Toco toucan

The toco toucan (Ramphastos toco), also known as the common toucan or giant toucan, is a species of bird in the toucan family Ramphastidae. It is the largest species of toucan and has a distinctive appearance, with a black body, a white throat, chest and uppertail-coverts, and red undertail-coverts. Its most conspicuous feature is its massive beak, which is yellow-orange with a black base and large spot on the tip. It is endemic to South America, where it has a wide distribution from the Guianas south to northern Argentina and Uruguay, and its range has recently been expanding southwards. Unlike other toucans, which inhabit continuous forests, toco toucans inhabit a variety of semi-open habitats at altitudes of up to 1,750 m (5,740 ft). They are especially common in the Brazilian cerrado, gallery forests, and the wetlands of the Pantanal.

Toco toucan
CITES Appendix II (CITES)[2]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Class: Aves
Order: Piciformes
Family: Ramphastidae
Genus: Ramphastos
Species:
R. toco
Binomial name
Ramphastos toco
Subspecies

See text

Toco toucans mainly feed on fleshy fruits, but also supplement their diets with insects, eggs, and nestlings of other birds. They feed on any available sugar-rich fruits, and show a high level of variation in their diet depending on the surrounding habitat. Breeding is seasonal, with the timing of the breeding season differing between regions. Nests are usually made in hollows in trees and contain two to four eggs; both parents incubate the eggs for 17–18 days before hatching. It is considered to be of Least Concern by BirdLife International.

Taxonomy and systematics

The toco toucan was described as Ramphastos Toco by the German zoologist Philipp Ludwig Statius Müller in 1776 on the basis of specimens from Cayenne, French Guiana.[3] In 1862, the German ornithologist Jean Cabanis described R. albogularis as a new species based on specimens from southern Brazil;[4] by 1870, it had been demoted to the status of a subspecies of R. toco.[5][6] The name of the genus, Ramphastos, derives from a misspelling of the Ancient Greek ῥαμφηστης, rhamphēstēs, meaning snouted, which is itself derived from the Greek ῥαμφη, rhampē, meaning bill; the name refers to the large bill of the toucans. The specific name toco comes from either Tucá or Tucán, the Guarani word for toucan, and may mean "bone-nose".[7] Toco toucan is the official common name designated by the International Ornithologists' Union (IOU).[8] It is called the tucanuçu in Portuguese and the tucán grande or tucán toco in Spanish.[9]

The toco toucan is one of eight species currently placed in the genus Ramphastos, and one of over forty species in the toucan family Ramphastidae.[8] In 1974, the German ornithologist Jürgen Haffer hypothesized that the Ramphastos toucans could be split into two clades (groups of organisms descending from a common ancestor): the "smooth-billed yelpers", comprising the chestnut-mandibled and yellow-throated toucans, and the "channel-keel-billed croakers", comprising the toco, red-breasted, keel-billed, Choco, and channel-billed toucans. He further postulated that the toco toucan was basal (closest to the root of the phylogenetic tree) within the group of channel-keel-billed croakers.[10] Later studies of mitochondrial DNA have largely confirmed the existence of these two clades, but have found the toco toucan to be basal within the family instead of being a part of the channel-keel-billed croakers.[11][12] The following cladogram shows phylogenetic relationships within Ramphastos, based on a 2009 study by José Patané and colleagues:[lower-alpha 1][12]

Ramphastos

Toco toucan (Ramphastos toco)

White-throated toucan (Ramphastos tucanus)

Yellow-throated toucan (Ramphastos ambiguus)

Keel-billed toucan (Ramphastos sulfuratus)

Red-breasted toucan (Ramphastos dicolorus)

Choco toucan (Ramphastos brevis)

Channel-billed toucan (Ramphastos vitellinus)

Subspecies

Two subspecies are recognized by the IOU:[8]

  • R. t. toco Statius Müller, 1776: found in the Guianas, north and northeastern Brazil, and south-eastern Peru.[8]
  • R. t. albogularis Cabanis, 1862:[lower-alpha 2] Found in Paraguay, southern and eastern Brazil, northern Bolivia, and northern Argentina.[8] Very similar to the nominate subspecies, but is said to be slightly smaller, with a shorter bill and whiter throat; however, measurements of individuals from both subspecies have found that differences in size are not consistent.[9]

Description
A partially leucistic toco toucan
Adult (left) and juvenile toucan

The toco toucan has conspicuously contrasting plumage with a mainly black body, a white throat, chest and uppertail-coverts, and red undertail-coverts. What appears to be a blue iris is actually thin blue skin around the eye. This blue skin is surrounded by another ring of bare, orange skin. The most noticeable feature, however, is its huge bill, which measures from 15.8 to 23 cm (6+1⁄4 to 9 in) in length, which is yellow-orange, tending to deeper reddish-orange on its lower sections and culmen, and with a black base and large spot on the tip.[13] It looks heavy, but as in other toucans it is relatively light because the inside largely is hollow. The tongue is nearly as long as the bill and very flat. This species is the largest toucan and the largest representative of the order Piciformes.[14] The total length of the species is 55–65 cm (21+1⁄2–25+1⁄2 in). Body weight in these birds can vary from 500 to 876 g (1 lb 1+5⁄8 oz to 1 lb 14+7⁄8 oz), with males averaging 723 g (1 lb 9+1⁄2 oz) against the smaller female, which averages 576 g (1 lb 4+3⁄8 oz). Among standard measurements, the wing chord is 22 to 26 cm (8+1⁄2 to 10 in), the tail is 14.1 to 17.9 cm (5+9⁄16 to 7+1⁄16 in) and the tarsus is 4.8 to 6.5 cm (1+7⁄8 to 2+9⁄16 in).[13] Other than the size difference, there are no external differences between the sexes. Juveniles are duller and shorter-billed than adults.

Bill
Further information: Toco toucan beak

The bill is serrated and is the largest relative to body size of all birds providing 30 to 50% of its body surface area, although another Neotropical species, the sword-billed hummingbird, has a longer bill relative to its body length.[15] It was called by Buffon a "grossly monstrous" appendage.[16] Diverse functions have been suggested. Charles Darwin suggested it was a sexual ornament: "toucans may owe the enormous size of their beaks to sexual selection, for the sake of displaying the diversified and vivid stripes of colour with which these organs are ornamented".[17] Further suggestions have included aid in peeling fruit, intimidating other birds when robbing their nests, social selection related to defense of territory, and as a visual warning.[15][18]

Research has shown that one function is as a surface area for heat exchange. The bill has the ability to modify blood flow and so regulate heat distribution in the body, allowing for the use of the bill as a thermal radiator.[15] In terms of surface area used for this function, the bill relative to the bird's size is amongst the largest of any animal and has a network of superficial blood vessels supporting the thin horny sheath on the bill made of keratin called the rhamphotheca.

Close-up of toco toucan beak

In its capacity to remove body heat, the bill is comparable to that of elephant ears. The ability to radiate heat depends upon air speed: if this is low, only 25% of the adult bird's resting heat production is radiated; if high, it radiates as much as four times this heat production. In comparison, the bill of a duck and the ears of an elephant can shed only about 9% of resting heat production. The bill normally is responsible for 30 to 60% of heat loss. The practice of toco toucans of placing their bills under their wings may serve to insulate the bill and reduce heat loss during sleep. It has been observed that "complexities of the vasculature and controlling mechanisms needed to adjust the blood flow to the bill may not be completely developed until adulthood."[15]

Vocalizations

Its voice consists of a deep, coarse croaking, often repeated every few seconds. It also has a rattling call and will bill-clack.

Distribution and habitat
Toco toucan flying in the Brazilian Pantanal

The toco toucan is endemic to South America, where it has a wide distribution from the Guianas south to northern Argentina and Uruguay. In the northern portion of its range, it has several disjunct populations, including in the Guianas, in northern Brazil near the Rio Branco, and along the mouth of the Amazon upstream to around Manaus in eastern Amazonas. It is further found from coastal Maranhão southwest to southwestern Brazil, Bolivia, and Pampas de Heath in far southeastern Peru, and south through Piauí and Bahia to northern Argentina and Uruguay.[9][19] It was previously thought to have gone extinct in Tucumán in northwestern Argentina by the late 1990s, but was rediscovered there in the early 2010s.[20] The species's range in the Amazon rainforest may be increasing due to deforestation.[9] Similarly, it has only been recorded from Uruguay recently; previously, the southward limit of its range was Lagoa dos Patos in Brazil. The recent expansion its range south of the 30th parallel may be caused by escapes of captive individuals or changes in ecological conditions.[21] Toco toucans are generally resident, but will sometimes move en masse in search of food.[9]

Unlike other toucans, toco toucans do not inhabit continuous, closed-canopy forests, instead preferring a variety of semi-open habitats such as gallery forests, savannas, forests adjoining water bodies, woodlands and secondary forest, chaco, plantations, orchards, and groves.[9] It is especially common in the Brazilian cerrado, a form of tropical savanna, and gallery forests. It is also common in the wetlands of the Pantanal.[22] It is known to inhabit altitudes of up to 1,750 m (5,740 ft).[1]

Behaviour and ecology

Toco toucans are typically seen when flying or feeding in treetops, hopping from branch to branch. Their flight is somewhat undulating because they switch between heavy flapping and gliding when flying. Toucans are less active during the day, occasionally resting in treetops. They are less gregarious than other toucans and usually feed alone or in small groups at fruiting trees. When foraging together, toucans fly from treetop to treetop in single file. Members of a group will often preen each other, although this behaviour usually ends after egg-laying occurs, as mates begin to exclusively preen each other.[9]

Diet
Toco toucan feeding on papaya

Toco toucans are generalist frugivores that primarily feed on fleshy fruits, but also commonly supplement their diets with insects, eggs, and nestlings of other birds. Foraging usually takes place in the canopy, but toucans will also visit the understory and ground to feed on fallen fruits.[9] Toucans are known to feed on a variety of fleshy fruit, most notably figs.[9] Other plants that fruit year-round and feature significantly in the species's diet include Cecropia pachystachya and Inga laurina.[22][23] Toucans will also opportunistically feed on any available sugar-rich fruits, and display a high level of variation in their diet depending on the surrounding habitat.[24] In deciduous forests with a year-round supply of figs, toucans do not show significant change from season to season in their diet.[25] In areas with significant variation in the availability of fleshy fruits, toucans are more common during the fruiting season, and subsequently move to other habitats when fruit availability declines.[24][25]

Plants that have been recorded as contributing majorly to the toco toucan's diet include Genipa americana, Ficus luschnatiana, and Virola sebifera in gallery forest,[22][24] Schefflera macrocarpa, Copaifera langsdorffii, Didymopanax morototoni, and Nectandra cissiflora in the cerrado,[24][26] and Guibourtia hymenaefolia and D. morototoni in semi-deciduous forests.[25] During the dry season when the availability of fruits declines, toco toucans will also feed on flowers of species such as Handroanthus chrysotrichus and Erythrina fusca.[27]

The insects most commonly consumed by toco toucans are caterpillars and termites.[9] Toco toucans also feed on small vertebrates such as lizards,[28] nestlings, and small birds.[9] In the wild, toucans prey on the nests of icterid blackbirds, tyrant flycatchers,[9] puffbirds,[29] and parrots,[24] and especially target the nests of yellow-rumped caciques.[9] The largest known birds whose nests are predated upon by the toco toucan are the hyacinth macaw and buff-necked ibis; the nestlings of the latter weigh 200–300 g (7.1–10.6 oz), compared to the toucan's body mass of around 540 g (19 oz), and are killed by breaking their necks with sideways movement of the toucan's beak.[30] In captivity, toucans will feed on a number of smaller birds that are placed with them, such as Toxostoma thrashers, house sparrows, Inca doves, cactus wrens, and Gambel's quails.[9]

Breeding

Toco toucans show several changes in behavior during their nesting season, becoming more secretive and solitary in habit.[31] Preening of toucans by birds outside of the mated pair stops soon after egg-laying; mates will continue to frequently preen each other or tap their beaks together. Males will sometimes display the red feathers on the underside of the tail while fanning out their undertail coverts; their large, conspicuous bill may also be used in breeding displays.[9]

Nesting is seasonal, but timing of the breeding season differs between regions. Breeding has been observed from September to January in Amazonas and Maranhão and from September to February in São Paulo, Rio de Janeiro, Paraná, and Paraguay. In eastern Argentina and Rio Grande do Sul, it occurs from October to February, and in Goiás, Minas Gerais, and Bahia, it lasts from November to February. It has been documented from December to June in Bolivia and western Argentina, and from May to June in Piauí.[9] Nesting occurs in cavities, which are usually made in trees such as coral cockspur and slash pines, but which can also be in stream banks or termite mounds. Nests are sometimes made in cavities excavated by other species of birds: recorded instances include nests in cavities made by campo flickers and cream-backed woodpeckers.[9][31] Successful breeding has also been observed in urban areas, although high levels of human-caused disturbances, such as construction activities, around the nest can lead to abandonment by the parents.[31] Captive birds have been observed cleaning their nests early on, but the nests are eventually filled with droppings and fruit seeds.[9]

Toucans usually breed annually in the wild, but have been reported breeding multiple times a year in captivity. Females lay two to four eggs, which are incubated by both parents for 17–18 days, after which they hatch. Hatchlings are initially fed mostly insects, with the proportion of fruit in their diet increasing as they age. Chicks fledge 43–52 days after hatching.[9]

  • Hatchlings
    Hatchlings
  • Young at nest; note the layer of fruit seeds at the bottom of the nest.
    Young at nest; note the layer of fruit seeds at the bottom of the nest.
  • Adult toucan feeding young
    Adult toucan feeding young

Interactions with other species

No specific predators of the toco toucan are known, but toucans in general are known to be hunted by monkeys and large birds of prey.[9]

Although toco toucans feed on other birds at low rates, they may be ecologically significant nest predators for species which nest in areas with few other terrestrial predators, such as cliffs.[30]

Several species of chewing lice, such as Austrophilopterus cancellosus and Myrsidea witti, are known to parasitize the toco toucan.[9] The species is also parasitized by the protozoans Plasmodium nucleophilum, P. huffi, P. pinottii,[32] Toxoplasma gondii,[33] and Giardia duodenalis,[34] Trichomonas parabasalids,[35] and trematodes.[36]

Relationship with humans

Conservation

Because it prefers open habitats, the toco toucan is likely to benefit from the widespread deforestation in tropical South America. It has a large range and except in the outer regions of its range, it typically is fairly common. It is therefore considered to be of Least Concern by BirdLife International.[1] Toco toucans are hunted for their meat and for the pet trade. The impact of this on the population is unknown.[9]

Aviculture

Like the keel-billed toucan, the toco toucan is sometimes kept in captivity, but has a high fruit diet and is sensitive to hemochromatosis (an iron storage disease).[37] Also, pet toco toucans must not be permitted to eat mouse (or rat) meat, due to a risk of bacterial infection.[38] There is an ongoing population management plan that should help to reverse the decreasing captive population of the toco toucan for Association of Zoos and Aquariums member institutions. This is the second management plan that is occurring since 2001.[39]

Notes
  1. The study treated the citron-throated toucan (R. citreolaemus) as a subspecies of the channel-billed toucan (R. vitellinus).[12]
  2. albogularis means "white-throated" and is derived from the Latin albus, meaning "white", and the Modern Latin gularis, meaning "-throated".[7]

References
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Wikimedia Commons has media related to Ramphastos toco.
  • Toco toucan videos on the Internet Bird Collection
  • Stamps (for Argentina, Bolivia, Brazil, French Guiana, Guyana, Paraguay)
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