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Soil-feeding species are found in 3 subfamilies of higher termites and constitute 67% of all genera. The habit, which may have evolved several times, is principally associated with lowland humid equatorial rainforests, but there are some savanna forms. Soil-feeders can generally be distinguished from wood-feeders by intestinal morphology, the stable isotope ratios of C and N, and by the higher activity of certain elements of the gut flora, notably methanogens and organisms able to ferment reduced and recalcitrant substrates, including aromatics. Soil-feeders emit more methane as free gas, but do not appear to fix N₂ in significant amounts. Organic material passing through the gut is further humified, with enrichment in total C, N and fulvic acid compared with parent soil, while humic acid is depleted. Mound materials and galleries made using faeces show enhanced cation exchange capacity, with a …