Corvida I

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Corvida Wagler, 1830

At this point the passerines divide into two branches, the Corvida and Passerida. Corvida is the smaller branch, so we consider it first. The use of “Corvida” for the clade sister to Passerida follows the logical terminology of Cracraft et al. (2004), although the term Corvoidea, or even “core Corvoidea” is in common use. The actual membership of Corvida is a bit different from Cracraft et al. as some families have been moved eleswhere in the phylogeny.

The taxonomy in Barker et al. (2004) suggests Corvida is divided into two superfamilies, Callaeoidea and Corvoidea. However, Irestedt and Ohlson (2009) argue that the families in Barker et al.'s Callaeoidea (Melanocharitidae, Cnemophilidae, Callaeidae, and by implication, Notiomystidae) are really basal Passerida. They show that if the analysis is done without the RAG-1 gene, these families end up in Passerida. Apparently RAG-1 has a strong signal that overwhelms conflicting evidence from other genes. Since the RAG-1 signal is not supported by other genes, they argue it is spurious. Although there is room for dispute here, I found Irestedt and Ohlson's argument sufficiently convincing to follow it here.

In early 2011, the most comprehensive study of the Corvida is the six-gene analysis by Jøsson et al. (2011b), which includes one of more representative from each of the corvid families except the monotypic Pityriaseidae (bristlehead).

Interestingly, they not only include Callaeidae in Passerida, but place it a little deeper than Irestedt and Ohlson. Jønsson et al. do have Cnemophilidae and Melanocharitidae as basal groups in Corvida, and probabaly sister to one another.

Although Jøsson et al.'s results were not entirely clear-cut, they found evidence for a grade of corvid groups, including 4 major groups and 4 minor groups. The major groups are here called: Campephagoidea, Orioloidea, Malaconotoidea, and Corvoidea. The last two are essentially the same as in Cracraft et al. (2004).

Psophodoidea

Psophodidae: Whipbirds & allies Bonaparte, 1854

7 genera, 20 species HBW-12, as Eupetidae

Psophodidae tree

This small Australasian group is the basal group in Corvida. It consists of a single family—Psophodidae. I have previously treated these taxa in various ways (e.g., based on Norman et al., 2009a). Most of them were recently divided into 3 families (the other two being Cinclosomatidae and Paramythiidae). They have been collected together and Eulacestoma has been moved from the Pachycephalidae based on Jønsson et al. (2011b).

Most other studies that include Ptilorrhoa also put it in a relatively basal position in Corvida, with mulitgene studies by Jønsson et al. (2007), Irestedt and Ohlson (2008), and Irestedt et al. (2008) putting it at the base. However, Norman et al., (2009a) place it (and Cinclosoma) closer to the Corvoidea.

Although there is some support for including Falcunculus, in Psophodidae (e.g, Barker et al., 2004; Dumbacher, 2008), the multigene analyses of Norman et al. (2009a) and Jønsson et al. (2011b) separate them. Note that Norman et al. (2009a) put Eulacestoma near or in the Pachycephalidae.

Sibley and Monroe's Cinclosomatinae included two extra genera: Eupetes and Ifrita, while HBW-12's corresponding Eupetidae (del Hoyo et al., 2007) also includes Melampitta. Barker et. al (2004) suggested that Melampitta belonged in the monarchs (Monarchidae), but with weak support. They and the mudnesters (Corcoracidae) were thought to form a clade. Reddy and Cracraft (2007), using the same genes, found Melampitta either with the monarchs or mudnesters. Dumbacher et al. (2008) found it related to Ifrita, but did not include the necessary taxa to say where they goes. Irestedt et al. (2008) considered Melampitta basal in the narrowly construed Corvoidea, while Norman et al. (2009a) include Ifrita in the monarchs. The most comprehensive study, Jønsson et al. (2011b) puts Ifrita in the monarchs and Melampitta in a clade containing the mudnesters and birds-of-paradise.

Jønsson et al. (2007) showed that the rail-babbler Eupetes was really related to Chaetops and Picathartes. It is now in its own family Eupetidae within Passerida.

The painted berrypeckers (Paramythiidae) were previously removed from the Melanocharitidae (Passerida) and placed in their own family. I'm not sure they (or the other groups here) are distinctive enough for this and have included them in Psophodidae.

The genus Androphobus is sometimes merged into Psophodes.

Toon et al. (2012) examined DNA from a number of Quail-thrushes. The arrangement of Cinclosoma is based on their work. They found that at least two subspecies should be regarded as full species. This involves splitting Western Quail-thrush, Cinclosoma marginatum from Chestnut-breasted Quail-thrush, Cinclosoma castaneothorax (both now monotypic) and Nullarbor Quail-thrush, Cinclosoma alisteri, from Cinnamon Quail-thrush, Cinclosoma cinnamomeum (including tirariense).

Campephagoidea

The first sizeable group is the Campephagoidea, consisting of Mohouidae and Campephagidae.

Mohouidae: Whitehead & allies Matthews, 1946

2 genera, 3 species Not HBW Family

The Mohouidae are New Zealand endemics that have previously been included in many different families. Writing in HBW-13, Boles (2007) mentions Paridae, Timaliidae, Orthonychidae, Campephagidae, Sylviidae, Maluridae, and Acanthizidae (=Pardalotidae). The Acanthizidae option was followed by Sibley and Monroe (1990), Dickinson et al. (2003) and version 2.0 of the IOC checklist (Jan 2009). Boles (2007) includes them in Pachycephalidae. However, the evidence for any of these is very weak. Jønsson et al. (2011b) is the first genetic study to suggest any real affinities for them (sister to Campephagidae), although the support for this could be stronger.

Boles merges Finschia into Mohoua. They appear to be very closely related, possibly congeneric, whether one looks at osteology (Olson, 1990) or DNA hybridization (Sibley and Ahlquist, 1987).

Campephagidae: Cuckooshrikes Vigors, 1825

11 genera, 92 species HBW-10

The ordering of the minivets (Pericrocotus) is based on Jønsson et al. (2010b). The results in Fuchs et al. (2007a) and Jønsson et al. (2008a, b) suggested some generic boundaries need to be redrawn for Coracina and related genera. Jønsson et al. (2010c) analyzed most of the revelant taxa, and the current organization is based on their results.

They found that several of the Coracina cuckooshrikes were actually closer to some of the trillers (Lalage). They advocate use of a broad genus Lalage including the species list here in Malindangia, Cyanograucalus, Celebesica, Analisoma, and Edolisoma. The extra genera here serve to highlight the major groups among Lalage and allies. McGregor's Cuckooshrike, Malindangia mcgregori, is basal among them. The rest are in two clades. One includes the species listed below in Lalage. Further division of this did not seem warranted. The other clade probably includes the rest of the Jønsson et al.'s broad Lalage.

Campephagidae tree The support for grouping Cyanograucalus (Hartlaub, 1861) and Celebesica (Strand, 1928, replacing the preoccupied Celebesia Riley, 1918) is relatively low, so I've put them in separate genera. Indeed, there is some chance they don't even group with the rest of Lalage. I've separated the three Analisoma (Mathews, 1928) because the cicadabirds (Edolisoma, Jacquinot and Pucheran, 1853) form a tight group that is relatively distant from the Analisoma.

Oreoicoidea

Oreoicidae: Crested Bellbird & allies Sibley & Ahlquist, 1985

3 genera, 3 species Not HBW Family

Oreoicidae tree This is another small Australasian group. Jønsson et al. (2011b) included all three of these species in their analysis. Their results show the Oreoicidae on a separate branch. In contrast, Norman et al. (2009a) found a small basal grouping in Malaconotoidea that includes Aleadryas and Oreoica, which Jønsson et al. (2008a) grouped with the former Pitohui, Ornorectes.

One casualty of the restructing of the Corvida is the shrike-thrush family, Colluricinclidae. The Howard-Moore checklist includes two genera. Of those, Colluricinla joins the whistlers (Pacycephalidae), while Pitohui is split into 4 parts (Ornorectes, Pitohui, Pseudorectes, Melanorectes) and scattered across three families and two superfamilies. Both the Crested Pitohui and Crested Bellbird come from the defunct Colluricinclidae.

Although it turns out that they are not so closely related, the pitohuis share an interesting characteristic. They're poisonous! (Dumbacher et al., 1992.) You can read more about these birds at Dumbacher's website. Exactly which species belong to the pitohuis has not been exactly clear, so we shouldn't be entirely surprised that they are split apart. A paper by Jønsson et al. (2008a) finds that the pitohuis are not that closely related to each other. Accordingly, they end up in four genera (Ornorectes, Pitohui, Pseudorectes, and Melanorectes), in Oreoicidae, Oriolidae, and Pachycephalidae (2 places).

Orioloidea

Another large grouping is next. The Orioloidea include the whistler, oriole, and vireo families (Pachycephalidae, Oriolidae, and Vireonidae).

Pachycephalidae: Whistlers Swainson, 1831

6 genera, 54 species HBW-12

Pachycephalidae tree Accordingly to the analysis of Norman et al. (2009a), Eulacestoma is basal, with Falcunculus branching off next. The distance between them and the rest of the Pachycephalidae seems substantial, and it might be reasonable to consider them separate families, or at least subfamilies. This is particularly true of Eulacestoma, where support was weak.

The shrike-thrushes have been included with the whistlers (Pachycephalidae) due to the analysis in Jønsson et al. (2008b, 2010a). They also found that the former Olive-flanked Whistler (Hylocitrea bonensis, now called Hylocitrea) is not only not a whistler, but belongs somewhere in Passerida, not Corvida.

Based on Jønsson et al. (2008b, 2010a), The Sangihe Shrike-thrush has been transferred to Coracornis. Using different data sets, Dumbacher et al. (2008) and Jønsson et al. (2008b, 2010a) found that the Morningbird, which is sometimes placed in Colluricincla, sometimes in Pitohui, actually belongs in Pachycephala. The order in Pachycephala roughly conforms to Jønsson et al. (2010a). There are indications in Jønsson et al. (2010a) that some species limits need revision, but the exact nature of this remains unclear.

Oriolidae: Orioles, Figbirds Vigors, 1825

4 genera, 38 species HBW-13

The affinities of the recently extinct Piopios of New Zealand have long been uncertain. They have been variously considered birds-of-paradise, bowerbirds, thrushes, whistlers, and others. Sometimes they have been placed in their own family. There had been weak genetic evidence that they were close to bowerbirds. Two recent papers have now resolved the puzzle. Johansson et al. (2011) and Zuccon and Ericson (2012) found they are orioles!

Click for Oriolidae tree
Click for Oriolidae tree

The arrangement here is primarily based on the analysis of Jønsson et al. (2010d), with some input from Johansson et al. (2011) and Zuccon and Ericson (2012). The genera are arranged as in Zuccon and Ericson (2012), which uses more genes, while Jønsson et al. (2010d) is followed for Oriolus. Jønsson et al. found evidence that Oriolus chinensis involves at least 3 species (there are 20 subspecies). It also is likely that steerii contains two or more species. The placement of crassirostris is a bit uncertain as it was not sampled.

The genetic results in Jøsson et al. (2010) also support splitting Sunda Golden-Oriole, Oriolus maculatus, and Asian Golden-Oriole, Oriolus diffusus, from Black-naped Oriole, Oriolus chinensis. Jøsson et al. only included four of the twenty subspecies (melanisticus grouped with chinensis). Presumably Asian Golden-Oriole is monotypic, and mostly migratory. Sunda Golden-Oriole probably includes andamanensis, macrosurus, maculatus, mundus, richmondi, sipora lamprochryseus, and insularis, while Black-naped Oriole includes the Philippine races (chinensis, yamamurae, sulensis, and melanisticus). I'm not sure which group the Wallacean races belong to (sangirensis, formosus, celebensis, stresemanni, frontalis, bonratensis, and broderipi), and they are left in the Black-naped group until further information becomes available.

Vireonidae: Vireos Swainson, 1837

6 genera, 62 species HBW-15

Vireonidae tree Next come the vireos. They were once thought to be closely related to the wood-warblers (Parulidae). When Sibley and Ahlquist discovered the corvid group, they found that the vireos (and shrikes) were members of it. Some thought it odd that the vireos, with an old world origin, had no remaining old world members. While it is true that it seemed odd, it is false that there are no old world members. They do have some old world relations. They've been hiding out among the babblers (Timaliidae). One is Erpornis zantholeuca (nee Yuhina zantholeuca). Rather than being a babbler like the other Yuhinas, it is actually a vireo relative. More recently, Reddy and Cracraft (2007) found that the shrike-babblers (Pteruthius) are also vireo relatives. Right now, the choices are either to call them all vireos or to have 3 families of vireos and allies. Since there are now more old world vireos known than before, Epornis and Pteruthius, they seem less special. Thus I take the former route for the present, keeping in mind that the discovery of additional vireo taxa might change things.

Reddy (2008), using the phylogenetic species concept, advocated splitting Pteruthius into 19 species. While this seems extreme, Reddy's evidence suggests that 5 species were too few. Rheindt and Eaton (2009) reexamined the issue taking into consideration plumage, morphology, and vocalizations. Combining this information with Reddy's genetic analysis, they suggest that Pteruthius contains 9 biological species. Their treatment is followed here.

There is some uncertainty about whether Vireolanius or Cyclarhis is closer to Hylophilus and Vireo. Although it is not clearcut, the balance of the evidence is that Cyclarhis is closer (see Cicero and Johnson, 2001; Reddy, 2008). It is also not entirely clear that Hylophilus and Vireo form separate clades (Cicero and Johnson, 2001).

Neosittoidea

Neosittidae: Sittellas Ridgway, 1904

1 genus, 3 species HBW-12

One common thread in many analyses of the Corvida is that the sittellas, which wee formerly considered whistlers, end up on a branch by themselves. That's still true. Jønsson (2008b) had the branch occur after the Corvoidea-Malaconotoidea split. Barker et al. (2004) considered them more basal in the Corvida. But they still end up on their own branch.

Malaconotoidea

The last two groups are big ones—the Malaconotoidea and the Corvoidea. Many of the shrike-like birds—helmet-shrikes, bush-shrikes, wood-shrikes, vanga-shrikes, butcherbirds—are included in Malaconotoidea, but the true shrikes are in Corvoidea.

The various analyses of the Malaconotoidea have not given consistent results, sometimes leading to a more expansive Malaconotoidea including the orioles, whistlers, and cuckooshrikes. The arrangement here is based on Jønsson et al. (2011b). Interestingly, the resulting division into Malaconotoidea and Corvoidea is the same as found by Cracraft et al., 2004.

Artamidae: Woodswallows, Butcherbirds Hartlaub, 1877

6 genera, 26 species HBW-14

Artamidae tree I now follow Christidis and Boles (2008) by including the woodswallows, butcherbirds, and currawongs in the same family. There is an increasing body of genetic evidence that they form a clade. Baker et al. (2004) and Moyle et al. (2004b) placed them in the same clade, while Norman et al. (2009a) added the boatbills as a basal branch. This arrangement is also supported by Jønsson et al. (2011b). The extra structure is handled here by ranking the boatbills (Machaerirhynchinae), woodswallows (Artaminae), and butcherbirds and currawongs (Cracticinae) as subfamilies. Peltops seems to be closer to the butcherbirds than the woodswallows (Jønsson et al., 2010c, 2011b), so it is included in Cracticinae.

Machaerirhynchinae: Boatbills Schodde & Mason, 1999

Artaminae: Woodswallows Hartlaub, 1877

Cracticinae: Butcherbirds & allies Chenu & des Murs, 1853 (1836)

Pityriaseidae: Bristlehead Mayr & Amadon, 1951

1 genus, 1 species HBW-14

Moyle et al. (2006b) found that the Bornean Bristlehead and Ioras together are sister to the bush-shrikes (Malaconotidae), with the whole shebang sister to our enlarged vanga family. This is also consistent with Barker et al. (2004), which doesn't examine the Bristlehead. If the genetic evidence were a little stronger, I would probably merge the Pityriaseidae into the Aegithinidae.

Incertae Sedis: Mottled Whistler

1 genus, 1 species HBW-12, in Pachycephalidae

This New Guinea endemic seems to be close to the ioras (Jønsson et al. 2011b). Norman et al. (2009a) placed Rhagologus near Batis, but didn't include the ioras in their analysis. It makes a bit more sense to put it near here as this New Guinean endemic woud a bit out of place among the African Platysteiridae.

Aegithinidae: Ioras G.R. Gray, 1869

1 genus, 4 species HBW-10

The ioras range from India to Borneo.

Platysteiridae: Wattle-eyes, Batises Sundevall, 1872

3 genera, 30 species HBW-11

Platysteiridae tree For version 2.01, I've followed the recommendation of Njabo et al. (2008) and merged Dyaphorophyia into Platysteira. They found that those two genera are more closely related to each other than to Lanioturdus. The order here reflects that.

Vangidae: Vangas Swainson, 1831

21 genera, 39 species HBW-14

Vangidae tree Both IOC 2.11 and Roberson's Family List 11 have a more traditional treatment of this group. They both use three families: Prionopidae (Helmetshrikes), Tephrodornithdae (Woodshrikes and allies), and Vangidae. The IOC list also places Megabyas and Bias in Platysteiridae, while Roberson includes them in Prionopidae in version 11 of his family list. Previously, I described traditional arrangements as “hard to justify on molecular grounds”. But now, thanks to Reddy et al. (2012), I can say it's just wrong. Their multi-gene analysis provides strong evidence that the Vangidae, as constituted here, are monophyletic. While there remains a possibility that another vanga or two may be hiding in some other family, Reddy et al. covered all the known suspects. The current arrangement of genera is based on their results.

Previously, the papers by Yamagishi et al. (2001), Fuchs et al. (2004, 2006b, 2007a), Moyle et al. (2006b), and Johansson et al. (2008a) help clarify the relation between the vangas (Vangidae), the wattle-eyes and batises (Platysteiridae), and the bush-shrikes (Malaconotidae).

Some genera have moved around, mostly in or out of the vangas. Even compared to the recent treatment of Dickinson et al. (2003), the vangas gained Prionops from Malaconotidae, Megabyas and Bias from Platysteiridae, Hemipus from Campephagidae, and the uncertainly placed Tephrodornis and Philentoma. Stepping back a couple of years: Tylas was sometimes considered a bulbul (although correctly identified as a vanga by Beecher, 1953), Newtonia was thought to be a Sylviioid or Muscicapoid, and Hypositta was considered a Parid or Sittid. For those interested in examing an old taxonomy of the Passerida, I recommend taking a look at the diagram on page 324 of Beecher. Johansson et al. (2008a) show convincingly that Mystacornis is a vangid (Crossley's Vanga, formerly Crossley's Babbler).

Malaconotidae: Bush-shrikes, Puffbacks Swainson, 1824

8 genera, 50 species HBW-14

Laniarius has been reordered using Nguembock et al. (2008c). Based on their work, the Black Boubou, Laniarius nigerrimus (erlangeri is a junior synonym), and East Coast Boubou, Laniarius sublacteus, have been split from Tropical Boubou, Laniarius aethiopicus. They also found that the Bulo Burti Boubou, Laniarius liberatus, was a color morph of the Black Boubou, Laniarius nigerrimus. Finally, Tropical Boubou, Laniarius aethiopicus, is split into Tropical Boubou, Laniarius major, and Ethiopian Boubou, Laniarius aethiopicus. I would expect more changes for this genus in the future.

Following the recommendations of Fuchs et al. (2004), Rhodophoneus has been submerged in Telophorus, as have the undergrowth species of Chlorophoneus (dohertyi, viridis, and quadricolor).

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