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Urbanization, despite its destructive effects on natural habitats, offers species an opportunity to colonize novel niches. Previous research found that urban Anolis lizards in Puerto Rico had increased adhesive toepad area and more ventral toepad scales, traits that are likely adaptive and genetically based. We further investigated these phenotypic changes using geometric morphometrics to measure differences in toe shape, toepad shape, and lamellar morphology. Our results indicate that the increased toepad area of urban Anolis cristatellus lizards in Puerto Rico is not simply an isometric increase in toe size. Toes of urban populations exhibit multiple disproportional changes compared to forest lizards, with a larger proportion of the toe length covered in adhesive toepad. In addition, the toepads of urban lizards increase more in length than width. Lastly, lizards in urban populations exhibit both increased number of lamellae as well as increased spacing between individual lamellae. We also observed regional variation, with urban specimens having significantly more disparity, suggesting similar processes of urban adaptation are likely happening in parallel across the island, yet with region-specific idiosyncrasies, possibly generating more variation in toepad morphology across urban specimens as compared to forest specimens. Considering the use of geometric morphometrics, we found that specimen preparation, specifically how flat and straight toes are during imaging, to be an important factor affecting our data, more so than specimen size or any other meaningful morphological variation. In addition, we found that landmark and semilandmark data can be used to directly estimate toepad area, offering the opportunity to streamline future studies. In conclusion, our results highlight the value of considering toepad morphology in more detail beyond adhesive pad area or number of lamellae. Geometric morphometrics tools may be employed to elucidate subtle differences in shape to better allow researchers to connect changes in morphology to ecology and adhesive performance.

 

Abstract Human activity drastically transforms landscapes, generating novel habitats to which species must adaptively respond. Consequently, urbanization is increasingly recognized as a driver of phenotypic change. The structural environment of urban habitats presents a replicated natural experiment to examine trait–environment relationships and phenotypic variation related to locomotion. We use geometric morphometrics to examine claw morphology of five species of Anolis lizards in urban and forest habitats. We find that urban lizards undergo a shift in claw shape in the same direction but varying magnitude across species. Urban claws are overall taller, less curved, less pointed and shorter in length than those of forest lizards. These differences may enable more effective attachment or reduce interference with toepad function on smooth anthropogenic substrates. We also find an increase in shape disparity, a measurement of variation, in urban populations, suggesting relaxed selection or niche expansion rather than directional selection. This study expands our understanding of the relatively understudied trait of claw morphology and adds to a growing number of studies demonstrating phenotypic changes in urban lizards. The consistency in the direction of the shape changes we observed supports the intriguing possibility that urban environments may lead to predictable convergent adaptive change.