Few recent studies have examined the histological basis for tooth attachment in squamates. In the past few years, a surge of interest in this topic has led to the intriguing suggestion that the major tissues derived from the tooth germ (enamel, dentine, cementum and alveolar bone), are conservative and are present in all amniotes. In this study, we describe the histology and development of the tooth attachment complex in Varanus rudicollis, the rough-neck monitor. We provide the first published evidence for the role of cementum and alveolar bone in tooth attachment in varanoid lizards. In Varanus, cementum is deposited on the external surface of the tooth root as well as at the base of the tooth, where it plays a role in the attachment of the tooth to the jawbone. Alveolar bone is also involved in tooth ankylosis. Our results support the hypothesis that the major tooth germ tissues are found in all amniotes. We provide insights into the structure and development of plicidentine, defined as infolding of the dentine around the tooth base. This feature is unique to varanoids among extant tetrapods and is the third tissue implicated in tooth attachment in Varanus. Plicidentine develops asymmetrically along the labial-lingual axis of a tooth. Varanus is characterized by the presence of both primary and higher-order lamellae, which anastomose to form a honeycomb-like surface that then interacts with the more basal attachment tissues.
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