Resilience of tropical tree cover: The roles of climate, fire, and herbivory.

Fires and herbivores shape tropical vegetation structure, but their effects on the stability of tree cover in different climates remain elusive. Here, we integrate empirical and theoretical approaches to determine the effects of climate on fire- and herbivore-driven forest-savanna shifts. We analyzed time series of remotely sensed tree cover and fire observations with estimates of herbivore pressure across the tropics to quantify the fire-tree cover and herbivore-tree cover feedbacks along climatic gradients. From these empirical results, we developed a spatially explicit, stochastic fire-vegetation model that accounts for herbivore pressure. We find emergent alternative stable states in tree cover with hysteresis across rainfall conditions. Whereas the herbivore-tree cover feedback can maintain low tree cover below 1,100 mm mean annual rainfall, the fire-tree cover feedback can maintain low tree cover at higher rainfall levels. Interestingly, the rainfall range where fire-driven alternative vegetation states can be found depends strongly on rainfall variability. Both higher seasonal and interannual variability in rainfall increase fire frequency, but only seasonality expands the distribution of fire-maintained savannas into wetter climates. The strength of the fire-tree cover feedback depends on the spatial configuration of tree cover: Landscapes with clustered low tree-cover areas are more susceptible to cross a tipping point of fire-driven forest loss than landscapes with scattered deforested patches. Our study shows how feedbacks involving fire, herbivores, and the spatial structure of tree cover explain the resilience of tree cover across climates.

Testing for criticality in ecosystem dynamics: the case of Amazonian rainforest and savanna fire.

We test for two critical phenomena in Amazonian ecosystems: self-organized criticality (SOC) and critical transitions. SOC is often presented in the complex systems literature as a general explanation for scale invariance in nature. In particular, this mechanism is claimed to underlie the macroscopic structure and dynamics of terrestrial ecosystems. These would be inextricably linked to the action of fire, which is conceived as an endogenous ecological process. We show that Amazonian savanna fires display the scale-invariant features characteristic of SOC but do not display SOC. The same is true in Amazonian rainforests subject to moderate drought. These findings prove that there are other causes of scale invariance in ecosystems. In contrast, we do find evidence of a critical transition to a megafire regime under extreme drought in rainforests; this phenomenon is likely to determine the time scale of a possible loss of Amazonian rainforest caused by climate change.

Biodiversity policy beyond economic growth.

Increasing evidence-synthesized in this paper-shows that economic growth contributes to biodiversity loss via greater resource consumption and higher emissions. Nonetheless, a review of international biodiversity and sustainability policies shows that the majority advocate economic growth. Since improvements in resource use efficiency have so far not allowed for absolute global reductions in resource use and pollution, we question the support for economic growth in these policies, where inadequate attention is paid to the question of how growth can be decoupled from biodiversity loss. Drawing on the literature about alternatives to economic growth, we explore this contradiction and suggest ways forward to halt global biodiversity decline. These include policy proposals to move beyond the growth paradigm while enhancing overall prosperity, which can be implemented by combining top-down and bottom-up governance across scales. Finally, we call the attention of researchers and policy makers to two immediate steps: acknowledge the conflict between economic growth and biodiversity conservation in future policies; and explore socioeconomic trajectories beyond economic growth in the next generation of biodiversity scenarios.

Fire forbids fifty-fifty forest.

Recent studies have interpreted patterns of remotely sensed tree cover as evidence that forest with intermediate tree cover might be unstable in the tropics, as it will tip into either a closed forest or a more open savanna state. Here we show that across all continents the frequency of wildfires rises sharply as tree cover falls below ~40%. Using a simple empirical model, we hypothesize that the steepness of this pattern causes intermediate tree cover (30‒60%) to be unstable for a broad range of assumptions on tree growth and fire-driven mortality. We show that across all continents, observed frequency distributions of tropical tree cover are consistent with this hypothesis. We argue that percolation of fire through an open landscape may explain the remarkably universal rise of fire frequency around a critical tree cover, but we show that simple percolation models cannot predict the actual threshold quantitatively. The fire-driven instability of intermediate states implies that tree cover will not change smoothly with climate or other stressors and shifts between closed forest and a state of low tree cover will likely tend to be relatively sharp and difficult to reverse.

The maximum entropy formalism and the idiosyncratic theory of biodiversity.

Why does the neutral theory, which is based on unrealistic assumptions, predict diversity patterns so accurately? Answering questions like this requires a radical change in the way we tackle them. The large number of degrees of freedom of ecosystems pose a fundamental obstacle to mechanistic modelling. However, there are tools of statistical physics, such as the maximum entropy formalism (MaxEnt), that allow transcending particular models to simultaneously work with immense families of models with different rules and parameters, sharing only well-established features. We applied MaxEnt allowing species to be ecologically idiosyncratic, instead of constraining them to be equivalent as the neutral theory does. The answer we found is that neutral models are just a subset of the majority of plausible models that lead to the same patterns. Small variations in these patterns naturally lead to the main classical species abundance distributions, which are thus unified in a single framework.

Rare, Intense, Big fires dominate the global tropics under drier conditions.

Wildfires burn large parts of the tropics every year, shaping ecosystem structure and functioning. Yet the complex interplay between climate, vegetation and human factors that drives fire dynamics is still poorly understood. Here we show that on all continents, except Australia, tropical fire regimes change drastically as mean annual precipitation falls below 550 mm. While the frequency of fires decreases below this threshold, the size and intensity of wildfires rise sharply. This transition to a regime of Rare-Intense-Big fires (RIB-fires) corresponds to the relative disappearance of trees from the landscape. Most dry regions on the globe are projected to become substantially drier under global warming. Our findings suggest a global zone where this drying may have important implications for fire risks to society and ecosystem functioning.

Comparison of two views of maximum entropy in biodiversity: Frank (2011) and Pueyo et al. (2007).

An increasing number of authors agree in that the maximum entropy principle (MaxEnt) is essential for the understanding of macroecological patterns. However, there are subtle but crucial differences among the approaches by several of these authors. This poses a major obstacle for anyone interested in applying the methodology of MaxEnt in this context. In a recent publication, Frank (2011) gives some arguments why his own approach would represent an improvement as compared to the earlier paper by Pueyo et al. (2007) and also to the views by Edwin T. Jaynes, who first formulated MaxEnt in the context of statistical physics. Here I show that his criticisms are flawed and that there are fundamental reasons to prefer the original approach.

Comment on "A keystone mutualism drives pattern in a power function".

Vandermeer and Perfecto (Reports, 17 February 2006, p. 1000) maintain that a mutualist ant disrupts the power law distribution of scale insect abundances. However, reanalysis of the data reveals that ants cause an increase in the range of the power law and modify its exponent. We present a tentative, but more realistic, model that is suitable for quantitative predictions.

Helical Lévy walks: adjusting searching statistics to resource availability in microzooplankton.

The searching trajectories of different animals can be described with a broad class of flight length (lj) distributions with P(lj) = lj-mu. Theoretical studies have shown that changes in these distributions (i.e., different mu values) are key to optimizing the long-term encounter statistics under certain searcher-resource scenarios. In particular, they predict the advantage of Lévy searching (mu approximately 2) over Brownian motion (mu > or = 3) for low-prey-density scenarios. Here, we present experimental evidence of predicted optimal changes in the flight-time distribution of a predator's walk in response to gradual density changes of its moving prey. Flight times of the dinoflagellate Oxyrrhis marina switched from an exponential to an inverse square power-law distribution when the prey (Rhodomonas sp.) decreased in abundance. Concomitantly, amplitude and frequency of the short-term helical path increased. The specific biological mechanisms involved in these searching behavioral changes are discussed. We suggest that, in a three-dimensional environment, a stronger helical component combined with a Lévy walk searching strategy enhances predator's encounter rates. Our results support the idea of universality of the statistical laws in optimal searching processes despite variations in the biological details of the organisms.