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    Patch dynamics integrate mechanisms for savanna tree–grass coexistence

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    Many mechanisms have been suggested to explain the coexistence of woody species and grasses in savannas. However, evidence from field studies and simulation models has been mixed. Patch dynamics is a potentially unifying mechanism explaining tree-grass coexistence and the natural occurrence of shrub encroachment in arid and semi-arid savannas. A patch-dynamic savanna consists of a spatial mosaic of patches. Each patch maintains a cyclical succession between dominance of woody species and grasses, and the succession of neighbouring patches is temporally asynchronous. Evidence from empirical field studies supports the patch dynamics view of savannas. As a basis for future tests of patch dynamics in savannas, several hypotheses are presented and one is exemplarily examined: at the patch scale, realistically parameterized simulation models have generated cyclical succession between woody and grass dominance. In semi-arid savannas, cyclical successions are driven by precipitation conditions that lead to mass recruitment of shrubs in favourable years and to simultaneous collapse of shrub cohorts in drought years. The spatiotemporal pattern of precipitation events determines the scale of the savanna vegetation mosaic in space and time. In a patch-dynamic savanna, shrub encroachment is a natural, transient phase corresponding to the shrub-dominated phase during the successional cycle. Hence, the most promising management strategy for encroached areas is a large-scale rotation system of rangelands. In conclusion, patch dynamics is a possible scale-explicit mechanism for the explanation of tree-grass coexistence in savannas that integrates most of the coexistence mechanisms proposed thus far for savannas

    The role of biotic factors during plant establishment in novel communities assessed with an agent-based simulation model

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    Background Establishment success of non-native species is not only influenced by environmental conditions, but also by interactions with local competitors and enemies. The magnitude of these biotic interactions is mediated by species traits that reflect competitive strength or defence mechanisms. Our aim was to investigate the importance of species traits for successful establishment of non-native species in a native community exhibiting biotic resistance in the form of competition and herbivory. Methods We developed a trait-based, individual-based simulation model tracking the survival of non-native plants in a native community. In the model, non-native plants are characterized by high or low values of competition and defence traits. Model scenarios included variation of initial number of non-natives, intensity of competitive interaction, density of herbivores and density as well as mixture of the native community. Results Traits related to competition had a much greater impact on survival of non-native species than traits related to defence. Survival rates of strong competitors never fell below 50% while survival of weak competitors averaged at about 10%. Weak competitors were also much more susceptible to competitive pressures such as community density, composition and competition intensity. Strong competitors responded negatively to changes in competition intensity, but hardly to composition or density of the native community. High initial numbers of non-native individuals decreased survival rate of strong competitors, but increased the survival rate of weak competitors. Survival under herbivore attack was only slightly higher for plants with high defensive ability than for those with low defensive ability. Surprisingly, though, herbivory increased survival of species classified as weak competitors. Discussion High survival rates of strong non-native competitors relate to a higher probability of successful establishment than for weak competitors. However, the reduced survival of strong competitors at high initial numbers indicates a self-thinning effect, probably mediated by a strongly competitive milieu. For weak competitors, our model emphasizes positive effects of high propagule pressure known from field studies. General effects of herbivory or defence abilities on survival were not supported by our model. However, the positive effect of herbivory on survival of weak competitors indicated side effects of herbivory, such as weakening resident competitors. This might play an important role for establishment of non-natives in a new community
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