Afroaves

Accipitrimorphae

Strigiformes

Picimorphae

Coliiformes

Leptosomiformes

Trogoniformes

Bucerotiformes

Coraciiformes

Piciformes

The 46 Orders

Paleognathae

Galloanserae

Columbimorphae

Otidimorphae

Strisores

Opisthocomiformes

Gruiformes

Mirandornithes

Ardeae

Charadriiformes

Telluraves

Afroaves

Australaves

PICIFORMES Linnaeus, 1758

Click for Piciformes tree
Click for Piciformes tree

Linnaeus used order Picae and the genus Picus. Seems like a slam-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 from Europe starting in the middle Eocene. Recognizable Pici fossils are first found in the early Oligocene.

GALBULI Vigors, 1825

The jacamars and puffbirds are more closely related to each other than to 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), who included most of the species in his analysis.

Galbulidae: Jacamars Vigors, 1825

5 genera, 19 species HBW-7

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

Bucconidae: Puffbirds Horsfield, 1821

12 genera, 36 species HBW-7

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

The Chestnut-capped Puffbird is placed in genus Cyphos (Spix 1824) rather than Argicus (Cabanis and Heine 1863) because Cyphos is not preoccupied by Cyphus under current ICZN rules.

PICI Linnaeus, 1758

Megalaimidae: Asian Barbets Blyth, 1852

2 genera, 34 species HBW-7 (split)

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

Click for Megalaimidae tree
Click for Megalaimidae tree

Moyle (2004) found that the Fire-tufted Barbet is embedded deep within the traditional Megalaima. I have put them all in the same genus as a result. The genus name Psilopogon (Muller 1835, type pyrolophus) replaces Megalaima (G.R. Gray 1842, type virens) 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 those recommendations are followed here. (1) The Sooty Barbet, Caloramphus hayii has been split from Brown Barbet, Caloramphus fuliginosus. (2) Blue-eared Barbet, Psilopogon duvaucelii has been split from Yellow-eared Barbet, Psilopogon australis. The later is considered monotypic. Since cyanotis and duvaucelii are thought to hybridize in Thailand, the remaining races are assigned to species duvaucelii. (3) The Golden-faced Barbet, Psilopogon chrysopsis (monotypic), has been split from Golden-whiskered Barbet, Psilopogon chrysopogon. (4) Finally, the Turquoise-throated Barbet, Psilopogon chersonesus, which is locally endenmic on the Kra Isthmus, has been split 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 splitting Psilopogon auricularis from Psilopogon franklinii. However, these taxa are believed to hybridize in Vietnam (Annam) and nearby Laos. Given that, I think additional information is needed to split them. They also note that Psilopogon asiatica likely contains at least one more species, but that further study is required to sort out the situation.

Lybiidae: African Barbets, Tinkerbirds Sibley & Ahlquist, 1985

The 4th edition of the Howard and Moore checklist (Dickinson and Remsen, 2013) separated Pogonornis (Billberg 1828,type dubius) from Lybius. However, although they are somewhat sparse, the results in Moyle (2004) suggest that more changes are needed. The genus Tricholaema is restricted to the Hairy-breasted Barbet. Two former Tricholaema move to Lybius (Spot-flanked and Black-throated Barbets). The other three former Tricholaema form the genus Notopogonius (Roberts 1922, type leucomelas).

11 genera, 42 species HBW-7 (split)

Capitonidae: New World Barbets Bonaparte, 1838

2 genera, 15 species HBW-7 (split)

The arrangement of species within Capito is based on the maximum likelihood tree in Armenta et al. (2005), which did not include the White-mantled Barbet or the recently discovered 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 them is fairly small, they have developed distinctive plumages are here treated as separate species.

Semnornithidae: Toucan-barbets Prum, 1988

1 genus, 2 species HBW-7 (split)

Ramphastidae: Toucans, Aracaris, Toucanets Vigors, 1825

6 genera, 44 species HBW-7

Ramphastidae Tree
Click for Ramphastidae Tree

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

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

The ordering of the Ramphastos toucans is based on Patané et al. (2009). The Toco Toucan is basal, and the rest fall into two sister groups, the smooth-billed yelping toucans (ambiguus through tucanus) and the channel-billed croaking toucans (sulfuratus through dicolorus). Although previous studies by Weckstein (2004, 2005) suggested the species limits needed adjustment, the analysis by Patané et al. (2009), which used more genes and taxa, did not concur.

The English name of Ramphastos ambiguus is changed to Yellow-throated Toucan. The point is that Black-mandibled properly applies only to the ambiguus group, but is not appropriate when swainsonii (Chestnut-mandibled Toucan) is included in the species. See SACC #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, and inscriptus (including humboldti); (2) torquatus (including nuchalis and erythrozonus), frantzii, sanguineus, and erythropygius; (3) azara (including flavirostris and mariae), bitorquatus and beauharnaesii; and (4) aracari, castanotis, and pluricinctus. Note that Pteroglossus bailloni, which has sometimes been considered a separate genus (Baillonius) is firmly embedded in Pteroglossus.

The subspecies P. azara mariae is sometimes separated as Brown-mandibled Aracari, leaving azara and flavirostris joined as Ivory-billed Aracari. However, Patel et al. (2011) found that mariae is more closely related to flavirostris than either is to azara. Since the genes don't match the usual grouping of these taxa, and since it is unclear whether they are separate biological species, I'm leaving them all lumped into Ivory-billed Aracari for now, although it is possible that 2 or 3 species are involved here.

I do not follow SACC concerning the Pteroglossus torquatus complex. Haffer (1967) found a narrow hybridization zone between P. torquatus and P. sanguineus (10-20 km across). This sort of hybridization zone is usually taken as evidence of biologically separate species. There is some weak 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 between P. sanguineus and P. 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 to be fairly limited. The result of all this is that I recognize Stripe-billed Aracari, Pteroglossus sanguineus, and Pale-mandibled Aracari, Pteroglossus erythropygius, as distinct species, pending more definitive data.

The sequence of the Andigena and Selenidera toucans is based on Lutz et al. (2013). The Yellow-eared Toucanet appears to group with the mountain-toucans rather than the other yellow-eared toucanets. Since no genus name is available, and Lutz et al. declined to create one, I'm listing it as "Selenidera" spectabilis. One alternative would be to include the mountain-toucans in Selenidera, another would be to move the Yellow-eared Toucanet into Andigena. Given distinctive appearance of the mountain-toucans, I think both are bad ideas. There is some question about whether the Guianan Toucanet belongs with the other yellow-eared toucanets (this is why Lutz et al. were reluctant to create a new genus).

Two molecular studies of the Aulacorhynchus toucanets have focused on different sets of species. Puebla-Olivares et al. (2008) concentrated on the emerald 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 closest relatives. Following their recommendataions, it has been split into Whitely's Toucanet, Aulacorhynchus whitelianus (subspecies duidae, whitelianus, and osgoodi) and Derby's Toucanet, Aulacorhynchus derbianus (subspecies nigrirostris and derbianus). They also examined samples of all three subspecies of the Groove-billed Toucanet, Aulacorhynchus sulcatus. Although A. s. calorhynchus is sometimes considered a separate species (Yellow-billed Toucanet), it was nested within the sulcatus clade. The samples of calorhynchus formed a monophyletic group, suggesting that speciation is underway, but further study will be needed to clarify whether is has yet reached the point where it should be considered a separate biological species.

Some authors had previously considered the Emerald Toucanet to include multiple species (e.g., Navarro et al., 2001). The paper by Puebla-Olivares et al. (2008) considerably strengthens the case, and I have split them accordingly. Except for the treatment of albivitta and griseigularis as separate species, Clements 6th edition also follows the split. SACC has not yet considered the issue.

The Middle American portion of the Emerald Toucanet complex is fairly clearcut. This includes A. cognatus through A. prasinus with maxillaris included in A. caeruleogularis and all of the other Middle American subspecis in A. prasinus. The only real question is whether to lump A. wagleri into A. prasinus.

The South American Emerald Toucanets are another matter. There are some key gaps in the taxa analyzed by Puebla-Olivares et al., and the situation concerning albivitta is rather confusing. The races lautus (believed to be a distinct species) and phaeolaemus (probably closest to griseigularis) were not analyzed. Moreover, the samples from NE Ecuador, believed to be albivitta, do not appear to be albivitta. This creates a hole at the boundary between albivitta and atrogularis. For the present, atrogularis includes cyanolaemus and dimidiatus as subspecies.

Conventional wisdom has been that griseigularis and phaeolaemus group with albivitta. But Puebla-Olivares et al. found griseigularis in a tight grouping with atrogularis and the Ecuadorian ‘albivitta’. What is one to think? It could be that there is an un-named white-throated subspecies in Ecuador and perhaps southern Colombia. It could equally be that there was a glitch somewhere in the collection or analysis of the Ecuadorian ‘albivitta’. In any event, it makes the situation unclear. I have left A. griseigularis as a separate species pending a resolution of the problem, although I expect it to be folded into either A. atrogularis or A. albivitta.

Indicatoridae: Honeyguides Swainson, 1837

4 genera, 17 species HBW-7

Picidae: Woodpeckers Leach, 1820

38 genera, 237 species HBW-7

Picadae tree
Click for Picidae genera

The details of woodpecker taxonomy are still being worked out, with species being shuffled around between genera (e.g., in Picoides and Veniliornis). The papers by Benz et al. (2006), Benz and Robbins (2011), Dufort (2016), Fuchs et al. (2006c, 2007b, 2008a), Fuchs and Pons (2015), Moore et al. (2006), Overton and 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), with assistance from Benz and Robbins (2011) for Celeus and Fuchs and Pons (2015) for Picoidini. 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 sister to the rest. Benz et al. (2006) showed that this is not quite true. The Antillean Piculet does not belong with the other piculets, but is the sister of the Picinae. The time-aligned phylogeny of Dufort (2016) suggests that is it only a slightly older branch than various pieces of Picidae. Accordingly, it is placed in the monotypic tribe Nesoctitini within Picidae.

There have been a couple of generic changes in the piculets. The African Piculet, Verreauxia africana, has been removed from Sasia, which is now restricted to Asia (see H&M-4 and Dufort, 2016). Also, the Speckled Piculet, Vivia innominata, has been removed from Picumnus, which now consists solely of New World species (see Dufort, 2016).

Fuchs et al. (2007b, 2008a) found that Hemicircus is sister to the rest of the Picinae. The TiF list now uses Dufort (2016) for the structure of the rest of the Picinae. It now seems likely there are two groups: Chrysocolaptini/Campephilini, and Picini/Melanerpini/Picoidini. Each of the tribes seems to be a relatively deep clade, and Melanerpini and Picoidini are sister tribes.

Picini tree The key point that Fuchs et al. (2013) resolved is the relation between Campephilus (Campephilini) and an Asian clade containing the flamebacks (Chrysocolaptini). Morphology has always suggested they are closely related, but genetic analyses were yielding conflicting results. Fuchs et al. provide evidence that this is due to a hybridization event that occured when the melanerpine and Campephilus woodpeckers colonized the Americas. They found that Blythipicus through Chrysocolaptes is sister to Campephilus.

Picini seems to look something like the tree to the right (Dufort, 2016). Here Celeus brachyurus has been placed in the monotypic genus Micropternus, Chrysophlegma has been separated from Picus, and Piculus rubiginosus and P. rivolii move to Colaptes. Notice that Campethera has been merged into Geocolaptes. This is because Dufort (2016) found that the Ground Woodpecker, Geocolaptes olivaceus, is embedded in Campethera. Fuchs et al. (2017) examined the entire genus and the species tree for Geocolaptes is based on their work. Since Geocolaptes (Burchell 1832, type olivaceus) has priority over Campethera (G.R. Gray 1841, type maculosa), the combined genus must take the name Geocolaptes. The order of species in Picus and Colaptes is now based Dufort (2016).

Based on Fuch et al. (2017), I have split Fine-banded Woodpecker, Geocolaptes taeniolaema, (including hausburgi) from Tullberg's Woodpecker, Geocolaptes tullbergi. Fuchs et al. (2017) have noted some other possible splits, but I find those splits less compelling in the absence of closer study.

Contrary to studies using fewer genes, Dryocopus appears to be paraphyletic. It seems likely that the division is between New World and Old World species, with the Old World Dryocopus being closer to Mulleripicus. The New World Dryocopus then take the name Hylatomus (Baird 1858, type pileatus). Note that the Helmeted Woodpecker has been transferred to Celeus from Hylatomus. See Benz et al. (2015) and Lammertink et al. (2016).

The genera Chryserpes, Melanerpes, Sphyrapicus, and Xiphidiopicus group together (see Dufort, 2016; Overton and Rhoads, 2006 for details). Here they are treated as tribe Melanerpini. Dufort (2016) found that the Hispaniolan Woodpecker does not belong to Melanerpes, but is basal in the tribe. It now takes the name Chryserpes striatus. The position of Xiphidiopicus is rather uncertain as there are problems with the only available genetic data.

Picini tree This 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 all species in Chloropicus, and the species tree follows their analysis.

There have been a number of changes in the Picoidini. Here I compare with H&M-3 (Dickinson, 2003) to better see what has happened. (1) The genus Picoides has been split into four pieces: Picoides, part of Dryobates, most of Leuconotopicus, and two members of Veniliornis. (2) The genus Dendrocopos has also been split into Yungipicus, Dendrocoptes, Leiopicus, Dendrocopos, and the rest of Dryobates. On the plus side, Dendrocopos also absorbed Hypopicus and Sapheopipo (see Winkler et al., 2005). (3) Veniliornis lost the Smoky-brown Woodpecker to Leuconotopicus, but gained the South American Picoides.

H&M-4 (Dickinson and Remsen, 2013) brought attention to the name Chloropicus (Malherbe 1845, type pyrrhogaster), which has priority over Dendropicos (Malherbe 1849, type fuscescens, subspecies lafresnayi), so the name Dendropicos is replaced by Chloropicus. With these changes, the TiF take on Picoidini is shown in the diagram.

Further Issues

H&M-4 have a different approach to Chloropicus, treating it as four genera: Chloropicus, Dendropicos, Mesopicos, and Ipophilus. Although the taxon sampling is rather sparse, Fuchs and Pons (2015) find that Ipophilus is embeded in Mesopicos and suggests both taxa are embeded in Dendropicos. A further complication is that the Yellow-crowned Woodpecker, Leiopicus mahrattensis, is also in this group. The tree using the most data suggests Leiopicus (Bonaparte 1854, type mahrattensis) and Chloropicus are sisters, and that the three form a clade. However, there is also some support in the paper for putting Leiopicus as basal among the three with Chloropicus and Dendropicos sister. The ongoing study of this clade by Fuchs, Bowie, Carre and Pons should resolve the question. Given the current uncertainties, I prefer to lump them for under the name Chloropicus.

A few species besides those mentioned above have changed genera. Note that the name Dendrocoptes (Cabanis and Heine 1863, type medius), has priority over Desertipicus (Kinnear and Bates 1935, type dorae). Also, the Checkered and Striped Woodpeckers (formerly Picoides mixtus and P. lignarius) moved to Veniliornis.

Splits

Chrysocolaptini: The former Greater Flameback has been split into 6 species (Collar, 2011). Three of the new species are monotypic: haematribon, xanthocephalus, and erythrocephalus; while strictus includes kangeanensis and both rufopunctatus and montanus are included in lucidus. The remaining subspecies belong to guttacristatus, which retains the name Greater Flameback.

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

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

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

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

The Sooty Woodpecker, Mulleripicus funebris, has been split into Sooty Woodpecker / Southern Sooty-Woodpecker, Mulleripicus fuliginosus, and Funereal 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-olive Woodpecker, Colaptes rubiginosus (which itself is sister to Black-necked Woodpecker, Colaptes atricollis). Further, from Dufort (2016), it seems that the Bronze-winged Woodpecker also includes the subspecies yucatanensis. As a result, it takes the scientific name Colaptes yucatanensis as yucatanensis (S. Cabot, 1844) has priority over aeruginosus (Malherbe, 1862). These two taxa may be separate species, but more study is need here and elsewhere in the Golden-olive complex.

Melanerpini: García-Trejo et al. (2009) found that the northern subspecies of the Golden-fronted Woodpecker, Melanerpes aurifrons, is more closely related to the Red-bellied Woodpecker, Melanerpes carolinus, than to other Golden-fronted races. Accordingly, they recommend splitting the other races as the tropical Melanerpes santacruzi, known as Velasquez's Woodpecker. It is possible that further splitting will be needed. The names Lesson's Woodpecker and Truxillo Woodpecker have been applied to some of the other tropical races. I had previously arranged the Centurus woodpeckers based on their work. Navarro-Sigüenza et al.\ (2017) have recently taken another look at the complex, using more genes. The current arrangement, as well as the recognition of Centurus (Swainson, 1837, type carolinus) is based on Navarro-Sigüenza et al.

Click for Picidae species tree
Click for Picidae species tree

Jynginae: Wrynecks Swainson, 1831

Picumninae: Piculets G.R. Gray, 1840

Picinae: Woodpeckers Leach, 1820

Nesoctitini: Antillean Piculet Wolters, 1976

Hemicircini Cabanis and Heine, 1863

Chrysocolaptini Bonaparte, 1854

Campephilini Blyth, 1852

Picini Leach, 1820

Melanerpini G.R. Gray, 1846

Picoidini Olphe-Galliard, 1888 (1846)

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