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.

The past, present, and future of predator-prey interactions in a warming world: Using species distribution modeling to forecast ectotherm-endotherm niche overlap.

Climate change has the potential to disrupt species interactions across global ecosystems. Ectotherm-endotherm interactions may be especially prone to this risk due to the possible mismatch between the species in physiological response and performance. However, few studies have examined how changing temperatures might differentially impact species' niches or available suitable habitat when they have very different modes of thermoregulation. An ideal system for studying this interaction is the predator-prey system. In this study, we used ecological niche modeling to characterize the niche overlap and examine biogeography in past and future climate conditions of prairie rattlesnakes (Crotalus viridis) and Ord's kangaroo rats (Dipodomys ordii), an endotherm-ectotherm pair typifying a predator-prey species interaction. Our models show a high niche overlap between these two species (D = 0.863 and I = 0.979) and further affirm similar paleoecological distributions during the last glacial maximum (LGM) and mid-Holocene (MH). Under future climate change scenarios, we found that prairie rattlesnakes may experience a reduction in overall suitable habitat (RCP 2.6 = -1.82%, 4.5 = -4.62%, 8.5 = -7.34%), whereas Ord's kangaroo rats may experience an increase (RCP 2.6 = 9.8%, 4.5 = 11.71%, 8.5 = 8.37%). We found a shared trend of stable suitable habitat at northern latitudes but reduced suitability in southern portions of the range, and we propose future monitoring and conservation be focused on those areas. Overall, we demonstrate a biogeographic example of how interacting ectotherm-endotherm species may have mismatched responses under climate change scenarios and the models presented here can serve as a starting point for further investigation into the biogeography of these systems.

Cryptic behavior and activity cycles of a small mammal keystone species revealed through accelerometry: a case study of Merriam's kangaroo rats (Dipodomys merriami).

BACKGROUND: Kangaroo rats are small mammals that are among the most abundant vertebrates in many terrestrial ecosystems in Western North America and are considered both keystone species and ecosystem engineers, providing numerous linkages between other species as both consumers and resources. However, there are challenges to studying the behavior and activity of these species due to the difficulty of observing large numbers of individuals that are small, secretive, and nocturnal. Our goal was to develop an integrated approach of miniaturized animal-borne accelerometry and radiotelemetry to classify the cryptic behavior and activity cycles of kangaroo rats and test hypotheses of how their behavior is influenced by light cycles, moonlight, and weather. METHODS: We provide a proof-of-concept approach to effectively quantify behavioral patterns of small bodied (< 50 g), nocturnal, and terrestrial free-ranging mammals using large acceleration datasets by combining low-mass, miniaturized animal-borne accelerometers with radiotelemetry and advanced machine learning techniques. We developed a method of attachment and retrieval for deploying accelerometers, a non-disruptive method of gathering observational validation datasets for acceleration data on free-ranging nocturnal small mammals, and used these techniques on Merriam's kangaroo rats to analyze how behavioral patterns relate to abiotic factors. RESULTS: We found that Merriam's kangaroo rats are only active during the nighttime phases of the diel cycle and are particularly active during later light phases of the night (i.e., late night, morning twilight, and dawn). We found no reduction in activity or foraging associated with moonlight, indicating that kangaroo rats are actually more lunarphilic than lunarphobic. We also found that kangaroo rats increased foraging effort on more humid nights, most likely as a mechanism to avoid cutaneous water loss. CONCLUSIONS: Small mammals are often integral to ecosystem functionality, as many of these species are highly abundant ecosystem engineers driving linkages in energy flow and nutrient transfer across trophic levels. Our work represents the first continuous detailed quantitative description of fine-scale behavioral activity budgets in kangaroo rats, and lays out a general framework for how to use miniaturized biologging devices on small and nocturnal mammals to examine behavioral responses to environmental factors.

The tailless gecko gets the worm: prey type alters the effects of caudal autotomy on prey capture and subjugation kinematics.

Prey capture and subjugation are complex behaviors affected by many factors including physiological and behavioral traits of both the predator and the prey. The western banded gecko (Coleonyx variegatus) is a small generalist predator that consumes both evasive prey items, such as spiders, wasps, and orthopterans, and non-evasive prey items, including larvae, pupae, and isopterans. When consuming certain prey (e.g., scorpions), banded geckos will capture and then rapidly oscillate, or shake, their head and anterior part of their body. Banded geckos also have large, active tails that can account for over 20% of their body weight and can be voluntarily severed through the process of caudal autotomy. However, how autotomy influences prey capture behavior in geckos is poorly understood. Using high-speed 3D videography, we studied the effects of both prey type (mealworms and crickets) and tail autotomy on prey capture and subjugation performance in banded geckos. Performance metrics included maximum velocity and distance of prey capture, as well as velocity and frequency of post-capture shaking. Maximum velocity and distance of prey capture were lower for mealworms than crickets regardless of tail state. However, after autotomy, maximum velocity increased for strikes on mealworms but significantly decreased for crickets. After capture, geckos always shook mealworms, but never crickets. The frequency of shaking mealworms decreased after autotomy and additional qualitative differences were observed. Our results highlight the complex and interactive effects of prey type and caudal autotomy on prey capture biomechanics.

Growing up in a rough world: scaling of frictional adhesion and morphology of the Tokay gecko (Gekko gecko).

Many geckos have the remarkable ability to reversibly adhere to surfaces using a hierarchical system that includes both internal and external elements. The vast majority of studies have examined the performance of the adhesive system using adults and engineered materials and substrates (e.g., acrylic glass). Almost nothing is known about how the system changes with body size, nor how these changes would influence the ability to adhere to surfaces in nature. Using Tokay geckos (Gekko gecko), we examined the post-hatching scaling of morphology and frictional adhesive performance in animals ranging from 5 to 125 grams in body mass. We quantified setal density, setal length, and toepad area using SEM. This was then used to estimate the theoretical maximum adhesive force. We tested performance with 14 live geckos on eight surfaces ranging from extremely smooth (acrylic glass) to relatively rough (100-grit sandpaper). Surfaces were attached to a force transducer, and multiple trials were conducted for each individual. We found that setal length scaled with negatively allometry, but toepad area scaled with isometry. Setal density remained constant across the wide range in body size. The relationship between body mass and adhesive performance was generally similar across all surfaces, but rough surfaces had much lower values than smooth surfaces. The safety factor went down with body mass and with surface roughness, suggesting that smaller animals may be more likely to occupy rough substrates in their natural habitat.

Ecomechanics and the Rules of Life: a Critical Conduit Between the Physical and Natural Sciences.

Nature provides the parameters, or boundaries, within which organisms must cope in order to survive. Therefore, ecological conditions have an unequivocal influence on the ability of organisms to perform the necessary functions for survival. Biomechanics brings together physics and biology to understand how an organism will function under a suite of conditions. Despite a relatively rich recent history linking physiology and morphology with ecology, less attention has been paid to the linkage between biomechanics and ecology. This linkage, however, could provide key insights into patterns and processes of evolution. Ecomechanics, also known as ecological biomechanics or mechanical ecology, is not necessarily new, but has received far less attention than ecophysiology or ecomorphology. Here, we briefly review the history of ecomechanics, and then identify what we believe are grand challenges for the discipline and how they can inform some of the most pressing questions in science today, such as how organisms will cope with global change.

The evolution of mechanical properties of conifer and angiosperm woods.

The material properties of the cells and tissues of an organism dictate, to a very large degree, the ability of the organism to cope with mechanical stress induced by externally applied forces. It is, therefore, critical to understand how these properties differ across diverse species and how they have evolved. Herein, a large data base (N = 84 species) for the mechanical properties of wood samples measured at biologically natural moisture contents (i.e., "green wood") was analyzed to determine the extent to which these properties are correlated across phylogenetically diverse tree species, to determine if a phylogenetic pattern of trait values exists, and, if so, to assess whether the rate of trait evolution varies across the phylogeny. The phylogenetic comparative analyses presented here confirm previous results that critical material properties are significantly correlated with one another and with wood density. Although the rates of trait evolution of angiosperms and gymnosperms (i.e., conifers) are similar, the material properties of both clades evolved in distinct selective regimes that are phenotypically manifested in lower values across all material properties in gymnosperms. This observation may be related to the structural differences between gymnosperm and angiosperm wood such as the presence of vessels in angiosperms. Explorations of rate heterogeneity indicate high rates of trait evolution in wood density in clades within both conifers and angiosperms (e.g., Pinus and Shorea). Future analyses are warranted using additional data given these preliminary results, especially because there is ample evidence of convergent evolution in the material properties of conifers and angiosperm wood that appear to experience similar ecological conditions.

Comparative analysis of Dipodomys species indicates that kangaroo rat hindlimb anatomy is adapted for rapid evasive leaping.

Body size is a key factor that influences antipredator behavior. For animals that rely on jumping to escape from predators, there is a theoretical trade-off between jump distance and acceleration as body size changes at both the inter- and intraspecific levels. Assuming geometric similarity, acceleration will decrease with increasing body size due to a smaller increase in muscle cross-sectional area than body mass. Smaller animals will likely have a similar jump distance as larger animals due to their shorter limbs and faster accelerations. Therefore, in order to maintain acceleration in a jump across different body sizes, hind limbs must be disproportionately bigger for larger animals. We explored this prediction using four species of kangaroo rats (Dipodomys spp.), a genus of bipedal rodent with similar morphology across a range of body sizes (40-150 g). Kangaroo rat jump performance was measured by simulating snake strikes to free-ranging individuals. Additionally, morphological measurements of hind limb muscles and segment lengths were obtained from thawed frozen specimens. Overall, jump acceleration was constant across body sizes and jump distance increased with increasing size. Additionally, kangaroo rat hind limb muscle mass and cross-sectional area scaled with positive allometry. Ankle extensor tendon cross-sectional area also scaled with positive allometry. Hind limb segment length scaled isometrically, with the exception of the metatarsals, which scaled with negative allometry. Overall, these findings support the hypothesis that kangaroo rat hind limbs are built to maintain jump acceleration rather than jump distance. Selective pressure from single-strike predators, such as snakes and owls, likely drives this relationship.

Jumping with adhesion: landing surface incline alters impact force and body kinematics in crested geckos.

Arboreal habitats are characterized by a complex three-dimensional array of branches that vary in numerous characteristics, including incline, compliance, roughness, and diameter. Gaps must often be crossed, and this is frequently accomplished by leaping. Geckos bearing an adhesive system often jump in arboreal habitats, although few studies have examined their jumping biomechanics. We investigated the biomechanics of landing on smooth surfaces in crested geckos, Correlophus ciliatus, asking whether the incline of the landing platform alters impact forces and mid-air body movements. Using high-speed videography, we examined jumps from a horizontal take-off platform to horizontal, 45° and 90° landing platforms. Take-off velocity was greatest when geckos were jumping to a horizontal platform. Geckos did not modulate their body orientation in the air. Body curvature during landing, and landing duration, were greatest on the vertical platform. Together, these significantly reduced the impact force on the vertical platform. When landing on a smooth vertical surface, the geckos must engage the adhesive system to prevent slipping and falling. In contrast, landing on a horizontal surface requires no adhesion, but incurs high impact forces. Despite a lack of mid-air modulation, geckos appear robust to changing landing conditions.

Ankle structure of the Tokay gecko (Gekko gecko) and its role in the deployment of the subdigital adhesive system.

The remarkable ability of geckos to adhere to smooth surfaces is often thought of in terms of external structures, including the branching setae that make contact with the surface producing van der Waals forces. Some geckos also exhibit unique movements of the distal segments of the limbs during locomotion and static clinging, including active digital hyperextension and considerable pedal rotation. During static clinging, geckos can exhibit considerable adduction/abduction of the pes while the crus and thigh remain firmly adpressed to the substratum. This decoupling of pedal adduction/abduction from ankle flexion/extension and pedal long-axis rotation is a significant departure from pedal displacements of a typical lizard lacking adhesive ability. The structure of the ankle is likely key to this decoupling, although no detailed comparison of this complex joint between pad-bearing geckos and other lizards is available. Here we compare the configuration of the mesotarsal joint of nongekkotan lizards (Iguana and Pristidactylus) with that of the Tokay gecko (Gekko gecko) using prepared skeletons, scanning electron microscopy, and micro-computed tomographic (µCT) scans. We focus on the structure of the astragalocalcaneum and the fourth distal tarsal. The mesotarsal joint exhibits a suite of modifications that are likely associated with the secondarily symmetrical pes of pad-bearing geckos. For example, the lateral process of the astragalocalcaneum is much more extensive in G. gecko compared with other lizards. The mesotarsal joint exhibits several other differences permitting dissociation of long-axis rotation of the pes from flexion-extension movement, including a reduced ventral peg on the fourth distal tarsal, an articulatory pattern dominated by a well-defined, expansive distomesial notch of the astragalocalcaneum, and an associated broad proximodorsal articulatory facet of the fourth distal tarsal. Pad-bearing geckos are capable of effectively deploying their intricate adhesive system across a broad array of body angles because of this highly modified ankle. Future research should determine whether the differences encountered in G. gecko (and their extent) apply to the Gekkota as a whole and should examine how the elements of the ankle move dynamically during locomotion across a range of taxa.

Linking ecomechanical models and functional traits to understand phenotypic diversity.

Physical principles and laws determine the set of possible organismal phenotypes. Constraints arising from development, the environment, and evolutionary history then yield workable, integrated phenotypes. We propose a theoretical and practical framework that considers the role of changing environments. This 'ecomechanical approach' integrates functional organismal traits with the ecological variables. This approach informs our ability to predict species shifts in survival and distribution and provides critical insights into phenotypic diversity. We outline how to use the ecomechanical paradigm using drag-induced bending in trees as an example. Our approach can be incorporated into existing research and help build interdisciplinary bridges. Finally, we identify key factors needed for mass data collection, analysis, and the dissemination of models relevant to this framework.

And thereby hangs a tail: morphology, developmental patterns and biomechanics of the adhesive tails of crested geckos (Correlophus ciliatus).

Among the most specialized integumentary outgrowths in amniotes are the adhesive, scale-like scansors and lamellae on the digits of anoles and geckos. Less well-known are adhesive tail pads exhibited by 21 gecko genera. While described over 120 years ago, no studies have quantified their possible adhesive function or described their embryonic development. Here, we characterize adult and embryonic morphology and adhesive performance of crested gecko (Correlophus ciliatus) tail pads. Additionally, we use embryonic data to test whether tail pads are serial homologues to toe pads. External morphology and histology of C. ciliatus tail pads are largely similar to tail pads of closely related geckos. Functionally, C. ciliatus tail pads exhibit impressive adhesive ability, hypothetically capable of holding up to five times their own mass. Tail pads develop at approximately the same time during embryogenesis as toe pads. Further, tail pads exhibit similar developmental patterns to toe pads, which are markedly different from non-adhesive gecko toes and tails. Our data provide support for the serial homology of adhesive tail pads with toe pads.

Tail Autotomy Alters Prey Capture Performance and Kinematics, but not Success, in Banded Geckos.

Tails are versatile structures with diverse forms and functions across vertebrates. They are involved in almost all behaviors critical to survival including locomotion, feeding, and predator avoidance. Although the tail's role in locomotion and stability has been widely studied, its role in prey capture is relatively unknown. Lizards are an ideal system to examine the tail's impact on prey capture as most are capable of autotomizing, or dropping, their tail in response to predation and intraspecific competition. Tail autotomy can lower reproduction, decrease locomotor performance, impart instability during jumping, and decrease social status. Desert banded geckos (Coleonyx variegatus) frequently capture evasive prey in nature and appear to use their tail during strikes. However, it is unclear if these tail movements are important for the strike itself, or if they simply draw attention to that part of the body. We used high-speed 3D videography to quantify prey capture performance and kinematics of C. variegatus striking at crickets before and after total caudal autotomy. Trials were conducted within 2 h of autotomy and then repeatedly over a 2-week period. Overall, prey capture success was unaffected by caudal autotomy. However, maximum strike velocity decreased significantly after autotomy, highlighting the importance of the tail during prey capture. Strike kinematics were altered after autotomy in several ways, including geckos adopting a more sprawled posture. Maximum pectoral girdle and mid-back height were significantly lower during post-autotomy strikes, whereas maximum pelvic girdle height was unaffected. However, individual variation was considerable. This downward pitching of the body after tail loss suggests that the tail is necessary for counterbalancing the anterior portion of the body and resisting the rotational inertia incurred after pushing off with the hindlimbs. Utilizing autotomy to test tail function in prey capture can provide valuable insight into how the tail is used in terrestrial predation across a wide variety of species and ecological niches.

Tail Control Enhances Gliding in Arboreal Lizards: An Integrative Study Using a 3D Geometric Model and Numerical Simulation.

The ability to glide through an arboreal habitat has been acquired by several mammals, amphibians, snakes, lizards, and even invertebrates. Lizards of the genus Draco possess specialized morphological structures for gliding, including a patagium, throat lappets, and modified hindlimbs. Despite being among the most specialized reptilian gliders, it is currently unknown how Draco is able to maneuver effectively during flight. Here, we present a new computational method for characterizing the role of tail control on Draco glide distance and stability. We first modeled Draco flight dynamics as a function of gravitational, lift, and drag forces. Lift and drag estimates were derived from wind tunnel experiments of 3D printed models based on photos of Draco during gliding. Initial modeling leveraged the known mass and planar surface area of the Draco to estimate lift and drag coefficients. We developed a simplified, 3D simulation for Draco gliding, calculating longitudinal and lateral position and a pitch angle of the lizard with respect to a cartesian coordinate frame. We used PID control to model the lizards' tail adjustment to maintain an angle of attack. Our model suggests an active tail improves both glide distance and stability in Draco. These results provide insight toward the biomechanics of Draco; however, future in vivo studies are needed to provide a complete picture for gliding mechanics of this genus. Our approach enables the replication and modification of existing gliders to better understand their performance and mechanics. This can be applied to extinct species, but also as a way of exploring the biomimetic potential of different morphological features.

The effects of temperature on the defensive strikes of rattlesnakes.

Movements of ectotherms are constrained by their body temperature owing to the effects of temperature on muscle physiology. As physical performance often affects the outcome of predator-prey interactions, environmental temperature can influence the ability of ectotherms to capture prey and/or defend themselves against predators. However, previous research on the kinematics of ectotherms suggests that some species may use elastic storage mechanisms when attacking or defending, thereby mitigating the effects of sub-optimal temperature. Rattlesnakes (Crotalus spp.) are a speciose group of ectothermic viperid snakes that rely on crypsis, rattling and striking to deter predators. We examined the influence of body temperature on the behavior and kinematics of two rattlesnake species (Crotalus oreganus helleri and Crotalus scutulatus) when defensively striking towards a threatening stimulus. We recorded defensive strikes at body temperatures ranging from 15-35°C. We found that strike speed and speed of mouth gaping during the strike were positively correlated with temperature. We also found a marginal effect of temperature on the probability of striking, latency to strike and strike outcome. Overall, warmer snakes are more likely to strike, strike faster, open their mouth faster and reach maximum gape earlier than colder snakes. However, the effects of temperature were less than would be expected for purely muscle-driven movements. Our results suggest that, although rattlesnakes are at a greater risk of predation at colder body temperatures, their decrease in strike performance may be mitigated to some extent by employing mechanisms in addition to skeletal muscle contraction (e.g. elastic energy storage) to power strikes.

Evolution of pedal digit orientation and morphology in relation to acquisition and secondary loss of the adhesive system in geckos.

Among geckos, the acquisition of the adhesive system is associated with several morphological changes of the feet that are involved in the operation of the adhesive apparatus. However, analyses using a comparative framework are lacking. We applied traditional morphometrics and geometric morphometric analysis with phylogenetic comparative methods to morphological data, collected from X-ray scans, to examine patterns of morphological evolution of the pes in association with the gain and loss of adhesive capabilities, and with habitat occupancy among 102 species of gecko. Padbearing gecko lineages tend to have shorter digits and greater inter-digital angles than padless ones. Arboreal and saxicolous species have shorter digits than terrestrial species. Our results suggest repeated shifts that converge upon a similar padbearing morphology, with some modifications being associated with the habitat occupied. We demonstrate that functional innovation and habitat can operate on, and influence, different components of foot morphology.

A Hierarchical View of Gecko Locomotion: Photic Environment, Physiological Optics, and Locomotor Performance.

Terrestrial animals move in complex habitats that vary over space and time. The characteristics of these habitats are not only defined by the physical environment, but also by the photic environment, even though the latter has largely been overlooked. For example, numerous studies of have examined the role of habitat structure, such as incline, perch diameter, and compliance, on running performance. However, running performance likely depends heavily on light level. Geckos are an exceptional group for analyzing the role of the photic environment on locomotion as they exhibit several independent shifts to diurnality from a nocturnal ancestor, they are visually-guided predators, and they are extremely diverse. Our initial goal is to discuss the range of photic environments that can be encountered in terrestrial habitats, such as day versus night, canopy cover in a forest, fog, and clouds. We then review the physiological optics of gecko vision with some new information about retina structures, the role of vision in motor-driven behaviors, and what is known about gecko locomotion under different light conditions, before demonstrating the effect of light levels on gecko locomotor performance. Overall, we highlight the importance of integrating sensory and motor information and establish a conceptual framework as guide for future research. Several future directions, such as understanding the role of pupil dynamics, are dependent on an integrative framework. This general framework can be extended to any motor system that relies on sensory information, and can be used to explore the impact of performance features on diversification and evolution.

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.

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

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.

Replicating the complexity of natural surfaces: technique validation and applications for biomimetics, ecology and evolution.

The surfaces of animals, plants and abiotic structures are not only important for organismal survival, but they have also inspired countless biomimetic and industrial applications. Additionally, the surfaces of animals and plants exhibit an unprecedented level of diversity, and animals often move on the surface of plants. Replicating these surfaces offers a number of advantages, such as preserving a surface that is likely to degrade over time, controlling for non-structural aspects of surfaces, such as compliance and chemistry, and being able to produce large areas of a small surface. In this paper, we compare three replication techniques among a number of species of plants, a technical surface and a rock. We then use two model parameters (cross-covariance function ratio and relative topography difference) to develop a unique method for quantitatively evaluating the quality of the replication. Finally, we outline future directions that can employ highly accurate surface replications, including ecological and evolutionary studies, biomechanical experiments, industrial applications and improving haptic properties of bioinspired surfaces. The recent advances associated with surface replication and imaging technology have formed a foundation on which to incorporate surface information into biological sciences and to improve industrial and biomimetic applications. This article is part of the theme issue 'Bioinspired materials and surfaces for green science and technology'.