Tyranni: Suboscines

Passeri: Oscines


Muscicapoidea and allies

The 46 Orders














Swallows, Bulbuls, and the rest

The swallows and bulbuls seem to be independent lineages, while the remaining Sylvioidea species are a bit more closely related to each other (Fregin et al., 2012).

Hirundinidae: Martins, Swallows Rafinesque, 1815

18 genera, 88 species HBW-9

The organization of the swallows is based on Sheldon et al. (2005), with Whittingham et al. (2002) and Dor et al. (2010) providing additional information. The arrangement within Tachycineta is based on Cerasale et al.'s (2012) analysis using the complete mitochondrial genome. Dor et al. (2012) give three slightly different arrangements based on 16 nuclear genes, two of which differ from Cerasale et al. only in the position of the Tree Swallow. I did not find the BEAST tree in Dor et al. to be all that convincing.

I've inserted subfamiles and tribes to make the organization a bit clearer. There is a basal division between the river martins (Pseudochelidoninae) and everything else (Hirundinae). Hirundinae divides into three groups, with the Psalidoprocnini likely closer to the Hirundinini. The tree shows this in detail:

Hirundinidae tree

The Rough-winged Swallows (genus Stelgidopteryx) are treated here as two species, but there is substantial genetic distance within both species (Babin, 2005), and they may be better treated as several species. Although "Ridgway's Swallow", Stelgidopteryx ridgwayi, is relatively distant from the Northern Rough-winged Swallow subspecies serripennis and fulvipennis, this distance is within the range of both the Southern Rough-winged Swallow and the remaining races of Northern Rough-winged Swallow. Further research, including more comprehensive genetic testing of the various races and at more locations, will be necessary to sort this out.

The Cave Swallow, Petrochelidon fulva, which includes two geographically-based distinct color morphs, is sometimes considered a candidate for a split. However, Kirchman et al. (2000) found substantial gene flow between the various populations. It was also striking how genetically distinct the Florida population had become in just a few years.

Pseudochelidoninae: River Martins Shelley, 1896

Hirundininae Rafinesque, 1815

Prognini Cassin, 1853

Psalidoprocnini Sharpe, 1870

Hirundinini Rafinesque, 1815

Pycnonotidae: Bulbuls G.R. Gray, 1840

28 genera, 153 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 dapplethroat 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 satisfactory. 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.

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).

Based on Manawatthana et al. (2017), Charlotte's Bulbul, Iole charlottae (inc. perplexa) has been split from Buff-vented Bulbul, Iole olivacea, and Cachar Bulbul, Iole cacharensis, has been split from Olive Bulbul, Iole virescens. Moreover, the subspecies lekhakuni and cinnamonomeoventris have been transferred from Gray-eyed Bulbul, Iole propinqua to Olive Bulbul, Iole virescens while myitkyinensis has moved the other way, from Olive Bulbul to Gray-eyed Bulbul.

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.

Aceh Bulbul, Pycnonotus snouckaerti, has been split from Orange-spotted Bulbul, Pycnonotus bimaculatus, based on Eaton & Collar (2015).

Crinigerinae: Greenbuls Bonaparte, 1854 (1831)

Pycnonotinae: Bulbuls G.R. Gray, 1840

Hyliidae, Aegithalidae, Cettiidae, and Phylloscopidae

These four families are fairly closely related.

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. Fregin et al. (2012) place them sister to the long-tailed tits, albeit with less support than one might like. If they really are sister to the Long-tailed Tits, it would probably make a lot of sense to merge the two families.

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. Sefc et al. (2003) found Hylia and Pholidornis more closely related than either is to Aegithalos or Phylloscopus.

Aegithalidae: Long-tailed Tits Reichenbach, 1849-50

3 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.

Johansson et al. (2016) found that Psaltria is embedded in the concinnus complex (which still needs to be sorted out). I have merged Psaltria into Aegithalos.

Cettiidae: Cettiid Warblers Coues, 1903

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 cettiid bush-warblers (Cettiidae). This group has an primarily south and east Asian distribution ranging into Oceania, the Western Palearctic, and tropical Africa. Although IOC and Clements use the term “Bush-Warblers and allies” for the family, I think it's a problem because there are nearly as many “Bush-Warblers” in Locustellidae as in Cettiidae.

Fregin et al. (2012) propose treating the genus Erythrocercus as a separate family, Erythrocercidae. One reason for separating them is that they have 12 rectrices rather than the 10 which the other Cettiidae have. Further, they are African rather than Asian (only one purely African species would be left in Cettiidae), and they are a fairly deep branch in the Sylvioidea tree. However, I am not yet persuaded we need another tiny family here. I'm even less persuaded by their suggestion of raising Scotocerca to family status. Both are treated as subfamilies here.

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 cettiid warbler. They found it sister to a clade containing Cettia and Abroscopus. This is reflected in the tree shown. However, the sampling of the cettiids was rather sparse and there may still be a residual question about its exact position within the Cettiidae.

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.

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).

Erythrocercinae: Bristle-flycatchers Fregin et al., 2012

Scotocercinae: Streaked Scrub Warbler Fregin et al., 2012

Cettiinae: Cettiid Warblers Alström, Olsson, & Ericson, 2014

Phylloscopidae: Leaf-Warblers Jerdon, 1863 (1854)

9 genera, 79 species Not HBW Family

These are followed by the leaf-warblers (Phylloscopidae), which is a new family consisting of about 80 species usually classified in two genera formerly belonging to the Sylviidae, Phylloscopus and Seicercus. 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. One way to handle the resulting changes is to move many species from Phylloscopus to Seicercus. This approach was previously taken in the TiF checklist. However, I think it is better to recognize the major clades as genera. Every genus recoginized differs in DNA by at least 12%, and is visually distinct from its nearest relative. Some subclades that also differ by 12% or more, but are not visually distinct, are not recognized as separate genera.

Both papers are consistent with a division into nine clades, each of which I recognize as a separate genus. A brief description of the typical member of each genus is given below. Some members of the various genera may differ from the description. As is commonly done, I use the term “phylloscops” to describe any of the eye-lined warblers typically placed in the genus Phylloscopus.

  1. Abrornis. Kinglet-like phylloscops: maculipennis through proregulus;
  2. Rhadina. Plain phylloscops with pale-edged tertials: orientalis, bonelli, and sibilatrix;
  3. Phyllosocpus. Plain phylloscops without wingbars: tytleri through canariensis;
  4. Pindalus. Reddish phylloscops: umbrovirens through laetus;
  5. Seicercus. Spectacled warblers: poliogenys through soror;
  6. Pycnosphrys: Reddish spectacled warblers: castaniceps, montis, and grammiceps;
  7. "Pycnosphrys": Fairly plain phylloscops, possibly with wingbars: cebuensis to ijimae;
  8. Acanthopneuste: Phylloscops with wingbars: xanthodryas through plumbeitarsus;
  9. Cryptigata: Capped phylloscops: ricketti to the end.

They also agree that clades 8 and 9 are sisters, as are (1)-(3) and (4)-(9). Olsson et al. (2004) has yet another topology, but the same clades appear. Previously, clades (1)-(4), and (7)-(9) were considered Phylloscopus while (5) and (6) were considered Seicercus.

There is another unresolved nomenclatural issue besides "Pycnosphrys". I'm not sure whether Rhadina (Billberg 1828) has priority over Phyllopneuste (Boie 1828). Both have the same type, sibilatrix.

I've split Japanese Leaf-Warbler, Acanthopneuste xanthodryas, and Kamchatka Leaf-Warbler, Acanthopneuste examinandus, from Arctic Warbler, Acanthopneuste 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 Acanthopneuste 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, Cryptigata calciatilis, see Alström et al. (2010).

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