induced bias that was worse for large species. Combined with the fact that extinction risk increases as body mass increases, the allometric scaling of bias we observed suggests the most threatened species are also likely to be those with the least accurate home-range estimates. As a correction, we tested whether data thinning or autocorrelation-informed home-range estimation minimized the scaling effect of autocorrelation on area estimates. Data thinning required an approximately93% data loss to achieve statistical independence with 95% confidence and was, therefore, not a viable solution. In contrast, autocorrelation-informed home-range estimation resulted in consistently accurate estimates irrespective of mass. When relating body mass to home range size, we detected that correcting for autocorrelation resulted in a scaling exponent significantly >1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum.
We investigated whether a given landscape structure affects the level of home range size adaptation in response to resource variability. We tested whether increasing resource variability forces herbivorous mammals to increase their home ranges. In 2014 and 2015 we collared 40 European brown hares (Lepus europaeus) with GPS-tags to record hare movements in two regions in Germany with differing landscape structures. We examined hare home range sizes in relation to resource availability and variability by using the normalized difference vegetation index as a proxy. Hares in simple landscapes showed increasing home range sizes with increasing resource variability, whereas hares in complex landscapes did not enlarge their home range. Animals in complex landscapes have the possibility to include various landscape elements within their home ranges and are more resilient against resource variability. But animals in simple landscapes with few elements experience shortcomings when resource variability becomes high. The increase in home range size, the movement related increase in energy expenditure, and a decrease in hare abundances can have severe implications for conservation of mammals in anthropogenic landscapes. Hence, conservation management could benefit from a better knowledge about fine-scaled effects of resource variability on movement behaviour.
In semi-arid environments characterized by erratic rainfall and scattered primary production, migratory movements are a key survival strategy of large herbivores to track resources over vast areas. Veterinary Cordon Fences (VCFs), intended to reduce wildlife-livestock disease transmission, fragment large parts of southern Africa and have limited the movements of large wild mammals for over 60 years. Consequently, wildlife-fence interactions are frequent and often result in perforations of the fence, mainly caused by elephants. Yet, we lack knowledge about at which times fences act as barriers, how fences directly alter the energy expenditure of native herbivores, and what the consequences of impermeability are. We studied 2-year ungulate movements in three common antelopes (springbok, kudu, eland) across a perforated part of Namibia's VCF separating a wildlife reserve and Etosha National Park using GPS telemetry, accelerometer measurements, and satellite imagery. We identified 2905 fence interaction events which we used to evaluate critical times of encounters and direct fence effects on energy expenditure. Using vegetation type-specific greenness dynamics, we quantified what animals gained in terms of high quality food resources from crossing the VCF. Our results show that the perforation of the VCF sustains herbivore-vegetation interactions in the savanna with its scattered resources. Fence permeability led to peaks in crossing numbers during the first flush of woody plants before the rain started. Kudu and eland often showed increased energy expenditure when crossing the fence. Energy expenditure was lowered during the frequent interactions of ungulates standing at the fence. We found no alteration of energy expenditure when springbok immediately found and crossed fence breaches. Our results indicate that constantly open gaps did not affect energy expenditure, while gaps with obstacles increased motion. Closing gaps may have confused ungulates and modified their intended movements. While browsing, sedentary kudu's use of space was less affected by the VCF; migratory, mixed-feeding springbok, and eland benefited from gaps by gaining forage quality and quantity after crossing. This highlights the importance of access to vast areas to allow ungulates to track vital vegetation patches.
Effects of fences and fence gaps on the movement behavior of three southern African antelope species
(2022)
Globally, migratory ungulates are affected by fences. While field observational studies reveal the amount of animal–fence interactions across taxa, GPS tracking-based studies uncover fence effects on movement patterns and habitat selection. However, studies on the direct effects of fences and fence gaps on movement behavior, especially based on high-frequency tracking data, are scarce. We used GPS tracking on three common African antelopes (Tragelaphus strepsiceros, Antidorcas marsupialis, and T. oryx) with movement strategies ranging from range residency to nomadism in a semi-arid, Namibian savanna traversed by wildlife-proof fences that elephants have regularly breached. We classified major forms of ungulate–fence interaction types on a seasonal and a daily scale. Furthermore, we recorded the distances and times spent at fences regarding the total individual space use. Based on this, we analyzed the direct effects of fences and fence gaps on the animals’ movement behavior for the previously defined types of animal–fence interactions. Antelope-fence interactions peaked during the early hours of the day and during seasonal transitions when the limiting resource changed between water and forage. Major types of ungulate–fence interactions were quick, trace-like, or marked by halts. We found that the amount of time spent at fences was highest for nomadic eland. Migratory springbok adjusted their space use concerning fence gap positions. If the small home ranges of sedentary kudu included a fence, they frequently interacted with this fence. For springbok and eland, distance traveled along a fence declined with increasing utilization of a fence gap. All species reduced their speed in the proximity of a fence but often increased their speed when encountering the fence. Crossing a fence led to increased speeds for all species. We demonstrate that fence effects mainly occur during crucial foraging times (seasonal scale) and during times of directed movements (daily scale). Importantly, we provide evidence that fences directly alter antelope movement behaviors with negative implications for energy budgets and that persistent fence gaps can reduce the intensity of such alterations. Our findings help to guide future animal–fence studies and provide insights for wildlife fencing and fence gap planning.
Subterranean termites create tunnels (macropores) for foraging that can influence water infiltration and may lead to preferential flow to deeper soil layers. This is particularly important in water limited ecosystems such as semi-arid, agriculturally utilized savannas, which are particularly prone to land degradation and shrub-encroachment. Using termite activity has been suggested as a restoration measure, but their impact on hydrology is neither universal nor yet fully understood. Here, we used highly replicated, small-scale (50 x 50 cm) rain-simulation experiments to analyse the interacting effects of either vegetation (grass dominated vs. shrub dominated sites) or soil texture (sand vs. loamy sand) and termite foraging macropores on infiltration patterns. We used Brilliant Blue FCF as colour tracer to make the flow pathways in paired experiments visible, on either termite-disturbed soil or controls without surface macropores in two semi-arid Namibian savannas (with either heterogeneous soil texture or shrub cover). On highly shrub-encroached plots in the savanna site with heterogeneous soil texture, termite macropores increased maximum infiltration depth and total amount of infiltrated water on loamy sand, but not on sandy soil. In the sandy savanna with heterogeneous shrub cover, neither termite activity nor shrub density affected the infiltration. Termite's effect on infiltration depends on the soil's hydraulic conductivity and occurs mostly under ponded conditions, intercepting run-off. In semi-arid savanna soils with a considerable fraction of fine particles, termites are likely an important factor for soil water dynamics.
Changing climatic conditions and unsustainable land use are major threats to savannas worldwide. Historically, many African savannas were used intensively for livestock grazing, which contributed to widespread patterns of bush encroachment across savanna systems. To reverse bush encroachment, it has been proposed to change the cattle-dominated land use to one dominated by comparatively specialized browsers and usually native herbivores. However, the consequences for ecosystem properties and processes remain largely unclear. We used the ecohydrological, spatially explicit model EcoHyD to assess the impacts of two contrasting, herbivore land-use strategies on a Namibian savanna: grazer- versus browser-dominated herbivore communities. We varied the densities of grazers and browsers and determined the resulting composition and diversity of the plant community, total vegetation cover, soil moisture, and water use by plants. Our results showed that plant types that are less palatable to herbivores were best adapted to grazing or browsing animals in all simulated densities. Also, plant types that had a competitive advantage under limited water availability were among the dominant ones irrespective of land-use scenario. Overall, the results were in line with our expectations: under high grazer densities, we found heavy bush encroachment and the loss of the perennial grass matrix. Importantly, regardless of the density of browsers, grass cover and plant functional diversity were significantly higher in browsing scenarios. Browsing herbivores increased grass cover, and the higher total cover in turn improved water uptake by plants overall. We concluded that, in contrast to grazing-dominated land-use strategies, land-use strategies dominated by browsing herbivores, even at high herbivore densities, sustain diverse vegetation communities with high cover of perennial grasses, resulting in lower erosion risk and bolstering ecosystem services.
Global change is shifting the timing of biological events, leading to temporal mismatches between biological events and resource availability. These temporal mismatches can threaten species' populations. Importantly, temporal mismatches not only exert strong pressures on the population dynamics of the focal species, but can also lead to substantial changes in pairwise species interactions such as host-pathogen systems. We adapted an established individual-based model of host-pathogen dynamics. The model describes a viral agent in a social host, while accounting for the host's explicit movement decisions. We aimed to investigate how temporal mismatches between seasonal resource availability and host life-history events affect host-pathogen coexistence, that is, disease persistence. Seasonal resource fluctuations only increased coexistence probability when in synchrony with the hosts' biological events. However, a temporal mismatch reduced host-pathogen coexistence, but only marginally. In tandem with an increasing temporal mismatch, our model showed a shift in the spatial distribution of infected hosts. It shifted from an even distribution under synchronous conditions toward the formation of disease hotspots, when host life history and resource availability mismatched completely. The spatial restriction of infected hosts to small hotspots in the landscape initially suggested a lower coexistence probability due to the critical loss of susceptible host individuals within those hotspots. However, the surrounding landscape facilitated demographic rescue through habitat-dependent movement. Our work demonstrates that the negative effects of temporal mismatches between host resource availability and host life history on host-pathogen coexistence can be reduced through the formation of temporary disease hotspots and host movement decisions, with implications for disease management under disturbances and global change.