Metaves I

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

NEOAVES Sibley et al., 2008

Neoaves is the sister clade to Galloanserae. It has two parts: Metaves and Coronaves.

METAVES Fain & Houde, 2004

Metaves contains about 10% of the avian tree: 946 extant species in 211 genera and 16 families. Many of the Metavian families have long puzzled ornithologists. They just didn't seem to fit comfortably anywhere on the avian taxonomic tree. One of the attractions of the Metaves hypothesis is that it packages together so many troublesome taxa. The Metaves hypothesis remains controverisal and is a current topic of research.

COLUMBIMORPHAE

Metaves includes two big groups. This is the smaller one.

Mirandornithes Sangster, 2005

Regardless of whether Metaves holds, the association of the Flamingos and Grebes is likely to remain (see Mayr, 2008). The name “Mirandornithes” was introduced for this clade by Sangster (2005). At least one analysis suggests that the Mesites are close to the the flamingos and grebes (Ericson et. al., 2006a), but I follow Hackett et al. and put them near the doves.

It's optional whether to lump the grebes and flamingos into one order, or treat them as two. I've switched to the conventional treatment with two orders. The large morphological differences and ancient split from the flamingos (e.g., Brown et al., 2008) support this.

PHOENICOPTERIFORMES Fürbringer, 1888

Phoenicopteridae: Flamingos Bonaparte, 1831

3 genera, 6 species HBW-1

PODICIPEDIFORMES Fürbringer, 1888

Podicipedidae: Grebes Bonaparte, 1831

7 genera, 23 species HBW-1

There's no molecular phylogeny of the grebes. However, Fjeldså (2004) provides a morphological phylogeny. Christidis and Boles (2008) have also adopted Fjeldså's phylogeny. For the present, I also use it.

Bochenski's (1994) osteological study of the grebes found that the Great Grebe is significantly different from the other Podiceps grebes. He created the genus Podicephorus for it. Fjeldså endorsed this view by putting Podicephorus in an unresolved trichotomy with Aechmophorus and the remaining Podiceps. The SACC has yet to consider the issue.

Tricolored Grebe, Tachybaptus tricolor, has been split from Little Grebe, Tachybaptus ruficollis. The subspecies vulcanorum is included in Tricolored Grebe. See Mlíkovský (2010).

Podicipedidae tree

Other Columbimorphae

PHAETHONTIFORMES Sharpe, 1891

The Tropicbirds are usually considered close to a variety of seabirds, but in an uncertain location (different studies give different results). Recently, they have become more controversial. Fain and Houde (2004), Ericson et al. (2006a), and Hackett et al. (2008) placed them in Metaves while a recent paper by Morgan-Richards et al. (2008) found them somewhat related to the hawks.

Phaethontidae: Tropicbirds Brandt, 1840

1 genus, 3 species HBW-1

PTEROCLIFORMES Huxley, 1868

Huxley (1868) first separated the sandgrouse as an order, using the name Pteroclomorphae, which modernizes to Pterocliformes.

I follow Hackett et al. (2008) where the sandgrouse, mesites, doves and pigeons form a clade. In contrast, Ericson et al. (2006a) placed the sandgrouse next to the doves and pigeons, but the mesites were close to flamingos and grebes. Gibb and Penny (2010), using only cytochrome-b, found the sandgrouse in a clade containing the falcons as well as the doves and pigeons. Analyses that do not use β-fibrogen, such as Gibb and Penny normally do not recover Metaves. Nonetheless, it calls into question whether the sandgrouse are sister to the doves and pigeons (or doves, pigeons, and mesites). The uncertainty about whether this grouping is actually a clade and the depth of the divisions between it members justify placing each of them in their own orders.

Pteroclidae: Sandgrouse Bonaparte, 1831

2 genera, 16 species HBW-4

There seems to be some controversy about how to spell the family name. Both Pteroclidae (Clements, HBW, Sibley-Monroe) and Pteroclididae (AOU, BLI, Howard-Moore, IOC) are in use, and Pterocleidae has also been used. The name indicates it is known for its wing, i.e., “-cles” takes the same meaning as in names such as Heracles. By analogy with Heraclidae/Heracleidae, it would then appear that either Pteroclidae or Pterocleidae would be correct. The first is the form used by Bonaparte when he established the family-group name in 1831 (as the subfamily Pteroclinae), and is used here.

MESITORNITHIFORMES Wetmore 1960

Sharpe had earlier tried to establish such a suborder (Mesitides), but based it on the preoccupied genus name Mesites. I haven't been able to find uses of Mesitornis as a type genus earlier than Wetmore.

Mesitornithidae: Mesites Wetmore, 1960 (1850)

2 genera, 3 species HBW-3

COLUMBIFORMES Latham, 1790

Columbidae: Doves, Pigeons Leach, 1820

46 genera, 332 species HBW-4

Click for genus-level Columbidae tree
Click for genus-level
Columbidae tree

I have based the organization of the Columbidae on the recent paper by Pereira et al. (2007), which provides a comprehensive DNA-based phylogentic tree. Although the dodos and Rodriguez Solitaire (genera Raphus and Pezophaps) have been traditionally considered a separate family in the Columbiformes, the DNA says otherwise. Shapiro et al. (2002) and Pereira et al. (2007) found that these two genera are buried deeply within the Columbidae, in the Raphinae. The Reunion Solitaire seems to have actually been an ibis! See Mourer-Chauviré et al. (1995).

The DNA testing shows 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. I divide it into two subfamilies: Phabinae and Raphinae. The dodos and solitaires are nested well within this third clade, in Raphinae.

The subfamily Columbinae has been studied in more detail by Johnson and Clayton (2000), Johnson et al. (2001), Gonzalez et al. (2009a), and Johnson and Weckstein (2011). Zenaidini and the species in Streptopelia and Columba, but not Patagioenas, have been rearranged accordingly. Johnson et al. (2001) also found that Nesoenas should be merged into Streptopelia, which I've done.

The work by Johnson and Weckstein (2011) showed that Geotrygon as usually constituted is paraphyletic with its members belonging to 3 separate clades. The Olive-backed Quail-Dove is sister to Leptotila. Johnson and Weckstein suggest that the relationship is fairly close, so I've moved it to Leptotila. However, if the relationship is as distant as implied by Pereira et al. (2007), it would be better to place it in a separate genus instead. The remaining quail-doves fall into at least two groups. Unfortunately, it is unclear where the type species (versicolor) of Geotyrgon goes. It is thought to be most closely related to caniceps and leucometopia, neither of which were included in the DNA analysis. That whole lot is thought to be close to montana, and I follow that presumption here. In that case, a different name is needed for the other Geotrygon clade, which is sister to Zenaida. So far as I can tell, none are available. For the present, I designate this group as "Geogrygon".

Jønsson et al. (2011a) studied the Gallicolumba ground-doves and bleeding-hearts. They found that Gallicolumba was paraphyletic, and recommended splitting it into Gallicolumba (bleeding-hearts) and Alopecoenas (Australasian ground-doves). However, their phylogenetic trees indicate there is still paraphyly even with the restricted Gallicolumba. That is handled here by further separating one of the clades as Pampusanna (Jacquinot & Pucheran 1853, type criniger; not Bonaparte's similarly named Pampusana).

Gibb and Penny (2010) investigated the fruit-doves and close relatives. They confirmed Shapiro et al.'s (2002) result that Alectroenas and Drepanoptila are embedded in fruit-dove genus Ptilinopus. Moreover, they tested most of the major subgenera and groups, allowing for a new arrangement of the enlarged Ptilinopus. An alternative way of handling the situation would be to retain Alectroenas and Drepanoptila and break Ptilinopus into 6 genera. The alternative requires changing many more scientific names, and I do not see sufficient reason to do that at this time.

The Black-banded Fruit-Dove, Ptilinopus alligator, has been split from Banded Fruit-Dove, Ptilinopus cinctus. Given present knowledge, it's fairly arbitrary whether to split or lump these allopatric forms. However, one is in Australia, the other in the Lesser Sundas, and the current tendency is for such forms to be split.

Based on Rheindt et al. (2011a), the Maroon-chinned Fruit-Dove, Ptilinopus subgularis is split into 3 species: Oberholser's Fruit-Dove, Ptilinopus epius, Banggai Fruit-Dove, Ptilinopus subgularis, and Sula Fruit-Dove, Ptilinopus mangoliensis.

Columbinae Leach, 1820

Zenaidini Bonaparte, 1853

Columbini Leach, 1820

Claravinae: American Ground-Doves Richmond, 1917 (1850)

Phabinae: Australasian Pigeons and Doves Bonaparte, 1853

Raphinae: Old World Doves and Pigeons Wetmore, 1930 (1835)

Raphini Wetmore, 1930 (1835)

Treronini: Green-Pigeons G.R. Gray, 1840

Turturini G.R. Gray, 1840

Ptilinopini: Fruit-Doves and Imperial-Pigeons Selby, 1835

EURYPYGIMORPHAE

Although the tree suggests that the Eurypgiformes belong with the Cypselomorphae, support for this seems to be fairly weak. For that reason I have put them in a separate superorder.

EURYPYGIFORMES Fürbringer, 1888

These two monotypic families form a strongly supported clade in Hackett et al. (2008). Their affinities have long been unclear. They have recently been grouped near the cranes, but that appears incorrect and they are likely a relatively basal group. The best estimate from Hackett et al. has them sister to the Cypselomorphae.

Rhynochetidae: Kagu Carus, 1868

1 genus, 1 species HBW-3

Eurypygidae: Sunbittern Selby, 1840

1 genus, 1 species HBW-3

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