Sylvioidea II

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Bulbuls and Cisticolas

These two clades appear to be independent lineages in the core Sylvioidea. I had formerly grouped them with the Pnoepygidae, but they have been moved to the Acrocephalus group based on Alström et al. (2011a) and Irestedt et al. (2011).

Pycnonotidae: Bulbuls G.R. Gray, 1840

28 genera, 150 species HBW-10

Pycnonotidae tree The bulbul family Pycnonotidae has seen some change in composition, but the biggest change has been an internal reorganization.

Genera formerly considered bulbuls include Nicator, now in its own family, Nicatoridae; Bernieria and Xanthomixis, now in Bernieridae; the vanga Tylas; and the sugarbird Arcanator. Finally, Lioptilus belongs in Sylviidae. There had been some question whether Neolestes was a bulbul or not. Zuccon and Ericson (2010b) showed that it is a bulbul.

Pasquet et al. (2001) found two clades, the African greenbuls and the mostly Asian bulbuls. They also argued in favor of restructing some genera. The more recent papers by Moyle and Marks (2006) and Johansson et al. (2007b) sampled a large fraction of the species and permitted a comprehensive phyogenetic reorganization of the family. That said, the coverage of Pycnonotinae was less satsifactory. This has been partially remedied by Oliveros (2009) and Oliveros and Moyle (2010), which add more data on Philippine bulbuls to the Moyle and Marks (2006) data set. Zuccon and Ericson (2010b) included additional genetic data in their analysis.

Several genera have been reorganized. The formerly large genus Andropadus has been split into several 4 genera, including Stelgidillas, Eurillas, and Arizelocichla. Atimastillas is split from Chlorocichla. The Red-tailed Leaf-love (formerly Pyrrhurus scandens) is included in Phyllastrephus. Criniger has been split into the African Criniger and Asian Alophoixus.

It's still not clear whether Pycnonotus is monophyletic (see HBW-10). However, it's getting closer. For some time, Pycnonotus has been overlumped. I've placed several former Pycnonotus species in Microtarsus, which appears as a basal group in Pycnonotinae (Moyle and Marks, 2006; Oliveros, 2009; Oliveros and Moyle, 2010). I had previously split part of Pycnonotus into Euptilosus, and Brachypodius, but it now seems clear they are better merged into Microtarsus. Finally, several species often placed in Pycnonotus are separated in the restored genus Rubigula.

I've also moved the Philippine members of Ixos into Hypsipetes, which has also absorbed Microscelis. The results of Oliveros (2009) and Oliveros and Moyle (2010) also allow an alternative treatment, placing the former Ixos in a separate genus. The Indian Ocean/Malagasy Hypsipetes have been rearranged in view of Warren et al. (2005). Further, the Moheli Bulbul, Hypsipetes moheliensis, has been split from the Comoros Bulbul (now Grand Comoro Bulbul), Hypsipetes parvirostris.

I have included the Bare-faced Bulbul, Pycnonotus hualon, described by Woxvold et al. (2009). Where exactly this goes in Pycnonotus is unclear, but after reading Woxvold et al. carefully, this is my best guess. As the authors note, Pycnonotus itself needs revision.

Oliveros (2009) and Oliveros and Moyle (2010) note several subspecies that appear to deserve species rank. Two species contain two or more separate clades. The IOC has provided English names for Ashy-fronted Bulbul, Pycnonotus cinereifrons (split from Olive-winged Bulbul, Pycnonotus plumosus), Palawan Bulbul, Alophoixus frater (split from Gray-cheeked Bulbul, Alophoixus bres), Visayan Bulbul, Hypsipetes guimarasensis, and Mindoro Bulbul, Hypsipetes mindorensis (both split from Philippine Bulbul, Hypsipetes philippinus). In all 4 cases, I'm presuming that each of the new species are monotypic. Further, there are indications that Pycnonotus urostictus atricaudatus, and one or more of the subspecies of Pycnonotus goiavier and Hypsipetes amaurotis should be considered distinct species, but these cases involve additional complexities.

The green-headed Montane Tiny-Greenbul, Phyllastrephus albigula, has been split from the gray-headed Lowland-Tiny Greenbul, Phyllastrephus debilis. See Fuchs et al. (2011a).

Crinigerinae: Greenbuls Bonaparte, 1854 (1831)

Pycnonotinae: Bulbuls G.R. Gray, 1840

Cisticolidae: Cisticolas Sundevall, 1872

27 genera, 156 species HBW-11

Cisticolidae tree The cisticola family has grown since Sibley and Ahquist (1990) separated it from the Sylviidae. Gains include Eremomela, Neomixis (except flavoviridis, now in Hartertula), Phyllolais. Poliolais and most of Orthotomus (except two species). Rhopophilus has been lost to the Sylviidae. Following Nguembock et al. (2008a), I've moved the Black-collared Apalis and Ruwenzori Apalis to Oreolais, which is sister to Artisornis. Like the Artisornis, the Oreolais have 10 rather than 12 rectrices. Also, Mrs. Moreau's Warbler and the recently split Rubeho Warbler are now listed in the genus Scepomycter.

The arrangment of the genera is based on Alström et al. (2006), Johansson et al. (2008b), and Nguembock et al. (2007, 2008a).

Hyliidae, Aegithalidae, Cettiidae, and Phylloscopidae

Hyliidae: Hylias Bannerman, 1923

2 genera, 2 species Not HBW Family

I've put the Hylias in a separate family based on Johanssen et al. (2008), although I'm not entirely confident about this. Classification of the Tit-Hylia (Pholidornis rushiae) has long been controversial. It has variously been placed in at least 7 other families: Sylviidae, Estrildidae, Dicaeidae, Nectariniidae, Remizidae, Hyliidae and Meliphagidae.

Aegithalidae: Long-tailed Tits Reichenbach, 1849-50

4 genera, 13 species HBW-13

Click for Aegithalidae tree
Click for Aegithalidae tree

Leptopoecile has been included in Aegithalidae rather than Sylviidae based on Sturmbauer et al. (1998), Alström et al. (2006), Johansson et al. (2008b), and Päckert et al. (2010).

The current arrangement of Aegithalos is based on the results of Päckert et al. (2010). However, they found that the last four species A. bonvaloti through A. sharpei are extremely close genetically, close enough to call into serious question whether they are separate species. Nonetheless, they appear distinct and, according to HBW-13, there is no evidence of interbreeding in areas of overlap. Päckert et al. also found quite large genetic differences between the various races of A. concinnus, suggesting several species are involved. This had previously been suspected based on plumage, but the presence of zones of intergradation has kept them lumped together. Unfortunately, Päckert et al. did not examine the entire concinnus complex, so the situation is not entirely clear, even without the complication of intergrades. Until more information is available, I leave the species limits unchanged.

Cettiidae: Cettid Warblers Alström et al., 2006

12 genera, 36 species Not HBW Family

Click for Cettiidae tree
Click for Cettiidae tree

We briefly moved back to familiar territory family-wise, with the swallows and then the long-tailed tits. The long-tailed tits seem to be the sister group of another new family, the cettid bush-warblers (Cettiidae). This group has an primarily African and Asian distribution

Irestedt et al. (2011) found that Neumann's Warbler, Hemitesia neumanni, is a member of the Cettiidae. It is most likely closest to the Urosphena stubtails, although the relationship is somewhat distant. Given that most of the Cettiidae are oriental, it's interesting that the African genus ended up among them. Irestedt et al. (2011) also provided stronger support for the basal position of Erythrocercus previously noted by Johansson et al. (2008b). Since Erythrocercus is also African, this opens the possibility that the Cettiidae may have originated in Africa.

There have been questions about which family Scotocerca inquieta belongs to. It has often been placed with the cisticolas, but Barhoum and Burns (2002) presented evidence suggesting that Scotocerca is part of Acrocephalidae, not Cisticolidae. However, there were issues about the sample used, and the result did not appear robust. Alström et al. (2011a) have carried out a more comprehensive analysis and found that it is actually a cettid warbler. They found it sister to a clade containing Cettia and Abroscopus. This is reflected in the tree shown. However, the sampling of the cettids was rather sparse and there may still be a residual question about its exact position within the Cettiidae.

The Phyllergates tailorbirds were formerly placed among the cisticolas in the genus Orthotomus (see Alström et al., 2006, 2011d; Fuchs et al., 2006a). The Odedi Bush-Warbler is a recent discovery (LeCroy and Barker, 2006). Its song had been heard by Jared Diamond in 1972, but the bird was not tracked down until recently. Hadden was able to mist-net the bird in 2000.

The arrangement of genera is now based on Alström et al. (2011d). This is generally consistent with my previous treatment based on Alström et al. (2006), Fuchs et al. (2006a), Johansson et al. (2008b), and Irestedt et al. (2011).

The genus Cettia as usually constituted is polyphyletic. Although my original guess was that it involved two different groups, Alström et al. (2011d) found that the situation was more complex. Accordingly, I've transferred the Pale-footed Bush-Warbler to Hemitesia, Gray-sided Bush-Warbler to Oligura, and temporarily designated Chestnut-crowned Bush-Warbler as “Cettia”. Only Cetti's Warbler is left in Cettia (as in the previous TiF version). The other former Cettia warblers are placed in Horornis (Hodgson 1845, type fortipes). I had previously put them in Horeites in spite of some uncertainty about whether the type species, brunnifrons, actually belonged with them. Well, it doesn't.

Alström et al. (2011d) suggest a somewhat different set of generic limits, with Hemitesia merged into Urosphena, and with Cettia including “Cettia” and Oligura. I decided to not follow that because (1) the genera I use seem to mark relatively deep divisions, and (2) there is some uncertainty about whether such a broad Cettia would also have to include Tesia (there's a reason I have a 4-fold polytomy there; see Fig. 3 and the supplementary material in Alström et al., 2011d).

Finally, the Manchurian Bush-Warbler is restricted to the race borealis, with canturians being demoted to a subspecies of Japanese Bush-Warbler, Horornis diphone. Genetically, borealis is closer to the Philippine Bush-Warbler than to the Japanese Bush-Warbler races cantans canturians, justifying treatment as a separate species. As borealis breeds in Manchuria, while canturians does not, I've retained the name Manchurian Bush-Warbler. Since canturians is quite close to cantans, and presumably to diphone, it is treated as a subspecies of the Japanese Bush-Warbler. That said, diphone itself has not been studied, and there are differences (e.g., song) that suggest canturians may be a separate biological species from cantans and diphone. A more comprehensive look at this complex would be helpful (including restricta).

Phylloscopidae: Leaf-Warblers Jerdon, 1863 (1854)

2 genera, 79 species Not HBW Family

These are followed by the leaf-warblers (Phylloscopidae), which is a new family consisting of about 70 species in two genera formerly belonging to the Sylviidae. The taxonomy here is primarily based on Johansson et al. (2007a) and Olsson et al. (2005), with Irwin et al. (2005) and Martens et al. (2008) filling in some details. The resulting changes moved many species from Phylloscopus to Seicercus. Besides the division into the new Phylloscopus and Seicercus, both papers agree on several clades within Seicercus. These are (1) S. umbrovirens to S. soror (labelled Seicercus on the tree); (2) S. castaniceps, montis, and grammiceps; (3) S. cebuensis to S. ijimae; (4) S. borealis through S. plumbeitarsus; and (5) S. cantator to the end. They also agree that clades 4 and 5 are sisters. Olsson et al. (2004) has yet another topology, but the same clades appear, including the basic split into the new Phylloscopus and Seicercus. Previously, clades 3-5, and part of 1 were considered Phylloscopus.

I've split Japanese Leaf-Warbler, Seicercus xanthodryas, and Kamchatka Leaf-Warbler, Seicercus examinandus, from Arctic Warbler, Seicercus borealis. Reeves et al. (2008) found substantial genetic differences between the Kamchatka/Sakhalin populations and borealis/kennicotti. They suggested separating them as “Pacific Warbler”. However, Saitoh et al. (2010) also included Japanese populations of in their analysis. They found that the Hokkaido birds grouped with those on Sakhalin Island and in Kamchatcka, but that the other Japanese populations were well-separated from the rest. Additional races are sometimes recognized, but they appear to belong to the borealis/kennicotti clade (Alström et al., 2011c). The English names are those of Alström et al. (2011c), which contains further information on the split. Note that there is a record of Kamchatka Leaf-Warbler from Amchitka Island, Alaska (Kenyon, 1961).

I've handled the Seicercus trochiloides complex in an unusual way. This includes nitidus, viridanus, trochiloides, obscuratus, plumbeitarsus and S. v. ludlowli (or at least part of it). Irwin et al. (2001) found two groups. The first comprised of nitidus and viridanus, the second including trochiloides, obscuratus, and plumbeitarsus in a trichotomy. Although ludlowli had its own clade, it was too closely related to viridis to separate. This gives us several choices: lump them all, split the first group (which are more distantly related) and lump the second, or split them all. I take the last option here.

In contrast, BLI and Clements lump them all, which is defensible. Sibley and Monroe, Dickinson, IOC, and HBW take a different route. They split trochiloides and plumbeitarsus. This is not consistent with Irwin et al.'s results. It conflicts with the status of obscuratus as an equal member of the trichotomy. It also includes taxa from both groups in trochiloides.

For information concerning the newly recognized Limestone Leaf-Warbler, Seicercus calciatilis, see Alström et al. (2010).

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