Hornbill

Hornbills (Bucerotidae) are a family of bird found in tropical and subtropical Africa, Asia and Melanesia. They are characterized by a long, down-curved bill which is frequently brightly-colored and sometimes has a casque on the upper mandible. Both the common English and the scientific name of the family refer to the shape of the bill, "buceros" being "cow horn" in Greek. In addition, they possess a two-lobed kidney. Hornbills are the only birds in which the first two neck vertebrae (the axis and atlas) are fused together; this probably provides a more stable platform for carrying the bill. The family is omnivorous, feeding on fruit and small animals. They are monogamous breeders nesting in natural cavities in trees and sometimes cliffs. A number of species of hornbill are threatened with extinction, mostly insular species with small ranges.

Distribution and Habitats
The Bucerotidae include about 55 living species, though a number of cryptic species may yet be split, as has been suggested for the Red-billed Hornbill. Their distribution includes Sub-Saharan Africa and the Indian Subcontinent to the Philippines and Solomon Islands, but no genus is found in both Africa and Asia. Most are arboreal birds, but the large ground-hornbills (Bucorvus), as their name implies, are terrestrial birds of open savanna. Of the 24 species found in Africa, 13 of these species are birds of the more open woodlands and savanna, and some species even occur in highly arid environments. The remaining species are found in dense forests. This contrasts with Asia, where a single species occurs in open savanna and the remainder are forest species. The Indian Subcontinent has 10 species of hornbills, of which 9 are found in India and adjoining countries, whereas the Sri Lanka Grey Hornbill is restricted to the island. The most common widespread species in the Indian Subcontinent, overall, is the Indian Grey Hornbill. According to the International Union for Conservation of Nature (IUCN), Indonesia has 13 hornbill species: 9 of them exist in Sumatra, and the rest exist in Sumba, Sulawesi, Papua and in Kalimantan has the same hornbill species as in Sumatra, except the Rangkong Papan (Buceros bicornis).

Description
Hornbills show considerable variation in size as a family, ranging in size from the Black Dwarf Hornbill (Tockus hartlaubi), at 102 grams (3.6 oz) and 30 cm (1 foot), to the Southern Ground Hornbill (Bucorvus leadbeateri), at up to 6.2 kg (13.6 lbs) and 1.2 m (4 feet). Males are always bigger than the females, though the extent to which this is true varies dependent upon species. The extent of sexual dimorphism also varies with body parts, for example the difference in body mass between males and females is between 1-17%, but the variation is 8-30% for bill length and 1-21% in wing length.

The most distinctive feature of the hornbills is the heavy bill, supported by powerful neck muscles as well as by the fused vertebrae. The large bill assists in fighting, preening, and constructing the nest, as well as catching prey. A feature unique to the hornbills is the casque, a hollow structure that runs along the upper mandible. In some species it is barely perceptible and appears to serve no function beyond reinforcing the bill. In other species it is quite large, is reinforced with bone, and has openings between the hollow centre allowing serve as a resonator for calls. In the Helmeted Hornbill the casque is not hollow but is filled with ivory and is used as a battering ram used in dramatic aerial jousts. Aerial casque-butting has also been reported in the Great Hornbill.

The plumage of hornbills is typically black, grey, white, or brown, although typically offset by bright colours on the bill, or patches of bare coloured skin on the face or wattles. Some species exhibit sexual dichromatism; in the Abyssinian Ground-hornbill, for example, pure blue skin on the face and throat denotes an adult female, and red and blue skin denotes an adult male. The calls of hornbills are loud, and vary distinctly between different species.

Hornbills possess binocular vision, although unlike most birds with this type of vision the bill intrudes on their visual field. This allows them to see their own bill tip and aids in precision handling of food objects with their bill. The eyes are also protected by large eyelashes which act as a sunshade.

Behaviour
Hornbills are diurnal, generally travelling in pairs or small family groups. Larger flocks sometimes form in the non-breeding season. The largest assemblages of hornbills form at some roosting sites, where as many as 2400 individual birds may be found.

Diet
Hornbills are omnivorous birds, eating fruit, insects and small animals. They cannot swallow food caught at the tip of the beak as their tongues are too short to manipulate it, so they toss it back to the throat with a jerk of the head. While both open country and forest species are omnivorous, species that specialise in feeding on fruit are generally found in forests while the more carnivorous species are found in open country. Forest living species of hornbills are considered to be important seed dispersers.

In some instances hornbills defend a fixed territory. Territoriality is related to diet; fruit sources are often patchily distributed and require long distance travel in order to find, thus species that specialise in fruit are less territorial.

Breeding
Hornbills generally form monogamous pairs, although some species engage in cooperative breeding. The female lays up to six white eggs in existing holes or crevices, either in trees or rocks. The cavities are usually natural, but some species may nest in the abandoned nests of woodpeckers and barbets. Nesting sites may be used in consecutive breeding seasons by the same pair. Before incubation, the females of all Bucerotinae—sometimes assisted by the male—begin to close the entrance to the nest cavity with a wall made of mud, droppings and fruit pulp. When the female is ready to lay her eggs, the entrance is just large enough for her to enter the nest, and after she has done so, the remaining opening is also all but sealed shut. There is only one narrow aperture, big enough for the male to transfer food to the mother and eventually the chicks. The function of this behaviour is apparently related to protecting the nesting site from rival hornbills. The sealing can be done in just a few hours, at most it takes a few days. Having sealed the nest it takes a further five days for the first egg to be laid. Clutch size varies from one or two eggs in the larger species to up to eight eggs for the smaller species. During the incubation period the female undergoes a complete and simultaneous moult. It has been suggested that the darkness of the cavity triggers a hormone involved in moulting. Non-breeding females and males go through a sequential moult. When the chicks and the female are too big to fit in the nest, the mother breaks out, then both parents feed the chicks. In some species the mother rebuilds the wall, whereas in others the chicks themselves rebuild the wall unaided. The ground-hornbills are conventional cavity-nesters instead.

Associations with other species
A number of hornbills have associations with other animal species. For example, some species of hornbills in Africa have a mutualistic relationship with Dwarf Mongooses, in which they forage together and warn each other of nearby birds of prey and other predators. Other relationships are commensal, for example following monkeys or other animals and eating the insects flushed up by them.

Taxonomy
There are two subfamilies: the Bucorvinae contain the 2 ground-hornbills in a single genus, whereas the Bucerotinae contain all other taxa. Traditionally they are within the order Coraciiformes (which includes also kingfishers, rollers, hoopoes and bee-eaters). In the Sibley-Ahlquist taxonomy, however, hornbills are separated from the Coraciiformes and make an order of their own, Bucerotiformes, with the subfamilies elevated to family level. Given that they are almost as distant from the rollers, kingfishers and allies as are the trogons, the arrangement chosen is more a matter of personal taste than any well-established taxonomic practice. All that can be said with reasonable certainty is that placing the hornbills outside the Coraciiformes and the trogons inside would be incorrect.

Species list in taxonomic order
This is a list of extant hornbill species, presented in taxonomic order.

Subfamily ''Bucerotinae


 * Genus Tropicranus (sometimes included in Tockus)
 * White-crested Hornbill Tropicranus albocristatus


 * Genus Tockus
 * Black Dwarf Hornbill Tockus hartlaubi
 * Red-billed Dwarf Hornbill Tockus camurus
 * Monteiro's Hornbill Tockus monteiri
 * Red-billed Hornbill Tockus erythrorhynchus
 * Eastern Yellow-billed Hornbill Tockus flavirostris
 * Southern Yellow-billed Hornbill Tockus leucomelas
 * Jackson's Hornbill Tockus jacksoni
 * Von der Decken's Hornbill Tockus deckeni
 * Crowned Hornbill Tockus alboterminatus
 * Bradfield's Hornbill Tockus bradfieldi
 * African Pied Hornbill Tockus fasciatus
 * Hemprich's Hornbill Tockus hemprichii
 * Pale-billed Hornbill, Tockus pallidirostris
 * African Grey Hornbill Tockus nasutus


 * Genus Ocyceros
 * Malabar Grey Hornbill Ocyceros griseus
 * Sri Lanka Grey Hornbill Ocyceros gingalensis
 * Indian Grey-Hornbill Ocyceros biostris


 * Genus Anthracoceros
 * Malabar Pied Hornbill Anthracoceros coronatus
 * Oriental Pied Hornbill Anthracoceros albirostris
 * Black Hornbill Anthracoceros malayanus
 * Palawan Hornbill Antracoceros marchei
 * Sulu Hornbill Anthracoceros montani


 * Genus Buceros
 * Rhinoceros Hornbill Buceros rhinoceros
 * Great Hornbill Buceros bicornis
 * Rufous Hornbill Buceros hydrocorax


 * Genus Rhinoplax (sometimes included in Buceros)
 * Helmeted Hornbill Rhinoplax vigil


 * Genus Anorrhinus
 * Austen's Brown Hornbill, Anorrhinus austeni
 * Tickell's Brown Hornbill, Anorrhinus tickelli
 * Bushy-crested Hornbill Anorrhinus galeritus


 * Genus Penelopides
 * Luzon Hornbill Penelopides manillae
 * Mindoro Hornbill Penelopides mindorensis
 * Visayan Hornbill Penelopides panini
 * Samar Hornbill Penelopides samarensis
 * Mindanao Hornbill Penelopides affinis
 * Sulawesi Hornbill Penelopides exarhatus


 * Genus Berenicornis (sometimes included in Aceros)
 * White-crowned Hornbill Berenicornis comatus


 * Genus Aceros
 * Rufous-necked Hornbill Aceros nipalensis
 * Wrinkled Hornbill Aceros corrugatus
 * Writhed Hornbill Aceros leucocephalus
 * Rufous-headed Hornbill Aceros waldeni
 * Knobbed Hornbill Aceros cassidix


 * Genus Rhyticeros (sometimes included in Aceros)
 * Wreathed Hornbill Rhyticeros undulatus
 * Narcondam Hornbill Rhyticeros narcondami''
 * Sumba Hornbill Rhyticeros everetti
 * Plain-pouched Hornbill Rhyticeros subruficollis
 * Papuan Hornbill Rhyticeros plicatus


 * Genus Bycanistes (sometimes included in Ceratogymna).
 * Trumpeter Hornbill Bycanistes bucinator
 * Piping Hornbill Bycanistes fistulator
 * Silvery-cheeked Hornbill Bycanistes brevis
 * Black-and-white-casqued Hornbill Bycanistes subcylindricus
 * Brown-cheeked Hornbill Bycanistes cylindricus
 * White-thighed Hornbill Bycanistes albotibialis


 * Genus Ceratogymna
 * Black-casqued Wattled Hornbill Ceratogymna atrata
 * Yellow-casqued Wattled Hornbill Ceratogymna elata

Subfamily ''Bucorvinae


 * Genus Bucorvus
 * Abyssinian Ground Hornbill Bucorvus abyssinicus
 * Southern Ground Hornbill Bucorvus leadbeateri

Cultural significance


Most species' casques are very light, containing a good deal of airspace. However, the Helmeted Hornbill has a solid casque made of a material called hornbill ivory, which is greatly valued as a carving material in China and Japan. It is often used as a medium for the art of netsuke.

Status
None of the African species of hornbills are seriously threatened, but many Asian hornbills are threatened due to hunting and habitat loss, as they tend to require primary forest. Among these threatened species, only the Plain-pouched Hornbill and Rufous-necked Hornbill are found in the Asian mainland; all others are insular in their distribution. In the Philippines alone, one species (Palawan Hornbill) is vulnerable and two species (Mindoro and Visayan Tarictic Hornbills) are endangered. The only critically endangered hornbills, the Rufous-headed Hornbill and Sulu Hornbill, are also restricted to the Philippines. The Ticao Tarictic Hornbill, a subspecies of the Visayan Tarictic Hornbill, is probably already extinct.