Floristic evidence for alternative biome states in tropical Africa.

The idea that tropical forest and savanna are alternative states is crucial to how we manage these biomes and predict their future under global change. Large-scale empirical evidence for alternative stable states is limited, however, and comes mostly from the multimodal distribution of structural aspects of vegetation. These approaches have been criticized, as structure alone cannot separate out wetter savannas from drier forests for example, and there are also technical challenges to mapping vegetation structure in unbiased ways. Here, we develop an alternative approach to delimit the climatic envelope of the two biomes in Africa using tree species lists gathered for a large number of forest and savanna sites distributed across the continent. Our analyses confirm extensive climatic overlap of forest and savanna, supporting the alternative stable states hypothesis for Africa, and this result is corroborated by paleoecological evidence. Further, we find the two biomes to have highly divergent tree species compositions and to represent alternative compositional states. This allowed us to classify tree species as forest vs. savanna specialists, with some generalist species that span both biomes. In conjunction with georeferenced herbarium records, we mapped the forest and savanna distributions across Africa and quantified their environmental limits, which are primarily related to precipitation and seasonality, with a secondary contribution of fire. These results are important for the ongoing efforts to restore African ecosystems, which depend on accurate biome maps to set appropriate targets for the restored states but also provide empirical evidence for broad-scale bistability.

Herbivore population regulation and resource heterogeneity in a stochastic environment.

Large-mammal herbivore populations are subject to the interaction of internal density-dependent processes and external environmental stochasticity. We disentangle these processes by linking consumer population dynamics, in a highly stochastic environment, to the availability of their key forage resource via effects on body condition and subsequent fecundity and mortality rates. Body condition and demographic rate data were obtained by monitoring 500 tagged female goats in the Richtersveld National Park, South Africa, over a three-year period. Identifying the key resource and pathway to density dependence for a population allows environmental stochasticity to be partitioned into that which has strong feedbacks to population stability, and that which does not. Our data reveal a density- dependent seasonal decline in goat body condition in response to concomitant density-dependent depletion of the dry-season forage resource. The loss in body condition reduced density-dependent pregnancy rates, litter sizes, and pre-weaning survival. Survival was lowest following the most severe dry season and for juveniles. Adult survival in the late-dry season depended on body condition in the mid-dry season. Population growth was determined by the length of the dry season and the population size in the previous year. The RNP goat population is thereby dynamically coupled primarily to its dry-season forage resource. Extreme environmental variability thus does not decouple consumer resource dynamics, in contrast to the views of nonequilibrium protagonists.