Early development of the actinopterygian head. I. External development and staging of the paddlefish Polyodon spathula

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Abstract
A large sample of embryological material of the North American paddlefish Polyodon spathula (Acipenseriformes: Polyodontidae) confirms that early development in Polyodon is very similar to that reported for the sister group of Polyodontidae, the sturgeons (Acipenseridae). Polyodon illustrates many basic aspects of acipenseriform (and actinopterygian) head development that have not been adequately described. In this paper, we provide an overview of external features of cranial development using scanning electron microscopy. The observations are correlated with staging schemes previously proposed for paddlefishes and other acipenseriforms. Events that occur after the start of neurulation (stage 19) to the start of feeding (stage 46) are emphasized. New information on the structure and folding of the mandibular and hyoid segments permits an understanding of the early development of the pharyngeal region. In addition, we offer new descriptions of the hatching gland, the olfactory organ, the sensory barbel, and the initiation of paddle outgrowth. We also comment on the mode of origin of the hypophysis, and refute the notion that it is derived from the lips of the anterior neuropore as suggested in older literature. This information sets the stage for future comparative and experimental studies of the embryology of basal actinopterygians.