Corvida

Mohouoidea

Orioloidea

Campephagoidea

Malaconotoidea

Corvoidea

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 46 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, comprising 800+ species. Since there are many more species in Passerida, so we place Corvida 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.

Current thinking is that the Corvida have their origin in Australia/New Guinea (the actual continent of Australia). In particular, their diversity may have been developed during the late Eocene/Oligocene in the New Guinea archipelago, and then certain taxa pumped out into Asia and the rest of the world. I particularly commend to your attention the discussion in Jønsson et al. (2011b) and secondarily in Aggerbeck et al. (2014).

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 late 2013 I restructured the Corvida based on Aggerbeck et al. (2014). They examined 22 nuclear loci, and were able to create a generally well-supported phylogeny for Corvida. Previously, the most comprehensive study of the Corvida was the six-gene analysis by Jønsson et al. (2011b), which includes one of more representative from each of the corvid families except the monotypic Pityriaseidae (bristlehead). Most of the well-supported portions of Jønsson et al.'s phylogeny are pretty much the same as Aggerbeck et al. However, Aggerbeck et al. were able to find good support for much more of their phylogeny.

Corvida tree

Both Aggerbeck et al. and Jønsson et al. include Callaeidae, Cnemophilidae, and Melanocharitidae in Passerida, although there is some disagreement about the exact ordering of taxa.

Aggerbeck et al. describe their results as a basal group (Mohouidae) plus three major clades. One of those clades includes a fairly deep division involving Campephagidae. I'm interpreting this a I'm treating the whole thing as five main clades: Mohouoidea, Orioloidea, Campephagoidea, Malaconotoidea, and Corvoidea. Each clade seems to be well-supported. The last two are essentially the same as in Cracraft et al. (2004).

Mohouoidea

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) and HBW include them in Pachycephalidae. However, the evidence for any of these is very weak.

Jønsson et al. (2011b) was the first genetic study to suggest any real affinities for them (sister to Campephagidae), although the support for this was weak. Although Jønsson et al. were unable to place Mohouidae reliably, Aggerbeck et al. (2014) found strong support for Mohouidae as the basal clade in Corvida.

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). Regardless of whether they are placed in one genus or two, the Pipipi is basal, and the Whitehead and Yellowhead are sister to one another (Aidala et al., 2013).

Pachycephaloidea

Pachycephaloidea is our first large grouping in Corvida. Pachycephaloidea includes the 3 small groups: the bellbird, shriketit and quail-thrush families (Oreoicidae, Falcunculidae, and Cinclosomatidae) as well as the bigger whistler family (Pachycephalidae).

Click for Pachycephaloidea tree
Click for Pachycephaloidea
species tree

The Pachycephaloidea range from south Asia, through Wallacea, New Guinea, and Australia, to New Zealand and the South Pacific.

Genetic studies have done a number on the family. Although the core of it, the whistlers and shrike-thrushes, has remained unscathed, other taxa associated with the family have moved in or out. The pitohuis have been split into 4 genera, with Pitohui itself moving to Oriolidae. The others are all currently gathered here in Pachycephalidae, although there has been question about whether they all belong.

The analysis of Norman et al. (2009a) had Eulacestoma basal in Pachycephaloidea, but weakly supported, with Falcunculus branching off next. However, Jønsson et al. (2011b) had Eulacestoma weakly associated with Psophodidae. Well, Aggerbeck et al. (2014) have booted Eulacestoma out. It now is in its own monotypic family in the Orioloidea. The shriketit (Falcunculidae: Falcunculus) remains here, and has been joined by the Cinclosoma quail-thrushes and Ptilorrhoa jewel-babblers (from Psophodidae, now treated as Cinclosomatidae).

The Falcunculidae/Cinclosomatidae clade seems to be sister to a small Australo-Papuan group, Oreoicidae (Oreoica, Ornorectes, and Aleadryas) that Jønsson et al. (2011b) placed on a separate branch in Corvida. Aggerbeck et al. (2014) included one of these species, and it ended up here. In contrast, Norman et al. (2009a) weakly supported placing Aleadryas and Oreoica in Malaconotoidea. In contrast, Jønsson et al. (2008a) put Aleadryas and the former Pitohui, Ornorectes, near Campehagidae. The Aggerbeck solution seems well-supported, even though only Oreoica was included, and I'm using that here.

I've separated Oreoicidae, Falcunculidae, and Cinclosomatidae in separate families. The divisions between them and Pachycelphidae are deep, dating back to the Oligocene (Aggerbeck et al., 2014). Further, while the Oreocidae, and even Falcunculidae could be interprested as variant whistlers, the quail-thrushes and jewel-babblers cannot. That gives us four families.

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 not even in Corvida. It belongs somewhere in Passerida.

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 (Oreoicidae), Pitohui (Oriolidae), Pseudorectes (Pachycephalidae), and Melanorectes (Pachycephalidae). All but Pitohui are in Pachycephaloidea.

Oreoicidae: Australo-Papuan Bellbirds Sibley and Ahlquist, 1985

3 genera, 3 species

I accept the name Oreoicidae (Sibley and Ahlquist, 1985a) over Oreoicidae (Schodde and Christidis, 2014). Schodde and Christidis object that there is not a proper definition or description of Oreoicini Sibley and Ahlquist. However, they treat it as monotypic, and I think that defining the tribe Oreoicini as consisting of Crested Bellbird is sufficient definition.

Falcunculidae: Shriketits Chenu and des Murs, 1853

1 genus, 3 species

Cinclosomatidae: Quail-thrushes and Jewel-babblers Matthews, 1921-2

2 genera, 11 species

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).

Pachycephalidae: Whistlers Swainson, 1831

5 genera, 59 species HBW-12

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 fact that the Morningbird is not in Colluricincla causes a change in the scientific name of the Sooty Shrike-thrush. With Colluricincla tenebrosa of Hartlaub and Finsch (1868) no longer using tenebrosa, the Sooty Shrike-thrush retakes the name tenebrosa (Rothschild 1911) which has priority over umbrina (Reichenow 1915). 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.

The current arrangement of the whistlers is based on Jønsson et al. (2008c, 2009, 2014) and Andersen et al. (2014b). I have not strictly followed the phylogeny in Anderseen et al., but have kept several lumped together several closely related taxa that may their phylogeny splits. Even so, I recognize 8 additional species:

  1. Bougainville Whistler, Pachycephala richardsi, split from Hooded Whistler, Pachycephala implicata.
  2. Louisiade Whistler, Pachycephala collaris, split from Bismarck Whistler, Pachycephala citreogaster.
  3. Rossell Whistler, Pachycephala rosseliana, split from Bismarck Whistler, Pachycephala citreogaster.
  4. Rennell Whistler, Pachycephala feminina, split from Oriole Whistler, Pachycephala orioloides.
  5. Santa Cruz Whistler, Pachycephala vanikorensis, comprised of subspecies utupuae and ornata from the White-throated Whistler, Pachycephala vitiensis, and vanikorensis from the Melanesian Whistler, Pachycephala caledonica.
  6. Timor Whistler, Pachycephala calliope, split from Yellow-throated Whistler, Pachycephala macrorhyncha
  7. Wetar Whistler, Pachycephala arthuri, split from Yellow-throated Whistler, Pachycephala macrorhyncha. Note that arthuri is often synonymized with calliope.
  8. Babar Whistler, Pachycephala sharpei, split from Yellow-throated Whistler, Pachycephala macrorhyncha.

Further, the remainging (i.e. Fijian) races of the White-throated Whistler, Pachycephala vitiensis, are lumped into the Fiji Whistler, Pachycephala graeffii. The name vitiensis has priority, so the Fiji Whistler is now Pachycephala vitiensis.

Several whistler subspecies also move around. The Yellow-throated Whistler, Pachycephala macrorhyncha, loses dammeriana which joins the Mangrove Golden Whistler, Pachycephala melanura. On the other hand, the Yellow Throated Whistler gains balim from the Australian Golden Whistler, Pachycephala pectoralis. The Mangrove Golden Whistler, Pachycephala melanura loses dahli, which joins the Golden-backed Whistler, Pachycephala aurea, which becomes Pachycephala dahli due to priority.

Both the Melanesian Whistler, Pachycephala chlorura, and Oriole Whistler, Pachycephala orioloides may require further splitting in the future.

Orioloidea

Orioloidea is the next large grouping in Corvida. Orioloidea include the oriole, and vireo families (Oriolidae and Vireonidae) as well as several small groups (Eulacestomidae, Neosittidae, Paramythiinae, and Psophodidae).

Eulacestomidae: Ploughbill Schodde and Christidis, 2014

1 genus, 1 species Not HBW Family

The Ploughbill is a New Guinea endemic. Its closest relatives are the sittellas. They seem different enough to justify treating them as separate families.

Neosittidae: Sittellas Ridgway, 1904

1 genus, 3 species HBW-12

The sittellas are nuthatch-like birds of Australia and New Guinea. Indeed, they were considered nuthatches as late as 1967. They aren't nuthatches, and aren't even members of Passerida. Like many Australasian birds, they are part of Corvida.

One common thread in most recent analyses of the Corvida is that the sittellas, which were formerly considered whistlers, end up on a branch by themselves. 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. Aggerbeck et al. (2014) have them sister to Eulacestoma, but even they find support for this to be somewhat weak.

Oriolidae: Orioles, Figbirds Vigors, 1825

4 genera, 40 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 Oriolus orioles are widespread in the Old World, including Australasia. The other Oriolidae are confined to Australo-Papua (Pitohui, Sphecotheres) or New Zealand (Turnagra).

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.

Based on Dumbacher and Fleischer (2001), the Variable Pitohui has been split into three species: Southern Variable Pitohui, Pitohui uropygialis, Raja Ampat Pitohui, Pitohui cerviniventris, and Northern Variable Pitohui, Pitohui kirhocephalus.

The genetic results in Jønsson et al. (2010) also support splitting Sunda Golden-Oriole, Oriolus maculatus, and Asian Golden-Oriole, Oriolus diffusus, from Black-naped Oriole, Oriolus chinensis. Jønsson 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.

Paramythiidae: Painted Berrypeckers P. L. Sclater, 1893

2 genera, 2 species HBW-13

This New Guinea endemic family was previously removed from the Melanocharitidae (Passerida) and placed in their own family in Corvida. It has been unclear whether they should be treated as a separate family, and the TiF list has sometimes included them in Psophodidae. Aggerbeck et al. (2014) found them to be a separate branch sister to Psophodidae and Vireonidae.

Sibley and Ahlquist (1987) removed the painted berrypeckers from Melanocharitidae (berrypeckers and longbills), but left them in place near the flowerpeckers and sunbirds (others had previously questioned the affinities of these species). [see also Sibley and Ahlquist (1990) and Sibley and Monroe (1990)]. Nonetheless, H&M3 (Dickinson, 2003) still had them united in a single family, but now inside what I call Corvida. They were separated again in Barker et al. (2004), this time with the Paramythiidae located near Oriolus (as they are) and with the Melanocharitidae as basal Corvida near Callaeatidae. As previously mentioned, Irestedt and Ohlson (2009) recognized that Melanocharitidae, Cnemophilidae, and Callaeidae, were really basal Passerida, and that the RAG-1 gene had given a misleading signal. Aggerbeck et al.'s many gene analysis (2014) supports this interpretion.

Psophodidae: Whipbirds & Wedgebills Bonaparte, 1854

2 genera, 6 species HBW-12, as Eupetidae

This small group of Australo-Papuan birds has been substantially reduced in size. I have previously treated these taxa and other possibily related taxa such as Paramythiidae in various ways (e.g., based on Norman et al., 2009a). The current version is based on Aggerbeck et al. (2014) and leaves a very narrowly circumscribed Psophodidae that is sister to Vireonidae.

Other birds that have previously been considered to belong in this family include Cinclosoma and Ptilorrhoa (currently in Cinclosomatidae) and Eulacestoma, which is now in its own family. There is also some support for including Falcunculus (Falcunculidae), in Psophodidae (e.g, Barker et al., 2004; Dumbacher, 2008), but the multigene analyses of Aggerbeck et al. (2014), Norman et al. (2009a) and Jønsson et al. (2011b) all separate them.

Sibley and Monroe's Cinclosomatinae included two additional 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. Jønsson et al. (2011b) put Ifrita in the monarchs and Melampitta in a clade containing the mudnesters and birds-of-paradise. Aggerbeck et al. (2014) have the mudnesters, Ifrita and Melampitta in a grade with the birds-of-paradise. In any event, they do not belong here.

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 genus Androphobus is sometimes merged into Psophodes.

Vireonidae: Vireos Swainson, 1837

8 genera, 66 species HBW-15

Click for Vireonidae tree
Click for Vireonidae
species 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. It is true that it seemed odd. But it is false that there are no old world members.

The old world vireos have been hiding out among the babblers (Timaliidae), living in southeast Asia and the Himalayas. The first to be noticed was Erpornis zantholeuca (nee Yuhina zantholeuca). Cibois et al. (2002) discovered it was not a babbler, but belonged somewhere in the Corvida. Rather than being a babbler like the other Yuhinas, it is actually a vireo relative (Barker et al., 2004).

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, Erpornis and Pteruthius, they seem less special. Thus I take the former route for the present, ranking them as subfamilies and keeping in mind that the discovery of additional hidden vireo taxa might change things. The arrangement of the known vireos is based on Slager et al. (2014).

Pteruthiinae: Shrike-babblers Informal?

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.

Erporninae: Erpornis Informal?

Vireoninae: Vireos, Greenlets and allies Swainson, 1837

The current arrangment of the Vireoninae (what was traditionally considered the vireo family) is based on Slager et al. (2014) (see also the preliminary results posted by Battey, 2014). They sampled most of the taxa. The conventional Hylophilus turned out to be paraphyletic. To solve this, one species moves to Vireo (Tepui Greenlet, now Vireo sclateri) and the remaining Hylophilus are split into three genera: Hylophilus (scrub greenlets), Tunchiornis (tawny-crowned greenlets, Slager and Klicka, 2014b), and Pachysylvia (canopy greenlets, Bonaparte 1850, type decurtata). Vireo remains unchanged except for the addition of the Tepui Greenlet (make that Tepui Vireo). The concatenated analysis placed the scrub greenlets (Hylophilus) sister to the peppershrikes (Cyclarhis), branching before the Shrike-Vireos (Vireolanius) which are basal to the remaining taxa.

It's unclear whether the two peppershrikes are reciprocally monophyletic. They are not in ND2, but are in the Z-linked genes studied by Slager et al. (2014). There is also a substantial amount of genetic distance within the Rufous-browed Peppershrike, Cyclarhis gujanensis, and more than one species might be involved. Further study is needed to clarify matters, especially as there is little vocal variation among the subspecies (Tubaro and Segura, 1994), with the main difference depending on climate and habitat (more high frequencies in closed habitats, and more bandwidth in the tropics).

Another potential split involves the Slaty-capped Shrike-Vireo, Vireolanius leucotis. The one sample from western Ecuador (subspecies mikettae, sampled from Lita, in Esmeraldas province) did not group with the birds east of the Andes in ND2.

The Golden Vireo (Golden Greenlet?) moves to Pachysylvia. The Red-fronted Greenlet, Tunchiornis rubrifrons, has been split from Tawny-crowned Greenlet, Tunchiornis ochraceiceps. Further research may support additional splits in the ochraceiceps complex, particularly between the Red-fronted Greenlets north and south of the Amazon.

When possible, the species tree includes available subgeneric names in italics for the various clades within Vireo. The four subgenera are spectacled vireos, Lanivireo (Baird 1858, type flavifrons); the eye-ringed vireos, Vireo (Vieillot 1808, type griseus); the eye-lined gilvus group, Melodivireo (Oberholser 1974, type gilvus); and the eye-lined olivaceus group, Vireosylva (Bonaparte, 1838, type olivaceus). And yes, that's the way Bonaparte spelled Vireosylva, so we're stuck with it.

Slager et al. (2014) found that the Central American Vireo, Vireo notius (inc. montanus), is basal in the Solitary Vireo group. It has accordingly been split from Plumbeous Vireo, Vireo plumbeus. Although genetic data is not available, the subspecies notius and montanus differ in appearance and are candidates for a further split.

Slager et al. (2014) found that Brown-capped Vireos of Mexico and northern Central America are genetically distinct from the remaining Brown-capped Vireos, Vireo leucophrys. Further research may lead to a split.

Warbling Vireo, Vireo gilvus, has been split into the monotypic Eastern Warbling-Vireo, Vireo gilvus, and Western Warbling-Vireo, Vireo swainsoni. They differ vocally, in morphometrics, molt and migration timing (Voelker and Rohwer, 1998) and the genetic distance supports their treatment as distinct species (Murray et al., 1994; Slager et al., 2014).

I have also split the South American Chivi Vireo, Vireo chivi, from Red-eyed Vireo, Vireo olivaceus. Slager et al. (2014) sampled Red-eyed Vireos from 6 locations and 18 Chivi Vireos and found that they are not sister taxa. Interestingly, the Black-whiskered Vireo, the local breeding member of the Red-eyed complex (flavoviridis through chivi) here in Miami is better thought of a a northern form of Chivi Vireo, rather than a southern form of Red-eyed. Both the Yellow-green and Yucatan Vireo developed from the Red-eyed complex prior to the Red-eyed/Chivi split.

Campephagoidea

The Campephagoidea contain a single family, Campephagidae. I've separated it from Orioloidea because the division between it and the Orioloidea families seems to be rather deep.

Campephagidae: Cuckooshrikes Vigors, 1825

11 genera, 92 species HBW-10

The Campephagidae are widespread in the southern portions of the Old World (Afrotropics, Indomalaya, Australasia), but present in only a limited area of the eastern palearctic.

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.

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. However, recent papers seem to agree on what is in, and what is not. The arrangement here is based on the multigene analyses of Aggerbeck et al. (2014) and Fuchs et al. (2012b). Interestingly, the resulting division into Malaconotoidea and Corvoidea is the same as found by Cracraft et al. (2004). It makes biogeographical sense in that the Australasian species in Malaconotoidea are grouped together in Artamide.

It's interesting to compare the osteological analysis of Manegold (2008) with Fuchs et al. (2012b), or to any of the other molecular analysis. It's something to keep in mind when looking at even the best hypothetical trees for fossil organisms.

Artamidae: Woodswallows, Butcherbirds Hartlaub, 1877

6 genera, 26 species HBW-14

This family includes the Papuan endemic Mottled Whistler, the Austral-Papuan Cracticinae, and the more widespread Artaminae (woodswallows) that range from India to Australasia.

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. The extra structure is handled by ranking the Mottled Whistler (Rhagologinae), woodswallows (Artaminae), and butcherbirds and currawongs (Cracticinae) as subfamilies. Support for any particular position of the Mottled Whistler and the boatbills is weak in all studies. The arrangement here is that of Aggerbeck et al. (2014). No one would be surprised if they changed places or ended up sisters.

Peltops seems to be closer to the butcherbirds than the woodswallows (Jønsson et al., 2010c, 2011b; Fuchs et al., 2012b, Kearns et al., 2013), so it is included in Cracticinae. Kearns et al. (2013) found that the actual species boundaries within Peltops do not seem to match current thinking. There are two species, but the division is not as expected.

The genus Gymnorhina (Australian Magpie) has been merged with Cracticus and Strepera has been reordered based on Kearns et al. (2013). The Black Butcherbird has been split into New Guinea Black-Butcherbird, Cracticus quoyi, and Australian Black-Butcherbird, Cracticus spaldingi. Kearns et al. (2011) found substantial genetic distance between these allopatric groups of taxa.

The gray (a.k.a. white-throated) Butcherbird group continues to be controversial. Kearns et al. (2013) found little genetic distance between them and relationships that did not match the traditional allopatric species in the group. Recognition of the Silver-backed Butcherbird, Cracticus argenteus, has always been controversial, but the Black-backed Butcherbird, Cracticus mentalis, has been considered a separate species. Kearns et al. (2013) found that some argenteus grouped with mentalis and some grouped with torquatus. All were genetically close, with a common ancestor probably about 200,000 years ago. Because of this I, had lumped them with the Gray Butcherbird, Cracticus torquatus. However, that is not the end of the story. Kearns et al. (2014) increased sampling of both individuals and nuclear genes. They found significant conflict between the mitochondrial and nuclear dna. However, the species tree made sense, and current geneflow is modest, so I have returned to treating them as three species.

Rhagologinae: Mottled Whistler Schodde and Christidis, 2014

Artaminae: Woodswallows Hartlaub, 1877

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

Machaerirhynchidae: Boatbills Schodde & Mason, 1999

1 genus, 2 species HBW-14

This small Australo-Papuan group seems to be sister to the remaining Malaconotoidea.

Aegithinidae: Ioras G.R. Gray, 1869

1 genus, 4 species HBW-10

The ioras range from India to Borneo.

Pityriaseidae: Bristlehead Mayr & Amadon, 1951

1 genus, 1 species HBW-14

Moyle et al. (2006b) was the first to find that the Bornean Bristlehead belonged to the Malaconotoidea. This arrangement with the Bristlehead sister to the Malaconotidae family is based on Fuchs et al. (2012b) and also supported by Aggerbeck et al. (2014).

Malaconotidae: Bush-shrikes, Puffbacks Swainson, 1824

8 genera, 50 species HBW-14

The overall structure of Malaconotidae follows Fuchs et al. (2012b). 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).

Platysteiridae: Wattle-eyes, Batises Sundevall, 1872

4 genera, 31 species HBW-11

Platysteiridae tree I had earlier followed the recommendation of Njabo et al. (2008) and merged Dyaphorophyia into Platysteira. However, further analysis by Fuchs et al. (2012b) revealed that things are not quite so simple. Their results call into question whether Batis itself is monophyletic. They found four deep clades separated by very short internodes. Two parts of Batis, the restricted Dyaphorophyia used here, and Platysteira (which includes part of the former Dyaphorophyia). Although their species tree says Batis is not monophyletic, I find that hard to believe and leave Batis as a single genus.

Njabo et al. (2008) found that the West African Wattle-eye, Platysteira hormophora, formerly considered a subspecies of Chestnut Wattle-eye, Platysteira castanea, is only distantly related to it.

Vangidae: Vangas Swainson, 1831

21 genera, 39 species HBW-14

Vangidae tree IOC 3.3 follows a more traditional treatment of this group. They use three families: Prionopidae (Helmetshrikes), Tephrodornithdae (Woodshrikes and allies), and Vangidae. The IOC list also places Megabyas and Bias in Platysteiridae. Previously, I described traditional arrangements as “hard to justify on molecular grounds”. But now, thanks to Reddy et al. (2012), Jønsson et al. (2012), and Fuchs et al. (2012b), 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, all the known suspects have been tested. The current arrangement of genera is based on Reddy et al. (2012) and the arrangement within some of the genera follows Jønsson et al. (2012).

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).

Prionopinae: Helmetshrikes Bonaparte, 1853

Tephrodornithinae: Woodshrikes, Shrike-flycatchers Informal

Some internet sources claim that Tephrodornithidae is due to Moyle et al. (2006b). Although they did notice the clade, they did not use the term Tephrodornithidae, and it remains an informal name.

Philentominae: Philentomas Informal

Although the Philentomas could be included in Vanginae, it makes sense to separate these southeast Asian species from the Madagascan vangas.

Vanginae: Vangas Swainson, 1831

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