An Interspecific Assessment of Bergmann’s Rule in Tenebrionid Beetles (Coleoptera, Tenebrionidae) along an Elevation Gradient

In endotherms, body size tends to increase with elevation and latitude (i.e., with decreasing temperatures) (Bergmann's rule). These patterns are explained in terms of heat balance since larger animals need to produce less heat relative to their size to maintain stable body temperatures. In ectotherms like most insects, where this mechanism cannot operate, a reverse pattern is frequently observed, as a higher surface area-to-volume ratio in colder climates may allow for more rapid heating and cooling. However, patterns of increasing body size with decreasing temperatures can also be observed in ectotherms if selection for more stable internal temperatures leads to smaller surface area-to-volume ratios. Data on tenebrionids from Latium (Central Italy) were used to model elevational variations in average values of body size (total length, mass and volume) and surface area-to-volume ratio. Analyses were performed by considering the whole fauna and two ecological groups separately: ground-dwelling species (geophilous) and arboreal (xylophilous) species. The surface area-to-volume ratios declined with increasing elevation in all cases, indicating that the need for heat conservation is more important than rapid heating and cooling. However, in xylophilous species (which typically live under bark), body size increased with increasing elevation, and in geophilous species, an opposite pattern was observed up to about 1000 m, followed by an increasing pattern. This suggests that a reduction in resource availability with elevation limits body size in geophilous species up to a certain elevation but not in xylophilopus species, which benefit from more climatically stable conditions and constant resources and need energy for overwintering.