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Institute
We analyzed relative sensitivities of small- and medium-sized carnivores to livestock husbandry (stocking rates and predator control) in Kalahari, South Africa, rangelands at a regional scale. We monitored small carnivores using track counts on 22 Kalahari farms across a land-use gradient ranging from low to high stocking rates and also interviewed each farm manager to identify farmers" perception of small carnivores as potential predators for livestock. We recorded 12 species of small- and medium-sized carnivores across 22 Kalahari farms. Stocking rate was the most important driving variable for local carnivore abundance. Abundance of all species was lowest on farms where stocking rate was high. Most farm managers perceived medium-sized carnivores, in particular, African wildcat (Felis silvestris lybica), black-backed jackal (Canis mesomelas), and caracal (Caracal caracal), as potential predators of livestock. Multiple regression analysis shows that black-backed jackal, African wildcat, and caracal were negatively affected by predator control measures, whereas bat-eared fox (Otocyon megalotis), cape fox (Vulpes chama), and small-spotted genet (Genetta genetta) were positively affected. Our results show a need for expanding research and conservation activities toward small- and medium-sized carnivores in southern African savannah rangelands. We, therefore, suggest developing a monitoring program combining passive tracking with indigenous knowledge of local Khoisan Bushmen to monitor carnivore populations, and we recommend additional predator removal experiments that manipulate predator densities.
Structural and animal species diversity in arid and semi-arid savannas of the southern Kalahari
(2004)
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.
Although the effects of grazing-induced savannah degradation on animal diversity are well documented, knowledge of how they affect space use or responding behaviour remains poor. In this study, we analysed space use of the spotted sand lizard (Pedioplanis l. lineoocellata) in degraded versus nondegraded habitats of southern Kalahari savannah habitats. Lizards were radio tracked, daily movement distances recorded and home range sizes calculated. In degraded Kalahari savannah habitats where plant diversity and perennial grass cover are low but shrub cover high, P. lineoocellata moves larger distances (40.88 +/- 6.42 m versus 27.43 +/- 5.08 m) and occupies larger home ranges (646.64 +/- 244.84 m(2) versus 209.15 +/- 109.84 m(2)) than in nondegraded habitats (high plant diversity, high perennial grass cover and low shrub cover). We assume that this increase in daily movement distances and home range sizes is a behavioural plasticity to limited food resources in degraded savannah habitats. Although P. lineoocellata is able to adjust to resource-poor savannah habitats, the increase in the lizard's movement activities is likely to result in a higher predation risk. This is supported by the lower availability of protective vegetation i.e. perennial grass cover. Hence, we conclude that despite behavioural plasticity of P. lineoocellata, overgrazing has a severe negative impact on the space use of P. lineoocellata.
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.
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.
Aim: Across the planet, grass-dominated biomes are experiencing shrub encroachment driven by atmospheric CO2 enrichment and land-use change. By altering resource structure and availability, shrub encroachment may have important impacts on vertebrate communities. We sought to determine the magnitude and variability of these effects across climatic gradients, continents, and taxa, and to learn whether shrub thinning restores the structure of vertebrate communities. Location: Worldwide. Time period: Contemporary. Major taxa studied: Terrestrial vertebrates. Methods: We estimated relationships between percentage shrub cover and the structure of terrestrial vertebrate communities (species richness, Shannon diversity and community abundance) in experimentally thinned and unmanipulated shrub-encroached grass-dominated biomes using systematic review and meta-analyses of 43 studies published from 1978 to 2016. We modelled the effects of continent, biome, mean annual precipitation, net primary productivity and the normalized difference vegetation index (NDVI) on the relationship between shrub cover and vertebrate community structure. Results: Species richness, Shannon diversity and total abundance had no consistent relationship with shrub encroachment and experimental thinning did not reverse encroachment effects on vertebrate communities. However, some effects of shrub encroachment on vertebrate communities differed with net primary productivity, amongst vertebrate groups, and across continents. Encroachment had negative effects on vertebrate diversity at low net primary productivity. Mammalian and herpetofaunal diversity decreased with shrub encroachment. Shrub encroachment also had negative effects on species richness and total abundance in Africa but positive effects in North America. Main conclusions: Biodiversity conservation and mitigation efforts responding to shrub encroachment should focus on low-productivity locations, on mammals and herpetofauna, and in Africa. However, targeted research in neglected regions such as central Asia and India will be needed to fill important gaps in our knowledge of shrub encroachment effects on vertebrates. Additionally, our findings provide an impetus for determining the mechanisms associated with changes in vertebrate diversity and abundance in shrub-encroached grass-dominated biomes.
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.
1. The complex, nonlinear response of dryland systems to grazing and climatic variations is a challenge to management of these lands. Predicted climatic changes will impact the desertification of drylands under domestic livestock production. Consequently, there is an urgent need to understand the response of drylands to grazing under climate change. 2. We enhanced and parameterized an ecohydrological savanna model to assess the impacts of a range of climate change scenarios on the response of a semi-arid African savanna to grazing. We focused on the effects of temperature and CO2 level increase in combination with changes in inter- and intra-annual precipitation patterns on the long-term dynamics of three major plant functional types. 3. We found that the capacity of the savanna to sustain livestock grazing was strongly influenced by climate change. Increased mean annual precipitation and changes in intra-annual precipitation pattern have the potential to slightly increase carrying capacities of the system. In contrast, decreased precipitation, higher interannual variation and temperature increase are leading to a severe decline of carrying capacities owing to losses of the perennial grass biomass. 4. Semi-arid rangelands will be at lower risk of shrub encroachment and encroachment will be less intense under future climatic conditions. This finding holds in spite of elevated levels of atmospheric CO2 and irrespective of changes in precipitation pattern, because of the drought sensitivity of germination and establishment of encroaching species. 5. Synthesis and applications. Changes in livestock carrying capacities, both positive and negative, mainly depend on the highly uncertain future rainfall conditions. However, independent of the specific changes, shrub encroachment becomes less likely and in many cases less severe. Thus, managers of semi-arid rangelands should shift their focus from woody vegetation towards perennial grass species as indicators for rangeland degradation. Furthermore, the resulting reduced competition from woody vegetation has the potential to facilitate ecosystem restoration measures such as re-introduction of desirable plant species that are only little promising or infeasible under current climatic conditions. On a global scale, the reductions in standing biomass resulting from altered degradation dynamics of semi-arid rangelands can have negative impacts on carbon sequestration.
We investigated sex-specific parental care behaviour of lesser spotted woodpeckers Picoides minor in the low mountain range Taunus, Germany. Observed parental care included incubation, nest sanitation as well as brooding and feeding of nestlings. Contributions of the two sexes to parental care changed in progress of the breeding period. During incubation and the first half of the nestling period, parental care was divided equally between partners. However, in the late nestling stage, we found males to feed their nestlings irrespective of brood size while females considerably decreased feeding rate with the number of nestlings. This behaviour culminated in desertion of small broods by females shortly before fledging. The fact that even deserted nests were successful indicates that males were able to compensate for the females' absence. Interestingly, the mating of one female with two males with separate nests could be found in the population, which confirms earlier findings of polyandry in the lesser spotted woodpecker. We conclude that biparental care is not essential in the later stage and one partner can reduce effort and thus costs of parental care, at least in small broods where the mate is able to compensate for that behaviour. Reduced care and desertion appears only in females, which might be caused by a combination of two traits: First, females might suffer higher costs of investment in terms of mortality and secondly, male-biased sex ratio in the population generally leads to higher mating probabilities for females in the following breeding season. The occurrence of polyandry seems to be a result of these conditions.
Seed traits matter
(2021)
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.
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.
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.
Seasonal differences in spatial distribution of small carnivores in fragmented savannah landscapes
(2008)
Scaling up ecohydrological processes : role of surface water flow in water-limited landscapes
(2009)
In this study, we present a stochastic landscape modeling approach that has the power to transfer and integrate existing information on vegetation dynamics and hydrological processes from the small scale to the landscape scale. To include microscale processes like ecohydrological feedback mechanisms and spatial exchange like surface water flow, we derive transition probabilities from a fine-scale simulation model. We applied two versions of the landscape model, one that includes and one that disregards spatial exchange of water to the situation of a sustainably used research farm and communally used and degraded rangeland in semiarid Namibia. Our simulation experiments show that including spatial exchange of overland flow among vegetation patches into our model is a precondition to reproduce vegetation dynamics, composition, and productivity, as well as hydrological processes at the landscape scale. In the model version that includes spatial exchange of water, biomass production at light grazing intensities increases 2.24-fold compared to the model without overland flow. In contrast, overgrazing destabilizes positive feedbacks through vegetation and hydrology and decreases the number of hydrological sinks in the model with overland flow. The buffer capacity of these hydrological sinks disappears and runoff increases. Here, both models predicted runoff losses from the system and artificial droughts occurring even in years with good precipitation. Overall, our study reveals that a thorough understanding of overland flow is an important precondition for improving the management of semiarid and arid rangelands with distinct topography.
Savanna rangelands worldwide are threatened by shrub encroachment, i.e. the increase of woody plant species at the cost of perennial grasses, causing a strong decline in the productivity of domestic livestock production. Although recent studies indicate that fire might be of great importance for semi-arid and arid savanna dynamics, it is largely not applied in the management of semi-arid rangelands especially with regard to woody plant control. We used the eco-hydrological savanna model EcoHyD to simulate the effects of different fire management strategies on semi-arid savanna vegetation and to assess their longterm suitability for semi-arid rangeland management. Simulation results show that prescribed fires, timed to kill tree seedlings prevented shrub encroachment for a broad range of livestock densities while the possible maximum long-term cattle densities on the simulated semi-arid rangeland in Namibia increased by more than 30%. However, when grazing intensity was too high, fire management failed in preventing shrub encroachment.
Our findings indicate that with regard to fire management a clear distinction between mesic and more arid savannas is necessary: While the frequency of fires is of relevance for mesic savannas, we recommend a fire management focussing on the timing of fire for semi-arid and arid savannas. (C) 2014 Elsevier Ltd. All rights reserved.
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.
Introduction: Many semiarid regions around the world are presently experiencing significant changes in both climatic conditions and vegetation. This includes a disturbed coexistence between grasses and bushes also known as bush encroachment, and altered precipitation patterns with larger rain events. Fewer, more intense precipitation events might promote groundwater recharge, but depending on the structure of the vegetation also encourage further woody encroachment.
Materials and Methods: In this study, we investigated how patterns and sources of water uptake of Acacia mellifera (blackthorn), an important encroaching woody plant in southern African savannas, are associated with the intensity of rain events and the size of individual shrubs. The study was conducted at a commercial cattle farm in the semiarid Kalahari in Namibia (MAP 250 mm/a). We used soil moisture dynamics in different depths and natural stable isotopes as markers of water sources. Xylem water of fifteen differently sized individuals during eight rain events was extracted using a Scholander pressure bomb.
Results and Discussion: Results suggest the main rooting activity zone of A. mellifera in 50 and 75 cm soil depth but a reasonable water uptake from 10 and 25 cm. Any apparent uptake pattern seems to be driven by water availability, not time in the season. Bushes prefer the deeper soil layers after heavier rain events, indicating some evidence for the classical Walter’s two-layer hypothesis. However, rain events up to a threshold of 6 mm/day cause shallower depths of use and suggest several phases of intense competition with perennial grasses. The temporal uptake pattern does not depend on shrub size, suggesting a fast upwards water flow inside. d2H and d18O values in xylem water indicate that larger shrubs rely less on upper and very deep soil water than smaller shrubs. It supports the hypothesis that in environments where soil moisture is highly variable in the upper soil layers, the early investment in a deep tap-root to exploit deeper, more reliable water sources could reduce the probability of mortality during the establishment phase. Nevertheless, independent of size and time in the season, bushes do not compete with potential groundwater recharge. In a savanna encroached by A. mellifera, groundwater will most likely be affected indirectly.
Moving in the Anthropocene
(2018)
Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission.