CORE PASSEROIDEA II

Nine-primaried Oscines

The nine-primaried oscines are called that because they appear to have nine primary feathers. Actually, they have ten primaries, but the tenth primary is reduced and usually hidden under the ninth primary covert (Hall, 2004). The concept was introduced by Wallace (1874), who referred to them as the Tanagroid Passeres. Wallace's grouping was not a natural group. It also included the waxwings, flowerpeckers (in which Wallace included pardalotes and white-eyes), and swallows.

The nine-primaried oscines repesent a recent development, with most of its families originating less than 15 million years old. Nonetheless, the nine-primaried oscines are extremely numerous, and account for almost 18% of all passeriformes. To organize this better, we will use family-level and genus-level groups that are younger than those in most other parts of the avian tree. However, things will stay pretty much unchanged for species.

The basal division in the nine-primaried oscines is between the Motacillidae and the “finches”, Sibley and Monroe's broadly-defined Fringillidae. Wagtails and pipits are cosmopolitan. They are typically open country insectivores.

Motacillidae: Wagtails, Longclaws & Pipits Horsfield, 1821

8 genera, 67 species HBW-9

Click for Motacillidae tree

The Motacillidae include two species that were long thought of as belonging to Sylviidae, the Sao Tome Shorttail (Amaurocichla), and the Madanga (Madanga).

The Sao Tome Shorttail (or Bocage's Longbill) was thought to be related to the Macrosphenus longbills. When the old Sylviidae were broken up, that put it in the crombec family, Macrosphenidae. However, Johansson et al. (2008b) results suggested by it was related to the wagtails and pipits, and possibly sister to the wagtails. Alström et al. (2015a) found it was not only related to the wagtails, but is actually part of Motacilla.

More surprisingly, Alström et al. (2015a) also found that the Madanga is actually a pipit, not a white-eye (Sylviioidea: Zosteropidae). In fact, it falls in the genus Anthus. Its closest relative seems to be the Alpine Pipit, Anthus gutturalis. Laurent Raty has pointed out a complication. In 1831, Lesson used the name Anthus ruficollis. The Madanga wasn't named until 1923 by Rothschild and Hartert, so Lesson's ruficollis has priority if both species are placed in Anthus. The fact that Lesson's ruficollis is a junior synonym of Anthus cervinus (orginally Motacilla cervina Pallas 1811) does not eliminate the conflict. Until either a new name is proposed or Lesson's name Anthus ruficollis is suppressed, the Madanga won't have a proper scientic name. For the present, I will refer to it as Anthus “ruficollis”.

The overall organization of the Motacillidae is based on a combination of Alström et al. (2015a), Drovetski et al. (2018), Harris et al. (2018), Outlaw and Voelker (2006b), Van Ells and Norambuena (2018), and Voelker and Edwards (1998). When there are disagreements, the more recent papers have been given heavier weight.

One interesting thing in Alström et al. (2015a) is that their multigene analysis yields a different placement of the longclaws compared with Outlaw and Voelker (2006b). The limited taxon sampling leaves some residual uncertainty, but it has caused me to separate some of Anthus in Corydalla (Vigors, 1825, type richardi) and Cinaedium (Sundevall, 1850, type lineiventre).

The wagtails, and particularly the yellow-colored wagtails, are more troublesome. Mitochondrial DNA analysis (Alström and Mild, 2003; Ödeen and Bjöklund, 2003; Outlaw and Voelker, 2006b, Pavlova et al., 2003) suggests that there are two species of Citrine Wagtail (citreola and werae) and three species of Yellow Wagtail (flava, taivana, and tschutschensis). The AOU has adopted the split of the Eastern Yellow Wagtail (tschutschensis), but takes no official position on the 3-way split. Christidis and Boles (2008) accept the 3-way split with Green-headed Yellow Wagtail, Motacilla taivana, being the third species.

These splits are not followed by Alström and Mild (2003) or by Ödeen and Björklund (2003), both of which also consider nuclear DNA. The nuclear DNA yields a different tree for the wagtails, where macronyx, taivana and tschutschensis form a clade that can be regarded as a species: Eastern Yellow Wagtail, Motacilla tschutschensis. This species is sister to the Gray Wagtail, Motacilla cinerea. The citrine wagtails end up as sisters, and can also be regarded as a single species. Ödeen and Björklund (2003) argue that the mitochondrial tree reflects the effects of past hybridization.

In the end, I have gone with the analysis by Harris et al. (2018). This treats the Citrine and Eastern Yellow Wagtails as sister species, with the Western Yellow Wagtail sister to both, and the whole lot sister to the Gray Wagtail. Drovetski et al. (2018) has a bit different arrangment.

There have also been questions concerning whether the White Wagtail is a single species. Some authorities have separated lugens as Black-backed Wagtail and yarrelli as Pied Wagtail. However, it is hard to make a genetic case for either of these (see Alström and Mild, 2003; Pavlova et al., 2005; Voelker, 2002), with yarrelli ending up near alba and alba and lugens intertwinned.

Australasian Pipit, Corydalla novaeseelandiae, has been split into Australian Pipit, Corydalla australis, and New Zealand Pipit, Corydalla novaeseelandiae. This split was recommended by Schodde and Mason (1999), but rejected by Christidis and Boles (2008) “in the absence of molecular evidence”. Tavares and Baker (2008) provided limited molecular evidence in the form of a barcode divergence of 4.1%, which is a good indication that they are separate species.

Based on Davies and Peacock (2014), the Kimberley Pipit, Anthus pseudosimilis, is considered conspecific with African Pipit, Anthus cinnamomeus, and the Long-tailed Pipit, Anthus longicaudatus, is considered conspecific with Buffy Pipit, Anthus vaalensis. In both cases, the original collections seem to involve several taxa.

The New World pipits have been rearranged based on Van Ells and Norambuena (2018). Further, their analysis resulted in two splits and one lump.

• Peruvian Pipit, Anthus peruvianus, has been split from Yellowish Pipit, Anthus lutescens
• Puna Pipit, Anthus brevirostris, has been split from Short-billed Pipit, Anthus furcatus.
• South Georgia Pipit, Anthus antarcticus, has been lumped into Correndera Pipit, Anthus correndera due to very small DNA differences between them.

1. Forest Wagtail, Dendronanthus indicus
2. Mountain Wagtail, Motacilla clara
3. Cape Wagtail, Motacilla capensis
4. Sao Tome Shorttail, Motacilla bocagii
5. Madagascan Wagtail, Motacilla flaviventris
6. Gray Wagtail, Motacilla cinerea
7. Western Yellow Wagtail, Motacilla flava
8. Eastern Yellow Wagtail, Motacilla tschutschensis
9. Citrine Wagtail, Motacilla citreola
10. White-browed Wagtail, Motacilla maderaspatensis
11. Mekong Wagtail, Motacilla samveasnae
12. Japanese Wagtail, Motacilla grandis
13. White Wagtail, Motacilla alba
14. African Pied Wagtail, Motacilla aguimp
15. Upland Pipit, Corydalla sylvana
16. Australian Pipit, Corydalla australis
17. New Zealand Pipit, Corydalla novaeseelandiae
18. Tawny Pipit, Corydalla campestris
19. Berthelot's Pipit, Corydalla berthelotii
20. Richard's Pipit, Corydalla richardi
21. Paddyfield Pipit, Corydalla rufula
22. Blyth's Pipit, Corydalla godlewskii
23. Plain-backed Pipit, Corydalla leucophrys
24. Wood Pipit, Corydalla nyassae
25. Long-billed Pipit, Corydalla similis
26. African Pipit, Corydalla cinnamomea
27. Malindi Pipit, Corydalla melindae
28. Buffy Pipit, Corydalla vaalensis
29. Long-legged Pipit, Corydalla pallidiventris
30. Sokoke Pipit, Cinaedium sokokense
31. Short-tailed Pipit, Cinaedium brachyurum
32. Bushveld Pipit, Cinaedium caffrum
33. Mountain Pipit, Cinaedium hoeschi
34. Striped Pipit, Cinaedium lineiventre
35. African Rock Pipit, Cinaedium crenatum
36. Golden Pipit, Tmetothylacus tenellus
37. Yellow-breasted Pipit, Hemimacronyx chloris
38. Sharpe's Longclaw, Hemimacronyx sharpei
39. Abyssinian Longclaw, Macronyx flavicollis
40. Fuelleborn's Longclaw, Macronyx fuelleborni
41. Cape Longclaw, Macronyx capensis
42. Yellow-throated Longclaw, Macronyx croceus
43. Pangani Longclaw, Macronyx aurantiigula
44. Rosy-throated Longclaw, Macronyx ameliae
45. Grimwood's Longclaw, Macronyx grimwoodi
46. Sprague's Pipit, Anthus spragueii
47. Yellowish Pipit, Anthus lutescens
48. Puna Pipit, Anthus brevirostris
49. Short-billed Pipit, Anthus furcatus
50. Peruvian Pipit, Anthus peruvianus
51. Pampas Pipit, Anthus chacoensis
52. Paramo Pipit, Anthus bogotensis
53. Hellmayr's Pipit, Anthus hellmayri
54. Correndera Pipit, Anthus correndera
55. Ochre-breasted Pipit, Anthus nattereri
56. Pechora Pipit, Anthus gustavi
57. Alpine Pipit, Anthus gutturalis
58. Madanga, Anthus "ruficollis"
59. Nilgiri Pipit, Anthus nilghiriensis
60. Tree Pipit, Anthus trivialis
61. Olive-backed Pipit, Anthus hodgsoni
62. Rosy Pipit, Anthus roseatus
63. Red-throated Pipit, Anthus cervinus
64. American Pipit / Buff-bellied Pipit, Anthus rubescens
65. Meadow Pipit, Anthus pratensis
66. Water Pipit, Anthus spinoletta
67. Eurasian Rock Pipit, Anthus petrosus

Sibley and Monroe's Fringillidae

Sibley and Monroe's (1990) Fringillidae included all of the remaining birds, over 1000 of them. Most authors use a finely-grained family structure for these species that roughly corresponds to Sibley and Monroe's tribes. The TIF taxonomy divides Sibley and Monroe's Fringillidae into 11.

Sibley and Monroe's Fringillidae: TiF Family Tree

Fringillidae: Finches, Euphonias Leach, 1819

60 genera, 228 species HBW-15

Not long ago, the chlorophonias and euphonias were considered tanagers. Many guidebooks still list them as such, but it is not so. The AOU recognized them as finches in the 44th checklist supplement (2003), placing them in the subfamily Euphoniinae. Accordingly, they are placed in Fringillidae as a subfamily (Groth, 1998; Klicka et al., 2000; Yuri and Mindell, 2002; Zuccon et al., 2012). The Hawaiian Honeycreepers were once considered a separate family (Drepanididae). They are now thought to form a clade buried inside Carduelinae (Yuri and Mindell, 2002; Arnaiz-Villena et al., 2007b; Lerner et al., 2011; Zuccon et al., 2012). They are treated here as a tribe within Carduelinae.

Click for Fringillidae genus tree

The arrangement of the finches is based on several sources. I originally relied on Groth (1998), Klicka et al. 2007, and Arnaiz-Villena et al. (1998, 1999, 2001, 2007a, b, 2008). Their studies have included a substantial number of finch species. However, some monotypic genera had been left out (e.g., Callacanthis, Chaunoproctus, Kozlowia, Neospiza). These have all been included in the analysis by Zuccon et al. (2012).

The tribes within the Carduelinae are now based on Lerner et al. (2011) and Zuccon et al. (2012). Except for the basal position of the Coccothraustini, there had previously been a lack of consensus on the relationships between the Cardueline tribes. See Yuri and Mindell (2002), Arnaiz-Villena et al. (2007a), Nguembock et al. (2009a), and Töpfer et al. (2011). That has changed with the publication of Lerner et al. (2011) and Zuccon et al. (2012). Although the taxon sampling is different, the two papers are completely consistent at the generic level except for the placement of the Pyrrhulini. Zuccon et al. put the Pyrhullini sister to the Carduelini while Lerner et al. place it sister to the Carpodacini/Drepanidini clade. Note that “European Serin” on Lerner et al.'s tree is a typo as noted in the supplementary material. It actually refers to the White-bellied Canary.

The combination of a large number of genes and attention to the problem of discerning the true species tree in spite of incongruent gene trees has led me to use the overall structure from Lerner et al. (2011). They use a large superset of the genes used by Zuccon et al., but only sample half as many taxa. The TiF tree of genera is 100% consistent with Lerner et al., and except for the placement of Pyrrhulini, is also consistent with Zuccon et al. It's nice to see so much consensus.

The Fringillidae start with a basal Palearctic group, Fringillinae. Fringillinae is comprised of three species, one of which has spread across the northern Palearctic. The remaining Fringillidae fall into two sister clades, Euphoniinae and Carduelinae. The common ancestor of the Euphoniinae/Carduelinae clade was likely an Old World species. Something must connect it with the Neotropical Euphoniinae, but that something is missing without a trace. Being geographically contiguous and more similar in appearance, the position of the Carduelinae is easy to understand. Euphoniinae is a mystery that unscores the importance of extinction in creating the bird distribution we currently see.

Fringillinae: Chaffinches Leach, 1819

The split of the Blue Chaffinch, Fringilla teydea, from Common Chaffinch, Fringilla coelebs, was based on on Suárez et al. (2009). The further split of Gran Canaria Blue Chaffinch, Fringilla polatzeki, from Blue Chaffinch, Fringilla teydea, is based on Suárez et al. (2009) and the analysis by Sangster et al. (2016b).

1. Brambling, Fringilla montifringilla
2. Common Chaffinch, Fringilla coelebs
3. Blue Chaffinch, Fringilla teydea
4. Gran Canaria Blue Chaffinch, Fringilla polatzeki

Euphoniinae: Euphonias, Chlorophonias Tschudi, 1846

Zuccon et al. (2012) found evidence that the Antillean Euphonia is more closely related to the chlorophonias than to most euphonias. The two other blue-headed species are usually considered closely related to musica (sometimes treated as one species under the name Blue-hooded Euphonia), so all three have been placed in the genus Cyanophonia (Bonaparte 1851, type musica).

Due mainly to plumage differences between the subspecies, the Antillean Euphonia, Cyanophonia musica has been split into Hispaniolan Euphonia, Cyanophonia musica, Puerto Rican Euphonia, Cyanophonia sclateri, and Lesser Antillean Euphonia, Cyanophonia flavifrons, following the 64th AOS Supplement.

1. Golden-rumped Euphonia, Cyanophonia cyanocephala
2. Antillean Euphonia, Cyanophonia musica
3. Elegant Euphonia, Cyanophonia elegantissima
4. Blue-naped Chlorophonia, Chlorophonia cyanea
5. Chestnut-breasted Chlorophonia, Chlorophonia pyrrhophrys
6. Yellow-collared Chlorophonia, Chlorophonia flavirostris
7. Blue-crowned Chlorophonia, Chlorophonia occipitalis
8. Golden-browed Chlorophonia, Chlorophonia callophrys
9. Jamaican Euphonia, Euphonia jamaica
10. Plumbeous Euphonia, Euphonia plumbea
11. Scrub Euphonia, Euphonia affinis
12. Yellow-crowned Euphonia, Euphonia luteicapilla
13. Purple-throated Euphonia, Euphonia chlorotica
14. Trinidad Euphonia, Euphonia trinitatis
15. Velvet-fronted Euphonia, Euphonia concinna
16. Orange-crowned Euphonia, Euphonia saturata
17. Finsch's Euphonia, Euphonia finschi
18. Violaceous Euphonia, Euphonia violacea
19. Thick-billed Euphonia, Euphonia laniirostris
20. Yellow-throated Euphonia, Euphonia hirundinacea
21. Green-throated Euphonia / Green-chinned Euphonia, Euphonia chalybea
22. Fulvous-vented Euphonia, Euphonia fulvicrissa
23. Spot-crowned Euphonia, Euphonia imitans
24. Olive-backed Euphonia, Euphonia gouldi
25. Golden-bellied Euphonia / White-lored Euphonia, Euphonia chrysopasta
26. Bronze-green Euphonia, Euphonia mesochrysa
27. White-vented Euphonia, Euphonia minuta
28. Tawny-capped Euphonia, Euphonia anneae
29. Orange-bellied Euphonia, Euphonia xanthogaster
30. Rufous-bellied Euphonia, Euphonia rufiventris
31. Chestnut-bellied Euphonia, Euphonia pectoralis
32. Golden-sided Euphonia, Euphonia cayennensis

Carduelinae Vigors, 1825

As mentioned above, how the tribes with the Carduelinae relate has been somewhat contentious. There is considerable agreement that the holarctic Coccothraustini are basal. The results of Nguembock et al. (2009a) suggested that Carpodacus and Pyrrhula were fairly closely related to each other, as did Töpfer et al. (2011). Neither had much to say about their relation to the Hawaiian Honeycreepers (Drepanidini). Zuccon et al. (2012) were unable to clearly resolve the position of the honeycreepers (see Fig. 2), but Lerner et al. (2011) place them sister to the rosefinches (Carpodacini). The two together are then sister to the bullfinches and many arid-zone finches (Pyrrhulini). Lerner et al. place the whole lot of them are sister to Carduelini, which includes the American red finches (previously separated as a tribe). As mentioned before, Zuccon et al. disagree, placing Pyrrhulini sister to Carduelini.

Coccothraustini: Grosbeak Finches Swainson, 1831

Zuccon et al. (2012) found that Eophona was embedded within Coccothraustes. A reasonable solution to this problem is to restore the two American species to Hesperiphona and leave the Asian species in a reduced Eophona.

1. Black-and-yellow Grosbeak, Mycerobas icterioides
2. Collared Grosbeak, Mycerobas affinis
3. Spot-winged Grosbeak, Mycerobas melanozanthos
4. White-winged Grosbeak, Mycerobas carnipes
5. Evening Grosbeak, Hesperiphona vespertina
6. Hooded Grosbeak, Hesperiphona abeillei
7. Hawfinch, Coccothraustes coccothraustes
8. Chinese Grosbeak, Eophona migratoria
9. Japanese Grosbeak, Eophona personata

Pyrrhulini: Bullfinches and Arid-zone FinchesVigors, 1825

Click for Pyrrhulini tree

Pyrrhulini consists of 3 clades. The first consists of the Pyrrhula bullfinches and Pine Grosbeak (Pinicola). The second and third are sister, and are the arid-zone finches indentified by Arnaiz-Villena et al. (2008). The second clade includes Bucanetes and Rhodopechys. The third consists of the mountain-finches and rosy-finches (Leucosticte), together with several monotypic genera: Procarduelis (Dark-breasted Rosefinch), Agraphospiza (Blanford's Rosefinch), Callacanthis (Spectacled Finch), and Pyrrhoplectes (Golden-naped Finch). Both Procarduelis and Agraphospiza are usually included in Carpodacus.

The bullfinches are arranged based on Töpfer et al. (2011). Kirwan and Gregory (2005) established the monotypic genus Eremopsaltria for the Mongolian Finch, separating it from Bucanetes. However, they acted under the misapprehension that the Mongolian Finch is more likely closer to the Common Rosefinch than to the Trumpeter Finch. Arnaiz-Villena et al. (2008) found a rather different result, with the two Bucanetes as sister taxa, and the rosefinch placed rather distantly from them. Zuccon et al. (2012) confirm this.

Arnaiz-Villena et al. (2008) found that the Dark-breasted Rosefinch, formerly Carpodacus nipalensis belongs in the same clade close to Leucosticte. Rather than folding it in Leucosticte, I restored the genus name Procarduelis (Blyth 1843). I had previously noted that Blanford's Rosefinch is thought to be close to nipalensis. Indeed, Zuccon et al. found it in the other portion of this clade. It is sufficently distant from its sister group (Callacanthis and Pyrrhoplectes) to deserve a separate genus. Zuccon et al. (2012) established the name Agraphospiza for it.

I follow AOU in listing three North American rosy-finches. Some have even suggested further splitting the Gray-crowned Rosy-Finch. They have also been all been lumped into one species in the past. Drovetski et al. (2009) find little genetic difference between the three, suggesting that they may be lumped again.

1. Pine Grosbeak, Pinicola enucleator
2. Brown Bullfinch, Pyrrhula nipalensis
3. White-cheeked Bullfinch, Pyrrhula leucogenis
4. Eurasian Bullfinch, Pyrrhula pyrrhula
5. Azores Bullfinch, Pyrrhula murina
6. Orange Bullfinch, Pyrrhula aurantiaca
7. Red-headed Bullfinch, Pyrrhula erythrocephala
8. Gray-headed Bullfinch, Pyrrhula erythaca
9. Trumpeter Finch, Bucanetes githagineus
10. Mongolian Finch, Bucanetes mongolicus
11. Eurasian Crimson-winged Finch, Rhodopechys sanguineus
12. African Crimson-winged Finch, Rhodopechys alienus
13. Blanford's Rosefinch, Agraphospiza rubescens
14. Spectacled Finch, Callacanthis burtoni
15. Golden-naped Finch, Pyrrhoplectes epauletta
16. Dark-breasted Rosefinch, Procarduelis nipalensis
17. Plain Mountain-Finch, Leucosticte nemoricola
18. Brandt's Mountain-Finch, Leucosticte brandti
19. Asian Rosy-Finch, Leucosticte arctoa
20. Gray-crowned Rosy-Finch, Leucosticte tephrocotis
21. Black Rosy-Finch, Leucosticte atrata
22. Brown-capped Rosy-Finch, Leucosticte australis

Carpodacini: Rosefinches Bonaparte, 1853

The arrangement of the rosefinches is now based on Päckert et al. (2020b), which improves on the analyses of Tietze et al. (2013) and Zuccon et al. (2012). Now that we had a decent understanding of how the rosefinches relate, and how old the taxa are, we can establish generic limits in a reasonable way, balancing age and coherence.

There is a deep division in the basal clade, which consists of the Common Rosefinch (Erythrina) and the Scarlet Finch (Haematospiza).

The extinct Bonin Grosbeak (aka Chaunoproctus) is basal in the main clade, and is sister to the remaining two dozen rosefinches. These split into two clades.

The first ranges from Blyth's Rosefinch to Taiwan Rosefinch. The Taiwan Rosefinch, Carpodacus formosanus, has been split from Vinaceous Rosefinch, Carpodacus vinaceus, based on Wu et al. (2011).

The second clade includes synoicus group, the former Kozlowia (C. roborowskii), the Rubicilla clade (Streaked and Great Rosefinches), the former Uragus (C. sibiricus), Pyrrhospiza (C. puniceus), and Pinicola subhimachala (aka Propyrrhula subhimachala). It ends with a group including the Carpodacus type species Pallas's Rosefinch, Carpodacus roseus (see Banks and Browning 1995), through C. dubius.

Tietze et al. (2013) found substantial genetic distance between the Sinai Rosefinch in the strict sense C. synoicus synoicus, and the Pale Rosefinch group (subspecies beicki, salimalii, and presumably stoliczkae). These have been suspected of being different species, and I have now split them.

I moved Sillem's Mountain-Finch (rediscovered in June 2012) from Leucosticte (Pyrrhulini) to Carpodacus based on a photo of what is believed to be the female. When Roselaar (1992) named Sillem's Rosefinch, he put it in Leucosticte. At the time, the female plumage was unknown. The other Leucosticte have female plumages that are only slightly different from the male, usually just duller. As Roselaar pointed out, if it has a distinctive female plumage, it would more likely related to Kozlowia (now part of Carpodacus) than to Leucosticte. More recently, Muzika (2014) has made the same point. Yann Muzika's photo of the probable female Sillem's shows a bird that is quite different from the adult male. See Kazmierczak and Muzika (2012) for an account of this rediscovery and Muzika (2014) for further observations.

A DNA analysis by Sangster et al. (2016a) has confirmed that Sillem's Mountain-Finch is a Carpodacus rosefinch, and as expected, is sister to Tibetan Rosefinch, Carpodacus roborowskii.

Tietze et al. (2013) also found a surprisingly small genetic distance between the Caucasian and Spotted Great-Rosefinches (rubicilla and severtzovi) and recommended lumping them. Although I have followed that advice, further study is in order. Rasmussen and Anderton (2005) recommended splitting them based on differences in plumage, habitat, and voice.

The genus Carpodacus has been divided into the following genera:

• Common Rosefinch, Carpodacus erythrinus, is transferred to Erythrina (Brehm 1828, monotypic)
• Scarlet Finch, Carpodacus sipahi, is transferred to Haematospiza (Blyth 1845, monotypic)
• Bonin Grosbeak, Carpodacus ferreorostris, is transferred to Chaunoproctus (Bonaparte 1850, monotypic)
• Sinai Rosefinch, Carpodacus synoicus, and Pale Rosefinch, Carpodacus stoliczkae need a genus name. For now, I'll list them as "Carpodacus"
• Sillem's Mountain-Finch, Carpodacus sillemi, and Tibetan Rosefinch, Carpodacus roborowskii, are transferred to Kozlowia (Bianchi 1907, type roborowskii)
• Red-fronted Rosefinch, Carpodacus punicea, and Crimson-browed Finch, Carpodacus subhimachala, are transferred to Propyrrhula (Hodgson 1844, type subhimachala)
• Long-tailed Rosefinch, Carpodacus sibiricus, is transferred to Uragus (Keyserling and Blasius 1840, monotypic)
• Pallas's Rosefinch, Carpodacus roseus and the three following species remain in Carpodacus (Kaup 1829, type roseus)
• Streaked Rosefinch, Carpodacus rubicilloides, and Great Rosefinch, Carpodacus rubicilla, are transferred to Rubicilla (Bonaparte and Schlegel 1850 type rubicilla)
• Blyth's Rosefinch, Carpodacus grandis, through Taiwan Rosefinch, Carpodacus formosanus (11 species) are transferred to Propasser (Hodgson 1844 type rodopeplus)

Click for Carpodacini tree

1. Common Rosefinch, Erythrina erythrina
2. Scarlet Finch, Haematospiza sipahi
3. †Bonin Grosbeak, Chaunoproctus ferreorostris
4. Sinai Rosefinch, "Carpodacus" synoicus
5. Pale Rosefinch, "Carpodacus" stoliczkae
6. Sillem's Mountain-Finch, Kozlowia sillemi
7. Tibetan Rosefinch, Kozlowia roborowskii
8. Red-fronted Rosefinch, Propyrrhula punicea
9. Crimson-browed Finch, Propyrrhula subhimachala
10. Long-tailed Rosefinch, Uragus sibiricus
11. Pallas's Rosefinch, Carpodacus roseus
12. Three-banded Rosefinch, Carpodacus trifasciatus
13. Himalayan White-browed Rosefinch, Carpodacus thura
14. Chinese White-browed Rosefinch, Carpodacus dubius
15. Streaked Rosefinch, Rubicilla rubicilloides
16. Great Rosefinch, Rubicilla rubicilla
17. Blyth's Rosefinch, Propasser grandis
18. Red-mantled Rosefinch, Propasser rhodochlamys
19. Pink-rumped Rosefinch, Propasser waltoni
20. Himalayan Beautiful-Rosefinch, Propasser pulcherrimus
21. Chinese Beautiful-Rosefinch, Propasser davidianus
22. Dark-rumped Rosefinch, Propasser edwardsii
23. Pink-browed Rosefinch, Propasser rodochroa
24. Sharpe's Rosefinch, Propasser verreauxii
25. Spot-winged Rosefinch, Propasser rodopeplus
26. Vinaceous Rosefinch, Propasser vinaceus
27. Taiwan Rosefinch, Propasser formosanus

Drepanidini Cabanis, 1847

The results in both Lerner et al. (2011) and Zuccon et al. (2012) support the Drepanidini as sister to Carpodacini (not just the Common Rosefinch, as might be inferred from the press coverage). Interestingly, the extinct Bonin Grosbeak is a relatively basal member of the Carpodacini. Perhaps it too is a relic of the spread of the ancestral Carpodacini/Drepanidini into the Pacific.

The order here is based on a combination of the DNA study of Lerner et al. (2011) and the osteological study of James (2004). Fleischer et al. (2001), Pratt (2001), Arnaiz-Villena et al. (2007b), and Reding et al. (2009) have also been consulted. It fits nicely with morphology (look at bill shapes in each clade) and subfossil taxa can be easily accomodated using James (2004). This is pretty similar to the previous version, but tweaked based on Lerner et al. (2011), which greatly decreases the conjecture needed, and increases my confidence in it.

The genera Palmeria, Himatione, and Vestiaria have been merged into Drepanis. The genetic distance between them seems to be small (Lerner et al., 2011) and there is evidence of hybridization between Vestiaria and Himatione (Knowlton et al., 2014). Olson (2012b) also argues that this is a reasonable treatment. Although I think the genera I use for the Hawaiian Honeycreepers are oversplit, I consider the AOU genera even more oversplit. I haven't gone further on the lumping because data on the extinct species is too limited.

Although Pratt (2009) has established the genus Manucerthia for the Hawaiian Creeper, Loxops mana, I have not adopted it. Lerner et al. (2011) make clear that the Hawaiian Creeper is sister to the other Loxops. Moreover, they shared a common ancestor about 2 million years ago and do not seem so different as to justify introducing an extra genus into a tribe that already has too many genera.

I have kept the genus Akialoa merged with Hemignathus as Akialoa appears to be paraphyletic (see the trees in James, 2004). However, portions of Hemignathus have been split off as Chlorodrepanis and Viridonia. This has allowed restoration of the name Hemignathus wilsoni for the Akiapolaau (Hemignathus munroi). When Chlorodrepanis virens is subsumed in Hemignathus, the name wilsoni belongs to a subspecies of virens. The situation with the Kauai Amakihi, Chlorodrepanis stejnegeri is similar. In Hemignathus, the name stejnegeri belongs to a subspecies of the Greater Akialoa, Hemignathus ellisianus stejnegeri, and the Kauai Amakihi uses the substitute name H. kauaiensis.

I follow the AOU decisions (56th supplement) to split the Apapane, Akepa, Greater Akialoa, and Nukupuu. Thus:

• Apapane, Drepanis (Himatione) sanguinea is split into
  • Laysan Honeycreeper, Drepanis fraithii (extinct)
  • Apapane, Drepanis sanguinea
• Akepa / Hawaii Akepa, Loxops coccineus is split into
  • Oahu Akepa, Loxops wolstenholmei (probably extinct)
  • Maui Akepa, Loxops ochraceus (probably extinct)
  • Hawaii Akepa, Loxops coccineus
• Nukupuu, Hemignathus lucidus, is split into
  • Kauai Nukupuu, Hemignathus hanapepe (probably extinct)
  • Oahu Nukupuu, Hemignathus lucidus (probably extinct)
  • Maui Nukupuu, Hemignathus affinis (possibly extinct)
• Greater Akialoa, Hemignathus ellisianus, is split into
  • Kauai Akialoa, Hemignathus stejnegeri (extinct)
  • Oahu Akialoa, Hemignathus ellisianus (extinct)
  • Maui-nui Akialoa, Hemignathus lanaiensis (extinct)

This changes clade sizes and I have also reordered the Hawaiian Honeycreepers as a result.

Click for Drepanidini species tree

1. Poo-uli, Melamprosops phaeosoma
2. Akikiki / Kaui Creeper, Oreomystis bairdi
3. Oahu Alauahio / Oahu Creeper, Paroreomyza maculata
4. Kakawahie / Molokai Creeper, Paroreomyza flammea
5. Maui Alauahio / Maui Creeper, Paroreomyza montana
6. Laysan Finch, Telespiza cantans
7. Nihoa Finch, Telespiza ultima
8. Palila, Loxioides bailleui
9. Kona Grosbeak, Chloridops kona
10. Lesser Koa-Finch, Rhodacanthis flaviceps
11. Greater Koa-Finch, Rhodacanthis palmeri
12. Ula-ai-hawane, Ciridops anna
13. Akohekohe / Crested Honeycreeper, Drepanis dolei
14. Laysan Honeycreeper, Drepanis fraithii
15. Apapane, Drepanis sanguinea
16. Iiwi, Drepanis coccinea
17. Black Mamo, Drepanis funerea
18. Hawaii Mamo, Drepanis pacifica
19. Greater Amakihi, Viridonia sagittirostris
20. Anianiau, Magumma parva
21. Hawaii Amakihi, Chlorodrepanis virens
22. Kauai Amakihi, Chlorodrepanis stejnegeri
23. Oahu Amakihi, Chlorodrepanis flava
24. Hawaii Creeper, Loxops mana
25. Akekee / Kauai Akepa, Loxops caeruleirostris
26. Oahu Akepa, Loxops wolstenholmei
27. Maui Akepa, Loxops ochraceus
28. Hawaii Akepa, Loxops coccineus
29. Ou, Psittirostra psittacea
30. Lanai Hookbill, Dysmorodrepanis munroi
31. Maui Parrotbill / Kiwikiu, Pseudonestor xanthophrys
32. Lesser Akialoa / Hawaii Akialoa, Hemignathus obscurus
33. Kauai Akialoa, Hemignathus stejnegeri
34. Oahu Akialoa, Hemignathus ellisianus
35. Maui-nui Akialoa, Hemignathus lanaiensis
36. Akiapolaau, Hemignathus wilsoni
37. Kauai Nukupuu, Hemignathus hanapepe
38. Oahu Nukupuu, Hemignathus lucidus
39. Maui Nukupuu, Hemignathus affinis

Carduelini: Canaries, Siskins and allies Vigors, 1825

This brings us to the Carduelini. It has become apparent that the North American Carpodacus finches are not that closely related to the true Carpodacus finches. Previously, the TiF list used the genus Burrica, also used as a subgenus by AOU (2nd-5th AOU checklists). However, the oldest available name seems to be Erythrospiza (Bonaparte 1830, type purpurea—compare Haemorhous Swainson, 1837, type purpureus or Burrica Ridgway, 1887, type mexicana). Erythrospiza appears to be sister to the rest of the Carduelini, and rather distantly related. Perhaps it would even make sense to treat it as a separate tribe (as done previously in the TiF list).

Click for Carduelini tree

I follow the recent paper by Zuccon et al. (2012) to arrange the various Cardueline genera. Lerner et al. (2011) gives essentially identical results for the taxa they included. Previous studies did a decent job at the generic level, and sometimes for groups of genera, but were unable to get strong support for any method of arranging them. Thus the papers by Arnaiz-Villena et al. (1998, 1999, 2001, 2007a, b, 2008) and Zamora et al. (2006) have been used in ordering the species and determing the exact generic boundaries. Unfortunately, the papers by Arnaiz-Villena and colleagues have been completely useless for determining deeper relations. One wonders whey they continue to reuse the same data rather than collecting more. Nguembock et al. (2009a) has also been helpful at higher levels, but I rely on Lerner et al. and Zuccon et al. for the overall structure.

The various studies of the finches have made it clear that the genera Carduelis and Serinus both required substantial surgery. The alternative for Carduelis would be to put almost all of the Carduelini into one genus! For Serinus, the alternative is less drastic, mainly because Carduelis has priority. Neither of these outcomes is particuarly desirable, so both Serinus and Carduelis have been divided into several pieces each.

As constituted here, Carduelini contains 4 major clades. As mentioned above, the first clade includes only a single genus: Erythrospiza — the American purple finches. The genus has been changed from Haemorhous (Swainson 1837) to Erythrospiza (Bonaparte 1830) type purpurea as the latter has priority.

The second clade includes the Chloris greenfinches (sometimes included in Carduelis), the Desert Finch (Rhodospiza), and the the Golden-winged Grosbeaks (Rhynchostruthus). Kirwan and Grieve (2007) argue that Rhynchostruthus includes three species.

The third Carduelini clade is exclusively Afrotropical. The Oriole Finch (Linurgus) is sister to the rest. These closely related birds are variously called canaries, seedeaters, serins, and siskins and are all placed in the genus Crithagra. The species-level phylogeny still needs some work, as might be guessed from the question marks, polytomies, and blue ink.

I recognize several subgenera. Besides Crithagra, there are Dendrospiza, Neospiza, Pseudochloroptila, and Ochrospiza. All are marked on the species tree.

Subgenus Crithagra includes most of the larger African canaries and allies.

The Sao Tome Grosbeak has usually been included in Neospiza. Stervander (2010) found that the Principe Seedeater is it's closest relative and both species together form subgenus Neospiza. In fact, Stervander found that the Sao Tome subspecies of the seedeater (thomensis) appeared more closely related to the Sao Tome Grosbeak than to the seedeater. The two birds are quite distinct, and it seems unreasonable to put them in one species. Perhaps the genetic results are a sign of introgression.

It's not entirely clear whether subgenus Pseudochloroptila belongs in subgenus Ochrospiza, so they are kept separate. Both the Yemen Serin, Crithagra menachensis, and Ankober Serin, Crithagra ankoberensis, remain somewhat mysterious. These have be considered conspecific by some, or in separate genera by others. I have little confidence in their placement here (hence the blue color on the tree), but it seems at least as reasonable as any other. This clade is expected to get a revision when more information becomes available.

The fourth Carduelini clade is more cosmopolitan. The basal piece includes the Palearctic Linnets (Linaria, formerly Carduelis), which are sister to the holarctic Redpolls (now Acanthis rather than Carduelis) and crossbills (Loxia). There is some uncertainty about the linnets. The mitochondrial dna consistently puts them near the siskins and American goldfinches rather than near the redpolls and crossbills. The nuclear dna does the opposite. Since they seem more akin to the redpolls and crossbills, and since the combined analysis puts them there, I have followed suit.

The redpolls themselves are an interesting case. It's not clear how many redpoll species there are. There is evidence that at least two are good biological species (e.g., Knox, 1988). In fact, Knox et al. (2001) support separation of Lesser Redpoll, Acanthis cabaret. However, genetic studies have failed to find any differences between Lesser and Common Redpolls (Ottvall et al., 2002). Worse, Marthinsen et al. (2008) found little genetic difference between any of the redpolls! There may be only one redpoll species. Indeed, Mason and Taylor's detailed study using SNP's (2015) found little genetic differentiation among the redpolls. At this point the balance of the evidence is that there is only one species involved. As a result, I've lumped them all as a single species, Redpoll, Acanthis flammea.

It's also not really clear how many red crossbills there are. I continue to follow the AOU and BOU taxonomy for the crossbills, but Benkman et al. (2009) make a case for considering the “type 9” crossbills of Idaho to be a separate species, South Hills Crossbill, Loxia sinesciuris. See also Parchman et al. (2006). However, it is doubtful there's any real genetic differentiation (Piertney et al., 2001). I have also lumped the Scottish Crossbill into the Red (Common) Crossbill.

The other piece of this final clade includes the true Carduelis and Serinus finches of the Palearctic. It starts with the Mountain Serin of Indonesia and the Philippines. The Mountain Serin is sometimes considered part of Serinus, but I've had it in its own genus, Chrysocorythus, for a while. Zuccon et al. (2012) found Chrysocorythus sister to the true Carduelis finches, now reduced to the European Goldfinch and the Corsican and Citril Finches. These two genera together are sister to the rest of the clade.

The now-reduced Serinus comes next, followed by the Tibetan Serin. It's in Chionomitris rather than Serinus.

Most of the remaining Carduelini are from the Americas. There are two classification schemes in current use. The AOU's NACC puts them all together in genus Spinus. The AOU's SACC separates them into three genera: The North American goldfinches, Astragalinus; the northern siskins Spinus; and the Middle and South American siskins, Sporagra. Astragalinus splits off first, and the final division is between Spinus and Sporagra.

Several Carduelis siskins have moved to Spinus (including Lesser Antillean Siskin). In another taxonomic note, the Pine Siskin, Spinus pinus, and Black-capped Siskin, Spinus atriceps, are quite close. Some have suggested they are conspecific. Although they don't comment on it, the genetic tree in Arnaiz-Villena et al. (2008) suggests another possibility. The subspecies perplexus may actually belong to S. atriceps, as had been suggested by Banks in 1982 (see a 2008 AOU proposal).

The placement of Spinus may be a little soft. E.g., in Nguembock et al. (2009a), it appeared in different places in the combined tree (their Figure 4) and some of the individual gene trees. This is the subject of some discussion in Nguembock et al. The arrangement here is based on Zuccon et al. (2012) and Beckman and Witt (2015).

These three genera were formerly considered part of Carduelis. The AOU currently treats them all as Spinus, and rejected a proposal to make these last two generic splits, also proposed by Nguembock et al.

One has to be careful with the genus name for the South American siskins. Both Sporagra (Reichenbach 1850, type magellanica) and Pyrrhomitris (Bonaparte 1850, type cucullata) come into consideration. The publication date of Sporagra seems to be June 1, 1850. The publication date for Pyrrhomitris is not as clear. Bonaparte's “Conspectus generum avium” was published in sections beginning in mid-1850. The first part was already available in mid-June, and likely published a bit earlier, perhaps earlier than Reichenbach. However, Pyrrhomitris was not included in the section I, nor was it even included in the first part of section II (published by Oct 15, 1850). It appeared in the second part of section II (dated Nov 10, 1850 and certainly published before Feb 3, 1851). If I understand the ICZN correctly, the parts should be treated as separate publications, in which case it appeared either in the later part of 1850 or early 1851. That would give priority to Reichenbach's Sporagra. In this case there is further evidence of which was published first. Bonaparte refers to Sporagra on page 516, the page before he establishes Pyrrhomitris.

Based on Beckman and Witt (2015), the Hooded Siskin, Sporagra magellanica, has been split into Lowland Hooded Siskin, Sporagra magellanica, and Andean Hooded Siskin, Sporagra capitalis. We lack complete information on the subspecies, but I've tentatively allocated boliviana, alleni, icterica, longirostris, and magellanica to the lowland group, S. magellanica, and capitalis, paula, peruana, urubambensis, santaecrucis, hoyi, and tucumana to the Andean group, S. capitalis. The genetic distances between the capitalis group and atrata, crassirostris, siemiradzkii is razor-thin, calling their species status into question. Even uropygialis appears to be closely related to the capitalis group. Further study is needed to sort out these taxa.

1. House Finch, Erythrospiza mexicana
2. Purple Finch, Erythrospiza purpurea
3. Cassin's Finch, Erythrospiza cassinii
4. Desert Finch, Rhodospiza obsoleta
5. Socotra Golden-winged Grosbeak, Rhynchostruthus socotranus
6. Arabian Golden-winged Grosbeak, Rhynchostruthus percivali
7. Somali Golden-winged Grosbeak, Rhynchostruthus louisae
8. European Greenfinch, Chloris chloris
9. Oriental Greenfinch / Gray-capped Greenfinch, Chloris sinica
10. Yellow-breasted Greenfinch, Chloris spinoides
11. Vietnamese Greenfinch, Chloris monguilloti
12. Black-headed Greenfinch, Chloris ambigua
13. Oriole Finch, Linurgus olivaceus
14. Thick-billed Seedeater, Crithagra burtoni
15. Protea Seedeater / Protea Canary, Crithagra leucoptera
16. Kipengere Seedeater, Crithagra melanochroa
17. Streaky Seedeater, Crithagra striolata
18. Yellow-browed Seedeater, Crithagra whytii
19. White-throated Canary, Crithagra albogularis
20. Brimstone Canary, Crithagra sulphurata
21. Yellow Canary, Crithagra flaviventris
22. Northern Grosbeak-Canary, Crithagra donaldsoni
23. Southern Grosbeak-Canary, Crithagra buchanani
24. Brown-rumped Seedeater, Crithagra tristriata
25. Reichard's Seedeater, Crithagra reichardi
26. Black-eared Seedeater, Crithagra mennelli
27. West African Seedeater, Crithagra canicapilla
28. Streaky-headed Seedeater, Crithagra gularis
29. Principe Seedeater, Crithagra rufobrunnea
30. Sao Tome Grosbeak, Crithagra concolor
31. Black-faced Canary, Crithagra capistrata
32. Papyrus Canary, Crithagra koliensis
33. Forest Canary, Crithagra scotops
34. African Citril, Crithagra citrinelloides
35. Western Citril, Crithagra frontalis
36. Southern Citril, Crithagra hyposticta
37. Cape Siskin, Crithagra totta
38. Drakensberg Siskin, Crithagra symonsi
39. Yemen Serin, Crithagra menachensis
40. Ankober Serin, Crithagra ankoberensis
41. White-bellied Canary, Crithagra dorsostriata
42. Yellow-fronted Canary, Crithagra mozambica
43. Lemon-breasted Canary, Crithagra citrinipectus
44. White-rumped Seedeater, Crithagra leucopygia
45. Black-throated Canary, Crithagra atrogularis
46. Yellow-rumped Seedeater, Crithagra xanthopygia
47. Reichenow's Seedeater, Crithagra reichenowi
48. Arabian Seedeater / Arabian Serin, Crithagra rothschildi
49. Yellow-throated Seedeater, Crithagra flavigula
50. Salvadori's Seedeater, Crithagra xantholaema
51. Twite, Linaria flavirostris
52. Common Linnet, Linaria cannabina
53. Yemen Linnet, Linaria yemenensis
54. Warsangli Linnet, Linaria johannis
55. Redpoll, Acanthis flammea
56. Hispaniolan Crossbill, Loxia megaplaga
57. White-winged Crossbill / Two-barred Crossbill, Loxia leucoptera
58. Parrot Crossbill, Loxia pytyopsittacus
59. Scottish Crossbill, Loxia scotica
60. Red Crossbill, Loxia curvirostra
61. Mountain Serin, Chrysocorythus estherae
62. European Goldfinch, Carduelis carduelis
63. Citril Finch, Carduelis citrinella
64. Corsican Finch, Carduelis corsicana
65. Island Canary / Atlantic Canary, Serinus canaria
66. European Serin, Serinus serinus
67. Syrian Serin, Serinus syriacus
68. Red-fronted Serin, Serinus pusillus
69. Ethiopian Siskin, Serinus nigriceps
70. Black-headed Canary, Serinus alario
71. Yellow-crowned Canary, Serinus flavivertex
72. Cape Canary, Serinus canicollis
73. Tibetan Serin, Chionomitris thibetana
74. Lawrence's Goldfinch, Astragalinus lawrencei
75. American Goldfinch, Astragalinus tristis
76. Lesser Goldfinch, Astragalinus psaltria
77. Antillean Siskin, Spinus dominicensis
78. Eurasian Siskin, Spinus spinus
79. Pine Siskin, Spinus pinus
80. Black-capped Siskin, Spinus atriceps
81. Black-headed Siskin, Sporagra notata
82. Red Siskin, Sporagra cucullata
83. Black-chinned Siskin, Sporagra barbata
84. Andean Siskin, Sporagra spinescens
85. Olivaceous Siskin, Sporagra olivacea
86. Yellow-bellied Siskin, Sporagra xanthogastra
87. Yellow-faced Siskin, Sporagra yarrellii
88. Lowland Hooded Siskin, Sporagra magellanica
89. Yellow-rumped Siskin, Sporagra uropygialis
90. Andean Hooded Siskin, Sporagra capitalis
91. Saffron Siskin, Sporagra siemiradzkii
92. Thick-billed Siskin, Sporagra crassirostris
93. Black Siskin, Sporagra atrata

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