Taxonomic scale dependency of Bergmann's patterns: A cross-scale comparison of hawkmoths and birds along a tropical elevational gradient

Abstract

Bergmanns rule predicts a larger body size for endothermic organisms in colder environments. The multiplicity of patterns and processes is expected because body size and temperature are two most fundamental factors on which many physiological, ecological and evolutionary processes depend, affecting all levels of biological organization, from individuals to communities. The confounding results from previous studies may be due to the differences in taxonomic (intraspecific, interspecific and community) and spatial (latitudinal vs elevational) scales. We compared Bergmanns patterns for endotherms (Aves) and ectotherms (Lepidoptera: Sphingidae) along a same 2.6 km elevational transect in the eastern Himalayas. Using a large data spanning 3,302 hawkmoths (76 morpho-species) and 15,746 birds (245 species), we compared the patterns at the intraspecific (hawkmoths only), interspecific and community scales. At the interspecific scale, we account for phylogenetic non-independence in body mass by using a heirarchical linear mixed effects model for hawkmoths, and a phylogenetic generalised least squares model for birds. We assess the importance of using abundance-weighted metrics at the community scales, after accounting for spatial auto-correlation in communities. Hawkmoths exhibited positive Bergmanns pattern at the intraspecific and abundance-weighted community scale. Intraspecific variation accounted for a substantial 33% variation at the community level. Contrary to this, birds exhibited a strong converse-Bergmanns pattern at interspecific and community scales, both with- and without-abundance. Overall, all metrics which incorporate local traits and/or species abundances show stronger correlations than when this information is lacking. The multiplicity of patterns at a single location provides the opportunity to disentangle the relative contribution of individual- and species-level processes by integrating data across multiple nested taxonomic scales for the same taxa. We suggest that future studies of Bergmanns patterns should explicitly address taxonomic- and spatial-scale dependency, with species relative abundance and intraspecific trait variation as essential ingredients especially at short elevational scales.