Hooded pitohui
The hooded pitohui is a species of bird in the genus Pitohui found in New Guinea. It was long thought to be a whistler but is now known to be in the Old World oriole family. Within the oriole family, this species is most closely related to the variable pitohuis in the genus Pitohui, and then the figbirds.
A medium-sized songbird with reddish-brown and black plumage, this species is one of the few known poisonous birds, containing a range of batrachotoxin compounds in its skin, feathers and other tissues. These toxins are thought to be derived from their diet and may function both to deter predators and to protect the bird from parasites. The close resemblance of this species to other unrelated birds also known as pitohuis which are also poisonous is an example of convergent evolution and Müllerian mimicry. Their appearance is also mimicked by unrelated non-poisonous species, a phenomenon known as Batesian mimicry. The toxic nature of this bird is well known to local hunters, who avoid it. It is one of the most poisonous species of pitohui, but the toxicity of individual birds can vary geographically.
The hooded pitohui is found in forests from sea level up to but is most common in hills and low mountains. A social bird, it lives in family groups and frequently joins and even leads mixed-species foraging flocks. Its diet is made up of fruits, seeds and invertebrates. This species is apparently a cooperative breeder, with family groups helping to protect the nest and feed the young. The hooded pitohui is common and is currently not at risk of extinction, with its numbers being stable.
Taxonomy and systematics
The hooded pitohui was described by the French ornithologist Charles Lucien Bonaparte in 1850. Bonaparte placed it in the genus Rectes which had been erected in the same year by Ludwig Reichenbach as an alternative name for the genus Pitohui, which had been described by René Lesson in 1831. No explanation was given for the preference of the newer name over the established older one, but it was common to prefer Latin names over non-Latin names, and to provide Latin names to those without. Richard Bowdler Sharpe encapsulated that attitude when he wrote in 1903 "Pitohui is doubtless an older name than Rectes, but can surely be laid aside as a barbarous word". Eventually however the principle of priority, which favours the first formal name given to a taxon, was applied, and Rectes was suppressed as the junior synonym of Pitohui.The hooded pitohui was placed in the genus Pitohui with five other species, and the genus was thought to reside within the Australasian whistler family. A 2008 examination of the genus, however, found it to be polyphyletic, with some purported members of the genus not actually falling within the whistlers. The hooded pitohui and the closely related variable pitohui were both found to be related to the Old World orioles. A 2010 study by the same team confirmed that the hooded pitohui and variable pitohui were orioles and indeed were sister species, and that together with the figbirds they formed a well defined basal clade within the family. As the variable pitohui was the type species for the genus Pitohui, the hooded pitohui was retained in that genus and the four remaining species were moved to other genera.
The hooded pitohui is monotypic, lacking any subspecies. Birds in the south east of New Guinea are sometimes separated into a proposed subspecies, P. d. monticola, but the differences are very slight and the supposed subspecies are generally regarded as inseparable.
Pitohui, the common name for the group and the genus name, is a Papuan term for rubbish bird, a reference to its inedibility. The specific name dichrous is from the Ancient Greek word, meaning. Alternate names for the hooded pitohui include the black-headed pitohui and lesser pitohui.
Physiology and description
The hooded pitohui is long and weighs. The adult has a black, head, chin, throat and upper breast and a black tail. The rest of the plumage is a reddish brown. The bill and legs are black, and the irises are either reddish brown, dark brown or black. Both sexes look alike. Juvenile birds look like adults, except that the of the tail and of the wing are tinged with brown.Toxicity
In 1990 scientists preparing the skins of the hooded pitohui for museum collections experienced numbness and burning when handling them. It was reported in 1992 that this species and some other pitohuis contained a neurotoxin called homobatrachotoxin, a derivative of batrachotoxin, in their tissues. This led to them being more broadly known outside New Guinea, considered by some to be the first documented poisonous birds, other than some reports of coturnism caused by consuming quail, and the first bird discovered with toxins in the skin. The same toxin had previously been found only in Central and South American poison dart frogs from the genera Dendrobates, Oophaga and Phyllobates. The batrachotoxin family of compounds are among the most toxic compounds by weight in nature, being 250 times more toxic than strychnine. Later research found that the hooded pitohui had other batrachotoxins in its skin, including batrachotoxinin-A cis-crotonate, batrachotoxinin-A and batrachotoxinin-A 3′-hydroxypentanoate.Bioassays of their tissue found that the skins and feathers were the most toxic, the heart and liver less toxic, and the skeletal muscles the least toxic parts of the birds. Of the feathers, the toxin is most prevalent in those covering the breast and belly. Microscopy has shown that the toxins are sequestered in the skin in organelles analogous to lamellar bodies and are secreted into the feathers. The presence of the toxins in muscle, heart and liver shows that hooded pitohuis have a form of insensitivity to batrachotoxins. A bird has been estimated to have up to 20 μg of toxins in its skin and up to 3 μg in its feathers. This can vary dramatically geographically and by individual, and some have been collected with no detectable toxins.
The poisonous pitohuis, including the hooded pitohui, are not thought to create the toxic compound themselves but instead sequester them from their diet. The extent of the toxicity varies both in the pitohuis across their range and also across the range of the unrelated blue-capped ifrit, another New Guinean bird found with toxic skin and feathers. Both of these facts suggest that the toxins are obtained from the diet. The presence of the toxins in the internal organs as well as the skins and feathers rules out the possibility that the toxins are applied topically from an unknown source by the birds.
One possible source has been identified in the forests of New Guinea: beetles of the genus Choresine, which contain the toxin and have been found in the stomachs of hooded pitohui. An alternative explanation, that the birds and beetles both get the toxin from a third source, is considered unlikely as the blue-capped ifrit is almost exclusively insectivorous.
Ecology
The function of the toxins to the hooded pitohui has been the source of debate and research since its discovery. The initial suggestion was that the toxins acted as a chemical deterrent to predators. Some researchers cautioned this suggestion was premature, and others noted that the levels of batrachotoxins were three orders of magnitude lower than in the poison dart frogs that do use it in this way.Another explanation for the purpose of the toxins is to mitigate the effects of parasites. In experimental conditions chewing lice were shown to avoid toxic feathers of hooded pitohui in favour of feathers with lower concentrations of toxin or no toxins at all. Additionally lice that did live in the toxic feathers did not live as long as control lice, suggesting that the toxins could lessen both the incidence of infestation and the severity. A comparative study of the tick loads of wild birds in New Guinea would seem to support the idea, as hooded pitohuis had considerably fewer ticks than almost all the 30 genera examined. The batrachotoxins do not seem to have an effect on internal parasites such as Haemoproteus or the malaria-causing Plasmodium.
A number of authors have noted that the two explanations, as a chemical defence against predators and as a chemical defence against ectoparasites, are not mutually exclusive, and evidence for both explanations exists. The fact that the highest concentrations of toxins are bound in the feathers of the breast and belly, in both pitohuis and ifrits, has caused scientists to suggest that the toxins rub off on eggs and nestlings providing protection against predators and nest parasites.
One argument in favour of the toxin acting as a defence against predators is the apparent Müllerian mimicry in some of the various unrelated pitohui species, which all have similar plumage. The species known as pitohuis were long thought congeneric, due to their similarities in plumage, but are now spread through three families, the oriole, whistlers and Australo-Papuan bellbirds. The similarity in appearance therefore presumably evolved as a shared aposematic signal to common predators of their distasteful nature. This signal is reinforced by the species' strong sour odor. There is also evidence that some other birds in New Guinea have evolved Batesian mimicry, where a non-toxic species adopts the appearance of a toxic species. An example of this is the non-toxic juvenile greater melampitta, which has plumage similar to the hooded pitohui.
There have also been experiments to test pitohui batrachotoxins on potential predators. They have been shown to irritate the buccal membranes of brown tree snakes and green tree pythons, both of which are avian predators in New Guinea. The unpalatability of the species is also known to local hunters, who otherwise hunt songbirds of the same size.
The existence of resistance to batrachotoxins and the use of those toxins as chemical defences by several bird families have led to competing theories as to its evolutionary history. Jønsson suggested that it was an ancestral adaptation in Corvoidea songbirds, and that further studies would reveal more toxic birds. Dumbacher argued instead that it was an example of convergent evolution.