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Abstract The lungs of squamate reptiles (lizards and snakes) are highly diverse, exhibiting single chambers, multiple chambers, transitional forms with two to three chambers, along with a suite of other anatomical features, including finger-like epithelial projections into the body cavity known as diverticulae. During embryonic development of the simple, sac-like lungs of anoles, the epithelium is pushed through the openings of a pulmonary smooth muscle mesh by the forces of luminal fluid pressure. This process of stress ball morphogenesis generates the faveolar epithelium typical of squamate lungs. Here, we compared embryonic lung development in brown anoles, leopard geckos, and veiled chameleons to determine if stress ball morphogenesis is conserved across squamates and to understand the physical processes that generate transitional-chambered lungs with diverticulae. We found that epithelial protrusion through the holes in a pulmonary smooth muscle mesh is conserved across squamates. Surprisingly, however, we found that luminal inflation is not conserved. Instead, leopard geckos and veiled chameleons appear to generate their faveolae via epithelial folding downstream of epithelial proliferation. We also found experimental and computational evidence suggesting that the transitional chambers and diverticulae of veiled chameleon lungs develop via apical constriction, a process known to be crucial for airway branching in the bird lung. Thus, distinct morphogenetic mechanisms generate epithelial diversity in squamate lungs, which may underpin their species-specific physiological and ecological adaptations. 

Abstract Among squamates, hemipenes are known to evolve rapidly and exhibit diverse shapes, sizes, and ornamentation. Croaking geckos (Aristelliger) are unique among geckos in exhibiting mineralized structures (hemibacula) in their hemipenes. We here describe the gross morphology of the hemibacula of each currently recognized species ofAristelliger, document hemibacular histology, and report on hemibaculum development. We confirm the presence of hemibacula in all currently recognized species and demonstrate that three distinct morphologies correspond to three putative clades in the genus. Histology revealed that hemibacula are superficially similar to chondroid bone and composed of mineralized dense connective tissue covered in a thin layer of epidermis with alcian‐positive cells embedded within a mineralized matrix. Additionally, we demonstrate that hemibacula do not develop until past the onset of sexual maturity and that hemibaculum length scales isometrically with body size. We hypothesize that hemibacula ofAristelligerdevelop via peramorphosis, a phenomenon also expressed in the cranial morphology of this genus. Additionally, we speculate on the functional significance of these enigmatic structures. 

Synopsis 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. 

Several species of geckos have independently evolved patagia, membranous features that facilitate gliding.Detailed morphological investigations of gecko patagia have largely been limited to gliding members of the genus Gekko(formerly in the genus Ptychozoon). Herein we describe the morphology of gliding patagia of the Flat-tailed HouseGecko (Hemidactylus platyurus), a species with an independent evolutionary origin of gliding patagia from Gekko andan important species for researching gliding biomechanics. We compared morphology of H. platyurus with a closelyrelated non-gliding species, the Common House Gecko (Hemidactylus frenatus). Using external examination and histologicaltechniques, we compared and contrasted three regions that exhibit patagia (trunk, femoral region, and tail)in H. platyurus but not in H. frenatus. We find that patagia in a gliding Hemidactylus, like patagia in gliding membersof the genus Gekko, are derived from expansion of lateral fat bodies, suggesting analogous processes to achieve similarphenotypic outcomes.