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.
Evolution and taxonomy
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 an 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.
Distribution and habitat
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.
Morphology and flight
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.
Diet and feeding
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 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.
* Order Trogoniformes
1. ^ Johansson, Ulf S. & Ericson, Per G. P. (2003): Molecular support for a sister group relationship between Pici and Galbulae (Piciformes sensu Wetmore 1960). Journal of Avian Biology 34: 185–197. PDF fulltext
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