Bergmann meets Scholander: geographical variation in body size and insulation in the koala is related to climate
Natalie J. Briscoe1,2*, Andrew Krockenberger3, Kathrine A. Handasyde1 and Michael R. Kearney1
1 Department of Zoology, The University of Melbourne, Melbourne, Australia,
2 National Environmental Research Program, School of Botany, The University of Melbourne, Melbourne, Australia,
3 Centre for Tropical Biodiversity and Climate Change, College of Marine and Environmental Science, James Cook University, Cairns, Australia
*Correspondence: Natalie J. Briscoe, National Environmental Research Program, School of Botany, The University of Melbourne, Melbourne 3010, Australia. E-mail:
Aim Body size often varies clinally, with dominant explanations centred on how body size influences heat exchange (e.g. Bergmann’s rule). However, for endotherms, pelage properties can also dramatically alter heat exchange – a point emphasized by Scholander in the 1950s but which has received little attention in biogeographical analyses. Here, we investigate how geographical variation in both body size and fur properties of the koala (Phascolarctos cinereus) is related to climate.
Location Eastern Australia.
Methods We measured head length and fur depths of koala museum specimens from across its geographical range, and quantified the relationship between fur depth and insulation. We used linear regression and regression tree analyses to test for associations between morphological traits and climate variables relating to four hypotheses: heat conservation (Bergmann’s rule), heat dissipation, fasting endurance/survival of extremes, and productivity.
Results Both body size and fur depth of koalas decrease substantially towards the tropics. Consistent with Scholander’s view, fur properties showed stronger associations with climate than body size. Males, the larger sex, had shorter fur than females in hot environments but not in more temperate regions, suggesting that shorter fur compensates for sexual size dimorphism. While fur depth and male body size were more strongly associated with variables relating to heat dissipation, female body size was most strongly associated with minimum temperatures.
Main conclusions Body size interacts strongly with other traits, such as fur properties, to influence how animals experience climate. Our results emphasize how the consideration of geographical variation in suites of functionally related traits can provide important insight into how species persist across broad environmental gradients.