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Claws are a common anatomical feature among limbed amniotes and contribute to a variety of functions including prey capture, locomotion, and attachment. Previous studies of both avian and non-avian reptiles have found correlations between habitat use and claw morphology, suggesting that variation in claw shape permits effective functioning in different microhabitats. How, or if, claw morphology influences attachment performance, particularly in isolation from the rest of the digit, has received little attention. To examine the effects of claw shape on frictional interactions, we isolated the claws of preserved specimens of Cuban knight anoles (Anolis equestris), quantified variation in claw morphology via geometric morphometrics, and measured friction on four different substrates that varied in surface roughness. We found that multiple aspects of claw shape influence frictional interactions, but only on substrates for which asperities are large enough to permit mechanical interlocking with the claw. On such substrates, the diameter of the claw's tip is the most important predictor of friction, with narrower claw tips inducing greater frictional interactions than wider ones. We also found that claw curvature, length, and depth influence friction, but that these relationships depend on the substrate's surface roughness. Our findings suggest that although claw shape plays a critical role in the effective clinging ability of lizards, its relative importance is dependent upon the substrate. Description of mechanical function, as well as ecological function, is critical for a holistic understanding of claw shape variation., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier GmbH. All rights reserved.)

Gekkotan lizards of the genus Hemidactylus exhibit derived digital morphologies. These include heavily reduced antepenultimate phalanges of digits III and IV of the manus and digits III-V of the pes, as well as enigmatic cartilaginous structures called paraphalanges. Despite this well-known morphological derivation, no studies have investigated the development of these structures. We aimed to determine if heterochrony underlies the derived antepenultimate phalanges of Hemidactylus. Furthermore, we aimed to determine if convergently evolved paraphalanges exhibit similar or divergent developmental patterns. Herein we describe embryonic skeletal development in the hands and feet of four gekkonid species, exhibiting a range of digital morphologies. We determined that the derived antepenultimate phalanges of Hemidactylus are the products of paedomorphosis. Furthermore, we found divergent developmental patterns between convergently evolved paraphalanges., (© 2022 Anatomical Society.)

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., (© 2021 Anatomical Society.)

The functional morphology of squamate fibrillar adhesive systems has been extensively investigated and has indirectly and directly influenced the design of synthetic counterparts. Not surprisingly, the structure and geometry of exemplar fibrils (setae) have been the subject of the bulk of the attention in such research, although variation in setal morphology along the length of subdigital adhesive pads has been implicated to be important in the effective functioning of these systems. Adhesive setal field configuration has been described for several geckos, but that of the convergent Anolis lizards, comprised of morphologically simpler fibrils, remains largely unexplored. Here, we examine setal morphology along the proximodistal axis of the digits of Anolis equestris and compare our findings to those for a model gecko, Gekko gecko. Consistent with previous work, we found that the setae of A. equestris are generally thinner, shorter, and present at higher densities than those of G. gecko and terminate in a single spatulate tip. Contrastingly, the setae of G. gecko are hierarchically branched in structure and carry hundreds of spatulate tips. Although the splitting of contacts into multiple smaller tips is predicted to increase the adhesive performance of a fiber compared to an unbranched one, we posited that the adhesive performance of G. gecko and A. equestris would be relatively similar when the configuration of the setal fields of each was accounted for. We found that, as in geckos, setal morphology of A. equestris follows a predictable pattern along the proximodistal axis of the pad, although there are several critical differences in the configuration of the setal fields of these two groups. Most notably, the pattern of variation in setal length of A. equestris is effectively opposite to that exhibited by G. gecko. This difference in clinal variation mirrors the difference in the direction in which the setal fields of anoles and geckos are peeled from the substrate, consistent with the hypothesis that biomechanical factors are the chief determinants of these patterns of variation. Future empirical work, however, is needed to validate this. Our findings set the stage for future comparative studies investigating the functional morphology of these convergent adhesive apparatuses. Such investigations will lead to an enhanced understanding of the interactions between form, function, and environment of fibril-based biological adhesive systems., (© 2020 Anatomical Society.)

External morphological metrics have featured prominently in comparative studies examining the morphological convergence that characterizes anoline ecomorphs. To what degree the appendicular-skeletal morphology of Greater Antillean island Anolis lizards tracks their diversity and ecological adaptation, however, remains relatively unexplored. Here we employ computed tomographic scanning techniques to visualize in situ the scapulocoracoid of ecomorph representatives (trunk-ground, trunk-crown, crown-giant, twig) from three islands (Jamaica, Hispaniola, and Puerto Rico), and compare its three-dimensional geometry using qualitative-descriptive and quantitative-morphometric techniques. In contrast to our previous, similarly-conducted study of the pelvic girdle of these same species, the form of the scapulocoracoid varies markedly both within and between species, with much of the variation relating to phylogenetic relationship, specimen size, and assigned ecomorph category. Morphometric variation that correlates with size and/or phylogenetic signal varies between species and cannot be eliminated from the data set without markedly reducing its overall variability. The discovered patterns of skeletal variation are consistent with the demands of locomotor mechanics imposed by the structural configuration of the microhabitat of these ecomorphs. Most pertinently the ecomorphs differ in the anteroposterior length of the coracoid, the dorsoventral height of the scapulocoracoid, the dorsoventral height of the scapula in relation to the height of the suprascapula, and the relative positioning of the borders of the scapulocoracoid fenestra. In the examined ecomorph categories these skeletal differences likely relate to microhabitat usage by permitting different degrees of tilting and displacement of the scapulocoracoid in the parasagittal plane and influencing the sizes of muscle origins and the vectors of their actions. These differences relate to the amount of humeral adduction applied during its protraction, and to the structural stability of the shoulder girdle during acrobatic maneuvers, thus influencing the perch diameter that can be effectively negotiated, a critical factor in the microhabitat structure of Anolis ecomorphs., (Crown Copyright © 2020. Published by Elsevier GmbH. All rights reserved.)

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., (© 2019 Wiley Periodicals, Inc.)

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., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

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., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

The study of gecko adhesion is necessarily interdisciplinary due to the hierarchical nature of the adhesive system and the complexity of interactions between the animals and their habitats. In nature, geckos move on a wide range of surfaces including soft sand dunes, trees, and rocks, but much of the research over the past two decades has focused on their adhesive performance on artificial surfaces. Exploring the complex interactions between geckos and their natural habitats will reveal aspects of the adhesive system that can be applied to biomimetic research, such as the factors that facilitate movement on dirty and rough surfaces with varying microtopography. Additionally, contrasting suites of constraints and topographies are found on rocks and plants, likely driving differences in locomotion and morphology. Our overarching goals are to bring to light several aspects of ecology that are important for gecko-habitat interactions, and to propose a framework for how they can inspire material scientists and functional ecologists. We also present new data on surface roughness and topography of a variety of surfaces, and adhesive performance of Phelsuma geckos on surfaces of varying roughness. We address the following key questions: (1) why and how should ecology be incorporated into the study of gecko adhesion? (2) What topographical features of rocks and plants likely drive adhesive performance? (3) How can ecological studies inform material science research? Recent advances in surface replication techniques that eliminate confounding factors among surface types facilitate the ability to address some of these questions. We pinpoint gaps in our understanding and identify key initiatives that should be adopted as we move forward. Most importantly, fine details of locomotor microhabitat use of both diurnal and nocturnal geckos are needed., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Recently-developed, molecularly-based phylogenies of geckos have provided the basis for reassessing the number of times adhesive toe-pads have arisen within the Gekkota. At present both a single origin and multiple origin hypotheses prevail, each of which has consequences that relate to explanations about digit form and evolutionary transitions underlying the enormous variation in adhesive toe pad structure among extant, limbed geckos (pygopods lack pertinent features). These competing hypotheses result from mapping the distribution of toe pads onto a phylogenetic framework employing the simple binary expedient of whether such toe pads are present or absent. It is evident, however, that adhesive toe pads are functional complexes that consist of a suite of integrated structural components that interact to bring about adhesive contact with the substratum and release from it. We evaluated the competing hypotheses about toe pad origins using 34 features associated with digit structure (drawn from the overall form of the digits; the presence and form of adhesive scansors; the proportions and structure of the phalanges; aspects of digital muscular and tendon morphology; presence and form of paraphalangeal elements; and the presence and form of substrate compliance-enhancing structures). We mapped these onto a well-supported phylogeny to reconstruct their evolution. Nineteen of these characters proved to be informative for all extant, limbed geckos, allowing us to assess which of them exhibit co-occurrence and/or clade-specificity. We found the absence of adhesive toe pads to be the ancestral state for the extant Gekkota as a whole, and our data to be consistent with independent origins of adhesive toe pads in the Diplodactylidae, Sphaerodactylidae, Phyllodactylidae, and Gekkonidae, with a strong likelihood of multiple origins in the latter three families. These findings are consistent with recently-published evidence of the presence of adhesively-competent digits in geckos generally regarded as lacking toe pads. Based upon morphology we identify other taxa at various locations within the gekkotan tree that are promising candidates for the expression of the early phases of adhesively-assisted locomotion. Investigation of functionally transitional forms will be valuable for enhancing our understanding of what is necessary and sufficient for the transition to adhesively-assisted locomotion, and for those whose objectives are to develop simulacra of the gekkotan adhesive system for biotechnological applications., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Amazonia harbors the greatest biological diversity on Earth. One trend that spans Amazonian taxa is that most taxonomic groups either exhibit broad geographic ranges or small restricted ranges. This is likely because many traits that determine a species range size, such as dispersal ability or body size, are autocorrelated. As such, it is rare to find groups that exhibit both large and small ranges. Once identified, however, these groups provide a powerful system for isolating specific traits that influence species distributions. One group of terrestrial vertebrates, gecko lizards, tends to exhibit small geographic ranges. Despite one exception, this applies to the Neotropical dwarf geckos of the genus Gonatodes. This exception, Gonatodes humeralis, has a geographic distribution almost 1,000,000 km 2 larger than the combined ranges of its 30 congeners. As the smallest member of its genus and a gecko lizard more generally, G. humeralis is an unlikely candidate to be a wide-ranged Amazonian taxon. To test whether or not G. humeralis is one or more species, we generated molecular genetic data using restriction-site associated sequencing (RADseq) and traditional Sanger methods for samples from across its range and conducted a phylogeographic study. We conclude that G. humeralis is, in fact, a single species across its contiguous range in South America. Thus, Gonatodes is a unique clade among Neotropical taxa, containing both wide-ranged and range-restricted taxa, which provides empiricists with a powerful model system to correlate complex species traits and distributions. Additionally, we provide evidence to support species-level divergence of the allopatric population from Trinidad and we resurrect the name Gonatodes ferrugineus from synonymy for this population., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Postnatal growth patterns within the vertebral column may be informative about body proportions and regionalization. We measured femur length, lengths of all pre-sacral vertebrae, and lengths of intervertebral spaces, from radiographs of a series of 21 Eublepharis macularius, raised under standard conditions and covering most of the ontogenetic body size range. Vertebrae were grouped into cervical, sternal, and dorsal compartments, and lengths of adjacent pairs of vertebrae were summed before analysis. Femur length was included as an index of body size. Principal component analysis of the variance-covariance matrix of these data was used to investigate scaling among them. PC1 explained 94.19% of total variance, interpreted as the variance due to body size. PC1 differed significantly from the hypothetical isometric vector, indicating overall allometry. The atlas and axis vertebrae displayed strong negative allometry; the remainder of the vertebral pairs exhibited weak negative allometry, isometry or positive allometry. PC1 explained a markedly smaller amount of variance for the vertebral pairs of the cervical compartment than for the remainder of the vertebral pairs, with the exception of the final pair. The relative standard deviations of the eigenvalues from the PCAs of the three vertebral compartments indicated that the vertebrae of the cervical compartment were less strongly integrated by scaling than were the sternal or dorsal vertebrae, which did not differ greatly between themselves in their strong integration, suggesting that the growth of the cervical vertebrae is constrained by the mechanical requirements of the head. Regionalization of the remainder of the vertebral column is less clearly defined but may be associated with wave form propagation incident upon locomotion, and by locomotory changes occasioned by tail autotomy and regeneration. Femur length exhibits negative allometry relative to individual vertebral pairs and to vertebral column length, suggesting a change in locomotor requirements over the ontogenetic size range., (© 2018 Wiley Periodicals, Inc.)

Ecological niche partitioning of Anolis lizards of the Greater Antillean islands has been the focus of many comparative studies, and much is known about external morphological convergence that characterizes anole ecomorphs. Their internal anatomy, however, has rarely been explored in an ecomorphological context, and it remains unknown to what degree skeletal morphology tracks the diversity and ecological adaptation of these lizards. Herein, we employ CT scanning techniques to visualise the skeleton of the pelvic girdle in situ, and 3D geometric morphometrics to compare the form of the ilium, ischium, and pubis within and between ecomorphs. We examine 26 species of anoles representing four ecomorphs (trunk-ground, trunk-crown, crown-giant, twig) from three islands (Jamaica, Hispaniola, and Puerto Rico). The subtle variations in pelvic girdle morphology discovered are directly associable with all three parameters that we set out to focus on: phylogenetic relationship, specimen size, and assigned ecomorph category. Morphometric variation that correlates with size and/or phylogenetic signal varies between species and cannot be eliminated from the data set without markedly reducing its overall variability. The discovered patterns of skeletal variation are consistent with the demands of locomotor mechanics pertinent to the structural configuration of the microhabitat of three of the four ecomorphs, with the fourth having no discernible distinctive features. This manifests itself chiefly in the relative anteroposterior extent and anteroventral inclination of the ilium and pubis, which differ between ecomorphs and are postulated to reflect optimization of the direction of muscle vectors of the femoral protractors and retractors. Our investigation of the form of the pelvic girdle of anoles allows us to generalize our findings to entire ecomorph categories within a broad phylogenetic and biogeographic context. Differences in the form and configuration of the postcranial skeleton are directly related to ecological patterns., (© 2018 Wiley Periodicals, Inc.)

We published a phylogenetic comparative analysis that found geckos had gained and lost adhesive toepads multiple times over their long evolutionary history (Gamble et al., PLoS One, 7, 2012, e39429). This was consistent with decades of morphological studies showing geckos had evolved adhesive toepads on multiple occasions and that the morphology of geckos with ancestrally padless digits can be distinguished from secondarily padless forms. Recently, Harrington & Reeder (J. Evol. Biol., 30, 2017, 313) reanalysed data from Gamble et al. (PLoS One, 7, 2012, e39429) and found little support for the multiple origins hypothesis. Here, we argue that Harrington and Reeder failed to take morphological evidence into account when devising ancestral state reconstruction models and that these biologically unrealistic models led to erroneous conclusions about the evolution of adhesive toepads in geckos., (© 2017 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2017 European Society For Evolutionary Biology.)

The remarkable adhesive capabilities of geckos have garnered attention from scientists and the public for centuries. Geckos are known to have an adhesive load-bearing capacity far in excess (by 100-fold or more) of that required to support their body mass or accommodate the loading imparted during maximal locomotor acceleration. Few studies, however, have investigated the ecological contexts in which geckos use their adhesive system and how this may influence its properties. Here we develop a modelling framework to assess whether their prodigious adhesive capacity ever comes under selective challenge. Our investigation is based upon observations of escape-induced aerial descents of canopy-dwelling arboreal geckos that are rapidly arrested by clinging to leaf surfaces in mid-fall. We integrate ecological observations, adhesive force measurements, and body size and shape measurements of museum specimens to conduct simulations. Using predicted bending mechanics of petioles and leaf midribs, we find that the drag coefficient of the gecko, the size of the gecko and the size of the leaf determine impact forces. Regardless of the landing surface, safety factors for geckos range from a maximum of just over 10 to a minimum of well under one, which would be the point at which the adhesive system fails. In contrast to previous research that intimates that gecko frictional adhesive capacity is excessive relative to body mass, we demonstrate that realistic conditions in nature may result in frictional capacity being pushed to its limit. The rapid arrest of the lizard from its falling velocity likely results in the maximal loading to which the adhesive system is exposed during normal activities. We suggest that such activities might be primary determinants in driving their high frictional adhesive capacity., (© 2017 The Author(s).)

Anoline lamellae terminate in an epidermal free margin carrying the majority of its setae. How the free margin is extruded from the body of the scale is not well understood. Two hypotheses have been advanced to account for it, one advocating distal migration of the outer epidermal layers relative to the body of the lamella, and the other proposing regression of the dermal core. Available evidence provides partial support for both. We assembled a series of specimens of Anolis grahami representing all shedding cycle stages, and prepared histological sections of the toe pads to allow measurement of appropriate lamellar components through the shedding cycle. Through its proliferative phases the lamellae increase markedly in length, with the distance between the distal tip of the dermal core and that of the lamella accounting for most of this, indicating that epidermal extrusion is responsible for production of the new free margin. The dermal core showed no evidence of regression. Concomitant with epidermal extrusion, the lacunar cells on the inner lamellar face hypertrophy and keep pace with the increasing thickness of the outer lamellar face resulting from the lengthening of the replacement setae. The integrated changes observed are consistent with continuity of functioning and alignment of the exposed setal carpet of the outer epidermal generation while ensuring that the new setal carpet is fully aligned and functional immediately after shedding. At shedding the original proportions of the lamellae are restored. Development of the new free margin results from a combination of distal displacement of Oberhäutchen cells along with arrested maturation of the epidermis in this region. Changes in length of the lamellae during the proliferative stages may impact the overall size of the adhesive toe pad, which may have consequences for assessments of the relationship between whole animal clinging ability and adhesive pad area. J. Morphol. 278:360-368, 2017. © 2017 Wiley Periodicals, Inc., (© 2017 Wiley Periodicals, Inc.)

Post-cloacal bones of gekkotans may be present as a single (medial) pair, two pairs (medial and lateral), or may be lacking. We, herein, demonstrate that the presence of a single medial pair is the ancestral condition for the Gekkota, that the lateral pair is of sporadic occurrence within and between families, except for the Eublepharidae where it is universal, and that absence is also of sporadic occurrence except for the Sphaerodactylidae where it is the ancestral condition. Adult male Tokay geckos (Gekko gecko) possess only the medial pair of bones, and these exhibit a regionally-specific expression of woven, fibrolamellar, and lamellar bone, and an enclosed medullary cavity. Females and small juvenile males lack bony elements but exhibit a conspicuous band of dense connective tissue located about the anterior and lateral margins of the cloacal sacs. As males grow and attain sexual maturity, the medial post-cloacal bones condense in this band of dense connective tissue, and are thus shown to be dermal ossifications, similar to osteoderms but with muscular associations (although this is also known for crocodylians). Based upon ontogenetic data we set forth a scenario to explain the loss of the medial post-cloacal bones in various lineages. Differential staining of the cloacal sacs failed to reveal any specialized glandular structures. Investigation of the post-cloacal spurs shows them to be associated with cellular connective tissue of a type similar to that found in the vicinity of the medial post-cloacal bones. This suggests that the lateral post-cloacal bones may also be dermal bones, but histological evidence is needed to corroborate this., (© 2015 Wiley Periodicals, Inc.)

Although the phenomenon of adhesion in geckos has been intensively studied for over 200 years, our understanding of how the morphological apparatus associated with this arose is less clear. Indeed, whether or not all of the intricate morphological hierarchy that is implicated in the attachment and removal of the adhesive setae originated at the same time is unknown. To explore whether setae may have arisen prior to the other parts of this structural hierarchy, we undertook morphological observations of Gonatodes, an ancestrally padless, sphaerodatyline genus known to exhibit the expression of incipient subdigital pads in some species. Focusing on this geographically and morphologically well-circumscribed genus, for which intraspecific relationships are adequately known and ecology is quite well documented, allowed us to deduce trends in digit proportions, shape, scalation, and skeletal structure, and associate these with the micro-ornamentation of the subdigital surfaces. Our findings indicate that in Gonatodes, setae capable of inducing adhesion are present without the modifications of the digital musculotendinous, circulatory and skeletal systems that are generally considered to be necessary for the operation of a functional adhesive apparatus. The acquisition of these latter characteristics (independently in many lineages of gekkotans, and incipiently so in Anolis) may have been preceded by a suite of modifications of the digits that enhanced static clinging in relation to sit-and-wait predation and the ability to take refuge on surfaces unavailable to other taxa. These possibilities await further testing., (© 2015 Wiley Periodicals, Inc.)

Osteoderms are bone-rich organs found in the dermis of many scleroglossan lizards sensu lato, but are only known for two genera of gekkotans (geckos): Tarentola and Gekko. Here, we investigate their sequence of appearance, mode of development, structural diversity and ability to regenerate following tail loss. Osteoderms were present in all species of Tarentola sampled (Tarentola annularis, T. mauritanica, T. americana, T. crombei, T. chazaliae) as well as Gekko gecko, but not G. smithii. Gekkotan osteoderms first appear within the integument dorsal to the frontal bone or within the supraocular scales. They then manifest as mineralized structures in other positions across the head. In Tarentola and G. gecko, discontinuous clusters subsequently form dorsal to the pelvis/base of the tail, and then dorsal to the pectoral apparatus. Gekkotan osteoderm formation begins once the dermis is fully formed. Early bone deposition appears to involve populations of fibroblast-like cells, which are gradually replaced by more rounded osteoblasts. In T. annularis and T. mauritanica, an additional skeletal tissue is deposited across the superficial surface of the osteoderm. This tissue is vitreous, avascular, cell-poor, lacks intrinsic collagen, and is herein identified as osteodermine. We also report that following tail loss, both T. annularis and T. mauritanica are capable of regenerating osteoderms, including osteodermine, in the regenerated part of the tail. We propose that osteoderms serve roles in defense against combative prey and intraspecific aggression, along with anti-predation functions., (© 2015 Wiley Periodicals, Inc.)

Innovations permit the diversification of lineages, but they may also impose functional constraints on behaviors such as locomotion. Thus, it is not surprising that secondary simplification of novel locomotory traits has occurred several times among vertebrates and could potentially lead to exceptional divergence when constraints are relaxed. For example, the gecko adhesive system is a remarkable innovation that permits locomotion on surfaces unavailable to other animals, but has been lost or simplified in species that have reverted to a terrestrial lifestyle. We examined the functional and morphological consequences of this adaptive simplification in the Pachydactylus radiation of geckos, which exhibits multiple unambiguous losses or bouts of simplification of the adhesive system. We found that the rates of morphological and 3D locomotor kinematic evolution are elevated in those species that have simplified or lost adhesive capabilities. This finding suggests that the constraints associated with adhesion have been circumvented, permitting these species to either run faster or burrow. The association between a terrestrial lifestyle and the loss/reduction of adhesion suggests a direct link between morphology, biomechanics, and ecology.

Animals can undergo significant weight change for a variety of reasons. Autotomy, the voluntary shedding of an appendage in response to a predator stimulus, provides an effective model for measuring the effects of rapid weight change on locomotor behavior and the responses to more gradual weight gain, particularly in lizards capable of both autotomizing and regenerating their tail. Although the general effects of autotomy on locomotor performance are commonly explored, we investigated changes in locomotor mechanics associated with tail loss and long-term regeneration for the first time by measuring morphology, 3D kinematics and ground reaction forces (GRFs) in the leopard gecko Eublepharis macularius. Tail autotomy resulted in a 13% anterior shift in the center of mass (CoM), which only partially recovered after full regeneration of the tail. Although no changes in body or forelimb kinematics were evident, decreases in hindlimb joint angles signify a more sprawled posture following autotomy. Changes in hindlimb GRFs resulted in an increase in weight-specific propulsive force, without a corresponding change in locomotor speed. Hindlimb kinematics and GRFs following autotomy recovered to pre-autotomy values as the tail regenerated. These results suggest an active locomotor response to tail loss that demonstrates the causal relationships between variations in morphology, kinematics and force., (© 2014. Published by The Company of Biologists Ltd.)

The lizard tail is well known for its ability to autotomize and regenerate. Physical contact of the tail by a predator may induce autotomy at the location at which the tail is grasped, and upon detachment the tail may undergo violent, rapid, and unpredictable movements that appear to be, to some degree, regulated by contact with the physical environment. Neither the mechanism by which tail breakage at a particular location is determined, nor that by which environmental feedback to the tail is received, are known. It has been suggested that mechanoreceptors (sensilla) are the means of mediation of such activities, and reports indicate that the density of sensilla on the tail is high. To determine the feasibility that mechanoreceptors are involved in such phenomena, we mapped scale form and the size, density, distribution, and spacing of sensilla on the head, body, limbs, and tail of the leopard gecko. This species has a full complement of autotomy planes along the length of the tail, and the postautotomic behavior of its tail has been documented. We found that the density of sensilla is highest on the tail relative to all other body regions examined; a dorsoventral gradient of caudal sensilla density is evident on the tail; sensilla are more closely spaced on the dorsal and lateral regions of the tail than elsewhere and are carried on relatively small scales; and that the whorls of scales on the tail bear a one to one relationship with the autotomy planes. Our results are consistent with the hypotheses of sensilla being involved in determining the site at which autotomy will occur, and with them being involved in the mediation of tail behavior following autotomy. These findings open the way for experimental neurological investigations of how autotomy is induced and how the detached tail responds to external environmental input., (© 2014 Wiley Periodicals, Inc.)

The breast-shoulder apparatus (BSA) is a structurally and kinematically complex region of lizards. Compared with the pelvic region it has received little attention, even though its morphological variation is known to be extensive. This variability has seldom been the focus of functional explanation, possibly because the BSA has been difficult to explore as a composite entity. In this study we apply geometric morphometric techniques to the analysis of the BSA in an attempt to more fully understand its configuration in relation to differential use in locomotion. Our approach centers upon the Jamaican radiation of anoline lizards (genus Norops) as a tractable, small monophyletic assemblage consisting of species representing several ecomorphs. We hypothesized that the different species and ecomorphs would exhibit variation in the configuration of the BSA. Our findings indicate that this is so, and is expressed in the component parts of the BSA, although it is subtle except for Norops valencienni (twig ecomorph), which differs greatly in morphology (and behavior) from its island congeners. We further found similarities in the BSA of N. grahami, N. opalinus (both trunk-crown ecomorphs), and N. garmani (crown giant). These outcomes are promising for associating morphology with ecomorphological specialization and for furthering our understanding of the adaptive response of the BSA to demands on the locomotor system., (Copyright © 2014 Wiley Periodicals, Inc.)

The striking morphological convergence of hair-like integumentary derivatives of lizards and arthropods (spiders and insects) demonstrates the importance of such features for enhancing purchase on the locomotor substrate. These pilose structures are responsible for the unique tractive abilities of these groups of animals, enabling them to move with seeming ease on overhanging and inverted surfaces, and to traverse inclined smooth substrates. Three groups of lizards are well known for bearing adhesion-promoting setae on their digits: geckos, anoles and skinks. Similar features are also found on the ventral subdigital and distal caudal skin of chameleons. These have only recently been described in any detail, and structurally and functionally are much less well understood than are the setae of geckos and anoles. The seta-like structures of chameleons are not branched (a characteristic of many geckos), nor do they terminate in spatulate tips (which is characteristic of geckos, anoles and skinks). They are densely packed and have attenuated blunt, globose tips or broad, blade-like shafts that are flattened for much of their length. Using a force transducer, we tested the hypothesis that these structures enhance friction and demonstrate that the pilose skin has a greater frictional coefficient than does the smooth skin of these animals. Our results are consistent with friction being generated as a result of side contact of the integumentary filaments. We discuss the evolutionary and functional implications of these seta-like structures in comparison with those typical of other lizard groups and with the properties of seta-mimicking synthetic structures.

Caudal autotomy is a widespread phenomenon among lizards, and similar processes occur in other groups of vertebrates and invertebrates. Many costs have been associated with autotomy, including the regeneration of lost biomass. For lizards, it is not known whether resources are preferentially directed toward caudal regeneration or whether regeneration occurs only when resources are abundant. Conflicting information is present in the literature, and an absence of controlled experiments prevents determination of what pattern of regeneration may occur under a given set of circumstances. We employed the leopard gecko, a fat-tailed species, to examine whether tail regeneration is a priority and, if so, whether it remains so when resources become limiting. We explored this through caudal autotomy and dietary manipulation under conditions that ensured that differences in diet were sufficient to permit differential growth. We examined juvenile leopard geckos because these animals are rapidly growing and allocation of energy is not compromised by reproductive investment. The effects of dietary resource availability and the demands of caudal regeneration were compared in intact and regenerating animals. Our evidence indicates that caudal regeneration is a priority, even when resources are limiting. We conclude that tail regrowth is a priority that is associated with long-term survival and possibly reproductive success.

The ability to shed an appendage occurs in both vertebrates and invertebrates, often as a tactic to avoid predation. The tails of lizards, unlike most autotomized body parts of animals, exhibit complex and vigorous movements once disconnected from the body. Despite the near ubiquity of autotomy across groups of lizards and the fact that this is an extraordinary event involving the self-severing of the spinal cord, our understanding of why and how tails move as they do following autotomy is sparse. We herein explore the histochemistry and physiology of the tail muscles of the leopard gecko (Eublepharis macularius), a species that exhibits vigorous and variable tail movements following autotomy. To confirm that the previously studied tail movements of this species are generally representative of geckos and therefore suitable for in-depth muscle studies, we quantified the three-dimensional kinematics of autotomized tails in three additional species. The movements of the tails of all species were generally similar and included jumps, flips, and swings. Our preliminary analyses suggest that some species of gecko exhibit short but high-frequency movements, whereas others exhibit larger-amplitude but lower-frequency movements. We then compared the ATPase and oxidative capacity of muscle fibers and contractile dynamics of isolated muscle bundles from original tails, muscle from regenerate tails, and fast fibers from an upper limb muscle (iliofibularis) of the leopard gecko. Histochemical analysis revealed that more than 90% of the fibers in original and regenerate caudal muscles had high ATPase but possessed a superficial layer of fibers with low ATPase and high oxidative capacity. We found that contraction kinetics, isometric force, work, power output, and the oscillation frequency at which maximum power was generated were lowest in the original tail, followed by the regenerate tail and then the fast fibers of the iliofibularis. Muscle from the original tail exhibited greater resistance to fatigue, followed by the regenerate tail and then the fast iliofibularis fibers. These results suggest that the relatively slow and oxidative fibers found within the tail musculature have a significant impact on contractile function, which translates into a trade-off between longevity of performance and power after autotomy.

Self-amputation (autotomy) of the tail is essential for the survival of many lizards. Accordingly, it has garnered the attention of scientists for more than 200 years. Several factors can influence the release of the tail, such as the size, sex, and age of the lizard; type of predator; ecology; and evolutionary history of the lineage. Once lost, the tail will writhe for seconds to minutes, and these movements likely depend on the size and physiology of the tail, habitat of the lizard, and predation pressure. Loss of the tail will, in turn, have impacts on the lizard, such as modified locomotor performance and mechanics, as well as escape behavior. However, the tail is almost always regenerated, and this involves wound healing, altered investment of resources, and tissue differentiation. The regenerated tail generally differs from the original in several ways, including size, shape, and function. Here we summarize recent findings of research pertaining to tail autotomy, and we propose a framework for future investigations.

The persistence of the notochord into the skeletally mature life stage is characteristic of gekkotans, but is otherwise of rare occurrence among amniotes. The taxonomic diversity of Gekkota affords the opportunity to investigate the structure and development of this phylogenetically ancestral component of the skeleton, and to determine its basic characteristics. The gekkotan notochord spans almost the entire postcranial long axis and is characterized by a moniliform morphology with regularly alternating zones of chordoid and chondroid tissue. Chordoid tissue persists in the region of intervertebral articulations and occupies the cavitations that lie between the centra of the amphicoelous vertebrae. Chondroid tissue is restricted to zones in which the diameter of the notochord is reduced, corresponding to mid-vertebral locations. In the tail, these zones of chondroid tissue are associated with the autotomic fracture planes. Chondroid tissue first manifests during late embryogenesis, appears to differentiate from pre-existing chordoid tissue, and has the histological and histochemical characteristics of cartilage. Our observations lend support to the hypothesis that cartilage can be derived directly from notochordal tissue, and suggest that the latter may be an evolutionary and developmental precursor to chordate cartilage. The persistence of chordoid tissue in the intervertebral regions of amphicoelous vertebrae is consistent with a suite of paedomorphic traits exhibited by gekkotans and suggests that the typical hydrostatic nature of notochordal tissue may play a role in mechanically governing patterns of displacement between adjacent amphicoelous vertebrae that lack extensive centrum-to-centrum contact., (Copyright © 2012 Wiley Periodicals, Inc.)

Geckos are well known for their extraordinary clinging abilities and many species easily scale vertical or even inverted surfaces. This ability is enabled by a complex digital adhesive mechanism (adhesive toepads) that employs van der Waals based adhesion, augmented by frictional forces. Numerous morphological traits and behaviors have evolved to facilitate deployment of the adhesive mechanism, maximize adhesive force and enable release from the substrate. The complex digital morphologies that result allow geckos to interact with their environment in a novel fashion quite differently from most other lizards. Details of toepad morphology suggest multiple gains and losses of the adhesive mechanism, but lack of a comprehensive phylogeny has hindered efforts to determine how frequently adhesive toepads have been gained and lost. Here we present a multigene phylogeny of geckos, including 107 of 118 recognized genera, and determine that adhesive toepads have been gained and lost multiple times, and remarkably, with approximately equal frequency. The most likely hypothesis suggests that adhesive toepads evolved 11 times and were lost nine times. The overall external morphology of the toepad is strikingly similar in many lineages in which it is independently derived, but lineage-specific differences are evident, particularly regarding internal anatomy, with unique morphological patterns defining each independent derivation.

Five nominal elements comprise the circumorbital series of bones in gekkotans: prefrontal, postfrontal, postorbital, jugal, and lacrimal. Determination of the homology of two of these, the postfrontal and postorbital, has been particularly problematic. Two conflicting hypothesis exist relating to these: either the postorbital is lost and the postfrontal remains or they fuse during development to form a combined element, the postorbitofrontal. Such a combined element apparently occurs in at least some members of all lizard clades. There is, however, no direct developmental evidence that supports either theory. To overcome that, we investigate the sequence and pattern of ossification in the circumorbital region in a developmental series of the Leopard gecko. We posit that both the postfrontal and postorbital appear during development. Contrary to previous predictions they neither fuses to each other, nor do either degenerate. Instead, the postfrontal shifts anteriorly and fuses with the frontal to become indistinguishable from it by the time of hatching, and the postorbital persists as a robust independent element bounding the frontoparietal suture. These observations accord, in part, with both hypotheses of homology of these elements and result in the recognition of a new pattern, placing in doubt the existence of the composite postorbitofrontal. The phylogenetic implications of these findings may prove to be far reaching if similar and conserved patterns of development are encountered in other clades.

Many animals lose and regenerate appendages, and tail autotomy in lizards is an extremely well-studied example of this. Whereas the energetic, ecological and functional ramifications of tail loss for many lizards have been extensively documented, little is known about the behaviour and neuromuscular control of the autotomized tail. We used electromyography and high-speed video to quantify the motor control and movement patterns of autotomized tails of leopard geckos (Eublepharis macularius). In addition to rhythmic swinging, we show that they exhibit extremely complex movement patterns for up to 30 min following autotomy, including acrobatic flips up to 3 cm in height. Unlike the output of most central pattern generators (CPGs), muscular control of the tail is variable and can be arrhythmic. We suggest that the gecko tail is well suited for studies involving CPGs, given that this spinal preparation is naturally occurring, requires no surgery and exhibits complex modulation.

Lizards commonly climb in complex three-dimensional habitats, and gekkotans are particularly adept at doing this by using an intricate adhesive system involving setae on the ventral surface of their digits. However, it is not clear whether geckos always deploy their adhesive system, given that doing so may result in decreased (i.e. reduction in speed) locomotor performance. Here, we investigate circumstances under which the adhesive apparatus of clinging geckos becomes operative, and examine the potential trade-offs between speed and clinging. We quantify locomotor kinematics of a gecko with adhesive capabilities (Tarentola mauritanica) and one without (Eublepharis macularius). Whereas, somewhat unusually, E. macularius did not suffer a decrease in locomotor performance with an increase in incline, T. mauritanica exhibited a significant decrease in speed between the level and a 10 degrees incline. We demonstrate that this results from the combined influence of slope and the deployment of the adhesive system. All individuals kept their digits hyperextended on the level, but three of the six individuals deployed their adhesive system on the 10 degrees incline, and they exhibited the greatest decrease in velocity. The deployment of the adhesive system was dependent on incline, not surface texture (600 grit sandpaper and Plexiglas), despite slippage occurring on the level Plexiglas substrate. Our results highlight the type of sensory feedback (gravity) necessary for deployment of the adhesive system, and the trade-offs associated with adhesion.

Squamates constitute a major vertebrate radiation, representing almost one-third of all known amniotes. Although speciose and morphologically diverse, they remain poorly represented in developmental studies. Here, we present an embryonic staging table of in ovo development for the basal gekkotan Eublepharis macularius (the leopard gecko) and advocate this species as a laboratory-appropriate developmental model. E. macularius, is a hardy and tractable species of relatively large body size (with concomitantly relatively large eggs and embryos), that is widely available and easy to maintain and propagate. Additionally, E. macularius displays a body plan appropriate to the study of the plesiomorphic quadrupedal condition of early pentadactylous terrestrial amniotes. Although not unexpected, it is worth noting that the morphological events characterizing limb development in E. macularius are comparable with those described for the avian Gallus gallus. Therefore, E. macularius holds great promise as a model for developmental studies focusing on pentadactyly and the formation of digits. Furthermore, it is also attractive as a developmental model because it demonstrates temperature-dependent sex determination. The staging table presented herein is based on an all-female series and represents the entire 52 day in ovo period. Overall, embryogenesis of E. macularius is similar to that of other squamates in terms of developmental stage attained at the time of oviposition, patterns of limb and pharyngeal arch development, and features of the appearance of scalation and pigmentation, indicative of a conserved developmental program., ((c) 2009 Wiley-Liss, Inc.)

Many gekkotans possess seta-bearing adhesive subdigital pads. Details of setal structure, however, are largely based upon putatively exemplary fibrils deemed typical of the species. Little is known of the pattern of configuration of the setae across the subdigital pads and how great, if any, the variance in structure and dimensions is. To understand setal fields as functional entities, as opposed to individual setae, it is necessary to consider this pattern. Additionally, gekkotans within individual radiations occupy different environments and potentially are substrate-specific in terms of the locomotor surface exploited. To investigate these issues, we herein examine the configuration and dimensions of seven species of the gekkotan genus Rhoptropus, and an outgroup taxon, Chondrodactylus bibronii. All of these taxa are rupicolous and the array of rock surfaces exploited by this cluster of taxa is extensive. Our results show that setal field configuration follows a predictable pattern, both from one digit to another within a species, and between homologous digits and anatomical locations between species. One species, Rhoptropus afer, a more terrestrial taxon, exhibits significantly shorter setae and a smaller subdigital pad area than do its congeners, but exhibits the same overall pattern of setal arrangement. Our findings have implications for the understanding of the evolution of adhesive structures, and for the principles used for generating and manufacturing biomimetic artificial microfibrillar arrays.

The sprawling pattern of locomotion in lizards is kinematically intriguing and is underpinned by a distinctive pattern of appendicular morphology. The statics of the sprawling posture dictate fundamental design principles, and these place constraints on the three-dimensional kinematics of the limbs and body axis as locomotion is effected. The fore and hind limbs accommodate these constraints and dictates in fundamentally similar, but positionally different ways, resulting in different kinematic profiles for these two appendages. Recent kinematic investigations have helped to clarify earlier generalizations about lizard locomotion and have revealed that kinematic patterns are more variable than was previously supposed. Such analyses, and attendant detailed studies of the anatomy of the locomotor system, promise a new synthesis and enhanced understanding of evolutionary patterns of locomotion of lizards and adjustment to various locomotor substrata and modes of progression.

Ptychozoon kuhli is known for its parachuting/gliding capabilities. In this contribution, we document the allometric scaling properties of its patagium, accessory flaps and folds, and its total body surface area and compare them to similar attributes of the agamine lizard Draco volans. Ptychozoon kuhli has passive patagia that lack skeletal support and muscular control. Patagial area in P. kuhli is smaller than that in D. volans for individuals of identical snout-vent length, but the accessory folds and flaps compensate for this shortfall and overall P. kuhli has equivalent total body surface area to D. volans for equally sized individuals. The structure of the patagium of Ptychozoon kuhli was investigated in terms of its scalation patterns and structural integrity, its relationship to the body wall and its mechanism of erection, and the distribution of mechanoreceptive sensilla across its surface. Scalation patterns and the internal architecture of the patagium indicate how its shape and form are maintained as it erects and becomes exposed to air flow. Its cross-sectional shape, together with that of the entire body indicates how it is able to behave as an airfoil. The distribution of sensilla across the patagial surface reflect positioning indicative of the monitoring of scale-to-scale contact on the dorsal surface, and possibly air pressure and flow on the ventral surface., (Copyright 2001 Wiley-Liss, Inc.)

The structural variation of the gekkonid larynx and trachea is examined within a representative subset of 17 species of Afro-Madagascan gekkonines to determine if there are underlying morphological correlates of vocalization. The documented morphology is compared to that of the tokay (Gekko gecko), which has previously been described. Data were obtained from gross anatomical observations, scanning electron microscopy, histological examinations and computer-generated, three-dimensional, skeletal reconstructions. Although there is limited variation among most Afro-Malagasy gekkonids, the larynges of Ptenopus garrulus and Uroplatus fimbriatus exhibit marked degrees of differentiation, suggesting that laryngeal and tracheal morphology may account for the documented vocal variability of gekkonid lizards. Cladistic analyses indicated that parallel adaptive trends characterize the laryngeal morphology of the examined taxa. Alternate designs and refinements to a model of gekkonid phonation are presented, and the evolution of acoustic communication in the Gekkonidae is considered., (Copyright 2000 Wiley-Liss, Inc.)

A novel technique for modeling microscopic anatomical structures in three dimensions was developed as part of a survey of gekkonid laryngeal skeletal morphology (Reptilia: Gekkonidae). Excised larynges were transversely sectioned at 10 microns and stained using standard procedures. With a projection microscope, outline drawings of the sectioned laryngeal cartilages were made at regular intervals, depending on the rate and degree of structural change observed while sampling. The drawing set was digitized with a flatbed scanner, and aligned using 'NIH Image' for Macintosh computers. Physical connectivity between successive outlines was provided by inserting one or more artificial slices between those that had been digitized, and draping a skin of rendered contours over all of the interstitial spaces present in the template. The Application Visualization System, a general purpose visualization package for UNIX-based computer systems, was used to visualize and render the resulting 'isosurfaces', which appear as solid three-dimensional objects and can be viewed from any perspective. Since isosurfaced reconstructions can be based on as little as 20% of the cross-sections available, this procedure has the potential to be a valuable research tool for future morphological work at the microscopic level.

The lungs of the New Caldeonian gecko Rhacodactylus leachianus were examined by means of gross dissection and light and electron microscopy. This tropical species, which is the largest living gecko, possesses two simple, single-chambered lungs. Right and left lungs are of similar size and shape. The lung volume (27.2 ml.100 g-1) is similar to that of the tokay (Gekko gecko) but differs in that the gas exchange tissue is approximately homogeneously distributed, and the parenchymal units (ediculae) are very large, approximately 2 mm in diameter. The parenchymal depth varies according to the location in the lung, being deepest near the middle of the lung and shallowest caudally. Scanning and transmission electron microscopy reveal an unusual distribution of ciliated cells in patches on the edicular walls as well as on the trabeculae. Secretory cells are very numerous, particularly in the bronchial epithelium, where they greatly outnumber the ciliated cells. The secretory cells form a morphological continuum characterized by small secretory droplets apically and large vacuoles basally. This continuum includes cells resembling type II pneumocytes but which are devoid of lamellar bodies. Type I pneumocytes similar to those of other reptiles cover the respiratory capillaries, where they form a thin, air-blood barrier together with the capillary endothelial cells and the fused basement laminae. The innervation, musculature, and vascular distribution in R. leachianus are also characterized. Apparent simplification of the lungs in this taxon may be related to features of its sluggish habits, whereas peculiarities of cell and tissue composition may reflect demands of its mesic habitat.