Despite the wide variety of adaptive modifications in the oral and facial regions of vertebrates, their early oropharyngeal development is considered strictly uniform. It involves sequential formation of the mouth and pharyngeal pouches, with ectoderm outlining the outer surface and endoderm the inner surface, as a rule. At the extreme anterior domain of vertebrate embryos, the ectoderm and endoderm directly juxtapose and initial development of this earliest ecto-endoderm interface, the primary mouth, typically involves ectodermal stomodeal invagination that limits the anterior expansion of the foregut endoderm. Here we present evidence that in embryos of extant non-teleost fishes, oral (stomodeal) formation is preceded by the development of prominent pre-oral gut diverticula (POGD) between the forebrain and roof of the forming mouth. Micro-computed tomography (micro-CT) imaging of bichir, sturgeon and gar embryos revealed that foregut outpocketing at the pre-oral domain begins even before the sequential formation of pharyngeal pouches. The presence of foregut-derived cells in the front of the mouth was further confirmed by in vivo experiments that allowed specific tracing of the early endodermal lining. We show that POGD in sturgeons contribute to the orofacial surface of their larvae, comprising oral teeth, lips, and sensory barbels. To our knowledge, this is the first thorough evidence for endodermal origin of external craniofacial structures in any vertebrate. In bichir and gar embryos, POGD form prominent cranial adhesive organs that are characteristic of the ancient bauplan of free-living chordate larvae. POGD hence seem arguably to be ancestral for all ray-finned fishes, and their topology, pharyngeal-like morphogenesis and gene expression suggest that they are evolutionarily related to the foregut-derived diverticula of early chordate and hemichordate embryos. The formation of POGD might thus represent an ancestral developmental module with deep deuterostome origins.
Cercosaurine lizards (subfamily Cercosaurinae of the family Gymnophthalmidae) represent a substantial component of the reptile fauna in the Neotropics. Several attempts have been made to reconstruct the phylogenetic relationships within this group, but most studies focused on particular genera or regions and did not cover the subfamily as a whole. In this study, material from the montane forests of Peru was newly sequenced. In combination with all cercosaurine sequences available on GenBank, an updated phylogeny of Cercosaurinae is provided. Monophyly was not supported for three of the currently recognised genera (Echinosaura, Oreosaurus, and Proctoporus). The genus Proctoporus is formed by five monophyletic groups, which should be used in future taxonomic revisions as feasible entities. Recognition of two previously identified undescribed clades (Unnamed clades 2 and 3) was supported and yet another undescribed clade (termed here Unnamed clade 4), which deserves recognition as an independent genus, was identified herein. Selvasaura brava, a new genus and new species of arboreal gymnophthalmid lizard is described from the montane forests of the Pui Pui Protected Forest, Provincia de Chanchamayo, Región Junín, Peru. The new species is characterised by its small size (SVL 42.1–45.9 mm), slender body, smooth head shields, presence of paired prefrontal shields, fused anteriormost supraocular and anteriormost superciliary shields, transparent not divided lower palpebral disc, slightly rugose subimbricate rectangular dorsal scales in adults (slightly keeled in juveniles), distinctly smaller but non-granular lateral scales, smooth squared to rectangular ventral scales, and hemipenial lobes large, distinct from the hemipenial body. Phylogenetic affinities of the new genus to the other cercosaurine genera, as well as basal phylogenetic relationships between the other cercosaurine genera remain unresolved.
In most vertebrates, pharyngeal arches form in a stereotypic anterior-to-posterior progression. To gain insight into the mechanisms underlying evolutionary changes in pharyngeal arch development, here we investigate embryos and larvae of bichirs. Bichirs represent the earliest diverged living group of ray-finned fishes, and possess intriguing traits otherwise typical for lobe-finned fishes such as ventral paired lungs and larval external gills. In bichir embryos, we find that the anteroposterior way of formation of cranial segments is modified by the unique acceleration of the entire hyoid arch segment, with earlier and orchestrated development of the endodermal, mesodermal, and neural crest tissues. This major heterochronic shift in the anteroposterior developmental sequence enables early appearance of the external gills that represent key breathing organs of bichir free-living embryos and early larvae. Bichirs thus stay as unique models for understanding developmental mechanisms facilitating increased breathing capacity.
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