Passeriformes

Corvids and Sub-oscines

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Most species of modern birds are passerines, songbirds. Although we have long known which birds are passerines and which are not, their relationships have been poorly understood. A comparison of Clements 5th edition (which uses an old taxonomy) and Howard-Moore 3rd edition (more recent, but not current) shows how much revision has been necessary. There are still many passerines that are classified in the wrong family (and genus) which makes it harder to determine proper family boundaries and relations. Recent work on passerine taxonomy has done much to clarify the situation.

It is important to recall that bird families should be natural groups. They should be monophyletic, meaning that they include the most recent common ancestors and all of its descendents. I also impose the requirement that a family should appear to be a natural group. The second requirement is somewhat subjective, but it will be appealed to several times below.

We now consider the passerines. The basal split is between the New Zealand wrens and all other songbirds (Barker et al., 2002; Barker et al., 2004). The common ancestor of the suboscines and the oscine passerines comes after the split between the New Zealand wrens, so we cannot put the New Zealand wrens in the suboscines. That forces them into their own family (this wasn't known when Gill wrote his text, and he considers them suboscines). The next split is between the oscines and suboscines. The oscine group is bigger, so we consider it the main trunk.

Suboscines

Like the passerines as a whole, the suboscines have generally been identifiable as suboscine, but teasing out the relationships between the suboscines has been difficult. The suboscine branch is comprised of three groups, an old world group (Eurylaimida), a new world group containing the ovenbirds (Furnariida), and a group consisting of the tyrant flycatchers and allies (Tyrannida). The old world suboscines, the pittas, asities, and broadbills have recently been reorganized (Moyle et al., 2006). The Sapayoa, Sapayoa aenigma, has finally found a new home in this group as the only new world representative of the Eurylaimida.

The main split among the Eurylaimida is between the pittas and the rest. Moyle et al. found that the broadbills were not a natural grouping. Some are more closely related to the Sapayoa and the asities than they are to the other broadbills. This list considers the broadbills to consist of two families, one of them sister to the asities, the other sister to the rest of the broadbills, asities, and Sapayoa.

The latest changes from the SACC have been used in reconstructing the Furnariida, although we follow a different ordering based on Chesser (2004; see also Irestedt et al., 2002; Rice, 2005). The Grallariidae (antpittas) have been separated from the Formicariidae (ant-thrushes) because the tapaculos are more closely related to the remaining ant-thrushes than the antpittas are. Similar issues lead to a separation of the Melanopareiidae (crescent-chests) and the Rhinocryptidae (tapaculos). The Pittasoma antpittas have been moved to the Conopophagidae. Finally, the Dendrocolaptidae (woodcreepers) have been folded into the Furnariidae (ovenbirds). This was necessary because the woodcreepers are nested within the Furnariidae. If they were separated, the Furnariidae would no longer be monophyletic. The SACC has introduced subfamilies: Sclerurinae (leaftossers and miners), Furnariinae (ovenbirds), and Dendrocolaptinae (woodcreepers) to reflect this.

That leaves the Tyrannida. Relationships here have been difficult to tease out and roughly 400 species remain together in the Tyrannidae. There is some inner structure in the Tyrannidae (see the discussions in Fitzpatrick, 2004a, b; Tello and Bates, 2007; Rheindt et al., 2008). Although Fitzpatrick's presentation of the Tyrannidae has considerable truth in it, it also involves some oversimplification and does not properly account for Sibley and Monroe's Pipromorphinae. The recent analyses of Tello and Bates uncovered a basal clade including the Royal Flycatcher. This has been dubbed the ``Myiobius assemblage''. At a minimum, it includes Myiobius, Terenotriccus, and Onychorhynchus. Tello and Bates and Rheindt et al. have also added more genera to Sibley and Monroe's Pipromorphinae. A number of genera remain difficult to place, and some do not seem to belong to any of the major clades. Due to the amount that remains unknown, it is premature to split the Tyrannidae into families or subfamilies. Even some of the species need adjustment. For example, La Sagra's and Stolid Flycatcher seem to be mutually paraphyletic and either need to be further split or regrouped (Joseph et al. 2004).

Considerable progress has been made classifying the other tyrannids (Chesser, 2004; Ericson et al. 2006b; Ohlson et al., 2007). The arrangement of families is based on Ericson et al. One key was the recognition that Sapayoa, sometimes considered a manakin, was not part of the Tyrannida. Another set of ringers in the manakins were the birds in the genus Schiffornis. These taxonomically troubling birds are more closely related to the becards (Pachyramphus) and tityras. The family Tityridae has been formed about them, gathering together several genera previously classified as Tyrannidae, Cotingidae, and Pipridae.

The genus Piprites remains troublesome, but may belong in the main Tyrannid group. The Phytotomidae (plantcutters) are moved into the Cotingidae. Oxyruncidae (sharpbill) is retained as a monotypic family for now, partly because DNA testing has been inconclusive, partly because it has often been regarded as its own family, and partly because it is an unusual tyrannid. There is some evidence in favor of including it in the Tityridae (see Ericson et al., 2006b).

Oscines: Corvids

Unlike the passerines and suboscines, the corvid assemblage has not been readily identifiable and taxa are still being moved between the corvid groups and the Passerida. Sibley, Ahlquist, and Monroe took the first step of gathering the corvids together (albeit imperfectly). This is reflected in the modified SAM list and the 3rd edition Howard-Moore checklist, which did a lot to group them in reasonable families. Checklists of a more traditional sort, such as Clements 5th edition list place the corvid assemblage all over the map.

Sibley and Ahlquist's view was that the remaining passerines split cleanly into a corvid group and a group containing everything else (Passerida). Further study of these families has shown that reality is more complex. Unlike the Passerida, the SAM corvids are not a monophyletic group. There are several groups that we might call paracorvids (Menurida, Climacterida, Meliphagida, Pomatostomida, possibly Orthonychida), that branch off separately from the main corvid group, Corvida (see Barker et al., 2004).

Many of the changes since the mid-90's are shown in a side-by-side family listing below.

Paracorvids in Four Parts

The basic structure of the tree appears in Barker et al. (2004). The parvorders Menurida, Climacterida, Meliphagida, and Pomatostomida branch off successively, leaving the main corvid group, Corvida.

The first branch, Menurida consists of the lyrebirds (Menuridae) and scrub-birds (Atrichornithidae). Ericson et al. (2002b) found the Menurida sister to the rest of the oscines, but did not include scrub-birds in his analysis. Morphological analyses had placed the scrub-birds next to the lyrebirds. A genetic analysis was recently carried out by Chesser and ten Have (2007). It concurs that the lyrebirds and scrub-birds are sister families, and also concurs with basic tree we present here.

The Climacterida are the next branch. There are two families here: Australian treecreepers (Climacteridae) and bowerbirds (Ptilonorhynchidae). Genetic evidence has shown that the probably extinct Piopio was actually a basal member of the bowerbirds. It had sometimes been classified with the Pachycephalidae and there had been questions about whether it related to the birds-of-paradise (Christidis et al., 1996).

There are five families in the Meliphagida: Australasian wrens (Maluridae), bristlebirds (Dasyornithidae), thornbills and gerygones (Acanthizidae), pardalotes (Pardalotidae) and honeyeaters (Meliphagidae). We follow the order found in Driskell and Christidis (2004), with each family successively branching off. The Acanthizidae, Dasyornithidae, and Pardalotidae have sometimes been placed in the same family under the name Pardalotidae. They are treated as separate families because they seem to be on separate branches of the Meliphagida tree. They can't be lumped together without also including the Meliphagidae.

The honeyeaters were substantially restructured by Sibley and Ahlquist (1990), losing the genera Cleptornis (Passerida), Oedistoma and Toxorhamphus (Melanocharitidae), and Promerops (Promeropidae), but gaining Epthianura and Ashbyia from the defunct Epthianuridae. Driskell and Christidis concur that Epthianura and Ashbyia are honeyeaters. It was later found that the Bonin Honeyeater, Apalopteron familiare belongs in the Passerida (Springer et al., 1995). More recently, Cracraft and Feinstein (2000) found that MacGregor's Bird-of-Paradise, Macgregoria pluchra, is actually a honeyeater, while Driskell et al. (2007) found that the Stitchbird, Notiomystis cincta belongs in the Corvida, where it becomes a monotypic family. The now-extinct Hawaiian honeyeaters (genera Moho and Chaetoptila) were probably not paracorvids, but rather passerids; details on this have not yet been published.

The last two paracorvid families are the logrunners (Orthonychidae) and Australian babblers (Pomatostomidae), which together form the Pomatostomida. They are placed together on a branch as in Barker et al. (2002). However, Barker et al. (2004) considered each a separate paracorvid branch, while Ericson et al. (2002b) has them grouped with the Meliphagida. In any event, they split off before the division between the Corvida and Passerida, which means they are in the paracorvids.

The Core Corvids: Corvida

This brings us to the Corvida. It divides into two superfamilies, ‘Callaeatoidea’ and Corvoidea. The ‘Callaeatoidea’ were assembled from a variety of misclassified taxa. In it are four families: Stitchbird (Notiomystidae), the New Zealand wattlebirds (Callaeatidae), satinbirds (Cnemophilidae), and berrypeckers (Melanocharitidae). This whole group has been run though a taxonomic blender. The monotypic family Notiomystidae was previously considered a honeyeater (Driskell et al., 2007). The division between the Stitchbird and other families in its clade seems quite ancient, so it is given its own family. The New Zealand wattlebirds were thought to be closely related to the vangas (Corvoidea). The satinbirds, Cnemophilidae, were formerly considered birds-of-paradise (Paradisaeidae). Finally, the berrypeckers were formerly placed in Passida, near Passeridae! They had themselves been augmented with the former honeyeaters Oedistoma and Toxorhamphus by Sibley, Ahlquist, and Monroe.

The Corvoidea proper start with the Neosittidae (sitellas), which were formerly considered part of the Pachycephalidae. Next come the vireos. The vireos were once thought to be closely related to the wood-warblers (Parulidae). When Sibley and Ahlquist discovered the corvid group, they found that the vireos (and shrikes) were members of it. Some thought it odd that the vireos, with an old world origin, had no remaining old world members. This has turned out to be incorrect. They do have some old world relations. They've been hiding out among the babblers (Timaliidae). One is the Erpornis zantholeuca (nee Yuhina zantholeuca). Rather than being a babbler like the other Yuhinas, it is actually a vireo relative. On his bird families website, Don Roberson placed it in its own family to emphasize its special status as the only old world vireo. Recently, Reddy and Cracraft (2007) found that the shrike-babblers (Pteruthiidae) are also vireo relatives. Right now, the choices are either to call them all vireos or to have 3 families of vireos and allies. Since there are now more old world vireos, Epornis and Pteruthius, they seem less special. Thus I take the former route for the present, keeping in mind that it might make sense to split the Vireonidae into 3 families.

The next branch is the cuckoo-shrikes (Campephagidae). This family was once considered related to bulbuls or waxwings. It has moved to the Corvoidea, losing the genera Hemipus and Tephrodornis to the vangas (Fuchs et al., 2006b, 2007) along the way.

The Pitohuiidae are worth noting. They're poisonous! (Dumbacher et al., 1992.) The pitohuis could have been placed in the Oriolidae, but their distinctive poisonous nature makes the a good candidate for a separate family. The other toxic bird, Ifrita kowaldi, does not seem to be closely related to the pitohuis, although its true affinities remain unknown (Dumbacher et al., 2000; Jønsson et al., 2008). You can read more about these interesting birds at Dumbacher's website. A new paper by Jønsson et al. (2008) finds that the genus Pitohui includes a number of birds that are not that closely related. In fact, this is true of other genera in the families in the range from Campephagidae though Oriolidae (e.g., Fuchs et al., 2007). I think the overall picture for these families will need some revision. The fact that several genera in here need their boundaries redrawn only adds to the current confusion.

In contrast, several recent papers by Fuchs et al. (2004, 2006b, 2007) and Moyle et al. (2007) have clarified the relations between the woodswallows (Artamidae), butcherbirds (Cracticidae), vangas (Vangidae), wattle-eyes and batises (Platysteiridae), bristlehead (Pityriaseidae), ioras (Aegithinidae), and bush-shrikes and puffbacks (Malaconotidae). All are on the same branch of the Corvoidea. The vangas gain several genera: Megabyas and Bias from the Platysteiridae, Hemipus from the Campephagidae, and Philentoma, Prionops, Tephrodornis from the former Prionopidae (the remainder forms the Platysteiridae). The also gain Newtonia and Mystacornis from the Sylviidae.

The painted berrypeckers (Paramythiidae) are removed from the Melanocharitidae (which itself has migrated from the Passeroidea) and placed in their own family. In principle, the Paramythiidae could be lumped with the Oriolidae, but like the pitohuis they are distinctive enough to get their own family.

There are a couple of interesting small changes in the Corvidae. Hume's Groundpecker (or Ground Jay), Pseudopodoces humilis was thought to be a corvid, the smallest of them. It is so listed even in Howard and Moore, but recent evidence (James et al., 2003; Gill et al., 2005) suggests it is actually a parid! The Crested Jay, Platylophus galericulatus, is another ringer. It's definitely not a corvid. It may be a starling (Clench, 1985).

The next section continues the discussion of the passerines.

PASSERIFORMES

Version 1.12: Feb. 7, 2008. Copyright © 2007, 2008 by John H. Boyd III << Non-Passerines     |     Passerida I >>