Galliformes

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

GALLIFORMES Temminck, 1820

The Galliformes are sister to the Anseriformes. Together, they are sister to Neoaves.

Galliformes
Click for genus-level tree
for Galliformes

The arrangement of Galliform families follows the recent genetic analyses of Crowe et al. (2006a), Cox et al. (2007) and Hackett et al. (2008). It is also consistent with Kaiser et al. (2007) and Kriegs et al. (2007), who analyze retropositions. Of the two, Kriegs et al. is more complete, and exhibits the same basic structure as the tree here. Since it is a different method of reading the genetic data, it represents significant additional support for the current arrangement of Galliform families. Nonetheless, there is still some question about whether the Numidade branch off before or after the Odontophoridae (e.g., Shen et al., 2010).

The Galliform tree takes the form of a cascade, with one group breaking off at a time until you get to the Phasianidae. It starts with the megapodes, then the Cracidae (guans, chachalacas, and curassows), guineafowl, and finally new world quail, all before we get to the Phasianidae. A number of genera have been rearranged or split compared with previous treatments. The genera decorated with question marks are uncertainly placed and could easily end up in a different subfamily.

Megapodiidae: Megapodes Lesson, 1831

7 genera, 22 species HBW-2

The arrangement of the megapodes is based on the 2-gene analysis of Birks and Edwards (2002). There is some uncertainty here. Kimball et al. (2011), apparently using the same data, swaps the position of Talegalla and Leipoa (also seen in the some of the single-gene Birks and Edwards analyses), and places Macrocephalon as sister to the megapodes instead of being sister to the mound-building group (Talegalla thruough Aepypodius).

Cracidae: Chachalacas, Curassows, Guans Rafinesque, 1815

9 genera, 54 species HBW-2

There has been a debate concerning whether the chachalacas and Horned Guan are more closely related to the guans or currasows. Pereira et al. (2002, 2009) found that the Horned Guan is closer to the currasows, and that the chachalacas are sister to the Horned Guan-Currasow clade. This topology has generally been supported by other studies, e.g., Kimball et al. (2011), although Frank-Hoeflich et al. (2006) found the opposite branching order using combined molecular, osteological, morphological, and behaviorial data (but see the response by Pereira et al., 2009).

Pereira et al. (2002) estimated divergence times for various genus-level nodes. They found several divisions dating back to the Oligocene. I've used subfamilies and tribes to highlight these divisions. Except for the Horned Guan, they correspond to the traditional division into chachalacas, curassows, and guans.

The genus Pipile has been submerged into Aburria (Grau et al., 2005). No comprehensive study of the guans is available. The order follows SACC. The chachalaca ordering is based on the analyses of Frank-Hoeflich et al. (2006). Interestingly, they found the Ortalis canicollis race pantanalensis on a separate branch from O. canicollis, whether this is an artifact or real is unclear. The order within the currasows is based on Pereira and Baker (2004), which also requires merging Mitu into Pauxi.

Based on Gastañaga et al. (2011), Sira Curassow, Pauxi koepckeae, from the Sira Mountains in Peru, has been split from Horned Curassow, Pauxi unicornis.

The SACC has split East Brazilian Chachalaca, Ortalis araucuan, and Scaled Chachalaca, Ortalis squamata, from Speckled Chachalaca, Ortalis guttata.

Penelopinae: Guans Bonaparte, 1831

Cracinae: Chachalacas and Curassows Rafinesque, 1815

Ortalidaini: Chachalacas Donegan, 2012

Oreophaseini: Horned Guan Bonaparte, 1853

Cracini: Curassows Rafinesque, 1815

Numididae: Guineafowl de Selys Longchamps, 1842

4 genera, 6 species HBW-2

The order of genera follows Crowe et al. (2006a, b). This coincides with the cytochrome-b tree in Crowe et al. (2006a). Kimball et al. (2011) give a slighty different arrangement that swaps Numida and Guttera.

Odontophoridae: New World Quail Gould, 1844

10 genera, 34 species HBW-2

Surprisingly, two Old World species thought to be unrelated members of Phasianidae have turned out to be sister taxa and basal members of the Odontophoridae. They are the Stone Partridge (Ptilopachus petrosus) and Nahan's Francolin (now Nahan's Partridge, Ptilopachus nahani). See Crowe et al. (2006a) and Cohen et al. (2012).

The New World Quail are arranged based on the tree in Eo et al. (2009). Only half of that tree seems to be based on DNA. If you examine the tree diagram, you'll find that half of the genera have question marks on them, indicating that I have no DNA evidence concerning them.

Phasianidae: Turkeys, Grouse, Pheasants, Partridges Horsfield, 1821

53 genera, 182 species HBW-2

The divisions within the Phasianidae remain a problem. The papers by Bao et al. (2010), Bonilla et al. (2010), Crowe et al. (2006a, b), Kan et al. (2010), Kimball and Braun (2008), Kimball et al. (2011), Kriegs et al. (2007), Liu et al. (2012) Meng et al. (2008), and Shen et al. (2010) give different answers about how the various components of Phasianidae are related. This is especially true if you consider the individual gene trees contained in Shen et al.'s supplementary material. And what is one to make of Liu et al. (2012)?

There does seem to be a broad, but not complete consensus that the Asiatic partridges (Rollulinae) are basal (see Liu et al. (2012) for a contrary view), and that some group containing Gallus is sister to a group containing the Phasianinae, but the placement of various pheasants (Pavoninae, and Polyplectroninae) and even the placement of the Tetraogallinae is problematic (ditto its internal organization). Sometimes these tribes group near the Gallinae in various configurations, sometimes they are not in the Gallinae-Phasianinae clade. Ultimately, I decided to treat all of the troublesome groups as separate subfamilies, but leave the branching order completely unresolved for the present. They are listed in order of size.

How to piece together the Phasianinae subfamily had been a little tricky, but a consistent picture has now emerged. The current order is based on Kimball et al. (2011), which includes most of the species in the subfamily. Other than including more taxa, it is quite similar to the results of Kimball and Braun (2008). Phasianinae is their “erectile clade”. I have divided it into tribes to better show the relationships of the various groups. As you can see by their membership, these tribes generally correspond to what we think of as natural groupings. Bao et al. (2010) is generally consistent with the present arrangement of the Phasianinae. Crowe et al. (2006a) suggested that the turkeys and Perdix partridges might be sister genera, but that has gotten little support from recent analyses.

Not surprisingly, part of the reconstruction of the Phasianidae involved moving some of the species to new genera. Crowe et al. (2006a) found two species that were quite wrongly placed: the Stone Partridge (Ptilopachus petrosus) and Nahan's Francolin (now Nahan's Partridge, Ptilopachus nahani), both of which ended up in the new world quail (Odontophoridae). When Crowe et al. (1992, with a different set of co-authors) had reorganized the francolins, they noted that Nahan's Francolin didn't appear to be a francolin. That reorganization is mostly supported by the new paper, but one other francolin proved problematic. The Crested Francolin, which they had already reassigned to the genus Peliperdix, is now Dendroperdix sephaena, although it remains in the junglefowl/francolin subfamily, Gallinae.

Eo et al. (2009) draws attention to several genera that are currently regarded as part of Phasianidae (Haematortyx, Melanoperdix, Rhizothera, Galloperdix). While it suggests closer scrutiny of these genera is in order, I don't buy the arrangement of them in the paper. I think it's an artifact of the supertree method. In fact, there is a little genetic data on one of them, Haematortyx. It was included in Crowe et al. (2006b) who put it sister to Polyplectron, a very different location than in Eo et al. Crowe et al. (2006a), does find that the Ptilopachus-Odontophoridae clade is sister to the Phasianidae, which is why I've moved Ptilopachus into Odontophoridae. They did not find Phasianidae nested within the Ptilopachus-Odontophoridae clade. I'm leaving the other genera highlighted by Eo et al. in Phasianidae for now, but we should keep in mind that some or all may too belong in Odontophoridae.

Rollulinae: Asiatic Partridges Bonaparte, 1850 (1848)

Bonaparte's name Rollulinae has priority over Arborophilinae, Crowe et al. (2006a). Note that the Rubeho Forest Partridge, Xenoperdix obscuratus has been split from Udzungwa Forest Partridge, Xenoperdix udzungwensis (Bowie and Fjeldså, 2005). See Wang et al. (2013) for the treatment of Caloperdix as sister to Rollulus.

Pavoninae: Peafowl and allies Rafinesque, 1815

Pavonini: Peafowl Rafinesque, 1815

Polyplectronini: Peacock-Pheasants Blyth, 1852

The arrangement of Polyplectron is based on Davison et al. (2012) and Kimball et al. (2011).

Gallini: Junglefowl, Francolins Brehm, 1831

Latham's Francolin, Peliperdix lathami, takes the basal position based on Kimball et al. (2011). The other 3 species sometimes placed in Peliperdix (coqui, albogularis, schlegelii) seem more closely related to the Scleroptila francolins, and are merged into that genus. Although the Crested Francolin, Francolinus sephaena, has been included in Peliperdix, indications are that it is in the francolin group (Crowe et al., 2006; Meng et al., 2008) perhaps closer to the Gray Francolin, Francolinus pondicerianus, than to the francolinus group (Kimball and Braun, 2011). The arrangement within the rest of Francolinus follows Forcina et al. (2012) I don't know for sure that Galloperdix belongs here, but it is often placed next to Bambusicola, as I have done here.

Tetraogallini: Old World Partridges and Quail, Spurfowl Bonaparte, 1854 (1846)

The Brown Quail is not closely related to the other Coturnix species (Seabrook-Davison et al., 2009; Kimball et al., 2011). Its closest relatives are Excalfactoria. However, it seems to be a fairly distant relative and I have placed it in Synoicus (Gould, 1843). Wang et al. (2013) found that Ammoperdix is a basal member of the Coturnix clade.

Phasianinae: Pheasants, Grouse, Turkeys Horsfield, 1821

The Vietnamese Pheasant, Lophura hatinhensis, is considered part of Edwards's Pheasant, Lophura edwardsi. It appears to be either a subspecies or color morph. In any event, it is not genetically distinct (Hennache et al., 2003). Also, the arrangement within Lophura is based on Randi et al., 2001.

Ithaginini: Blood Pheasant Wolters, 1976

Lophophorini: Monals and Tragopans G.R. Gray, 1841

Tetraonini: Grouse and Turkeys Leach, 1820

The Turkeys are now consiered embedded in Tetraonini. Interestingly enough, the Koklass Pheasant appears to be basal in this group (Kimball and Braun, 2014).

Phasianini: Pheasants, Perdix Partridges Horsfield, 1821

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