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AFROAVES IV

Piciformes

Afroaves

Accipitrimorphae

Strigiformes

Picimorphae

Coliiformes

Leptosomiformes

Trogoniformes

Bucerotiformes

Coraciiformes

Piciformes

The 47 Orders

Palaeognathae

Galloanserae

Mirandornithes

Columbaves

Otidimorphae

Columbimorphae

Elementaves

Opisthocomimorphae

Gruimorphae

Ardeae

Strisores

Telluraves

Afroaves

Australaves

PICIFORMESLinnaeus, 1758

Click for Piciformes tree
Click for Piciformes tree

Linnaeus used order Picae and the genusPicus. Seems like aslam-dunk! Then why do others attribute Piciformes to Meyer and Wolf, 1810?

The Piciformes fossil record is shorter than that of the other orders in Afroaves. The Sylphornithidae are thought to be stem Piciformes. They are known fromEurope starting in the middle Eocene. Recognizable Pici fossils are first found inthe early Oligocene.

GALBULIVigors, 1825

The jacamars and puffbirds are more closely related to each other thanto the rest of the Piciformes. They are sometimes placed in their own order,Galbuliformes. The higher-level taxonomy of the Piciformes is based on Ericson et al. (2006a), Johansson and Ericson (2003) and Moyle(2004). Details of the Galbulidae and Bucconidae are from Witt (2004), whoincluded most of the species in his analysis.

Galbulidae: JacamarsVigors, 1825

5 genera, 19 speciesHBW-7

The Spot-tailed Jacamar,Galbula rufoviridis (includingheterogyna), has been split from the Rufous-tailed Jacamar,Galbula ruficauda. There are indications that further splitsmight be needed. See Witt (2004).

Bucconidae: PuffbirdsHorsfield, 1821

12 genera, 36 speciesHBW-7

The Striolated Puffbird,Nystalus striolatus, has been split into Western Striolated-Puffbird,Nystalus obamai, andEastern Striolated-Puffbird,Nystalus striolatus. See Whitney et al. (2013e) andSACC #617 and#679.

The Chestnut-capped Puffbird is placed in genusCyphos (Spix 1824)rather thanArgicus (Cabanis and Heine 1863) becauseCyphosis not preoccupied byCyphus under current ICZN rules.

PICILinnaeus, 1758

Megalaimidae: Asian BarbetsBlyth, 1852

2 genera, 34 speciesHBW-7 (split)

Barbet and Toucan taxonomy has not been completely resolved(see Barker and Lanyon, 2000; Moyle, 2004). The five families used hererepresent one possibility. Another is to roll all five into one family,Ramphastidae, as in the Howard and Moore checklist (Dickinson andRemsen, 2013). Another is to keep four of the five familes and merge thetoucan-barbets into either the toucans (most likely) or New Worldbarbets (possibly). The position of bothCaloramphus andTrachyphonus among the other barbets has also not been completelyresolved.

Click for Megalaimidae tree
Click forMegalaimidae tree

Moyle (2004) found that the Fire-tufted Barbet is embedded deep within the traditionalMegalaima. I have put them all in the same genus as a result.The genus namePsilopogon (Muller 1835, typepyrolophus) replacesMegalaima (G.R. Gray 1842, typevirens) due to priority.Although this changes the type genus, the family name remains Megalaimidae.

More recently, den Tex and Leonard (2013) analyzed all of the Megalaimidae.The current species tree is based on their analysis. They recommended elevating several taxa to species level. Four of thoserecommendations are followed here. (1) The Sooty Barbet,Caloramphushayii has been split from Brown Barbet,Caloramphus fuliginosus.(2) Blue-eared Barbet,Psilopogon duvaucelii has been split fromYellow-eared Barbet,Psilopogon australis. The later is consideredmonotypic. Sincecyanotis andduvaucelii are thought to hybridizein Thailand, the remaining races are assigned to speciesduvaucelii.(3) The Golden-faced Barbet,Psilopogon chrysopsis(monotypic), has been split from Golden-whiskered Barbet,Psilopogonchrysopogon. (4) Finally, the Turquoise-throated Barbet,Psilopogonchersonesus, which is locally endenmic on the Kra Isthmus, has beensplit from the Blue-throated Barbet,Psilopogon asiatica. The name"Golden-faced Barbet" is used in the absence of any established English name.

Den Tex and Leonard (2013) also recommend splittingPsilopogon auricularisfromPsilopogon franklinii. However, these taxa are believed to hybridizein Vietnam (Annam) and nearby Laos. Given that, I think additional information isneeded to split them. They also note thatPsilopogon asiatica likely containsat least one more species, but that further study is required to sort outthe situation.

Lybiidae: African Barbets, TinkerbirdsSibley & Ahlquist, 1985

The 4th edition of the Howard and Moore checklist (Dickinson and Remsen,2013) separatedPogonornis (Billberg 1828,typedubius) fromLybius. However, although they are somewhat sparse, the results inMoyle (2004) suggest that more changes are needed. The genusTricholaema is restricted to the Hairy-breasted Barbet. Two formerTricholaema move toLybius (Spot-flanked and Black-throatedBarbets). The other three formerTricholaema form the genusNotopogonius (Roberts 1922, typeleucomelas).

11 genera, 42 speciesHBW-7 (split)

Capitonidae: New World BarbetsBonaparte, 1838

2 genera, 15 speciesHBW-7 (split)

The arrangement of species withinCapito is based on the maximum likelihood treein Armenta et al. (2005), which did not include the White-mantled Barbet or the recentlydiscovered Sira Barbet (Seeholzer et al., 2012). Interestingly, this is the second recently discovered barbet in Peru, the other being the closely related Scarlet-banded Barbet (O'Neill et al., 2000). Although the genetic distance between themis fairly small, they have developed distinctive plumages are here treated as separate species.

Semnornithidae: Toucan-barbetsPrum, 1988

1 genus, 2 speciesHBW-7 (split)

Ramphastidae: Toucans, Aracaris, ToucanetsVigors, 1825

6 genera, 44 speciesHBW-7

Ramphastidae Tree
Click for Ramphastidae Tree

Moyle (2004) laid out the overall structure of the toucans used here, as shownin the tree diagram (Barker and Lanyon, 2000, is a bit different).

The age of the Ramphastidae crown-group is rather uncertain. Patelet al. (2011) estimate it at about 11 million years ago (lateMiocene), while Lutz et al. (2013) estimate it about about 21 millionyears (early Miocene). Lutz et al. estimate the common ancestor withSemnornisat 29 mya (±5) while Patel et al. put it at 13 mya (±3).

The ordering of theRamphastos toucans is based on Patané et al.(2009). The Toco Toucan is basal, and the rest fall into two sistergroups, the smooth-billed yelping toucans (ambiguus throughtucanus) and the channel-billed croaking toucans(sulfuratus throughdicolorus). Although previous studiesby Weckstein (2004, 2005) suggested the species limits neededadjustment, the analysis by Patané et al. (2009), which used moregenes and taxa, did not concur.

The English name ofRamphastos ambiguus is changed to Yellow-throatedToucan. The point is that Black-mandibled properly applies only to theambiguus group, but is not appropriate whenswainsonii(Chestnut-mandibled Toucan) is included in the species. SeeSACC #663.

The aracari (Pteroglossus) sequence is based on Patel et al. (2011), which builds on Kimura et al. (2004) and Pereira and Wajntal (2008). The aracaris fall into several clades: (1)bailloni,viridis, andinscriptus (includinghumboldti);(2)torquatus (includingnuchalis anderythrozonus),frantzii,sanguineus, anderythropygius;(3)azara (includingflavirostris andmariae),bitorquatus andbeauharnaesii;and (4)aracari,castanotis, andpluricinctus.Note thatPteroglossus bailloni, which has sometimes beenconsidered a separate genus (Baillonius) is firmly embedded inPteroglossus.

The subspeciesP. azara mariae is sometimesseparated as Brown-mandibled Aracari, leavingazara andflavirostrisjoined as Ivory-billed Aracari. However, Patel et al. (2011) found thatmariaeis more closely related toflavirostris than either is toazara.Since the genes don't match the usual grouping of these taxa, and since itis unclear whether they are separate biological species, I'm leavingthem all lumped into Ivory-billed Aracari for now, although it is possible that2 or 3 species are involved here.

I do not follow SACC concerning thePteroglossus torquatus complex.Haffer (1967) found a narrow hybridization zone betweenP. torquatusandP. sanguineus (10-20 km across). This sort of hybridization zoneis usually taken as evidence of biologically separate species. There is someweak support from Patel et al. (2011) who found no evidence of hybridization.However, their sample size is small and this cannot be taken as strong evidence.The genetic differences do not compel either way.Ridgely and Greenfield (2001) report limited hybridization betweenP. sanguineus andP. erythropygius near Playa de Oro, Ecuador, and Short and Horne (2001) report more extensive hybridization near Gualea.The exact size of the hybridization zone is not known, but it appears tobe fairly limited. The result of all this is that I recognizeStripe-billed Aracari,Pteroglossus sanguineus, andPale-mandibled Aracari,Pteroglossus erythropygius, as distinct species, pending more definitive data.

The sequence of theAndigena andSelenidera toucans is basedon Lutz et al. (2013). The Yellow-eared Toucanet appears to group with the mountain-toucans rather than the other yellow-eared toucanets. Since no genusname is available, and Lutz et al. declined to create one, I'm listing itas"Selenidera" spectabilis. One alternative would be to include themountain-toucans inSelenidera, another would be to move theYellow-eared Toucanet intoAndigena. Given distinctive appearanceof the mountain-toucans, I think both are bad ideas. There is somequestion about whether the Guianan Toucanet belongs with the otheryellow-eared toucanets (this is why Lutz et al. were reluctant to create anew genus).

Two molecular studies of theAulacorhynchus toucanets have focused ondifferent sets of species. Puebla-Olivares et al. (2008) concentrated on theemerald toucanet complex, while Bonaccorso et al. (2011) focus on the others.

Bonaccorso et al. (2011) found that the Chestnut-tipped Toucanet(Aulacorhynchus derbianus) contains two groups that are not each other's closestrelatives. Following their recommendataions, it has been split into Whitely'sToucanet,Aulacorhynchus whitelianus (subspeciesduidae,whitelianus, andosgoodi) and Derby's Toucanet,Aulacorhynchusderbianus (subspeciesnigrirostris andderbianus). They alsoexamined samples of all three subspecies of the Groove-billed Toucanet,Aulacorhynchus sulcatus. AlthoughA. s. calorhynchus is sometimesconsidered a separate species (Yellow-billed Toucanet), it was nested within thesulcatus clade. The samples ofcalorhynchus formed a monophyletic group,suggesting that speciation is underway, but further study will be needed to clarifywhether is has yet reached the point where it should be considered a separatebiological species.

Some authors had previously considered the Emerald Toucanet to includemultiple species (e.g., Navarro et al., 2001). The paper by Puebla-Olivares etal. (2008) considerably strengthens the case, and I have split them accordingly. Except for the treatment ofalbivitta andgriseigularis asseparate species, Clements 6th edition also follows the split. SACC has not yetconsidered the issue.

The Middle American portion of the Emerald Toucanet complex is fairlyclearcut. This includesA. cognatus throughA. prasinus withmaxillaris included inA. caeruleogularis and all of the otherMiddle American subspecis inA. prasinus. The only real question is whether to lumpA. wagleri intoA. prasinus.

The South American Emerald Toucanets are another matter. There aresome key gaps in the taxa analyzed by Puebla-Olivares et al., and the situation concerningalbivitta is rather confusing. The raceslautus(believed to be a distinct species) andphaeolaemus (probably closest togriseigularis) were not analyzed. Moreover, the samples from NE Ecuador,believed to bealbivitta, do not appear to bealbivitta. Thiscreates a hole at the boundary betweenalbivitta andatrogularis. For the present,atrogularis includescyanolaemusanddimidiatus as subspecies.

Conventional wisdom has been thatgriseigularis andphaeolaemusgroup withalbivitta. But Puebla-Olivares et al. foundgriseigularis in atight grouping withatrogularis and the Ecuadorian‘albivitta’. What is one to think? It could be that there isan un-named white-throated subspecies in Ecuador and perhaps southernColombia. It could equally be that there was a glitch somewhere in thecollection or analysis of the Ecuadorian‘albivitta’.In any event, it makes the situation unclear. I have leftA. griseigularisas a separate species pending a resolution of the problem, although I expect itto be folded into eitherA. atrogularis orA. albivitta.

Indicatoridae: HoneyguidesSwainson, 1837

4 genera, 17 speciesHBW-7

Picidae: WoodpeckersLeach, 1820

39 genera, 237 speciesHBW-7

Picadae tree
Click for Picidae genera

The details of woodpecker taxonomy are still being worked out, with speciesbeing shuffled around between genera (e.g., in Picoides and Veniliornis). Thepapers by Benz et al. (2006), Benz and Robbins (2011), Dufort (2016), Fuchs etal. (2006c, 2007b, 2008a), Fuchs and Pons (2015), Moore et al. (2006), Overtonand Rhoads (2006), Webb and Moore (2005), and Weibel and Moore (2002a, 2002b)have all proven helpful.

Starting with version 3.02, I am primarily following Dufort (2016), withassistance from Shakya et al. (2017). If you examine the two papers you willfind some discord in the overall arrangement of taxa. I've generally givenpreference to Dufort et al. on the grounds that they generally consider moredata. The diagram sums it up.

Woodpecker Phylogeny

That the wrynecks were sister to the remaining woodpeckers has been known for a while, as has the fact that the piculets were sisterto the rest. Benz et al. (2006) showed that this is not exactly true. TheAntillean Piculet does not belong with the other piculets, but is the sister ofthe rest ot the Picinae. The time-aligned phylogeny of Dufort (2016) suggeststhat is it only a slightly older branch than various pieces of Picidae. Accordingly, it is placed in the monotypic tribe Nesoctitini within Picidae.But there is still more! The rest of the piculets are not monophyletic. Rather,some of the Old World piculets (Sassinae) are sister to Picinae, while the others,almost all from Latin America, are on an older branch (Picumninae).

The situation with the Old World piculets is complex. The AfricanPiculet,Verreauxia africana, has been removed fromSasia, whichis now restricted to Asia (see H&M-4 and Dufort, 2016). Also, the SpeckledPiculet,Vivia innominata, has been removed fromPicumnus, whichnow consists solely of New World species (see Dufort, 2016). Thus one Old Worldpiculet (Vivia) is sister to the Latin American piculets, while the other threeare on a separate branch that is sister to Picinae.

Fuchs et al. (2007b, 2008a) found thatHemicircus is sister to theremaining Picinae. Whether it is closer to the Antillean Piculet, or to the remainderof Picinae is not entirely clear. The TiF list now uses Dufort (2016) for thestructure of the rest of the Picinae, which takes the latter position. It nowseems likely there are two groups: Chrysocolaptini/Campephilini, andPicini/Melanerpini/Picoidini. Each of the tribes seems to be a relatively deepclade, and Melanerpini and Picoidini are sister tribes.

Picini treeThe key point that Fuchs et al. (2013) resolved is the relation betweenCampephilus (Campephilini) and an Asian clade containing the flamebacks(Chrysocolaptini). Morphology has always suggested they are closely related, butgenetic analyses were yielding conflicting results. Fuchs et al. provideevidence that this is due to a hybridization event that occured when themelanerpine andCampephilus woodpeckers colonized the Americas. Theyfound thatBlythipicus throughChrysocolaptes is sister toCampephilus.

Picini seems to look something like the tree to the right (Dufort, 2016). HereCeleus brachyurus has been placed in the monotypic genusMicropternus,Chrysophlegma has been separated fromPicus,andPiculus rubiginosus andP. rivolii move toColaptes. Notice thatCampethera has been merged intoGeocolaptes. This isbecause Dufort (2016) found that the Ground Woodpecker,Geocolaptesolivaceus, is embedded inCampethera. Fuchs et al. (2017)examined the entire genus and the species tree forGeocolaptesis based on their work. SinceGeocolaptes (Burchell 1832, typeolivaceus) has priority overCampethera (G.R. Gray 1841, typemaculosa), the combined genus must take the nameGeocolaptes. Theorder of species inPicus andColaptes is now based Dufort(2016).

Based on Fuch et al. (2017), I have split Fine-banded Woodpecker,Geocolaptes taeniolaema, (includinghausburgi) from Tullberg'sWoodpecker,Geocolaptes tullbergi. Fuchs et al. (2017) have noted someother possible splits, but I find those splits less compelling in the absence ofcloser study.

Contrary to studies using fewer genes,Dryocopus appears to beparaphyletic. It seems likely that the division is between New World and OldWorld species, with the Old WorldDryocopus being closer toMulleripicus. The New WorldDryocopus then take the nameHylatomus (Baird 1858, typepileatus). Note that the HelmetedWoodpecker has been transferred toCeleus fromHylatomus. See Benzet al. (2015) and Lammertink et al. (2016).

The generaChryserpes,Melanerpes,Sphyrapicus, andXiphidiopicus group together (see Dufort, 2016; Overton and Rhoads, 2006for details). Here they are treated as tribe Melanerpini. Dufort (2016) foundthat the Hispaniolan Woodpecker does not belong toMelanerpes, but isbasal in the tribe. It now takes the nameChryserpes striatus.The position ofXiphidiopicus is rather uncertain as there are problemswith the only available genetic data.

Picini treeThis brings us to the rest of the woodpeckers, the Picoidini. Dufort (2016)has decent coverage of this clade, and when combined with Fuchs and Pons (2015),I was able to make a tentative tree. Fuchs et al. (2017) analyzed DNA from allspecies inChloropicus, and the species tree follows their analysis.

There have been a number of changes in the Picoidini. Here I compare withH&M-3 (Dickinson, 2003) to better see what has happened.(1) The genusPicoides has been split into four pieces:Picoides,part ofDryobates, most ofLeuconotopicus, and two membersofVeniliornis.(2) The genusDendrocopos has also been split intoYungipicus,Dendrocoptes,Leiopicus,Dendrocopos, and the rest ofDryobates. On the plus side,Dendrocopos also absorbedHypopicus andSapheopipo (see Winkler et al., 2005).(3)Veniliornis lost the Smoky-brown Woodpecker toLeuconotopicus,but gained the South AmericanPicoides.

H&M-4 (Dickinson and Remsen, 2013) brought attention to the nameChloropicus (Malherbe 1845, typepyrrhogaster), which has priorityoverDendropicos (Malherbe 1849, typefuscescens, subspecieslafresnayi), so the nameDendropicos is replaced byChloropicus. With these changes, the TiF take on Picoidini is shown inthe diagram.

Further Issues

H&M-4 have a different approach toChloropicus, treating it as four genera:Chloropicus,Dendropicos,Mesopicos, andIpophilus. Although the taxon sampling is rather sparse, Fuchs and Pons (2015)find thatIpophilus is embeded inMesopicos and suggests bothtaxa are embeded inDendropicos. A further complication is thatthe Yellow-crowned Woodpecker,Leiopicus mahrattensis, is also in thisgroup. The tree using the most data suggestsLeiopicus (Bonaparte 1854,typemahrattensis) andChloropicus are sisters, and that the threeform a clade. However, there is also some support in the paper for puttingLeiopicus as basal among the three withChloropicus andDendropicos sister. The ongoing study of this clade by Fuchs, Bowie,Carre and Pons should resolve the question. Given the current uncertainties, Iprefer to lump them for under the nameChloropicus.

A few species besides those mentioned above have changed genera. Note that the nameDendrocoptes (Cabanis and Heine 1863, typemedius), has priority overDesertipicus (Kinnear and Bates 1935,typedorae). Also, the Checkered and Striped Woodpeckers (formerlyPicoides mixtus andP. lignarius) moved toVeniliornis.

Splits

Chrysocolaptini: The former Greater Flamebackhas been split into 6 species (Collar, 2011). Three of the new species aremonotypic:haematribon,xanthocephalus, anderythrocephalus;whilestrictus includeskangeanensis and bothrufopunctatus andmontanus are included inlucidus. Theremaining subspecies belong toguttacristatus, which retains the nameGreater Flameback.

Campephilini:Ivory-billed Woodpecker,Campephilus principalis has been split intoAmerican Ivory-billed Woodpecker,Campephilus principalis, and CubanIvory-billed Woodpecker,Campephilus bairdii, based on Fleischer et al.(2007) and Dufort (2016).

Picini: Shakya et al. (2017) found that the Olive-backed Woodpecker,Dinopiumrafflesii, is sister toGecinulus. As a result, I've placed it inthe monotypic genusChloropicoides (Malherbe 1848-49).

Following Collar (2011), the Philippine race of Common Flameback,Dinopium javanense, has been elevated to species status asSpot-throated Flameback,Dinopium everetti.

The Red-backed Flameback,Dinopium psarodes, has been split from theBlack-rumped Flameback,Dinopium benghalense. Fernando et al. (2016)makes the strongest case for the split. Note that the racejaffnensefrom northern Sri Lanka remains a subspecies of Black-rumped Flameback,Dinopium benghalense. There seems to be a stable hybrid zone betweenthem. Fernando and Seneviratne (2015) and Freed et al. (2015) have moreinformation on this.

The Iberian Green Woodpecker,Picus sharpei, has been split from theEuropean Green-Woodpecker,Picus viridis, based on Perktas et al. (2011)and Pons et al. (2011). The TiF list tries to use the biological speciesconcept when possible. As pointed out by Perktas et al., the case forbiological species status for the Zagros Green-Woodpecker remains weak, so itremains a subspecies,Picus viridis innominatus.

The Sooty Woodpecker,Mulleripicus funebris, has been split into SootyWoodpecker / Southern Sooty-Woodpecker,Mulleripicus fuliginosus, andFunereal Woodpecker / Northern Sooty-Woodpecker,Mulleripicus funebris,based on Dufort (2016).

As recommended by Benz and Robbins (2011), I've split Ochre-backed Woodpecker,Celeus ochraceus, from Blond-crested Woodpecker,Celeus flavescens.

Moore et al. (2011) provided evidence that the Bronze-winged Woodpecker,Colaptes aeruginosus, of NE Mexico is sister to the Gray-crowned Woodpecker,Colaptes auricularis, rather than being a subspecies of the Golden-oliveWoodpecker,Colaptes rubiginosus (which itself is sister to Black-neckedWoodpecker,Colaptes atricollis). Further, from Dufort (2016), it seems thatthe Bronze-winged Woodpecker also includes the subspeciesyucatanensis. As a result, it takes the scientific nameColaptesyucatanensis asyucatanensis (S. Cabot, 1844) has priority overaeruginosus (Malherbe, 1862). These two taxa may be separate species, butmore study is need here and elsewhere in the Golden-olive complex.

Melanerpini:García-Trejo et al. (2009) found that the northern subspecies of theGolden-fronted Woodpecker,Melanerpes aurifrons, is more closely relatedto the Red-bellied Woodpecker,Melanerpes carolinus, than to otherGolden-fronted races. Accordingly, they recommend splitting the other races asthe tropicalMelanerpes santacruzi, known as Velasquez's Woodpecker. Itis possible that further splitting will be needed. The names Lesson'sWoodpecker and Truxillo Woodpecker have been applied to some of the othertropical races. I had previously arranged theCenturus woodpeckers basedon their work. Navarro-Sigüenza et al.\ (2017) have recently taken anotherlook at the complex, using more genes. The current arrangement, as well as therecognition ofCenturus (Swainson, 1837, typecarolinus) is basedon Navarro-Sigüenza et al.

Click for Picidae species tree
Click for Picidae species tree

Jynginae: WrynecksSwainson, 1831

Picumninae: PiculetsG.R. Gray, 1840

Sasiinae: Old World PiculetsInformal

Picinae: WoodpeckersLeach, 1820

Nesoctitini: Antillean PiculetWolters, 1976

HemicirciniCabanis and Heine, 1863

ChrysocolaptiniBonaparte, 1854

CampephiliniBlyth, 1852

PiciniLeach, 1820

MelanerpiniG.R. Gray, 1846

PicoidiniOlphe-Galliard, 1888 (1846)

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Taxonomy in Flux:Version 3.49, June 19, 2024 (August 9, 2019).Copyright © 2008–2024 by John H. Boyd III
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