The "tooth systems" of Lissotriton vulgaris (Amphibia: Urodela) with special regard to delayed metamorphosis

Abstract

A study on the cranial morphology, especially on the tooth bearing (dental) systems of several preserved developmental stages (from early premetamorphic larvae, in which most skull elements were already present and ossified or ossified in part, to transformed adult) of the smooth newt Lissotriton vulgaris (Salamandridae) was undertaken. We used μCT (to visualize the ossified elements in general and their relationships to each other) and an overall Alizarinred staining (to at best visualize teeth, replacement teeth, tooth buds and, after removing the oral mucosa, the course of dental laminae). Specimens cleared and stained with Alzarinred and Alcianblue were shown to be less suitable for our questions. In one case we used histological sections to follow the course of dental laminae, and in a second case scanning electron microscopy to show the structure of teeth in detail. The general sequence, growth, and changes of the bony elements including the “dental systems”, especially around metamorphosis, known from several other salamandrids are largely confirmed. Concerning the “tooth systems”, metamorphic events include the late appearance of the maxillae, resorption of the coronoids and palatines including their tooth-patches, remodelling of the vomer, i.e. resorption of the vomerine larval tooth-patch, formation of the edentate vomerine plate, and outgrowth of the monstichously dentate vomerine bar (typical for salamandrids). We show evidence that the larval vomer is not completey resorbed and that, unlike what has been described for Salamandra salamandra, the development of the vomerine bar is probably preceded by a shift of the dental lamina towards the middle of the palate, leaving a broad area between larval vomer and dental lamina. We hypothesize that the connective tissue in this area ossifies later and extends posteriorly forming the vomerine bar. It is noteworthy that in nearly all larvae vomer and intact pterygopalatinum were very close together either on one side or on both sides leading in overwintered larvae to the fusion of the vomer and the palatinal portion of the pterygopalatinum, primarily on one side. The zone of fusion is always characterized by a buccal notch. We think that in L. vulgaris the formation of “vomeropterygopalatina” is supported by the close proximity of the two bones and that these bones may fuse due to an imbalance between differentiation- and growth rate (indirectly caused by low temperatures). Approximation and especially fusion of the two bones correspond with the extension of the vomerine dental lamina into the area of the palatine, which temporally provides the latter with teeth. Overwintered larvae show further deviations concerning growth and differentiation of the mouth roof, which can be also interpreted as signs of delayed metamorphosis. They retain, for example, a largely intact dentate palatine, but with some regression of its tooth-patch, while the larval vomer is enlarged anteriorly and posteriorly and its number of teeth has increased; and the largely intact pterygopalatinal bony bridge. Further, maxillae begin to ossify. All larvae obviously have reached a late premetamorphic larval stage before the delay has started.