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AFROAVES III

Trogoniformes, Bucerotiformes & Coraciiformes

Afroaves

Strigiformes

Accipitrimorphae

Coliiformes

Picimorphae

Leptosomiformes

Trogoniformes

Bucerotiformes

Coraciiformes

Piciformes

The 47 Orders

Palaeognathae

Galloanserae

Mirandornithes

Columbaves

Otidimorphae

Columbimorphae

Elementaves

Opisthocomimorphae

Gruimorphae

Ardeae

Strisores

Telluraves

Afroaves

Australaves

COLIIFORMESMurie, 1872

The Coliiformes are a relict Afrotropical group consisting of 6 species in 2genera. It wasn't always so. There's a fairly extensive fossil record from Europe andNorth America. Primitive Coliiformes (Sandcoleidae) are known from the earlyEocence (48-56 mya). By the late Eocene, only modern forms are known (Mayr, 2009).

Coliidae: MousebirdsSundevall, 1836

2 genera, 6 speciesHBW-6

  1. Blue-naped Mousebird,Urocolius macrourus
  2. Red-faced Mousebird,Urocolius indicus
  3. Speckled Mousebird,Colius striatus
  4. White-headed Mousebird,Colius leucocephalus
  5. Red-backed Mousebird,Colius castanotus
  6. White-backed Mousebird,Colius colius

LEPTOSOMIFORMESSharpe, 1891

The Cuckoo Roller was originally considered a cuckoo, some affinities withthe rollers were noted, and it has more recently been considered its own family,Leptosomidae. Mayr (2008a) discusses some of thedifferences between the Cuckoo Roller and the true rollers. Darren Naish'scomments on the Cuckoo Rollerat Tetrapod Zoology are also worth reading.

This is another relict group, consisting of a single species from Madagascarand the Comoros. Fossils are again know from Europe and North America in theearly and middle Eocene.

Leptosomidae: Cuckoo RollerBlyth, 1838

1 genus, 1 speciesHBW-6

TROGONIFORMESAOU, 1886

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

One unusual feature of trogons is their heterodactyl foot. In somecases this allows fossils to be easily identified as trogoniformes. The trogonlineage seems to date to the late Paleocene (Jarvis et al., 2014; Prum et al., 2015)and early trogoniform fossils have been found from the London Clay and DanishFur Formations, both from the early Eocene (Mayr, 2009). There may also bean early Eocene fossil from the Green River Formation in North America.When we see a possible Laurasian distribution like this we should keep in mindthat there were land bridges between Europe and America in the Eocene(maps may be found in Brikiatis, 2014). We should also keep in mind that the fossil record of Paleogene birds is much better in Europe and North America thanin other locations, and that the pattern we see could reflect patterns ofpreservation and discovery rather than the actual distribution. Nonetheless, ourbest guess is that trogons originated in Laurasia, likely in Europe.

In spite of their ancient lineage, only one early trogon left any surviving descendants.The crown group of trogons diversified starting in the Miocene, roughly 20 mya (Oliveros,2015; Prum et al., 2016).

The geographic origin of the crown trogons is unknown, and investigation oftheir origin has been hampered by a lack of a solidly established phylogeny.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 avariety of arrangements. Oliveros's Ph.D. dissertation (2015) seems to have resolved the phylogenetic problems. The African trogons (Apaloderma) are the basal branch, the next split is between the Asian trogons and the New World trogons, which are sister clades.

So how do we get a distribution like that, keeping in mind that stem trogonshad a European or perhaps Laurasian distribution? We also have the informationthat the Trogoniformes are sister to the Bucerotiformes-Coraciiformes-Piciformesclade and that the Leptosomiformes are basal to all four orders. This fits inwell with the picture of Trogoniformes as having a Laurasian distribution.Oliveros also considers the Coliiformes as part of the picture, but Suh et al. (2015)ans Suh (2016) suggest that the Coliiformes are more distant relatives, leavingthe Strigiformes as the next nearest clade.

From a Laurasian distribution, it is easy to imagine some trogons moving toAfrica, and then an Asian clade splitting across the Bering Striat (orlandbridge) to the Americas, followed by a southward movement perhaps driven byclimate change. This is basically the scenario presented by Oliveros (2015).

Trogonidae: TrogonsLesson, 1828

7 genera, 44 speciesHBW-6

Click for Trogonidae tree
Click for Trogonidae tree

The AOU's South American checklist committee had considered making somechanges inTrogon species in 2003, partly due to the treatment inRidgely and Greenfield (2001) and Hilty (2003). For more information,read the discussion forviridis,violaceus, andmelanurus on the SACC site. They found that the relevant data,if it existed, was not collected together in a way that allowed them tojudge it properly. The publication of DaCosta and Klicka (2008) haschanged the balance of evidence here. Although the SACC had not then actedon it, I changed generic limits accordingly.

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

The SACC has passed (Apr. 8, 2009) a set of proposals concerning splittingmesurus frommelanurus,chionurus fromviridis,andcaligatus fromviolaceus.I've updated the table with the SACC English names. The NACC has also tentativelyratifed the relevant portions of these changes. The SACC has nowrecognizedthe Amazonian Trogon,Trogon ramonianus, as a separate species (Oct 6, 2010).They also adopted the English name Guianan Trogon for the now monotypicT. violacenous. Note that there are still some unresolved issues concerningthe exact distribution of these species.

SpeciesSubspecies
Collared/Orange-bellied Trogon
*Collared Trogon,T. collarisextimus, hoethinus, virginalis,
subtropicalis, exoptatus,
collaris, castaneus
*Orange-bellied Trogon,T. puellapuella, underwoodi,
aurantiiventris, flavidor
Black-tailed Trogon Complex
Lattice-tailed Trogon,T. clathratusclathratus
*Ecuadorian Trogon,T. mesurusmesurus
*Black-tailed Trogon,T. melanurusmacroura?, eumorphus,
occidentalis, melanurus
Blue-tailed Trogon,T. comptuscomptus
White-tailed Trogon Complex
Black-headed Trogon,T. melanocephalusmelanocephalus
Citreoline Trogon,T. citreoluscitreolus, sumichrasti
*Green-backed Trogon,T. viridisviridis, melanopterus?
*White-tailed Trogon,T. chionuruschionurus
Baird's Trogon,T. bairdiibairdii
Violaceous Trogon Complex
*Gartered Trogon,T. caligatus(sallaei akabraccatus),concinnus,
caligatus
*Amazonian Trogon,T. ramonianusramonianus, crissalis
Surucua Trogon,T. surrucuraaurantius, surrucura
*Guianan Trogon,T. violaceusviolaceus
Blue-crowned Trogon,T. curucuiperuvianus (bolivianus),
curucui, behni

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

These changes combine the Middle American Collared Trogons withOrange-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 notmatch the AOU because they don't includepuella in the Orange-bellied Trogon. I do, andit has priority.Thus the range of Orange-bellied extendsfrom Mexico to western Panama. The Collared Trogons of South America and Eastern Panama retain the nameCollared Trogon. The Black-tailed Trogons west of the Andes in Ecuadorand Peru become Ecuadorian Trogon. The White-tailed Trogon can be found from Panama to W Ecuador. The rest of the White-tailed Trogons are groupedas the Green-backed Trogon, even in Trinidad (the population insoutheast Brazil may be split at some point). Gartered Trogon ispresent from Mexico into S. America west of the Andes, and in the northinto western Venezuela. The Amazonian Trogon is in the Amazon Basin. Finally, the Guianan Trogon is present in eastern Venezuela, theGuianas, Trinidad, and northeastern Brazil.

BUCEROTIFORMESFürbringer, 1888

The Bucerotiformes are another ancient order.According to Mayr (2009), the only documented early fossil Bucerotiformesseem more closely related to the hoopoes and woodhoopoes. The Messelirrisoridaeare 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: HoopoesLeach, 1820

1 genus, 4 speciesHBW-6

Phoeniculidae: WoodhoopoesBonaparte, 1831

2 genera, 9 speciesHBW-6

Bucorvidae: Ground-HornbillsBonaparte, 1854

1 genus, 2 speciesNot HBW Family

Bucerotidae: HornbillsRafinesque, 1815

16 genera, 59 speciesHBW-6

Click for Bucerotidae tree
Click forBucerotidae tree

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

I have followed Hübner et al. (2003) and Gonzalez et al. (2013a) insplittingTockus into the whistlers (Lophoceros) and cluckers(Tockus). Besides the vocal distinctions, the split bewteen these clades isquite deep. Viseshakul et al. (2011) et al. estimate it at around 45 million years.Note the genus nameLophoceros (Ehrenberg 1833)has priority overRhynchaceros (Gloger 1842) as suggested by bothHübner et al. (2003) and Gonzalez et al. (2013a). (They subsequently correctedthis 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 inTockus. Here it has been placed sister toTropicranus in the monotypic genusHorizocerus (Oberholser 1899). The Sulawesi Hornbill,Penelopides exarhatus, doesn't belong withPenelopides. Instead, it joins Wrinkled, Walden's, and Writhed Hornbills. Viseshakul et al. (2011)had previously shown they did not belong inAceros, and I hadseparated them asCranobrontes (Riley 1921, type speciesleucocephalus). But now, sinceexarhatus has joined them,they must take the nameRhabdotorrhinus (Meyer and Wiglesworth 1898,typeexarhatus), which has priority overCranobrontes.

The genusAceros has been reduced to a single species. Besideslosing 3 species toRhabdotorrhinus, the other formerAceros hornbill, the Knobbed Hornbill, has been merged intoRhyticeros. It seems tobe rather distant from the otherRhyticeros hornbill and couldreasonably 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 includedin Viseshakul et al. (2011), which also supports this treatment.

CORACIIFORMESForbes, 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 threegroups: 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 ofthe three groups unresolved, while Hackett et al. (2008) considered thebee-eaters the basal group, with the roller and kingfisher groups sister. Themorphological analysis reported in Mayr (2009) had the bee-eaters grouped withthe kingfisher group, and the roller group basal.

Early fossil Coraciiformes include the families Primobucconidae, Eocoraciidae,Geranopteridae, and the speciesQuasisyndactylus longibranchis. As withmany other Afroavian orders, the earliest fossils are from the early Eocene inthe Northern Hemisphere (Clarke et al., 2009; Ksepka and Clarke, 2010), in thiscase including both North America and Europe (Clarke et al., 2009). Although thetodies are now restricted to the Caribbean, there are fossils from the Oligoceneof Europe as well as North America (Mayr, 2009).

MEROPIForbes, 1884

Meropidae: Bee-eatersRafinesque, 1815

3 genera, 27 speciesHBW-6

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

CORACIIForbes, 1884

Brachypteraciidae: Ground-RollersBonaparte, 1854

4 genera, 5 speciesHBW-6

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

Coraciidae: RollersRafinesque, 1815

2 genera, 12 speciesHBW-6

HALCYONIForbes, 1884

Todidae: TodiesVigors, 1825

1 genus, 5 speciesHBW-6

Todidae tree

The tody sequence is based on Overton and Rhoads (2004). They alsoconsidered 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: MotmotsG.R. Gray, 1840 (1832-33)

6 genera, 15 speciesHBW-6

The motmot sequence is based on Witt (2004).

The Blue-crowned Motmot complex has been split based onWitt (2004) and Stiles (2009). Witt found that theAndean (Highland) Motmot,Momotus aequatorialis (includingchlorolaemus), is not truly part of the Blue-crowned complex, but is sister to the combinedBlue-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.

NameAssociated SubspeciesRange
Blue-capped Motmot,Momotus coerulicepscoerulicepsNE Mexico: Nuevo Leon & Tamaulipas
Lesson's Motmot,Momotus lessoniigoldmani,exiguus,lessoniiMiddle America: Veracruz to W Panama (Chiriqui)
Amazonian Motmot,Momotus momotamicrostephanus,momota,ignobilis,simplex,cametensis,paraensis,marcgravianus,nattereri,pilcomajensisAmazon Basin
Silver-banded Motmot,Momotus argenticinctusargenticinctusW Ecuador and NW Peru, W of Andes
Trinidad Motmot,Momotus bahamensisbahamensisTrinidad & Tobago
Whooping Motmot,Momotus subrufescenssubrufescens,spatha,osgoodi,conexus*,reconditus*,olivaresi*C & E Panama and NW South America (Chocó, Caribbean slope, Magdalena Valley)
* = There is a case for submergingconexus,reconditus,olivaresi intosubrufescens.
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 iswhy I'm treating it as a separate species.

Alcedinidae: KingfishersRafinesque, 1815

19 genera, 116 speciesHBW-6

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

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

The Silvery Kingfisher,Ceyx argentatus, has been split into SouthernSilvery-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 splitinto 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 atDimorphic Dwarf-Kingfisher, and continuing to the end of Alcedininae.

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

Although data is lacking, the rearrangment increased the doubt about whether the twotaxa 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 resequencedTodiramphus 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 KingfishersRafinesque, 1815

Cerylinae: Water KingfishersReichenbach, 1851

Halcyoninae: Tree KingfishersVigors, 1825

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Taxonomy in Flux:Version 3.49 June 19, 2024 (December 11, 2016).Copyright © 2008–2024 by John H. Boyd III
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