RESUMO
The anatomy and development of the eyelids in squamate reptiles are still relatively unknown, considering its variation within the group. The neotropical Gymnophthalmini are traditionally characterized by having lost the eyelids, but their structure is not well described. In this study, the embryonic development and the adult morphology of the gymnophthalmid eye, with special attention to the eyelids, the nictitating membrane, and the spectacle are described. The eye in some Gymnophthalmini is covered by a spectacle, formed by the embryonic fusion of the dorsal and ventral eyelids, a character possibly synapomorphic to the tribe. The genus Tretioscincus, which floats either as sister to all other Gymnophthalmini, or is nested within the group, is unique in showing functional and movable eyelids. Thus, the presence of functional eyelids can be either considered as the primitive condition for the gymnophthalmini or as a re-acquisition of the character, showing the importance of a well-established phylogenetic hypothesis for understanding morphological evolution.
Assuntos
Pálpebras/anatomia & histologia , Lagartos/anatomia & histologia , Adaptação Fisiológica , Animais , Evolução Molecular , Pálpebras/embriologia , Pálpebras/fisiologia , Lagartos/embriologia , Lagartos/fisiologia , Filogenia , Especificidade da EspécieRESUMO
Calcified spines in the hemipenial surface occur convergently in several gymnophthalmid lizard species and in advanced snakes. Based on the pronounced degrees of limb reduction in these distantly related lineages, such hemipenial structures were suggested to play a functional role in couple-anchoring during copulation, partly assuming the function of the limbs during mating. Herein, we assessed the hemipenial morphology of virtually all the valid genera of the family Gymnophthalmidae to test for a phylogenetic correlation between limb reduction and the presence of calcified hemipenial spines. The occurrence of calcified structures was mapped on the two most comprehensive phylogenies of the family. We concluded that spiny hemipenes are by no means necessarily associated with reduction of limbs. Conversely, the presence of well-developed hemipenial spines in specific limb-reduced taxa does not allow one to disregard the possibility that in some instances such structures might indeed be functionally associated with couple-anchoring, improving the success of mating.
Assuntos
Evolução Biológica , Extremidades/anatomia & histologia , Lagartos/anatomia & histologia , Pênis/anatomia & histologia , Animais , Calcificação Fisiológica , Copulação , Extremidades/fisiologia , Lagartos/fisiologia , Masculino , Pênis/fisiologia , Filogenia , Especificidade da EspécieRESUMO
The tetrapod limb is a highly diverse structure, and reduction or loss of this structure accounts for many of the limb phenotypes observed within species. Squamate reptiles are one of the many tetrapod lineages in which the limbs have been greatly modified from the pentadactyl generalized pattern, including different degrees of reduction in the number of limb elements to complete limblessness. Even though limb reduction is widespread, the evolutionary and developmental mechanisms involved in the formation of reduced limb morphologies remains unclear. In this study, we present an overview of limb morphology within the microteiid lizard group Gymnophthalmidae, focusing on digit arrangement. We show that there are two major groups of limb-reduced gymnophthalmids. The first group is formed by lizard-like (and frequently pentadactyl) species, in which minor reductions (such as the loss of 1-2 phalanges mainly in digits I and V) are the rule; these morphologies generally correspond to those seen in other squamates. The second group is formed by species showing more drastic losses, which can include the absence of an externally distinct limb in adults. We also present the expression patterns of Sonic Hedgehog (Shh) in the greatly reduced fore and hindlimb of a serpentiform gymnophthalmid. Our discussion focuses on identifying shared patterns of limb reduction among tetrapods, and explaining these patterns and the morphological variation within the gymnophthalmids based on current knowledge of the molecular signaling pathways that coordinate limb development.
Assuntos
Membro Anterior/embriologia , Proteínas Hedgehog/genética , Membro Posterior/embriologia , Lagartos/embriologia , Dedos do Pé/embriologia , Animais , Evolução Biológica , Padronização Corporal/genética , Osso e Ossos/embriologia , Membro Anterior/anatomia & histologia , Membro Posterior/anatomia & histologia , Lagartos/classificação , Filogenia , Transdução de Sinais/genética , Dedos do Pé/anatomia & histologiaRESUMO
Squamates are found in a wide range of habitats and show a corresponding diversity of morphologies that can often be correlated with locomotor mode. The evolution of a snake-like body form, frequently associated with fossoriality, from a typical lacertiform morphology involves changes in the morphology of vertebrae, girdles, and limbs; the changes are mainly manifested by the reduction or loss of limbs and body elongation. In this study, we describe the axial and appendicular skeletons of six closely related gymnophthalmid species. Three of them show a lizard-like morphology, with a four-digit forelimb and a five-digit hindlimb, and the other three show a snake-like morphology associated with a burrowing habit, with reduced limbs and a longer body in comparison to the former three species. We show that vertebral morphology is similar among the six species, with the differences being accounted for by an increase in the number of vertebrae and by the structural reduction of girdles and limbs in the snake-like species. Skeletal morphology provides valuable information on locomotion type, physiology, diet, and other biological features. The burrowing morphology usually involves accentuated reduction of girdle and limb elements, reflecting an undulating type of locomotion in which the limbs play little or no role in propelling the body; in contrast, well-developed limbs and girdles indicate a greater reliance on the limbs for body propulsion. Limb reduction is frequent among vertebrates, but many different phenotypes are found in species exhibiting some kind of reduction, indicating that different mechanisms and evolutionary pressures may be involved in generating the diverse morphologies.
Assuntos
Osso e Ossos/anatomia & histologia , Lagartos/anatomia & histologia , Animais , Evolução Biológica , Ecossistema , Extremidades/anatomia & histologia , Membro Anterior/anatomia & histologia , Membro Posterior/anatomia & histologia , Lagartos/fisiologia , Ossos Pélvicos/anatomia & histologia , Costelas/anatomia & histologia , Serpentes/anatomia & histologia , Especificidade da Espécie , Coluna Vertebral/anatomia & histologiaRESUMO
The evolutionary history of the lizard family Gymnophthalmidae is characterized by several independent events of morphological modifications to a snake-like body plan, such as limb reduction, body elongation, loss of external ear openings, and modifications in skull bones, as adaptive responses to a burrowing and fossorial lifestyle. The origins of such morphological modifications from an ancestral lizard-like condition can be traced back to evolutionary changes in the developmental processes that coordinate the building of the organism. Thus, the characterization of the embryonic development of gymnophthalmid lizards is an essential step because it lays the foundation for future studies aiming to understand the exact nature of these changes and the developmental mechanisms that could have been responsible for the evolution of a serpentiform (snake-like) from a lacertiform (lizard-like) body form. Here we describe the post-ovipositional embryonic development of the fossorial species Nothobachia ablephara and Calyptommatus sinebrachiatus, presenting a detailed staging system for each one, with special focus on the development of the reduced limbs, and comparing their development to that of other lizard species. The data provided by the staging series are essential for future experimental studies addressing the genetic basis of the evolutionary and developmental variation of the Gymnophthalmidae.
Assuntos
Desenvolvimento Embrionário , Extremidades/embriologia , Lagartos/embriologia , Animais , Evolução Biológica , Feminino , Masculino , Microscopia Eletrônica de VarreduraRESUMO
The development of the cartilaginous and bony elements that form the skull and axial and appendicular skeleton is described in detail for the post-ovipositional embryonic development of the fossorial gymnophthalmid species Calyptommatus sinebrachiatus and Nothobachia ablephara. Both species have a snake-like morphology, showing an elongated body and reduced or absent limbs, as well as modifications in skull bones for burrowing, such as complex articulation surfaces and development of bony extensions that enclose and protect the brain. Similar morphological changes have originated independently in several squamate groups, including the one that led to the snake radiation. This study characterizes the patterns of chondrogenesis and osteogenesis, with special emphasis on the features associated with the burrowing habit, and may be used for future comparative analyses of the developmental patterns involved in the origin of the convergent serpentiform morphologies.
Assuntos
Osso e Ossos/embriologia , Condrogênese , Extremidades/embriologia , Lagartos/embriologia , Osteogênese , Animais , Desenvolvimento EmbrionárioRESUMO
Squamates (lizards, snakes and amphisbaenians) are represented by a large number of species distributed among a wide variety of habitats. Changes in body plan related to a fossorial habit are a frequent trend within the group and many morphological adaptations to this particular lifestyle evolved convergently in nonrelated species, reflecting adaptations to a similar habitat. The fossorial lifestyle requires an optimal morphological organization for an effective use of the available resources. Skeleton arrangement in fossorial squamates reflects adaptations to the burrowing activity, and different degrees of fossoriality can be inferred through an analysis of skull morphology. Here, we provide a detailed description of the skull morphology of three fossorial gymnophthalmid species: Calyptommatus nicterus, Scriptosaura catimbau, and Nothobachia ablephara.