Emberizoidae III

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

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 Emberizoidae.

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

11 genera, 50 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 and both Amaurospiza and Cyanocompsa into Cyanoloxia.

The species flagged in blue lack molecular data, but are believed to be in the proper genus, although I wouldn't be terribly surprised if the yellow grosbeaks ended up in a slightly different spot.

Three species of Hepatic Tangers are recognized here: Lowland Hepatic-Tanager / Red Tanager, Piranga flava, Northern Hepatic-Tanager / Hepatic Tanager, Piranga hepatica, and Highland Hepatic-Tanager / Tooth-billed Tanager, Piranga lutea. These have frequently been considered distinct species and are so treated in many regional guidebooks. Burns (1998) found genetic differences between the three in the range typical of species, not subspecies. The tepui form haemalea remains part of lutea due to insufficient evidence.

Thraupidae: Tanagers Cabanis, 1847

94 genera, 374 species HBW-16

Thraupidae tree
Thraupidae family tree

We finally reach the last family on the list! It may not stay that way. The tanagers are the subject of an extensive ongoing revision that may ultimately split them into several families....maybe.

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 that are often considered sparrows mostly end up in the tanagers. They also gain the saltators and some other cardinals.

The result of this shifting genera is a very large, very heterogeneous tanager family. Further structure is required to organize this heterogenity. At the very least, the Thraupidae should be divided into subfamilies.

Right now, it is not clear exactly how that should be done, and the latest paper (Barker et al., 2013) suggests that it will not be easy to sort out. If we try to sort out the monophyletic groups, we end up 13-16 clades. I think 16 of them should be distinguished, and I rank them as tribes. Most of these tribes are easily identified in Barker et al., (2013), with one hidden due to a node with 18-21% bootstrap support. Most are also easily visible in the comprehensive analyses of Klicka et al., (2007) and Weir et al. (2009) Most of the tribes can also be identified by combining the more restricted analyses of Burns (1997), Burns et al. (2003) and Yuri and Mindell (2002).

Most of these tribes, or their cores, are either strongly supported in Barker et al. (2013). In the major case where they are not, the species-level detailed analysis (e.g., Sedano and Burns, 2009) makes them clear.

The tribes are real. Unfortunately, there is little solid information about how they relate to one another. The various relatively comprehensive analysis give conflicting results, and usually have weak support. Barker et al. (2013) is a champion in the weak support department, with many nodes in the Dacninae receiving single digit support values (out of 100).

Making do with the information available, I've divided the tribes into three groups, which I rank as subfamilies: Saltatorinae, Thraupinae, and Dacninae. I expect this to be revised at some point, but given the amount of data Barker et al. (2013) threw at the problem, I'm not expecting the deeper part of the tanager tree to be well-understood soon.

There is huge uncertainty about the saltators. Barker et al. (2013) place them in the Dacninae clade, but with weak support. Klicka et al. (2007) found they were a well-supported clade and placed them basally in the tanagers. In their broad analysis, Weir et al. (2009) found Saltator basal in Dacninae and Saltatricula basal in the Cardinalidae. A second analysis, focusing on the tanagers, ended up with the saltators forming a basal clade. In view of all this, I'm leaving them rather uncertainly in the basal position among the tanagers.

Support for the other two groups is not strong either. Barker et al. (2013) show about 50% bootstrap support for the core Thraupinae, but it drops to about 10% when the Incazpizini are included. They don't separate Dacninae, but Dacninae plus Saltatorinae gets 7% support. Nonetheless, except for the saltators, the same division was found by Klicka et al. (2007) and Weir et al. (2009).

Due to the lack of any credible arrangement of the tribes in Dacninae, I've arranged them by size, least speciose to most speciose. The situation in the Thraupinae is somewhat better due to Sedano and Burns (2009). However, the results from Barker et al. (2013) suggest some skepticism is due here too.

Thraupidae Subfamilies

Saltatorinae: Saltators Bonaparte, 1853

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.

Saltatricula is not particularly close to the Saltator saltators. The Black-throated Saltator has been moved to Saltatricula. Although I don't have an alternative name for it, the Rufous-bellied Saltator, “Saltator” rufiventris, is not really a saltator, and so has been moved to the Thraupini, as shown on the Thraupinae diagram. Finally, Pitylus has been merged into Saltator.

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 three main groups: Incaspizini, Coerebini, and the Thaupini, Cissopini, Pipraeideini clade. 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).

It's not at all clear that the Incaspizini belong in the Thraupinae. The only real evidence for it is from Barker et al. (2013), and the support for this placement is quite low, although support for the clade itself is high. They could actually be basal in Dacninae, or even more basal in Thraupidae.

Incaspizini

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. 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).

Coeribini

I've changed several of the generic boundaries in Tholospiza to reflect the genetic tree found by Burns et al. (2002). 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). Darwin's finches are quite closely related and introgression makes it hard to discern the actual relationships between them. I've paid particular attention to the microsatellite results of Petren et al. (2005). See also Tonnis et al. (2005), and the discussion in Grant and Grant (2008). The full tree is shown in the species list.

The current tree for the remainder of the Thraupinae is based largely on the fairly complete phylogeny provided by Sedano and Burns (2010). There are some differences in the Barker et al. (2013) tree, but these differences are poorly supported. Moreover, Sedano and Burns have much better taxon sampling. The results are not unexpected. There is a lot of overlap with Burns and Naoki (2004), and some of the remaining changes had been rumored for a while.

Thraupini

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 unlabelled clade consists of the 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.

Cissopini

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.

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.

Pipraeideini

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 Sedano and Burns (2010) 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. Most of Buthraupis has moved to Chlorornis and Cnemathraupis, with B. wetmorei joining Anisognathus). Anisognathus has also absorbed the Blue-capped Tanager, formerly Thraupis cyanocephalus. Pipraeideini also includes the Rufous-belled Saltator, which needs a new genus name.

Thraupinae Species List

Incaspizini: Inca Finches and allies Informal

Coerebini: Tholospiza, the Domed-Nest Clade d'Orbigny & Lafresnaye, 1838

Thraupini: Tangara Tanagers Cabanis, 1847

Cissopini Sundevall, 1872

Pipraeideini: Mountain-Tanagers Informal

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 brush-finches. 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.

This section of the tree has been less studied than the Thraupinae, and presents a murkier picture. It's rather like a jigsaw puzzle, but with some of the pieces missing. The situation is gradually improving. Most of the pieces that are here can be found in Burns (1997), Yuri and Mindell (2002), Burns et al. (2003), Klicka et al. (2007), Burns and Racicot (2009), Weir et al. (2009), Shultz and Burns (2013), and Barker et al. (2013).

Since the large-scale phylogenies have substantial disagreement and low support, I've arranged the tribes according to size with the smallest first.

Charitospizini

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), so we don't really know where it goes.

Nemosiini

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).

Emberizoidini

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, although I have doubts about whether that is correct.

Hemithraupini

Except for two of the Hemithraupis tanagers, all the members of this colorful group have been sequences (see Barker et al., 2013; Weir et al. 2009). Barker et al. found that Orchesticus, Parkerthraustes, and the flashy Plushcap (Catamblyrhynchus) are basal members of the Hemithraupini, albeit with mediocre support. 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.

Dacnini

Dacinini includes the dacnises, the Cyanerpes honeycreepers, and the Swallow Tanager. The order within Dacnis is based on Weir et al. (2009).

Tachyphonini

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. They suggest the lumping Ramphocelus-Lanio group into one or two genera. The modifications here are slightly less drastic, leaving three genera. The monotypic genera Eucometis, Rhodospingus, and Trichothraupis have been merged into Lanio, as has Coryphospingus and 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.

Sporophilini

The position of Sporophilini is particularly uncertain. It sometimes shows up as basal to Dacninae and Thraupinae combined. The order of the seedeaters is uncertain, and generic limits will almost certainly be changed eventually (see Lijtmaer et al., 2004; Weir et al., 2009). One option would be to merge both Dolospingus and Oryzoborus into Sporophila.

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).

Poospizini

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 Burn suggest that their genetic results would support, but do not compel, splitting Black-and-chestnut Warbling-Finch, P. whitii, from Black-and-rufous Warbling-Finch, Poospiza nigrorufa. The true Poospiza group is sister to the remaining Poospizini.

Based on García-Moreno et al. (2001, 2003), White-browed Hemispingus, “Hemispingus” auricularis, is split from Black-capped Hemispingus, “Hemispingus” 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, “Hemispingus” reyi, which is the basal member of this “Hemispingus” group. There does not seem to be an available name for the group, so I use “Hemispingus” for now.

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 1861, 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. For now, I distinguish it as “ruficeps”. 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 and has no available name. Thus I refer to the Bay-chested Warbling-Finch as “Poospiza” thoracica. 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!

Conirostrini

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.

Diglossini

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. Resolving this clade requires a little guesswork. Rowettia and Nesospiza are closely related to Sicalis (Ryan et al., 2005, supplement). However, Ryan et al. did not include Melanodera or any of the core Phrygilus species. Barker et al. (2013) was missing any of the true Phrygilus sierra finches. Campagna et al. found the true Phrygilus finches and Melanodera to be sister genera and also closely related to Sicalis. But Campagna et al. did not include Rowettia and Nesospiza.

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.

Dacninae Species List

Charitospizini Informal

Nemosiini Bonaparte, 1854

Emberizoidini Informal

Hemithraupini Sundevall, 1872

Dacnini Sundevall, 1836

Tachyphonini Bonaparte, 1853

Sporophilini Ridgway, 1901 (1853)

Poospizini Wolters, 1980

Conirostrini Edwards, 1986

Diglossini P.L. Sclater, 1875

Previous Page