2c507aa86336ac12336d5baefe982cfe SJP-145-881_article-193019_en_1.pdf 49887dda0c5aaa184f3ca6bcfbde6bbe470756f4 SJP-145-881_article-193019_en_1.pdf 327c80ae6ed4b18b4625451f5b9bd8204bdb05db88d01a0b63aadaff2ae1ad70 SJP-145-881_article-193019_en_1.pdf Title: Lizards under pressure: skull nanoindentation in an insular reptile for palaeobiological inference Subject: Computational approaches are increasingly used to study the relationship between form and function in extinct and living organisms. However, one major challenge remains: assumptions about bone material properties, especially when studying fossils. In this work, direct mechanical testing and computer-based modelling are combined to assess cranial bone stiffness in the lizard genus Gallotia, endemic to the Canary Islands (Spain), including extinct species. Using linear mixed models, bone stiffness was found to be uniform across skull regions, with a trend towards lower stiffness in larger species. Simulations of biting loads showed that differences in bone properties had a limited influence on mechanical performance, supporting the use of average values in functional models. Using this framework, bone stiffness in extinct Gallotia giants was estimated, and its significance was explored. The results suggest a link between skull flexibility, body size, and diet, offering a practical approach to reconstructing biomechanical traits in extinct vertebrates across evolutionary timescales. 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1 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseNone Language: en-US Outlines: Item: Title: Lizards under pressure: skull nanoindentation in an insular reptile for palaeobiological inference Item: Title: Abstract Item: Title: Introduction Item: Title: Materials and methods Children: Item: Title: Nanoindentation Item: Title: Statistical analyses and linear mixed models (LMMs) Item: Title: Young’s modulus (E) extrapolation on fossil Gallotia specimens Item: Title: Finite element analysis Item: Title: Results Children: Item: Title: Nanoindentation analysis Item: Title: Homogeneity of Young’s modulus across cranial areas (LMM 1) Item: Title: Relationship between Young’s modulus and dorsal cranial length (LMM 2) Item: Title: Finite element analysis: differences between defined materials Item: Title: Differences between species Item: Title: Discussion Children: Item: Title: Nanoindentation analysis Item: Title: Diet and food resources Item: Title: Protection from predation Item: Title: Conclusion Item: Title: Acknowledgements Item: Title: References Item: Title: Additional information Item: Title: Supplementary material 1 Children: Item: Title: Summary of the number of indentations and sample size for each zone across five individuals: G_Stehlini-040322_3, G_Stehlini-040322_1, G_Sthelini-492-22, G_Atlantica-020322_4, and G_Galloti 121-1 Item: Title: Supplementary material 2 Children: Item: Title: Results from the nanoindentation arrays on the areas of interest in the cranium and mandible of G. atlantica 020322-4 Item: Title: Supplementary material 3 Children: Item: Title: Results from the nanoindentation arrays on the areas of interest in the cranium and mandible of G. stehlini 040322-3 Item: Title: Supplementary material 4 Children: Item: Title: Results from the nanoindentation arrays on the areas of interest in the cranium and mandible of G. stehlini GAST492-22 Item: Title: Supplementary material 5 Children: Item: Title: Results from the nanoindentation arrays on the areas of interest in the cranium and mandible of G. stehlini (040322-1) Item: Title: Supplementary material 6 Children: Item: Title: Results from the nanoindentation arrays on the areas of interest in the cranium and mandible of G. stehlini (040322-3) Item: Title: Supplementary material 7 Children: Item: Title: Results of all statistical tests performed with the complete dataset comprising 39,284 measurements per area across all species Item: Title: Supplementary material 8 Children: Item: Title: Scripts for the statistical analysis of data in R, including those used for the two linear mixed models (LMM1 and LMM2) and for the descriptive statistics applied in the methodology Item: Title: Supplementary material 9 Children: Item: Title: Supplementary text, figures and tables Info: Title: Lizards under pressure: skull nanoindentation in an insular reptile for palaeobiological inference Author: Author Subject: Computational approaches are increasingly used to study the relationship between form and function in extinct and living organisms. However, one major challenge remains: assumptions about bone material properties, especially when studying fossils. In this work, direct mechanical testing and computer-based modelling are combined to assess cranial bone stiffness in the lizard genus Gallotia, endemic to the Canary Islands (Spain), including extinct species. Using linear mixed models, bone stiffness was found to be uniform across skull regions, with a trend towards lower stiffness in larger species. Simulations of biting loads showed that differences in bone properties had a limited influence on mechanical performance, supporting the use of average values in functional models. Using this framework, bone stiffness in extinct Gallotia giants was estimated, and its significance was explored. The results suggest a link between skull flexibility, body size, and diet, offering a practical approach to reconstructing biomechanical traits in extinct vertebrates across evolutionary timescales. 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However, one major challenge remains: assumptions about bone material properties, especially when studying fossils. In this work, direct mechanical testing and computer-based modelling are combined to assess cranial bone stiffness in the lizard genus Gallotia, endemic to the Canary Islands (Spain), including extinct species. Using linear mixed models, bone stiffness was found to be uniform across skull regions, with a trend towards lower stiffness in larger species. Simulations of biting loads showed that differences in bone properties had a limited influence on mechanical performance, supporting the use of average values in functional models. Using this framework, bone stiffness in extinct Gallotia giants was estimated, and its significance was explored. The results suggest a link between skull flexibility, body size, and diet, offering a practical approach to reconstructing biomechanical traits in extinct vertebrates across evolutionary timescales. Laia Garcia-Escolà et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Author Bone mechanical properties finite element analysis Gallotia inference insular nanoindentation Young’s modulus 2026-08-14T11:41:06+03:00 2026-08-14T11:41:12+03:00 2026-08-14T11:41:12+03:00 Adobe InDesign 21.2 (Windows) uuid:97a33202-9a19-4b3c-ac78-ff3c96ef017c xmp.did:9F28E084BD8CE31184EC999A5D291C94 xmp.id:09a2b713-17a0-df42-83da-ac3a0c49cba9 proof:pdf xmp.iid:586b76da-c645-e94f-b1ec-8efdbe1408a1 xmp.did:4444FC51923DE811ADBCA6C78E600139 xmp.did:9F28E084BD8CE31184EC999A5D291C94 default converted from application/x-indesign to application/pdf Adobe InDesign 21.2 (Windows) / 2026-08-14T11:41:06+03:00 Adobe PDF Library 18.0 False Pages: Page: Label: 881 Sequence: 1 Annotations: Annotation: Subtype: Link Rect: 145, 663, 156, 652 Annotation: Subtype: Link Rect: 256, 663, 268, 652 Annotation: Subtype: Link Rect: 342, 663, 353, 652 Annotation: Subtype: Link Rect: 450, 663, 461, 652 Annotation: Subtype: Link Rect: 126, 649, 137, 637 Annotation: Subtype: Link Rect: 289, 649, 300, 637 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