Core Passeroidea II

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 44 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Passerid Clade

While the Estrilid clade is primarily old world and southern, its Passerid sister clade is cosmopolitan. More precisely, the nine-primaried oscinces are cosmopolitan, while their sister group, the Passeridae, are restricted to the Old World.

Passeridae: Sparrows, Snowfinches

8 genera, 41 species HBW-15

The Passeridae are seed-eating birds, primarily distributed in the southern portion of the palearctic, but ranging into the afrotropics. Several passerids are closely associated with humans, especially the ubiquitous House Sparrow, which has spread worldwide.

Passeridae tree Dickinson et al. (2003) consider Plocepasser, Histurgops, Pseudonigrita, and Philetairus to be Passeridae. However, Groth (1998) found them to be in the Ploceidae.

The most astonishing change in the Passeridae is due to Fjeldså et al. (2010). They found that the Cinnamon Ibon, long thought to be an abberant white-eye, is actually a sparrow. This is a canopy bird from the cloud-forest of Mindanao, in the Philippines. It's a long way from anything we think of as sparrows. It's also a long way physically. The nearest native populations of any sparrows are on the Asian mainland. Nonetheless, it is a sparrow. Indeed, Fjeldså et al. report than its skull is similar to a sparrow, and that it has many other features in common with the other Passeridae.

The order of species in Passer does not seem to be quite correct, but Allende et al. (2001) do not consider enough species for reorganization of Passer to be helpful at this time. Qu et al. (2006) was helpful concerning the snowfinch genera. Note that the true affinities of the Pale Rockfinch are uncertain, although it is sometimes included in Petronia.

Different sources treat the Rufous Sparrow (Passer motitensis) complex differently. Here it is treated as 5 species: iagoensis, motitensis (including benguellensis and subsolanus), rufocinctus (including cordofanicus and shellyi), insularis, and hemileucus. See Kirwan (2008) for arguments that the small pale hemileucus should be split from insularis. As in the IOC list, the name Great Sparrow is used for the narrowly defined Passer motitensis.

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). This sometimes occurs in birds outside the nine-primaried oscine group, but is most characteristic of this group.

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, Pipits

7 genera, 67 species HBW-9

Motacillidae tree The overall organization of the pipits and wagtails is based on Outlaw and Voelker (2006b). Their results agree with topology (c) in Voelker and Edwards (1998), which presents several alternatives. Most of the alternatives also have Anthus sister to the other pipits, and Dendroanthus in a basal position. There is some ambiguity about whether it is closer to the pipits or wagtails or maybe basal to both.

Johansson et al. (2008b) suggest that Bocage's Longbill (Amaurocichla) is related to the wagtails and pipits. It may be closer to the wagtails. I don't have an obvious place to put Alpine Pipit, so it is also flagged in blue. There have been questions about the status of the Kimberley Pipit, Anthus pseudosimilis, but I include it, as does Tyler in HBW-9 (2004).

Within the pipits, I've also followed Outlaw and Voelker (2006b). Using the same data, Voelker (1999) found a slightly different tree, which is also used by Alström and Mild (2003).

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.

Pending further analysis, I accept this interpretation. However, the nuclear DNA trees are not fully resolved. The tree below is a compromise between the two. I follow the nuclear tree where it is resolved, and use the mitochondrial results to break ties. The result even makes a certain amount of sense both biogeographically and in terms of plumage. well

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.

Greater 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 includes 9 families in the greater Fringillidae.

Fringillidae: Finches, Euphonias

62 genera, 217 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). The Hawaiian Honeycreepers, formerly considered their own family Drepanididae, are actually a clade within the Carduelinae (Yuri and Mindell, 2002; Arnaiz-Villena et al., 2007b).

Fringillidae tree The arrangement of the finches is based on several sources. I mainly relied on Groth (1998), Klicka et al. 2007, and Arnaiz-Villena et al. Arnaiz-Villena et al. (1998, 1999, 2001, 2007a, b, 2008). Their studies have included a substantial number of finch species. However, some monotypic genera have not been sequenced yet (e.g., Callacanthis, Chaunoproctus, Kozlowia, Neospiza). The tribes within the Carduelinae derive from the results of Yuri and Mindell (2002), Arnaiz-Villena et al. (2007a), and Nguembock et al. (2009a). Except for the basal position of the Coccothraustini, there is a lack of consensus on the relations between the Cardueline tribes.

The Fringillidae start with a basal palearctic group, Fringillinae. Fringilliane 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 almost certainly an Old World species. Something must connect it with the neotropic Euphoniinae, but that something is missing. Being geographically contiguous and more similar in appearance, the position of the Carduelinae is easy to understand. Euphoniinae is a mystery.

As mentioned above, how the tribes with the Carduelinae relate is somewhat murky. The holarctic Coccothraustini seem to be basal. The results of Nguembock et al. (2009a) suggest that the Carpodacini and Pyrrhulini are more closely related to each other than to the rest. The information that is available is pretty equivocal on the overall relationships of these tribes. On biogeographic grounds, it seems more likely that the Hawaiian honeycreepers have a North American origin.

Fringillinae: Chaffinches

Euphoniinae: Euphonias, Chlorophonias

Carduelinae

Coccothraustini: Grosbeak Finches

Drepanidini

Taxonomy within the honeycreepers is somewhat contentious. The ordering is based on a combination of Fleischer et al. (2001), Arnaiz-Villena et al. (2007b), and Reding et al. (2009), using additional morphological and osteological information from Pratt (2001) and James (2004). These don't fit together that smoothly. As a result, the tree I supply is somewhat conjectural.

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.

Click for Drepanidini tree
Click for Drepanidini tree

Pyrrhulini: Bullfinches and Mountain-Finches

The deciding factor in treating Pyrrhulini and Carpodacini as separate tribes is that I'm not entirely convinced they are each other closest relatives.

The extinct Bonin Grosbeak is rather uncertainly placed at the base of Pyrrhulini. The rest of the Pyrrhulini is on firmer ground. There are two clades. The first includes the Golden-naped Finch, Pyrrhoplectes epauletta. It belongs near the Pyrrhula (Nguembock et al., 2009a), as does Pinicola enucleator (Arnaiz-Villena et al., 2001, 2008). Pinicola subhimachala seems to belong in Carpodacini (Groth 1998, 2000).

The remaining Pyrrhulini also seem to comprise a clade. Callacanthis, Rhodopechys, and Bucanetes are generally considered closely related, with the last two sometimes congeneric. Kirwan and Gregory (2005) argue in favor of three genera. Their position in the phylogenetic tree is based on Arnaiz-Villena et al. (2008). Of the three genera, they only studied Bucanetes. Arnaiz-Villena et al. also found that the Dark-breasted Rosefinch, formerly Carpodacus nipalensis belongs in the same clade close to Leucosticte. Blanford's Rosefinch is thought to be very close to nipalensis, so I have moved it also. Rather than folding them in Leucosticte, I restore the genus name Procarduelis, which belongs to nipalensis.

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.

Click for Pyrrhulini/Carpodacini tree
Click for
Pyrrhulini/Carpodacini tree

Carpodacini: Rosefinches

The American Carpodacus finches move to Burricini. The Crimson-browed Finch has been moved from Pinicola based on Groth (1998, 2000). The genus name Propyrrhula is restored for it (Pinicola belongs to the Pine Grosbeak). This leaves us with a group of mostly Asian species, with distributions centered around the Himalayan region.

The remaining Carpodacini have been considered part of Carpodacus. The entire tribe could be thought of as Carpodacus in a broad sense. An earlier version of the TIF list merged Haematospiza and Uragus into Carpodacus. That has been undone. By resurrecting the genera Erythrina and Rubicilla, I am able to both better reflect the phylogenetic tree and retain the genera Haematospiza, Kozlowia, and Uragus.

Note that the correct type species of Carpodacus is Pallas's Rosefinch, Carpodacus roseus (Banks and Browning, 1994).

Burricini: Purple Finches

The North American Carpodacus finches are not that closely related to the true Carpodacus finches and a new genus is needed. House Finches have sometimes been put in a separate subgenus Burrica (2nd-5th AOU checklists), so I use that for the American purple finches. This is the only genus in Burricini. Where exactly Burricini fits remains unclear.

Carduelini: Canaries, Siskins and allies

This brings us to the Carduelini. Carduelini tree The breakup into genera is primarily based on Nguembock et al. (2009a), while 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. The genera Carduelis and Serinus both require substantial surgery. The alternative for Carduelis would be to put all of the Carduelini into one genus! For Serinus, the alternative is a bit less drastic, it would only put Carduelini II and III into the same genus. Neither of these outcomes is particuarly desirable, so both Serinus and Carduelis have been divided into several pieces each.

There are three major clades in the Carduelini. The first clade, Carduelini I, starts with Neospiza. The true position of Neospiza is unclear. It is here because it might be related to Linurgus. But, it might not even be a Fringillid and has sometimes been considered a weaver.

Kirwan and Grieve (2007) argue that Rhynchostruthus includes three species. Although molecular data is not available for Rhynchostruthus, James (2004) places it here with Rhodospiza and Chloris. Zamora et al. (2006) found much the same relations between Rhodospiza and the Chloris greenfinches. Chloris is one of the genera carved out of Carduelis.

The second and third clade are sisters. The first of these, Carduelini II, is primarily African. I include a species-level phylogeny, although it's a bit shaky (notice all the question marks and polytomies). Carduelini II contains four genera separated from Serinus: Crithagra, Dendrospiza, Pseudochloroptila, and Ochrospiza. It's not entirely clear whether Pseudochloroptila belongs in Dendrospiza, so it is kept separate.

The third clade, Carduelini III, is more cosmopolitan. The palearctic Linnets (Linaria, formerly Carduelis) are basal here. After them we again have two pieces. The first piece starts off with the Mountain Serin of Indonesia and the Philippines. It's unclear where the Mountain Serin of fits in. It's sometimes considered part of Serinus, but here gets its own genus, Chrysocorythus. This seemed the most likely spot. I put the Tibetian Serin next. It's now placed in Chionomitris rather than Serinus, sister to Spinus. Several Carduelis siskins that 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 the recent AOU proposal).

It should be noted that the placement of Spinus is less certain than it may appear in Nguembock et al.'s (2009a) combined tree (their Figure 4), as some of the individual genes yield different results (e.g., sister to Astragalinus + Sporagra). This is the subject of some discussion in Nguembock et al.

This group of birds is sister to the holarctic Redpolls (now Acanthis rather than Carduelis) and crossbills (Loxia). It's not clear how many redpoll species there are. The European birds have sometimes been separated as Lesser Redpoll, Acanthis cabaret, (e.g., BOU) but genetic studies have failed to find any differences between them and Common Redpolls (Ottvall et al., 2002). Worse, Marthinsen et al. (2008) found little genetic difference between any of the redpolls!

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).

The last piece includes the true Carduelis and Serinus finches of the palearctic. The latter is sister to a collection of American finches formerly considered part of Carduelis. These former Carduelis finches are the North and Middle America genus Astragalinus (called Pseudomitris by Nguembock et al., 2009a) and the Middle and South American genus Sporagra (AOU's proposed Pyrrhomitris).

AOU rejected a proposal to make these last two generic splits, also proposed by Nguembock et al. However, they have chosen different genus names than Nguembock et al. The AOU's suggested Astragalinus (Cabanis 1851) appears to have clear priority over Nguembock et al.'s Pseudomitris (Cassin 1865).

The case of the South American siskins is not so easy. 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. If I'm wrong about how to handle the parts, Bonaparte's Pyrrhomitris might possibly have priority, even though Bonaparte refers to Sporagra on page 516, the page before he establishes Pyrrhomitris. Given all that, the name Sporagra appears to be correct.

Click for Carduelini tree, part I
Click for Carduelini tree
part I

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