RT Journal Article T1 Complementarity in Allen’s and Bergmann’s rules among birds A1 Baldwin, Justin W. A1 García Porta, Joan A1 Botero, Carlos A. AB Biologists have long noted that endotherms tend to have larger bodies (Bergmann’s rule) and shorter appendages (Allen’s rule) in colder environments. Nevertheless, many taxonomic groups appear not to conform to these ‘rules’, and general explanations for these frequent exceptions are currently lacking. Here we note that by combining complementary changes in body and extremity size, lineages could theoretically respond to thermal gradients with smaller changes in either trait than those predicted by either Bergmann’s or Allen’s rule alone. To test this idea, we leverage geographic, ecological, phylogenetic, and morphological data on 6,974 non-migratory terrestrial bird species, and show that stronger family-wide changes in bill size over thermal gradients are correlated with more muted changes in body size. Additionally, we show that most bird families exhibit weak but appropriately directed changes in both traits, supporting the notion of complementarity in Bergmann’s and Allen’s rules. Finally, we show that the few families that exhibit significant gradients in either bill or body size, tend to be more speciose, widely distributed, or ecologically constrained. Our findings validate Bergmann’s and Allen’s logic and remind us that body and bill size are simply convenient proxies for their true quantity of interest: the surface-to-volume ratio. PB Springer Nature SN 2041-1723 YR 2023 FD 2023-07-15 LK https://hdl.handle.net/20.500.14352/121883 UL https://hdl.handle.net/20.500.14352/121883 LA eng NO Baldwin, J.W., Garcia-Porta, J. & Botero, C.A. Complementarity in Allen’s and Bergmann’s rules among birds. Nat Commun 14, 4240 (2023). https://doi.org/10.1038/s41467-023-39954-9 NO Thanks to Jonathan Losos for comments on an early draft. C.A.B. was funded by NSF, award number DEB 184147. J.G.-P. was supported by a Beatriu de Pinós fellowship from the Generalitat de Catalunya (2020 BP 00147) and by the program “Atracción de Talento Investigador Modalidad I” from the Comunidad de Madrid (2022-T1/AMB-24171). NO Generalitat de Catalunya NO Comunidad de Madrid DS Docta Complutense RD 27 jun 2025