Tracing the developmental origin of a lizard skull: Chondrocranial architecture, heterochrony, and variation in lacertids.

The sand lizard, Lacerta agilis, is a classical model species in herpetology. Its adult skull anatomy and its embryonic development are well known. The description of its fully formed primordial skull by Ernst Gaupp, in 1900, was a key publication in vertebrate morphology and influenced many comparative embryologists. Based on recent methodological considerations, we restudied the early cranial development of this species starting as early as the formation of mesenchymal condensations up to the fully formed chondrocranium. We traced the formation of the complex chondrocranial architecture in detail, clarified specific homologies for the first time, and uncovered major differences to old textbook descriptions. Comparison with other lacertid lizards revealed a very similar genesis of the primordial skull. However, we detected shifts in the developmental timing of particular cartilaginous elements, mainly in the nasal region, which may correlate to specific ecological adaptation in the adults. Late timing of nasal elements might be an important innovation for the successful wide range distribution of the well-known sand lizard.