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Passeroidea

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Passeroidea

The last superfamily, Passeroidea, contains approximately 15% of all bird species. The exact membership of this group is still being worked out. In the past, Laniidae and Vireonidae (now in Corvoidea), Hirundinidae (now Sylvioidea), and Bombycillidae (now Muscicapoidea) have been included. Even Gill, who wrote after publication of the Sibley-Monroe checklist, includes Melanocharitidae (now Corvida) and Alaudidae (Sylvioidea) in the Passeroidea. He did not include Irenidae (fairy bluebirds) and Chloropseidae (leafbirds), but considered them part of Corvida.

The Passeroidea start off with several families that split off in ones or twos—sugarbirds, flowerpeckers and sunbirds, fairy bluebirds and leafbirds. I put the sugarbirds (Promeropidae) as the basal family in Passeroidea (Beresford et al., 2005). There is uncertainty about whether they even belong in Passeroidea, and there is some support for moving them near the Hyliotas (Fuchs et al., 2006a). As mentioned above, until recently the fairy bluebirds and leafbirds were considered corvids.

The Passeroidea include a new family, Urocynchramidae. This family contains one species, the Pink-tailed Bunting (Przevalski's Bunting), which was previously thought to be an Emberizid bunting (Groth, 2000).

The remainder of the Passeroidea splits into two groups. I've marked the smaller one as the Ploceid group. This includes the Olive Warbler, accentors, weavers, whydahs and indigobirds, and the estrildid finches. Not long ago, the Olive Warbler was considered one of the wood warblers, although there was some question as to whether it was really a warbler. The genes tell the tale, and the tale is that it is not a warbler (Groth, 1998). Instead, its closest relatives are the accentors, none of which are native to the Americas.

As illustrated on the tree, the Ploceid group is sister to the remaining Passeroidea. The next exact ordering of the next two families is made clear by the genetic code. The nine-primaried oscines and the Motacillidae (wagtails and pipits) share a genetic trait that other birds lack. They have DNA that codes for an extra three amino acids in a particular location on the c-myc gene (Ericson et al., 2000). Old world sparrows don't have the insertion. Wagtails and pipits do. Nine-primaried oscines do. That means the wagtails and pipits are more closely related to the nine-primaried oscines than the Passeridae are. Moreover, the traditional view that the Passeridae are closer to the nine-primaried oscines than the Ploceidae are is also correct. Summing up, Passeridae is the sister group to the wagtails, pipits and nine-primaried oscines. Equally, the wagtails and pipits are sister to the nine-primaried oscines.

Nine-primaried Oscines

The nine-primaried oscines don't actually have nine primaries. The tenth primary usually reduced and is hidden by the ninth covert (Hall, 2005). This group is basically Sibley and Monroe's Fringillidae. It includes the Fringillidae, Drepanididae, Emberizidae, Parulidae, Thraupidae, Cardinalidae, and Icteridae from the modified SAM list (see also Bledsoe, 1998). There are roughly a thousand closely-related species in this group and its evolution has not been fully worked out. However, one thing that stands out is the essential correctness of the family boundaries on the SAM list (original and modified), and how this contrasts with the more traditional family boundaries followed used by the AOU, Clements, and Howard-Moore lists. One key difference is which family gets the various neotropical finches. Are they tanagers or sparrows? Sibley, Ahlquist, and Monroe say most are tanagers, and they seem to be mostly right.

A group including Barker, Burns, Klicka, Lanyon, and Lovette are putting together a comprehensive dataset concerning these species. The overall picture will be much clearer when their analysis is finished. Some portions of it have been published, and other pieces have been presented but not yet published.

The reorganization starts with the recognition that even the larger traditional families: Fringillidae, Emberizidae, Parulidae, Thraupidae, Cardinalidae, and Icteridae, contain genera and species that don't belong to them. Many of these taxa have been identified in the last two decades.

Olive Warbler Out

We start with one that has already been mentioned: the Olive Warbler. The SAM list removed it from the Parulidae, listing it separately within the Fringilliade. They didn't go far enough. It is not only not one of the Parulidae but is not even a nine-primaried oscine (Groth, 1998). Rather, it is akin to the accentors. Accordingly, it is placed next to the accentors in its own family, Peucedramidae.

Fringillidae = Finches + Euphonias + Hawaiian Honeycreepers

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, as are the Hawaiian Honeycreepers, Drepanidinae (Groth, 1998; Klicka et al., 2000; Yuri and Mindell, 2002).

Calcariidae: Longspurs and Snow Buntings

Genetic evidence shows the longspurs and snow buntings are not sparrows (Emberizidae). Rather, they should be placed in their own family, Calcariidae, and that this family branches off next (Klicka et al., 2003).

Five Families of Emberizids

This leaves us with five major groups: the wood-warblers, blackbirds, sparrows, cardinals, and tanagers. These naturally divide into two groups: one containing the warblers, sparrows, and blackbirds, the other containing the cardinals and tanagers.

Parulidae: Warblers

A number of warblers true affinities have long been questioned. The Yellow-breasted and Wrenthrush are two that come immediately to mind. Klicka et al. (2007a) show that the Granatellus Chats are cardinals rather than warblers. This paper also suggests to me that the remainder of the questionable warblers are in fact warblers! However, they are distinct from the rest of the warblers in the sense that all appear basally in the warbler group. I'm a little concerned about this conclusion since I understand that Klicka and others working on this feel they are not warblers at all. Nonetheless, the information they've published so far (especially Klicka et al. 2007a and Lovette and Bermingham 2002) don't seem to back this up. Accordingly, I place Teretistris, Xenoligea, Microligea, Leucopeza (Semper's Warbler), Zeledonia (Wrenthrush), and Icteria (Yellow-breasted Chat) with the warblers. This also makes Spindalis a warbler (Stripe-headed Warbler?).

There are some indications that the tanager genera Nesospingus (Puerto Rican tanager), Calyptophilus (chat-tanagers), and Phaenicophilus (palm-tanagers) are closely related to Spindalis, but the evidence is not clear-cut (they could be sparrows or tanagers), and for now we leave them as tanagers. If they are not tanagers, it would mean that all birds with “tanager” in their name in the United States and Greater Antilles, are not really tanagers! However, there are some tanagers in the Greater Antilles that were previously considered sparrows.

Icteridae: Blackbirds

Amazingly, the blackbirds get away unscathed by all these changes. No species are added, nor are any taken away. Of course, that may change with further analysis.

Emberizidae: Sparrows

As mentioned above, most of the neotropical finches belong in the tanagers, as do the Gubernatrix and Paroaria cardinals. The neotropical finches that remain are the Brush-Finches, as well as the Large-footed, Yellow-thighed, and Yellow-green Finches, All that the sparrows get in return are the Urothraupis and Chlorospingus Bush-Tanagers and the Tanager Finch Oreothraupis arremonops. This reduces the family to about half its former size. It is not entirely clear that the old world sparrows (mainly Emberiza) belong to the same family as the new world sparrows (e.g., Alström et al., 2008). Whether the sparrows are unified or in separate families, both parts also require some internal reorganization (Klicka and Spellman, 2007b; Alström et al., 2008).

Cardinalidae: Cardinal-Grosbeaks

Several genera within the cardinals and tanagers need reclassification (Burns, 1998; Klicka et al., 2007a). The Piranga, Habia, and Chlorothraupis tanagers are really Cardinalidae. Yes, that means all of our North American tanagers are really cardinals. The Cardinalidae also gain the Amaurospiza sparrows and Granatellus warblers. However, the Cardinalidae lose the saltators, along with Parkerthraustes and Porphyrospiza, all of which are really tanagers. The number of species in the family remains about the same.

Thraupidae: Tanagers

Besides losing the euphonias and chlorophonias, and North American Tanagers, the Thraupidae may ultimately lose Mitrospingus (see below). However, the tanager-finches that are often considered Emberizidae (but not in SAM-style lists) end up in the tanagers, bringing Thraupidae to over 370 species. For the present, the tanagers must remain a very large, very heterogeneous 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, although there seem to be about a dozen major clades (Klicka et al., 2007a; see also Burns, 1997; Burns et al., 2003).

One of the better established clades is the Thylospiza of Burns et al. (2002). The Thylospiza consist of the Bananaquit, the Galapagos finches, the other island finches, the Orangequit, and the grassquits (excepting the Blue-black Grassquit). Except for the Bananaquit, all were previously considered sparrows! The Bananaquit was considered to be in its own family, which was once thought to include the American honeycreepers. Interestingly enough, there seems to be another tanager clade that includes the honeycreepers, but without the Bananaquit (Burns et al., 2003).

Most of the neotropical finches also join the tanagers. There is a seedeater and seed finch group. Another clade includes the grass-finches and pampa-finches, while a third contains the warbling finches and Hemispingus (Lougheed et al., 2000). The flower-piercers, Peg-billed Finch, sierra finches and yellow finches form a clade with the conebills. The Paroaria cardinal finches are in a clade with the Diuca finches and others. As mentioned above, the main group of neotropical finches that remains in the Emberizidae are the brush-finches.

There are also some more classic tanager clades. One includes the Mountain-Tanagers and other tanagers of the Andean slopes. Another primarily consists of the main tanager genus Tangara (Burns and Naoki, 2004), augmented by several species previously thought to be in Thraupis. There is a clade that includes the Tachyphonus and Ramphocelus Tanagers. The Plushcap, formerly considered a monotypic family, may be a member of this clade. Finally, the Saltators are in their own group.

Mitrospingus

The latest data (Klicka, 2007a) suggest that Mitrospingus tanagers belong in their own family within the warbler/sparrow/blackbird group. However, this is not yet decisive and we will leave Mitrospingus in the tanagers pending further information. However, it may be sister to the warblers, sparrows, and blackbirds.

The Basic Sequence

The Fringillidae are sister to the rest of the nine-primaried oscines. The new Longspur-Snow Bunting family (Calcariidae) comes next. It is one of the two families in this group that have old world representatives. The remainder breaks into two groups, the cardinal-tanager group and the warbler-sparrow-blackbird group. I place the warblers next, followed by the blackbirds and sparrows. Since the cardinal-tanager group is bigger, it goes at the end, with the cardinals first and tanagers last.

Roberson's Family List 7th edition

The list of families above is very similar to Roberson's family list. We both end up with 227 families, but there are a few differences. He considers the Frogmouths two families, lists Erpornis separately from the Vireos, does not lump the Prionopidae (helmet-shrikes) with the vangas, lumps the Zosteropidae with the Timaliidae, and considers the Rhabdornithidae (Philippine creepers) separate from the starlings. My reading of the Fuchs et al. series (2004, 2006b, 2007) is that the helmet-shrikes are vangas. I read Cibois (2003) as consistent with keeping Zosteropidae separate from the Timaliidae. Finally, Lovette and Rubenstein (2007) seems to have put an end to Rhabdornithidae.

I handle the suboscines differently, with two extra families in the Furnariida: the Melanopareiidae (crescent-chests) and Grallariidae (antpittas) as well as treating Oxyruncidae (sharpbill) as a separate family in the Tyrannida. I suspect it belongs in the Tityridae, but the evidence is currently equivocal (see the discussion in Ericson et al., 2006b and Ohlson et al., 2007).

Version 1.1: Dec. 18, 2007. Copyright © 2007 by John H. Boyd III << Passerida I