Reassessing the alternative ecosystem states proposition in the African savanna-forest domain.

Ecologists are being challenged to predict how ecosystems will respond to climate changes. According to the Multi-Colored World (MCW) hypothesis, climate impacts may not manifest because consumers such as fire and herbivory can override the influence of climate on ecosystem state. One MCW interpretation is that climate determinism fails because alternative ecosystem states (AES) are possible at some locations in climate space. We evaluated theoretical and empirical evidence for the proposition that forest and savanna are AES in Africa. We found that maps which infer where AES zones are located were contradictory. Moreover, data from longitudinal and experimental studies provide inconclusive evidence for AES. That is, although the forest-savanna AES proposition is theoretically sound, the existing evidence is not yet convincing. We conclude by making the case that the AES proposition has such fundamental consequences for designing management actions to mitigate and adapt to climate change in the savanna-forest domain that it needs a more robust evidence base before it is used to prescribe management actions.

Frightened of giants: fear responses to elephants approach that of predators.

Animals are faced with a variety of dangers or threats, which are increasing in frequency with ongoing environmental change. While our understanding of fearfulness of such dangers is growing in the context of predation and parasitism risk, the extent to which non-trophic, interspecific dangers elicit fear in animals remains less appreciated. We provide an experimental test for fear responses of savannah ungulates to a dominant and aggressive megaherbivore, the African bush elephant (Loxodonta africana), and contrast responses to an apex predator known to elicit fear in this system. Using an automated behavioural response system, we contrast vigilance and run responses of ungulates to elephant, leopard (Panthera pardus), and control (red-chested cuckoo Cuculus solitarius) vocalizations. Overall, we find that ungulates responded to elephant calls, both in terms of an increase in run and vigilance responses relative to controls. The magnitude of most behavioural responses (four of six considered) to elephant vocalizations were not significantly different than responses to leopards. These results suggest that megaherbivores can elicit strong non-trophic fear responses by ungulates and call to broaden frameworks on fear to consider dominant species, such as megaherbivores, as key modifiers of fear-induced interactions.

Limited climatic space for alternative ecosystem states in Africa.

One of the foundational premises of ecology is that climate determines ecosystems. This has been challenged by alternative ecosystem state models, which illustrate that internal ecosystem dynamics acting on the initial ecosystem state can overwhelm the influence of climate, and by observations suggesting that climate cannot reliably discriminate forest and savanna ecosystem types. Using a novel phytoclimatic transform, which estimates the ability of climate to support different types of plants, we show that climatic suitability for evergreen trees and C4 grasses are sufficient to discriminate between forest and savanna in Africa. Our findings reassert the dominant influence of climate on ecosystems and suggest that the role of feedbacks causing alternative ecosystem states is less prevalent than has been suggested.

Climate change decouples dominant tree species in African savannas.

To understand how two dominant African savanna trees will continue to respond to climate changes, we examined their regeneration niche and adult tree distributions. Specifically, we wanted to (1) determine if distributional patterns were shifting, (2) predict future distributions under different climate change scenarios and (3) evaluate the realism of predicted future distributions. We randomly placed 40 grids into 6 strata across a climate gradient in the kingdom of Eswatini. Within these grids, we sampled adult and seedling marula (Scelerocarya birrea) and knobthorn (Senegalia nigrecens) trees and used the data to model their abundance. Next, we quantified shifts in distributional patterns (e.g., expansion or contraction) by measuring the current and projected areas of overlap between seedling and adult trees. Finally, we predicted future distributions of abundance based on predicted climate conditions. We found knobthorn seedlings within a small portion of the adult distribution, suggesting it was unlikely to track climate changes. Alternatively, finding marula seedlings on and beyond one edge of the adult distribution, suggested its range would shift toward cooler climates. Predicted future distributions suggest suitable climate for both species would transition out of savannas and into grasslands. Future projections (2041-2070) appeared consistent with observed distributions of marula, but knobthorn predictions were unrealistic given the lack of evidence for regeneration outside of its current range. The idiosyncratic responses of these species to climate change are likely to decouple these keystone structures in the coming decades and are likely to have considerable cascading effects including the potential rearrangement of faunal communities.

Mass sterilization of a common palm species by elephants in Kruger National Park, South Africa.

Chronic herbivory by elephants rarely eliminates any species of woody savanna plants because these plants are typically vigorous basal resprouters after damage by fire or herbivory. In some instances, resprouting after elephant herbivory even increases stem numbers per unit area compared to protected areas. It is thus difficult to know whether an area has been severely degraded by elephant herbivory or not because although trees may be severely reduced in size, they will still be present and may even be relatively dense. By using an elephant exclosure in the Kruger National Park, South Africa, we demonstrate that this resprouting ability masks the fact that entire populations of a widespread African palm, Hyphaene petersiana, are prevented from reaching sexual maturity by chronic elephant herbivory. Besides sterilizing these palms and thus preventing their evolution and seed dispersal, the absence of the palm fruits, flowers and tall stems has other negative biodiversity impacts on their associated fauna. We suggest that to determine sustainable elephant impacts on savanna plants, conservation managers also use the reproductive condition of savanna plants rather than their presence, height or stem density.

Ecology of grazing lawns in Africa.

Grazing lawns are a distinct grassland community type, characterised by short-stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co-evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low-biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall-grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short-grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall-grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.