Thraupid Group

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 44 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Thraupid Group

Cardinalidae: Cardinals, Grosbeaks

11 genera, 50 species HBW-16

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

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

96 genera, 370 species HBW-16

Thraupidae 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, and they are losing Habia and Piranga (including all of the North American tanagers) to the cardinals. You'll notice that I've also removed Mitrospingus and Lamprospiza. 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. I think they are too heterogeneous. At the very least, the Thraupidae should be divided into subfamilies. Right now, it is not clear exactly how that should be done. If we try to sort out the monophyletic groups, we end up with 15 or so clades by combining Burns (1997), Burns et al. (2003), Yuri and Mindell (2002), and Klicka et al., (2007). Some are candidates for family status.

This version of the list takes a more modest approach, dividing the tanagers into three subfamilies. The saltators form a basal clade. Except for some species of unknown affinities, the remaining tanagers can usefully be broken into two large groups: Thraupinae and Dacninae, with a few left over as Incertae sedis. By abuse of notation, the identifiable clades are treated as tribes. For now, my working hypothesis is that the Thraupidae are organized according to the tree above. As more becomes known, there will doubtless be some rearrangement and (hopefully) consolidation. One or two tribes may even move to a different subfamily.

As time goes on, the three subfamily choice is looking better. Sedano and Burns (2010) have recently finished a more comprehensive examination of the Thraupinae. It fits comfortably into this framework, and provides supporting detail down to the species level.

Thraupidae Species List

Saltatorinae: Saltators

The saltators come first. Saltatricula is not particularly close to the other 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

Thraupinae tree The Thraupinae are shown in more detail at the right. The Coerebini are not the old honeycreepers, 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).

I've changed several of the generic boundaries in Tholospiza to reflect the genetic tree found by Burns et al. (2002). 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 below.

The current tree for the remainder of the Thraupinae is based largely on the fairly complete phylogeny provided by Sedano and Burns (2010). 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.

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.

Continuing down the tree, I have retained Chlorochysini as a tribe rather than merge it into Cissopini. This is because the support Sedano and Burns found for a sister relationship with Cissopini is a just bit weaker than the other major divisions in their tree. Chlorochysini includes only two genera, Chlorochrysa and the former cardinal Parkerthraustes. If they are merged, they would use the established name Cissopini rather than the informal name Chlorochysini.

The Paroria and Gubernatrix cardinals (often considered sparrows) are in Cissopini with several other tanagers and finches. No DNA information is available concerning Gubernatrix, so it's position here is more uncertain. The species limits of Paroaria are a bit non-standard. Based on Dávalos et al. (2009), I've separated Masked Cardinal, Paroaria nigrogenis from Red-capped Cardinal and merged Yellow-billed Cardinal into Red-capped Cardinal, as suggested many years ago by Hellmayr (1938). Thus Red-capped Cardinal includes the subspecies gularis, cervicalis, capitata, and fuscipes.

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. I've merged most of Buthraupis into Chlorornis, 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.

Coerebini: Tholospiza, the Domed-Nest Clade

Thraupini: Tangara Tanagers

Chlorochrysini

Cissopini

Pipraeideini: Mountain-Tanagers

Dacninae

Dacninae 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 even the studied parts present a murkier picture. It's rather like a jigsaw puzzle, but with some of the pieces missing. Most of the pieces that are here can be found in Burns (1997), Yuri and Mindell (2002), Burns et al. (2003), and Klicka et al. (2007).

This version of Dacninae starts with the Dacnini, the dacnises and most honeycreepers. Evidence on whether they belong here is mixed. Alternatives include grouping them with Nemosiini and Hemithraupini.

The Dacnini are followed by the grass- and pampa-finches, Emberizoidini. After that comes a big neotropical finch group, Poospizini. The phylogentic picture here is clouded by the the fact that neither Hemispingus nor Poospiza is monophyletic (see García-Moreno et al., 2001; Lougheed et al., 2000). Further, part of Phrygilus is here too. It's not altogether clear how to straighten them out. Doubly so as there are no papers that provide a comprehensive view of the group.

The name Corydospiza seems to apply to the Phrygilus refugees, while Compsospiza is available for two of the Poospiza. Creurgops seems to float around in the various papers. One plausible location is after Compsospiza, sister to the remaining Poospizini. Another part of Hemispingus groups with Cnemoscopus. I couldn't find any availble names for it. Cypsnagra though the true Hemispingus is likely sister to the rest of Poospiza (which may still need some adjustment). The remaining finches in Poospizini are often considered close to Poospiza.

The rest of Dacninae is in a bit better shape, although it's still not entirely clear how the clades fit together. While Nemosiini and Hemithraupini are both well-supported in Burns et al. (2003), that they are sisters is only weakly supported. Burns and Racicot (2009) sorted out a big chunk of the Tachyphonini. 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 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.

The position of Sporophilini is rather uncertain. It sometimes shows up as basal to Dacninae and Thraupinae combined. The order of the seedeaters is uncertain, and generic limits will probably be changed eventually (see Lijtmaer et al., 2004).

The conebills are in their own clade, Conirostrini. That brings us to another group of neotropical finches, Diglossini. The Geospizopsis sierra-finches were formerly part of Phrygilus. Not surprisingly, Acanthidops and Haplospiza group together, near the flower-piercers (Diglossa and Diglossopis). I couldn't find a name for the next group of Phrygilus, which are followed by the true Phygilus. Rowettia and Nesospiza are closely related to Sicalis (Ryan et al., 2005, supplement). Ridgely and Tudor (1989) suggest Melanodera and possibly Phrygilus are close relatives, so I've grouped the first three together, next to the true Phrygilus.

Dacnini

Emberizoidini

Poospizini

Nemosiini

Hemithraupini

Tachyphonini

Sporophilini

Conirostrini

Diglossini

Incertae sedis: Thraupidae

At the end of the list are 9 tanager species I have trouble putting in a definite place on the tree.

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! The Plushcap, which has often been placed in its own family, seems to belong in Dacninae somewhere (Yuri and Mindell, 2002), but pinning it down is a problem. Although some suspect it is also not a tanager, Seutin and Bermingham (1997) indicate Rhodinocichla belongs somewhere in the Dacninae.

I have little information about where to put Cyanicterus, Orchesticus, or Orthogonys. I suspect they belong in Dacninae. Compsothraupis is sometimes included in Sericossypha, but Storer argued their similarities are convergence, a view that was endorsed by Ridgely and Tudor (1989).

Finally, Charitospiza and Coryphaspiza might be tanagers, or might be sparrows.

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