Anomalogonates I

The 46 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

‘ANOMALOGONATAE’

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 Sibley-Monroe 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, Leptosomidae. It seems to be a relatively basal family in the ‘Anomalogonatae’ (which also puts it in its own order), although even Hackett et al. (2008) were unable to confidently determine where it fits. Mayr (2008) discusses some of the differences between the Cuckoo Roller and the rollers. See also Darren Naish's interesting post on the Cuckoo Roller at Tetrapod Zoology.

The owls, mousebirds, trogons are also placed in separate orders, as their affinities with the other ‘Anomalogonatae’ are relatively distant. The hornbills, which are split into Bucorvidae (ground-hornbills) and Bucerotidae (hornbills) form a grouping with the hoopoes and woodhoopoes. The bee-eaters, rollers, ground-rollers, todies, motmots, and kingfishers form the Coraciiformes. The last group is the the Piciformes, which I take to include the Galbuliformes. They will be considered on the next page.

COLIIFORMES Murie, 1872

Coliidae: Mousebirds Sundevall, 1836

2 genera, 6 species HBW-6

STRIGIFORMES Wagler, 1830

Tytonidae: Barn Owls Mathews, 1912 (1866)

2 genera, 18 species HBW-5

The Eastern Barn-Owl has been split as a separate species as in Christidis and Boles (2008). This had previously been called Tyto javanica based on Wink et al. (2004b). However, there seems to have been a problem with the DNA sample, and javanica is most likely a subspecies of Common Barn-Owl, Tyto alba. Thus the Eastern Barn-Owl becomes Tyto delicatula. Some other Barn Owl splits may be needed. In particular, the American subspecies could be split at Tyto furcata.

Following H&M 4 (Dickinson and Remsen, 2013) and König and Weick (2008), the poorly known Congo Bay-Owl has been moved to Tyto from Phodilus.

Strigidae: Typical Owls Leach, 1820

30 genera, 214 species HBW-5

The overall organization is based on Wink et al. (2008) and Fuchs et al. (2008). The typical owls are divided into three subfamilies: Ninoxinae, Surniinae, and Striginae. These are further divided into tribes, although that is almost overkill as the tribes are barely larger than the main genus, if at all.

Within Ninoxinae, the position of Uroglaux and Sceloglaux is unresolved, although they are believed to be near Ninox on morphological grounds.

Strigidae tree There are four main clades in Surniinae. There's no molecular evidence concerning Xenoglaux, but it is believed close to Micrathene, which seems to be basal in Surniinae. Heteroglaux is most likely sister to Athene, which it has sometimes been considered part of.

The treatment of the Glaucidium complex is a bit novel. The basic structure comes from Wink et. al. (2008). Although their taxon sampling is a bit sparse, it suggest five clades in the Glaucidium complex. I've applied the old name Smithiglaux (Bonaparte 1854) to the basal clade, which consists of two African species. The next clade gets the name Taenioglaux, which has recently been revived (e.g., König and Weick, 2008) for Taenioglaux plus Smithiglaux. Next comes the monotypic Surnia, then Glaucidium itself. Glaucidium consists of an Old World clade of owlets (Glaucidium proper) and a New World clade of pygmy-owls. The latter could take the name Phalaenopsis (Bonaparte 1854). For now, I am treating it as a subgenus to maintain consistency with AOU, although a separate genus would make the phylogeny more transparent.

The rest of the owls are in the subfamily Striginae. Otus has been split into the Old World Otini scops-owls and New World Megascopini based on genetic data (e.g., Wink and Heidrich, 1999; Wink et al., 2004b, 2008; Fuchs et al. 2008). Surprisingly, they are not sister groups. Otini consists of the Old World Otus and Pyrroglaux. It is not clear that the latter belongs here, but it seems more likely than the alternatives. The Giant Scops-Owl, Mimizuku gurneyi, is considered part of Otus (Miranda et al., 1997, 2011). I've rearranged Otus some based on Fuchs et al. (2008), Pons et al. (2013), Wink et al. (2008) and HBW-5 (del Hoyo et al., 1999), but a true species-level phylogeny is not available.

The three extinct owls from the Mascarenes are in the genus Mascarenotus, which is thought to be closely related to Otus, or may even be embedded in it.

The Megascopini come after the Asionini. Margarobyas is probably close to Megascops. Note that the genus name Gymnoglaux has been replaced by Margarobyas. This is because Gymnoglaux is actually a synonym of Megascops (see Olson and Suárez, 2008).

There has been resistance to separating the Flammulated Owl from Otus with Megascops, probably because it is too different from the screech-owls. Nonetheless, the genetic data group it with Megascops, albeit distantly. For that reason it gets its own genus Psiloscops (Coues 1899) as in König and Weick. I place it basally in Megascopini.

After Pulsatricini, comes Strigini. Jubula is conventionally place here, although its exact affinities remain unclear. My treatment of Strix is a bit unusual. Wink et al. (2008) found two clades in Strix. One is a Holarctic/Old World clade, the other consists of New World species. Some of the New World species have sometimes been separated as Ciccaba, but it has often been objected that they are not distinct from other New World Strix. Here I put all of the New World Strix in an expanded Ciccaba. That leaves the holarctic Great Gray Owl as the only Strix present in the New World. Finally, the African Wood-Owl, Strix woodfordii, which has sometimes been considered close to Ciccaba in the narrow sense, belongs in the Old World clade.

The last clade is Bubonini. DNA is showing that the old generic limits don't work, and the tendency has been to merge everything into Bubo. I think this hides the phylogeny too much. An improvement would be to use two genera, Bubo as constituted here and Ketupa. However, given that some of these species (and even former genera) are mophologically distinct, I think a better solution is to break the clade sister to Bubo into three genera.

We then have 4 genera in Bubonini: Bubo, Nyctaetus (Le Maout 1853), Scotopelia and Ketupa. Ketupa has been expanded with some species that may be closely related (more data is needed). Scotopelia retains its traditional limits. Shelley's and Verreaux's Eagle-Owls are in their own clade, Nyctaetus. It's unclear which clade the Dusky and Akun Eagle-Owls belong in, so I've left them in Bubo for now. Although the Snowy Owl has been considered to be in its own genus, Nyctea, it is actually in Bubo proper, sister to the Great Horned Owl.

The Bermuda Saw-whet Owl, Aegolius gradyi, has been added as it seems to have become extinct in historical times (Olson, 2012).

Northern Boobook, Ninox japonica, and Chocolate Boobook, Ninox randi, have been split from Brown Hawk-Owl, Ninox scutulata, following King (2002).

Rasmussen et al. (2012) re-examined the Philippine Hawk-Owl complex. Their fieldwork turned up two new species in the complex. They argue that the complex consists of 7 species, and I follow their recommendations. This means that Philippine Hawk-Owl is replaced by:

Guatemalan Pygmy-Owl, Glaucidium cobanense has been split from Northern Pygmy-Owl, Glaucidium gnoma, due to substantial differences in vocalizations (Eisermann and Howell, 2011).

Socotra Scops-Owl, Otus socotranus, has been separated from the African Scops-Owl, Otus senegalensis, due to substantial differences in vocalizations (e.g., König and Weick, 2008). In fact, its calls are more similar to the Oriental Scops-Owl, Otus sunia. Pons et al. (2013) found it was closely related to O. sunia and O. insularis. The Arabian Scops-Owl, Otus pamelae, has also been separated from the African Scops-Owl, Otus senegalensis (Pons et al., 2013). The Negros Scops-Owl, Otus nigrorum, and Everett's Scops-Owl, Otus everetti have been split from Philippine Scops-Owl, Otus megalotis. See Miranda et al. (2011). Finally, the newly discovered Rinjani Scops-Owl, Otus jolandae, has been added to the list (Sangster et al., 2013).

The Mexican Barred Owl, Ciccaba sartorii has been split from Northern Barred Owl, Ciccaba varia due to difference in both DNA and vocalizations. It is more closely related to the Fulvous Owl than to the Northern Barred Owl (Barrowclough et al., 2011).

Ninoxinae: Hawk-owls Marks et al., 1999

Surniinae: Owlets and Pygmy-Owls Bonaparte, 1838

Subgenus Phalaenopsis

Striginae Leach, 1820

Otini: Scops-Owls Bonaparte, 1854

Asionini: Eared Owls Vigors, 1825

Megascopini: Screech-Owls Wink et al., 2008? Informal?

Both Megascopini and Pulsatrigini have been used by Wink et al. (2008), but do not seem to have been designated as new taxa, which is now required by the ICZN (since 1999). The spelling of Pulsatrigini has been corrected to Pulsatricini.

Pulsatricini Wink et al., 2008? Informal?

Strigini: Wood-Owls Leach, 1820

Bubonini: Eagle-Owls Vigors, 1825

LEPTOSOMIFORMES Sharpe, 1891

Leptosomidae: Cuckoo Roller Blyth, 1838

1 genus, 1 species HBW-6

TROGONIFORMES AOU, 1886

This has been attributed to Fürbringer, 1888, but it was already used in 1886 in the 1st AOU checklist.

Trogonidae: Trogons Lesson, 1828

7 genera, 44 species HBW-6

Standard Tree
Click for Trogonidae tree
Click for Trogonidae tree

One unusual feature of trogons is their heterodactyl foot. In some cases this allows fossils to be easily identified as trogoniformes. Early trogoniform fossils have been found from the London Clay and Danish Fur Formations, both from the early Eocene (Mayr, 2009). The geographic origin of the trogons is unknown. It was formerly thought to be Africa, but as Apaloderma is not a basal clade, there is no real supporting evidence. Its nearest relative seems to be cuckoo-roller family, Leptosomidae. Since the cuckoo-rollers were widespread in the Northern Hemisphere during the Eocene (Mayr, 2009), this does not give us any useful information as to origin. The crown group dates to about 30 million years ago. Depending on the trogon tree, southeast Asia and the Americas are the most likely places the crown group originated.

The uncertainty is because the internal arrangement of Trogonidae is currently quite uncertain. Papers by Espinosa de los Monteros (1998, 2000), Johansson and Ericson (2005), Moyle (2005), Ornelas et al. (2009), and Hosner et al. (2010) have put forth a variety of arrangements. For the present, it is reasonably certain that the quetzal genera Euptilotis and Pharomachrus are sisters. Surprisingly, the African Apaloderma trogons appear to be sister to the Apalharpactes trogons of Java and Sumatra (Hosner et al., 2010). Although support for this is somewhat weak, it is also supported by a 3-bp insertion in the G3PDH, as well as the green plumage they both share (something absent in Harpactes). Beyond that, the studies lack consistency.

There are two main problems: establishing the root of the tree and the position of the Caribbean trogons, Priotelus. The weakly supported tree in Hosner et al. (2010) has additional differences.

The order here is based on Ornelas et al. (2009), which uses the most complete set of genes. The data is a superset of that in Espinosa de los Monteros (1998, 2000), Johansson and Ericson (2005), and Moyle (2005). The Apalharpactes-Apaloderma clade is basal, followed by Harpactes, and then the New World trogons. For the species-level tree, I also consulted DaCosta and Klicka (2008) for Trogon, and Hosner et al. (2010) for Harpactes (including Duvaucelius).

Alternate Tree
Alternate Trogonidae tree
Alternate Trogonidae
tree (after Moyle, 2005)

I also present an alternate arrangement based on Moyle (2005). This arrangement is also biogeographically simpler than the others, although I'm not sure that is a virtue.

In the alternate arrangement, the New World trogons, Euptilotis and Pharomachrus are the basal clade. The next branch is Priotelus, the Caribbean trogons. The remainder consists of the Old World clade and Trogon itself. Hosner et al. (2010) found that Duvaucelius is embedded in Harpactes, so all of them are placed in Harpactes. This group of Asian trogons may be sister to the Apalharpactes-Apaloderma. Finally, the genus Trogon breaks into two clades: rufus through aurantiiventris, and clathratus through surrucura. The generic name Trogonurus could be applied to the first clade.

The AOU's South American checklist committee had considered making some changes in Trogon species in 2003, partly due to the treatment in Ridgely and Greenfield (2001) and Hilty (2003). For more information, read the discussion for viridis, violaceus, and melanurus on the SACC site. They found that the relevant data, if it existed, was not collected together in a way that allowed them to judge it properly. The publication of DaCosta and Klicka (2008) has changed the balance of evidence here. Although the SACC had not then acted on it, I changed generic limits accordingly.

The species and species groups affected are White-tailed Trogon (T. viridis), Violaceous Trogon (T. violaceus), Black-tailed Trogon (T. malanurus), and Collared/Orange-bellied Trogon (T. collaris and aurantiiventris). The table below summarizes the splits, including genera. The species affected are maked with an asterisk. The question marks on T. macroura (sometimes called Large-tailed Trogon) and T. melanopterus reflect the possibility of future splits.

The SACC has passed (Apr. 8, 2009) a set of proposals concerning splitting mesurus from melanurus, chionurus from viridis, and caligatus from violaceus. I've updated the table with the SACC English names. The NACC has also tentatively ratifed the relevant portions of these changes. The SACC has now recognized the Amazonian Trogon, Trogon ramonianus, as a separate species (Oct 6, 2010). They also adopted the English name Guianan Trogon for the now monotypic T. violacenous. Note that there are still some unresolved issues concerning the exact distribution of these species.

Species Subspecies
Collared/Orange-bellied Trogon
*Collared Trogon, T. collaris extimus, hoethinus, virginalis,
subtropicalis, exoptatus,
collaris, casganeus
*Orange-bellied Trogon, T. puella puella, underwoodi,
aurantiiventris, flavidor

Black-tailed Trogon Complex
Lattice-tailed Trogon, T. clathratus clathratus
*Ecuadorian Trogon, T. mesurus mesurus
*Black-tailed Trogon, T. melanurus macroura?, eumorphus,
occidentalis, melanurus
Blue-tailed Trogon, T. comptus comptus

White-tailed Trogon Complex
Black-headed Trogon, T. melanocephalus melanocephalus
Citreoline Trogon, T. citreolus citreolus, sumichrasti
*Green-backed Trogon, T. viridis viridis, melanopterus?
*White-tailed Trogon, T. chionurus chionurus
Baird's Trogon, T. bairdii bairdii

Violaceous Trogon Complex
*Gartered Trogon, T. caligatus (sallaei aka braccatus), concinnus,
caligatus
*Amazonian Trogon, T. ramonianus ramonianus, crissalis
Surucua Trogon, T. surrucura aurantius, surrucura
*Guianan Trogon, T. violaceus violaceus
Blue-crowned Trogon, T. curucui peruvianus (bolivianus),
curucui, behni

All of these forms are apparently field-identifiable both by plumage and voice (see Hilty, 2003; Howell and Webb, 1995; Ridgely and Gwynne, 1989; Ridgely and Greenfield, 2001; Stiles and Skutch, 1989), although some of the differences have not been clearly documented.

These changes combine the Middle American Collared Trogons with Orange-bellied Trogon [I'd previously tried using Jalapa Trogon (used by Sibley and Monroe; 1990, 1993), but have decided to revert to Orange-bellied]. Note that the scientific name does not match the AOU because they don't include puella in the Orange-bellied Trogon. I do, and it has priority. Thus the range of Orange-bellied extends from Mexico to western Panama. The Collared Trogons of South America and Eastern Panama retain the name Collared Trogon. The Black-tailed Trogons west of the Andes in Ecuador and Peru become Ecuadorian Trogon. The White-tailed Trogon can be found from Panama to W Ecuador. The rest of the White-tailed Trogons are grouped as the Green-backed Trogon, even in Trinidad (the population in southeast Brazil may be split at some point). Gartered Trogon is present from Mexico into S. America west of the Andes, and in the north into western Venezuela. The Amazonian Trogon is in the Amazon Basin. Finally, the Guianan Trogon is present in eastern Venezuela, the Guianas, Trinidad, and northeastern Brazil.

BUCEROTIFORMES Fürbringer, 1888

Upupidae: Hoopoes Leach, 1820

1 genus, 4 species HBW-6

Phoeniculidae: Woodhoopoes Bonaparte, 1831

2 genera, 9 species HBW-6

Bucorvidae: Ground-Hornbills Bonaparte, 1854

1 genus, 2 species Not HBW Family

Bucerotidae: Hornbills Rafinesque, 1815

16 genera, 59 species HBW-6

Click for Bucerotidae tree
Click for Bucerotidae tree

Hornbill phylogeny follows Gonzalez et al. (2013a). They analyzed all hornbill species using both nuclear and mitochondrial DNA. At the genus level, their results match Viseshakul et al. (2011), which used only the cytochrome-b gene. Viseshakul et al. found that much of the hornbill diversity is truly ancient. The genera all seem to have existed as separate lineages for over 30 million years. This caused some problems for them fully resolving the hornbill tree. Gonzalez et al. found a somewhat different arrangement with better support. It is possible that consideration of additional genes would lead to some further adjustment. In particular, I have doubts that Berenicornis (White-crowned Hornbill) is correctly placed yet.

I have followed Hübner et al. (2003) and Gonzalez et al. (2013a) in splitting Tockus into the whistlers (Lophoceros) and cluckers (Tockus). Besides the vocal distinctions, the split bewteen these clades is quite deep. Viseshakul et al. (2011) et al. estimate it at around 45 million years. Note the genus name Lophoceros (Ehrenberg 1833) has priority over Rhynchaceros (Gloger 1842) as suggested by both Hübner et al. (2003) and Gonzalez et al. (2013a). (They subsequently corrected this in Gonzalez et al. 2013b.)

There were two surprises at the species level in Gonzalez et al. (2013). The Black Dwarf-Hornbill, Tockus hartlaubi, does not belong in Tockus. Here it has been placed sister to Tropicranus in the monotypic genus Horizocerus (Oberholser 1899). The Sulawesi Hornbill, Penelopides exarhatus, doesn't belong with Penelopides. Instead, it joins Wrinkled, Walden's, and Writhed Hornbills. Viseshakul et al. (2011) had previously shown they did not belong in Aceros, and I had separated them as Cranobrontes (Riley 1921, type species leucocephalus). But now, since exarhatus has joined them, they must take the name Rhabdotorrhinus (Meyer and Wiglesworth 1898, type exarhatus), which has priority over Cranobrontes.

The genus Aceros has been reduced to a single species. Besides losing 3 species to Rhabdotorrhinus, the other former Aceros hornbill, the Knobbed Hornbill, has been merged into Rhyticeros. It seems to be rather distant from the other Rhyticeros hornbill and could reasonably be put in its own genus, Cranorrhinus (Cabanis and Heine, 1860).

The Red-billed Hornbills are split based on Kemp and Delport (2002) and Delport et al. (2004). Some of the red-billed hornbills were included in Viseshakul et al. (2011), which also supports this treatment.

CORACIIFORMES Forbes, 1884

Click for Coraciiformes tree
Click for Coraciiformes tree

The genus-level tree of the Coraciiformes is based on Ericson et al. (2006), Hackett et al. (2008), Kirchman et al. (2001), Marks et al. (2007), and Moyle (2006).

Meropidae: Bee-eaters Rafinesque, 1815

3 genera, 27 species HBW-6

The bee-eaters follow Marks et al. (2007).

Brachypteraciidae: Ground-Rollers Bonaparte, 1854

4 genera, 5 species HBW-6

Kirchman et al. (2001) discuss the split of Geobiastes from Brachypteracias. However, they do not come to a definitive conclusion regarding how the Ground-Rollers are related.

Coraciidae: Rollers Rafinesque, 1815

2 genera, 12 species HBW-6

Todidae: Todies Vigors, 1825

1 genus, 5 species HBW-6

Todidae tree

The tody sequence is based on Overton and Rhoads (2004). They also considered the todies and motmots sister families, as did Johansson and Ericson (2003). I follow Ericson et al. (2006a) and Hackett et al. (2008), which consider motmots and kingfishers sister families.

Momotidae: Motmots G.R. Gray, 1840 (1832-33)

6 genera, 15 species HBW-6

The motmot sequence is based on Witt (2004).

The Blue-crowned Motmot complex has been split based on Witt (2004) and Stiles (2009). Witt found that the Andean (Highland) Motmot, Momotus aequatorialis (including chlorolaemus), is not truly part of the Blue-crowned complex, but is sister to the combined Blue-crowned complex plus the Russet-crowned Motmot, Momotus mexicanus.

The Blue-crowned complex proper is split into 6 species based on a combination of Witt (2004) and Stiles (2009). The races are allocated as follows, with approximate species ranges.

Name Associated Subspecies Range

Blue-crowned Motmot, Momotus coeruliceps coeruliceps NE Mexico: Nuevo Leon & Tamaulipas
Blue-diademed Motmot, Momotus lessonii goldmani, exiguus, lessonii Middle America: Veracruz to W Panama (Chiriqui)
Amazonian Motmot, Momotus momota microstephanus, momota, ignobilis, simplex, cametensis, paraensis, marcgravianus, nattereri, pilcomajensis Amazon Basin
Silver-banded Motmot, Momotus argenticinctus argenticinctus W Ecuador and NW Peru, W of Andes
Trinidad Motmot, Momotus bahamensis bahamensis Trinidad & Tobago
Whooping Motmot, Momotus subrufescens subrufescens, spatha, osgoodi, conexus*, reconditus*, olivaresi* C & E Panama and NW South America (Chocó, Caribbean slope, Magdalena Valley)

* = There is a case for submerging conexus, reconditus, olivaresi into subrufescens.
Momotidae tree

All but the Silver-banded Motmot are recognized by Stiles. However, Witt found it to be sister to the Trinidad/Whooping group, which is why I'm treating it as a separate species.

Alcedinidae: Kingfishers Rafinesque, 1815

19 genera, 107 species HBW-6

I've followed the taxonomic recommendations of Moyle et al. (2007) for the river kingfishers (Alcedininae). Two subspecies of the Malachite Kingfisher, Corythornis cristata, are sometimes considered full species. They are the Sao Tome Kingfisher (C. cristata thomensis) and the Principe Kingfisher (C. cristata nais). Recent work by Melo and Fuchs (2008) suggests they should continue to be considered subspecies.

The Oriental Dwarf-Kingfisher is here considered to be two species: Black-backed Kingfisher, Ceyx erithaca, and Rufous-backed Kingfisher, Ceyx rufidorsa (including motleyi). See Lim et al. (2010a) for more.

Based on Andersen et al. (2013), the former Variable Dwarf-Kingfisher has been split into 15 allopatric species. Most (all?) of these are field identifiable and the genetic distance between most of them is substantial. They are the species starting at Dimorphic Dwarf-Kingfisher, and continuing to the end of Alcedininae.

Alcedininae: River Kingfishers Rafinesque, 1815

Cerylinae: Water Kingfishers Reichenbach, 1851

Halcyoninae: Tree Kingfishers Vigors, 1825

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