Sally Archibald

The Global Extent and Determinants of Savanna and Forest as Alternative Biome States

Savanna and forest are alternative states governed by fire at intermediate rainfall levels. Theoretically, fire–tree cover feedbacks can maintain savanna and forest as alternative stable states. However, the global extent of fire-driven discontinuities in tree cover is unknown, especially accounting for seasonality and soils. We use tree cover, climate, fire, and soils data sets to show that tree cover is globally discontinuous. Climate influences tree cover globally but, at intermediate rainfall (1000 to 2500 millimeters) with mild seasonality (less than 7 months), tree cover is bimodal, and only fire differentiates between savanna and forest. These may be alternative states over large areas, including parts of Amazonia and the Congo. Changes in biome distributions, whether at the cost of savanna (due to fragmentation) or forest (due to climate), will be neither smooth nor easily reversible.

Savanna vegetation-fire-climate relationships differ among continents.

Surveying Savannas Savannas are structurally similar across the three major continents where they occur, leading to the assumption that the factors controlling vegetation structure and function are broadly similar, too. Lehmann et al. (p. 548) report the results of an extensive analysis of ground-based tree abundance in savannas, sampled at more than 2000 sites in Africa, Australia, and South America. All savannas, independent of region, shared a common functional property in the way that moisture and fire regulated tree abundance. However, despite qualitative similarity in the moisture–fire–tree-biomass relationships among continents, key quantitative differences exist among the three regions, presumably as a result of unique evolutionary histories and climatic domains. Evolution cannot be overlooked when aiming to predict the potential global impacts on savanna dynamics in a warming world. Ecologists have long sought to understand the factors controlling the structure of savanna vegetation. Using data from 2154 sites in savannas across Africa, Australia, and South America, we found that increasing moisture availability drives increases in fire and tree basal area, whereas fire reduces tree basal area. However, among continents, the magnitude of these effects varied substantially, so that a single model cannot adequately represent savanna woody biomass across these regions. Historical and environmental differences drive the regional variation in the functional relationships between woody vegetation, fire, and climate. These same differences will determine the regional responses of vegetation to future climates, with implications for global carbon stocks.

Defining pyromes and global syndromes of fire regimes

Fire is a ubiquitous component of the Earth system that is poorly understood. To date, a global-scale understanding of fire is largely limited to the annual extent of burning as detected by satellites. This is problematic because fire is multidimensional, and focus on a single metric belies its complexity and importance within the Earth system. To address this, we identified five key characteristics of fire regimes—size, frequency, intensity, season, and extent—and combined new and existing global datasets to represent each. We assessed how these global fire regime characteristics are related to patterns of climate, vegetation (biomes), and human activity. Cross-correlations demonstrate that only certain combinations of fire characteristics are possible, reflecting fundamental constraints in the types of fire regimes that can exist. A Bayesian clustering algorithm identified five global syndromes of fire regimes, or pyromes. Four pyromes represent distinctions between crown, litter, and grass-fueled fires, and the relationship of these to biomes and climate are not deterministic. Pyromes were partially discriminated on the basis of available moisture and rainfall seasonality. Human impacts also affected pyromes and are globally apparent as the driver of a fifth and unique pyrome that represents human-engineered modifications to fire characteristics. Differing biomes and climates may be represented within the same pyrome, implying that pathways of change in future fire regimes in response to changes in climate and human activity may be difficult to predict.

Evolution of human-driven fire regimes in Africa

Human ability to manipulate fire and the landscape has increased over evolutionary time, but the impact of this on fire regimes and consequences for biodiversity and biogeochemistry are hotly debated. Reconstructing historical changes in human-derived fire regimes empirically is challenging, but information is available on the timing of key human innovations and on current human impacts on fire; here we incorporate this knowledge into a spatially explicit fire propagation model. We explore how changes in population density, the ability to create fire, and the expansion of agropastoralism altered the extent and seasonal distribution of fire as modern humans arose and spread through Africa. Much emphasis has been placed on the positive effect of population density on ignition frequency, but our model suggests this is less important than changes in fire spread and connectivity that would have occurred as humans learned to light fires in the dry season and to transform the landscape through grazing and cultivation. Different landscapes show different limitations; we show that substantial human impacts on burned area would only have started ∼4,000 B.P. in open landscapes, whereas they could have altered fire regimes in closed/dissected landscapes by ∼40,000 B.P. Dry season fires have been the norm for the past 200–300 ky across all landscapes. The annual area burned in Africa probably peaked between 4 and 40 kya. These results agree with recent paleocarbon studies that suggest that the biomass burned today is less than in the recent past in subtropical countries.

Southern African fire regimes as revealed by remote sensing

Here we integrate spatial information on annual burnt area, fire frequency, fire seasonality, fire radiative power and fire size distributions to produce an integrated picture of fire regimes in southern Africa. The regional patterns are related to gradients of environmental and human controls of fire, and compared with findings from other grass-fuelled fire systems on the globe. The fire regime differs across a gradient of human land use intensity, and can be explained by the differential effect of humans on ignition frequencies and fire spread. Contrary to findings in the savannas of Australia, there is no obvious increase in fire size or fire intensity from the early to the late fire season in southern Africa, presumably because patterns of fire ignition are very different. Similarly, the importance of very large fires in driving the total annual area burnt is not obvious in southern Africa. These results point to the substantial effect that human activities can have on fire in a system with high rural population densities and active fire management. Not all aspects of a fire regime are equally impacted by people: fire-return time and fire radiative power show less response to human activities than fire size and annual burned area.

A continent-wide assessment of the form and intensity of large mammal herbivory in Africa.

How herbivores affect ecosystems Abiotic forces, such as fire and water, have powerful effects on ecosystem structure and function. Animals that eat plants also have strong effects in natural systems, but their impacts are harder to assess. Hempson et al. measured how herbivores affect vegetation across Africa (see the Perspective by Gill). Four distinct herbivory regimes emerge from the analysis, characterized by forest antelopes, arid-region gazelles, high-diversity savannah fauna, and bulk feeders (such as elephants), which have had equivalent impact to those of fire and water on shaping ecosystems. Science, this issue p. 1056; see also p. 1036 Census data reveal the ways that large herbivores impact vegetation and ecosystems across a continent. [Also see Perspective by Gill] Megafaunal extinctions and a lack of suitable remote sensing technology impede our understanding of both the ecological legacy and current impacts of large mammal herbivores in the Earth system. To address this, we reconstructed the form and intensity of herbivory pressure across sub-Saharan Africa ~1000 years ago. Specifically, we modeled and mapped species-level biomass for 92 large mammal herbivores using census data, species distributions, and environmental covariates. Trait-based classifications of these species into herbivore functional types, and analyses of their biomass surfaces, reveal four ecologically distinct continental-scale herbivory regimes, characterized by internally similar forms and intensities of herbivory pressure. Associations between herbivory regimes, fire prevalence, soil nutrient status, and rainfall provide important insights into African ecology and pave the way for integrating herbivores into global-scale studies.

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.

Managing the human component of fire regimes: lessons from Africa

Human impacts on fire regimes accumulated slowly with the evolution of modern humans able to ignite fires and manipulate landscapes. Today, myriad voices aim to influence fire in grassy ecosystems to different ends, and this is complicated by a colonial past focused on suppressing fire and preventing human ignitions. Here, I review available evidence on the impacts of people on various fire characteristics such as the number and size of fires, fire intensity, fire frequency and seasonality of fire in African grassy ecosystems, with the intention of focusing the debate and identifying areas of uncertainty. Humans alter seasonal patterns of fire in grassy systems but tend to decrease total fire emissions: livestock have replaced fire as the dominant consumer in many parts of Africa, and fragmented landscapes reduce area burned. Humans alter the season and time of day when fires occur, with important implications for fire intensity, tree–grass dynamics and greenhouse gas (GHG) emissions. Late season fires are more common when fire is banned or illegal: these later fires are far more intense but emit fewer GHGs. The types of fires which preserve human livelihoods and biodiversity are not always aligned with the goal of reducing GHG concentrations. Current fire management challenges therefore involve balancing the needs of a large rural population against national and global perspectives on the desirability of different types of fire, but this cannot happen unless the interests of all parties are equally represented. In the future, Africa is expected to urbanize and land use to intensify, which will imply different trajectories for the continents fire regimes. This article is part of the themed issue ‘The interaction of fire and mankind.

Natural Hazards in a Changing World: A Case for Ecosystem-Based Management

Communities worldwide are increasingly affected by natural hazards such as floods, droughts, wildfires and storm-waves. However, the causes of these increases remain underexplored, often attributed to climate changes or changes in the patterns of human exposure. This paper aims to quantify the effect of climate change, as well as land cover change, on a suite of natural hazards. Changes to four natural hazards (floods, droughts, wildfires and storm-waves) were investigated through scenario-based models using land cover and climate change drivers as inputs. Findings showed that human-induced land cover changes are likely to increase natural hazards, in some cases quite substantially. Of the drivers explored, the uncontrolled spread of invasive alien trees was estimated to halve the monthly flows experienced during extremely dry periods, and also to double fire intensities. Changes to plantation forestry management shifted the 1∶100 year flood event to a 1∶80 year return period in the most extreme scenario. Severe 1∶100 year storm-waves were estimated to occur on an annual basis with only modest human-induced coastal hardening, predominantly from removal of coastal foredunes and infrastructure development. This study suggests that through appropriate land use management (e.g. clearing invasive alien trees, re-vegetating clear-felled forests, and restoring coastal foredunes), it would be possible to reduce the impacts of natural hazards to a large degree. It also highlights the value of intact and well-managed landscapes and their role in reducing the probabilities and impacts of extreme climate events.

Competing consumers: contrasting the patterns and impacts of fire and mammalian herbivory in Africa

Fire and herbivory are the two consumers of above-ground biomass globally. They have contrasting impacts as they differ in terms of selectivity and temporal occurrence. Here, we integrate continental-scale data on fire and herbivory in Africa to explore (i) how environmental drivers constrain these two consumers and (ii) the degree to which each consumer affects the other. Environments conducive to mammalian herbivory are not necessarily the same as those conducive to fire, although their spheres of influence do overlap—especially in grassy ecosystems which are known for their frequent fires and abundance of large mammalian herbivores. Interactions between fire and herbivory can be competitive, facultative or antagonistic, and we explore this with reference to the potential for alternative ecosystem states. Although fire removes orders of magnitude more biomass than herbivory their methane emissions are very similar, and in the past, herbivores probably emitted more methane than fire. We contrast the type of herbivory and fire in different ecosystems to define ‘consumer-realms’. This article is part of the themed issue ‘Tropical grassy biomes: linking ecology, human use and conservation’.