Ancestral frogs underwent anatomical shifts including elongation of the hindlimbs and pelvis and reduction of the tail and vertebral column that heralded the transition to jumping as a primary mode of locomotion. Jumping has been hypothesized to have evolved in a step-wise fashion with basal frogs taking-off with synchronous hindlimb extension and crash-landing on their bodies, and then their limbs move forward. Subsequently, frogs began to recycle the forelimbs forward earlier in the jump to control landing. Frogs with forelimb landing radiated into many forms, locomotor modes, habitats, and niches with controlled landing thought to improve escape behavior. While the biology of take-off behavior has seen considerable study, interspecific comparisons of take-off and landing behavior are limited. In order to understand the evolution of jumping and controlled landing in frogs, data are needed on the movements of the limbs and body across an array of taxa. Here, we present the first description and comparison of kinematics of the hindlimbs, forelimbs and body during take-off and landing in relation to ground reaction forces in four frog species spanning the frog phylogeny. The goal of this study is to understand what interspecific differences reveal about the evolution of take-off and controlled landing in frogs. We provide the first comparative description of the entire process of jumping in frogs. Statistical comparisons identify both homologous behaviors and significant differences among species that are used to map patterns of trait evolution and generate hypotheses regarding the functional evolution of take-off and landing in frogs.
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.
Summary While most frogs maximize jump distance as an escape behaviour, toads have traded jump distance for endurance with a strategy of hopping repeatedly. This strategy has enabled toads to expand across the continents as one of the most diverse groups of anurans. Multiple studies have revealed physiological endurance adaptations for sustained hopping in toads, however, the kinematics of their sequential hopping behaviour, per se, has not been studied. We compared kinematics and forces of single hops and multiple hopping sequences and quantified field performance of hopping behaviours in free ranging toads of three species and discovered a novel aspect of locomotion adaptation that adds another facet to their exceptional terrestrial locomotor abilities. We found that bouts of repeated hopping are actually a series of bounding strides where toads rotate on their hands and then land on their extended their feet and jump again without stopping. In addition, free‐ranging toads appear to use bounding locomotion more frequently than single hops. Bounding in toads has the advantage of maintaining velocity and producing longer jump distances. In comparison to single hops, cyclic bounding steps reduce energy expenditure and appear to provide limb loading dynamics better suited for potential cycling of elastic energy from stride to stride than would be possible with repeated single hops. This is the first case of the common use of a bounding gait outside of mammals. Bounding adds a key terrestrial locomotor trait to the toad's phenotype that may help explain their history of global expansion and the challenges to modern faunas as introduced toads rapidly invade new ecosystems today.
1. For arboreal lizards, the ability to cling or adhere to the substrate is critical for locomotion during prey capture, predator escape, thermoregulation and social interactions. Thus, selection on traits related to clinging is likely strong.2. Correlations between morphology, performance and habitat use have been documented in arboreal lizards, providing a framework for using functional traits to predict habitat use in the field.3. We tested the hypothesis that clinging performance predicts habitat use in an actively assembling community of introduced lizards in Hawaiʻi comprised of anoles (Anolis carolinensis, A. sagrei) and day geckos (Phelsuma laticauda).4. We measured morphological traits (toepad area and lamellae number) and tested clinging performance on two artificial and eight natural substrates in the laboratory. We measured habitat use in 10 m × 10 m outdoor enclosures where habitat availability was controlled and the lizard species assemblage was manipulated to reflect all species combinations. The enclosure experiment generated more than 9,000 habitat use observations from 360 lizards. 5. Morphological traits that predict performance in Anolis were not predictive in Phelsuma, indicating that direct measures of performance are necessary for comparisons between the genera. Measuring clinging performance on multiple substrates provided key insights intopatterns of habitat use. While all three species performed best on an artificial smooth substrate (acrylic), performance on natural substrates predicted which texture (rough vs. smooth) was most often used by each species. 7. Performance predicted perch height use: species with the greatest clinging performance (A. carolinensis and P. laticauda) across substrates perched twice as high as A. sagrei. 8. We did not observe habitat shifts in the height or texture of perches used by any species in response to experimental manipulation of the lizard species assemblage. 9. Our results highlight the inextricable link between ecology, morphology and performance, the importance of measuring functional traits in ecologically relevant ways, and the potential for resource partitioning to be influenced by differences in the ability to attach to different substrates.
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