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The wreathed hornbill (''Rhyticeros undulatus'') is an
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by th ...
tropical bird of the
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 coloured and sometimes has a casque on the upper mandibl ...
family Bucerotidae, also called bar-pouched wreathed hornbill due to its distinctive blue-black band on its lower throat sac. It is named after its characteristic long, curved bill that develops ridges, or wreaths, on the casque of the upper mandible in adults. Males are black with a
rufous Rufous () is a color that may be described as reddish- brown or brownish-red, as of rust or oxidised iron. The first recorded use of ''rufous'' as a color name in English was in 1782. However, the color is also recorded earlier in 1527 as a ...
crown, a white upper breast and face, and a yellow featherless throat. Females are uniformly black with a blue throat and are slightly smaller than males. The wreathed hornbill ranges across the foothills and
evergreen forest An evergreen forest is a forest made up of evergreen trees. They occur across a wide range of climatic zones, and include trees such as conifers and holly in cold climates, eucalyptus, Live oak, acacias, magnolia, and banksia in more temperate zon ...
s of
Northeast India , native_name_lang = mni , settlement_type = , image_skyline = , image_alt = , image_caption = , motto = , image_map = Northeast india.png , ...
and
Bhutan Bhutan (; dz, འབྲུག་ཡུལ་, Druk Yul ), officially the Kingdom of Bhutan,), is a landlocked country in South Asia. It is situated in the Eastern Himalayas, between China in the north and India in the south. A mountainou ...
to
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
,
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...
and the
Greater Sunda Islands The Greater Sunda Islands ( Indonesian and Malay: ''Kepulauan Sunda Besar'') are four tropical islands situated within Indonesian Archipelago, in the Pacific Ocean. The islands, Borneo, Java, Sulawesi and Sumatra, are internationally recognised ...
. It is a
frugivore A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
and feeds mainly on large fruits, which it swallows whole leaving the seeds intact. This feeding behaviour plays an important ecological role for the long-distance
seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
in forest ecosystems. The wreathed hornbill is threatened by hunting,
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological process ...
and
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated ...
. It has been listed as Vulnerable on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biolog ...
since 2018, as the global population is estimated to decrease due to these threats.


Taxonomy

English ornithologist George Shaw first described the species in 1811 as ''Buceros undulatus'', based on a
zoological specimen A zoological specimen is an animal or part of an animal preserved for scientific use. Various uses are: to verify the identity of a (species), to allow study, increase public knowledge of zoology. Zoological specimens are extremely diverse. Exampl ...
collected in
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
. It was placed in the genus ''Rhytidoceros'' by
Daniel Giraud Elliot Daniel Giraud Elliot (March 7, 1835 – December 22, 1915) was an American zoologist and the founder of the American Ornithologist Union. Life He was born in New York City on March 7, 1835, to George and Rebecca Elliot. In 1858, he married Ann ...
in 1882; in '' Rhyticeros'' by
Allan Octavian Hume Allan Octavian Hume, CB ICS (4 June 1829 – 31 July 1912) was a British civil servant, political reformer, ornithologist and botanist who worked in British India. He was the founder of the Indian National Congress. A notable ornithologist, H ...
in 1878; and in '' Aceros'' by James L. Peters in 1931. The wreathed hornbill is a
monotypic taxon In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
and is widely recognized as a member of ''Rhyticeros'', the genus of hornbills with low wreathed casques. Analysis of the
mitochondrial genome Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
s of eight hornbill species revealed that the wreathed hornbill is closely related with ''Aceros'' and the
Visayan hornbill The Visayan hornbill (''Penelopides panini'') is a hornbill found in rainforests on the islands of Panay, Negros, Masbate, and Guimaras, and formerly Ticao, in the Philippines. It formerly included all other Philippine tarictic hornbills as su ...
(''Penelopides panini'').


Description

The wreathed hornbill is black with black legs and feet, but has a pale yellow bill, bare reddish skin around the eyes, a black band around the gular skin and a white tail. The female's gular skin is blue, whereas the male's is bright yellow; his neck is white and his nape dark reddish passing into black.
Sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
in the colouration of
plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
and inflatable gular skin is less pronounced in young wreathed hornbills, but becomes more apparent as both sexes mature. It is a medium-sized bird with a body length of ; adult males weigh from , and females weigh from .


Bill morphology

All members of the
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 coloured and sometimes has a casque on the upper mandibl ...
family have large, heavy bills with characteristic casques that distinguish them from other avian taxa. Hornbill casques contain foamy air-filled cavities surrounded by multiple layers of
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, hor ...
tiles that act as an external shell. The extra weight of the casque is supported by the fusion of the first two
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e. The bill has distinctive serrated casques near the base of the upper mandible, which are particularly unique because of its ridges. As the birds mature, forward growth causes the foremost ridge to break off. It is hypothesized that specialized casque characteristics evolved due to multiple selection pressures, including its use for vocalization enhancement, physical pounding, mating displays, and as visual cues for age and social status.


Distribution and habitat

The wreathed hornbill inhabits tropical
evergreen forest An evergreen forest is a forest made up of evergreen trees. They occur across a wide range of climatic zones, and include trees such as conifers and holly in cold climates, eucalyptus, Live oak, acacias, magnolia, and banksia in more temperate zon ...
s in the region from southern
Bhutan Bhutan (; dz, འབྲུག་ཡུལ་, Druk Yul ), officially the Kingdom of Bhutan,), is a landlocked country in South Asia. It is situated in the Eastern Himalayas, between China in the north and India in the south. A mountainou ...
,
Northeast India , native_name_lang = mni , settlement_type = , image_skyline = , image_alt = , image_caption = , motto = , image_map = Northeast india.png , ...
,
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
and through
mainland Southeast Asia Mainland Southeast Asia, also known as the Indochinese Peninsula or Indochina, is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to the west an ...
to
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
, where it is restricted to
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent is ...
,
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
,
Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and ea ...
and a few smaller islands. It has been recorded up to an elevation of . In Bhutan, two individuals were sighted in
Sarpang Sarpang, also transliterated as Sarbhang or Sarbang, is a thromde or town in Sarpang District in southern Bhutan Bhutan (; dz, འབྲུག་ཡུལ་, Druk Yul ), officially the Kingdom of Bhutan,), is a landlocked country in So ...
district in spring 1986. In Northeast India, it inhabits unlogged
primary forest An old-growth forestalso termed primary forest, virgin forest, late seral forest, primeval forest, or first-growth forestis a forest that has attained great age without significant disturbance, and thereby exhibits unique ecological featu ...
s and selectively logged forests in the foothills of the
Eastern Himalayas ] The Eastern Himalayas extend from eastern Nepal across Northeast India, Bhutan, the Tibet Autonomous Region to Yunnan in China and northern Myanmar. The climate of this region is influenced by the monsoon of South Asia from June to September. It ...
from
Nameri National Park Nameri National Park is a national park in the foothills of the eastern Himalayas in the Sonitpur District of Assam, India, about 35 km from Tezpur. Nameri is about 9 km from Chariduar, the nearest village. Nameri shares its northern boundary ...
in
Assam Assam (; ) is a state in northeastern India, south of the eastern Himalayas along the Brahmaputra and Barak River valleys. Assam covers an area of . The state is bordered by Bhutan and Arunachal Pradesh to the north; Nagaland and Manipur to ...
to Namdapha National Park in
Arunachal Pradesh Arunachal Pradesh (, ) is a state in Northeastern India. It was formed from the erstwhile North-East Frontier Agency (NEFA) region, and became a state on 20 February 1987. It borders the states of Assam and Nagaland to the south. It shares i ...
. During the breeding season, it lives in lowland areas, but migrates to higher elevations in the non-breeding season. In Myanmar, 62 wreathed hornbills were sighted in the
Mali River The Mali River (''Mali Hka'') is a river that originates in the hills of Kachin State, in the northern border of Myanmar. It flows approximately 320 km, when it meets with the N'Mai River and forms the Ayeyarwady River. History Constructio ...
valley at elevations of in winter 1999. A flock of about 50 individuals was sighted in Hponkanrazi Wildlife Sanctuary in September 2004.


Behaviour and ecology

The wreathed hornbill is a social bird and most active in the early mornings when foraging for fruits; 21 individuals studied in Pakke Tiger Reserve during the non-breeding season rested at midday and resumed activity in the early afternoons. It lives in larger flocks than other hornbill species and remains in groups during both breeding and non-breeding seasons. Flock size observed in Namdapha National Park ranged from a mean of 4.7 to 6.6 individuals with a density of in March and April to in December. In Arunachal Pradesh, the wreathed hornbill density is much lower in habitats where local people hunt, ranging from . It travels long distances to forage for fruit, sometimes flying between islands, but it does not migrate. A male wreathed hornbill was fitted with a
GPS tracking unit A GPS tracking unit, geotracking unit, satellite tracking unit, or simply tracker is a navigation device normally on a vehicle, asset, person or animal that uses satellite navigation to determine its movement and determine its WGS84 UTM ge ...
in Pakke Tiger Reserve during the breeding season; it moved a mean minimum daily distance of and dispersed seeds for up to from its nesting site.
Home range A home range is the area in which an animal lives and moves on a periodic basis. It is related to the concept of an animal's territory which is the area that is actively defended. The concept of a home range was introduced by W. H. Burt in 1943. He ...
s of the wreathed hornbill and other hornbill species overlap in several range countries.


Vocalizations

The main call of the wreathed hornbill is a loud double
yelp Yelp Inc. is an American company that develops the Yelp.com website and the Yelp mobile app, which publish crowd-sourced reviews about businesses. It also operates Yelp Guest Manager, a table reservation service. It is headquartered in San Fra ...
that has been described to sound like “coo-cuk” or “wuff-wurff”. This call is short and typically repeated in a sequence of three or more from a perching position or during flight. The second part of the call is higher pitched and louder than the first and can be heard from farther distances. When calling, the wreathed hornbill displays its brightly coloured throat sac by sharply thrusting its head backwards and upwards. This movement is thought to be a form of communication between individuals. It makes various other low-pitched, bisyllabic barking sounds.


Diet and feeding

The wreathed hornbill mainly feeds on large fruit with figs making up the majority of its diet. It consumes more drupaceous fruits than other hornbill species. It also occasionally eats small animals like
insect Insects (from Latin ') are pancrustacean Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, Thorax (ins ...
s,
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
s,
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Ga ...
s and
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchoce ...
s, particularly during the breeding season. It forages in the forest
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
across its range, sometimes descending to eat small prey or to retrieve fallen fruit. Tropical fruit abundance can greatly vary by location and season, therefore the wreathed hornbill flies long distances to track areas with high fruit availability. As it moves across its range, it disperses the consumed seeds. The
seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
behaviour of the wreathed hornbill plays a critical role in maintaining tropical forest diversity. The feeding technique used by the wreathed hornbill is called ballistic food transport. It holds the large food item at the tip of its bill and throws its head back quickly, swallowing it whole. This behaviour is likely to reduce resource
competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indiv ...
by exploiting fruit that is too large for smaller
frugivore A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
s. Another advantage to ballistic food transport is that the fruit seeds are protected from damage when they are swallowed whole. The entire seed passes through the
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans a ...
and is excreted intact, increasing its chances of
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
and forming a new plant.


Reproduction

The wreathed hornbill is a
monogamous Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time (serial monogamy) — as compared to the various forms of non-monogamy (e.g., pol ...
breeder that demonstrates biparental care. It nests in the cavities of large trees located in coastal forests, illegally logged areas, or among foothills up to an elevation of . It searches for existing oval-shaped tree cavities for nesting because it cannot excavate its own. The nest height ranges from above ground. Females use droppings or mud to cover the nest hole to protect the
clutch A clutch is a mechanical device that engages and disengages power transmission, especially from a drive shaft to a driven shaft. In the simplest application, clutches connect and disconnect two rotating shafts (drive shafts or line shafts). ...
and fledglings from predators but leave a gap large enough for males to deliver food. They unload the mud when they need to leave the nest. The average clutch size of the wreathed hornbill is 1–3 eggs, which the female incubates and guards alone for 40 days. She raises only one of the chicks and remains with it in the nest until it
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerab ...
s at the end of the nesting cycle, which lasts 111–137 days. Both parents remain with the juvenile for multiple months after it has left the nest.


Threats

Major threats to wreathed hornbill populations include hunting, use of
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lam ...
s,
illegal logging Illegal logging is the harvest, transportation, purchase or sale of timber in violation of laws. The harvesting procedure itself may be illegal, including using corrupt means to gain access to forests; extraction without permission, or from a p ...
and
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological process ...
. In Arunachal Pradesh, the local Mishmi and
Adi people Adi or ADI may refer to: Names and titles * Adi (mythology), an Asura in Hindu faith who appears in the Matsya Purāṇa * Adi (name), a given name in Hebrew and a nickname in other languages * Adi (title), a Fijian title used by females of ch ...
shoot 50–70 wreathed hornbills during the migratory season; they eat their meat, use their
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, ...
s as medicine to cure stomach problems, their feathers to decorate their homes and their casques to adorn their headgear. Local people also hunt wreathed hornbills for sport and use their fat for medical purposes. The wreathed hornbill has been listed as a
vulnerable species A vulnerable species is a species which has been categorized by the International Union for Conservation of Nature as being threatened with extinction unless the circumstances that are threatening its survival and reproduction improve. Vulner ...
on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biolog ...
since 2018, as these threats will likely lead to a decline of the global population in the near future.


Conservation

The wreathed hornbill is internationally protected under
CITES Appendix II CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
and is a Class II national protected species in China. In Pakke Tiger Reserve, a community-based ''Hornbill Nest Adoption Program'' was initiated in collaboration with the Arunachal Pradesh Forest Department in 2011; local
Nyishi people The Nyishi community is the largest ethnic group in Arunachal Pradesh in north-eastern India. In Nyishi, ''Nyi'' refers to "a human" and the word ''shi'' denotes "highland".The Nyishis are mentioned as the Daflas in the contemporary Ahom docume ...
from nine villages worked as nest protectors and one youth as field coordinator; more than 90 urban citizens supported the project financially. In the breeding seasons of 2012 and 2013, the nest protectors located eight wreathed hornbill nests, of which two had fledgings.


References


External links

* * * * * * {{Taxonbar, from=Q31829 Birds of Myanmar Birds of Northeast India Birds of Southeast Asia wreathed hornbill wreathed hornbill Taxa named by George Shaw