Attachment Beyond the Adhesive System: The Contribution of Claws to Gecko Clinging and Locomotion.

Attachment is imperative for many biological functions, such as holding position and climbing, but can be challenged by natural conditions. Adhesive toe pads and claws have evolved in multiple terrestrial lineages as important dynamic attachment mechanisms, and some clades (e.g., geckos) exhibit both features. The functional relationship of these features that comprise a complex attachment system is not well-understood, particularly within lizards (i.e., if pads and claws are redundant or multifunctional). Geckos exhibit highly adept frictional adhesive toe pads that continue to fuel biological inquiry and inspiration. However, gecko claws (the ancestral lizard clinging condition) have received little attention in terms of their functional or evolutionary significance. We assessed claw function in Thecadactylus rapicauda using assays of clinging performance and locomotor trials on different surfaces (artificial and natural) and inclines with claws intact, then partially removed. Area root mean square height (Sq), a metric of 3D surface roughness, was later quantified for all test surfaces, including acrylic, sandpaper, and two types of leaves (smooth and hairy). Maximum clinging force significantly declined on all non-acrylic surfaces after claw removal, indicating a substantial contribution to static clinging on rough and soft surfaces. With and without claws, clinging force exhibited a negative relationship with Sq. However, claw removal had relatively little impact on locomotor function on surfaces of different roughness at low inclines (≤30°). High static and dynamic safety factor estimates support these observations and demonstrate the species' robust frictional adhesive system. However, maximum station-holding capacity significantly declined on the rough test surface after partial claw removal, showing that geckos rely on their claws to maintain purchase on rough, steeply inclined surfaces. Our results point to a context-dependent complex attachment system within geckos, in which pads dominate on relatively smooth surfaces and claws on relatively rough surfaces, but also that these features function redundantly, possibly synergistically, on surfaces that allow attachment of both the setae and the claw (as in some insects). Our study provides important novel perspectives on gecko attachment, which we hope will spur future functional studies, new evolutionary hypotheses, and biomimetic innovation, along with collaboration and integration of perspectives across disciplines.