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1. Convergent developmental patterns underlie the repeated evolution of adhesive toe pads among lizards

   https://doi.org/10.1093/biolinnean/blab164  

   Griffing, Aaron H; Gamble, Tony; Cohn, Martin J; Sanger, Thomas J (January 2022, Biological Journal of the Linnean Society)

   Abstract How developmental modifications produce key innovations, which subsequently allow for rapid diversification of a clade into new adaptive zones, has received much attention. However, few studies have used a robust comparative framework to investigate the influence of evolutionary and developmental constraints on the origin of key innovations, such as the adhesive toe pad of lizards. Adhesive toe pads evolved independently at least 16 times in lizards, allowing us to examine whether the patterns observed are general evolutionary phenomena or unique, lineage-specific events. We performed a high-resolution comparison of plantar scale development in 14 lizard species in Anolis and geckos, encompassing five independent origins of toe pads (one in Anolis, four in geckos). Despite substantial evolutionary divergence between Anolis and geckos, we find that these clades have undergone similar developmental modifications to generate their adhesive toe pads. Relative to the ancestral plantar scale development, in which scale ridges form synchronously along the digit, both padded geckos and Anolis exhibit scansor formation in a distal-to-proximal direction. Both clades have undergone developmental repatterning and, following their origin, modifications in toe pad morphology occurred through relatively minor developmental modifications, suggesting that developmental constraints governed the diversification of the adhesive toe pad in lizards. 

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2. Developmental Patterns Underlying Variation in Form and Function Exhibited by House Gecko Toe Pads

   https://doi.org/10.1093/icb/icae008  

   Griffing, Aaron_H; Gamble, Tony; Behere, Ashmika; Higham, Timothy_E; Keller, Greta_M; Resener, John; Sanger, Thomas_J (March 2024, Integrative And Comparative Biology)

   Synopsis Adhesive toe pads have evolved numerous times over lizard evolutionary history, most notably in geckos. Despite significant variation in adult toe pad morphology across independent origins of toe pads, early developmental patterns of toe pad morphogenesis are similar among distantly related species. In these distant phylogenetic comparisons, toe pad variation is achieved during the later stages of development. We aimed to understand how toe pad variation is generated among species sharing a single evolutionary origin of toe pads (house geckos—Hemidactylus). We investigated toe pad functional variation and developmental patterns in three species of Hemidactylus, ranging from highly scansorial (H. platyurus), to less scansorial (H. turcicus), to fully terrestrial (H. imbricatus). We found that H. platyurus generated significantly greater frictional adhesive force and exhibited much larger toe pad area relative to the other two species. Furthermore, differences in the offset of toe pad extension phase during embryonic development results in the variable morphologies seen in adults. Taken together, we demonstrate how morphological variation is generated in a complex structure during development and how that variation relates in important functional outcomes. 

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3. Evolution of the gekkotan adhesive system: Does digit anatomy point to one or more origins?

   https://doi.org/doi:10.1093/icb/icz006  

   Russell, A. P.; T. Gamble, T. (January 2019, Integrative and comparative biology)

   Synopsis Recently-developed, molecularly-based phylogenies of geckos have provided the basis for reassessing the number of times adhesive toe-pads have arisen within the Gekkota. At present both a single origin and multiple origin hypotheses prevail, each of which has consequences that relate to explanations about digit form and evolutionary transitions underlying the enormous variation in adhesive toe pad structure among extant, limbed geckos (pygopods lack pertinent features). These competing hypotheses result from mapping the distribution of toe pads onto a phylo- genetic framework employing the simple binary expedient of whether such toe pads are present or absent. It is evident, however, that adhesive toe pads are functional complexes that consist of a suite of integrated structural components that interact to bring about adhesive contact with the substratum and release from it. We evaluated the competing hypotheses about toe pad origins using 34 features associated with digit structure (drawn from the overall form of the digits; the presence and form of adhesive scansors; the proportions and structure of the phalanges; aspects of digital muscular and tendon morphology; presence and form of paraphalangeal elements; and the presence and form of substrate compliance-enhancing structures). We mapped these onto a well-supported phylogeny to reconstruct their evolution. Nineteen of these characters proved to be informative for all extant, limbed geckos, allowing us to assess which of them exhibit co- occurrence and/or clade-specificity. We found the absence of adhesive toe pads to be the ancestral state for the extant Gekkota as a whole, and our data to be consistent with independent origins of adhesive toe pads in the Diplodactylidae, Sphaerodactylidae, Phyllodactylidae, and Gekkonidae, with a strong likelihood of multiple origins in the latter three families. These findings are consistent with recently-published evidence of the presence of adhesively-competent digits in geckos generally regarded as lacking toe pads. Based upon morphology we identify other taxa at various locations within the gekkotan tree that are promising candidates for the expression of the early phases of adhesively-assisted locomotion. Investigation of functionally transitional forms will be valuable for enhancing our understanding of what is necessary and sufficient for the transition to adhesively-assisted locomotion, and for those whose objectives are to develop simulacra of the gekkotan adhesive system for biotechnological applications. 

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4. The Integrative Biology of Gecko Adhesion: Historical Review, Current Understanding, and Grand Challenges

   https://doi.org/10.1093/icb/icz032  

   Russell, Anthony P; Stark, Alyssa Y; Higham, Timothy E (May 2019, Integrative and Comparative Biology)

   Abstract Geckos are remarkable in their ability to reversibly adhere to smooth vertical, and even inverted surfaces. However, unraveling the precise mechanisms by which geckos do this has been a long process, involving various approaches over the last two centuries. Our understanding of the principles by which gecko adhesion operates has advanced rapidly over the past 20 years and, with this knowledge, material scientists have attempted to mimic the system to create artificial adhesives. From a biological perspective, recent studies have examined the diversity in morphology, performance, and real-world use of the adhesive apparatus. However, the lack of multidisciplinarity is likely a key roadblock to gaining new insights. Our goals in this paper are to 1) present a historical review of gecko adhesion research, 2) discuss the mechanisms and morphology of the adhesive apparatus, 3) discuss the origin and performance of the system in real-world contexts, 4) discuss advancement in bio-inspired design, and 5) present grand challenges in gecko adhesion research. To continue to improve our understanding, and to more effectively employ the principles of gecko adhesion for human applications, greater intensity and scope of interdisciplinary research are necessary. 

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5. Notes on patagium morphology in the Gliding Flat-tailed House Gecko (Hemidactylus platyurus)

   https://doi.org/10.17161/randa.v31i1.22398  

   Petty, Amber; Gamble, Tony; Griffing, Aaron (January 2024, Reptiles & Amphibians)

   Several species of geckos have independently evolved patagia, membranous features that facilitate gliding.Detailed morphological investigations of gecko patagia have largely been limited to gliding members of the genus Gekko(formerly in the genus Ptychozoon). Herein we describe the morphology of gliding patagia of the Flat-tailed HouseGecko (Hemidactylus platyurus), a species with an independent evolutionary origin of gliding patagia from Gekko andan important species for researching gliding biomechanics. We compared morphology of H. platyurus with a closelyrelated non-gliding species, the Common House Gecko (Hemidactylus frenatus). Using external examination and histologicaltechniques, we compared and contrasted three regions that exhibit patagia (trunk, femoral region, and tail)in H. platyurus but not in H. frenatus. We find that patagia in a gliding Hemidactylus, like patagia in gliding membersof the genus Gekko, are derived from expansion of lateral fat bodies, suggesting analogous processes to achieve similarphenotypic outcomes. 

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