Afroaves

Accipitrimorphae

Strigiformes

Picimorphae

Coliiformes

Leptosomiformes

Trogoniformes

Bucerotiformes

Coraciiformes

Piciformes

The 46 Orders

Paleognathae

Galloanserae

Columbea

Otidae

Gruae

Ardeae

Telluraves

Afroaves

Australaves

TROGONIFORMES AOU, 1886

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

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).

Trogonidae: Trogons Lesson, 1828

7 genera, 44 species HBW-6

Standard Tree
Click for Trogonidae tree
Click for Trogonidae tree

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

The Bucerotiformes are another ancient order. According to Mayr (2009), the only documented early fossil Bucerotiformes seem more closely related to the hoopoes and woodhoopoes. The Messelirrisoridae are known from the London Clay (UK, early Eocene) and Messel (Germany, middle Eocene). He also mentions there may be a not yet described Bucerotiform from the Green River Formation (Wyoming, early Eocene).

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 family-level arrangement of the Coraciiformesis based on Ericson et al. (2006a) and Hackett et al. (2008). The Coraciiform families fall into three groups: the bee-eaters (Meropidae); the ground-rollers and rollers (Brachypteraciidae and Coraciidae); the kingfishers, motmots, and todies (Alceidiae, Momotidae, and Todidae). Ericson et al. left the relationships of the three groups unresolved, while Hackett et al. (2008) considered the bee-eaters the basal group, with the roller and kingfisher groups sister. The morphological analysis reported in Mayr (2009) had the bee-eaters grouped with the kingfisher group, and the roller group basal.

Early fossil Coraciiformes include the families Primobucconidae, Eocoraciidae, Geranopteridae, and the species Quasisyndactylus longibranchis. As with many other Afroavian orders, the earliest fossils are from the early Eocene in the Northern Hemisphere (Clarke et al., 2009; Ksepka and Clarke, 2010), in this case including both North America and Europe (Clarke et al., 2009). Although the todies are now restricted to the Caribbean, there are fossils from the Oligocene of Europe as well as North America (Mayr, 2009).

MEROPI Forbes, 1884

Meropidae: Bee-eaters Rafinesque, 1815

3 genera, 27 species HBW-6

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

CORACII Forbes, 1884

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

HALCYONI Forbes, 1884

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, 116 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.

The Silvery Kingfisher, Ceyx argentatus, has been split into Southern Silvery-Kingfisher, Ceyx argentatus, and Northern Silvery-Kingfisher, Ceyx flumenicola. See Collar (2011) and Andersen et al. (2013).

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.

Andersen et al. (2015a) examined many of the Todiramphus kingfishers. As a result, the Micronesian Kingfisher, Todiramphus cinnamominus, has been split into Rusty-capped Kingfisher, Todiramphus pelewensis, Guam Kingfisher, Todiramphus cinnamominus (including the extinct miyakoensis), and Pohnpei Kingfisher, Todiramphus reichenbachii.

Although data is lacking, the rearrangment increased the doubt about whether the two taxa united as Tuamotu Kingfisher, Todiramphus gambieri, are really conspecific. They have been split into Mangareva Kingfisher, Todiramphus gambieri, and Niau Kingfisher, Todiramphus gertrudae.

The Collared Kingfisher, Todiramphus chloris, has been split into Collared Kingfisher, Todiramphus chloris, Pacific Kingfisher, Todiramphus sacer, Melanesian Kingfisher, Todiramphus tristrami, Mariana Kingfisher, Todiramphus albicilla, Torresian Kingfisher, Todiramphus sordidus, and Islet Kingfisher, Todiramphus colonus. See the current IOC for the allocation of subspecies.

I've also resequenced Todiramphus based on Andersen et al. (2015a). Seven taxa were not analyzed. Of them, the positioning of Talaud Kingfisher, Todiramphus enigma, Sombre Kingfisher, Todiramphus funebris, and Cinnamon-banded Kingfisher, Todiramphus australasia, is particularly speculative.

Alcedininae: River Kingfishers Rafinesque, 1815

Cerylinae: Water Kingfishers Reichenbach, 1851

Halcyoninae: Tree Kingfishers Vigors, 1825

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