Reproductive biology

Hornbill reproduction tends to coincide with rainfall and increased food supply. In seasonal African savannas, Tockus species begin courtship and reproduction with the rains, when invertebrates and fruits are plentiful. The opposite occurs on Sulawesi where lack of rainfall stimulates reproduction in the Sulawesi red-knobbed hornbill, so the burst in fruit supply occurs immediately after fledging. In aseasonal Bornean rainforests, reproduction appears to be supra annual, tied to highly cyclical peaks in food supply. Breeding in these populations may be controlled by the rate at which pairs regain condition between reproductive cycles. In fig-rich forests of North Sulawesi, hornbills breed every year, usually returning to the same nest tree.

The hornbill's unique nesting behavior is the feature that has most fascinated students of nature. All hornbills are hole-nesters, preferring natural cavities in trees or rock crevices. Unlike any other group of birds, the female hornbill seals the entrance to her nest cavity, leaving only a narrow slit through which she, and later her chicks, receive food from her mate. In most species, the male ferries mud to the female who then works for several days to seal the cavity entrance. Where mud is a rare commodity, the female uses her own feces as building material.

Nest sealing is believed to have evolved as a form of predator defense, for protection against other intruding hornbills, and to enforce male fidelity. Nest sealing has been described as an example of male chauvinism in which the male cloisters his female, forcing her to depend on him for survival. In reality, the female incarcerates herself and later frees herself, forcing the male to provide for her and their offspring. Because the male is busy provisioning his family, he is incapable of maintaining two nests, and the female can be sure of his complete attention.

The onset of breeding begins with courtship. When in flight, courting pairs act as though they are attached by an invisible rubber band, reacting swiftly to each other's movements. They perch in cozy proximity, engage in mutual preening, and exchange food gifts as a demonstration of their ardor. Other clues of the onset of breeding include the intensification in color of the exposed fleshy areas around the face and throat, reflecting hormonal changes. Nest inspection increases in frequency until copulation occurs and the female enters the nest cavity.

The number of eggs, their size, and the length of incubation are all correlated with body size. Clutch size ranges from two to three eggs in large hornbills and up to eight for smaller hornbills. Incubation runs from 23-49 days in small and large species, respectively. Eggs hatch in intervals and the emerging chicks are naked and translucent pink with closed eyes. Feather growth begins within a few days and as chicks develop, the skin blackens and begging calls change from feeble cheeps to loud, insistent calls.

The timing of female emergence varies tremendously; some females accompany their chicks from the nests and others leave well before chicks fledge. Research on Monteiro's hornbill suggests that females emerge to ensure survival when their body condition reaches its lowest point.

Male hornbills can be impressive providers. Although many Tockus species carry items to the nest one-by-one, most hornbills collect multiple food items, stuffed into a bulging gullet before delivering a load to the nest. A Sulawesi red-knobbed hornbill once delivered 162 fruits in one trip, a load equivalent to nearly 20% of his body weight.

Nesting success is high for those species studied. In southern Africa, chicks fledged from 90-92% of the nests of four Tockus species and in Thailand, 80% of great hornbill nests monitored fledged young. Sulawesi red-knobbed hornbills averaged 80% nesting success over three years, but this figure plummeted to 62% during the 1997 El Nino/ENSO fires. Smaller hornbills fledge up to four chicks, but large hornbills rarely fledge more than one chick per year.

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