Chemoreception in crocodilians: anatomy, natural history, and empirical results.

Anatomical and behavioral aspects of chemoreception in crocodilians are reviewed. Extant crocodilians possess valvular nostrils regulated by smooth muscles; paired nasal cavities, each with three conchae; and flaps closing the rear of the buccopharyngeal passage to channel inspired air through the nasopharyngeal duct. Olfaction is enhanced by gular pumping, whereby the floor of the pharynx is alternately lowered and raised to pulse air through the nasal cavity. Taste buds occur on the tongue, palate, and pharyngeal walls. Crocodilians in nature or in semi-natural enclosures locate distant carrion or concealed meat, implying chemoattraction to food. The American alligator exhibits underwater head-waving and mouth-opening in response to aqueous and chloroform extracts of meat. Alligators increase gular pumping in response to the scents of various meats, indicating that they detect food chemicals by olfaction. Paired gular and paracloacal glands on the integument of crocodilians are thought to produce pheromones. Juvenile crocodilians exhibit heightened gular pumping in response to airborne skin gland secretions. Observations of adults suggest that gular and paracloacal glands produce chemical signals used in mating and/or nesting activities, but behavioral responses to skin gland exudates are too poorly documented to ascribe pheromonal properties.