More than the sum of its parts? Assessing litter heterogeneity effects on the decomposition of litter mixtures through leaf chemistry

Different components of functional biodiversity, such as functional type richness and composition, have been reported to affect the decomposition of litter mixtures. In spite of the numerous reports of these effects, mechanisms underlying patterns of decomposition in litter mixtures are still unclear. We analyzed whether mixture decomposition was affected by: (a) the number of species in the mixture (mixture richness); and (b) the mixture's functional composition (% of fast- vs. slow-decomposing species included in the mixture). We then tested if variation between observed and expected values of decomposition in mixtures was associated to: (c) the initial litter characteristics of the component species (initial nitrogen, lignin, cellulose and hemicellulose content of litters); and (d) the chemical heterogeneity of the mixtures (variation in the same chemical compounds between the components in each mixture). When up to 5 species representing different functional types were included, both species richness and functional composition showed statistically significant non-additive, and in general positive, effects on litter mixture decomposition. The positive effect of mixture richness on decomposition did not disappear, but was much less marked, when considering mixture with slow-decomposing species only. Although the main driver of decomposition in a mixture is still the average decomposability of the component species (itself largely determined by litter quality), the species interactions in a mixture add a consistent source of variability that is worth considering when predicting the decomposability of a given mixture. We showed that a greater positive difference between observed decomposition rates and that expected from its component species alone was found in mixtures with higher mean nitrogen content and a higher heterogeneity in non-labile compounds. Our results offer quantitative proof that litter chemical heterogeneity, as well as the mean quality of the mixture, can affect the decomposability in litter mixtures.