Paracorvids

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Paracorvids

Unlike the passerines and suboscines, the corvid assemblage has not been readily identifiable and taxa are still being moved between the corvid groups and the Passerida. Sibley, Ahlquist, and Monroe took the first step of gathering the corvids together (albeit imperfectly). This is reflected in Gill's family list (1995) and the 3rd edition Howard-Moore checklist, which did a lot to group them in reasonable families. Checklists of a more traditional sort, such as Clements 5th edition list place the corvid assemblage all over the map.

Sibley and Ahlquist's view was that the remaining passerines split cleanly into a corvid group (Corvida) and a group containing everything else (Passerida). Further study has shown that reality is more complex. Unlike the Passerida, their version of the corvids was not a monophyletic group. Nonetheless, there seems to be a core group of corvids that is sister to the Passerida. In between are several groups that we might call paracorvids (Menurida, Climacterida, Meliphagida, Pomatostomida, Orthonychida). They branch off separately before the split between the Corvida proper and Passerida (see Ericson et al., 2002a; Barker et al., 2004; Irestedt and Ohlson, 2008).

It is striking how the initial corvid radiation was confined to Australasia. We see this in the distribution of the paracorvid groups. All of the Menurida, Climacterida, and Pomatostomida are Australasian. Only Meliphagida has any species outside the area. Even there, two of the four families (Maluridae and Dasyornithidae) are also entirely Australasian. Further, only one of Pardalotidae crosses Wallace's line—the Golden-bellied Gerygone. That leaves the Meliphagidae, which have spread widely across Australasia and Oceania, with several species coming near Wallace's line. Even so, only one of them, the Indonesian Honeyeater, manages to ranges even barely into Indo-Malaya.

The recent paper by Jønsson et al. (2011b) offers some evidence concerning a possible relation between the emergence of the proto-Papuan archipelago and the corvid radiation. The New Scientist has a summary.

Menurida Sharpe, 1891

The first paracorvid branch, Menurida, is endemic to Australia. It consists of the lyrebirds (Menuridae) and scrub-birds (Atrichornithidae). Ericson et al. (2002b) found the Menurida sister to the rest of the oscines, but did not include scrub-birds in his analysis. Morphological analyses had placed the scrub-birds next to the lyrebirds. A genetic analysis was recently carried out by Chesser and ten Have (2007). It concurs that the lyrebirds and scrub-birds are sister families, and also concurs with basic tree we present here. For a discussion of the history of lyrebird and scrub-bird taxonomy, see the Ericson et al. and Chesser and ten Have papers, respectively.

Menuridae: Lyrebirds Lesson, 1828

1 genus, 2 species HBW-9

Atrichornithidae: Scrub-birds Stejneger, 1885 (1875)

1 genus, 2 species HBW-9

Climacterida Informal

The Climacterida, Meliphagida, Pomatostomida, and Orthonychida don't appear to have been formally named. They tend to be lumped together under terms such as “basal Corvida”. Nonetheless, it's been clear to many that there are 3-4 distinct clades here. I find it convenient for them to have names, so I've been using the first 3 since 2007, and started regarding the last as a parvorder in Feb. 2011.

Climacterida The Climacterida are the next branch. There are two families here: Australasian treecreepers (Climacteridae) and bowerbirds (Ptilonorhynchidae). These families are endemic to Australasia.

The overall taxonomy is based on Ericson et al. (2002b) for Climactrida, and Christidis et al. (1996) and Kusmierski et al. (1997) for the bowerbirds.

Climacteridae: Australasian Treecreepers de Selys-Longchamps, 1839

2 genera, 7 species HBW-12

Ptilonorhynchidae: Bowerbirds G.R. Gray, 1841

6 genera, 20 species HBW-14

Based on Zwiers et al. (2008), Sericulus ardens, has been split from Sericulus aureus. Interestingly, they are not each other's closest relatives. The name Flame Bowerbird follows S. ardens while S. aureus becomes Masked Bowerbird.

Meliphagida Informal

There are four families in the Meliphagida: Australasian wrens (Maluridae), bristlebirds (Dasyornithidae), gerygones and allies (Pardalotidae), and honeyeaters (Meliphagidae). We follow the order in Gardner et al. (2010). The Dasyornithidae have sometimes been included in Pardalotidae. However, the DNA shows that the Dasyornithidae are a separate branch of the Meliphagida.

Maluridae: Australasian Wrens Swainson, 1831

5 genera, 29 species HBW-12

Maluridae
Click for Maluridae tree

The Maluridae are another family that is restricted to Australia and New Guinea. The arrangement here is based on Lee et al. (2011), Christidis et al. (2010) (Amytornis), and Driskell et al. (2011) (the other genera). Although these are not in 100% agreement, the differences are minor. When applicable, I've followed Lee et al. (2011), which uses considerably more data. Gardner et al. (2010) has sparser taxon sampling and includes only a little genetic data for these species. They give a slightly different tree.

Driskell et al. (2011) found that the broad-billed fairywrens were more closely related to Sipodotus and Clytomyias than to the rest of Malurus. They recommend putting them in a separate genus, Chenorhamphus (Oustalet 1878, type grayi), which I have done here.

Driskell et al. (2011) found that the Lovely Fairywren, M. amabilis is nested within the Variegated Fairywren, M. lamberti. In particular, they found that M. lamberti assimilis is more closely related to the allopatric M. amabilis than to M. lamberti lamberti. However, assimilis reportedly interbreeds with lamberti where they meet. Further study seems warranted. One possibility is that the situation here is analogous to that of the Chihuahuan Raven and the California clade of Common Ravens. Another is that they should all be one species.

Based on Black et al. (2010) and Christidis et al. (2010), Thick-billed Grasswren, Amytornis textilis, is split into Western Grasswren, Amytornis textilis, and Thick-billed Grasswren, Amytornis modestus.

Dasyornithidae: Bristlebirds Sibley & Ahlquist, 1985

1 genus, 3 species HBW-12

The bristlebirds are endemic to Australia.

Pardalotidae: Gerygones and allies Strickland, 1842

16 genera, 69 species HBW-13

Pardalotidae
Click for genus-level tree for Pardalotidae

The Acanthizidae have been merged with the Pardalotidae. Although early genetic results had suggested that the pardalotes were more closely related to the honeyeaters than to the gerygones and thornbills, that seems to be incorrect. Relatively complete analyses, such as Garnder et al. (2010), Jønsson et al. (2011b) and Nyári (2011), have found that the pardalotes from a clade with the former Acanthizidae. Indeed, the pardalotes may even be nested inside!

Traditionally, the differences between these taxa have been considered relatively small, with all of them sometimes placed in a single family with the bristlebirds. We now know that the bristlebirds do not belong in this group, but that the remainer form a natural group of broadly similiar species. Such a group is best treated as a single family, and the name Pardalotidae has priority.

Although the Pardalotidae are primarily Australasian, with ranges east and south of Wallace's line, there is one exception—the Golden-bellied Gerygone. It ranges north to the Philippines and west to Malaysia and Sumatra.

The genus-level phylogenetic tree is mostly based on Gardner et al. (2010) and Nyári (2011), with some help from Norman et al. (2009b), Nicholls et al. (2000), and Christidis et al. (1988). A number of species, especially those outside of Australia and New Guinea, still remain to be tested. They are marked with question marks on the tree.

Christidis and Boles (2008) was also consulted during the process. I've decided to use their generic limits, putting the heathwrens in Hylacola and the Speckled Warbler in Chthonicola.

Recent work by Norman et al. has led to some adjustment of the boundaries of Pardalotidae. Norman et al. (2009a) showed that the mohouas are not part of Pardalotidae, but rather belong in Corvoidea. In a second paper, Norman et al. (2009b) showed that the Goldenface, Pachycare flavogriseum, does belong in Pardalotidae, not Pachycephalidae, Petroicidae, or anyplace else that had previously been suggested.

Meliphagidae: Honeyeaters Vigors, 1825

46 genera, 183 species HBW-13

Meliphagidae Although they have spread widely. The Meliphagidae are primarily Australasian. Some of them have colonized various islands in Oceania, and several have spread into Wallacea, including the Lesser Sundas, Moluccas, and Sulawesi, but only one occurs outside Australasia and Oceania. That is the Indonesian Honeyeater. It barely crosses Wallace's line into Bali, which is considered part of Indo-Malaya (aka the Oriental Region).

The honeyeaters were substantially restructured by Sibley and Ahlquist (1990), losing the genera Cleptornis (Passerida), Oedistoma and Toxorhamphus (Melanocharitidae), and Promerops (Promeropidae), but gaining Epthianura and Ashbyia from the defunct Epthianuridae. Sibley and Ahlquist's addition of the Epthianura and Ashbyia to the honeyeaters has been supported by the more recent studies of Driskell and Christidis (2004) and Nyári and Joseph (2011).

Spring et al. (1995) found that the Bonin Honeyeater, Apalopteron familiare is not a honeyeater, but rather belongs in the Passerida (more precisely, Zosteropidae). More recently, Cracraft and Feinstein (2000) found that MacGregor's Bird-of-paradise, Macgregoria pulchra, is actually a honeyeater, while Ewen et al. (2006) and Driskell et al. (2007) found that the Stitchbird, Notiomystis cincta belongs near the Callaeidae, where it becomes a monotypic family. The now-extinct Hawaiian honeyeaters (genera Moho and Chaetoptila) were formerly considered to be part of this family, but they have recently been found to be related to waxwings (Fleischer et al., 2008).

Between Driskell and Christidis (2004), Cracraft and Feinstein (2000), Norman et al. (2007), Higgins et al. (2008 = HBW-13), Gardner et al. (2010), and Nyári and Joseph (2011), there is enough information for a reasonable genus-level and even species-level taxonomy. The overall structure of the family is based on Driskell and Christidis (2004), which is similar to Gardner et al. (2010) and Nyári and Joseph (2011). In the true Meliphagidae, Driskell and Christidis found five major clades. They found the spinebills to be basal, but that is less clear after looking at Nyári and Joseph (2011). The various analyses have not resulted in consistent relation between the clades, so I'm treating them as a 5-way polytomy, with each clade ranked as a subfamily and listed in order of number of species.

The two spinebills lead off (Acanthorhynchinae) followed by Epthianurinae. Epthianurinae contains the Australian Chats as well as various honeyeaters. In their study of MacGregor's “Bird-of-paradise”, Cracraft and Feinstein (200) included representatives of four subfamilies. This is enough to show that Macgregoria is in the Epthianurinae. Consideration of plumage suggests it is sister to Melipotes.

Myzomelinae is next. The genera other than Sugomel and Myzomela sometimes show up elsewhere on the tree, but Driskell and Cristidis (2004) note that they share two indels with Sugomel and Myzomela, so this is probably the correct place for them. The position of Gliciphila, sometimes placed in Phylidonyris, is a little uncertain. It's definitely not in Phylidonyris, but it could be closer to the Glycichaera-Ptiloprora group or to the Sugomel-Myzomela group. Note that Sugomel nigrum is often placed in Certhionyx, but Driskell and Christidis showed this was mistaken.

Philemoninae contains the friarbirds and allied honeyeaters. Some of its members have moved to new genera. Like Sugomel, Cissomela pectoralis is often placed in Certhionyx. Two species are moved from Lichenostomus and placed in genus Nesoptilotis (Mathews 1913, type flavicollis): White-eared Honeyeater, Nesoptilotis leucotis, and Yellow-throated Honeyeater, Nesoptilotis flavicollis. These are rather different from the rest of the former Lichenostomus, so it's not surprising that they end up in a different genus. I had originally made this change based on Higgins et al. (2008) and Gardner et al. (2010), before DNA data was available for both species. It is also supported by the complete analysis of Lichenostomus of Nyári and Joseph (2011).

Gardner et al. (2010) had shown that Lichenostomus was polyphyletic, even after removing Nesoptilotis. Nyári and Joseph (2011) carried out a complete analysis of Lichenostomus using the mitochondrial ND2 and nuclear &beta-fibrinogen-7 genes. Their results were generally consistent with Gardner et al. (2010), but revealed some additional surprises that require further dismemberment of Lichenostomus and the use of three more genera: Bolemoreus, Caligavis, and Stomiopera. The Bridled Honeyeater (frenatus) and Eungella Honeyeater (hindwoodi) turn out to be sister to the wattlebirds plus Acanthagenys. Nyári and Joseph (2011) created the new genus Bolemoreus (type frenata) to accomodate them.

If you look at the tree, you'll see that the first few genera in Meliphaginae are shown in magenta. This is to highlight the fact that there is some uncertainty about whether they belong in Meliphaginae (Driskell and Christidis, 2004) as Nyári and Joseph (2011) put them in Acanthorhynchinae.

The other pieces of the former Lichenostomus run in a grade from Pitlotula to the miners (Manorina). For the first clade, Singing through Yellow-tinted Honeyeaters, the name Ptilotula (Mathews 1912, type flavescens) has priority Paraptilotis (Mathews 1912, type fuscus) because Ptilotula was chosen as a subgenus name by Schodde and Mason (1999) in preference to Paraptilotis. Then come the White-gaped and Yellow Honeyeaters, for which the name Stomiopera is available (Reichenbach 1852, type unicolor). This group is followed by three more former Lichenostomus honeyeater, now placed in Caligavis (Iredale 1956, type obscura). The Lichenostomus grade is interrupted by Purnella albifrons, sometimes placed in Phylidonyris. Then we finally reach the remaining Lichenostomus, now reduced to two species.

Based on Toon et al. (2010), Swan River Honeyeater / Western White-naped Honeyeater, Melithreptus chloropsis has been split from White-naped Honeyeater, Melithreptus lunatus. They also found evidence that the White-throated Honeyeater, Melithreptus albogularis, may contain more than one species, but more study is necessary to clarify the situation.

I've grouped three genera at the end. I don't have any good reason to put them in any particular place, so they end up incertae sedis. This includes the two Glycifohia, which are sometimes included in Phylidonyris. It might make sense to add the Gray Honeyeater, Conopophila whitei, to this group. It has been placed in the monotypic genus Lacustroica, and Christidis and Boles (2008) express some doubt as to whether it belongs in Conopophila.

Acanthorhynchinae: Spinebills Mathews, 1946

Epthianurinae: Australian Chats & allies Legge, 1887

Myzomelinae: Myzomela & allies G.R. Gray, 1840

Philemoninae: Friarbirds & allies Lesson, 1828

Meliphaginae: Honeyeaters, Wattlebirds, Miners Vigors, 1825

Incertae Sedis: Meliphagidae

Orthonychida Informal

The last two paracorvid families are the logrunners (Orthonychidae) and Australasian babblers (Pomatostomidae). These may be deep separated branches (see Barker et al., 2004; Jønsson et al., 2011b). This is emphasized by treating them as separate parvorders. Alternatively, they may actually be sister groups (see Barker et al., 2002; Irestedt and Ohlson, 2008), in which case the would together form the Pomatostomida. In any event, they split off before the division between the Corvida and Passerida, which means they are in the paracorvids.

Orthonychidae: Logrunners G.R. Gray, 1840

1 genus, 3 species HBW-12

The logrunners are found in Australia and New Guinea.

Pomatostomida Informal

Pomatostomidae: Australasian Babblers des Murs, 1860

2 genera, 5 species HBW-12

As befits their name, the Australasian babblers are found in Australia and New Guinea.

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