1. Within-clade allometric relationships represent standard laws of scaling between energy and size, and their outliers provide new avenues for physiological and ecological research. According to the metabolic-level boundaries hypothesis, metabolic rates as a function of mass are expected to scale closer to 0.67 when driven by surface-related processes (e.g. heat or water flux), while volume-related processes (e.g. activity) generate slopes closer to one. 2. In birds, daily energy expenditure (DEE) scales with body mass (M) in the relationship log DEE = 2.35 + 0.68 × log M , consistent with surface-level processes driving the relationship. However, taxon-specific patterns differ from the scaling slope of all birds. 3. Hummingbirds have the highest mass-specific metabolic rates among all vertebrates. Previous studies on a few hummingbird species, without accounting for the phylogeny, estimated that the DEE-body mass relationship for hummingbirds was log DEE = 1.72 + 1.21 × log M . In Contrast to the theoretical expectations, this slope >1 indicates that larger hummingbirds are less metabolically efficient than smaller hummingbirds.4. We collected DEE and mass data for 12 hummingbird species, which, combined with published data, represented 17 hummingbird species in eight of nine hummingbird clades over a sixfold size range of body size (2.7-17.5 g).
5.After accounting for phylogenetic relatedness, we found DEE scales with body mass as log DEE = 2.04 + 0.95 × log M . This slope of 0.95 is lower than previously estimated for hummingbirds, but much higher than the slope for all birds (0.68). The high slopes of torpor, hovering and flight potentially explain the high interspecific DEE slope for hummingbirds compared to other endotherms. K E Y W O R D S birds, body mass, daily energy expenditure, doubly labelled water, field metabolic rate, metabolic-level boundaries hypothesis, scaling, tropics 1 | INTRODUC TI ON Allometric scaling relationships describe how traits scale with one another (e.g. metabolic rate with body size). These relationships have intrigued scientists for almost two centuries because of their potential for explaining how metabolic processes constrain the range and proportions of body sizes (Sarrus & Rameaux, 1839). Scaling patterns can be compared among individuals, species or multispecies | 1255 Journal of Animal Ecology SHANKAR et Al.