Emberizidae: Buntings
1 genus, 42 species HBW-16
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.
Fringillaria group
- Brown-rumped Bunting, Emberiza affinis
- Golden-breasted Bunting, Emberiza flaviventris
- Socotra Bunting, Emberiza socotrana
- Somali Bunting, Emberiza poliopleura
- Cabanis's Bunting, Emberiza cabanisi
- Cape Bunting, Emberiza capensis
- Lark-like Bunting, Emberiza impetuani
- Cinnamon-breasted Bunting, Emberiza tahapisi
- Striolated Bunting, Emberiza striolata
- House Bunting, Emberiza sahara
Melophus group
- Crested Bunting, Emberiza lathami
- Black-headed Bunting, Emberiza melanocephala
- Red-headed Bunting, Emberiza bruniceps
Emberiza group
- Corn Bunting, Emberiza calandra
- Chestnut-eared Bunting, Emberiza fucata
- Jankowski's Bunting, Emberiza jankowskii
- Meadow Bunting, Emberiza cioides
- Rock Bunting, Emberiza cia
- Godlewski's Bunting, Emberiza godlewskii
- Gray-necked Bunting, Emberiza buchanani
- Cinereous Bunting, Emberiza cineracea
- Ortolan Bunting, Emberiza hortulana
- Cretzschmar's Bunting, Emberiza caesia
- Cirl Bunting, Emberiza cirlus
- White-capped Bunting, Emberiza stewarti
- Yellowhammer, Emberiza citrinella
- Pine Bunting, Emberiza leucocephalos
Cynchramus group
- Slaty Bunting, Emberiza siemsseni
- Yellow-throated Bunting, Emberiza elegans
- Ochre-rumped Bunting / Japanese Reed-Bunting, Emberiza yessoensis
- Pallas's Bunting / Pallas's Reed-Bunting, Emberiza pallasi
- Reed Bunting / Common Reed-Bunting, Emberiza schoeniclus
- Little Bunting, Emberiza pusilla
- Rustic Bunting, Emberiza rustica
- Yellow-breasted Bunting, Emberiza aureola
- Chestnut Bunting, Emberiza rutila
- Tibetan Bunting, Emberiza koslowi
- Yellow Bunting, Emberiza sulphurata
- Black-faced Bunting, Emberiza spodocephala
- Yellow-browed Bunting, Emberiza chrysophrys
- Tristram's Bunting, Emberiza tristrami
- Gray Bunting, Emberiza variabilis
Passerellidae: American Sparrows
33 genera, 119 species HBW-16 (split)
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| Click for genus-level tree for Passerellidae |
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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 internal arrangement of the Chlorospingini is rather clear. At least it's clear if you use the current species limits. However, these limits may not be correct. In particular, there may be additional species hiding within the Common Bush-Tanager (Weir et al, 2008).
Needless to say, 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 Klicka and Spellman (2007), 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 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. 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. According to AOU, the type is kieneri, although Ridgway, gives the type as biarcuata. I'll follow AOU on this, but wonder why the difference. 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 Pyrgisoma 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
- Ashy-throated Bush-Tanager, Chlorospingus canigularis

Click for Passerellidae I tree:
Chlorospingini &
Ammodramini
- Short-billed Bush-Tanager / Yellow-whiskered Bush-Tanager, Chlorospingus parvirostris
- Yellow-throated Bush-Tanager, Chlorospingus flavigularis
- Sooty-capped Bush-Tanager, Chlorospingus pileatus
- Common Bush-Tanager, Chlorospingus ophthalmicus
- Tacarcuna Bush-Tanager, Chlorospingus tacarcunae
- Yellow-green Bush-Tanager, Chlorospingus flavovirens
- Pirre Bush-Tanager, Chlorospingus inornatus
- Dusky Bush-Tanager, Chlorospingus semifuscus
Ammodramini
- Tumbes Sparrow, Rhynchospiza stolzmanni
- Stripe-capped Sparrow, Rhynchospiza strigiceps
- Grasshopper Sparrow, Ammodramus savannarum
- Grassland Sparrow, Ammodramus humeralis
- Yellow-browed Sparrow, Ammodramus aurifrons
- Black-striped Sparrow, Arremonops conirostris
- Green-backed Sparrow, Arremonops chloronotus
- Olive Sparrow, Arremonops rufivirgatus
- Tocuyo Sparrow, Arremonops tocuyensis
- Cinnamon-tailed Sparrow, Peucaea sumichrasti
- Rufous-winged Sparrow, Peucaea carpalis
- Botteri's Sparrow, Peucaea botterii
- Cassin's Sparrow, Peucaea cassinii
- Bachman's Sparrow, Peucaea aestivalis
- Stripe-headed Sparrow, Peucaea ruficauda
- Black-chested Sparrow, Peucaea humeralis
- Bridled Sparrow, Peucaea mystacalis
Spizellini
- Lark Sparrow, Chondestes grammacus

Click for Passerellidae II tree:
Spizellini, Arremonini,
Passerellini - Lark Bunting, Calamospiza melanocorys
- Zapata Sparrow, Torreornis inexpectata
- Black-throated Sparrow, Amphispiza bilineata
- Five-striped Sparrow, Amphispiza quinquestriata
- Clay-colored Sparrow, Spizella pallida
- Chipping Sparrow, Spizella passerina
- Brewer's Sparrow, Spizella breweri
- Black-chinned Sparrow, Spizella atrogularis
- Field Sparrow, Spizella pusilla
- Worthen's Sparrow, Spizella wortheni
Arremonini: Scrub Sparrows
- Sooty-faced Finch, Lysurus crassirostris
- Olive Finch, Lysurus castaneiceps
- Chestnut-capped Brush-Finch, Lysurus brunneinucha
- Green-striped Brush-Finch, Lysurus virenticeps
- Stripe-headed Brush-Finch, Buarremon torquatus
- Saffron-billed Sparrow, Arremon flavirostris
- Pectoral Sparrow, Arremon taciturnus
- Half-collared Sparrow, Arremon semitorquatus
- Sao Francisco Sparrow, Arremon franciscanus
- Black-capped Sparrow, Arremon abeillei
- Orange-billed Sparrow, Arremon aurantiirostris
- Golden-winged Sparrow, Arremon schlegeli
Passerellini: Juncos and allies
- American Tree Sparrow, Passerella arborea
- Fox Sparrow, Passerella iliaca
- Volcano Junco, Junco vulcani
- Dark-eyed Junco, Junco hyemalis
- Yellow-eyed Junco, Junco phaeonotus
- Rufous-collared Sparrow, Zonotrichia capensis
- Harris's Sparrow, Zonotrichia querula
- White-throated Sparrow, Zonotrichia albicollis
- White-crowned Sparrow, Zonotrichia leucophrys
- Golden-crowned Sparrow, Zonotrichia atricapilla
Passerculini
- Vesper Sparrow, Pooecetes gramineus

Click for Passerellidae III tree:
Passerculini & Pipilonini - Sage Sparrow, Artemisospiza belli
- Le Conte's Sparrow, Ammospiza leconteii
- Seaside Sparrow, Ammospiza maritimus
- Nelson's Sparrow, Ammospiza nelsoni
- Saltmarsh Sparrow, Ammospiza caudacutus
- Striped Sparrow, Oriturus superciliosus
- Savannah Sparrow, Passerculus sandwichensis
- Baird's Sparrow, Centronyx bairdii
- Henslow's Sparrow, Nemospiza henslowii
- Sierra Madre Sparrow, Xenospiza baileyi
- Song Sparrow, Melospiza melodia
- Lincoln's Sparrow, Melospiza lincolnii
- Swamp Sparrow, Melospiza georgiana
Pipilonini: Towhees
- Large-footed Finch, Pezopetes capitalis
- Rusty-crowned Ground-Sparrow, Pyrgisoma kieneri
- Canyon Towhee, Pyrgisoma fusca
- White-throated Towhee, Pyrgisoma albicollis
- California Towhee, Pyrgisoma crissalis
- Abert's Towhee, Pyrgisoma aberti
- White-eared Ground-Sparrow, Melozone leucotis
- Prevost's Ground-Sparrow, Melozone biarcuata
- Rusty Sparrow, Aimophila rufescens
- Rufous-crowned Sparrow, Aimophila ruficeps
- Oaxaca Sparrow, Aimophila notostictus
- Collared Towhee, Pipilo ocai
- Green-tailed Towhee, Pipilo chlorurus
- Spotted Towhee, Pipilo maculatus
- Eastern Towhee, Pipilo erythrophthalmus
- Tanager Finch, Oreothraupis arremonops
- Yellow-thighed Finch, Pselliophorus tibialis
- Yellow-green Finch, Pselliophorus luteoviridis
- Moustached Brush-Finch, Atlapetes albofrenatus
- Ochre-breasted Brush-Finch, Atlapetes semirufus
- Tepui Brush-Finch, Atlapetes personatus
- White-naped Brush-Finch, Atlapetes albinucha
- Santa Marta Brush-Finch, Atlapetes melanocephalus
- Pale-naped Brush-Finch, Atlapetes pallidinucha
- Rufous-capped Brush-Finch, Atlapetes pileatus
- Yellow-headed Brush-Finch, Atlapetes flaviceps
- Dusky-headed Brush-Finch, Atlapetes fuscoolivaceus
- Tricolored Brush-Finch, Atlapetes tricolor
- White-rimmed Brush-Finch, Atlapetes leucopis
- Yellow-breasted Brush-Finch / Rufous-naped Brush-Finch, Atlapetes latinuchus
- Antioquia Brush-Finch, Atlapetes blancae
- Rufous-eared Brush-Finch, Atlapetes rufigenis
- Apurimac Brush-Finch, Atlapetes forbesi
- Black-spectacled Brush-Finch, Atlapetes melanopsis
- Slaty Brush-Finch, Atlapetes schistaceus
- White-winged Brush-Finch, Atlapetes leucopterus
- White-headed Brush-Finch, Atlapetes albiceps
- Pale-headed Brush-Finch, Atlapetes pallidiceps
- Bay-crowned Brush-Finch, Atlapetes seebohmi
- Rusty-bellied Brush-Finch, Atlapetes nationi
- Cuzco Brush-Finch, Atlapetes canigenis
- Vilcabamba Brush-Finch, Atlapetes terborghi
- Black-faced Brush-Finch / Gray-eared Brush-Finch, Atlapetes melanolaemus
- Rufous-naped Brush-Finch / Bolivian Brush-Finch, Atlapetes rufinucha
- Fulvous-headed Brush-Finch, Atlapetes fulviceps
- Yellow-striped Brush-Finch, Atlapetes citrinellus
