Fire spread and the issue of community-level selection in the evolution of flammability.

Whether plants can evolve to promote flammability is controversial. Ecologically, fire only spreads in landscapes when many plants are flammable, but collective behaviours among large groups are difficult to evolve at the individual level. Here, we formulate a model that examines how flammability can spread from rarity, combining individual-level costs and payoffs of flammability with landscape-level fire spread, sufficiently generic to analogize flammability among grasses, Mediterranean systems, and others. We found that fire-prone and fire-suppressing landscapes, composed of flammable and non-flammable plants, respectively, were alternatively stable in some environments, and flammability therefore only increased from rarity in environments when fire-proneness was the only stable state. Thus, fire-vegetation feedbacks alone probably did not drive the evolution and spread of flammability. However, evolution of flammability did promote fire-proneness in temporally and spatially heterogeneous environments: when flammable plants already occupied some substantial fraction of a fire-prone landscape, a positive feedback with fire could maintain flammability in a decreasingly favourable environment, and fire feedbacks could expand the distribution of flammability traits from fire-prone into fire-suppressing areas in a heterogeneous landscape. Thus, fire feedbacks could potentially have promoted the widespread invasion and persistence of flammability traits to their current widespread prominence.