Basal Passeroidea

Core Passeroidea


Tyranni: Suboscines

Passeri: Oscines


Muscicapoidea and allies

The 46 Orders














Thraupid Group

We've now reached the last set of families, the Mitrospingidae, Cardinalidae, and Thraupidae. It might make sense to to treat them all as subfamilies of an expanded Thraupidae. The genetic distance between them is small. Barker et al. (2013) estimate they have a common within the last 13 million years. In comparison, most families represent separate lineages stretching back 20 million years or more. Moreover, it's difficult to distinguish whether birds are cardinal-grosbeaks or tanagers (e.g., the Piranga tanagers, the saltators, the Paroaria cardinals, all of Mitrospingidae) suggests that it might be better to reduce both Mitrospingidae and Cardinalidae to subfamilies of Thraupidae.

Moreover, I am not the first to suggest downgrading them. Sibley and Monroe (1993) also noted the close relationship and went much further in combining families, treating the tanagers and cardinals as tribes within a broad subfamily Emberizinae which is equivalent to what is ranked here as the epifamily Icteroidae.

Mitrospingidae: Mitrospingus and allies Barker et al., 2013

3 genera, 4 species Not HBW Family

Although they are generally considered tanagers, there have been several papers indicating that Lamprospiza and Mitrospingus are not tanagers. Burns (1997) already found they lie outside the tanagers, and perhaps sister to Chlorospingus, now known to be a sparrow. Taxon sampling outside the tanagers was sparse, so Burns could not accurately place them. Yuri and Mindell (2002) provided additional evidence that they were not tanagers. The same was true of Burns et al. (2003). Klicka et al. (2007) had good sampling of all the relevant groups. They found Mitrospingus sister to the Emberizid group.

The most recent and most comprehensive study including them is Barker et al. (2013). Their results suggest that the two genera belong to the Thraupid group, and that Orthogonys belongs near them. Barker et al. established the family name Mitrospingidae for these three genera.

Cardinalidae: Cardinals, Grosbeaks Ridgway, 1901

12 genera, 56 species HBW-16

Click for Cardinalidae tree
Click for Cardinalidae tree

The Cardinalidae have been reorganized using Klicka et al. (2007). The saltators, Parkerthraustes, and Porphyrospiza are out. They join the tanagers. In return, the cardinals get Piranga, Habia, and Chlorothraupis from the tanagers. They also gain the Granatellus chats from the warblers and the Amaurospiza blue seedeaters that are sometimes considered sparrows, sometimes tanagers.

Both Klicka et al. (2001) and Klick et al. (2007) found that Lazuli Bunting, Passerina amoena, is sister to the Blue Grosbeak, P. caerulea, rather than to the Indigo Bunting, P. cyanea. This is apparently an artifact due to a combination of recent divergence and relatively large populations. In such cases, a longer time is necessary for all genes to fully reflect the species tree. This has been studied in detail by Carling and Brumfield (2008), who found that the Lazuli and Indigo Buntings are sister species. As is well-known, they do hybridize in the contact zones of the western US. Carling and Brumfield (2009) and Carling et al. (2010) have also studied this phenomenon. Among other things, their further studies support the classification of Lazuli and Indigo Buntings as distinct and monotypic species.

Some of the generic boundaries needed adjustment. I have followed Klicka et al.'s suggestions of folding Chlorothraupis into Habia. Based on Bryson et al. (2014), Cyanocompsa has been merged into Cyanoloxia, except for the Blue Bunting, which is now in Passerina. The species flagged in blue lack molecular data, but are believed to be in Habia.

Following IOC and Ridgely (Ridgely and Greenfield, 2001; Ridgely and Tudor, 2009), the Olive Tanager, Habia frenata, has been split from from Carmiol's Tanager, Habia carmioli.

The arrangement of Piranga is based on Manthey et al. (2016). Hepatic Tanger has been split into four species: Northern Hepatic-Tanager / Hepatic Tanager, Piranga hepatica, Highland Hepatic-Tanager / Tooth-billed Tanager, Piranga lutea, Blood-red Tanager, Piranga haemalea, and Lowland Hepatic-Tanager / Red Tanager, Piranga flava, These have frequently been considered distinct species and some are so treated in many regional guidebooks. Burns (1998) found genetic differences between the three in the range typical of species, not subspecies. Manthey et al. (2016) clarified the position of Piranga haemalea.

The order within Pheucticus is based on Pulgarín-R et al. (2013). They also found that two of the yellow grosbeaks were not monophyletic. The stronger case for a split is the Orange-colored Grosbeak, Pheucticus aurantiacus, which I have split from the (Mexican) Yellow Grosbeak, Pheucticus chrysopeplus. The Orange-colored Grosbeak is monotypic, distinctly orange, and can be found in Chiapis, Mexico and Guatemala. There is a weaker case for splitting Pheucticus uropygialis from the Black-backed Grosbeak, Pheucticus aureoventris (monotypic). There's not an identification problem. The rump is variously yellow or mottled in the uropygialis group, black in aureoventris. However, the genetic distance is much smaller and there are questions about whether crissalis (part of the uropygialis group) interbreeds with chrysogaster in Ecuador. It could be that this complex is best treated as one species rather than three. For the present, I leave it at the conventional two.

Within the blue cardinalids, I've followed Bryson et al. (2014) in splitting Ecuadorian Seedeater, Amaurospiza aequatorialis, from Blue Seedeater, Amaurospiza concolor (which now takes the name Cabanis's Seedeater), and splitting Rothschild's Grosbeak, Cyanoloxia rothschildii, from Blue-black Grosbeak, Cyanoloxia cyanoides. The latter gives me an armchair tick as I have seen Rothschild's at Alta Floresta (Brazil) and Blue-black in the Canal Zone (Panama). I additionally split the Turquoise-fronted Bunting, Passerina indigotica, of western Mexico from Blue Bunting, Passerina parellina.

Thraupidae: Tanagers Cabanis, 1847

95 genera, 385 species HBW-16

Thraupidae tree
Thraupidae family tree

We finally reach the last family on the list! The tanagers have been the subject of an extensive ongoing revision. Kevin Burns and his co-workers have spent the last twenty years studying the tanagers. E.g., Burns (1997, 1998); Burns et al. (2002, 2003); Burns and Naoki (2004); Burns and Racicot (2009); Mauck and Burns (2009); Mallarinoa et al. (2012); Sedano and Burns (2010); Barker et al. (2013); Mason and Burns (2013); Ryan et al. (2013); Shultz and Burns (2013). This work has culminated in six-gene study Burns et al. (2014), which is the primary source for the TiF phylogeny of the tanagers.

In recent years, the tanagers lost the euphonias and chlorophonias to the finches, Habia and Piranga (including all of the North American tanagers) to the cardinals, the Mitrospingidae, and the Phaenicophilidae. However, the tanagers have also gained many species. The tanager-finches were historically often considered sparrows mostly end up in the tanagers. The tanagers also gained the saltators, some other cardinals, and the bananaquit.

The result of this shifting genera is a very large, very heterogeneous tanager family. This is especially remarkable as the tanagers are a relatively recent radiation. There is a basal division into two parts, Thraupinae and Dacninae. I have found it useful to further divide these into 18 tribes (15 of these are the “subfamilies” of Burns et al., 2014; 3 are further subdivisions of these).

Thraupidae Subfamilies

Thraupinae Cabanis, 1847

Thraupinae tree
Thraupinae subfamily tree
Click for Thraupinae species tree

The Thraupinae are shown in more detail at the right. There are six groups: Catamblyrhynchini and Porphyrospizini (formerly Incaspizini), and the Orchesticini, Thaupini, Cissopini, Pipraeideini clade.

Catamblyrhynchini: Plushcap

This contains a single species, Plushcap. It's closest relatives appear to be the Porphyrospizini, but the relationship is rather distant.

Porphyrospizini: Inca-Finches

I had previously tentatively placed Incaspiza and its allies basally in Poospizini. However, Barker et al. (2013) was the first to sequence any of the Incaspiza, finding that the Great Inca-Finch grouped with the Blue Finch and Mourning Sierra Finch.

The Rhopospina are rather distant from the other sierra finches (Campanga et al., 2011, Klicka et al., 2007). Ridgely and Tudor (1989) long ago grouped the Band-tailed, Carbonated and Mourning Sierra Finches based on plumage. Burns et al. (2014) found that the Blue Finch (Porphyrospiza) also belongs in this clade. The display flights may also indicate a connection. Campagna et al. (2011) found that they are each other's closest relatives, and quite distant from any of the other sierra finches. This small clade takes the name Rhopospina (Cabanis 1851, type fruticeti) which has priority over Corydospiza (Sundevall 1872, type alaudina) and Porphyrospiza (PL Sclater and Salvin 1873, type caerulescens).

The rest of Thraupinae is in the main clade. This includes four successive branches: Orchesticini, Pipraeideini, Cissopini, and Thraupini.

Orchesticini: Grosbeak-Tanagers

This is a small group of two species, Parkerthraustes and Orchesticus that Burns et al. (2014) found to be basal to the main clade, and not particularly closely related to them. The relationships of these two species have been murky. Weir et al. (2009), using only cytochrome-b, found Parkerthraustes basal in Saltatorinae. Klicka et al. (2007), using more genes, found it sister to Chlorochrysa, but with weak support. Finally, the more comprehensive analysis of Barker et al. (2013) put it sister to Orchesticus with very strong support, and the two of them were weakly attached to the Hemithraupini clade.

Pipraeideini: Mountain-Tanagers

That brings us to Pipraeideini. These are almost all mountain-tanagers, birds of the Andes. Here again I use an informal name for the tribe as none have been established in the literature.

Based on the topology of Sedano and Burns (2010) and SACC decision #569 several of the genus boundaries have been changed. The Blue-and-yellow Tanager, formerly Thraupis bonariensis, is now in Pipraeidea. The genus Delothraupis (Chestnut-bellied Mountain-Tanager) has been merged into Dubusia. It is sister to the former Rufous-bellied Saltator, Dubusia rufiventris, which is now called Rufous-bellied Mountain-Tanager, Pseudosaltator rufiventris due to SACC Proposal #722.

The genus Buthraupis has been dismembered, leaving only the Hooded Mountain-Tanager, Buthraupis montana. The Grass-green Tanager is now in the monotypic Chlorornis (Reichenbach, 1850), and the Masked Mountain-Tanager is in the monotypic Tephrophilus (Moore 1934), while the Black-chested and Golden-backed Mountain-Tanagers have been put in Cnemathraupis (Penard, 1919, type eximia). The Blue-capped Tanager, formerly Thraupis cyanocephalus, is now in the monotypic Sporathraupis (Bonaparte, 1850). The Blue-winged Mountain-Tanager, Anisognathus somptuosus, and Black-chinned Mountain-Tanager, Anisognathus notabilis, have been placed in genus Compsocoma (Cabanis, 1853, type somptuosa victorini).

Cissopini: Cardinal-Tanagers

Both Weir et al. (2009) and Sedano and Burns (2010) put Chlorochrysa in Cissopini, and we follow that here. Note that Klicka et al. (2007) found a somewhat different arrangement, with Chlorochrysa sister to Parkerthraustes, and the pair basal to several tanager tribes.

Klicka et al. (2014) found that the Yellow-green Bush-tanager, formerly Chlorospingus flavovirens, was a tanager, and not part of the sparrow genus Chlorospingus. However, they did not include enough tanagers in their analysis to determine its affinities. This has now been remedied by Avendaño et al. (2016), who found it is sister to the Blue-and-gold Tanager, Bangsia arcaei. As a result, the Yellow-green Bush-tanager is now Bangsia flavovirens.

The Paroria and Gubernatrix cardinals (often considered sparrows) are in Cissopini with several other tanagers and finches. The only DNA information available concerning Gubernatrix is from Barker et al. (2013). The species limits of Paroaria are a bit non-standard. Based on Dávalos and Porzecanski (2009), I've separated Masked Cardinal, Paroaria nigrogenis, from Red-capped Cardinal, P. gularis, and moved the subspecies cervicalis of eastern Boliva and NW Mato Grosso into Yellow-billed Cardinal, P. capitata. (An alternative would be to lump capitata with gularis as Red-capped Cardinal, as suggested many years ago by Hellmayr (1938)). Thus Yellow-billed Cardinal includes the subspecies cervicalis, capitata, and fuscipes. Based on Lopes and Gonzaga (2013), I have also split the Crimson-fronted Cardinal into the black-throated Xingu Cardinal, Paroria xinguensis and crimson-throated Araguaia Cardinal, Paroria baeri.

Thraupini: Tangara Tanagers

In the current phylogeny, Thraupini is the biggest group in Thraupinae. It consists solely of the genus Tangara. That means it includes some of the most attractive birds on the planet. You may be wondering why it's called Thraupini when the only genus present is Tangara. Most of the genus Thraupis, including the terminologically important type species (formerly Thraupis ornata) have been subsumed in Tangara as a result of Sedano and Burns (2010). Although Tangara has priority (by seniority) over Thraupis at the genus-level, an official ruling means that Thraupis has priority at the family (and tribal) level. Thus the tribe containing Tangara ornata is known as the Thraupini.

If you example the species-level tree, you'll see that Tangara contains two clades, which could legitimately be called Tangara and Thraupis. I've chosen to retain these as subgenera, but hope that the AOU will do the sensible thing and use these these as genus names. Arguably I should go ahead and use them here, but at present I prefer that one of the AOU committees take the lead.

A close examination of the species-level Thraupini tree also reveals the numbers 1-13 labelling most of the clades in Tangara. These indicate the numbered clades identified by Isler and Isler (1987) using traditional taxonomic methods. Except for clades 3 and 9, they match up precisely with the genetic data. The clade labelled “Th” consists of the Tangara species formerly placed in Thraupis.

The Black-headed Tanager had to take an alternate name, Tangara argentea, because the Azure-shouldered Tanager, formerly Thraupis cyanoptera, has first claim on Tangara cyanoptera.

Thraupinae Species List

Catamblyrhynchini: Plushcap Ridgway, 2014

Porphyrospizini: Inca Finches and allies Burns et al., 2014

Orchesticini: Grosbeak-Tanagers Burns et al., 2014

Pipraeideini: Mountain-Tanagers Informal

Cissopini: Cardinal-Tanagers Sundevall, 1872

Thraupini: Tangara Tanagers Cabanis, 1847

Dacninae Sundevall, 1836

Dacninae tree
Dacninae subfamily tree

The rest of the Neotropical finches are part of Dacninae (a few remain outside the tanagers). This part of the tanagers is full of seedeaters, seedfinches, grass-finches, warbling finches, sierra finches, pileated finches, yellow finches, etc, etc, but not the brushfinches. It also contains the flower-piercers. The Dacninae does contain birds other than Neotropical finches. It includes the conebills, honeycreepers, and even some tanagers such as the striking Silver-beaked Tanager.

As with the Thraupinae, the arrangement here is based on Burns et al. (2014).

Nemosiini: Flocking-dwelling Tanagers

Barker et al. (2013) found that the previously untested Cyanicterus and Compsothraupis tanagers belong in this small clade which has 100% bootstrap support. There had been question about whether the similarity between Compsothraupis (Scarlet-throated Tanager) and Sericossypha (White-capped Tanager) was convergence or inheritance. Storer argued that it was convergence, a view endorsed by Ridgely and Tudor (1989). The jury is now in. Barker et al. found they are sister species and so I have merged Compsothraupis (Richmond 1915) into Sericossypha (Lesson 1844).

Conirostrini: Conebills

The conebills are in their own clade, Conirostrini. There is general consensus that the Conirostrini are sister to Diglossini. They could be treated as a single tribe under the name Diglossini. Since the conebills are distinctive and the division is relatively deep (Weir et al., 2009), I leave them separate.

The Giant Conebill is usually placed in the genus Oremanes as Oremanes fraseri (Sclater 1860). However, it turns out to be nested in Conirostrum, and so has been changed. This creates a bit of a nomenclatural complication as fraseri is preoccupied by Conirostrum cinereum fraseri (Sclater 1859). The next oldest available name for the Giant Conebill appears to be binghami (Chapman 1919).

Diglossini: Highland Tanagers

That brings us to the last group, the Diglossini finches. The arrangement here draws heavily on Campagna et al. (2011) and Mauck and Burns (2009). Barker et al. (2013) allowed me compeletely resolve the tree in a reasonable way.

Diglossini contains two clades. The first consists of yellow finches. Ryan et al. (2013) have eliminated the guesswork here. They found that Rowettia and Melanodera were each other's closest relatives, with Nesospiza sister to the pair. The whole clade is sister to the true Phrygilus sierra finches. Although this was the most likely arrangement, there was some support for Melanodura and Nesospingus as sisters, with Rowettia sister to the pair. Either way, the Atlantic islands of Tristan de Cuha and Gough have twice been colonized by South American finches.

The arrangement within Sicalis is based on Ryan et al. (2013), who included all but one species in their analysis. Their analysis also raised doubts about whether the Stripe-tailed Yellow-Finch belongs in Sicalis. In fact, most genes placed it outside Sicalis, but there was no agreement on whether it is closer to the true Phrygilus, basal to both, or even closer to the Melanodera/Rowettia/Nesospiza clade. For now, it is placed in the monotypic Pseudochloris (Sharpe 1888).

The Monte Yellow-Finch, Sicalis mendozae, has been split from Greenish Yellow-Finch, Sicalis olivascens. See Areta et al. (2012) and the discussion of SACC proposal 539.

The other clade includes the flowerpiercers, Tit-like Dacnis, and various gray finches. It's a bit unclear exactly where Catemenia fits. I'm following Barker et al. (2013) who place it sister to Diglossa, although Campagna et al. (2011) give a slightly different result (both are strongly supported). The two Idiopsar sierra finches have also been removed from Phrygilus. It might be better to put them in their own genus, sister to Idiopsar, but no genus name is available for them. The Geospizopsis sierra finches were formerly part of Phrygilus. Campagna et al. found them close to Haplospiza, and presumably Acanthidops. Note that all of the flower-piercers are in Diglossa, including those formerly placed in Diglossopis (see Mauck and Burns, 2009).

There may be a case for merging Acanthidops into Haplospiza. Weir et al. (2009) found that Acanthidops was more closely related to H. unicolor than to H. rustica. This is based on a single gene (cytochrome-b), so it is probably a bit premature to make the change.

Hemithraupini: Yellow-and-black Tanagers

All the members of this colorful group have been sequenced (see Barker et al., 2013; Burns et al., 2014; Weir et al. 2009). The Scarlet-and-white Tanager is sometimes put in a separate genus, Erythrothlypis. Weir et al. found it sister to the Black-and-yellow Tanager, so I leave them both in Chrysothlypis.

Tachyphonini: Ornamented Tanagers

Besides the complete analysis of Burns et al. (2014), other relevant papers include Burns and Racicot (2009), sorted out a big chunk of the Tachyphonini (which could equally be called Ramphocelini as both were established by Bonaparte in the same publication). Barker et al. (2013) found strong support for this clade, with mediocre support for including Volatinia, Creurgops, and Conothraupis as basal members. Note however that the two Creurgops show up next to the Poospzini in Weir et al. (2009), and in one of the two analyses in Burns et al. (2003).

The genus Tachyphonus was found to be paraphyletic, with part more closely related to Lanio, and part more closely related to Ramphocelus. The monotypic genus Eucometis has been merged into Trichothraupis, where it joins a couple of ex-Tachyphonus. Another monotypic genus, Rhodospingus, has joined Lanio, as has part of Tachyphonus.

Burns and Racicot did not consider whether or not the Lemon-rumped Tanager (R. flammigerus icteronotus) and Flame-rumped Tanager (R. f. flammigerus) should be split. They did not have samples from flammigerus, but they did find substantial genetic diversity within icteronotus. There is a hybrid zone in Colombia, which has been examined by Morales-Rozo et al. (2014). I don't see the case for a split.

Charitospizini: Coal-crested Finch

This tribe contains a single species, Coal-crested Finch. Barker et al. (2013) found that the Coal-crested Finch, Charitospiza eucosma did not belong with any of the other tribes. They put it in the middle of the Dacninae, but the node connecting it had quite low support (about 15% bootstrap). Burns et al. (2014) placed it on a separate branch, basal to a clade containing Dacini and the rest of the Dacinae.

Dacnini: Blue Tanagers

Dacinini includes the dacnises, the Cyanerpes honeycreepers, and the Swallow Tanager. The order within Dacnis is based on Burns et al. (2014).

Based on IOC and Ridgely and Greenfield (2001), Yellow-tufted Dacnis, Dacnis egregia (inc. aequatorialis) has been split from Black-faced Dacnis, Dacnis lineata.

Emberizoidini: Grassland Tanagers

This clade includes the grass and pampa finches. Embernagra and Emberizoides have grouped together in several studies. Barker et al. (2013) found that Coryphaspiza belongs with them to form a strongly supported clade. They also found some support for grouping them with the saltators, which is also supported by Burns et al. (2014).

Saltatorini: Saltators

Saltatricula is not particularly close to the Saltator saltators. The Black-throated Saltator has been moved to Saltatricula. The Rufous-bellied Saltator, “Saltator” rufiventris, is not really a saltator, and so has been moved to Dubusia in the Pipraeideini, as shown on the Thraupinae diagram. Finally, Pitylus has been merged into Saltator. Chaves et al. (2013) was the first analysis to include all of the saltator species. The order of the saltators reflects the results in Burns et al. (2014).

The Grayish Saltator has been split into three species based on Chaves et al. (2013). The three species are Middle American Saltator, Saltator grandis (all Middle American races); Plumbeous Saltator, Saltator plumbeus (presumed to include plumbeus and brewsteri); and Grayish Saltator, Saltator coerulescens (all other races). There may be more than three species here. In particular, plumage suggests that Brewster's may need to be separated from Plumbeous, but there is currently no genetic data.

Poospizini: Warbler-Tanagers

Poospizini is a big Neotropical finch group. The name Nephelornithini is equally applicable.

Until recently, the phylogentic picture here is clouded by the fact that neither Hemispingus nor Poospiza is monophyletic (see García-Moreno et al., 2001; Lougheed et al., 2000). Further, part of Phrygilus seems to be here too (see Klicka et al., 2007). The situation has improved with the appearance Shultz and Burns (2013), which has sorted out the various Hemispingus and Poospiza species. The result is that both Poospiza and Hemispingus are dismembered, with Poospiza spread across four different genera and Hemispingus across six. The generic name Hemispingus may disapper as a result.

The Gray-hooded Bush-Tanager (Cnemoscopus) is sister to a pair of hemispinguses, the Black-headed and Drab Hemispinguses. These are moved to the genus Pseudospingus (Berlepsch and Stolzmann 1896, type xanthophthalmus).

Although early indications were that the Cochabamba and Tucuman Mountain-Finches were separated from the other Poospiza, the more complete taxon sampling in Shultz and Burns (2013) shows that they are actually embedded in the main body of Poospiza. This means that the genus Compsospiza (Berlepsch 1893, type garleppi) must again be submerged in Poospiza (Cabanis 1847, type nigrorufa). Shultz and Burns suggest that their genetic results would support splitting Black-and-chestnut Warbling-Finch, P. whitii, from Black-and-rufous Warbling-Finch, Poospiza nigrorufa. The additional evidence from Jordan et al. (2017) is sufficient to justify the split. See also SACC Proposal 753.

The true Poospiza group is sister to the remaining Poospizini.

Based on García-Moreno et al. (2001, 2003), White-browed Hemispingus, Kleinothraupis auricularis, is split from Black-capped Hemispingus, Kleinothraupis atropileus. Although García-Moreno et al. were able to identify the main clades in the former Hemispingus, they weren't able to discern their relationships. They had trouble with the Gray-capped Hemispingus, Kleinothraupis reyi, which is the basal member of this Kleinothraupis group. The name Kleinothraupis was created by Burns et al. (2016) as there was not an available name for the group.

The Piura Hemispingus, Sphenopsis piurae, including macrophrys, is split from Black-eared Hemispingus, Sphenopsis melanotis. These two and frontalis form a small clade for which the name Sphenopsis (PL Sclater 1862, type frontalis) is available. Some authorities split Western Hemispingus (S. ochraceus) from Black-eared. What litte genetic data is available does not support this (García-Moreno et al., 2001).

Sphenopsis is sister to Thlypopsis, which has absorbed Pyrrhocoma and the Superciliaried Hemispingus. This creates a couple of nomenclatural problems which cannot be avoided. The first is that two of these species have the specific epithet ruficeps, the Chestnut-headed and Rust-and-yellow Tanagers. The Rust-and-yellow has priority (d'Orbigny and Lafresnay, 1837) over the Chestnut-headed (Strickland, 1844). However, there seem to be no junior names for the Chestnut-headed. Burns et al. (2016) have now created the name Thlypopsis pyrrhocoma for it. The other problem is that this group now contains the type species of three genera (Hemispingus, Pyrrhocoma, and Thlypopsis). All were named by Cabanis in the same publication in 1851, and none have priority over the others. A first reviser action is needed to resolve this. As most of these species have been in Thlypopsis, I will use that until the nomenclatural issue is resolved.

Another stray Poospiza is in a clade by itself which was named Castanozoster by Burns et al. (2016). Thus I refer to the Bay-chested Warbling-Finch as Castanozoster thoracicus. Two more Poospiza take the name Poospizopsis (Berlepsch 1893, type caesar). They are sister to Cypsnagra and Donacospiza. These three genera together are sister to Urothraupis, Nephelornis, and a collection of more Poospiza and a Hemispingus. The name Microspingus (Taczanowski 1874, type trifasciatus) is available for this group, which is at the end of the Poospizini.

Beldsoe (1988) showed that Urothraupis belongs with the tanagers, but gives little guidance as to where in the tanagers. It has sometimes been considered close to the Chlorospingus bush tanagers, but that is incorrect since they are not tanagers!

Coeribini: Dome-nesting Tanagers

The Coerebini are not the old honeycreeper family, but are a clade of mainly island species including West Indian “quits” and bullfinches, and Darwin's Finches. The term Tholospiza (dome finch) was introduced by Burns et al. (2002) to avoid confusion with the old honeycreeper family (Coeribidae), but Coeribini has priority.

I've changed several of the generic boundaries in Coeribini to reflect the genetic tree found by Burns et al. (2014). The Barker et al. (2013) version is similar. The Puerto Rican and Greater Antillean Bullfinches move from Loxigilla to join the Cuban Bullfinch in Melopyrrha. Only the Yellow-faced Grassquit remains in Tiaris. The rest of Tiaris moves to Loxigilla, as does the St. Lucia Black Finch (formerly the only member of Melanospiza). The Dull-colored and Sooty Grassquits are in a clade of their own, named Asemospiza by Burns et al. (2016).

The Grand Cayman Bullfinch, Melopyrrha taylori, has been split from the Cuban Bullfinch, Melopyrrha nigra (see Garrido et al., 2014). They differ in plumage, morphology, size, and song.

Darwin's finches are quite closely related and introgression makes it hard to discern the actual relationships between them. Previous versions wereh based on Petren et al. (2005), Tonnis et al. (2005), and the discussion in Grant and Grant (2008). After considering Farrington et al. (2014) and Lamichhaney et al. (2015), I have rearranged Darwin's Finches. I also merged Camarhynchus into Geospiza, and split some taxa.

This leads to a net gain of 3 species. However, there is a case to be made for lumping conirostris into magnirostris, propinqua into scandens, and acutirostris into fortis. If the distribution table below is correct, each pair is allopatric, and the divergence times fairly short. The three pairs occupy the last 6 rows of the table. The full tree is shown in the species list.

Distribution of Darwin's Finches in the Galápagos Islands
Species Sey Bal Isa Fer Stg Rab Pnz SCrz Dap SFe SCrst Esp Flo Gen Mar Pnt Dar Wol
C. fusca B B B E B B B B B
C. olivacea B B B B B B
Pl. crassirostris B B B (E) E B (E) B B B B
G. difficilis (E) B B E (E) E B
G. psittacula B B B (E) E B B B B B
G. pauper B
G. parvula B B B (E) B B B B B
G. heliobates B E
G. pallida B (B) B B B B
G. septentrionalis B B
G. fuliginosa B B B B B B B B B B B B B B B
G. fortis B B B B B B B B B B B B B B
G. acutirostris B
G. magnirostris B B B B B B B B E E B B B B B
G. conirostris B
G. scandens B B B B B (B) B B B B B B B
G. propinqua B

Sporophilini: Seedeaters

Mason and Burns (2013) showed that both Dolospingus and Oryzoborus are embedded in Sporophila, as expected. As a result, both Dolospingus and Oryzoborus have been merged into Sporophila. The arrangement of the Sporophilini now follows Mason et al. (2016).

Following an SACC decision, Capped Seedeater, Sporophila bouvreuil, has been split into Pearly-bellied Seedeaster, Sporophila pileata, and Copper Seedeater, Sporophila bouvreuil. See Machado and Silveira (2010; 2011). Further, the yellow-billed Tropeiro Seedeater, Sporophila beltoni, has been split from the dark-billed Plumbeous Seedeater, Sporophila plumbea (Repenning and Fontana, 2013).

Based on Mason et al. (2016), the White-collared Seedeater has been split into Cinnamon-rumped Seedeater, Sporophila torqueola (Western and Central Mexico) and Morellet's Seedeater / White-collared Seedeater, Sporophila morelleti (the rest of the range). These are not even sister taxa.

Dacninae Species List

Nemosiini: Flock-dwelling Tanagers Bonaparte, 1854

Conirostrini: Conebills Edwards, 1986

Diglossini: Highland Tanagers P.L. Sclater, 1875

Hemithraupini: Yellow-and-black Tanagers Sundevall, 1872

Tachyphonini: Ornamented Tanagers Bonaparte, 1853

Charitospizini: Coal-crested Finch Burns et al., 2014

Dacnini: Blue Tanagers Sundevall, 1836

Emberizoidini: Grassland Tanagers Burns et al., 2014

Saltatorini: Saltators Bonaparte, 1853

Poospizini: Warbler-Tanagers Wolters, 1980

Coerebini: Dome-Nesting Tanagers d'Orbigny & Lafresnaye, 1838

Sporophilini: Seedeaters Ridgway, 1901 (1853)

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