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Ensuring ecosystem resilience is an intuitive approach to safeguard the functioning of ecosystems and hence the future provisioning of ecosystem services (ES). However, resilience is a multi-faceted concept that is difficult to operationalize. Focusing on resilience mechanisms, such as diversity, network architectures or adaptive capacity, has recently been suggested as means to operationalize resilience. Still, the focus on mechanisms is not specific enough. We suggest a conceptual framework, resilience trinity, to facilitate management based on resilience mechanisms in three distinctive decision contexts and time-horizons: i) reactive, when there is an imminent threat to ES resilience and a high pressure to act, ii) adjustive, when the threat is known in general but there is still time to adapt management, and iii) provident, when time horizons are very long and the nature of the threats is uncertain, leading to a low willingness to act. Resilience has different interpretations and implications at these different time horizons, which also prevail in different disciplines. Social ecology, ecology, and engineering are often implicitly focussing on provident, adjustive, or reactive resilience, respectively, but these different notions and of resilience and their corresponding social, ecological, and economic trade-offs need to be reconciled. Otherwise, we keep risking unintended consequences of reactive actions, or shying away from provident action because of uncertainties that cannot be reduced. The suggested trinity of time horizons and their decision contexts could help ensuring that longer-term management actions are not missed while urgent threats to ES are given priority.

Fertilization causes species loss and species dominance changes in plant communities worldwide. However, it still remains unclear how fertilization acts upon species functional traits, e.g. seed mass. Seed mass is a key trait of the regeneration strategy of plants, which influences a range of processes during the seedling establishment phase. Fertilization may select upon seed mass, either directly by increased nutrient availability or indirectly by increased competition. Since previous research has mainly analyzed the indirect effects of fertilization, we disentangled the direct and indirect effects to examine how nutrient availability and competition influence the seed mass relationships on four key components during seedling establishment: seedling emergence, time of seedling emergence, seedling survival and seedling growth. We conducted a common garden experiment with 22 dry grassland species with a two-way full factorial design that simulated additional nutrient supply and increased competition. While we found no evidence that fertilization either directly by additional nutrient supply or indirectly by increased competition alters the relationship between seed mass and (time of) seedling emergence, we revealed that large seed mass is beneficial under nutrient-poor conditions (seedlings have greater chances of survival, particularly in nutrient-poor soils) as well as under competition (large-seeded species produced larger seedlings, which suffered less from competition than small-seeded species). Based on these findings, we argue that both factors, i.e. nutrient availability and competition intensity, ought to be considered to understand how fertilization influences seedling establishment and species composition with respect to seed mass in natural communities. We propose a simple conceptual model, in which seed mass in natural communities is determined by competition intensity and nutrient availability. Here, we hypothesize that seed mass shows a U-shaped pattern along gradients of soil fertility, which may explain the contrasting soil fertility-seed mass relationships found in the recent literature.

Density regulation influences population dynamics through its effects on demographic rates and consequently constitutes a key mechanism explaining the response of organisms to environmental changes. Yet, it is difficult to establish the exact form of density dependence from empirical data. Here, we developed an individual-based model to explore how resource limitation and behavioural processes determine the spatial structure of white stork Ciconia ciconia populations and regulate reproductive rates. We found that the form of density dependence differed considerably between landscapes with the same overall resource availability and between home range selection strategies, highlighting the importance of fine-scale resource distribution in interaction with behaviour. In accordance with theories of density dependence, breeding output generally decreased with density but this effect was highly variable and strongly affected by optimal foraging strategy, resource detection probability and colonial behaviour. Moreover, our results uncovered an overlooked consequence of density dependence by showing that high early nestling mortality in storks, assumed to be the outcome of harsh weather, may actually result from density dependent effects on food provision. Our findings emphasize that accounting for interactive effects of individual behaviour and local environmental factors is crucial for understanding density-dependent processes within spatially structured populations. Enhanced understanding of the ways animal populations are regulated in general, and how habitat conditions and behaviour may dictate spatial population structure and demographic rates is critically needed for predicting the dynamics of populations, communities and ecosystems under changing environmental conditions.

Arid savannas are regarded as one of the ecosystems most likely to be affected by climate change. In these dry conditions, even top predators like raptors are affected by water availability and precipitation. However, few research initiatives have addressed the question of how climate change will affect population dynamics and extinction risk of particular species in arid ecosystems. Here, we use an individual-oriented modeling approach to conduct experiments on the population dynamics of long lived raptors. We investigate the potential impact of precipitation variation caused by climate change on raptors in arid savanna using the tawny eagle (Aquila rapax) in the southern Kalahari as a case study. We simulated various modifications of precipitation scenarios predicted for climate change, such as lowered annual precipitation mean, increased inter-annual variation and increased auto-correlation in precipitation. We found a high impact of these modifications on extinction risk of tawny eagles, with reduced population persistence in most cases. Decreased mean annual precipitation and increased inter-annual variation both caused dramatic decreases in population persistence. Increased auto-correlation in precipitation led only to slightly accelerated extinction of simulated populations. Finally, for various patterns of periodically fluctuating precipitation, we found both increased and decreased population persistence. In summary, our results suggest that the impacts on raptor population dynamics and survival caused by climate change in arid savannas will be great. We emphasize that even if under climate change the mean annual precipitation remains constant but the inter-annual variation increases the persistence of raptor populations in arid savannas will decrease considerably. This suggests a new dimension of climate change driven impacts on population persistence and consequently on biodiversity. However, more investigations on particular species and/or species groups are needed to increase our understanding of how climate change will impact population dynamics and how this will influence species diversity and biodiversity.

The spatial distribution of plants is often thought to be an indicator of underlying biotic and abiotic processes. However, there are relatively few examples of spatial patterns being analysed to detect an underlying ecological process. Using the spacing patterns being analysed to detect an underlying ecological process. Using the spacing of savanna trees in the southern Kalahari as an example, we applied methods of computer simulation modelling and point pattern analysis in an evaluation of their potential for identifying relevant pattern generating processes from snapshot pattern. We compared real tree patterns from the southern Kalahari, derived from aerial photographs, with patterns produced from computer simulation experiments in an investigation of the following questions: does the present pattern of tree distributions allow us to characterize (1) the relative importance of the major driving forces (e.g., competition for moisture, grass fire, herbivory), (2) the spatial dimensions and structures of the underlying processes, and (3) the actual dynamic status of the ecological system (a phase of decline, increase or constancy with respect to tree abundance)? The simulation experiments are based on a well established, spatially explicit, grid-based model that simulates the vegetation dynamics of the major life forms under a realistic rainfall scenario of the southern Kalahari and under the impact of grass fires, herbivory and the formation of localized clumps with increased tree seed availability. For a realistic range of parameters the simulation model produces long-term coexistence of trees and grasses with tree densities that correspond with long-term coexistence of trees and grasses with tree densities that correspond with densities observed in the field. Both real tree distributions derived from acrial photographs and tree pattern produced by the model are characterized by a tendency towards even spacing at small scales, clumping at intermediate scales and randomness or clumping at large scales. However, increasing the spatio-temporal correlation in the formation of seed patches in the model caused an increase in the tendency towards clumping in the tree distribution whereas an increase in seed patch numbers led to a decrease in clumping. Within single simulation runs the tree pattern could change in response to the variable rainfall sequences and the corresponding differences in grass fire frequency: periods of slightly increasing tree numbers caused by higher precipitation were characterized by an increase in tree clumping whereas periods of slightly decreasing tree numbers showed a tendency towards random or even tree spacing. Simulating the transition of an open savanna to a savanna woodland showed that the tree pattern in the transitional phase can be diagnostic of the underlying process: If the transition was caused by improved moisture conditions the transitional phase was characterized by increased clumping in the tree pattern. In contrast, a transition caused by an increase in the number of localized tree seed patches led to a characteristic even spacing of trees. Even though the simulated savanna clearly showed non-equilibrium dynamics, simulation results indicate that the tree population in the simulated area of the southern Kalahari is in a state of long-term tree-grass coexistence with the persisting structure of an open savanna system.