Muscicapoidea II

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 44 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Cinclidae, Turdidae, and Muscicapidae

The dippers (Cinclidae) seem to come first (see Barker et al., 2004; Beresford et al., 2005; Treplin et al., 2008; but for a contrary view see Ericson and Johansson, 2003; Voelker and Spellman, 2004).

The last clade in the Muscicapoidea is the Turdidae-Muscicapidae clade. The biggest change here is that a number of genera including the old world chats and wheatears are transferred from Turdidae to Muscicapidae. This is apparent from Cibois and Cracraft (2004), and made quite clear by Voelker and Spellman (2004).

Dippers

Cinclidae: Dippers

1 genus, 5 species HBW-10

Turdidae: Thrushes

20 genera, 167 species HBW-10

The overall organization of the Turdidae follows Klicka et al. (2005) and Voelker and Klicka (2008). The arrangement of Myadestes follows Miller et al. (2007). Voelker et al. (2007) was used for the large genus Turdus while Voelker and Klicka (2008) covers the true Zoothera. The treatment of the Olive Thrush complex is based on Bowie et al. (2005).

Turdidae tree Klicka et al. identified several major clades in Turdidae, but I have not translated them into subfamilies and tribes, mainly because I'm not sure how they fit together. The basal group includes the bluebirds and solitaires. After that come the Australasian Zoothera. A big chunk of what had been Zoothera is not closely related to the other part. The Afroasiatic Zoothera are now Geokichla (for details, see Voelker and Outlaw, 2008). The true Zoothera form another clade, as does Catharus and allies. There is also a small clade consisting of Chlamydochaera and Cochoa which are allied to Geokichla and Turdus.

As part of the Turdus reorganization, Cichlherminia, Platycichla, and Nesocichla have been merged into Turdus. The genera Myophonus, Brachypteryx, and Alethe have moved to Muscicapidae. Grandala has moved the other way, from Muscicapidae to Turdidae (Jønsson and Fjeldså, 2006a).

Muscicapidae: Old World Flycatchers, Chats

58 genera, 316 species HBW-11

Muscicapidae tree The Muscicapidae have had another major reorganization in version 2.10, as I have found some additional papers concerning them. The fundamental problem remains: Too few taxa have been sampled! The fact that some generic boundaries need to be redrawn only compounds the problem (e.g., see Seki, 2006), and there are problems all the way down to the species level (e.g., Stonechats). The starting point is the assignment of genera to the two subfamilies is based on Voelker and Spellman (2004), to which I have added information from a number of additional papers: Beresford (2003), Cibois and Cracraft (2004), Illera et al. (2008), Lei et al. (2007), Moyle et al. (2005), Outlaw and Voelker (2006), Outlaw et al. (2007), Pan et al. (2006), Seki (2006), Voelker and Spellman (2004), and Wink et al. (2002). From these, I came up with the tree on the right. Earlier versions relied on Jønsson and Fjeldså's supertree (2006a), but it has little direct impact on this version.

The relationships amoung the Turdidae (traditionally including robins, chats, and thrushes) and the Muscicapidae (flycatchers) have long been confusing. Sibley and Monroe grouped them into a large Muscicapidae family with the thrushes in Turdinae, flycatchers as tribe Muscicapini (in subfamily Muscicapinae), and robins and chats in tribe Saxicolini (also Muscicapinae). A similar arrangement is followed here, with families Turdidae and Muscicapidae, the latter divided into Muscicapinae and Saxicolinae. However, some of the genera invovled had hopped from one group to another.

Muscicapinae

This is pretty obvious when we look at the Muscicapinae. The basal group consists of one genus from Sibley and Monroe's Turdinae (Alethe) and five from their Saxicolini (Saxicoloides, Trichixos, Copsychus, Cercotrichas, and Erythropygia). Muscicapini is conspicuously absent. The situation is actually a little more complex than it seems as Beresford (2003) split Alethe, with part (Pseudalethe) moving to Saxicolinae. None remained in Turdidae.

The other half of the Muscicapinae are traditional members of the group. The genera from Empidornis to Melaenornis are sometimes all considered part of Melaenornis. There has been little in the way of DNA studies involving these species, so the arrangement is fairly standard.

The Philippine Magpie-Robin, Copsychus mindanensis, was recently split from Oriental Magpie-Robin, Copsychus saularis, by Sheldon et al. (2009). It appears to be basal among the Magpie-Robins.

Saxicolinae

I've put the former Turdidae Brachypteryx and Myophonus at the beginning of Saxicolinae, but I don't really know where they go (hence the blue on the diagram). Voelker and Spellman (2004) found they belonged in Saxicolinae, but did not pin down where. Except for the position of the blue-flycatchers (which were not pinned down), the placement of the Saxicolinae clades follows Voelker and Spellman. Using less data, Lei et al. (2007) considered the blue-flycatchers part of Muscicapinae (as did Sibley and Monroe), but Voelker and Spellman found the contrary. It seems reasonable to put them in a basal position within Saxicolinae. According to Outlaw and Voelker (2006a), who removed the two Anthipes flycatchers from Ficedula, they belong near Niltava/Cyornis.

The forest-robins were studied by Roy et al. (2000), Beresford (2003), and Voelker et al. (2010a). Voelker et al. found the European Robin sister to Stiphrornis. Beresford found that several members of Alethe were not closely related to the other Alethe. In fact, it turns out they do not belong in Muscicapinae at all. Rather, they belong with the forest-robins and take the genus name Pseudalethe. Beresford also found evidence that both Sheppardia and Cossypha were paraphyletic. Recently, Voelker et al. (2010a) focused on Sheppardia and the related species in Cossypha. Their analysis is the basis of the arrangement here. The Gray-winged Robin-Chat, formerly Cossypha polioptera, has been transferred to Sheppardia. Three other members of Cossypha, caffra, anomala, and archeri, have been placed in a separate genus sister to Sheppardia.

Voelker et al. (2010a) suggest this sister genus should be called Callene. The name Callene is due to Blyth (1847). He introduced it as a substitute for Cinclidium. He felt this was necessary as Cinclidium was already in use in botany for a type of moss. These days, the same genus name can and is used in both botany and zoology without conflict, and Cinclidium has regained its name. Although it subsequently became a larger genus (and later shrank), when Blyth proposed Callene, Cinclidium contained only one species, Cinclidium frontale. That means the type of Callene is Cinclidium frontale. Thus Callene does not apply to the clade consisting of caffra, anomala, and archeri. So far as I can determine, the oldest name having any of these as type species is Caffronis (Roberts, 1922; type C. caffra). That is the name I use for these three species.

Outlaw and Voelker (2006a) made a comprehensive study of the Ficedula flycatchers, covering most of the species (a less complete treatment is Lei et al., 2007). As a result, two were removed entirely to Anthipes. Further, Muscicapella has been moved into Ficedula. This means that the Pygmy Blue-Flycatcher, Muscicapella hodgsoni, is now Ficedula hodgsoni. This creates a name conflict with the Slaty-backed Flycatcher, which becomes Ficedula erithacus. Særte et al. (2001) show that the Atlas Flycatcher is a separate species, and the Olive-backed Forest-Robin, Stiphrornis pyrrholaemus, was recently discovered (Schmidt et al., 2008). The forktails have been studied by Moyle et al. (2005).

From here on, most genera have been considered part of Saxicolinae. Generic limits among the palearctic robins have long been in flux. Seki (2006) found that the eastern Erithacus (sometimes considered Luscinia) were not closely related to the European Robin. Rather, they are close to some, but not all Luscinia. I've used the old name Larvivora to refer to this group, which is sister to Tarsiger. The true Luscinia seem to be sister to the whole bunch. I also don't know whether Hodgsonius, Cinclidium, or Myiomela belong with the robins or not. However, they are often placed near them, so it seems like a reasonable choice.

The Redstarts are often grouped together. There is some question whether Chaimarrornis and Rhyacornis are good genera. Pan et al. (2007) found them embedded in Phoenicurus. Perhaps they should be merged, but I'd like to see a more complete analysis first. The recent paper by Outlaw et al. (2010) suggests that the redstarts and palearctic robins form a clade, but does not give details.

The rest fall into two groups, the Monticola rock-thrushes and the chat/wheatear clade. The evidence on whether they are sister is mixed, and Outlaw et al. (2010) show a different arrangement (without details). The Monticola rock-thrushes do not belong in Turdidae (see Wink et al., 2002; Outlaw et al., 2007). They include the Madagascan Pseudocossyphus rock-thrushes. The former Pseudocossyphus rock-thrushes are sometimes treated as 4 species, but Zuccon and Ericson (2010a) found little genetic distinction between M. sharpei, bensoni, and erythronotus, so I've followed their recommendation and lumped them all into M. sharpei. The other former Pseudocossyphus, M. imerinus is retained as a separate species. Zuccon and Ericson (2010a) also found that the White-winged Cliff-Chat belongs in Monticola, not Thamnolaea. Moreover, M. solitarius breaks into two clades which are not each other's closest relatives. The second group consists of the races madoci, philippensis, and pandoo, so they take the name Monticola philippensis. The English name Red-bellied Rock-Thrush has been previously used for philippensis, and I use it here. Nonetheless, it is not a very good name as pandoo, which interbreeds widely with philippensis, is not red-bellied. Perhaps something like “Variable Rock-Thrush” would be better.

The chat/wheatear clade has undergone a fair amount of reorganization. It includes the Buff-streaked Chat, Campicoloides bifasciatus, which Illera et al. (2008) removed from Saxicola. The clade probably includes the Mountain Wheatear, which Outlaw et al. (2010) found is not part of Oenanthe. For the present, it is placed in the monotypic genus Dromolaea (Cabanis, 1850). The Moorland Chat has been moved to Pinarochroa from Cercomela as in Outlaw et al. They also dismembered Cercomela, with the species invovled being distributed between Oenanthe and the revived genus Emarginata (Shelly, 1896, type E. sinuata). The arrangement of Oenanthe is based primarily on Outlaw et al. (2010), but Aliabadian et al. (2007) was also consulted concerning species not included by Outlaw et al.

The treatment of Saxicola here is based on Wink et al. (2002), Illera et al. (2008), Woog et al. (2008), and Zink et al. (2009). This involves splitting the Common Stonechat into at least seven species: Stejneger's, Siberian, Canary Islands, European, African, Madagascan, and Reunion Stonechats. There may still be additional species hiding in the stonechat complex. For ABA-listers, East Siberian Stonechat is known to occur in the ABA area. The status of West Siberian Stonechat in the ABA area is currently unclear to me.

It is still quite unclear where the remaining three genera go, so I've put them in a separate Incertae sedis group. They are thought to be in Saxicolinae, but there isn't much real evidence. One or more might belong to Turdidae, or even to a less closely related family. Still, the odds are they go somewhere in Muscicapidae.

Muscicapinae

Saxicolinae

Incertae sedis

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