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Peer-reviewedCC BY 4.0Royal Society Open Science

Modularity without convergence: integrated evolution of the salamander autopodium

  • Giacomo Rosa1,2
  • Emma Centomo3
  • Andrea Costa1
  • Elisa Damonte1
  • Julien Renet4
  • Antonio Romano5
  • Luca Roner3,6
  • Sebastiano Salvidio1
  • Sara Ferrando1,2
6 affiliations
  1. 1University of Genova, Department of Earth, Environment and Life Sciences (DISTAV) 1 , Corso Europa 26, I-16132 Genova, Italy
  2. 2National Biodiversity Future Center (NBFC) 2 , Piazza Marina 61, I- 90133 Palermo (PA), Italy
  3. 3Research and Museum Collection Office, Conservation Biology Unit, MUSE - Science Museum 3 , Corso del Lavoro e della Scienza 3, 38122 Trento, Italy
  4. 4Fauna studium, Scientific Consulting 4 , 25 Avenue du Jabron, 04200 Sisteron, France
  5. 5Italian National Research Council (CNR), Institute of Bioeconomy – Biology, Agriculture and Food Sciences Department (IBE) 5 , Via dei Taurini, 19, 00185 Roma, Italy
  6. 6Departamento de Biología de Organismos y Sistemas, Unidad de Ecología, Universidad de Oviedo 6 , C/ Valentín Andŕes Alvarez s/n, 33006, Oviedo, Spain
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Abstract

Morphological modularity is known to favour evolutionary diversification by enabling different body regions to vary in response to functional and ecological demands. Despite their remarkable diversity in limb morphology and ecology, amphibians—and salamanders in particular—remain significantly under-represented in empirical tests of modularity. Using geometric morphometrics and phylogenetic comparative methods, we investigate patterns of shape variation, integration and modularity in the salamander autopodium. We first validated a landmark-based method to outline foot shape, demonstrating that image orientation does not affect shape analyses and that species can be reliably distinguished. We then applied this approach to a larger comparative dataset encompassing 149 salamander taxa across major lineages and ecological strategies. Our findings demonstrate that the salamander autopodium is functionally modular: the distal region (digits and interdigital region) shows more evolutionary lability than the proximal pes, while remaining integrated within a coordinated evolutionary system. Furthermore, the combinations of allometric, ecological and life-history factors account for a small but significant percentage of shape variation, with a low evolutionary signal in foot shape. Our results suggest that modular organization of the salamander foot may allow for morphological variation while still being limited by a common evolutionary and developmental history. By focusing on an understudied group and morphological system, this study provides new insight on how modularity influences limb evolution in tetrapods.

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Cite

BibTeXRIS
BibTeX
@article{rosa2026modularity,
  title = {{Modularity without convergence: integrated evolution of the salamander autopodium}},
  author = {Giacomo Rosa and Emma Centomo and Andrea Costa and Elisa Damonte and Julien Renet and Antonio Romano and Luca Roner and Sebastiano Salvidio and Sara Ferrando},
  journal = {Royal Society Open Science},
  year = {2026},
  month = oct,
  doi = {10.1098/rsos.261083},
  url = {https://royalsocietypublishing.org/rsos/article/13/10/261083/483790/Modularity-without-convergence-integrated},
  note = {CC BY 4.0},
}
RIS
TY  - JOUR
TI  - Modularity without convergence: integrated evolution of the salamander autopodium
AU  - Giacomo Rosa
AU  - Emma Centomo
AU  - Andrea Costa
AU  - Elisa Damonte
AU  - Julien Renet
AU  - Antonio Romano
AU  - Luca Roner
AU  - Sebastiano Salvidio
AU  - Sara Ferrando
PY  - 2026
DA  - 2026/10/21
JO  - Royal Society Open Science
DO  - 10.1098/rsos.261083
UR  - https://royalsocietypublishing.org/rsos/article/13/10/261083/483790/Modularity-without-convergence-integrated
AB  - Morphological modularity is known to favour evolutionary diversification by enabling different body regions to vary in response to functional and ecological demands. Despite their remarkable diversity in limb morphology and ecology, amphibians—and salamanders in particular—remain significantly under-represented in empirical tests of modularity. Using geometric morphometrics and phylogenetic comparative methods, we investigate patterns of shape variation, integration and modularity in the salamander autopodium. We first validated a landmark-based method to outline foot shape, demonstrating that image orientation does not affect shape analyses and that species can be reliably distinguished. We then applied this approach to a larger comparative dataset encompassing 149 salamander taxa across major lineages and ecological strategies. Our findings demonstrate that the salamander autopodium is functionally modular: the distal region (digits and interdigital region) shows more evolutionary lability than the proximal pes, while remaining integrated within a coordinated evolutionary system. Furthermore, the combinations of allometric, ecological and life-history factors account for a small but significant percentage of shape variation, with a low evolutionary signal in foot shape. Our results suggest that modular organization of the salamander foot may allow for morphological variation while still being limited by a common evolutionary and developmental history. By focusing on an understudied group and morphological system, this study provides new insight on how modularity influences limb evolution in tetrapods.
SN  - 2054-5703
ER  -