Evolution and systematics

The hoopoe (Upupa epops) has long been considered to be related to the hornbills (Bucerotidae), and Tertiary fossil evidence supports this view. Its closest relatives are the wood-hoopoes and scimitarbills (Phoeniculidae), with which the hoopoe shares characters such as feather structure, pterylosis, tongue structure, bill morphology, skeletal features, and egg-white proteins. DNA-DNA hybridization studies suggest that the hoopoe diverged from the hornbills, and the wood-hoopoes and scimitarbills from the hoopoe.

The hoopoe differs from all other Coraciiform birds in having no expansor secondarium muscle and in the newly hatched chick being downy. The bird also has a tufted uropy-gial gland that produces a foul smell to repel nest intruders. Such characteristics have led some authors to place it in a separate order, the Upupiformes.

Although it is usually regarded as a polytypic single species, some authors regard the Madagascar hoopoe (subspecies mar-ginata) as a full species on the basis of its distinctive advertising call and large size. The nine subspecies are separated mainly on size, color, and wing pattern. Subfossils of a large, probably flightless hoopoe (Upupa antaois) are known from St. Helena island in the South Atlantic.

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