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  1. Abstract

    The subdigital adhesive pads of Caribbean Anolis lizards are considered to be a key innovation that permits occupation of novel ecological niches. Although previous work has demonstrated that subdigital pad morphology and performance vary with habitat use, such investigations have only considered the macroscale aspects of these structures (e.g. pad area). The morphological agents of attachment, however, are arrays of hair-like fibres (setae) that terminate in an expanded tip (spatula) and have not been examined in a similar manner. Here we examine the setal morphology and setal field configuration of ecologically distinct species of the monophyletic Jamaican Anolis radiation from a functional and ecological perspective. We find that anoles occupying the highest perches possess greater setal densities and smaller spatulae than those exploiting lower perches. This finding is consistent with the concept of contact splitting, whereby subdivision of an adhesive area into smaller and more densely packed fibres results in an increase in adhesive performance. Micromorphological evidence also suggests that the biomechanics of adhesive locomotion may vary between Anolis ecomorphs. Our findings indicate that, in a similar fashion to macroscale features of the subdigital pad, its microstructure may vary in relation to performance and habitat use in Caribbean Anolis.

     
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  2. Abstract

    The roughness and wettability of surfaces exploited by free-ranging geckos can be highly variable and attachment to these substrates is context dependent (e.g., presence or absence of surface water). Although previous studies focus on the effect of these variables on attachment independently, geckos encounter a variety of conditions in their natural environment simultaneously. Here, we measured maximum shear load of geckos in air and when their toes were submerged underwater on substrates that varied in both surface roughness and wettability. Gecko attachment was greater in water than in air on smooth and rough hydrophobic substrates, and attachment to rough hydrophilic substrates did not differ when tested in air or water. Attachment varied considerably with surface roughness and characterization revealed that routine measurements of root mean square height can misrepresent the complexity of roughness, especially when measured with single instruments. We used surface roughness power spectra to characterize substrate surface roughness and examined the relationship between gecko attachment performance across the power spectra. This comparison suggests that roughness wavelengths less than 70 nm predominantly dictate gecko attachment. This study highlights the complexity of attachment in natural conditions and the need for comprehensive surface characterization when studying biological adhesive system performance.

     
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  3. Abstract

    It has been nearly 20 years since Autumn and colleagues established the central role of van der Waals intermolecular forces in how geckos stick. Much has been discovered about the structure and function of fibrillar adhesives in geckos and other taxa, and substantial success has been achieved in translating natural models into bioinspired synthetic adhesives. Nevertheless, synthetics still cannot match the multidimensional performance observed in the natural gecko system that is simultaneously robust to dirt and water, resilient over thousands of cycles, and purportedly competent on surfaces that are rough at drastically different length scales. Apparent insensitivity of adhesion to variability in roughness is particularly interesting from both a theoretical and applied perspective. Progress on understanding the extent to which and the basis of how the gecko adhesive system is robust to variation in roughness is impeded by the complexity of quantifying roughness of natural surfaces and a dearth of data on free-ranging gecko substrate use. Here we review the main challenges in characterizing rough surfaces as they relate to collecting relevant estimates of variation in gecko adhesive performance across different substrates in their natural habitats. In response to these challenges, we propose a practical protocol (borrowing from thermal biophysical ecological methods) that will enable researchers to design detailed studies of structure–function relationships of the gecko fibrillar system. Employing such an approach will help provide specific hypotheses about how adhesive pad structure translates into a capacity for robust gecko adhesion across large variation in substrate roughness. Preliminary data we present on this approach suggest its promise in advancing the study of how geckos deal with roughness variation. We argue and outline how such data can help advance development of design parameters to improve bioinspired adhesives based on the gecko fibrillar system.

     
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  4. Abstract

    The remarkable ability of geckos to adhere to a wide-variety of surfaces has served as an inspiration for hundreds of studies spanning the disciplines of biomechanics, functional morphology, ecology, evolution, materials science, chemistry, and physics. The multifunctional properties (e.g., self-cleaning, controlled releasability, reversibility) and adhesive performance of the gekkotan adhesive system have motivated researchers to design and fabricate gecko-inspired synthetic adhesives of various materials and properties. However, many challenges remain in our attempts to replicate the properties and performance of this complex, hierarchical fibrillar adhesive system, stemming from fundamental, but unanswered, questions about how fibrillar adhesion operates. Such questions involve the role of fibril morphology in adhesive performance and how the gekkotan adhesive apparatus is utilized in nature. Similar fibrillar adhesive systems have, however, evolved independently in two other lineages of lizards (anoles and skinks) and potentially provide alternate avenues for addressing these fundamental questions. Anoles are the most promising group because they have been the subject of intensive ecological and evolutionary study for several decades, are highly speciose, and indeed are advocated as squamate model organisms. Surprisingly, however, comparatively little is known about the morphology, performance, and properties of their convergently-evolved adhesive arrays. Although many researchers consider the performance of the adhesive system of Anolis lizards to be less accomplished than its gekkotan counterpart, we argue here that Anolis lizards are prime candidates for exploring the fundamentals of fibrillar adhesion. Studying the less complex morphology of the anoline adhesive system has the potential to enhance our understanding of fibril morphology and its relationship to the multifunctional performance of fibrillar adhesive systems. Furthermore, the abundance of existing data on the ecology and evolution of anoles provides an excellent framework for testing hypotheses about the influence of habitat microstructure on the performance, behavior, and evolution of lizards with subdigital adhesive pads.

     
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