Spatial and temporal variability in Acacia population dynamics

Includes bibliographical references (p. 130-141).Variability in fire, herbivory, and climate facilitate the coexistence of trees and grasses in savannas and impact upon savanna structure, which also varies substantially both spatially and temporally. These features can shape savannas at an ecosystem and even at a global scale, but mechanisms for the effects of fire, herbivory, and climate variability on tree cover are often demographic at the tree population level. Sapling growth in particular has repeatedly been shown to be the limiting step, or 'bottleneck', in the establishment of trees in savannas. I set out to investigate how spatial and temporal variability in fire, herbivory, and climate shape population dynamics of a suite of common African savanna trees, the Acacia, in a landscape context. I carried out my field work in Hluhluwe iMfolozi Park in K waZulu Natal, South Africa, during 2006 and 2007. Fire, herbivory, and the grass layer were primary determinants of distributions and co-occurrence of Acacia species

Dispersal limitation and fire feedbacks maintain mesic savannas in Madagascar

Madagascar is regarded by some as one of the most degraded landscapes on Earth, with estimates suggesting that 90% of forests have been lost to indigenous Tavy farming. However, the extent of this degradation has been challenged: paleoecological data, phylogeographic analysis, and species richness indicate that pyrogenic savannas in central Madagascar predate human arrival, even though rainfall is sufficient to allow forest expansion into central Madagascar. These observations raise a question—if savannas in Madagascar are not anthropogenic, how then are they maintained in regions where the climate can support forest? Observation reveals that the savanna–forest boundary coincides with a dispersal barrier—the escarpment of the Central Plateau. Using a stepping-stone model, we show that in a limited dispersal landscape, a stable savanna–forest boundary can form because of fire–vegetation feedbacks. This phenomenon, referred to as range pinning, could explain why eastern lowland forests have not expanded into the mesic savannas of the Central Highlands. This work challenges the view that highland savannas in Madagascar are derived by human-lit fires and, more importantly, suggests that partial dispersal barriers and strong nonlinear feedbacks can pin biogeographical boundaries over a wide range of environmental conditions, providing a temporary buffer against climate change

Disease and fire interact to influence transitions between savanna-forest ecosystems over a multi-decadal experiment

Global change is shifting disturbance regimes that may rapidly change ecosystems, sometimes causing ecosystems to shift between states. Interactions between disturbances such as fire and disease could have especially severe effects, but experimental tests of multi-decadal changes in disturbance regimes are rare. Here, we surveyed vegetation for 35 years in a 54-year fire frequency experiment in a temperate oak savanna-forest ecotone that experienced a recent outbreak of oak wilt. Different fire regimes determined whether plots were savanna or forest by regulating tree abundance (r(2) = 0.70), but disease rapidly reversed the effect of fire exclusion, increasing mortality by 765% in unburned forests, but causing relatively minor changes in frequently burned savannas. Model simulations demonstrated that disease caused unburned forests to transition towards a unique woodland that was prone to transition to savanna if fire was reintroduced. Consequently, disease-fire interactions could shift ecosystem resilience and biome boundaries as pathogen distributions change

Historical and future global burned area with changing climate and human demography

Wildfires influence terrestrial carbon cycling and represent a safety risk, and yet a process-based understanding of their frequency and spatial distributions remains elusive. We combine satellite-based observations with an enhanced dynamic global vegetation model to make regionally resolved global assessments of burned area (BA) responses to changing climate, derived from 34 Earth system models and human demographics for 1860–2100. Limited by climate and socioeconomics, recent BA has decreased, especially in central South America and mesic African savannas. However, future simulations predict increasing BA due to changing climate, rapid population density growth, and urbanization. BA increases are especially notable at high latitudes, due to accelerated warming, and over the tropics and subtropics, due to drying and human ignitions. Conversely, rapid urbanization also limits BA via enhanced fire suppression in the immediate vicinity of settlements, offsetting the potential for dramatic future increases, depending on warming extent. Our analysis provides further insight into regional and global BA trends, highlighting the importance of including human demographic change in models for wildfire under changing climate

The role of browsers in maintaining the openness of savanna grazing lawns

In savannas, ruminant herbivores can have divergent impacts on tree recruitment and subsequent woody cover. Whereas heavy grazing by cattle results in woody thickening, intensive grazing by wildlife instead tends to be associated with lower woody cover. To disentangle why woody cover is low in areas heavily grazed by wildlife, we investigated (a) whether grazing lawns attract indigenous mammalian browsers, and if a preference for short-grass habitat decreases with browser body mass as predator susceptibility decreases; and (b) whether browser attraction to grazing lawns translates into the suppression of woody plants, including seedlings and saplings, thus maintaining the openness of heavily grazed short-grass areas. In Kruger National Park, South Africa, we contrasted browser abundance (using dung counts) on grazing lawns and on low-herbivory sites characterised by tall grass. Additionally, a herbivore exclosure experiment was set up to investigate the combined impact of browser removal and grass height habitat type on seedling survival and sapling growth of a dominant woody plant species. Finally, in Hluhluwe-iMfolozi Park (HiP), we examined the long-term (10 years) impact of browser removal on the growth rates of a range of woody species, monitored across ten sites along a gradient of herbivory ranging from grazing lawn to tall grass. Steenbok and impala selected short- over tall grass as preferred browsing sites, while elephant preferred tall grass. Browser abundance on short grass decreased with browser body mass, indicating that predator avoidance might be a key factor driving mesoherbivores to utilise grazing lawns. Seedling survival was lowest on grazing lawns when browsers were present, with mortality occurring in two out of every three seedlings. Similarly, sapling growth was lowest on grazing lawns, although browser removal had no significant effect. Evidence for increased browser impact on grazing lawns was clearest from our long-term herbivore exclosure experiment in HiP, which demonstrated that browsers strongly modify the growth rates of woody plants in short-grass habitats. Synthesis. These results provide support for the hypothesis that browsers, particularly browsing mesoherbivores and mixed feeders, are attracted to short-grass habitats, and that they help maintain grazing lawn openness by suppressing seedling survival and woody plant growth where grass is kept short by grazers.SUPPORTING INFORMATION: Table S1. Proportion short grass calculated as the mean proportion of grass height below 10 cm, recorded at ten sites in HiP (Hluhluwe and iMfolozi Game Reserves). The mean number of grazer dung piles per year (species: buffalo, blue wildebeest, impala, warthog, white rhino and zebra) are shown for each site. Values in brackets represent standard errors.Figure S1. The effect of grass height and exclosure status (fenced vs. unfenced) on tree height gain (mean ± SE). Browsing impact on tree height gain was higher in areas with short grass i.e. browsers utilised trees in short-grass ecosystems more than in tall-grass ecosystems. Results of linear regression analyses are displayed for the unfenced treatment of two dominant woody plant species: a) D. cinerea (R2 = 0.38, p = 0.05) and b) A. nilotica (R2 = 0.44, p = 0.11), and for two woody plant functional types: c) fine-leaved (R2 = 0.33, p = 0.09) and d) broad-leaved (R2 = 0.60, p = 0.02), across 10 sites in Hluhluwe-iMfolozi Park for the period 2000-2009.DATA AVAILABILITY STATEMENT: Data are available from the Dryad Digital Repository https://doi-org.uplib.idm.oclc.org/10.5061/dryad.76hdr7st3 (Voysey et al., 2020).The USAID/NAS program ‘Partnerships for Enhanced Engagement in Research' (sub-grant 2000004946, Cycle 3) and the South African National Research Foundation, Department of Science and Technology, Innovation and Priority Research Masters Scholarship.http://www.wileyonlinelibrary.com/journal/jechj2022Plant Production and Soil Scienc

Soil carbon in tropical savannas mostly derived from grasses

Tropical savannas have been increasingly targeted for carbon sequestration by afforestation, assuming large gains in soil organic carbon (SOC) with increasing tree cover. Because savanna SOC is also derived from grasses, this assumption may not reflect real changes in SOC under afforestation. However, the exact contribution of grasses to SOC and the changes in SOC with increasing tree cover remain poorly understood. Here we combine a case study from Kruger National Park, South Africa, with data synthesized from tropical savannas globally to show that grass-derived carbon constitutes more than half of total SOC to a soil depth of 1 m, even in soils directly under trees. The largest SOC concentrations were associated with the largest grass contributions (>70% of total SOC). Across the tropics, SOC concentration was not explained by tree cover. Both SOC gain and loss were observed following increasing tree cover, and on average SOC storage within a 1-m profile only increased by 6% (s.e. = 4%, n = 44). These results underscore the substantial contribution of grasses to SOC and the considerable uncertainty in SOC responses to increasing tree cover across tropical savannas

Disease and fire interact to influence transitions between savanna-forest ecosystems over a multi-decadal experiment.

Global change is shifting disturbance regimes that may rapidly change ecosystems, sometimes causing ecosystems to shift between states. Interactions between disturbances such as fire and disease could have especially severe effects, but experimental tests of multi-decadal changes in disturbance regimes are rare. Here, we surveyed vegetation for 35 years in a 54-year fire frequency experiment in a temperate oak savanna-forest ecotone that experienced a recent outbreak of oak wilt. Different fire regimes determined whether plots were savanna or forest by regulating tree abundance (r2  = 0.70), but disease rapidly reversed the effect of fire exclusion, increasing mortality by 765% in unburned forests, but causing relatively minor changes in frequently burned savannas. Model simulations demonstrated that disease caused unburned forests to transition towards a unique woodland that was prone to transition to savanna if fire was reintroduced. Consequently, disease-fire interactions could shift ecosystem resilience and biome boundaries as pathogen distributions change

Decadal changes in fire frequencies shift tree communities and functional traits

Global change has resulted in chronic shifts in fire regimes. Variability in the sensitivity of tree communities to multi-decadal changes in fire regimes is critical to anticipating shifts in ecosystem structure and function, yet remains poorly understood. Here, we address the overall effects of fire on tree communities and the factors controlling their sensitivity in 29 sites that experienced multi-decadal alterations in fire frequencies in savanna and forest ecosystems across tropical and temperate regions. Fire had a strong overall effect on tree communities, with an average fire frequency (one fire every three years) reducing stem density by 48% and basal area by 53% after 50 years, relative to unburned plots. The largest changes occurred in savanna ecosystems and in sites with strong wet seasons or strong dry seasons, pointing to fire characteristics and species composition as important. Analyses of functional traits highlighted the impact of fire-driven changes in soil nutrients because frequent burning favoured trees with low biomass nitrogen and phosphorus content, and with more efficient nitrogen acquisition through ectomycorrhizal symbioses. Taken together, the response of trees to altered fire frequencies depends both on climatic and vegetation determinants of fire behaviour and tree growth, and the coupling between fire-driven nutrient losses and plant traits

Precipitation gradients drive high tree species turnover in the woodlands of eastern and southern Africa

Savannas cover one-fifth of the Earth's surface, harbour substantial biodiversity, and provide a broad range of ecosystem services to hundreds of millions of people. The community composition of trees in tropical moist forests varies with climate, but whether the same processes structure communities in disturbance-driven savannas remains relatively unknown. We investigate how biodiversity is structured over large environmental and disturbance gradients in woodlands of eastern and southern Africa. We use tree inventory data from the Socio-Ecological Observatory for Studying African Woodlands (SEOSAW) network, covering 755 ha in a total of 6780 plots across nine countries of eastern and southern Africa, to investigate how alpha, beta, and phylogenetic diversity varies across environmental and disturbance gradients. We find strong climate-richness patterns, with precipitation playing a primary role in determining patterns of tree richness and high turnover across these savannas. Savannas with greater rainfall contain more tree species, suggesting that low water availability places distributional limits on species, creating the observed climate-richness patterns. Both fire and herbivory have minimal effects on tree diversity, despite their role in determining savanna distribution and structure. High turnover of tree species, genera, and families is similar to turnover in seasonally dry tropical forests of the Americas, suggesting this is a feature of semiarid tree floras. The greater richness and phylogenetic diversity of wetter plots shows that broad-scale ecological patterns apply to disturbance-driven savanna systems. High taxonomic turnover suggests that savannas from across the regional rainfall gradient should be protected if we are to maximise the conservation of unique tree communities