Longer wing bones in warmer climates suggest a role of thermoregulation in bird wing evolution

The tendency for animals in warmer climates to be longer-limbed (Allen’s Rule) is widely attributed to the demands of thermoregulation. The role played by thermoregulation in structuring bird wings, however, has been overshadowed by the selective demands placed on wings by flight. Using computer vision, we measure wing bone length from photographs of museum skeletal specimens for 1,520 species of passerine birds. We then model the relationship between wing bone length and temperature, accounting for allometry, the demands of flight efficiency and maneuverability, and a range of ecological and environmental variables.  We find that wing bones are longer in warmer climates. Our models, largely as a result of allometric effects, explain nearly all the variation in wing bone length in our data, with a marginal R2 = 0.80 and a conditional R2 > 0.99.  Thus, across 1,520 species of birds, higher temperatures are associated with longer wing bones, as predicted by Allen’s Rule. The vascularized musculature along these bones is maximally uncovered when birds actively hold their wings away from their bodies to aid in cooling or during flight. Conversely, the musculature along the wing bones is insulated by feathering when at rest, such that wings play a minor role in heat exchange when individuals are less active and may need to retain heat. While our analyses do not directly establish the mechanistic basis underlying the pattern we recover, given the asymmetry in the role of wings in thermoregulation, we interpret the positive relationship between temperature and wing bone length to reflect increased demand for heat dissipation in warmer climates. Our findings highlight the role of thermoregulation in shaping even the most critical features of vertebrate anatomy.

Methods
The data were collected by photographing museum skeletal specimens and then segmenting, identifying, and measuring the wing bones using Skelevision (Weeks et al., 2023).

Weeks, B.C., Z. Zhou, B.K. O'Brien, R. Darling, M. Dean, T. Dias, G. Hassena, M. Zhang, and D.F. Fouhey. 2023. A deep neural network for high-throughput measurement of functional traits on museum skeletal specimens. Methods in Ecology and Evolution 14(2): 347-359.