The Non-Passerines

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Metaves

Now you can see all 227 families below, so more discussion of Metaves is possible. There are several interpretations of the tree in Ericson et al. (2006a). After reading the paper and the supplementary material, I compromised on this version of Metaves. Metaves breaks into two parts. One, which I refer to as the ‘Columbimorphae’, contains what look like a rather heterogeneous collection of families: sandgrouse, pigeons, kagu, sunbittern, hoatzin, tropicbirds, mesites, flamingos, grebes. The other group contains nightjars and allies, hummingbirds, and swifts. Following Huxley 1867, it is often called the Cypselomorphae.

In fact, there had been hints that some of the ‘Columbimorphae’ were related, but even Sibley, Ahlquist, and Monroe missed the big picture. The Hoatzin may be basal in this group. Then there is split between pigeons, doves, and sandgrouse on one side and a rather heterogeneous group including mesites, flamingos, and kagu on the other. The extinct dodos and solitaires (Raphidae) are now considered to be embedded within the Columbidae (Pereira et al., 2007) rather than being a separate family. The latter grouping splits into a kagu/sunbittern group, and a group including flamingos, grebes, mesites, and tropicbirds. The kagu/sunbittern and flamingo/grebe pairings had been noticed previously.

Pereira et al. (2007) have recently done an extensive study of Columbiforme DNA. The Metaves hypothesis would suggest that the Cypselomorphae would make good outgroups for such a study, and by chance, they form a big part of Pereira's outgroups. Although the dodos and Rodriguez and Reunion solitaires (genera Raphas and Pezophaps) have been traditionally considered a separate family in the Columbiformes, the DNA says otherwise. Pereira et al. found that these two genera are buried deeply within the Columbidae.

The DNA testing revealed three major clades. A basal clade includes a subclade consisting of the New World genera Geotrygon, Leptotila, Zenaida, together with a subclade of New World pigeons (including the Passenger Pigeon), as well as typical pigeons, cuckoo-doves, and turtle-doves. It is sister to the other two clades together. One of them consists of the New World Ground-Doves. The other contains all other doves. The dodos and solitaires are nested well within this third clade.

The other half of Metaves is the Cypselomorphae. It includes nightjars and relatives (potoos, oilbird, frogmouths) and an interesting grouping of owlet-nightjars, swifts, treeswifts, and hummingbirds. The position of the owlet-nightjars is surprising as they had previously been thought closer to the main nightjar group. The exact relation of the other Cypselomorphae remains uncertain (e.g., Barrowclough et al., 2006; Mariaux and Braun, 1996; Mayr, 2002). Given the lack of solid information, I've put the Apodiformes (with owlet-nightjars) in one order, the nightjars and potoos in Caprimulgiformes, and the oilbird and frogmouths in their own orders, Steatornithiformes and Podargiformes.

In comparison, in the modified SAM list, the families in Metaves are scattered, but many are recognized as problematic. The modified SAM list has the Podicipediformes (grebes) and Phoenicopteriformes (flamingos) as isolated orders with unclear affinities. The Phaethontidae (tropicbirds) are in the Pelecaniformes, The Rhynochetidae (kagu), Eurypygidae (sunbittern), and Mesitornithidae (mesites) are considered Gruiformes while the Opisthocomidae (hoatzin) is placed in the Cuculiformes. The Pteroclidae (sandgrouse) are considered Charadriiformes and the Columbiformes (pigeons and doves) are an isolated order.

The other half of Metaves fares better. In 1995 Gill had already noted the Caprimulgiformes may be related to the Apodiformes.

Coronaves I: Waterbirds, Waders, and Cuckoos

Coronaves is variously regarded as dividing into 3 or 4 groups. I treat this as six subgroups within two major groups. The first major group, Coronaves I, includes the greater shorebird grouping (Charadriimorphae) and a group of mostly aquatic and semi-aquatic bird families (plus cuckoos and turacos), here called ‘Natatores’. (Natatores was a name used in the 19th century for waterbirds; although it has been attributed to Baird 1858, its ornithological use substantially predates him.) The terms Conglomerati and Cracrafti have been used for similar grouping that also include raptors.

There is a lot of evidence for the Charadriimorphae grouping (e.g., Ericson et al., 2003a; Paton et al., 2003; Cracraft et al., 2004). Other than arguments about the Herring Gull complex, the taxonomy of this order is now pretty well worked out. Many studies have found that gulls and alcids are closely related to the shorebirds. A recent development is the recognition that the buttonquails are part of it (Paton et al., 2003; Paton and Baker, 2006; Fain and Houde, 2007). The differences from the modified SAM list are the loss of the sandgrouse (to Metaves), the addition of the buttonquail, splitting the terns as a separate family, and some reordering of the families.

The sister group to the Charadriimorphae, the ‘Natatores’, is primarily an assemblage of water and wading birds. This includes most pelagic species (except tropicbirds), the big waders (but not sunbittern or flamingos), cranes and allies, bustards, and surprisingly, cuckoos. The un-named clade containing the Gaviiformes (loons) through Ciconiiformes (storks) is on solid ground. The rest of the ‘Natatores’ are more controversial. It's not clear that all of these birds belong in this superorder, or that everything related has been included. In particular, the inclusion of the cuckoos might be incorrect.

Again, we compare the modified SAM list. Big changes have hit both Pelecaniformes and Ciconiiformes. Most of the Ciconiiformes (except the Ciconiidae, storks) have been regrouped with the pelicans in a new Pelecaniformes. The remaining Pelecaniformes (except the Phaethontidae, tropicbirds, lost to Metaves) are renamed Phalacrocoraciformes. Finally, only the Ciconiidae (storks) are left in the Ciconiiformes. The Gaviiformes, Sphenisciformes, and Procellariiformes see little change.

There are also taxonomic issues in the families inside the Pelecaniformes. The Boat-billed Heron was previously considered to be the only member of the Cochlearidae. The status of two other monotypic families, the Shoebill and Hammerkop, has also been a perennial issue. The analysis of Ericson et al. (2006a) indicates that both are relatives of the pelicans. Indeed, the tree allows them to be lumped into the same family. We keep them separate not only because of their uniqueness, but also because the division between them seems to be ancient.

The Gruiformes are another area of major change, with three families lost to Metaves, one lost to Charadriiformes, one lost to Falconiformes, and one (Otididae, bustards) becoming its own order sister to the Gruiformes. On the plus side, the Musophagidae (turacos) find a new home in the Gruiformes, at least for now. Gill suggests the Cuculiformes might be related to the Musophagidae, and we see they also are placed in the ‘Natatores’.

Coronaves II: Land Birds

The other major piece of Coronaves consists of land birds, including raptors. Coronaves II has four parts: Passerines; parrots and cockatoos; falcons and seriemas; and everything else. The last group is further divided into hawks, American vultures, and a group I'm calling the ‘true Anomalogonatae’.

A notable feature of this taxonomy is that the American Vultures are classified in their own order, Cathartiformes. I have done this partly to reflect the considerable uncertainty about where they fit in. Contrary to Sibley-Monroe and 1990's-style checklists, they are not closely related to storks (see Cracraft et al., 2004; Gibb et al., 2007; Slack et al., 2007). Keep in mind that the vultures also appear no more related to the Accipitriformes than they are to anything else in this superorder.

I have followed Ericson et al. (2006a) concerning the placement of the hawks, falcons, and American vultures (Accipitriformes, Falconiformes, and Cathartiformes). However, this is controversial and several alternatives tree should be considered. Slack et al. (2007) put all three orders in Coronaves I. Gibb et al's (2007) results are more ambiguous. They present a tree showing the all three in a group that includes other members of both parts of Coronaves. They also present a network diagram that could be consistent with placing the hawks and falcons in Coronaves II. In that case, the hawks would be basal in Coronaves II. The fact that Gibb et al. include more birds from Coronaves II (an owl and parrot) in their analysis than Slack et al. did may account for this.

One big difference from the modified SAM list and recent AOU lists is the split of the Falconiformes, Accipitriformes, and Cathartiformes. Also, the Cariamidae (seriemas) are moved from the Gruiformes to the Falconiformes. The “terror birds” of ancient South America (Phorusrhacidae) are thought to be related to the seriemas (Alvarenga and Höfling, 2003).

The original Anomalogonatae were named by Garrod (1874), with the Strigiformes added later by Beddard (1898). Most of them have remained together in most taxonomic lists since then. One important defining character was the lack of an ambiens muscle (also lost by some unrelated birds). The ‘true Anomalogonatae’ are Beddard's Anomalogonatae, minus the Passeriformes and Cypselomorphae. This leaves a core group consisting of the Leptosomatiformes, Strigiformes, Coliiformes, Troganiformes, Upupiformes, Coraciiformes, and Piciformes. Ericson et al. (2006a) found it a monophyletic group, a group that includes a common ancestor and all descendants.

Other than their ordering and placement in the Anomalogonatae, the treatment of much of the remaining non-passerine families is close to that of the modified SAM list. Changes over time have mostly involved whether to consider certain groups families or sub-families. One interesting case is the Cuckoo Roller. It was originally considered a cuckoo, some affinities with the rollers were noted, and it has more recently been considered its own family, Leptosomatidae. It seems to be a basal family in the Anomalogonatae (which also puts it in its own order).

The owls, mousebirds, trogons are also placed in separate orders, as their affinities with the other Anomalogonatae remain unclear. The hornbills, which are split into Bucorvidae (ground-hornbills) and Bucerotidae (hornbills) form a grouping with the hoopoes and woodhoopoes. The rollers, bee-eaters, todies, motmots, and kingfishers form the Coraciiformes. The consensus seems to be to leave the three kingfisher subfamilies as subfamilies, and I follow that. That brings us to the Piciformes.

The classification of the Piciformes follows the AOU's South American Classification Committee rather than the AOU checklist. This means the Capitonidae (American barbets), Semnornithidae (toucan-barbets), and Ramphastidae (toucans) have family status and that requires the Asian and African barbets each have their own family (Moyle, 2004; Johansson and Ericson, 2003). All of the new world forms are more closely related to each other than to the old world barbets. The tree makes this clear.

This completes the non-passerines, of which there are over 4000 species. The other 5700+ species are Passerines, members of a single order, the Passeriformes. They are discussed further starting in the next section.

The Birds in 227 Families

PALEOGNATHES

STRUTHIONIFORMES

• Rheas (Rheidae)
• Ostrich (Struthionidae)
• Cassowaries (Casuariidae)
• Emus (Dromaiidae)
• Kiwis (Apterygidae)

TINAMIFORMES

• Tinamous (Tinamidae)

NEOGNATHES