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, making Turdidae and Muscicapidae sisters (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 remainder of the Muscicapoidea is the Turdidae-Muscicapidae clade, containing almost 500 species. This whole group and its relation to the Sylviioidea has long been contentitous. Ornithologists had great difficulty finding clear-cut ways to distinguish the various groups. This remained true until the DNA era. E.g., the 6th edition AOU checklist includes Sylviinae, Muscicapinae, Monarchinae, Turdinae, and Timaliinae as subfamilies of Muscicapidae! It was also typical to put old world robins, chats and wheatears in the Turdidae (or Turdinae), and various flycatchers in Muscicapidae (Muscicapinae). Sibley and Ahlquist (1990) and Sibley and Monroe (1990) began the process of untangling them by separating the true thrushes in Turdinae and reconsituting Muscicapinae as containing a flycatcher group (Muscicapini) and a robin-chat-wheatear group (Saxicolini). They didn't get it entirely right, but the current arrangement still has these general features. However, the membership of each group has been altered. Note that the Sibley and Monroe subfamilies Muscicapinae and Turdinae are treated as full families, and their tribes have been promoted to subfamilies.

Although the general arrangement of Sibley, Ahlquist, and Monroe has continued to be supported (e.g., Cibois and Cracraft, 2004; Voelker and Spellman, 2004; Zuccon and Ericson, 2010c; Sangster et al. 2010), the details have changed. Indeed, there's been substantial internal restructuring in both the Turdidae and Muscicapidae, with new genera appearing and old genera disappearing under the onslaught of the DNA-based studies.

Cibois and Cracraft already found a Cercotrichas Scrub-Robin and Copsychus Shama grouped in Muscicapinae, while a Ficedula flycatcher ended up with the Saxicolinae. Voelker and Spellman found much the same, and additionally provided a list of taxa erroneously placed in Turdidae, Muscicapinae, and Saxicolinae. Further work has given us a more detailed framework, but within that framework there are a number of surprises and apparently conflicting results.

Dippers

Cinclidae: Dippers

1 genus, 5 species HBW-10

Turdidae: Thrushes

19 genera, 166 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). There is some question about the relationships within Turdus itself. Voelker et al. (2007) and Nylander et al. (2008) found rather different results, although they are broadly similar.

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, Heinrichia and Alethe have moved to Muscicapidae. Grandala has moved the other way, from Muscicapidae to Turdidae (Jønsson and Fjeldså, 2006a).

There are two reasons I currently give preference to the Voelker et al. (2007) topology for Turdus rather than use Nylander et al. (2008). One is simply history. I used Voelker et al. first, and don't see a compelling case for change (both topologies have a number of soft nodes in them). The second is that some taxa that seem to be very similar get scattered by Nylander et al. This affects the blackbird complex (Nylander et al. have merula, intermedius, mandarinus, maximus, and simillimus in 4 different clades) and the olive thrushes (3 groups). They may ultimately prove correct, but I'd like stronger evidence before implementing such a thing, especially where there is evidence against it of equal strength.

Muscicapidae: Old World Flycatchers, Chats

56 genera, 318 species HBW-10 & 11

Muscicapidae tree The Muscicapidae have been a very troublesome family for the TiF list. They have undergone several reorganizations as new data become available. The lastest is version 2.60 which incorporates Sangster et. al. (2010). I'm happy to say that their results are generally congruent with Zuccon and Ericson (2010c), so that most of the large-scale features of the 2.54 reorganization are retained. Even the 2.54 changes were less extensive than the 2.10 reorganization, and it seems were are converging on the true phylogenetic tree. Most of the differences that exist between Sangster et al. (2010) and Zuccon and Ericson (2010c) are within the major clades, not between them.

Both Sangster et al. (2010) and Zuccon and Ericson (2010c) are multi-gene analyses with large samples of species. Both use about the same amount of DNA from 4 genes, with a 2 gene overlap between the papers. Zuccon and Ericson sampled 66 species of Muscicapidae, while Sangster et al. examined 124 species of Muscicapidae.

There are still problems because some generic boundaries need to be redrawn (e.g., see Seki, 2006), and there are issues all the way down to the species level (e.g., Stonechats). The starting point is now Sangster et al. (2010), supplemented by Zuccon and Ericson (2010c). To this 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), Outlaw et al. (2010), Pan et al. (2006), Seki (2006), Sheldon et al. (2009), Voelker (2010), Voelker and Spellman (2004), Wink et al. (2002), and Zuccon and Ericson (2010a). 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. This is pretty obvious when we look at the Muscicapinae.

Muscicapinae

One genus from Sibley and Monroe's Turdinae (Alethe) is basal. 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. There is a conflict here between Sangster et al. (2010) and Zuccon and Ericson (2010c). I've used the Zuccon and Ericson solution (placing Alethe basally in Muscicapinae rather than including it in Copsychini because they sampled both species of Alethe while Sangster et al. sampled only one. It is well-known that sampling two taxa from a genus is more likely to give reliable results.

The remainder of the subfamily splits into two groups, Copsychini and Muscicapini. Copsychini contains five genera that Sibley and Monroe put in Saxicolini (Saxicoloides, Trichixos, Copsychus, Cercotrichas, and Erythropygia). It turns out that some of the Cercotrichas form a basal clade. These have been separated in the genus Tychaedon (Richmond, 1917, type signata). The remaining Cercotrichas are sister to the Shamas and Magpie-Robins. The Indian Robin Saxicoloides is clearly nested within Copsychus, so I've merged it into Copsychus.

There is a little more uncertainty about the relationship of the other shamas to Rufous-tailed Shama, formerly in genus Trichixos. Both Sangster et al. (2010) and Zuccon and Ericson Trichixos to be the sister to the other shamsas, but Lim et al. (2010b) put the Rufous-tailed Shama closer to the magpie-robins. I've grouped it with the shamas, with both magpie-robins and shamas in Copsychus. An alternative naming convention would be to apply Copsychus to the magpie-robins (including the Indian Robin) and use Kittacincla (Gould 1830, type malabaricus) for the shamas.

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.

We now turn to Muscicapini. Using the terminology of Dickinson (2003), Muscicapini is restricted to Empidornis, Fraseria, Melaenornis, Muscicapa, and Myioparus. Neither Zuccon and Ericson (2010c) nor Sangster et al (2010) sample enough species to clarify the structure of Muscicapini. However, it appears that a group of Melaenornis flycatchers, including Empidornis and Sigelus, is basal. We list these all as Melaenornis. Next is the Pale Flycatcher. It's not clear if any other species group with it. If not, it will need a new genus name (I'm temporarily using “Bradornis”).

The remaining Muscicapini fall into three groups. The first includes some former Melaenornis which I've separated in Bradornis (Smith 1847, type mariquensis). There is a name complication here as both Sooty Flycatcher and Chat Flycatcher are named infuscatus. The Chat Flycatcher was originally called Saxicola infuscata by A. Smith in 1839, while the Sooty Flycatcher was dubbed Batails infuscatus by Cassin in 1855. Thus Chat Flycatcher gets the name infuscatus.

But what about the Sooty Flycatcher? It was originally named Artomyias fuliginosa by J. and E. Verreaux in 1855. Once upon a time this conflicted with Muscicapa fuliginosa Sparrman 1787, but that species is now in Rhipidura, far away from any of the Muscicapidae. If we retained the Sooty Flycatcher in Muscicapa, there'd be another fuliginosa conflict, as the name was also applied by Hodgson in 1845 to what is now Muscicapa sibirica cacabata. The name cacabata was coined by Penard in 1919 in order to avoid the conflict with Rhipidura. Since I'm treating the Sooty Flycatcher as part of Bradornis, it is safe to call it fuliginosa.

The second group includes one of the Fraseria, both Myioparus, and at least a couple of Muscicapa. These are all placed in Fraseria. Finally, there are the remaining Muscicapa. As this group includes the type species (striata), it retains the name Muscicapa. It seems likely that griseisticta through segregata is the sister clade to the remaining Muscicapa. The name Hemichelidon (Hodgson 1845, type sibirica) could be applied to this group, highlighting the geographic differences between the two clades.

I'm not at all sure where Humblotia fits, so I've placed it at the end with the other incertae sedis species.

Alethini: True Alethes

Copsychini: Scrub-Robins, Magpie-Robins, Shamas

Muscicapini: Old World Flycatchers

Niltavinae: Blue Flycatchers

Both Zuccon and Ericson (2010c) and Sangster et al. (2010) found a clade containing the blue flycatchers. Sangster et al. recommended treating it as a subfamily and proposed the name Niltavinae.

Sangster et al. considered several species of Rhinomyias and discovered it was not a natural group (there seem to be at least 4 pieces). Much of Rhinomyias seems to be embedded in Cyornis. Those taxa are merged here. It's a little surprising to include the drab Rhinomyias in Cyornis. Further analysis should help us make better sense of this. The Streak-breasted Jungle-Flycatcher, Rhinomyias additus, has been transferred to Eumyias and several other members of Rhinomyias have been transferred to Vauriella in Saxicolinae.

The White-tailed Flycatcher appears to be basal in this group. Whether it is alone or whether other species are with it is not yet clear. We use the temporary genus “Cyornis” until this situation is straightened out.

The remainder of Niltavinae breaks into two parts. The first consists of Cyanoptila, Eumyias, and Niltava. Although Lei et al. (2007), using less data, considered this group part of Muscicapinae (as did Sibley and Monroe), Voelker and Spellman (2004), Sangster et al. (2010), and Zuccon and Ericson (2010c) found the contrary. The second clade includes Anthipes (previously removed from Ficedula by Outlaw and Voelker, 2006a), the Cyornis blue-flycatchers and jungle-flycatchers (transferred from Rhinomyias).

Cossyphinae: African Robins

The African Robins, Cossyphinae, are the most problematic part of the Muscicapidae. There are many studies concerning them: Roy et al. (2000), Beresford (2003), and Voelker et al. (2010a) focus on the group, and it is an important component of the large-scale analyses of Zuccon and Ericson (2010c) and Sangster et al. (2010). Moreover, a number of other papers have focused on individual species or small groups of species. When all of this is put together, the result is confusion. There is no argreement on how the genera are arranged, and there are still questions about which species belong to which genera and about which species are actually species. The only really consistent point is the list of taxa that belong to Cossyphinae.

With that in mind, I'm now using Sangster et al. (2010) as the basic framework. It is not really satisfactory as it doesn't provide a lot of resolution. Moreover, comparison with Zuccon and Ericson (2010c) makes one wonder if even the terminal clades are right.

Even the name of this group is a problem. Sangster et al. (2010) refer to this subfamily as Erithacinae, citing G.R. Gray 1846 (priority seems to date to 1831 under a different name). Still, the name Cossyphinae dates to Vigors 1825 (as Cossyphina), and should have priority.

The sequence starts with Xenocopsychus and Dessonornis (A. Smith 1836, type humeralis). Although Sangster et al. only included archeri and humeralis, Voelker et al. (2010a) also grouped caffra and anomala with archeri. Using substantially less data, Beresford (2003) grouped caffra and archeri, but excluded anomala. Including humeralis allows us to use the name Dessonornis. Finally, Xenocopsychus has been thought to be close to humeralis.

Voelker et al. (2010a) advocated use of the name 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. If humeralis were not there, the oldest name having any of these as type species is Caffronis (Roberts, 1922; type C. caffra).

The remainder is organized as a four-fold polytomy, which is a way of say the data is pretty inconclusive. One piece of the polytomy is the only non-African species in the group, the Eurasian Robin.

A second piced consists of Pogonocichla, Oreocossypha, Cossyphicula, and Swynnertonia. Beresford found evidence that all three form a clade. It might be reasonable to place these species in the same genus, but it's probably better to wait for more evidence.

Beresford (2003) 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, part of Cossyphinae. Sangster et al. (2010) group them with a somewhat restructered Cossypha.

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, Dessonornis.

The last of the 4 clades includes Stiphrornis, Cichladusa, and the revised Sheppardia. Only Voelker et al. (2010a) analyzed Cichladusa, finding that it is near Sheppardia, but not so close to Stiphrornis.

The IOC list has lumped the various Stiphrornis forest-robins into a single species. I consider that unreasonable based on current data. The current data shows at least 5 taxa (xanthogaster, sanghensis, gabonensis, erythrothorax, and pyrrholaemus) that are all 5-6% distant from each other in terms of DNA, while members of the same taxa are separated by much smaller distances, often less that 0.5%. This type of genetic separation usually indicates species status, and I have not seen any compelling evidence to the contrary. It not a slam dunk only because the geographic sampling of these taxa is currently rather sparse. See Beresford and Cracraft (1999) and Schmidt et al. (2008).

Saxicolinae: Robins, Chats, Wheatears

Another important discrepancy between Zuccon and Ericson (2010c) and Sangster et al. (2010) is the basal group in Saxicolinae. Sangster et al. included the White-browed Jungle-Flycatcher, Rhinomyias insignis. in their analysis. It grouped with a former member of the Turdidae, the Great Shortwing, Heinrichia calligyna. The two were pulled into a group with more former Turdidae, Brachypterx, and some of the northern robins. Of course, most of the Rhinomyias are left in the Muscicapinae with Cyornis. In 1980, Wouters had suggested that insignis and several other Rhinomyias be placed in a new genus, Vauriella. I follow that here.

The former robins were variously placed in Luscinia and Erithacus. However, neither of those type species is in this group and they take the name Larvivora (Hodgson 1837, type cyane). Sangster et al. (2010) suggest that the the Rufous-headed Robin belongs here. I am not convinced, but will go with their suggestion as I am not personally familiar with these taxa. Larvivora is sister to the Brachypteryx shortwings.

This group is followed by the nightingales and allies. Here the White-throated Robin, Irania gutturalis, is basal. Then we have the White-bellied Redstart, Hodgsonius phoenicuroides and the Bluethroat on one side, and the two nightingales on the other. I've put the Bluethroat in its own genus, Cyanecula (Brehm 1828).

The next clade is somewhat larger, containing two main groups. The first is another collection of former Luscinia, these placed in Calliope (Gould 1836, type calliope). These are sister to the Myiomela blue-robins and Tarsiger bush-robins. The other half of the clade includes the Myophonus whistling-thrushes (former Turdidae) and the Enicurus forktails (former Muscicapinae). The forktails have been studied by Moyle et al. (2005), and the arrangement here draws on that work.

After this, several genera branch off one at a time: the Ficedula flycatchers, Phoenicurus redstars, Monticola rock-thrushes, and Saxicola chats. The Saxicolinae end with the wheatears and allies.

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.

The redstarts are often grouped together. Both Pan et al. (2006) and Voelker (2010) found that Phoenicurus is paraphyletic with respect to Chaimarrornis and Rhyacornis. There are two alternatives, either split the first five as Adelura (Bonaparte 1854, type caeruleocephala), or lump them all into Phoenicurus. Since Chaimarrornis and Rhyacornis have often been considered part of Phoenicurus, and since the genetic distances are relatively small, implying the whole clade is only 5-6 million years old (Voelker, 2010), I've merged them all into Phoenicurus. I've also rearranged them based on Voelker (2010). Pan et al. (2006) found schisticeps belongs in the Adelura group. The position of Moussier's Redstart, Phoenicurus moussieri, has been clarified by Sangster et al. (2010).

There is some question about whether the Monticola rock-thrushes are sister to the chats and 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 also 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). Note that Ruaha Chat, Pentholaea collaris, has been split from Arnot's Chat, Pentholaea arnotti, based on Glen et al. (2010). The arrangement of the various small genera is a compromise between Outlaw et al. (2010), Sangster et al. (2010), and Zuccon and Ericson (2010c).

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.

Incertae sedis

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