Summary

1 Savanna ecosystems are characterized by the codominance of two different life forms: grasses and trees. An operational understanding of how these two different life forms coexist is essential for understanding savanna function and for predicting its response to future environmental change.



2 The existing model, which proposes that grasses and trees coexist by a separation of rooting niches, is not supported by recent empirical investigations. Our aim was to define an alternative mechanism of grass–tree coexistence in savanna ecosystems. The model we have built concentrates on life history–disturbance interactions between grasses and trees.



3 The model demonstrates coexistence for a wide range of environmental conditions, and exhibits long periods of slow decline in adult tree numbers interspersed with relatively infrequent recruitment events. Recruitment is controlled by rainfall, which limits seedling establishment, and fire, which prevents recruitment into adult size classes. Decline in adult tree numbers is the result of continuing, but low levels, of adult mortality. Both aspects of the dynamics are consistent with an established non-equilibrium mechanism of coexistence (the storage effect).



4 A sensitivity analysis indicated that data on tree resprouting ability, stem growth rates and the relationship between seedling establishment and wet season drought are essential for predicting both the range of conditions for which coexistence is possible and the response of savanna ecosystems to environmental change.



5 Our analysis suggests that understanding grass–tree interactions in savanna requires consideration of the long-term effects of life history–disturbance interactions on demography, rather than the fine-scale effects of resource competition on physiological performance.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-2745.2000.00435.x

Fire, resprouting and variability: a recipe for grass–tree coexistence in savanna

Fire management for biodiversity conservation: Key research questions and our capacity to answer them

Tropical savanna and forest are recognized to represent alternate stable states, primarily determined by feedbacks with fire. Vegetation-fire dynamics in each of these vegetation types are largely determined by the influence of the vegetation on fire behavior, as well as the effects of fire behavior on tree mortality, topkill (defined here as complete death of the aerial biomass, regardless of whether the plant recovers by resprouting), and rate of growth of resprouts. We studied the effect of fire on three savanna-forest boundaries in central Brazil. Fire intensity was greater in savanna than forest, as inferred by a twofold greater height of stem charring. Despite lower fire intensity, forest tree species exhibited higher rates of topkill, which was best explained by their thinner bark, relative to savanna species. Following topkill, there was no tendency for sprouts of savanna trees to grow faster than those of forest species, contrary to expectations, nor was whole-plant mortality higher in forest than in savanna. This contrasts with observations of high rates of postburn mortality in many other tropical forests. The low tree mortality in these transitional forests suggests that the dynamic of these natural savanna-forest boundaries is fundamentally different from that of forest boundaries originating from deforestation in the humid tropics. The forests studied here appear to be much more resilient to occasional incursion of fire from the savanna, despite being unable to invade frequently burned savanna. The thin bark of forest species makes them particularly susceptible to the "fire trap," whereby repeated topkill of small trees prevents recruitment into adult size classes. Rapid growth will be particularly important for forest species to escape the fire trap, so we predict that, where fire is frequent, forests should be restricted to high-resource sites. Here, Mg2+ and Ca2+ concentrations had particularly strong effects on postburn growth rates, suggesting that these elements may most strongly limit the distribution of forest in these fire-prone savannas.

Tree topkill, not mortality, governs the dynamics of savanna–forest boundaries under frequent fire in central Brazil

One of the most complex and contentious issues in Australian ecology concerns the environmental impact of Aboriginal landscape burning. This issue is not only important for the development of a comprehensive understanding of the dynamics and evolution of the Australian biota, but is central to the formulation of appropriate strategies for the conservation of the nation's biodiversity. Ethnographic evidence leaves little doubt that Aboriginal burning played a central role in the maintenance of the landscapes subsequently colonized by Europeans. Both 19th century European colonists and anthropologists in the 20th century documented the indispensability of fire as a tool in traditional Aboriginal economies, which have aptly been described as ‘fire-stick farming’. Aborigines used fire to achieve short-term outcomes such as providing favourable habitats for herbivores or increasing the local abundance of food plants, but it is not clear whether or not Aborigines had a predictive ecological knowledge of the long-term consequences of their use of fire. A large body of ecological evidence suggests that Aboriginal burning resulted in substantial changes in the geographic range and demographic structure of many vegetation types. Aboriginal burning was important in creating habitat mosaics that favoured the abundance of some mammal species and in the maintenance of infrequently burnt habitats upon which the survival of specialized fauna depends. Aboriginal fire regimes were probably critical for the maintenance of at least one species of tree (Callitris intratropica) in the monsoon tropics.

The question of the original impact of humans on the Australian environment is fundamentally speculative because of vague, disputed time frames proposed for the waves of colonization and shifting settlement patterns of Aborigines in the late Quaternary period. There is an inherent circular argument concerning the cause and effect of climate change, vegetation change, and burning through the late Quaternary. Charcoal and pollen evidence from long sedimentary cores is ambiguous and cannot be used to demonstrate unequivocally the initial impact of Aboriginal people on the landscapes of Pleistocene Australia. The sparse available evidence does not support the hypotheses that Aboriginal burning was primarily responsible for the extinction of Pleistocene megafauna; was critical for the maintenance of habitats of small mammals that have become extinct following European colonization; initiated widespread accelerated soil erosion rates in either the Pleistocene or Holocene; or forced the evolutionary diversification of the Australian biota. Burning may have caused the extinction of some fire-sensitive species of plants and animals dependent upon infrequently burnt habitats, and it must have maintained structurally open vegetation such as grasslands and also extended the range of fire-adapted species, such as Eucalyptus, into environments climatically suitable for rain forest. Palaeoecological research concerning prior impacts of Aborigines must give way to focused studies of the role of different anthropogenic fire regimes in contemporary ecosystems that have not been destroyed by European colonization. Such research is crucial for comprehending the role of Aboriginal burning in the maintenance of Australia's unique, rich biodiversity.

The impact of Aboriginal landscape burning on the Australian biota

Provision of watering points in the Australian arid zone: a review of effects on biota

A general decline in populations of some savanna mammal species has occurred since the mid-1980s in a fairly pristine national park environment in the Australian wet-dry tropics. Terrestrial native rodents have exhibited the greatest decline and marsupials the least. During the same period, waterholes have dried up and the previous strong association of mammals with riparian vegetation has diminished.



A regional index of the level of groundwater was developed from 16 unexploited bores from across the northern half of the Northern Territory. Predicted mean groundwater values over 28 years showed a strong increase throughout the 1970s and a similar decrease throughout the 1980s. Using data since 1986, strong correlations between mammal numbers and groundwater levels for the preceding two years, and much weaker relationships with rainfall, were obtained. In turn, the groundwater levels were best correlated with an eight-year running mean of rainfall for both Darwin and Alice Springs and an eleven-year running mean for the Southern Oscillation Index. Extrapolating backward in time, cumulative pressure variation at Darwin showed two extended periods of predicted low ground-water values, 1900s–1920s and 1940s. Both periods were preceded by the last records of a number of now-extinct mammals from central Australia. A third trough in the early 1960s suggests another period of extinction in central Australia, previously unappreciated due to the lack of survey work in the 1950s. Further, by this analysis, the past 20 years has been the best period for mammals since weather records commenced.



Analysis of continental-scale distributions of mammals showed evidence of fluctuations suggesting rocky areas are important refuges for some species during periods of low groundwater levels. In contrast to those in savanna habitats, the mammals of the extensive wetland areas fluctuated in harmony with rainfall on a short-term basis except where habitat is flooded. It is the species which occur only in savanna but not also in rocky or wetland habitats which are most at risk.



A number of other species-vulnerability characteristics were identified: riparian vegetation specificity; the degree to which they fall within the critical size range (35 g-5. 5 kg); degree of semelparity; smallness of population size; and smallness of geographic range. Those savanna species with annual life histories strongly associated with riparian areas are likely to be in greatest jeopardy. The species which appear to be most vulnerable in northwestern Australia are Conilurus penicillatus, Mesembriomys gouldii, Mesembriomys macrurus, Antechinus bellus, Phascogale tapoatafa and Rattus tunneyi.



Our recent benign climatic history has coincided with the great public environmental awakening and upsurge in environmental research and management efforts. This historical accident has probably led us to overstate the negative effects of human impact and also our ability to change the course of biological history by ameliorating human impact. Our distorted view of these factors has further led us to over-emphasize research of a spatial nature to the detriment of temporal studies. We must redress this balance.

Rainfall, groundwater and refuges: Predicting extinctions of Australian tropical mammal species

The native annual Sorghum populations of the Australian wet-dry tropics are highly resilient to dry season fires. During the early wet season, however, fires that occur after the new grass population has emerged can cause catastrophic population crashes. We examined savanna plots that had been burnt in this way, and compared them with adjacent unburnt plots. We found that Sorghum densities in the burnt plots were lower on average by a factor of 10, but that some fires had reduced the density only to one-third of the unburnt plots. It is not clear whether these differences relate directly to site or seasonal factors, or to differences in the way the burning was carried out. Other vegetation components responded to the fires differently: forbs (dicotyledonous herbs) increased in cover, while perennial grasses, woody plants, and overall species richness, were not significantly affected. The amount of leaf litter declined. A population model for Sorghum based on the demography of unburnt populations predicted that they should recover from a wet season burn, taking 7–16 years to return to normal densities. However, the actual field populations did not seem to be recovering, suggesting that wet season fires not only lower densities, but may also fundamentally change population processes in these annual grasses.

Modelling the recovery of an annual savanna grass following a fire-induced crash

Fiddling while Rome burns: Nero much misunderstood. Reply to Bowman (1992)

(1993). The Importance of Pandanus spiralis to Birds. Emu - Austral Ornithology: Vol. 93, No. 1, pp. 53-58.

The Importance of Pandanus spiralis to Birds

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