|White-tailed Trogon, Trogon viridis|
The trogons and quetzals are birds in the order Trogoniformes which contains only one family, the Trogonidae. The family contains 39 species in eight genera. The fossil record of the trogons dates back 49 million years to the mid-Eocene. They might constitute a member of the basal radiation of the order Coraciiformes. The word "trogon" is Greek for "nibbling" and refers to the fact that these birds gnaw holes in trees to make their nests.
Trogons are residents of tropical forests worldwide, with the greatest diversity in the Neotropics. The genus Apaloderma contains the three African species, Harpactes and Apalharpactes are Asian, and the remaining four genera are found in Central and South America.
They feed on insects and fruit, and their broad bills and weak legs reflect their diet and arboreal habits. Although their flight is fast, they are reluctant to fly any distance. Trogons are generally not migratory, although some species undertake partial local movements.
Trogons have soft, often colourful, feathers with distinctive male and female plumage. They are the only type of animal with a heterodactyl toe arrangement.
The trogons are insectivorous, usually hunting from a perch. They nest in holes dug into trees or termite nests, laying 2-4 white or pastel-coloured eggs.
The position of the trogons within the class Aves has been a long-standing mystery. A variety of relations have been suggested, including the parrots, cuckoos, toucans, jacamars and puffbirds, rollers, owls and nightjars. More recent morphological evidence has suggested a relationship with the Coraciiformes. The unique arrangement of the toes on the foot (see morphology and flight) has led many to consider the trogons to have no close relatives, and to place them in their own order, possibly with the similarly atypical mousebirds as their closest relatives.
The family is thought to have an Old World origin (but see Moyle (2005) for an alternative theory), notwithstanding the current richness of the family, which is more diverse in the Neotropical New World. The oldest fossils attributed to the order were found in the Messel pit deposits from the mid-Eocene in Germany (49 mya). Other trogoniform fossils have been found in Oligocene rocks from Switzerland and Miocene France. The oldest New World fossil of a trogon is from the comparatively recent Pleistocene. DNA evidence seems to support and African origin for the trogons, with the African genus Apaloderma seemingly basal in the family, and the other two lineages, the Asian and American, breaking off between 20-36 million years ago. The trogons are split into three subfamilies, each reflecting one of these splits, Aplodermatinae is the African subfamily and contains a single genus, Apaloderma; Harpactinae is the Asian subfamily and contains two genera, Harpactes and Apalharpactes. Apalharpactes, consisting of two species in the Java and Sumatra, has only recently been accepted as a separate genus from Harpactes.
The remaining subfamily, the Neotropical Trogoninae, contains the remaining four genera, Trogon, Priotelus, Pharomachrus and Eupilotis. The two Caribbean species of Priotelus were formerly different ones (Temnotrogon on Hispaniola), and are extremely ancient. The two quetzal genera, Pharomachrus and Eupilotis are possibly derived from the final and most numerous genus of trogons in the Neotropics, Trogon. A 2008 study of the genetics of Trogon suggested the genus originated in Central America and radiated into South America after the formation of the Isthmus of Panama (as part of the Great American Interchange), thus making trogons relatively recent arrivals in South America.
The majority of trogons are birds of tropical and subtropical forests. They have a cosmopolitan distribution in the worlds wet tropics, being found in the Americas, Africa and Asia. A few species are distributed into the temperate zone, with one species, the Elegant Trogon, reaching the south of the United States specifically southern Arizona and the surrounding area. The Narina Trogon of Africa is slightly exceptional in that it utilises a wider range of habitats than any other trogon, ranging from dense forest to fairly open savannah, and from the Equator to southern South Africa. It is the most widespread and successful of all the trogons. The Eared Quetzal of Mexico is also able to use more xeric habitats, but preferentially inhabits forests. Most other species are more restricted in their habitat, with several species being restricted to undisturbed primary forest. Within forests they tend to be found in the mid story, occasionally in the canopy.
Some species, particularly the quetzals, are adapted to cooler montane forest. There are a number of insular species; these include a number of species found in the Greater Sundas, one species in the Philippines as well as two monophyletic genera endemic to Cuba and Hispaniola respectively. Outside of South Asia and the Caribbean, however, trogons are generally absent from islands, especially oceanic ones.
Trogons are generally sedentary, with no species known to undertake long migrations. A small number of species are known to make smaller migratory movements, particularly montane species which move to lower altitudes during different seasons. This has been demonstrated using radio tracking in the Resplendent Quetzal in Costa Rica and evidence has been accumulated for a number of other species. The Narina Trogon of Africa is thought to undertake some localised short distance migrations over parts of its range, for example birds of Zimbabwe's plateau savannah depart after the breeding season. A complete picture of these movements is however lacking. Trogons are difficult to study as their thick tarsi (feet bones) make ringing studies difficult.
The trogons as a family are fairly uniform in appearance, they have compact bodies with long tails (very long in the case of the quetzals), and short necks. Trogons range in size from the 23 cm, 40 gram Scarlet-rumped Trogon to the 40 cm, 210 gram Resplendent Quetzal (not including the male quetzal's 3-foot long tail streamers). Their legs and feet are weak and short, and trogons are essentially unable to walk beyond a very occasional shuffle along a branch. They are even incapable of turning around on a branch without using their wings. The ratio of leg muscle to body weight in trogons is only 3%, the lowest known ratio of any bird. The arrangement of toes on the feet of trogons is also unique amongst birds, although essentially resembling the zygodactyl’s two forward two backward arrangement of parrots and other near-passerines, the actual toes are arranged with usually inner hallux being the outer hind toe, an arrangement that is referred to as heterodactylous. The strong bill is short and the gape wide, particularly in the fruit eating quetzals, with a slight hook at the end. There is also a notch at the end of the bill and many species have slight serrations in the mandibles. The skin is exceptionally tender, making preparation of study skins difficult for museum curators. The skeletons of trogons are surprisingly slender, particularly the skulls which are very thin. The plumage of many species is iridescent, although most of the Asian species are not. The African trogons are generally green on the back with red bellies. The New World trogons similarly have green or deep blue upperparts but are more varied in their lowerparts. The Asian species tend towards red underparts and brown backs.
The wings are short but strong, with the wing muscle ratio being around 22% of the body weight. In spite of the strength of their flight, trogons do not fly often or for great distances, generally flying no more than a few hundred metres at a time. Only the montane species tend to make long distance flights. Shorter flights tend to be direct and swift, but longer flights are slightly undulating. Their flight can be surprisingly silent (for observers), although that of a few species is reportedly quite noisy.
The calls of trogons are generally loud and uncomplex, consisting of monosyllabic hoots and whistles delivered in varying patterns and sequences. The calls of the quetzals and the two Caribbean genera are the most complex. Amongst the Asian genera the Sumatran Trogon (Apalharpactes) has the most atypical call of any trogon, research has not yet established whether the closely related Javan Trogon has a similar call. The calls of the other Asian genus, Harpactes, are remarkably uniform. In addition to the territorial and breeding calls given by males and females during the breeding seasons, trogons have been recorded as having aggression calls given by competing males and alarm calls.
Trogons are generally inactive outside of infrequent feeding flights. Amongst birdwatchers and biologists it has been noted that "[a]part from their great beauty [they] are notorious ... for their lack of other immediately engaging qualities". Their lack of activity is possibly a defence against predation; trogons on all continents have been reported to shift about on branches to always keep their less brightly coloured backs turned towards observers, whilst their heads, which like owls can turn through 180 degrees, keep a watch on the watcher. Trogons have reportedly been preyed upon by hawks and predatory mammals; one report was of a Resplendent Quetzal taken whilst brooding young by a margay.
Trogons feed principally on insects, other arthropods, and fruit; to a lesser extent some small vertebrates such as lizards are taken. Amongst the insect prey taken one of the more important types are caterpillars; along with cuckoos trogons are one of the few birds groups to regularly prey upon them. The extent to which each food type is taken varies depending on geography and species. The three African trogons are exclusively insectivorous, whereas the Asian and American genera consume varying amounts of fruit. Diet is somewhat correlated with size, with larger species feeding more on fruit and smaller species focusing on insects.
Prey is almost always obtained on the wing. The most commonly employed foraging technique is a sally-glean flight, where a trogon flies from an observation perch to a target on another branch or in foliage. Once there the birds hovers or stalls and snatches the item before returning to its perch to consume the item. This type of foraging is commonly used by some types of bird to obtain insect prey; in trogons and quetzals it is also used to pluck fruit from trees. Insect prey may also be taken on the wing, with the trogon pursuing flying insects in a similar manner to drongos and Old World flycatchers. Frogs, lizards and large insects on the ground may also be pounced on from the air. More rarely some trogons may shuffle along a branch to obtain insects, insect eggs and very occasionally nestling birds. Violaceous Trogons will consume wasps and wasp larvae encountered while digging nests.
Trogons are territorial and monogamous. Males will respond quickly to playbacks of their calls and will repel other members of the same species and even other hole nesting species from around their nesting sites. Males attract females by singing, and, in the case of the Resplendent Quetzal, undertaking display flights. Some species have been observed in small flocks of 3–12 individuals prior to and sometimes during the breeding season, calling and chasing each other, but the function of these flocks is unclear.
Trogons are cavity nesters. Nests are dug into rotting wood or termite nests, with one species, the Violaceous Trogon, nesting in wasp nests. Nest cavities can either be deep upward slanting tubes that lead to fully enclosed chambers, or much shallower open niches (from which the bird is visible). Nests are dug with the beak, incidentally giving the family its name. Nest digging may be undertaken by the male alone or by both sexes. In the case of nests dug into tree trunks, the wood must be strong enough not to collapse but soft enough to dig out. Trogons have been observed landing on dead tree trunks and slapping the wood with their tails, presumably to test the firmness.
The nests of trogons are thought to usually be unlined. Between two to four eggs are laid in a nesting attempt. These are round and generally glossy white or lightly coloured (buff, grey, blue or green), although they get increasingly dirty during incubation. Both parents incubate the eggs (except in the case of the Bare-cheeked Trogon, where apparently the male takes no part), with the male taking one long incubation stint a day and the female incubating the rest of the time. Incubation seems to begin after the last egg is laid. The incubation period varies by species, usually lasting between 16–19 days. On hatching the chicks are altricial, blind and naked. The chicks acquire feathers rapidly in some of the montane species, in the case of the Mountain Trogon in a week, but more slowly in lowland species like the Black-headed Trogon, which may take twice as long. The nestling period varies by species and size, with smaller species generally taking 16 to 17 days to fledge, whereas larger species may take as long as 30 days, although 23–25 days is more typical.
Trogons and quetzals are considered to be "among the most beautiful of birds", yet they are also often reclusive and seldom seen. Little is known about much of their biology, and much of what is known about them comes from the research of neotropical species by the ornithologist Alexander Skutch. Trogons are nevertheless popular birds with birdwatchers, and there is a modest ecotourism industry in particular to view quetzals in Central America.
Phylogeny of Genera based on Moyle (2005)
TROGON, a word apparently first used as English 1 by G. Shaw (Mus. Leverianum, p. 177) in 1792, and now for many years accepted as the general name of certain birds forming the family Trogonidae of modern ornithology. The trogons are birds of moderate size: the smallest is hardly bigger than a thrush and the largest less bulky than a crow. In most of them the bill is very wide at the gape, which is invariably beset by recurved bristles. They seize most of their food, whether caterpillars or fruits, on the wing, though their alar power is not exceptionally great, their flight being described as short, rapid and spasmodic. Their feet are weak and of a unique structure, the second toe, which in most birds is the inner anterior one, being reverted, and thus the trogons stand alone, since in all other birds that have two toes before and two behind it is the outer toe that is turned backward. The plumage is very remarkable and characteristic. There is not a species which has not beauty beyond most birds, and the glory of the group culminates in the quezal. But in others golden green and steely blue, rich crimson 2 and tender pink, yellow varying from primrose to amber, vie with one another in vivid coloration, or contrasted, as happens in many species, with a warm tawny or a sombre slaty grey - to say nothing of the delicate freckling of black and white, as minute as the markings of a moth's wing - the whole set off by bands of white, producing an effect hardly equalled in any group. The plumage is further remarkable for the large size of its contour-feathers, which are extremely soft and so loosely seated as to come off in scores at a touch, and there is no down. The tail is generally a very characteristic feature, the rectrices, though in some cases pointed, being often curiously squared at the tip, and when this is the case they are usually 1 Trogonem (the oblique case) occurs in Pliny (H. N. x. 16) as the name of a bird of which he knew nothing, save that it was mentioned by Hylas, an augur, whose work is lost; but some would read Trygonem (turtle-dove). In 1752 MOhring (Av. Genera, p. 85) applied the name to the "Curucui" (pronounced "Suruqua," vide Bates, Nat. Amazons, i. 254) of Marcgrav (Hist. nat. Brasiliae, p. 211), who described and figured it in 1648 recognizably. In 1760 Brisson (Ornithologie, iv. 164) adopted Trogon as a generic term, and, Linnaeus having followed his example, it has since been universally accepted.
Anatole Bogdanoff determined the red pigment of the feathers of Pharomacrus auriceps to be a substance which he called "zooxanthine" (Comptes rendus, Nov. 2, 1857, xlv. 690).
barred ladder-like with white and black.' According to J. Gould, they are larger and more pointed in the young than in the old, and grow squarer and have the white bands narrower at each succeeding moult. He also asserts that in the species which have the wing coverts freckled, the freckling becomes finer with age. So far as has been observed, the nidification of these birds is in holes of trees, wherein are laid without any bedding two roundish eggs, generally white, but certainly in one species (quezal) tinted with bluish green.
The trogons form a very well-marked family, belonging to the coraciiform birds, and probably to be placed in that assemblage near the colies (see Mouse Bird) and swifts (q.v.). The remains of one, T. gallicus, have been recognized by A. Milne-Edwards (Ois. foss. de la France, ii. 395, pl. 177, figs. 18 -22) from the Miocene of the Allier. This fortunate discovery seems to account for the remarkable distribution of the trogons at the present day. While they chiefly abound, and have developed their climax of magnificence, in the tropical parts of the New World, they yet occur in the tropical parts of the Old. The species now inhabiting Africa, forming the group Hapaloderma, can hardly be separated generically from those of the Neotropical Trogon, and the difference between the Asiatic forms, if somewhat greater, is still comparatively slight. It is plain then that the Trogons are an exceptionally persistent type; indeed in the whole class few similar instances occur, and perhaps none that can be called parallel. The extreme development of the type in the New World just noticed also furnishes another hint. While in some of the American trogons (Pharomacrus, for instance) the plumage of the females is not very much less beautiful than that of the males, there are others in which the hen birds retain what may be fairly deemed a more ancient livery, while the cocks flaunt in brilliant attire. Now the plumage of both sexes in all but one 4 of the Asiatic trogons, Harpactes, resembles rather that of the young and of those females of the American species which are modestly clothed. The inference from this fact would seem to be that the general coloration of the Trogons prior to the establishment, by geographical estrangement, of the two types was a russet similar to that now worn by the adults of both sexes in the Indian region, and by a portion only of the females in the Neotropical. The Ethiopian type, as already said, very closely agrees with the American, and therefore would be likely to have been longer in connexion therewith. Again, while the adults of most of the American trogons (Pharomacrus and Euptilotis excepted) have the edges of the bill serrated, their young have them smooth or only with a single notch on either side near the tip, and this is observable in the Asiatic trogons at all ages. At the same time the most distinctive features of the whole group, which are easily taken in at a glance, but are difficult to express briefly in words, are equally possessed by both branches of the family, showing that they were in all likelihood - for the possibility that the peculiarities may have been evolved apart is not to be overlooked - reached before the geographical sundering of these branches (whereby they are now placed on opposite sides of the globe) was effected.
About sixty species of trogons are recognized, which J. Gould in the second edition of his Monograph of the family (1875) divides into seven genera. Pharomacrus, Euptilotis and Trogon inhabit the mainland of tropical America, no species passing to the northward of the Rio Grande nor southward of the forest district of Brazil, while none occur on the west coast of Peru or Chile. Prionotelus and Tmetotrogon, each with one species, are peculiar respectively to Cuba and Haiti. The African form Hapaloderma has two species, one found only on the west coast, the other of more general range. The Asiatic trogons, Harpactes (with eleven species according to the same authority), occur from Nepal to Malacca, in Ceylon, and in Sumatra, Java and Borneo, while one species is peculiar to some of the Philippine Islands. (A. N.)
Species: T. aurantiiventris - T. bairdii - T. citreolus - T. clathratus - T. collaris - T. comptus - T. curucui - T. elegans - T. massena - T. melanocephalus - T. melanurus - T. mexicanus - T. personatus - T. rufus - T. surrucura - T. violaceus - T. viridis
Trogon Brisson, 1760
Ornithologie 1 p.42 4 p.164