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

Abstract

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.

Post-fire reproduction of herbs at a savanna-gallery forest boundary in Distrito Federal, Brazil

“…Fires in gallery forest generally cause extensive topkill to small trees (Hoffmann et al, 2009) and leaf-fall to larger diameter trees (personal observation), allowing increased light to the forest floor. Thus, we might expect that this higher light availability would stimulate understory species to reproduce.…”

Flammable biomes dominated by eucalypts originated at the Cretaceous–Palaeogene boundary

“…Consequently, in non-epicormic resprouters, the woody biomass decays and the stored carbon is released after the fire. Such trees either resprout from the base or are killed and regenerate from seed 35,36 . They recover more slowly than eucalypts after fire 37 and are forced to accumulate aboveground wood from scratch.…”

How do small savanna trees avoid stem mortality by fire? The roles of stem diameter, height and bark thickness

“…Although bark thickness is implicated in fire resilience, it has received much less attention as an explanation for the general fire resilience of savanna trees than either the escape diameter or escape height hypotheses (cf. Gignoux et al 1997, Pinard and Huffman 1997, Nefabas and Gambiza 2007, but see Hoffmann et al 2009). …”