The Elaenia Elaenias

Yellow-bellied elaenia Elaenia flavogaster, photographed by Félix Uribe.

We are all aware that there are some truly stunning birds out there: majestic eagles and vultures, vibrant parrots and hummingbirds, eye-catching cranes and pelicans. But those of us who spend a lot of time contemplating the nature of bird diversity, whether as bird-watchers or ornithologists, will soon admit that the greater proportion of this diversity is composed of what are affectionately or not-so-affectionately referred to as Little Brown Jobs. In particular, the tyrant flycatchers or Tyrannidae of the Americas are one family of birds that is notorious for including some of the littlest, the brownest, and the jobbiest.

Elaenia is a genus of about twenty or so species of tyrannid found in Central and South America (Sibley & Monroe, 1990, listed eighteen, but phylogenetic studies suggest that some of these should be divided into more than one species—Rheindt et al. 2009). The name 'elaenia' does double service for these guys as both genus and vernacular name, though the members of some related genera are also labelled in the vernacular as 'elaenias'. As a result, Ridgely & Tudor (2009), without a trace of apparent irony, referred to the species of this genus as 'Elaenia elaenias'.

Mottled-backed elaenia Elaenia gigas, showing its distinctive divided crest, photographed by Nick Athanas.

The various species of Elaenia elaenias are notoriously difficult to distinguish, and none are particularly eye-catching. They are mostly greenish, though the slaty elaenia Elaenia strepera is dark grey, and the brownish elaenia E. pelzelni is (surprisingly) brown. Underparts may be white, or they may be yellow. One species in particular is labelled as the yellow-bellied elaenia E. flavogaster, but in this case it is not any more strikingly yellow than a number of other species, leading one to suspect whether its vernacular name is any sort of moral judgement. A number of species have some degree of white streak on the crown, and some have a small crest of feathers (the mottle-backed elaenia E. gigas has a well-developed, bifurcated crest). Elaenias are best distinguished by their calls, but that of course requires the bird in question to be calling.

Great elaenia Elaenia dayi, photographed by Thiago Orsi.

Though members of the tyrant flycatcher family in both affinities and appearance, elaenias consume a fair proportion of fruit as well as insects. In at least some species, fruit make up by far the greater part of the diet (Marini & Cavalcanti 1998). Different species often have different preferred habitats, and the relationship between habitat and phylogeny was examined by Rheindt et al. (2008). Two savannah-dwelling species, the plain-crested elaenia Elaenia cristata and the rufous-crowned elaenia E. ruficeps, appear to be the sister clade to the remaining species that mostly inhabit riparian habitats or montane and temperate forests (Elaenia species are largely absent from lowland tropical forest). The forest species fall into two clades nested among the riparian species. The great elaenia E. dayi, which happens to be the largest Elaenia species by a noticeable margin, inhabits the stunted montane forests of the south Venezuelan tepuis (if you've seen the film Up, this is the habitat in which that film is mostly set). Migratory habits, on the other hand, are less correlated with phylogeny than habitat preferences. A number of Elaenia species migrate between temperate breeding grounds and tropical wintering grounds, but migratory species may be closely related to sedentary species that inhabit the tropics all year round. Indeed, some species are mostly sedentary but have somewhat migratory populations in more temperate parts of their range.

REFERENCES

Marini, M. Â., & R. B. Cavalcanti. 1998. Frugivory by Elaenia flycatchers. Hornero 15: 47-50.

Rheindt, F. E., L. Christidis & J. A. Norman. 2008. Habitat shifts in the evolutionary history of a Neotropical flycatcher lineage from forest and open landscapes. BMC Evolutionary Biology 8: 1193.

Rheindt, F. E., L. Christidis & J. A. Norman. 2009. Genetic introgression, incomplete lineage sorting and faulty taxonomy create multiple cases of polyphyly in a montane clade of tyrant-flycatchers (Elaenia, Tyrannidae). Zoologica Scripta 38: 143-153.

Ridgely, R. S., & G. Tudor. 2009. Field Guide to the Songbirds of South America: The Passerines. University of Texas Press.

Sibley, C. G., & B. L. Monroe Jr. 1990. Distribution and Taxonomy of Birds of the World. Yale University Press.

Barn Owls and Such

European barn owl Tyto alba, photographed by Nuno Barreto.

I have no idea where the 'wise old owl' stereotype originally came from. Perhaps it simply originated from their appearance: their broad faces, sedate manner, and slightly supercilious half-lidded gaze (those last two, of course, only applying under the circumstances most people would actually see an owl: as a half-asleep night-dweller rudely awakened during the day). Whatever the cause for their associations, owls are one group of birds that have commonly featured in popular culture. The Eurasian barn owl Tyto alba is one owl species that has long held a particular association with humans. Owls mostly do not build their own nests, but make use of suitable hollows and crannies that they find ready-made. The preferred food of barn owls is small mammals such as rats and mice (though they will not turn up their beaks at alternative fare such as reptiles or large insects when their favourite is not available). Put these two facts together, and human constructions (i.e. barns) can be paradise for a barn owl: ready-made secluded nesting spots in the roof-space, and a steady supply of rodents attracted to stored foodstuffs and/or refuse.

Greater sooty owl Tyto tenebricosa arfaki, photographed by Nik Borrow.

The European barn owl is just one species in the genus Tyto, within which König & Weick (2010) recognised 25 species from around the world. While some Tyto species, like T. alba, are found over a wide range, others are found in restricted localities (many on particular oceanic islands). Some are very poorly known: the Taliabu masked owl Tyto nigrobrunnea from Indonesia was described from a single specimen in 1939, with only a handful of sight records since to attest to its continued existence. The Itombwe owl Tyto prigoginei of central Africa was similarly unknown between its initial description in 1952 and the mistnet capture of a live female in 1996. Prior to its transfer to Tyto by König & Weick (2010), this last species was included in the genus Phodilus, the bay owls, which is the living sister group to Tyto. Together, these two genera form the family Tytonidae, separate from all other owls in the family Strigidae. Tytonid owls differ from strigid owls in a features such as having the inner and central toes of the foot similar in length (versus the inner toe being distinctly shorter than the central one in strigids), with the central toe being serrated on the underside. The species of Tyto have a distinctly heart-shaped facial disc (that of Phodilus species is almost reminiscent of Hello Kitty). Many Tyto species, such as T. alba, prefer open habitats, but some, such as the sooty owls Tyto multipunctata and T. tenebricosa of eastern Australia and New Guinea, inhabit rainforests. The smallest Tyto species are T. prigoginei at about 24 cm total length and the Galapagos barn owl T. punctatissima at about 26 cm, and the largest is the Tasmanian grass owl T. castanops, reaching up to 55 cm in length and about 1.25 kg in weight.

Tasmanian grass owl Tyto castanops, photographed by Murray Lord.

Other extinct species would have also probably broken the 1 kg mark. Tyto species have a long fossil record, going back to the Middle Miocene European species T. sanctialbani (Kurochkin & Dyke 2011). Tytonids of now-extinct genera had been abundant in Europe before that time, but Mlíkovský (1998) suggested that they had become temporarily extinct there in the Early Miocene, owing to a gap in the fossil record. Tyto sanctialbani was similar in size to the modern T. alba (Mlíkovský 1998), but a number of giant fossil barn owls are known from islands around the world. The largest include Tyto pollens and T. riveroi in the West Indies (the Bahamas and Cuba, respectively), and T. robusta and T. gigantea from Gargano. Gargano is a peninsula of southern Italy that was a separate island during the Late Miocene to Early Pliocene, at which time it was home to a distinctive endemic fauna including such animals as the absolutely insane small ruminant Hoplitomeryx, which possessed both a crown of five spike-shaped horns and long dagger-like canines. It has been suggested that this over-exuberant armature had evolved as a defence against Gargano's main predators, an assemblage of raptors including the aforementioned Tyto species. The larger of the two, T. gigantea, was about twice the size of a living European barn owl, and perhaps larger than any living owl (Ballmann 1976), though it was more gracile in build than the largest living Bubo species. Ballmann provides measurements for leg bones of T. gigantea and not wing bones, but if we assume similar proportions to a modern barn owl then we'd be looking at a wingspan for T. gigantea of about two metres. That, I submit, is enough to scare seven colours of crap out of any number of small mammals.

REFERENCES

Ballmann, P. 1976. Fossile Vögel aus dem Neogen der Halbinsel Gargano (Italien), zweiter Teil. Scripta Geol. 38: 1-59, 7 pls.

König, C., & F. Weick. 2010. Owls of the World, 2nd ed. Christopher Helm: London.

Kurochkin, E. N., & G. J. Dyke. 2011. The first fossil owls (Aves: Strigiformes) from the Paleogene of Asia and a review of the fossil record of Strigiformes. Paleontological Journal 45 (4): 445-458.

Mlíkovský, J. 1998. A new barn owl (Aves: Strigidae) from the early Miocene of Germany, with comments on the fossil history of the Tytoninae. J. Ornithol. 139: 247-261.

Riroriro

The grey warbler or riroriro Gerygone igata, photographed by Peter Bray.

The eighteen recognised species of the genus Gerygone are an assemblage of small, drab-coloured birds found mostly in the Australo-Papuan region, with G. sulphurea found in the Malay Peninsula, Indonesia and the Philippines, and G. flavolateralis found in New Caledonia and Vanuatu. These are another group of birds that have tended to draw the short straw in the vernacular name stakes: G. igata, one of the most abundant of New Zealand's native birds, is usually identified by the uninspiring 'grey warbler'. Personally, I prefer the more onomatopoeiac Maori name for these lively little birds: 'riroriro' (it has been suggested in some circles that it could possibly be referred to as the 'grey gerygone'; this proposition shall be treated with the scorn that it deserves). The riroriro and its congeners feed on small insects that they mostly glean from leaves or small branches, generally in the middle to upper canopies (Ford 1985). A certain amount of their prey is caught in the air, while the riroriro and the brown warbler G. mouki of eastern Australia also forage in lower vegetation than other species. The riroriro is also the only Gerygone species known to forage on the ground (Keast & Recher 1997).

Gerygone species build hanging purse-shaped nests; this is a brown warbler Gerygone mouki photographed by Peter.

Somewhat unusually for a decently-speciose passerine genus, the circumscription of Gerygone has been fairly stable in recent years, and the genus has mostly been supported as monophyletic. The only exception of recent times has been the New Guinean G. cinerea, recently reclassified by Nyári & Joseph (2012) as a species of Acanthiza. In the early 1900s, some authors divided Gerygone species between smaller genera (for instance, the Australian ornithologist Gregory Mathews, who never met a genus he couldn't break down). One species so separated was the Chatham Island warbler G. albofrontata, which is something of an island giant compared to other Gerygone species, weighing about 12 g while other species are about 6 to 7 g (Keast & Recher 1997). Unfortunately, the Chatham Island warbler was not included in the phylogenetic analysis of Gerygone by Nyári & Joseph (2012), but it was not identified as significantly separate from other Gerygone species in the morphological analysis by Ford (1985).

The Chatham Island warbler Gerygone albofrontata, from here.

REFERENCES

Ford, J. 1985. Phylogeny of the acanthizid warbler genus Gerygone based on numerical analyses of morphological characters. Emu 86: 12-22.

Keast, A., & H. F. Recher. 1997. The adaptive zone of the genus Gerygone (Acanthizidae) as shown by morphology and feeding habits. Emu 97: 1-17.

Nyári, Á. S., & L. Joseph. 2012. Evolution in Australasian mangrove forests: multilocus phylogenetic analysis of the Gerygone warblers (Aves: Acanthizidae). PLoS One 7(2): e31840.

The Parulidae: Not-warblers, Not-ovenbirds and Not-redstarts

Black-crested warbler Myiothlypis nigrocristata, photographed by Mikko Pyhälä.

There is no denying the current status of English as the de facto lingua franca of the world*. And yet, I feel that a complaint must be laid at the feet of the Brits: they're a bit unimaginative when it comes to animal names. Many a British explorer, upon being presented with some hitherto unfamiliar product of the natural world, proceeded to label it with the name of whatever inhabitant of his native Europe he felt bore some vague resemblance. And hence, even today, there are significant groups of animals such as the Parulidae that are almost without a vernacular name to genuinely call their own.

*The potential irony of this sentence is not lost on me.

The Parulidae are a family of birds found throughout the Americas, though in the northern United States and Canada they are represented by migratory species that retreat further south in the cold months. Many of the migratory species have males with brightly coloured breeding plumage and are consequently idolised by North American bird watchers; non-migratory species, on the other hand, tend to have similarly subdued males and females (Update: see comments below). Members of the Parulidae are generally referred to as 'warblers' or 'wood warblers', despite not being at all closely related to the European warblers. Instead, parulids are members of the 'nine-primaried oscines', the passerine clade that also includes such birds as finches, buntings, sparrows, cardinals and tanagers. Within the nine-primaried oscines, parulids are closely related to the Icteridae, another American clade containing its fair share of representatives doomed to masquerade under stolen names (Barker et al. 2013).

Ovenbird Seiurus aurocapilla on its nest, photographed by M. C. Donald.

Though the nine-primaried oscines as a whole are fairly stable in their membership, recent years have seen a fair bit of shuffling back and forth between the clade's constituent families. As a result of this shuffling, the name 'Parulidae' has come to be associated with a core clade that excludes a number of more uncertainly placed taxa previously included in the family, such as the Central American wrenthrush Zeledonia coronata. A recent comprehensive study of the molecular phylogeny of the core parulids by Lovette et al. (2010) also resulted in a proposed shifting of many generic boundaries within the clade. According to Lovette et al., the basalmost member of the Parulidae is the ovenbird Seiurus aurocapilla, a migratory but monomorphic, relatively large parulid of North and Central America. Just to confuse matters, the name 'ovenbird' has also been used for an unrelated group of South American birds of the genus Furnarius. To be charitable, this is not a case of inappropriate name-saking, but refers to the construction by both groups of domed nests resembling an old earthernware oven. The next member of the parulids to split off was the worm-eating warbler Helmitheros vermivorus, a relatively long-billed species that migrates between the eastern United States and Central America.

Swainson's warbler Limnothlypis swainsonii, photographed by Greg Lavaty.

Next comes a clade of eight species classified in the genera Parkesia, Vermivora, Mniotilta, Limnothlypis and Protonotaria. The black-and-white warbler Mniotilta varia is noted for its distinctive feeding behaviour: it crawls along branches like a nuthatch or creeper, gleaning insects from the bark. The prothonotary warbler Protonotaria citrea is a bright yellow species that Kurt Vonnegut devoted some time to in Jailbird: "The song of a prothonotary warbler is notoriously monotonous, as I am the first to admit...Still—they are capable of expressing heartbreak—within strict limits, of course" (I personally feel the same about skylarks). The waterthrushes of the genus Parkesia are larger, terrestrially-feeding species.

Chestnut-sided warbler Setophaga pensylvanica, photographed by Cephas.
Other North American species are placed by Lovette et al. in the larger genera Geothlypis, Oreothlypis and Setophaga. The last genus contains the species previously included in Dendroica, but the recognition that the American redstart Setophaga ruticilla (again, no relation to the European redstart) is nested within Dendroica leads to the use of the older name. These genera include some of the most colorful parulids. The remaining genera Myiothlypis, Basileuterus, Cardellina and Myioborus form a mostly Neotropical clade. Myioborus species are also known as redstarts, presumably by comparison with the European birds as not one of them actually possesses a red tail. The name 'whitestart' has supposedly been proposed instead, but the only time that name appears to see use is when it is referred to by someone explaining why they are not using it...

REFERENCES

Barker, K. F., K. J. Burns, J. Klicka, S. M. Lanyon & I. J. Lovette. 2013. Going to extremes: contrasting rates of diversification in a recent radiation of New World passerine birds. Systematic Biology 62 (2): 298-320.

Lovette, I. J., J. L. Pérez-Emán, J. P. Sullivan, R. C. Banks, I. Fiorentino, S. Córdoba-Córdoba, M. Echeverry-Galvis, F. K. Barker, K. J. Burns, J. Klicka, S. M. Lanyon & E. Bermingham. 2010. A comprehensive multilocus phylogeny for the wood-warblers and a revised classification of the Parulidae (Aves). Molecular Phylogenetics and Evolution 57: 753-770.

Whistling for Whistlers

Male and juvenile golden whistler Pachycephala pectoralis, photographed by S. Lloyd. Female golden whistlers resemble the juveniles.

The name Pachycephalidae (or some orthographic variant thereof such as Pachycephalinae) has long been used to refer to a group of small insectivorous birds from the Australo-Papuan region (summarised by Boles 1979 as 'large-headed stout-bodied birds with poorly developed rictal bristles'). In the past, taxa included in the Pachycephalidae included the New Zealand genera Mohoua and Turnagra, the crested shriketit Falcunculus frontatus and the crested bellbird Oreoica gutturalis. However, as has often been the case with the more generalised groups of passerine birds, recent authors have tended to whittle the family's contents down as molecular studies have scattered the constituent taxa about the family tree. In its most recent iterations, Pachycephalidae generally corresponds to the taxa included in clade 'CC5CC6b' of Jetz et al. (2012), comprising primarily the genera Pachycephala and Colluricincla, plus a small number of satellite taxa.

Bare-throated whistler Pachycephala nudigula, photographed by Lars Peterssen.

The genus Pachycephala is currently used for the whistlers, a group of about forty species of mostly long-tailed, stout-billed birds, often (but not always) with sexually dimorphic coloration, with a contrasting pectoral band (most often black in the males) dividing the throat from the chest. The exact number of species in the genus is somewhat uncertain due to disagreements about the status of several constituent populations: the golden whistler P. pectoralis complex, for instance, includes over 65 named taxa variously recognised as species or subspecies (Jønsson et al. 2010). Species of Pachycephala are found from southeast Asia through to Fiji and Tonga. Some of you may recall whistlers featuring in Dougal Dixon's Life after Man as giving rise to an island radiation including nut-cracking, wood-pecking and predatory species (my only question being, who was the Hart that the plesiomorphic species in the radiation is supposed to be named after?)

Grey shrike-thrush Colluricincla harmonica, photographed by Sammy Sam.

Colluricincla, the shrike-thrushes, includes at least four species found in Australia and New Guinea (excluding a couple of species better included in Pachycephala). They are larger, slenderer and more narrow-headed than the whistlers, with less dimorphic coloration. The grey shrike-thrush C. harmonica of Australia has a high reputation as a singer. Some recent authors have also suggested inclusion of two further New Guinean species, the rusty pitohui Pitohui ferrugineus and white-bellied pitohui P. incertus, in Colluricincla. The genus Pitohui has been used to include a group of about half a dozen New Guinean species in the Pachycephalidae, which are relatively large and brash as pachycephalids go. They are variously reddish, black, or some combination of the two (the white-bellied pitohui has, as its name indicates, a yellowish-white underside). They have become most notorious in recent years for the discovery that, with the apparent exception of P. incertus, they are in fact toxic, with their skin and feathers containing batrachotoxins comparable to those found in the arrow-poison frogs of South America. Like the arrow-poison frogs, the birds probably do not generate the toxin themselves, but accumulate it from a diet of toxin-carrying melyrid beetles. However, molecular studies have indicated that the genus Pitohui as previously recognised is polyphyletic. The hooded pitohui P. dichrous and the variable pitohui P. kirhocephalus, the latter of which is the type species of the genus, are more closely related to the Oriolidae than the Pachycephalidae, while three species remain close to Pachycephala and Colluricincla. As well as the two species mentioned previously (which may be included in Colluricincla or maintained as a distinct genus for which the name Pseudorectes is available), the black pitohui 'Pitohui' nigrescens should be included in the Pachycephalidae as its own genus Melanorectes. As well as being closer to black than the reddish Pseudorectes species, Melanorectes nigrescens apparently possesses a 'peculiar musky smell' (Rothschild & Hartert 1913).

Rusty pitohui Pseudorectes ferrugineus, photographed by Dubi Shapiro.

The remaining two species of the Pachycephalidae are placed by Jønsson et al. (2010) in the genus Coracornis: the maroon-backed whistler C. raveni of Sulawesi and the Sangihe shrike-thrush C. sanghirensis of Sangihe, north of Sulawesi (as the vernacular name suggests, this species was previously included in Colluricincla). Neither of these species appears to be particularly well-known: in particular, C. sanghirensis was first described in 1881 but not observed again for over 100 years until 1985 (Rozendaal & Lambert 1999). So long was the gap between observations that several authors had, in this time, assumed that the original specimen must have been mislabelled and come from somewhere in New Guinea rather than Sangihe. This misinterpretation was encouraged by the belief that the species was directly related to the little shrike-thrush Colluricincla megarhyncha, found on the wrong side of the Wallace line to easily colonise Sangihe. As it is, C. sanghirensis is restricted to only a small area of forest on Sangihe at an altitude above 600 m, and was regarded by Rozendaal & Lambert (1999) as critically endangered. Its identification with Coracornis rather than Colluricincla was done by Jønsson et al. (2010) on the basis of molecular analysis; when comparing C. sanghirensis with Colluricincla, Rozendaal & Lambert (1999) do not appear to have also considered Coracornis raveni.

Sangihe shrike-thrush Coracornis sanghirensis, photographed by Marc Thibault.

REFERENCES

Boles, W, E. 1979. The relationships of the Australo-Papuan flycatchers. Emu 79: 107-110.

Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. O. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444-448.

Jønsson, K. A., R. C. K. Bowie, R. G. Moyle, L. Christidis, J. A. Norman, B. W. Benz & J. Fjeldså. 2010. Historical biogeography of an Indo-Pacific passerine bird family (Pachycephalidae): different colonization patterns in the Indonesian and Melanesian archipelagos. Journal of Biogeography 37: 245-257.

Rothschild, W., & E. Hartert. 1913. List of the collections of birds made by Albert S. Meek in the lower ranges of the Snow Mountains, on the Eilanden River, and on Mount Goliath during the years 1910 and 1911. Novitates Zoologicae 20 (3): 473-527.

Rozendaal, F. G., & F. R. Lambert. 1999. The taxonomic and conservation status of Pinarolestes sanghirensis Oustalet 1881. Forktail 15: 1-13.

Birds of the Sun

Handsome sunbird Aethopyga bella, photographed by Tonee Despojo. This species was only recently separated at species level from the lovely sunbird Aethopyga shelleyi; one of the distinguishing features of the two is the purple ear-patch in A. bella.

The sunbirds are definitely forerunners in the tally of the world's most brilliantly coloured birds. This family of long-billed nectar-feeders, found in tropical regions of the Old World, is often compared to the New World hummingbirds. Like hummingbirds, the males of most sunbirds shimmer with brilliant iridescent colours (the exceptions are the spiderhunters of the genus Arachnothera); the females are much more restrained, generally shades of olive-green or brown. However, though hummingbirds are committed aerialists (as befits their relationship with the swifts and nightjars), sunbirds are, as Passeriformes, more likely to feed while perched on a stem alongside their chosen flower. Also, while sunbirds are primarily nectar feeders, they also feed to a fair extent on small insects (this is also true of hummingbirds).

Male (above) and female (below) of fork-tailed sunbird Aethopyga christinae. Male photographed by Frankie Chu, female by Neil Fifer.

Sunbirds are also a rather less diverse group than hummingbirds, both in number of species and in external appearance. Because of their structural similarity, authors have differed in the number of genera recognised in the family, but one group that has generally been differentiated is the Aethopyga sunbirds of southern Asia. Aethopyga species tend to be smaller than other sunbirds, with relatively short but strongly downcurved bills. The male has the central tail-feathers elongate (Ali & Ripley 1999). Aethopyga species are also distinguished from other sunbirds by the structure of the tongue. As with other sunbirds, the tongue is elongate, with the sides curved inwards to form a double tube. Differing from others, the end of the tongue is divided into two inwardly open tubes but with a basal bifurcated plate connecting the tubes:

Tongues of sunbirds of different genera showing differences in morphology, from Cheke & Mann (2001).

Cheke & Mann (2001) listed seventeen species of Aethopyga, with an eighteenth species being added by Mann (2002). Several of these species are also currently recognised as polytypic, with multiple subspecies. Though Mann's (2002) 'new' species was simply derived from the elevation of previously-recognised subspecies, one entirely new species of Aethopyga, A. linaraborae from Mindanao in the Phillippines, was only described as recently as 1997. No large scale analysis of the interrelationships between Aethopyga species appears to have been published as yet, but centres of diversity are the Philippines and the Himalayas.

Elegant sunbird Aethopyga duyvenbodei, photographed by Marc Thibault. Having been informed by their vernacular names that Aethopyga sunbirds are, in turn, handsome, lovely and elegant, it is all the sadder to say that this last species from Sangihe, near Sulawesi, is regarded as endangered.

REFERENCES

Ali, S., & S. D. Ripley. 1999. Handbook of the Birds of India and Pakistan, together with those of Bangladesh, Nepal, Sikkim, Bhutan and Sri Lanka, 2nd ed., vol. 10. Flowerpeckers to Buntings. Oxford University Press.

Cheke, R. A., & C. F. Mann. 2001. Sunbirds: A Guide to the Sunbirds, Flowerpeckers, Spiderhunters and Sugarbirds of the World. A & C Black Publishers.

Mann, C. F. 2002. Systematic notes on Asian birds. 28. Taxonomic comments on some south and south-east Asian members of the family Nectariniidae. Zool. Verh. Leiden 340: 179-189.

A King among Parrots

Moluccan King parrot Alisterus amboinensis, photographed by Helsinki***.

For today's post, I'm looking at the King parrots of the genus Alisterus. There are three recognised species in this genus: the Australian King parrot Alisterus scapularis of eastern Australia, the green-winged or Papuan King parrot A. chloropterus of central and eastern New Guinea, and the Amboina or Moluccan King parrot A. amboinensis of eastern Indonesia and western Papua. However, each species is divided into subspecies, and some subspecies are quite distinct from each other. For instance, Alisterus scapularis shows distinct sexual dimorphism: the male has a bright red head and breast while the female has a green head and breast. In A. amboinensis, both sexes have red heads. In A. chloropterus, the nominate subspecies has a green-headed female like that of A. scapularis, but the northwesternmost subspecies A. chloropterus moszkowskii has a red-headed female like that of A. amboinensis (Forshaw & Knight 2010). In relation to other parrots, Alisterus belongs to the tribe Psittaculini that extends into eastern and southern Asia and the Mascarenes, among which it forms a clade with the other Australian genera Aprosmictus and Polytelis (Mayr 2010) (and hybrids have even been recorded between A. scapularis and species of these two genera—Rutgers & Norris 1972).

Female (left) and male Australian King parrot Alisterus scapularis, photographed by Peter Firminger.

You might be wondering why, among an entire order of particularly regal birds, it is this particular genus that is honoured with the title of 'King' (I know I certainly did). As it turns out, the reason appears to be that Alisterus is not, properly speaking, the 'king of parrots', but 'King's parrot', named after Philip Gidley King, governor of New South Wales from 1800 to 1806 ('Stentoreus' 2004; I might as well also point out for the benefit of those not familiar with Australian history that the original 'New South Wales' was considerably larger than the current state by that name, taking in the entire eastern seaboard of Australia).

Papuan King parrot Alisterus chloropterus, photographed by Mehd Halaouate.

King parrots are generalist feeders on fruit and seeds, which has not always endeared them to horticulturalists. They nest in deep holes in hollow trees: while the entrance to an Alisterus scapularis nest may be more than nine metres high, the actual nest may be nearly at ground level (Rutgers & Norris 1972). They lay 3-6 eggs between October and December.

REFERENCES

Forshaw, J. M., & F. Knight. 2010. Parrots of the World. Princeton University Press.

Mayr, G. 2010. Parrot interrelationships—morphology and the new molecular phylogenies. Emu 110: 348-357.

Rutgers, A., & K. A. Norris. 1972. Encyclopaedia of Aviculture vol. 2. Blandford Press: London.

More than Four and Twenty Blackbirds

The Tristan thrush or Starchy, Turdus eremita, an endemic bird of the remote Tristan da Cunha group of islands in the South Atlantic, scavenging on a dead penguin (starchies have decidedly more catholic tastes than other thrushes). Photo by Lex.

Turdus, the thrushes, is a large cosmopolitan genus of birds found throughout the world except for Australia. The extent of the genus' circumscription has varied between authorities, though most recent authors exclude the ground-thrushes of the genus Zoothera. Conversely, phylogenetic studies have indicated that the previously monotypic genera Cichlherminia lherminieri of the Caribbean and Nesocichla eremita from Tristan da Cunha should be subsumed within Turdus (Voelker et al., 2007). At present, it seems unlikely that Turdus will be further subdivided; as the basalmost species in the genus is likely to be the mistle thrush Turdus viscivorus, which happens to also be the type of the genus, any subdivision would require that Turdus be reduced to a single species and all other species placed in new genera.


The blackbird Turdus merula, a species found throughout northern Eurasia (and introduced to New Zealand). Only the males are black; the females are dark mottled brown and have grey rather than yellow beaks. Photo by Bence Mate.

Of the 60+ species remaining in Turdus, many are widespread and divided into a number of subspecies that may or may not be promoted to separate species by future researchers. As an extreme example, a study on variation between geographically separated populations of the island thrush Turdus poliocephalus, whose distribution extends from Sumatra and the Philippines east to Norfolk Island* and Vanuatu, suggested that there may be grounds for dividing them between nearly forty diagnostic taxonomic units (Peterson, 2007).

*At least, it did. The Tasman Sea populations of T. poliocephalus have, unfortunately, since shuffled off this mortal coil.


The St Lucia forest thrush, Turdus lherminieri sanctaeluciae. Like T. eremita, this is a distinctive species that was previously placed in its own genus. Photo by Jean-Michel Fenerole.

The base coloration of most species of Turdus can be described as 'mottled brown', though notable exceptions (at least as males) include the grey and red American robin T. migratorius and the blackbird T. merula. Most members of the genus are more highly regarded for their voices rather than their looks, an attribute honoured in both the vernacular and scientific names of the song thrush Turdus philomelos* ("lover of song"). As with other speciose songbird clades, variation in song has turned out to be significant in separating closely related species. Both the Príncipe thrush T. xanthorhynchus (Melo et al., 2010) and the black-throated thrush T. atrogularis (Sangster et al., 2009) differ in their songs (among other things) from species with which they were previously considered conspecific.

*Older references may one of the names Turdus musicus or Turdus ericetorum for this species. Both these names have since been suppressed by the ICZN. The history of Turdus musicus is particularly turgid, as authorities had disagreed over whether the name should be applied to the song thrush or to the redwing (now Turdus iliacus) (Mayr & Vaurie, 1957). Both T. musicus and T. iliacus appeared in Linnaeus' 1758 Systema Naturae. Unfortunately (whether because he was unclear on the distinction between the species, or by a simple composition error), Linnaeus confused the two species' descriptions: under T. musicus, he gave a description of the redwing but provided sources referring to the song thrush, while the entry for T. iliacus attached a description of the song thrush to references referring to the redwing! (The significance of Linnaeus' sources to his descriptions has previously been discussed in the sperm whale nomenclature post.) Mayr & Vaurie's (1957) application buried the name Turdus musicus and designated a neotype to fix Turdus iliacus firmly to the redwing.

REFERENCES

Mayr, E., & C. Vaurie. 1957. Proposed use of the plenary powers to suppress the specific name "musicus" Linnaeus, 1758, as published in the combination "Turdus musicus" and to approve a neotype for "Turdus iliacus" Linnaeus, 1758, the Eurasian redwing (class Aves). Bulletin of Zoological Nomenclature 13 (6): 177-181.

Melo, M., R. C. K. Bowie, G. Voelker, M. Dallimer, N. J. Collar & P. J. Jones. 2010. Multiple lines of evidence support the recognition of a very rare bird species: the Príncipe thrush. Journal of Zoology 282 (2): 120-129.

Peterson, A. T. 2007. Geographic variation in size and coloration in the Turdus poliocephalus complex: a first review of species limits. Scientific Papers, Natural History Museum, The University of Kansas 40: 1-17.

Sangster, G., A. B. van den Berg, A. J. van Loon & C. S. Roselaar. 2009. Dutch avifaunal list: taxonomic changes in 2004–2008. Ardea 97 (3): 373–381.

Voelker, G., S. Rohwer, R. C. K. Bowie & D. C. Outlaw. 2007. Molecular systematics of a speciose, cosmopolitan songbird genus: defining the limits of, and relationships among, the Turdus thrushes. Molecular Phylogenetics and Evolution 42: 422-434.

Name the Bug # 7 - Apalopteron familiare hahasima

Apalopteron familiare hahasima - photo by Outsuka Hiroyuki.

Apalopteron familiare, the Bonin honeyeater, is a small bird of the Japanese Ogasawara (or Bonin) island group in the north-west Pacific. Phylogenetically speaking, it's been shuffled around a bit over the years - originally described as a species of bulbul, it seems to have been regarded by many authors as some sort of timaliid (babbler) until Deignan identified it in 1958 as belonging to the Meliphagidae (honeyeaters), mainly on the basis of its branched, brushed tongue. There it stayed until molecular analysis re-identified it as a member of the white-eyes (Driskell & Christidis, 2004). There are two funny things about this - (1) Apalopteron really does look like a white-eye (albeit a large one), so identifying it as one is hardly surprising; (2) further phylogenetic analysis has also placed the white-eyes (previously regarded as a separate family) within the Timaliidae (Gelang et al., 2009), making Apalopteron's older position correct after all.

The Ogasawara islands are arranged in three groups - from north to south, these are the Mukojima, Chichijima and Hahajima groups. Shima (which often becomes voiced to -jima when used in a compound) is Japanese for "island", while chichi is "father" and haha is "mother" (other islands in the group include ani and otōto [elder and younger brother] and ane and imōto [elder and younger sister]). Apalopteron familiare was originally found on all three groups, but currently survives only in the Hahajima group. As on many Pacific islands, the Ogasawara fauna has been pretty heavily hit since the arrival of humans due to habitat disturbal and introduced predators - among the endemic species to have become extinct in the group are a heron (Nycticorax crassirostris), pigeon (Columba versicolor), thrush (Turdus terrestris), finch (Chaunoproctus ferrugineus) and bat (Pipistrellus sturdeei) (Iwahashi, 1992). The type subspecies of Bonin honeyeater (Apalopteron familiare familiare) became extinct on Mukojima by the 1940s (Morioka & Sakane, 1978)*. Kawakami et al. (2008) found that dispersal between populations of Apalopteron on separate islands was very low, which would further increase its vulnerability.

*It is possible that a bit of revision may be required here. Suzuki & Morioka (2005) indicated that the type locality for Apalopteron familiare was probably Chichijima rather than Mukojima. Chichijima is considerably closer to Hahajima than it is to Mukojima, raising the question of whether the correct subspecies has been identified as the type.

REFERENCES

Driskell, A. C., & L. Christidis. 2004. Phylogeny and evolution of the Australo-Papuan honeyeaters (Passeriformes, Meliphagidae). Molecular Phylogenetics and Evolution 31 (3): 943-960.

Gelang, M., A. Cibois, E. Pasquet, U. Olsson, P. Alström & P. G. P. Ericson. 2009. Phylogeny of babblers (Aves, Passeriformes): major lineages, family limits and classification. Zoologica Scripta 38: 225-236.

Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

Kawakami, K., S. Harada, T. Suzuki & H. Higuchi. 2008. Genetic and morphological differences among populations of the Bonin Islands white-eye in Japan. Zoological Science 25 (9): 882-887.

Morioka, H., & T. Sakane. 1978. Observations on the ecology and behavior of Apalopteron familiare (Aves, Meliphagidae). Memoirs of the National Science Museum, Tokyo 11: 169-188, pl. 7-8.

Suzuki, T., & H. Morioka. 2005. Distribution and extinction of the Ogasawara Islands honeyeater Apalopteron familiare on Chichijima, Ogasawara Islands. Journal of the Yamashina Institute for Ornithology 37 (1): 45-49.

The Tomb of the Unknown Honeyeater

Some of the birds referred to in this post. Clockwise from top left - Bombycilla garrulus, the Bohemian waxwing, Bombycillidae; Chaetoptila angustipluma, a Hawaiian honeyeater; two true honeyeaters (Meliphagidae), Anthochaera carunculata (red wattlebird) and Prosthemadera novaeseelandiae (tui); and Moho nobilis, the Hawai'i 'o'o, a Hawaiian honeyeater. Painting by John Anderton.

Fleischer, R. C., H. F. James & S. L. Olson. 2008. Convergent evolution of Hawaiian and Australo-Papuan honeyeaters from distant songbird ancestors. Current Biology 18: 1-5.

GrrlScientist brought my attention yesterday to an interesting new publication on the phylogeny of the Hawaiian honeyeaters. Not, I hasten to explain, the Hawaiian honeycreepers, the Drepanidini clade of birds unique to Hawaii that has become famed for their remarkable adaptive radiation into a whole range of ecological niches, but a smaller clade of five species, Chaetoptila angustipluma and the four species of 'o'o (Moho), that is also unique to Hawaii.

The honeyeaters of the family Meliphagidae are a sizable, fairly heterogenous assemblage of songbirds (Oscines) that are found throughout the Australo-Papuan region, with outliers on various Pacific islands such as New Zealand and Samoa. Despite including a diversity of morphologies, meliphagids are well established as a family, united by features such as a brushed tongue used for taking nectar from flowers (hence, of course, the name "honeyeaters"). The five Hawaiian species share many of these features, and are fairly similar in appearance to Australasian meliphagids, so have always been regarded as meliphagids themselves. The study being discussed here found in a DNA phylogenetic analysis that this was not the case.

Conducting a molecular analysis of Hawaiian honeyeaters is a remarkable achievement in itself because, tragically, not one of the five species remains alive today. All became extinct in the last two centuries. The last surviving species was the smallest, the Kauai 'o'o (Moho braccatus), the last male of which was sighted in 1987 (just to turn the pathos up a notch, a short video of this last individual can be seen here). The Hawaiian honeyeaters therefore join an all-too-long list of birds extinct on the Hawaiian islands since human colonisation, such as the moa-nalo. In the absence of living specimens, Fleischer et al. had to extract DNA from museum specimens, but were able to do so for all five species.

Phylogenetic analysis of these samples showed that, as mentioned above, Chaetoptila and Moho were not related to the true meliphagids. As I've explained elsewhere, recent molecular analyses have consistently identified three large clades within the songbirds (as well as a smattering of smaller clades), the Meliphagoidea (including the meliphagids), Corvoidea and Passerida (including most Northern Hemisphere songbirds). The Hawaiian honeyeaters are not members of the Meliphagoidea, but instead belong to an entirely different clade, the Passerida. Within the Passerida, they belong to an assemblage that includes the Holarctic waxwings (Bombycilla), North American silky flycatchers (Ptilogonatinae) and Caribbean palmchat (Dulus dominicus). Most authors have united these birds in the family Bombycillidae, and the name was recently used in this sense by Spellman et al. (2008). While Fleischer et al. (2008) establish a new family Mohoidae for the Hawaiian honeyeaters, that clade would belong within Bombycillidae in the broad sense. Relationships of the Bombycillidae within the Passerida remain largely unresolved.

The Hawaiian honeyeaters have not been the first birds to abscond from the Meliphagidae in recent years. I have previously discussed the discovery that the New Zealand stitchbird (Notiomystis cincta) is related to the New Zealand wattlebirds, and perhaps a basal member of the Corvoidea. The South African sugarbirds of the genus Promerops, long unsettled as meliphagids, belong to the Passerida and are basal members of the Passeroidea assemblage that includes finches and sparrows (Beresford et al., 2005). The Bonin honeyeater (Apalopteron familiare) is also a member of the Passerida, and falls within the family of white-eyes, Zosteropidae (Driskell & Christidis, 2004) - which I wasn't too surprised to hear because, if you ignore the "meliphagid" brushed tongue, Apalopteron really does look like a big white-eye. Still, the Hawaiian honeyeaters are probably the most typically "meliphagid-like" birds to be recognised as non-meliphagids.

In another interesting recurring theme in oscine phylogeny, the reclassification of Hawaiian honeyeaters, while morphologically unexpected, makes a certain degree of biogeographic sense. Most colonisation of the Hawaiian islands seems to have been derived from North America rather than the western part of the Pacific, with Hawaiian honeycreepers, warblers, geese and violets, among others, all having demonstrated North American (and often northern North American) affinities. As pointed out by a commentator at GrrlScientist's post linked to above, the only Hawaiian bird that still possesses western Pacific affinities is the monarch flycatcher Chasiempis sandwichensis, whose Monarchidae affinities were supported by Filardi & Moyle (2005).

That Hawaiian honeyeaters are such a distinct lineage makes their loss all the more tragic. An extra dose of tragedy that verges on the comic surrounds the most distinct of the mohoids, Chaetoptila angustipluma. Those of my readers who have heard of it before may have noticed that I have deliberately avoided using the vernacular name given to this bird, the kioea. My reason for doing so is that there is reason to doubt whether this name properly belongs to Chaetoptila at all. "Kioea" is actually the Hawaiian name for the bristle-thighed curlew (Numenius tahitiensis), a migratory wading bird and not very much like a honeyeater at all. References to "kioea" as a seabird include the Kumulipo, the epic poem that recited the genealogy of the Hawaiian royal family:

Hanau ke Kioea ka makua,
Puka kana keiki he Kukuluae'o, lele.

The Kioea was born and became parent,
Its offspring was a Kukuluaeo [stilt, Himantopus knudseni], and flew.

--Hawaiian text from here, 1897 translation by Queen Liliuokalani.

Peale (1848) provided no common name for Chaetoptila angustipluma when he first described it (as Entomiza angustipluma - Entomyza is a meliphagid genus). Bryan & Greenway (1944) gave the name "kioea" for this species, but with a question mark, and they also used the name elsewhere for the curlew - I haven't been able to find whether this is the first recorded association between the name and Chaetoptila. Not only has Chaetoptila been cruelly forced out of existence, but it has potentially been subjected to the indignity of a name that is not its own. It truly is the unknown honeyeater.

REFERENCES

Beresford, P., F. K. Barker, P. G. Ryan & T. M. Crowe. 2005. African endemics span the tree of songbirds (Passeri): molecular systematics of several evolutionary ‘enigmas’. Proceedings of the Royal Society of London Series B – Biological Sciences 272: 849-858.

Bryan, E. H., Jr & J. C. Greenway Jr. 1944. Check-list of the birds of the Hawaiian islands. Bulletin of the Museum of Comparative Zoology 94 (2): 92-140.

Driskell, A. C., & L. Christidis. 2004. Phylogeny and evolution of the Australo-Papuan honeyeaters (Passeriformes, Meliphagidae). Molecular Phylogenetics and Evolution 31 (3): 943-960.

Filardi, C. E., & R. G. Moyle. 2005. Single origin of a pan-Pacific bird group and upstream colonization of Australasia. Nature 438 (7065): 216-219.

Peale, T. R. 1848. Mammalia and Ornithology. C. Sherman: Philadelphia.

Spellman, G. M., A. Cibois, R. G. Moyle, K. Winker & F. K. Barker. 2008. Clarifying the systematics of an enigmatic avian lineage: what is a bombycillid?. Molecular Phylogenetics and Evolution 49 (3): 1036-1040.

The Shrikes of the South

Black-backed magpie (Gymnorhina tibicen), a member of the butcherbird family Cracticidae. Australian magpies are not closely related to the Eurasian true magpies (which are members of the crow family). They are best known for their vibrant warbling songs, and thinly veiled homicidal tendencies. Phot by Don Herbison-Evans.

Like them or loath them, there can be little argument that the introduction of molecular methods in the latter part of the last century revolutionised the study of phylogeny and evolution. In many cases, the results of molecular studies supported the theories already proposed about which taxa are related to which, and how. In other cases, molecular data came up with results that strongly contradicted what we thought we already knew. And in some cases, molecular studies gave results that had never been suggested before, but seemed perfectly reasonable in hindsight.


Black-headed gonolek (Laniarius erythrogaster), a member of the African bush-shrike family Malaconotidae. Photo by Derek Ramsey.

I have spoken elsewhere about the new picture of oscine (songbird) phylogeny which has arisen from molecular studies of the group. I'd recommend reading the second and third paragraphs of the post I've just linked to for the background to what I'm just about to talk about. Let it suffice to say that most of the Holarctic families of oscines belong to a clade called Passerida, which is nested within a series of mostly Australo-Papuan clades. Of these Australo-Papuan clades, the most diverse is the Corvoidea, which also includes a number of taxa that have dispersed outside of Oceania such as the crows and orioles (the proper Old World orioles, that is, not the American birds known as 'orioles' which are not orioles at all but members of the Passerida). Also included in the Corvoidea are the shrikes, Old World birds with hook-tipped bills that are more predatory than your average songbird.


White-breasted woodswallows (Artamus leucorynchus). Photo by Romy Ocon.

In 2004, Barker et al. published a phylogeny of the oscines that resolved a number of clades within the Corvoidea. Among the most interesting clades identified by this study was one that united the bush shrikes (Malaconotidae) and helmet-shrikes (Prionopidae) of Africa (both previously counted as subfamilies of the Laniidae) with the woodswallows (Artamus) and butcherbirds (Cracticidae) of Australia. This was definitely one of the third class of molecular results that was unexpected but sensible, as the cracticids are in many ways the shrikes of Australia. The new clade, which lacks a name but which for convenience I'll dub the 'malaconotoid clade', therefore combines many of the world's shrike-like birds in one convenient package. The notable exceptions are the true Eurasian shrikes of the stripped-down family Laniidae, which are Corvoidea but whose affinities seem to lie elsewhere as the sister group of the crows, and the Asian shrike-babblers of the genus Pteruthius, which Reddy and Cracraft (2007) showed to be corvoids related to the American vireos.


Bornean bristlehead (Pityriasis gymnocephala). Authors had disagreed continuously over the years about whether Pityriasis was related to the shrikes or the cracticids - a somewhat ironic argument since the recognition of the malaconotoid clade. Photo by James Eaton - photos of this retiring bird are few and far between, making this all the more impressive.

Comparison with the other corvoids suggests that the malaconotoids had an Australian origin, a suggestion corroborated by the fact that the basalmost split in the group seems to be between the Australian taxa on one side and the African taxa on the other (Barker et al., 2004; Moyle et al., 2006). As with all good scientific theories, this brings up a further question - how did the malaconotoids get from Australia to Africa? To answer this question, it turns out that a collection of small southern Asian families also fall into the malaconotoid clade - the Platysteiridae, the ioras of the genus Aegithina, and the unusual Bornean bristlehead (Pityriasis gymnocephala). The reasonable suggestion might therefore be made that southern Asia was used as a corridor by the malaconotoids on their way to Africa. Unfortunately, the evidence is a little more equivocal in this regard. It is true that the Asian taxa sit on the African side of the malaconotoid clade (Moyle et al., 2006 - Fuchs et al., 2006, found Aegithina to be sister to the Australian taxa, but with low support), but there is no clear division between taxa from the two continents. Instead, the African and Asian taxa are mixed together, suggesting more than one dispersal between the continents and also unclear whether dispersal was from Asia to Africa or vice versa.



The other significant dispersal in the history of the malaconotoids was from Africa to Madagascar, where an ancestral malaconotoid gave rise to the vangas (Vangidae). The vangas were one of the very few oscine groups to reach Madagascar, and once there they formed an island radiation comparable to the honeycreepers of Hawaii or the finches of the Galapagos. While there are only twenty-one species of vanga, the group is spectacularly diverse ecologically, as shown above in a figure from Yamagishi et al. (2001) - so much so, in fact, that many of the species had been assigned to separate families and were only recognised as vangas recently (see Don Roberson's page on the family for further details). Notable in this regard are the handsome sickle-billed vanga (Falculea palliata), the coral-billed nuthatch (Hypositta corallirostris) which bears an uncanny resemblance to the unrelated true nuthatches, and the large-billed helmetbird (Euryceros prevostii).

REFERENCES

Barker, F. K., A. Cibois, P. Schikler, J. Feinstein & J. Cracraft. 2004. Phylogeny and diversification of the largest avian radiation. Proceedings of the National Academy of Sciences of the USA 101: 11040-11045.

Fuchs, J., J. Fjeldså & E. Pasquet. 2006. An ancient African radiation of corvoid birds (Aves: Passeriformes) detected by mitochondrial and nuclear sequence data. Zoologica Scripta 35 (4): 375-385.

Moyle, R. G., J. Cracraft, M. Lakim, J. Nais & F. H. Sheldon. 2006. Reconsideration of the phylogenetic relationships of the enigmatic Bornean bristlehead (Pityriasis gymnocephala). Molecular Phylogenetics and Evolution 39 (3): 893-898.

Reddy, S., & J. Cracraft. 2007. Old World shrike-babblers (Pteruthius) belong with New World vireos (Vireonidae). Molecular Phylogenetics and Evolution 44 (3): 1352-1357.

Yamagishi, S., M. Honda, K. Eguchi & R. Thorstrom. 2001. Extreme endemic radiation of the Malagasy vangas (Aves: Passeriformes). Journal of Molecular Evolution 53: 39-46.

On Hybrid Birds

Fuller, E. 1995. The Lost Birds of Paradise. Swan-Hill Press.

The 19 "lost" birds of paradise that Errol Fuller describes in this book are forms that are mostly only known from very few specimens, often with very little supporting information. What makes them "lost", however, is that despite all but one of them being described as new species, all of them were later reinterpreted as hybrids between better-known species. Fuller's motivating question is whether these specimens are indeed hybrids, or represent valid species that might occupy unknown restricted ranges somewhere in the depths of New Guinea, or may perhaps have slipped into extinction without ever getting the recognition they deserved.

Most of the specimens reached Europe through the plume trade. Specimens of birds of paradise were purchased from native collecters and then shipped back to the West for use in the fashion industry (the first country to ban the import of birds of paradise for plumes, according to Fuller, was the US in 1913, a development that probably had less to do with the developing conservation movement than with the increasing unfashionability of wearing plumes*). As a result, the available collection data on most specimens is decidedly hazy - few bear more specific information than "Dutch New Guinea" (the western part of New Guinea that is now controlled by Indonesia). Even if a more specific locality is recorded, it is often unreliable - specimens could be passed through a number of different native tribes before eventually reaching the European traders. Throughout the book, we get introduced to many of the figures involved in the collection and study of these mystery birds.

*To add further complexity, a major factor in the decline of popularity of plumes was actually the rise in popularity of the motor-car - ornate plumed hats being decidedly impractical for wearing in open-topped cars.

'Paradisaea mirabilis', a possible hybrid of Paradisaea minor (lesser bird of paradise) and Seleucidis melanoleuca (twelve-wired bird of paradise). 1902 lithograph by Bruno Geisler.

Throughout Fuller's book, we get introduced to many of the personages involved in the collection and study of the specimens (including the spectacularly named Captain Neptune Blood*). A significant number passed through the collection of Lord Walter Rothschild, a somewhat eccentric enthusiast who amassed one of the world's largest private natural history collections (Darren Naish wrote a piece two years ago on Rothschild and his unusual enthusiasm for cassowaries), though many of Rothschild's bird of paradise specimens were included in the collection he was blackmailed into selling to the American Museum of Natural History**. The "hybrid" specimens were largely identified as such in 1930 by Erwin Stresemann. Fuller accuses Stresemann of overzealousness in embracing his hybrid theory of origin for "species" known only from one or two specimens, essentially assuming from the start that all such species must be hybrids of known species and identifying "parents" from the available options no matter how poorly supported.

*Seriously.

**Probably not, I hasten to add, by the American Museum of Natural History.

'Diphyllodes gulielmitertii', almost certainly a hybrid between Diphyllodes magnificus and Cicinnurus regius. Unlike most of the other forms described by Fuller, this hybrid occurs fairly commonly, with more than two dozen known specimens. Lithograph by J. Gould and W. Hart.

Unfortunately, many of Fuller's reinterpretations of the supposed hybrids end up falling a little flat. Fuller accepts a hybrid origin for some forms, refutes it for others, but in many cases it is debatable whether his interpretations are any better than Stresemann's. Because all Fuller has to go on is examination of specimens, most of his arguments for valid species status amount to little more than replying to Stresemann's statement that "Species A has features intermediate between those of B and C, and is therefore a hybrid between the two" with "No it doesn't, so it isn't". In two cases where Fuller does accept hybrid status, 'Loborhamphus ptilorhis' and 'Lamprothorax wilhelminae', the reasons for linking them to their supposed parents seem decidedly unconvincing (which, of course, does not eliminate the possibility that they could still be hybrids between other species), while 'Cicinnurus lyogyrus', which Fuller hesitatingly accepts as a hybrid of the king bird of paradise (Cicinnurus regius) and the magnificent bird of paradise (Diphyllodes magnificus) seems more likely to be simply an aberrant variant of Cicinnurus regius. In contrast, 'Janthothorax bensbachi', which Fuller suggests is a valid species, seems more likely to be a hybrid. Probably DNA analysis of the specimens would be the only way to convincingly decide the question one way or another - Fuller suggests this would be difficult because the close relationships of the parent species would make results unconvincing, but resolution of molecular analyses has decidedly improved since 1995. The main barrier would be that DNA extraction from museum specimens, especially ones that have been in storage since the 1800s (and were probably not exactly fresh when they first reached the museum) is a difficult process, with little guarantee of success.

Loborhamphus nobilis, regarded by Stresemann (1930) as a hybrid between Paradigalla carunculata and Lophorina superba, but by Fuller as a probable valid species. Unlike the other bird of paradise species mentioned in this post, the less sexually dimorphic Paradigalla species are not polygamous breeders, and form permanent pair bonds. They therefore strike me as less likely to produce hybrids.

The Lost Birds of Paradise is certainly a lavishly illustrated book, reproducing paintings by Gould and other spectacular bird illustrators (some of which I've taken the liberty of re-reproducing), as well as numerous photos and drawings. The distribution and subjects of these illustrations are often a little erratic, however - what's with the naked man in the bath on page 76? - and this same erraticism extends to the text. I can't escape the impression that most of the essays on the various birds were composed separately, with little cross-checking between chapters when the book was compilated. The story of Stresemann's 'overzealous' revisions is repeated a number of times in different chapters, for instance, while many chapters include rather tangential passages on matters related to birds of paradise in general, but not necessarily directly relevant to the specific form the chapter is devoted to (not surprisingly, this is particularly noticeable in some forms known only from single specimens for which otherwise Fuller probably just wouldn't have had that much to say). Probably this eclecticism is most marked in the chapter on 'Paradisaea mixta', in which we are treated to a lengthy quotation from the autobiography of Errol Flynn (complete with full-page photograph) and a description of his experiences trying to start a career collecting birds of paradise in New Guinea some years before he became an actor. And what does this have specifically to do with 'Paradisaea mixta'? As it happens, absolutely nothing.

Still, The Lost Birds of Paradise is easily readable, and at least highlights that the identity of many of the "hybrid" birds of paradise is not as firm as might be thought. A commentor on one discussion thread makes the comment that Fuller obviously really wants there to be overlooked species of birds of paradise, which may lead him to be a bit more hasty in his judgements than he probably should be. Nevertheless, New Guinea, especially the western half, is a surprisingly unexplored place, and as the recent discovery/rediscovery of unknown or near-unknown mammal species there shows, it would be wise to not rule anything out just yet.