RESUMO
Advanced endoglyptodont snakes share a complex but homologous venom delivery system associated with the upperjaw and its dentition. Recently, a remarkable novel lower jaw venom delivery system was described for the Neotropicaldipsadine radiation of goo-eating snakes. While most dipsadines are opistoglyphous and exhibit large, mainly serousvenom glands associated with the upper jaw and supralabial glands, goo-eating dipsadine snakes are aglyphous and lackserous upper labial venom glands. Here, we provide new morphological and histological information on the oral glandsand maxillary dentition of representatives of the major lineages of dipsadines that help trace the evolutionary steps thatshaped the venom delivery system of dipsadines. We performed a maximum likelihood analysis on a molecular datasetthat includes 443 terminals and seven loci. Our results show that goo-eating dipsadines form a monophyletic assemblagethat includes the genusAdelphicosfor the first time, along withGeophis,Atractus,Ninia,Chersodromus,Tropidodipsas,Sibon, andDipsas. We also provide the first evidence of a complete shift from an upper jaw to a lower jaw venomdelivery system associated with their specialized feeding behaviour. Unlike other dipsadines who exhibit typicalendoglyptodont anteroposteriorly ridged posterior maxillary teeth, goo-eating dipsadines have uniform lateromediallyridged teeth throughout their maxilla. Our results indicate that the loss of the endoglyptodont venom delivery systemoccurred in the most recent common ancestor of goo-eating dipsadines, probably resulting from the loss of theembryonic posterior maxillary lamina responsible for the development of the venom delivery system.
RESUMO
Geophis belongs to the goo-eating dipsadine assemblage of snakes that are known to feed exclusively on earthworms, snails, and slugs. Although the unusual feeding strategies of the goo-eating dipsadines are well known (but poorly documented), little attention has been paid to their internal anatomy. Here, we describe a new and noteworthy morphological and histochemical condition of the infralabial glands in three species of Geophis (G. brachycephalus, G. nasalis and G. semidoliatus), all earthworm feeders. Their infralabial glands are constituted of two distinct parts: an anterolateral portion composed of mucous and seromucous cells that stretches from the tip of the dentary to the corner of the mouth, and a tubular posteromedial portion that is exclusively seromucous. The anterolateral portion receives fibers of the levator anguli oris muscle that attaches on its posterodorsal extremity while the posteromedial portion extends posteriorly to the corner of the mouth where it receives fibers of the adductor mandibulae externus medialis muscle. Furthermore, the posteromedial portion of the infralabial gland is constituted by large acini filled with secretion that is periodic acid-Schiff positive. These acini release their secretion directly into a large lumen located in the middle of the glandular portion. In the three species examined, the supralabial glands show a traditional configuration, being constituted of mucous and seromucous cells and retaining an enlarged part in its caudal region that resembles a Duvernoy's gland. The presence in Geophis of an expanded lumen in part of the infralabial gland that is compressed by an adjacent muscle suggests a more specialized role for the secretion produced by these glands that may not be related to envenomation but rather to prey transport and mucus control.
