Adhesive Toe Pads Research Articles

Microhabitat and adhesive toepads shape gecko limb morphology.

Different substrates pose varied biomechanical challenges that select specific morphologies, such as long limbs for faster running and short limbs for balanced posture while climbing narrow substrates. We tested how gecko locomotion is affected by the microhabitat they occupy and by a key adaptation-adhesive toepads-through analyzing how those are related to limb morphology. We collected microhabitat and toepads data for over 90% of limbed gecko species, and limb measurements for 403 species from 83 of the 121 limbed gecko genera, which we then used in phylogenetic comparative analyses. Our data highlight the association of adhesive toepads with arboreality, but a phylogenetic analysis shows that this relationship is not significant, suggesting that these traits are phylogenetically constrained. Comparative analyses reveal that pad-bearing species possess shorter hindlimbs and feet, more even limb lengths, and lower crus: thigh ratios, than padless geckos, across microhabitats. Saxicolous geckos have the longest limbs and limb segments. This is probably influenced by selection for long strides, increased takeoff velocity, and static stability on inclined surfaces. Terrestrial geckos have more even hind- and forelimbs than arboreal geckos, unlike patterns found in other lizards. Our findings underline the difficulty to infer on microhabitat-morphology relationships from one taxon to another, given their differing ecologies and evolutionary pathways. We emphasize the importance of key innovation traits, such as adhesive toepads, in shaping limb morphology in geckos and, accordingly, their locomotion within their immediate environment.

Ecologically‐related variation of digit morphology in Cyrtodactylus (Gekkota, Squamata) reveals repeated origins of incipient adhesive toepads

Abstract Exploitation of different locomotor substrates in different ecological niches has driven the evolution of specialized morphological structures, and similar ecological demands, such as the structure of the microhabitat, often lead to convergent or parallel evolution. The evolution of adhesive toepads in geckos remains understudied because of the paucity of phylogenetically‐informed investigations of candidate clades exhibiting purported incipient expression of these (i.e. species having evolved some, but not all, parts of the complex adhesive system of pad‐bearing geckos). Using Cyrtodactylus, a speciose genus with well‐established ecotypes, we tested the hypothesis that microhabitats that require more climbing will lead to the acquisition of incipient adhesive morphology. We measured subdigital scale area, a proxy for adhesive toepad evolution, and quantified subdigital scale shape for 77 of the 354 described species, including at least one representative of each ecotype. Subdigital scale area increased from terrestrial through generalist and saxicoline (rock‐dwelling) to arboreal ecotypes, with subdigital scale shape evolving from ancestral conditions for padless lizards to lateromedially expanded lamella‐like scales only in the arboreal ecotypes. This significant link between phenotype and environment supports the contention that scansorial, and particularly arboreal, Cyrtodactylus ecotypes have evolved incipiently expressed adhesive toepads. This highlights the suitability of this genus as a model system for studying the ecology and evolution of adhesive toepads as well as being a promising candidate for research on adaptive radiations. Read the free Plain Language Summary for this article on the Journal blog.

Toe pad morphology and adhesion in the miniaturized gecko, Chatogekko amazonicus (Gekkota: Sphaerodactylidae).

Chatogekko amazonicus is a miniaturized gecko from northern South America and is among the smallest of toe pad bearing lizards. The toe pads of C. amazonicus are miniscule, between 18% and 27% of the plantar surface area. We aimed to investigate the relationship between adhesive toe pad morphology, body size, and adhesive capabilities. Using scanning electron microscopy, we determine that the adhesive pads of C. amazonicus exhibit branched setae similar to those of other geckos, but that are generally much smaller. When compared with other gecko taxa, we show that C. amazonicus setae occupy a similar range of seta length: snout-vent length ratio and aspect ratio as other gekkonoid species (i.e. Gekkonidae, Phyllodactylidae, and Sphaerodactylidae). We demonstrate that C. amazonicus-even with its relatively small toe pads-is capable of climbing a smooth glass surface at a nearly vertical angle. We suggest that sphaerodactylids like C. amazonicus offer an excellent system for studying toe pad morphology and function in relation to miniaturization.

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

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.

Early burst of parallel evolution describes the diversification of gecko toe pads

IntroductionSimilar traits appearing in distantly related organisms have intrigued scientists for generations. While anole lizards of the Caribbean are often touted as a classic example of repeated evolution, the adhesive toe pads of gecko lizards are an equally striking yet underappreciated example of relatedly evolved traits. The strikingly diverse toe pads of gecko lizards (Gekkota) have been gained and lost multiple times throughout the clade’s evolutionary history. In addition, distantly related genera have repeatedly evolved remarkably similar morphologies. This complicated combination of divergent and repeated evolution represents a useful system for understanding the evolution of complex structures, including repeated adaptation.MethodsUsing geometric morphometrics, we evaluated parallel morphological differences across families and expanded existing approaches fitting models of trait evolution to use geometric morphometric data. Adapting the use of phylogenetic independent contrasts for shape data, we conducted a node height test to investigate how toe pad shape has evolved across geckos.ResultsWe found multiple examples of significant parallel differences in toe pad morphology and support for a model of early burst morphological evolution.DiscussionOur results suggest the diversification of Gekkotan toe pads included repeated parallel changes from padless ancestral morphologies to derived pad bearing morphologies. This morphological diversification occurred rapidly early in the diversification of gecko families and genera and slowed more recently, during diversification within genera.

Geometric Morphometrics Reveal Shape Differences in the Toes of Urban Lizards.

SynopsisUrbanization, 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.

Variation in density, but not morphology, of cutaneous sensilla among body regions in nine species of Australian geckos.

Skin sense organs, cutaneous sensilla, are a well-known feature of the integument of squamate reptiles and particularly geckos. They vary widely in morphology among species and are thought to be mechanosensitive, associated with prey capture and handling, tail autotomy and placement of the adhesive toepads in pad-bearing species. Some authors suggest that they may also sense abiotic environmental features, such as temperature or humidity. Here, we describe the morphology and distribution of cutaneous sensilla among body regions of nine Australian gecko species, in four genera. We hypothesised that if sensilla morphology was distinct, or sensilla density high, around the mouth, on the tail and on extremities, sensilla were likely used for these direct tactile functions. We found that sensilla morphology was uniform among body regions within species, but varied among species, while sensilla densities varied among species and body regions. In all species studied, sensilla density was highest on the labials and the dorsal tail scales and low on the feet, head and body, providing strong support for the hypothesis that sensilla serve tactile mechanoreceptive functions for prey capture and handling and for predator avoidance, but not for toepad placement. We suggest sensilla density may be explained by mechanoreception, whereas structure may be influenced by other factors.

Development and function explain the modular evolution of phalanges in gecko lizards.

Selective regimes favouring the evolution of functional specialization probably affect covariation among phenotypic traits. Phalanges of most tetrapods develop from a conserved module that constrains their relative proportions. In geckos, however, biomechanical specializations associated with adhesive toepads involve morphological variation in the autopodium and might reorganize such modular structures. We tested two hypotheses to explain the modular architecture of hand bones in geckos, one based on developmental interactions and another incorporating functional associations related to locomotion, and compared the empirical support for each hypothetical module between padded and padless lineages. We found strong evidence for developmental modules in most species, which probably reflects embryological constraints during phalangeal formation. Although padded geckos exhibit a functional specialization involving the hyperextension of the distal phalanges that is absent in padless species, the padless species are the ones that show a distal functional module with high integration. Some ancestrally padless geckos apparently deviate from developmental predictions and present a relatively weak developmental module of phalanges and a strongly integrated distal module, which may reflect selective regimes involving incipient frictional adhesion in digit morphology. Modularity of digit elements seems dynamic along the evolutionary history of geckos, being associated with the presence/absence of adhesive toepads.

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

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.

Morphology and Distribution of Cutaneous Sensory Organs on the Digits of <i>Anolis carolinensis</i> and <i>A. sagrei</i> (Squamata: Dactyloidae) in Relation to the Adhesive Toepads and Their Deployment

Cutaneous sensory organs are characteristic of many squamate lineages. Such organs may occur on the surface of scales as button-like, circular protuberances set off from their surroundings by a noticeable boundary, often taking the form of a moat or furrow. They may be relatively unadorned, clad with the surface micro-ornamentation of the scales on which they are carried, or they may carry one or more bristles of varying length and surface ornamentation. Such bristles may extend away from the body of the organ to interface with the surrounding environment or to contact adjacent scales. Cutaneous sensory organs have been physiologically demonstrated to have a mechanoreceptive function but have also been posited to potentially be involved with additional sensory modalities. Their distribution and structure across the body surface has been shown to be unequal, with some regions being much more extensively endowed than others, indicative of regional differential sensitivity. The digits of Anolis (Iguania: Dactyloidae) carry adhesive toepads that are convergent with those of geckos (Gekkota). Geckos exhibit a high density of cutaneous sensory organs on their toepads and their form and distribution has been associated with the operation and control of the toepads during locomotion. Investigation of the form and topographical distribution of cutaneous sensory organs on the toepads of Anolis shows them to be convergent in these attributes with those of geckos and quite distinct from those of the ancestrally padless Iguana (Iguania: Iguanidae). Their location at scale margins and the direction of their bristles towards adjacent scales indicates that the cutaneous sensory organs play an important role in proprioception during toepad deployment in Anolis.

Does Chytridiomycosis Affect Tree Frog Attachment?

The pandemic disease chytridiomycosis, caused by the fungus Batrachochytrium dendrobatidis (Bd), is a major threat to amphibian biodiversity. For most species, the exact mechanisms of chytridiomycosis that lead to negative population dynamics remain uncertain, though mounting evidence suggests that sublethal effects could be an important driver. In this review, we propose that tree frog attachment is a promising case to study the sublethal effects of a Bd infection on amphibians. A synthesis of the current knowledge on the functional morphology of the adhesive toe pads of tree frogs, on the underlying mechanisms of tree frog attachment, and on the epidermal pathology of chytridiomycosis substantiates the hypothesis that Bd-induced epidermal alterations have the potential to disrupt tree frog attachment. We highlight a series of (biomechanical) experiments to test this hypothesis and to shed some light on the sublethal disease mechanisms of chytridiomycosis. The knowledge generated from such an approach could contribute to future research on Bd epidemiology and ultimately to the conservation of the biodiversity of arboreal anurans.

Novel Tests of the Key Innovation Hypothesis: Adhesive Toepads in Arboreal Lizards.

The evolution of key innovations-unique features that enable a lineage to interact with the environment in a novel way-may drive broad patterns of adaptive diversity. However, traditional tests of the key innovation hypothesis, those which attempt to identify the evolutionary effect of a purported key innovation by comparing patterns of diversity between lineages with and without the key trait, have been challenged on both conceptual and statistical grounds. Here, we explore alternative, untested hypotheses of the key innovation framework. In lizards, adhesive toepad structures increase grip strength on vertical and smooth surfaces such as tree trunks and leaves and have independently evolved multiple times. As such, toepads have been posited as a key innovation for the evolution of arboreality. Leveraging a habitat use data set applied to a global phylogeny of 2692 lizard species, we estimated multiple origins of toepads in three major clades and more than 100 origins of arboreality widely across the phylogeny. Our results suggest that toepads arise adaptively in arboreal lineages and are subsequently rarely lost while maintaining arboreal ecologies. Padless lineages transition away from arboreality at a higher rate than those with toepads, and high rates of invasion of arboreal niches by nonarboreal padbearing lineages provide further evidence that toepads may be a key to unlocking evolutionary access to the arboreal zone. Our results and analytical framework provide novel insights to understand and evaluate the ecological and evolutionary consequences of key innovations.[Arboreality; ecological transition; key innovation; macroevolution; phylogenetic comparative methods.].

Ecological Opportunity from Innovation, not Islands, Drove the Anole Lizard Adaptive Radiation.

Islands are thought to facilitate adaptive radiation by providing release from competition and predation. Anole lizards are considered a classic example of this phenomenon: different ecological specialists ("ecomorphs") evolved in the Caribbean Greater Antilles (Cuba, Hispaniola, Jamaica, and Puerto Rico), resulting in convergent assemblages that are not observed in mainland Latin America. Yet, the role of islands in facilitating adaptive radiation is more often implied than directly tested, leaving uncertain the role of biogeography in stimulating diversification. Here, we assess the proposed "island effect" on anole diversification using Bayesian phylogenetic comparative methods that explicitly incorporate rate heterogeneity across the tree and demonstrate two cases of would be false positives. We discovered that rates of speciation and morphological evolution of island and mainland anoles are equivalent, implying that islands provide no special context for exceptionally rapid diversification. Likewise, rates of evolution were equivalent between island anoles that arose via in situ versus dispersal-based mechanisms, and we found no evidence for island-specific rates of speciation or morphological evolution. Nonetheless, the origin of Anolis is characterized by a speciation pulse that slowed over time-a classic signature of waning ecological opportunity. Our findings cast doubt on the notion that islands catalyzed the anole adaptive radiation and instead point to a key innovation, adhesive toe pads, which facilitated the exploitation of many arboreal niches sparsely utilized by other iguanian lizards. The selective pressures responsible for arboreal niche diversification differ between islands and the mainland, but the tempo of diversification driven by these discordant processes is indistinguishable. [Anolis; Caribbean; key innovation; morphological evolution; RevBayes; speciation.].

Setal Field Transects, Evolutionary Transitions and Gecko–Anole Convergence Provide Insights Into the Fundamentals of Form and Function of the Digital Adhesive System of Lizards

Recent years have witnessed a multitude of studies focusing on gekkotan adhesion. Intense interest in this phenomenon was triggered by the discovery of the manner and magnitude of the forces generated by the hair-like filaments (setae) on the toe pads and inspired the development of the next generation of smart, reversible synthetic adhesives. Most studies pursuing these goals have concentrated on the generalized form and properties of gekkotan setae outlined in those key early studies, resulting in the fabrication of synthetic filaments of uniform dimensions. Although there are over 1,800 species of extant geckos, and hundreds of species of anoles (a separate lizard lineage that has convergently evolved adhesive toe pads), most investigations have used relatively few species as the source of basic information, the Tokay gecko (Gekko gecko) being the most prominent among these. Such exemplar taxa generally exhibit structurally intricate setae and morphologically complex configurations of the adhesive apparatus. Setal structure taken to be characteristic of these taxa is generally reported by singular statements of maximal length, diameter, density and branching pattern. Contemporaneous work focusing on the configuration of setae at locations across the toe pads and upon the evolutionary origin of adhesively competent digits in anoles and specific lineages of geckos, however, has revealed extensive variation of setal structure within individuals, information about how setae may have arisen from non-adhesive filamentous precursors, and how newly adhesively competent digits have been integrated into pre-existing patterns of locomotor mechanics and kinematics. Such observations provide insights into what is minimally necessary for adhesively competent digits to function and reveal the simplest configuration of components that make this possible. We contend that information gleaned from such studies will assist those seeking to employ the principles of fibrillar-based adhesion, as exemplified by lizards, for bio-inspired applications.

Parallel evolution of toepads in rock-dwelling lineages of a terrestrial gecko (Gekkota: Gekkonidae:Heteronotia binoei)

AbstractSelection for effective locomotion can lead to specialized morphological structures. Adhesive toepads, which have arisen independently in different lizard clades, facilitate the use of vertical and inverted substrates. Their evolution is poorly understood because functionally intermediate morphological configurations between padless and pad-bearing forms are rare. To shed light on toepad evolution, we assessed the subdigital morphology of phylogenetically distinct lineages of the Bynoe’s gecko species complex (Heteronotia binoei). Most populations of H. binoei are terrestrial, but two relatively distantly related saxicoline (rock-dwelling) lineages have enlarged terminal subdigital scales resembling toepads. We reconstructed the ancestral terminal subdigital scale size of nine lineages of H. binoei in eastern Australia, including these two saxicoline lineages. Additionally, we compared the subdigital microstructures of four lineages: the two saxicoline lineages and their respective terrestrial sister-lineages. Surprisingly, all four lineages had fully developed setae, but the setae of the two saxicoline lineages were significantly longer, branched more often and were more widely spaced than the terrestrial sister-lineages. We conclude that the saxicoline lineages represent examples of parallel evolution of enlarged adhesive structures in response to vertical substrate use, and their morphology represents a useful model as an intermediate state in toepad evolution.

Design of Tree-Frog-Inspired Adhesives.

SynopsisThe adhesive toe pads of tree frogs have inspired the design of various so-called ‘smooth’ synthetic adhesives for wet environments. However, these adhesives do not reach the attachment performance of their biological models in terms of contact formation, maintenance of attachment, and detachment. In tree frogs, attachment is facilitated by an interconnected ensemble of superficial and internal morphological components, which together form a functional unit. To help bridging the gap between biological and bioinspired adhesives, in this review, we (1) provide an overview of the functional components of tree frog toe pads, (2) investigate which of these components (and attachment mechanisms implemented therein) have already been transferred into synthetic adhesives, and (3) highlight functional analogies between existing synthetic adhesives and tree frogs regarding the fundamental mechanisms of attachment. We found that most existing tree-frog-inspired adhesives mimic the micropatterned surface of the ventral epidermis of frog pads. Geometrical and material properties differ between these synthetic adhesives and their biological model, which indicates similarity in appearance rather than function. Important internal functional components such as fiber-reinforcement and muscle fibers for attachment control have not been considered in the design of tree-frog-inspired adhesives. Experimental work on tree-frog-inspired adhesives suggests that the micropatterning of adhesives with low-aspect-ratio pillars enables crack arresting and the drainage of interstitial liquids, which both facilitate the generation of van der Waals forces. Our analysis of experimental work on tree-frog-inspired adhesives indicates that interstitial liquids such as the mucus secreted by tree frogs play a role in detachment. Based on these findings, we provide suggestions for the future design of biomimetic adhesives. Specifically, we propose to implement internal fiber-reinforcements inspired by the fibrous structures in frog pads to create mechanically reinforced soft adhesives for high-load applications. Contractile components may stimulate the design of actuated synthetic adhesives with fine-tunable control of attachment strength. An integrative approach is needed for the design of tree-frog-inspired adhesives that are functionally analogous with their biological paradigm.

Vertical Fibrous Morphology and Structure-Function Relationship in Natural and Biomimetic Suction-Based Adhesion Discs

Summary Recent years have seen rapid development in bio-inspired materials for adhesion. However, it remains challenging to realize robust adhesion on rough aquatic surfaces. An inspiring natural model is the remora fish, which has evolved to retain powerful adhesion to hosts using a dorsal suction disc. We find that the remora suction disc has a unique fibrous architecture of vertically oriented collagen fibers that enable anisotropic mechanical properties and enhanced adhesion performance. In the engineered prototype, vertically oriented nylon fibers are embedded into the soft silicone matrix using electrostatic flocking. The anisotropic mechanical properties are validated in both natural and biomimetic suction discs. Furthermore, the biomimetic suction disc demonstrates an enhanced adhesion function with a maximum 62.5% increase in pull-off force and a 340% increase in attachment time compared with the silicone control. This work can shed light on natural adhesion mechanisms and inspire novel designs for aquatic soft adhesives and actuators.

Nonlinear variation in clinging performance with surface roughness in geckos

Understanding the challenges faced by organisms moving within their environment is essential to comprehending the evolution of locomotor morphology and habitat use. Geckos have developed adhesive toe pads that enable exploitation of a wide range of microhabitats. These toe pads, and their adhesive mechanisms, have typically been studied using a range of artificial substrates, usually significantly smoother than those available in nature. Although these studies have been fundamental in understanding the mechanisms of attachment in geckos, it is unclear whether gecko attachment simply gradually declines with increased roughness as some researchers have suggested, or whether the interaction between the gekkotan adhesive system and surface roughness produces nonlinear relationships. To understand ecological challenges faced in their natural habitats, it is essential to use test surfaces that are more like surfaces used by geckos in nature. We tested gecko shear force (i.e., frictional force) generation as a measure of clinging performance on three artificial substrates. We selected substrates that exhibit microtopographies with peak‐to‐valley heights similar to those of substrates used in nature, to investigate performance on a range of smooth surfaces (glass), and fine‐grained (fine sandpaper) to rough (coarse sandpaper). We found that shear force did not decline monotonically with roughness, but varied nonlinearly among substrates. Clinging performance was greater on glass and coarse sandpaper than on fine sandpaper, and clinging performance was not significantly different between glass and coarse sandpaper. Our results demonstrate that performance on different substrates varies, probably depending on the underlying mechanisms of the adhesive apparatus in geckos.

Tree frog adhesion biomimetics: opportunities for the development of new, smart adhesives that adhere under wet conditions.

Enlarged adhesive toe pads on the tip of each digit allow tree frogs to climb smooth vertical and overhanging surfaces, and are effective in generating reversible adhesion under both dry and wet conditions. In this review, we discuss the complexities of the structure of tree frog toe pads in relation to their function and review their biomimetic potential. Of particular importance are the (largely) hexagonal epithelial cells surrounded by deep channels that cover the surface of each toe pad and the array of nanopillars on their surface. Fluid secreted by the pads covers the surface of each pad, so the pads adhere by wet adhesion, involving both capillarity and viscosity-dependent forces. The fabrication and testing of toe pad mimics are challenging, but valuable both for testing hypotheses concerning tree frog toe pad function and for developing toe pad mimics. Initial mimics involved the fabrication of hexagonal pillars mimicking the toe pad epithelial structure. More recent ones additionally replicate the nanostructures on their surface. Finally we describe some of the biomimetic applications that have been developed from toe pad mimics, which include both bioinspired adhesives and friction-generating devices. This article is part of the theme issue 'Bioinspired materials and surfaces for green science and technology (part 2)'.

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.