Piciformes

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

PICIFORMES Linnaeus, 1758

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

Click for Piciformes tree
Click for Piciformes tree

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, 18 species HBW-7

Bucconidae: Puffbirds Horsfield, 1821

12 genera, 35 species HBW-7

Megalaimidae: Asian Barbets Blyth, 1852

2 genera, 30 species HBW-7 (split)

Barbet and Toucan taxonomy has not been completely resolved (see Barker and Lanyon, 2000; Moyle, 2004). The five families here are one possibility. The toucan-barbets might belong to either the toucans or New World barbets. The position of both Caloramphus and Trachyphonus among the other barbets has also not been completely resolved.

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 is remains Megalaimidae.

Lybiidae: African Barbets, Tinkerbirds Sibley & Ahlquist, 1985

9 genera, 42 species HBW-7 (split)

Semnornithidae: Toucan-barbets Prum, 1988

1 genus, 2 species HBW-7 (split)

Capitonidae: New World Barbets Bonaparte, 1838

2 genera, 14 species HBW-7 (split)

Ramphastidae: Toucans, Aracaris, Toucanets Vigors, 1825

5 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 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 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.

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

33 genera, 232 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), Fuchs et al. (2006c, 2007b, 2008a), Moore et al. (2006), Overton and Rhoads (2006), Webb and Moore (2005), and Weibel and Moore (2002a, 2002b) have all proven helpful.

This version is based on a closer look at the above and the summary of the Melanerpini by Raty. A certain amount of uncertainty remains, but I think this is a pretty good representation of the current state of woodpecker taxonomy. The diagram sums it up.

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 the Antillean Piculet does not belong with the other piculets, but is the sister of the Picinae. Accordingly, it is placed in the monotypic subfamily Nesoctitinae.

For the inner structure of the Picinae, we follow Fuchs et al. (2007b, 2008a). They found that Hemicircus is sister to the rest of the Picinae.

For now, Chrysocolaptini is treated separately from Picini due to its uncertain relation with Campephilini. Note that 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.

Picini tree Picini seems to look something like the tree to the right (Fuchs, 2008a). 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. The order of species in Picus is based on a combination of Fuchs et al. (2008a) and Pons et al. (2011) while the ordering of Colaptes is based on Moore et al. (2011). Moore et al. also provide evidence that the Bronze-winged Woodpecker, Colaptes aeruginosus, is sister to the Gray-crowned Woodpecker, Colaptes auricularis, rather than being a subspecies of the Golden-olive Woodpecker, Colaptes rubiginosus (itself sister to Black-necked Woodpecker, Colaptes atricollis).

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.

Although I've left the American Dryocopus alone, they may have to be put in a separate genus (Ceophloeus, I think) as the old world Dryocopus may be more closely related to Mulleripicus.

Following the recommendation of Benz and Robbins (2011), I've split Ochre-backed Woodpecker, Celeus ochraceus, from Blond-crested Woodpecker, Celeus flavescens.

The other piece of the Picinae includes Campephilus and the Melanerpini. Campephilus could be placed basally in Melanerpini. I've kept it separate partly because there is some remaining ambiguity about it's relationship with the Chrysocolaptini.

Melanerpini tree This brings us to the Melanerpini. Melanerpes, Sphyrapicus, and Xiphidiopicus group together (see Overton and Rhoads, 2006 for details). That clade is sister to the remaining Melanerpini. The various papers have to be pieced together to make a complete picture of the rest of Melanerpini. This introduces some uncertainty about exactly how they fit together. Picoides is split into Picoides, Dryobates, and Leuconotopicus; and Dendrocopos is split into Dendrocopos, Yungipicus, and Leiopicus. Hypopicus and Sapheopipo are merged into Dendrocopos (see Winkler et al., 2005) while Dendrocopos minor moves to Dryobates, Veniliornis fumigatus moves to Leuconotopicus, and Picoides mixtus and P. lignarius move to Veniliornis. With these changes, Melanerpini looks like the diagram.

A recent paper by García-Trejo et al. (2009) finds 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. Based on their work, I have also rearranged the carolinus group, which starts at M. pucherani. The results of Overton and Rhoads (2006) suggest that the Puerto Rican, Hispaniolan, and presumably Jamaican Woodpeckers belong near carolinus. The entire Red-bellied complex needs further study to before it is completely sorted out.

Jynginae: Wrynecks Swainson, 1831

Picumninae: Piculets G.R. Gray, 1840

Nesoctitinae: Antillean Piculet Wolters, 1976

Picinae: Woodpeckers Leach, 1820

Hemicircini Cabanis and Heine, 1863

Chrysocolaptini Bonaparte, 1854

Picini Leach, 1820

Campephilini Blyth, 1852

Melanerpini G.R. Gray, 1846

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