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.

Experimental demonstration of chaotic instability in biological nitrification.

Biological nitrification (that is, NH(3) --> NO(2)(-) --> NO(3)(-)) is a key reaction in the global nitrogen cycle (N-cycle); however, it is also known anecdotally to be unpredictable and sometimes fails inexplicably. Understanding the basis of unpredictability in nitrification is critical because the loss or impairment of this function might influence the balance of nitrogen in the environment and also has biotechnological implications. One explanation for unpredictability is the presence of chaotic behavior; however, proving such behavior from experimental data is not trivial, especially in a complex microbial community. Here, we show that chaotic behavior is central to stability in nitrification because of a fragile mutualistic relationship between ammonia-oxidizing bacteria (AOB) and nitrite-oxidizing bacteria (NOB), the two major guilds in nitrification. Three parallel chemostats containing mixed microbial communities were fed complex media for 207 days, and nitrification performance, and abundances of AOB, NOB, total bacteria and protozoa were quantified over time. Lyapunov exponent calculations, supported by surrogate data and other tests, showed that all guilds were sensitive to initial conditions, suggesting broad chaotic behavior. However, NOB were most unstable among guilds and displayed a different general pattern of instability. Further, NOB variability was maximized when AOB were most unstable, which resulted in erratic nitrification including significant NO(2)(-) accumulation. We conclude that nitrification is prone to chaotic behavior because of a fragile AOB-NOB mutualism, which must be considered in all systems that depend on this critical reaction.