Comparative studies on the eye morphology of lungless salamanders, family Plethodontidae, and the effect of miniaturization

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Abstract
In seven species of lungless salamanders, family Plethodontidae, ranging from medium to very small in body size and from small to very large in cell size, the morphology of the eye and the retina were investigated. Haller's rule was only partially corroborated. While the smallest species had the relatively largest eyes, the largest two species, having the largest cells, showed the third and fourth largest eyes of the series. An effect of cell size was also found with regard to eye morphology. Small species with small cells as well as large species with large cells had relatively small retinae and relatively large lenses. In contrast, small eyes with relatively large cells had absolutely and relatively large retinae and relatively small lenses. The retinae of all investigated plethodontids showed a morphology typical for land vertebrates with two fiber and three nuclear layers. Rods, cones and double cones could clearly be distinguished. A fovea or area centralis was not found. Retina ganglion cell and photoreceptor counts show that the number of these elements was lower than in salamandrids. However, determination of the resolution power of miniaturized eyes based on morphological and behavioral data shows that this does not seem to constitute a functional disadvantage. The morphological and functional properties and constraints of eyes of miniaturized salamanders are discussed.