Emberizoidae I

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 44 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Emberizidae: Buntings

1 genus, 42 species HBW-16

Emberizidae tree
Click for species-level tree
for Emberizidae

There is some uncertainty concerning whether the Emberiza buntings and American sparrows are sister clades, and that is why I have them in separate families. For example, Klicka et al. (2003) has Emberizidae embedded in the Icteridae while Passerellidae is sister to Parulidae. Taxonomy within Emberizidae follows Alström et al. (2008a), with Melophus and Latoucheornis included in Emberiza. They identify four major clades, which could be regarded as subgenera: Fringillaria, Melophus, Emberiza, and Cynchramus.

Should anyone feel inclined to further divide this genus, there are available names for the main subclades of these clades. For one such possibility, see the species-level tree for Emberiza. Although there are some differences, the groupings are in general agreement with the morphological groups of Byers et al. (1995). The question marks on the tree indicate species not analyzed by Alström et al. (2008).

Fringillaria group
Melophus group
Emberiza group
Cynchramus group

Passerellidae: American Sparrows

33 genera, 127 species HBW-16 (split)

Passerellidae tree
Click for genus-level tree
for Passerellidae

The sparrow family has been carved up in the ongoing reorganization of the nine-primaried oscines. Most of the neotropical finches have joined the tanagers. The Gubernatrix and Paroaria cardinals have been moved to the tanagers. The neotropical finches that remain are the Atlapetes brush-finches, as well as the Large-footed, Yellow-thighed, and Yellow-green Finches. Although I've moved them to the tanagers near Poospiza, it remains uncertain whether the Inca-Finches are sparrows or tanagers. In return, the sparrows gain the Chlorospingus bush-tanagers and the Tanager Finch. That alone would reduce the family to about half its former size, but there is more. They also lose the Emberiza.

What remains is a somewhat more homogeneous family and it is possible to give it a coherent organization. Everything seems to fall into 8 clades, which I've treated at the tribe level. I haven't used subfamilies partly because I don't think it's clear yet how the tribes fit together, and partly because I don't think the divisions are particularly deep.

We start with the sister tribes Chlorospingini and Ammodramini. We follow the topology from DaCosta et al. (2009), where Chlorospingini + Ammodramini are sister to the remaining Passerellidae. It is not entirely clear whether this is correct. One alternative is in Klicka et al. (2007), where Ammodramini branches basally, followed by Chlorospingini. Another alternative, based on Carson and Spicer (2003), would be to put Ammodramini, Chlorospingini, and Spizellini in one subfamily and the remaining Passerellidae in another. This arrangement also receives some support from Yuri and Mindell (2002).

The treatment of the Common Bush-Tanager complex is based on García-Moreno et al. (2004), Sánchez-González et al. (2007), Bonaccorso et al. (2008), and Weir et al. (2008). This involves breaking up the Common Bush-Tanager into 9 species. Four of these are primarily Mexican: White-fronted Bush-Tanager, C. albifrons; Wetmore's Bush Tanager, C. wetmorei; Brown-headed Bush-Tanager, C. ophthalmicus; and Dwight's Bush-Tanager, C. dwighti. The color plate in Sánchez-González et al. (2007) illustrates these forms as well as Dusky-headed Bush-Tanager, C. postocularis. Other taxa that appear to deserve species status are the Central American Dotted Bush-Tanager, C. punctulatus; the Buff-breasted Bush-Tanager, C. cinereocephalus, of Peru; and the Yellow-breasted Bush-Tanager, C. flavopectus. The list of subspecies below presumes that Isler and Isler's (1987) flavopectus group stays together, although only two of its subspecies were analyzed by Weir et al. (2008). I've also presumed that Isler and Isler's venezuelanus group stays together. Weir found that some of them group with several more southern races. For the present, it seems reasonable to put the whole lot of them in a single species and call it Common Bush-Tanager, C. venezuelanus. There may still be additional species hiding within the Common Bush-Tanager complex.

The Common Bush-Tanager Complex
Species Subspecies
Wetmore's Bush-Tanager wetmorei
White-fronted Bush-Tanager albifrons, persimilis
Brown-headed Bush-Tanager ophthalmicus
Dwight's Bush-Tanager dwighti
Dusky-headed Bush-Tanager postocularis, honduratius
Dotted Bush-Tanager punctulatus, regionalias, ‘novicius’*
Common Bush-Tanager venezuelanus*, jacqueti, falconensis*, ponsi*, eminens*, peruvianus, bolivianus, fulvigularis, argentinus
Buff-breasted Bush-Tanager cinereocephalus
Tacarcuna Bush-Tanager tacarcunae
Pirre Bush-Tanager inornatus
Yellow-breasted Bush-Tanager flavopectus*, trudis*, exitelis*, macarenae*, nigriceps*, phaeocephalus, hiaticolus
Dusky Bush-Tanager semifuscus, livingstoni*
Subspecies based on Dickinson et al. (2003).
*An asterisk indicates taxa not sampled by Weir et al. (2008). The subspecies ‘novicius’ may be a hybrid form.

As for Ammodramini, some genera have required reorganization. Ammodramus and Aimophila have been affected the most. The papers by Klicka and Spellman (2007) and DaCosta et al. (2009) show what to do. Ammodramus itself is reduced to the Grasshopper Sparrow together with a couple of neotropical relatives. These are sister to Arremonops. Together, they are sister to a big chunk of what was Aimophila. As the Aimophila type species is not included, I have revived Audubon's 1839 name for them, Peucaea (type aestivalis). Two more of the former Aimophila are sister to the rest of Ammodramini. Following DaCosta et al.'s suggestion, they take the genus name Rhyncospiza (Ridgway 1898, type stolzmanni).

The next sparrow tribe is the Spizellini. As shown by Carson and Spicer (2003), this clade includes most of the Spizella. Within Spizellini, the position of the Zapata Sparrow is not completely clear. It's thought to be sister to Amphispiza, but genetic data is not available. This version of Amphispiza includes the Five-striped Sparrow and the Black-throated Sparrow, but not the Sage Sparrow. The arrangement within Spizella follows Canales-Del Castillo et al. (2010), with Brewer's Sparrow somewhat surprisingly sister to Worthen's Sparrow. The Timberline Sparrow, Spizella breweri taverneri, is sometimes thought to be a separate species. Although there is some differentiation between it and Brewer's Sparrow, it may be best thought of as an incipient species within Brewer's Sparrow. See Klicka et al. (1999).

Our next tribe consists of a group of neotropical sparrows. The genera Lysurus and Buarremon are sometimes merged into Arremon, especially after genetic data showed that the old arrangement was wrong. I thought it made a bit more sense to transfer two of the Buarremon to Lysurus. That leaves only torquatus in Burarremon, but it is a fairly distant relative of the narrowly circumscribed Arremon.

The Junco/Zonotrichia clade Passerellini is next. It is now clear that the American Tree Sparrow is not part of Spizella. Rather, it is most closely related to the Fox Sparrow, Passerella iliaca (Carson and Spicer, 2003). Accordingly, I've moved it to genus Passerella, which forms a clade with the juncos and Zonotrichia. Passerella iliaca itself remains controversial, and may end up being split into four species.

The rest of Ammodramus ends up in the Passerculini. The “marshland” Ammodramus form a clade. As Klicka and Spellman (2007) recommend, they get the name Ammospiza. Although Henslow's and Baird's Sparrows may be each other's closest relatives, it is also possible they are not. I've resurrected the old generic names Centronyx (for Baird's) and Nemospiza (for Henslow's). The Sage Sparrow gets a brand new genus name. Klicka and Spellman (2007) discovered that it is not related to the other Amphispiza and proposed the genus name Artemisospiza for it (think sagebrush, Artemesia).

DaCosta et al. (2009) resolved a big chunk of the uncertainty concerning the Pipilonini. The Melozone Ground-Sparrows end up in two (or three?) separate groups. One part is sister to the brown towhees. DaCosta et al. suggest the genus name Pyrgisoma for it. The AOU proposal suggested the type is kieneri, although Ridgway, gives the type as biarcuata. Apparently, treating kieneri as type dates from a time when biarcuata was considered a subspecies of kieneri. However, when separate, kieneri becomes the type species of Kieneria (Bonaparte 1855) and biarcuata is the type of Pyrgisoma. Since biarcuata ends up in the genus Melozone, Kieneria is the name used here. The brown towhees have always been considered different, and there have been suggestions they should get their own genus. That has never happened, so my best option seems to be to put them into Kieneria too.

The other part of Melozone retains the name, although it is unclear whether the two species are actually sisters (looking at them, it is hard to believe they are not sisters). These species form a clade with the remaining Aimophila. How these two clades relate to the Large-footed Finch, Pezopetes capitalis, remains unclear. It is closely related, and may be basal to both, or maybe not. This is left unresolved here.

The rest of the Pipilonini consists of the Pipilo towhees together with the Atalapetes brush-finches and allies. The Pselliophorus finches belong here, and appear to be embedded in Atlapetes. It has also been suggested that the Tanager Finch, Oreothraupis arremonops, is part of Atlapetes. At present there is no good comprehensive molecular phylogeny for Atlapetes, so I have not made significant changes to the linear order here.

As you can see, it's been possible to put together reasonable species trees for most of the sparrow tribes.

Chlorospingini: Bush-Tanagers

Ammodramini

Spizellini

Arremonini: Scrub Sparrows

Passerellini: Juncos and allies

Passerculini

Pipilonini: Towhees

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