Abstract
Bergmann’s rule, defined as the tendency for endotherms to be larger in colder environments, is a biophysical generalization of body size variation that is frequently tested along latitudinal gradients, even though latitude is only a proxy for temperature variation. We test whether variation in temperature and aridity determine avian body size conformity to Bergmann’s rule independent of latitude differences, using the ubiquitous Common Bulbul Pycnonotus barbatus, along a West African environmental gradient. We trapped 538 birds in 22 locations between latitudes 6 and 13°N in Nigeria, and estimated average body surface area to mass ratio per location. We then modelled body surface to mass ratio using general linear models, with latitude, altitude and one of 19 bioclimatic variables extracted from http://www.worldclim.org/bioclim as predictors. We sequentially dropped latitude and altitude from each model to obtain the R2 of the resultant models. Finally, we compared the R2 of univariate models, where bioclimatic variables predicted body surface area to mass ratio significantly (14 out of 19), to multivariate models including latitude, altitude and a bioclimatic variable, using the Wilcoxon matched pairs test. We found that multivariate models did not perform better than univariate models with only bioclimatic variables. Six temperature and eight precipitation variables significantly predicted variation in body surface area to mass ratio between locations; in fact, 50% (seven out of 14) of these better explained variation in body surface area to mass ratio than the multivariate models. Birds showed a larger body surface area relative to body mass ratio in hotter environments independent of latitude or altitude, which conforms to Bergmann’s rule. Yet, a combination of morphometric analyses and controlled temperature-exposure experiments is required to prove the proposed relationship between relative body surface area and thermoregulation in endotherms.
Highlights
Bergmann’s rule, defined as the tendency for endotherms to be larger in colder environments, is a well-known biophysical generalization for thermoregulation along environmental gradients (Salewski and Watt 2017)
Body surface area to mass ratio of Common Bulbuls across different environments was determined by local bioclimatic variables independent of latitude (Table 1; Figs. 3, 4)
Birds had larger body surface area to mass ratio in hotter, arid and more seasonal environments compared to colder, wetter and less seasonal ones in Nigeria independent of latitude (Figs. 1, 3, 4)
Summary
Bergmann’s rule, defined as the tendency for endotherms to be larger in colder environments, is a well-known biophysical generalization for thermoregulation along environmental gradients (Salewski and Watt 2017). It seems likely that the exploration of absolute body size patterns (Graves 1991; Meiri and Dayan 2003; Ashton 2002) rather than relative body surface area in relation to thermoregulation (Salewski and Watt 2017) and the use of latitude as surrogate for temperature variation (Meiri and Dayan 2003) are responsible for some of the inconsistencies observed. Bergmann’s rule, on the other hand, postulates structural adaptation of the entire animal body to environmental temperature, based on a biophysical principle that more heat is lost to the external environment as body surface area increases relative to volume (Harley et al 2009; Salewski and Watt 2017). Bergmann’s rule should be interpreted based on two measures of body size: the ratio of the area to a cubic measure of body size
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