Flammability across the gymnosperm phylogeny: the importance of litter particle size

Fire is important to climate, element cycles and plant communities, with many fires spreading via surface litter. The influence of species on the spread of surface fire is mediated by their traits which, after senescence and abscission, have 'afterlife' effects on litter flammability. We hypothesized that differences in litter flammability among gymnosperms are determined by litter particle size effects on litterbed packing. We performed a mesocosm fire experiment comparing 39 phylogenetically wide-ranging gymnosperms, followed by litter size and shape manipulations on two chemically contrasting species, to isolate the underlying mechanism. The first-order control on litter flammability was, indeed, litter particle size in both experiments. Most gymnosperms were highly flammable, but a prominent exception was the non-Pinus Pinaceae, in which small leaves abscised singly produced dense, non-flammable litterbeds. There are two important implications: first, ecosystems dominated by gymnosperms that drop small leaves separately will develop dense litter layers, which will be less prone to and inhibit the spread of surface litter fire. Second, some of the needle-leaved species previously considered to be flammable in single-leaf experiments were among the least flammable in litter fuel beds, highlighting the role of the litter traits of species in affecting surface fire regimes.