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Keeping cool on hot days
(2023)
Long-lived organisms are likely to respond to a rapidly changing climate with behavioral flexibility. Animals inhabiting the arid parts of southern Africa face a particularly rapid rise in temperature which in combination with food and water scarcity places substantial constraints on the ability of animals to tolerate heat. We investigated how three species of African antelope-springbok Antidorcas marsupialis, kudu Tragelaphus strepsiceros and eland T. oryx-differing in body size, habitat preference and movement ecology, change their activity in response to extreme heat in an arid savanna. Serving as a proxy for activity, dynamic body acceleration data recorded every five minutes were analyzed for seven to eight individuals per species for the three hottest months of the year. Activity responses to heat during the hottest time of day (the afternoons) were investigated and diel activity patterns were compared between hot and cool days. Springbok, which prefer open habitat, are highly mobile and the smallest of the species studied, showed the greatest decrease in activity with rising temperature. Furthermore, springbok showed reduced mean activity over the 24 h cycle on hot days compared to cool days. Large-bodied eland seemed less affected by afternoon heat than springbok. While eland also reduced diurnal activity on hot days compared to cool days, they compensated for this by increasing nocturnal activity, possibly because their predation risk is lower. Kudu, which are comparatively sedentary and typically occupy shady habitat, seemed least affected during the hottest time of day and showed no appreciable difference in diel activity patterns between hot and cool days. The interplay between habitat preference, body size, movement patterns, and other factors seems complex and even sub-lethal levels of heat stress have been shown to impact an animal's long-term survival and reproduction. Thus, differing heat tolerances among species could result in a shift in the composition of African herbivore communities as temperatures continue to rise, with significant implications for economically important wildlife-based land use and conservation.
Seasonal differences in spatial distribution of small carnivores in fragmented savannah landscapes
(2008)
Monitoring animal populations in changing environments is crucial to wildlife conservation and management, but restrictions in resources are a recurring problem for wildlife managers and researchers throughout Africa. Land-use- induced shrub encroachment in Kalahari savannah rangelands has led to fragmentation of the landscape. Mammalian carnivores are particularly vulnerable to local extinction in fragmented landscapes, but their low numbers and their often nocturnal and secretive habits make them difficult to monitor. In this study, we tested the applicability of a passive tracking method and compared two measurement methods and index calculations for monitoring small carnivores across a grazing gradient in the southern Kalahari. During the four years of monitoring in a five-year period, we used the knowledge of indigenous Khoisan Bushmen for the identification of carnivore tracks on 640 sand transects (5 m x 250 m). Our results showed that this simple and inexpensive observation method enabled detailed monitoring of 10 small carnivore species across the grazing gradient. A binary index calculated an index based on presence/absence of a species' tracks on each transect, whereas the track intrusion index used the number of track intrusions to each transect for each species in its calculations. For less common species, the two indices were similar in trend and magnitude, because the number of intrusions to each transect was typically 1 or 0. Usually, the two indices showed relatively strong correlations. However, species with patchy distributions of higher numbers presented difficulties for the binary index to monitor trends, but not for the track intrusion index.
Understanding mechanisms to predict changes in plant and animal communities is a key challenge in ecology. The need to transfer knowledge gained from single species to a more generalized approach has led to the development of categorization systems where species' similarities in life strategies and traits are classified into ecological groups (EGs) like functional groups/types or guilds. While approaches in plant ecology undergo a steady improvement and refinement of methodologies, progression in animal ecology is lagging behind. With this review, we aim to initiate a further development of functional classification systems in animal ecology, comparable to recent developments in plant ecology. We here (i) give an overview of terms and definitions of EGs in animal ecology, (ii) discuss existing classification systems, methods and application areas of EGs (focusing on terrestrial vertebrates), and (iii) provide a "roadmap towards an animal functional type approach" for improving the application of EGs and classifications in animal ecology. We found that an animal functional type approach requires: (i) the identification of core traits describing species' dependency on their habitat and life history traits, (ii) an optimization of trait selection by clustering traits into hierarchies, (iii) the assessment of "soft traits" as substitute for hardly measurable traits, e.g. body size for dispersal ability, and (iv) testing of delineated groups for validation including experiments.
In fragmented landscapes, the survival of species and the maintenance of populations with healthy genetic structures will largely depend on movement/dispersal of organisms across matrix areas. In this article, we highlight that effects of fragmentation and climate change occur simultaneously and may enhance or mitigate each other. We systematically analyzed the effect of increasing interannual variation in rainfall on the genetic structure of two neighbouring small mammal subpopulations in a fragmented savanna landscape. The effect of interannual rainfall variation is analyzed for two contrasting scenarios that differ in mean annual rainfall and are both close to a dispersal threshold. Scenario 1 (low mean annual rainfall) lies slightly below this threshold and scenario 2 (high mean annual rainfall) slightly above, i.e. the amount of rainfall in an average rainfall year prevents dispersal in scenario 1, but promotes gene flow in scenario 2. We show that the temporal dynamics of the matrix was crucial for gene flow and the genetic structure of the neighbouring small mammal subpopulations. The most important result is that the increase in rainfall variability could both increase and decrease the genetic difference between the subpopulations in a complex pattern, depending on the scenario and on the amount of variation in rainfall. Finally, we discuss that the relevance of the matrix as temporarily suitable habitat may become a key aspect for biodiversity conservation. We conclude to incorporate temporal changes in matrix suitability in metapopulation theory since local extinctions, gene flow and re-colonization are likely to be affected in fragmented landscapes with such dynamic matrix areas.
Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground- dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.
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.
In this field experiment we investigate the impact of land use induced savanna degradation on movement behaviour of the spotted sand lizard (Pedioplanis l. lineoocellata) in the southern Kalahari. Foraging behaviour of lizards was tested in a factorial design (low vs. high prey availability) in degraded and non-degraded habitats.
An interaction between habitat structure and prey availability affected movement behaviour. In degraded habitats with low prey availability and in non-degraded habitats with high prey availability the spotted sand lizard moved more like an active forager. In contrast, in degraded habitats with high prey availability and in non-degraded habitats with low prey availability lizards moved like sit-and-wait foragers. Interestingly, the behavioural flexibility of the spotted sand lizard seems to buffer extreme conditions and negative effects of land use impacts.
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.
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.
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.
Studies investigating the effect of aboveground herbivory on plants often use clipping to simulate the effects of herbivores, for practical reasons. However, herbivore movements and transfer of oral secretions during herbivory may cause a different response in plant physiology and morphology compared to clipping.
While studies have compared effects of real herbivory vs. clipping on biomass production, plant physiology, and shoot morphology, no study has compared such effects on root morphology.
Therefore, we investigated the effect of herbivory by grasshoppers, herbivory simulated by clipping, and no herbivory on root morphological traits of ten grassland plant species. Root morphological traits were differently affected by the two herbivory treatments. Grasshopper herbivory significantly changed root morphology toward thinner roots with increased specific root length and root area, and decreased root tissue density compared to untreated control plants.
Clipping had mostly similar, but weaker effects on root morphology than grasshopper herbivory.
On the species level, grasshopper herbivory led to strongest changes in root morphology in almost all cases. In contrast, depending on the species, clipping resulted in varying root morphological trait values similar to grasshopper-damaged plants, or in some cases, more closely aligned with control plants.
Though clipping was partly able to mimic the effects of herbivory by grasshoppers, results also indicate that, depending on the species, grasshopper herbivory had different but mostly stronger effects. We, therefore, recommend that future studies apply herbivory with real herbivores to better reflect natural responses in plants and related processes that root morphological traits mediate.
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.
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.
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.
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.
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.
Improving our understanding of biodiversity and ecosystem functioning and our capacity to inform ecosystem management requires an integrated framework for functional biodiversity research (FBR). However, adequate integration among empirical approaches (monitoring and experimental) and modelling has rarely been achieved in FBR. We offer an appraisal of the issues involved and chart a course towards enhanced integration. A major element of this path is the joint orientation towards the continuous refinement of a theoretical framework for FBR that links theory testing and generalization with applied research oriented towards the conservation of biodiversity and ecosystem functioning. We further emphasize existing decision-making frameworks as suitable instruments to practically merge these different aims of FBR and bring them into application. This integrated framework requires joint research planning, and should improve communication and stimulate collaboration between modellers and empiricists, thereby overcoming existing reservations and prejudices. The implementation of this integrative research agenda for FBR requires an adaptation in most national and international funding schemes in order to accommodate such joint teams and their more complex structures and data needs.
Movement of organisms is one of the key mechanisms shaping biodiversity, e.g. the distribution of genes, individuals and species in space and time. Recent technological and conceptual advances have improved our ability to assess the causes and consequences of individual movement, and led to the emergence of the new field of ‘movement ecology’. Here, we outline how movement ecology can contribute to the broad field of biodiversity research, i.e. the study of processes and patterns of life among and across different scales, from genes to ecosystems, and we propose a conceptual framework linking these hitherto largely separated fields of research. Our framework builds on the concept of movement ecology for individuals, and demonstrates its importance for linking individual organismal movement with biodiversity. First, organismal movements can provide ‘mobile links’ between habitats or ecosystems, thereby connecting resources, genes, and processes among otherwise separate locations. Understanding these mobile links and their impact on biodiversity will be facilitated by movement ecology, because mobile links can be created by different modes of movement (i.e., foraging, dispersal, migration) that relate to different spatiotemporal scales and have differential effects on biodiversity. Second, organismal movements can also mediate coexistence in communities, through ‘equalizing’ and ‘stabilizing’ mechanisms. This novel integrated framework provides a conceptual starting point for a better understanding of biodiversity dynamics in light of individual movement and space-use behavior across spatiotemporal scales. By illustrating this framework with examples, we argue that the integration of movement ecology and biodiversity research will also enhance our ability to conserve diversity at the genetic, species, and ecosystem levels.
While several empirical and theoretical studies have clearly shown the negative effects of climate or landscape changes on population and species survival only few of them addressed combined and correlated consequences of these key environmental drivers. This also includes positive landscape changes such as active habitat management and restoration to buffer the negative effects of deteriorating climatic conditions. In this study, we apply a conceptual spatial modelling approach based on functional types to explore the effects of both positive and negative correlations between changes in habitat and climate conditions on the survival of spatially structured populations. We test the effect of different climate and landscape change scenarios on four different functional types that represent a broad spectrum of species characterised by their landscape level carrying capacity, the local population turnover rates at the patch level (K-strategies vs. r-strategies) and dispersal characterstics. As expected, simulation results show that correlated landscape and climatic changes can accelerate (in case of habitat loss or degradation) or slow down (in case of habitat gain or improvement) regional species extinction. However, the strength of the combined changes depends on local turnover at the patch level, the overall landscape capacity of the species, and its specific dispersal characteristics. Under all scenarios of correlated changes in habitat and climate conditions we found the highest sensitivity for functional types representing species with a low landscape capacity but a high population growth rate and a strong density regulation causing a high turnover at the local patch level.
The relative importance of habitat loss or habitat degradation, in combination with climate deterioration, differed among the functional types. However, an increase in regional capacity revealed a similar response pattern: For all types, habitat improvement led to higher survival times than habitat gain, i.e. the establishment of new habitat patches. This suggests that improving local habitat quality at a regional scale is a more promising conservation strategy under climate change than implementing new habitat patches. This conceptual modelling study provides a general framework to better understand and support the management of populations prone to complex environmental changes.
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.
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.
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.
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.
The abandonment of military areas leads to succession processes affecting valuable open-land habitats and is considered to be a major threat for European butterflies. We assessed the ability of hyper spectral remote sensing data to spatially predict the occurrence of one of the most endangered butterfly species (Hipparchia statilinus) in Brandenburg (Germany) on the basis of habitat characteristics at a former military training area. Presence-absence data were sampled on a total area of 36 km(2), and N = 65 adult individuals of Hipparchia statilinus could be detected. The floristic composition within the study area was modeled in a three-dimensional ordination space. Occurrence probabilities for the target species were predicted as niches between ordinated floristic gradients by using Regression Kriging of Indicators. Habitat variance could be explained by up to 81 % with spectral variables at a spatial resolution of 2 x 2 m by transferring PLSR models to imagery. Ordinated ecological niche of Hipparchia statilinus was tested against environmental predictor variables. N = 6 variables could be detected to be significantly correlated with habitat preferences of Hipparchia statilinus. They show that Hipparchia statilinus can serve as a valuable indicator for the evaluation of the conservation status of Natura 2000 habitat type 2330 (inland dunes with open Corynephorus and Agrostis grasslands) protected by the Habitat Directive (Council Directive 92/43/EEC). The authors of this approach, conducted in August 2013 at Doberitzer Heide Germany, aim to increase the value of remote sensing as an important tool for questions of biodiversity research and conservation.
Military areas are valuable habitats and refuges for rare and endangered plants and animals. We developed a new approach applying innovative methods of hyperspectral remote sensing to bridge the existing gap between remote sensing technology and the demands of the nature conservation community. Remote sensing has already proven to be a valuable monitoring instrument. However, the approaches lack the consideration of the demands of applied nature conservation which includes the legal demands of the EU Habitat Directive. Following the idea of the Vital Signs Monitoring in the USA, we identified a subset of the highest priority monitoring indicators for our study area. We analyzed continuous spectral response curves and tested the measurability of N=19 indicators on the basis of complexity levels aggregated from extensive vegetation assemblages. The spectral differentiability for the floristic as well as faunistic indicators revealed values up to 100% accuracy. We point out difficulties when it comes to distinguishing faunistic habitat requirements of several species adapted to dry open landscapes, which in this case results in OVERALL ACCURACY of 67, 87-95, and 35% in the error matrix. In summary, we provide an applicable and feasible method to facilitating monitoring military areas by hyperspectral remote sensing in the following. (C) 2014 Elsevier Ltd. All rights reserved.
Many semi arid savannas are prone to degradation, caused for example, by overgrazing or extreme climatic events, which often lead to shrub encroachment. Overgrazing by livestock affects vegetation and infiltration processes by directly altering plant composition (selective grazing) or by impacting soil physical properties (trampling). Water infiltration is controlled by several parameters, such as macropores (created by soil-burrowing animals or plant roots) and soil texture, but their effects have mostly been studied in isolation. Here we report on a study, in which we conducted infiltration experiments to analyze the interconnected effects of invertebrate-created macropores, shrubs and soil texture (sandy soil and loamy sand) on infiltration in two Namibian rangelands. Using structural equation modeling, we found a direct positive effect of shrub size on infiltration and indirectly via invertebrate macropores on both soil types. On loamy sands this effect was even stronger, but additionally, invertebrate-created macropores became relevant as a direct driver of infiltration. Our results provide new insights into the effects of vegetation and invertebrates on infiltration under different soil textures. Pastoralists should use management strategies that maintain a heterogeneous plant community that supports soil fauna to sustain healthy soil water dynamics, particularly on soils with higher loam content. Understanding the fundamental functioning of soil water dynamics in drylands is critical because these ecosystems are water-limited and support the livelihoods of many cultures worldwide.
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.
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.
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.
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.
Genetic structure and dispersal in a small South African Rodent : is dispersal female-biased?
(2009)
Dispersal greatly determines genetic structure of populations, although it is influenced by landscape heterogeneity, quality of the matrix, resource distribution and local population densities and dynamics. To get insights into some of those processes we analysed the genetic structure of the hairy-footed gerbil Gerbillurus paeba (Rodentia, Murinae, Gerbillinae) in the southern Kalahari (South Africa). Samples were taken from 20 populations covering an area of about 2200 km2. Genetic data were related to landscape characters and population dynamics. We used newly developed microsatellites and found at all loci some indication for the presence of null alleles. However, null alleles seem to have little influence on the general results of our analyses. Altogether we found even nearby populations of G. paeba to be significantly differentiated, although assignment tests revealed 24% of individuals as immigrants. Genetic structure was independent of landscape heterogeneities at all spatial scales. Autocorrelation analyses (range 50-90 km) revealed significant genetic structure within populations on distances <3 km. We found some indication for female-biased dispersal. Our study suggests that dispersing individuals have little influence on the long-term genetic structure and that drift is the major cause of genetic diversity. The observed genetic pattern likely derives from strong population fluctuations of G. paeba. The landscape structure has little influence on the genetic differentiation between populations.
How predictable is the next move of an animal? Specifically, which factors govern the short- and long-term motion patterns and the overall dynamics of land-bound, plant-eating animals in general and ruminants in particular? To answer this question, we here study the movement dynamics of springbok antelopes Antidorcas marsupialis. We propose several complementary statistical-analysis techniques combined with machine-learning approaches to analyze—across multiple time scales—the springbok motion recorded in long-term GPS tracking of collared springboks at a private wildlife reserve in Namibia. As a result, we are able to predict the springbok movement within the next hour with a certainty of about 20%. The remaining about 80% are stochastic in nature and are induced by unaccounted factors in the modeling algorithm and by individual behavioral features of springboks. We find that directedness of motion contributes approximately 17% to this predicted fraction. We find that the measure for directedeness is strongly dependent on the daily cycle of springbok activity. The previously known daily affinity of springboks to their water points, as predicted from our machine-learning algorithm, overall accounts for only about 3% of this predicted deterministic component of springbok motion. Moreover, the resting points are found to affect the motion of springboks at least as much as the formally studied effects of water points. The generality of these statements for the motion patterns and their underlying behavioral reasons for other ruminants can be examined on the basis of our statistical-analysis tools in the future.
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.
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.
Home range estimation is routine practice in ecological research. While advances in animal tracking technology have increased our capacity to collect data to support home range analysis, these same advances have also resulted in increasingly autocorrelated data. Consequently, the question of which home range estimator to use on modern, highly autocorrelated tracking data remains open. This question is particularly relevant given that most estimators assume independently sampled data. Here, we provide a comprehensive evaluation of the effects of autocorrelation on home range estimation. We base our study on an extensive data set of GPS locations from 369 individuals representing 27 species distributed across five continents. We first assemble a broad array of home range estimators, including Kernel Density Estimation (KDE) with four bandwidth optimizers (Gaussian reference function, autocorrelated‐Gaussian reference function [AKDE], Silverman's rule of thumb, and least squares cross‐validation), Minimum Convex Polygon, and Local Convex Hull methods. Notably, all of these estimators except AKDE assume independent and identically distributed (IID) data. We then employ half‐sample cross‐validation to objectively quantify estimator performance, and the recently introduced effective sample size for home range area estimation ( N̂ area
) to quantify the information content of each data set. We found that AKDE 95% area estimates were larger than conventional IID‐based estimates by a mean factor of 2. The median number of cross‐validated locations included in the hold‐out sets by AKDE 95% (or 50%) estimates was 95.3% (or 50.1%), confirming the larger AKDE ranges were appropriately selective at the specified quantile. Conversely, conventional estimates exhibited negative bias that increased with decreasing N̂ area. To contextualize our empirical results, we performed a detailed simulation study to tease apart how sampling frequency, sampling duration, and the focal animal's movement conspire to affect range estimates. Paralleling our empirical results, the simulation study demonstrated that AKDE was generally more accurate than conventional methods, particularly for small N̂ area. While 72% of the 369 empirical data sets had >1,000 total observations, only 4% had an N̂ area >1,000, where 30% had an N̂ area <30. In this frequently encountered scenario of small N̂ area, AKDE was the only estimator capable of producing an accurate home range estimate on autocorrelated data.
Decisions for the conservation of biodiversity and sustainable management of natural resources are typically related to large scales, i.e. the landscape level. However, understanding and predicting the effects of land use and climate change on scales relevant for decision-making requires to include both, large scale vegetation dynamics and small scale processes, such as soil-plant interactions. Integrating the results of multiple BIOTA subprojects enabled us to include necessary data of soil science, botany, socio-economics and remote sensing into a high resolution, process-based and spatially-explicit model. Using an example from a sustainably-used research farm and a communally used and degraded farming area in semiarid southern Namibia we show the power of simulation models as a tool to integrate processes across disciplines and scales.
The interaction between ecological and hydrological processes is particularly important in arid and semi-arid regions. Often the interaction between these processes is not completely understood and they are studied separately. We developed a grid-based computer model simulating the dynamics of the four most common vegetation types (perennial grass, annuals, dwarf shrubs and shrubs) and related hydrological processes in the region studied. Eco-hydrological interactions gain importance in rangelands with increasing slope, where vegetation cover obstructs run-off and decreases evaporation from the soil. Overgrazing can influence these positive feedback mechanisms. In this study, we first show that model predictions of cover and productivity of the vegetation types are realistic by comparing them with estimates obtained from field surveys. Then, we apply a realistic range in slope angle combined with two land use regimes (light versus heavy grazing intensity). Our simulation results reveal that hydrological processes and associated productivity are strongly affected by slope, whereas the magnitude of this impact depends on overgrazing. Under low stocking rates, undisturbed vegetation is maintained and run-off and evaporation remain low on flat plains and gentle slope. On steep slopes, run-off and evaporation become larger, while water retention potential decreases, which leads to reduced productivity. Overgrazing, however, reduces vegetation cover and biomass production and the landscape"s ability to conserve water decreases even on flat plains and gentle slopes. Generally, the abundance of perennial grasses and shrubs decreases with increasing slope and grazing. Dominance is shifted towards shrubs and annuals. As a management recommendation we suggest that different vegetation growth forms should not only be regarded as forage producers but also as regulators of ecosystem functioning. Particularly on sloping range lands, a high percentage of cover by perennial vegetation insures that water is retained in the system.
The need to implement sustainable resource management regimes for semi-arid and arid rangelands is acute as non- adapted grazing strategies lead to irreversible environmental problems such as desertification and associated loss of economic support to society. In these sensitive ecosystems, traditional sectoral, disciplinary approaches will not work to attain sustainability: achieving a collective vision of how to attain sustainability requires interactive efforts among disciplines in a more integrated approach. Therefore, we developed an integrated ecological-economic approach that consists of an ecological and an economic module and combines relevant processes on either level. Parameters for both modules are adjusted for an arid dwarf shrub savannah in southern Namibia. The economic module is used to analyse decisions of different virtual farmer types on annual stocking rates depending on their knowledge how the ecosystem works and climatic conditions. We used a dynamic linear optimisation model to simulate farm economics and livestock dynamics. The ecological module is used to simulate the impact of the farmers' land-use decision, derived by the economic module, on ecosystem dynamics and resulting carrying capacity of the system for livestock. Vegetation dynamics, based on the concept of State-and-transition models, and forage productivity for both modules is derived by a small- scale and spatially explicit vegetation model. This mechanistic approach guarantees that data collected and processes estimated at smaller scales are included in our application. Simulation results of the ecological module were successfully compared to simulation results of the optimisation model for a time series of 30 years. We revealed that sustainable management of semi-arid and arid rangelands relies strongly on rangeland managers' understanding of ecological processes. Furthermore, our simulation results demonstrate that the projected lower annual rainfall due to climate change adds an additional layer of risk to these ecosystems that are already prone to land degradation.
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.
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.
Savannahs are often branded by livestock grazing with resulting land degradation. Holistic management of livestock was proposed to contribute to biodiversity conservation by simulating native wildlife grazing behaviour. This study attempts the comparison of the impact of a holistic management regime to a wildlife grazing management regime on grass and ground-dwelling beetle species diversity on neighboring farms in Namibian rangeland. Results show that the response of biodiversity in species richness and composition to holistic management of livestock differs substantially from wildlife grazing with a positive impact. From a total of 39 identified species of ground-dwelling beetles (Coleoptera: Tenebrionidae, Carabidae) from 29 genera, eight species were found to be indicators for holistic management of livestock and three were found to be indicators for wildlife grazed rangeland. Observations suggest that holistic management of livestock may contribute to biodiversity conservation, but the differential effect of grazing management on species assemblages suggests that livestock grazing cannot replace native wildlife herbivory. Implications for insect conservation An adaptive management strategy such as holistic management used in this study shows the potential to support high beetle biodiversity. Holistic management of livestock thus aspects in favour for a sustainable form of grazing management for insect conservation even though it does not functionally replace grazing by native wildlife.
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.
In semi-arid savannas, unsustainable land use can lead to degradation of entire landscapes, e.g. in the form of shrub encroachment. This leads to habitat loss and is assumed to reduce species diversity. In BIOTA phase 1, we investigated the effects of land use on population dynamics on farm scale. In phase 2 we scale up to consider the whole regional landscape consisting of a diverse mosaic of farms with different historic and present land use intensities. This mosaic creates a heterogeneous, dynamic pattern of structural diversity at a large spatial scale. Understanding how the region-wide dynamic land use pattern affects the abundance of animal and plant species requires the integration of processes on large as well as on small spatial scales. In our multidisciplinary approach, we integrate information from remote sensing, genetic and ecological field studies as well as small scale process models in a dynamic region-wide simulation tool. <hr> Interdisziplinäres Zentrum für Musterdynamik und Angewandte Fernerkundung Workshop vom 9. - 10. Februar 2006.
In most stochastic models addressing the persistence of small populations, environmental noise is included by imposing a synchronized effect of the environment on all individuals. However, buffer mechanisms are likely to exist that may counteract this synchronization to some degree. We have studied whether the flexibility in the mating system, which has been observed in some bird species, is a potential mechanism counteracting the synchronization of environmental fluctuations. Our study organism is the lesser spotted woodpecker Picoides minor (Linnaeus), a generally monogamous species. However, facultative polyandry, where one female mates with two males with separate nests, was observed in years with male-biased sex ratio. We constructed an individual-based model from data and observations of a population in Taunus, Germany. We tested the impact of three behavioural scenarios on population persistence: (1) strict monogamy; (2) polyandry without costs; and (3) polyandry assuming costs in terms of lower survival and reproductive success for secondary males. We assumed that polyandry occurs only in years with male-biased sex ratio and only for females with favourable breeding conditions. Even low rates of polyandry had a strong positive effect on population persistence. The increase of persistence with carrying capacity was slower in the monogamous scenario, indicating strong environmental noise. In the polyandrous scenarios, the increase of persistence was stronger, indicating a buffer mechanism. In the polyandrous scenarios, populations had a higher mean population size, a lower variation in number of individuals, and recovered faster after a population breakdown. Presuming a realistic polyandry rate and costs for polyandry, there was still a strong effect of polyandry on persistence. The results show that polyandry and in general flexibility in mating systems is a buffer mechanism that can significantly reduce the impact of environmental and demographic noise in small populations. Consequently, we suggest that even behaviour that seems to be exceptional should be considered explicitly when predicting the persistence of populations