A classification of the bird species of South America

South American Classification Committee
American Ornithologists' Union
(Part 5)

 

Part 5. Trogoniformes to Piciformes (below)

Part 1. Struthioniformes to Ciconiiformes (click)
Part 2. Falconiformes to Charadriiformes (click)
Part 3. Columbiformes to Caprimulgiformes (click)
Part 4. Apodiformes (click)
Part 6. Suboscine Passeriformes, A (Eurylaimidae and Furnariidae) (click)
Part 7. Suboscine Passeriformes, B (Thamnophilidae to Rhinocryptidae) (click)
Part 8. Suboscine Passeriformes, C (Tyrannidae to Pipridae) (click)
Part 9. Oscine Passeriformes, A (Vireonidae to Sturnidae) (click)
Part 10. Oscine Passeriformes, B (Motacillidae to Emberizidae) (click)
Part 11. Oscine Passeriformes, C (Cardinalidae to end) (click)

Hypothetical List (click)
Hybrids and Dubious Taxa (click)
Literature Cited (click)


TROGONIFORMES 1

TROGONIDAE (TROGONS)
Pharomachrus pavoninus Pavonine Quetzal 16
Pharomachrus auriceps
Golden-headed Quetzal 16, 16b
Pharomachrus fulgidus
White-tipped Quetzal 17, 18
Pharomachrus antisianus
Crested Quetzal 17, 19
Trogon massena
Slaty-tailed Trogon 11, 12
Trogon comptus
Blue-tailed Trogon 11, 13
Trogon melanurus
Black-tailed Trogon 11
Trogon viridis White-tailed Trogon 1a, 2, 3, 4
Trogon violaceus
Violaceous Trogon 8, 8a, 8b
Trogon curucui
Blue-crowned Trogon 6, 7, 7a, 7b
Trogon surrucura
Surucua Trogon 5, 6
Trogon rufus
Black-throated Trogon 6, 7
Trogon collaris
Collared Trogon 6, 7, 9
Trogon personatus
Masked Trogon 6, 10, 10a




1. The monophyly of the Trogoniformes has never been questioned; its relationships to other birds, however, are uncertain. Traditional classifications have considered the Trogonidae to be more closely related to the Coraciiformes than to other orders, or to a group that includes Coraciiformes + Piciformes; see reviews in Sibley & Ahlquist (1990), Espinosa de los Monteros (2000), and Mayr (2003); some genetic data (Sibley & Ahlquist 1990) are consistent with this view. A recent analysis of morphological data (Mayr 2003b, Mayr & Clarke 2003) suggest that the Steatornithidae and the Trogoniformes and might be sister taxa. Some genetic (Espinosa de los Monteros 2000) data suggest a sister relationship with the Coliiformes (mousebirds), whereas other genetic data (Fain & Houde 2004) support a sister relationship with the Old World Bucerotidae. The most comprehensive genetic survey (Hackett et al. 2008) found strong support for their traditional position: they are members of a group ot orders that conists of the Coraciiformes, Piciformes, Bucerotiformes, and Upupiformes. Recent genetic data (Moyle 2005) suggest that the quetzals (Pharomachrus + Euptilotis) might be basal to all other trogons, including Old World genera.
SACC proposal passed to invert linear sequence of genera.
1a. Within the genus Trogon, genetic data (Moyle 2005, DaCosta and Klicka 2008) provide strong support for two major groups: (1) those with brown-backed females (collaris, personatus, and rufus, along with Middle American aurantiiventris, elegans, and mexicanus) and (2) those with gray-backed females (the rest). Within the latter group, two additional groups are strongly supported: (3) those with blue-headed males and strongly contrasting black-and-white tail patterns (viridis, surrucura, violaceus, and curucui, along with Middle American melanocephalus, citreolus, and bairdii) and (4) those with green-headed males and mostly dark, unpatterned tails (massena, comptus, melanurus, and Middle American clathratus).
SACC proposal passed to change linear sequence of species.
2. Ridgely & Greenfield (2001) considered the subspecies chionurus of the Chocó region to be a separate species from Trogon viridis; followed by Hilty (2003);
SACC proposal to recognize this split did not pass because of insufficient published data. Genetic data (DaCosta & Klicka 2008) suggest that chionurus is more closely related to T. bairdii than either are to Amzonian T. viridis. Proposal needed. Sibley & Monroe (1990) considered Trogon viridis to form a superspecies with Central American T. bairdii, and suggested that they might be conspecific.
3. Collar (2001) also included Middle American Trogon melanocephalus and T. citreolus in a superspecies with T. viridis and T. bairdii, and genetic data (DaCosta & Klicka 2008) indicate that these two form a sister group to T. viridis + T. bairdii.
4. Trogon viridis was formerly (e.g.,
Pinto 1937, Peters 1945) called Trogon strigilatus, but see Zimmer (1948).
5. The subspecies aurantius was formerly (e.g., Cory 1919,
Pinto 1937) treated as a separate species from Trogon surrucura; they were treated as conspecific by Peters (1945), and this treatment has been followed in subsequent classifications.
6. Trogon collaris, T. personatus, T. rufus, T. surrucura, and T. curucui were formerly (e.g., Cory 1919,
Pinto 1937) placed in a separate genus, Trogonurus, but this was merged into Trogon by Peters (1945). Genetic data (Moyle et al. 2005, DaCosta & Klicka 2008) indicate that "Trogonurus" is not a monophyletic group (see Note 1a).
7. Species names used in Cory (1919),
Pinto (1937), and other literature before Peters (1945) used incorrect names that were sorted out by Schneider (1938); the name curucui was applied to T. collaris and to T. rufus, whereas T. curucui was called T. variegatus.
7a. The western subspecies bolivianus was formerly
(e.g., Cory 1919) considered a separate species from Trogon curucui. Peters (1945) treated them as conspecific, and this treatment has been followed in subsequent classifications.
7b. "Trogon variegatus," known from throughout much of range of T. curucui and treated as a valid species by Cory (1919); was considered by Peters (1945) <
a color variant?> and a synonym of nominate curucui. See Hybrids and Dubious Taxa.
8. The subspecies ramonianus and caligatus were formerly (e.g., Cory 1919,
Pinto 1937) considered separate species from Trogon violaceus, but Peters (1945) considered them all conspecific. Ridgely & Greenfield (2001) considered caligatus of Middle America and northwestern South America to be a separate species from Trogon violaceus, and this was followed by Hilty (2003); SACC proposal to recognize this split did not pass because of insufficient published data. Genetic data (DaCosta & Klicka 2008) indicate that caligatus is basal to a group that includes Amazonian T. violaceus, T. curucui, and T. surrucura (and that Amazonian violaceus may be paraphyletic with respect to the latter two species). Proposal needed.
8a. Trogon violaceus was formerly (e.g., Cory 1919,
Pinto 1937) placed in a separate genus, Chrysotrogon, but this was merged into Trogon by Peters (1945). Genetic data (Moyle 2005, DaCosta & Klicka 2008) provide no support for recognition of this monotypic genus.
9. The subspecies puella of Middle American was formerly (e.g., Cory 1919) considered a separate species from Trogon collaris; they were considered conspecific by Peters (1945), and this treatment has been followed in subsequent classifications. Genetic data (DaCosta & Klicka 2008) indicate that puella is more closely related to Middle American T. aurantiiventris than either is to Amazonain T. collaris.
Proposal needed.
10. Ridgely & Greenfield (2001) suggested that the higher-elevation subspecies temperatus deserves recognition as a separate species from lower elevation Trogon personatus, as originally designated by Chapman (1923) and so treated by Meyer de Schauensee (1964); their voices also differ; their apparent elevational parapatry in Ecuador would be sufficient evidence for recognition as separate species, and so the details of this situation need to be examined and published. Zimmer (1948), however, considered specimens from northern Ecuador and southern Colombia to show signs of intergradation.
10a. The subspecies assimilis of the W. Andes was formerly (e.g., Cory 1919) considered a separate species from Trogon personatus; Peters (1945) treated them as conspecific, and this treatment has been followed in subsequent classifications..
11. Trogon massena, T. comptus, and T. melanurus form a closely related group, along with Central American T. clathratus (Collar 2001), but geographic overlap prevents considering them as a superspecies (Meyer de Schauensee 1966); see also Zimmer (1948) for discussion of overlap and confusing character distribution of these three in western Colombia; they were formerly (e.g., Cory 1919,
Pinto 1937) placed in a separate genus, Curucujus, but this was merged into Trogon by Peters (1945). Genetic data (Moyle 2005, DaCosta & Klicka 2008) provide strong support for recognition of this group as monophyletic, but to treat it as a separate genus would require recognition of at least one additional genus with broadly defined Trogon.
12. <?Hellmayr 1929> considered the South American subspecies australis as a separate species from Middle American Trogon massena; Zimmer (1948) suspected that australis might actually be a subspecies of T. melanurus. The subspecies macroura of northwestern Colombia and Panama was formerly (e.g., REF<?Hellmayr 1929>) considered a species separate from Trogon melanurus, and it may deserve recognition as a separate species (Zimmer 1948). Ridgely & Greenfield (2001) considered mesurus of western Ecuador and northwestern Peru to be a separate species from T. melanurus;
SACC proposal to recognize this split did not pass because of insufficient published data. Genetic data (DaCosta & Klicka 2008) indicate that melanurus may be paraphyletic with respect to T. massena and T. comptus. Proposal needed.
13. Called "White-eyed Trogon" in Sibley & Monroe (1990) and "Chocó Trogon" in Ridgely & Greenfield (2001).
SACC proposal to change English name did not pass.
16. Sibley & Monroe (1990) and Collar (2001) considered Pharomachrus auriceps and P. pavoninus to form a superspecies, and they were formerly (e.g., Peters 1945, Zimmer 1948) considered conspecific. The subspecies hargitti of the Venezuelan Andes has been occasionally treated as a subspecies of (e.g., REF), or synonym of (e.g, Peters 1945), P. pavoninus instead of P. auriceps. The subspecies P. a. heliactin of western Ecuador has been considered (e.g., Peters 1945) a subspecies of P. pavoninus when auriceps treated as a subspecies of P. pavoninus; Fjeldså & Krabbe (1990) stated that heliactin may be a a separate species; Zimmer (1948) and Collar (2001), however, considered heliactin indistinguishable from P. a. auriceps.
16b. "Pharomachrus xanthogaster," known only from the type specimen from "Bogotá," was treated as a valid species by Cory (1919), but he suspected that it was a color variant of P. auriceps; Peters (1945) considered it a synonym of auriceps. See Hybrids and Dubious Taxa.
17. Pharomachrus fulgidus and P. antisianus may form a superspecies (REF); (REFS) considered them conspecific. Other authors suspect that P. fulgidus may be part of the P. auriceps-P. pavoninus superspecies (Collar 2001). <
incorp. Berlioz 1956>
18. The subspecies festatus of the Santa Marta Mountains was formerly
(e.g., Cory 1918) considered a separate species from Pharomachrus fulgidus, but Peters (1945) treated them as conspecific.
19. Sibley & Monroe (1990) and Collar (2001) considered Pharomachrus antisianus to form a superspecies with Middle American P. mocinno, but did not include fulgidus. Peters (1945), Zimmer (1948), and Phelps & Phelps (1958a) considered antisianus to be conspecific with Middle American P. mocinno.


CORACIIFORMES 1

ALCEDINIDAE (KINGFISHERS)
Megaceryle torquata Ringed Kingfisher 2, 3, 4, 5
Megaceryle alcyon
Belted Kingfisher (NB) 2, 3
Chloroceryle amazona
Amazon Kingfisher 6
Chloroceryle americana
Green Kingfisher 6
Chloroceryle inda
Green-and-rufous Kingfisher 6
Chloroceryle aenea
American Pygmy Kingfisher 6, 7


1. The monophyly of the Coraciiformes is controversial (see reviews in Sibley & Ahlquist 1990, Johansson et al. 2001).
The most comprehensive genetic survey (Hackett et al. 2008) found strong support for the monophyly of the traditional Coraciiformes only if limited to the families Meropidae, Coraciidae, Brachypteraciidae, Todidae, Momotidae, and Alcedinidae (and thus excluding hornbills, hoopoes, and Leptosomus). Sibley-Ahlquist (1990) divided the Alcedinidae into three families, which consist of the three traditional subfamilies elevated to family rank (Alcedinidae, Dacelonidae, and Cerylidae) because of DNA-DNA hybridization data indicated deep divergences among these three; New World kingfishers were placed in the Cerylidae.
2. Megaceryle torquata and M. alcyon have been placed in the genus Ceryle in many classifications (e.g., Meyer de Schauensee 1970, AOU 1983, 1998), but most classifications have followed Fry (1980) in restricting Ceryle to Old World C. rudis (e.g., Sibley & Monroe 1990, Fry & Fry 1992, Woodall 2001). Recent genetic data (Moyle 2006) indicate that Old World Ceryle rudis is the sister to Chloroceryle, and so Megaceryle must be recognized if Chloroceryle is maintained as a genus.
3. Megaceryle torquata and M. alcyon were considered sister species by Fry (1980) in a superspecies complex that included Old World M. maxima and M. lugubris. Moyle's (2006) data are consistent with their status as sister species.
4. The name Streptoceryle was formerly
(e.g., Ridgway 1914, Cory 1919) used in place of Megaceryle, but see (REF).
5. Ceryle is masculine, so the correct spelling of the species name is torquatus (David & Gosselin 2002b) when that genus is used; Megaceryle, however, is feminine, so the species name remains torquata when that genus is used (David & Gosselin 2002b).
6. Fry (1980) and Fry & Fry (1992) proposed that plumage similarities indicate that Chloroceryle amazona and C. americana are sister species, as are C. inda and C. aenea. Genetic data (Moyle 2006), however, indicate that C. americana and C. inda are sisters, that C. aenea is sister to this pair, and that C. amazona is basal in the genus.
7. Chloroceryle aenea was formerly known as "Pygmy Kingfisher", but most sources (e.g., AOU 1983, 1998, Stiles & Skutch 1989, Sibley & Monroe 1990, Fry et al. 1992, Ridgely & Greenfield 2001, Woodall 2001, Hilty 2003) now call this "American Pygmy Kingfisher" to avoid confusion with African taxa Ceyx pictus ("African Pygmy Kingfisher") and C. madagascariensis ("Madagascar Pygmy Kingfisher").



MOMOTIDAE (MOTMOTS) 1
Hylomanes momotula
Tody Motmot
Electron platyrhynchum Broad-billed Motmot 2
Baryphthengus martii
Rufous Motmot 3
Baryphthengus ruficapillus
Rufous-capped Motmot 3
Momotus momota
Blue-crowned Motmot 4



1. The monophyly of the Momotidae has never been seriously questioned. Several data sets (e.g., Olson 1976, Mayr 1998, Espinosa de los Monteros 2000, Johansson et al. 2001, Overton & Rhoads 2004) indicate that the Momotidae and the West Indian Todidae are sister families, but Ericson et al. (2004) and Hackett et al. (2008) supported a sister realtionship between te Alcedinidae and Momotidae . Although generic limits in the family have remained fairly constant (see Snow 2001), the relationships of genera within the Momotidae have not been subjected to any modern analyses. <
incorp Maurer & Raikow 1981>
2. Because they lack racket tips on their tails, Meyer de Schauensee (1966) suggested that the pyrrholaemum subspecies group east of the Andes might deserve treatments as a separate species from Electron platyrhynchum .
3. Baryphthengus martii,
formerly considered a subspecies of B. ruficapilla (e.g., Peters 1945, Meyer de Schauensee 1970), is now generally considered a separate species, following Sick (1993); however, no formal analysis has ever been published, although at one time (e.g., Ridgway 1914, Cory 1918) they were considered not only separate species but martii was placed in a separate genus, Urospatha; they form a superspecies (Sibley & Monroe 1990).
4. Momotus momota may consist of several species-level taxa (e.g., Ridgely & Greenfield 2001). The subspecies aequatorialis, venezuelae, subrufescens, microstephanus, and argenticinctus were all formerly
(e.g., Ridgway 1914, Cory 1919) considered separate species from M. momota, as were two Middle American taxa. Chapman (1923) recognized four species in South America: M. subrufescens (including "venezuelae") of the Caribbean rim of northern South America, M. bahamensis of Trinidad, M. aequatorialis of the Andes, and M. momota (including microstephanus) of the rest of South America, including argenticinctus of western Ecuador and northwestern Peru. Peters (1945) considered them all conspecific, and this was followed by Meyer de Schauensee (1970) and AOU (1983, 1998). Fjeldså & Krabbe (1990) proposed that the Andean form aequatorialis was a separate species from M. momota, and this was followed by Ridgely & Greenfield (2001), Dickinson (2003), and Schulenberg et al. (2007), thus returning to the classification of Cory (1919) and Chapman (1923, 1926). However, no formal analysis has ever been published, and the published evidence in support of treating aequatorialis as a species-level taxon is weak. SACC proposal passed for treating aequatorialis as conspecific with M. momota (and we hope that this decision stimulates further research on the M. momota complex).



GALBULIFORMES 1

GALBULIDAE (JACAMARS) 1a
Galbalcyrhynchus leucotis
White-eared Jacamar 2
Galbalcyrhynchus purusianus
Purus Jacamar 2
Brachygalba albogularis
White-throated Jacamar 3
Brachygalba lugubris
Brown Jacamar 3, 4, 4a
Brachygalba goeringi
Pale-headed Jacamar 3
Brachygalba salmoni
Dusky-backed Jacamar 3
Jacamaralcyon tridactyla
Three-toed Jacamar 5
Galbula albirostris
Yellow-billed Jacamar 6, 7
Galbula cyanicollis
Blue-cheeked Jacamar 6, 8
Galbula ruficauda
Rufous-tailed Jacamar 9, 10
Galbula galbula
Green-tailed Jacamar 9
Galbula tombacea
White-chinned Jacamar 9
Galbula cyanescens
Bluish-fronted Jacamar 9
Galbula pastazae
Coppery-chested Jacamar 9
Galbula chalcothorax
Purplish Jacamar 11
Galbula leucogastra
Bronzy Jacamar 11
Galbula dea
Paradise Jacamar 11a
Jacamerops aureus
Great Jacamar 12


1. Evidence from genetics (Sibley & Ahlquist 1990, Johansson et al. 2001, Johansson & Ericson 2003, Cracraft et al. 2004, Ericson et al. 2006, Hackett et al. 2008) and morphology (e.g., Sibley 1956, Simpson and Cracraft 1981, Swierczewski and Raikow 1981) strongly indicate that the Galbulidae and the Bucconidae are sister taxa, a relationship identified over 250 years ago. The monophyly of each has never been seriously questioned (see reviews in Sibley & Ahlquist 1990, Rasmussen & Collar 2002, Tobias 2002). They are usually considered to be a suborder, Galbulae, of the Piciformes, but some evidence (Sibley and Ahlquist 1972, 1985, 1986, Olson 1983, 1985, Burton 1984, Mayr 1998, Höfling & Alvarenga 2001) indicates that they might be more closely related to the Coraciiformes. The original the genetic evidence for this relationship (Sibley & Ahlquist 1990) is actually ambiguous (Harshman 1994). The Galbulae are here treated as a separate order, following AOU (1998), until their relationships are resolved. Recent genetic evidence (Johansson & Ericson 2003, Ericson et al. 2006) supports the traditional placement of the Galbulidae and Bucconidae in the Piciformes, whereas other recent genetic data fail to support this (Fain & Houde 2004).
1a. Within-family relationships in the Galbulidae have not been subjected to any modern analyses; see Tobias et al. (2002) for a summary of literature that supports the traditional linear sequence of genera used here.
2. Galbalcyrhynchus leucotis and G. purusianus were formerly (e.g., Cory 1919,
Pinto 1937) treated as separate species, but Peters (1948) and Meyer de Schauensee (1970) considered them conspecific ("Chestnut Jacamar"). Haffer (1974) noted that they are parapatric in the Río Ucayali area with no sign of interbreeding and that they differ in plumage to the same degree as other jacamars currently ranked as species; they constitute a superspecies (Haffer 1974, Sibley & Monroe 1990, Tobias et al. 2002).
3. The four Brachygalba species form a superspecies (Haffer 1967, 1974, Sibley & Monroe 1990, Tobias et al. 2002).
4. The subspecies phaeonota and melanosterna (with naumburgi) were formerly (e.g., Todd 1943, Peters 1948) each considered separate species from Brachygalba lugubris, but they were all treated as conspecific by Meyer de Schauensee (1966). The subspecies fulviventris (with caquetae) and melanosterna were also formerly (e.g., Cory 1919) each considered separate species from B. lugubris, but they were all treated as conspecific by Peters (1948).
5. Jacamaralcyon and Brachygalba are presumably sister genera (Haffer 1974).
6. Galbula albirostris and G. cyanicollis were formerly considered conspecific (e.g., Peters 1948, Meyer de Schauensee 1970), but Haffer (1974) noted that they are parapatric in eastern Peru area with no sign of interbreeding; they constitute a superspecies (Haffer 1974, Tobias et al. 2002); they had formerly (e.g., Cory 1919,
Pinto 1937) been considered separate species, and in fact, albirostris was formerly (e.g., Cory 1919) placed in a separate, monotypic genus, Psilopornis (which was merged into Galbula by Pinto 1937 and Peters 1948).
7. The subspecies chalcocephala may represent separate species from Galbula albirostris (Tobias et al. 2002); it shows no signs of intergradation with nominate albirostris where their ranges approach (Haffer 1974).
8. Called "Blue-necked Jacamar" by Sibley & Monroe (1990) and Tobias et al. (2002).
proposal needed?
9. Galbula ruficauda, G. galbula, G. tombacea, G. cyanescens, and G. pastazae are considered to form a superspecies (Haffer 1974, Sibley & Monroe 1990, Tobias et al. 2002); evidence for ranking them at species level rather weak (except perhaps for pastazae), but there is no sign of hybridization among them where their ranges are in contact.
10. The subspecies rufoviridis (with heterogyna) is geographically separated from northern subspecies by Galbula galbula and other members of the superspecies; thus, whether G. ruficauda, as presently constituted, is monophyletic warrants study; rufoviridis was formerly (e.g., Cory 1919,
Pinto 1937) considered a separate species. Trans-Andean melanogenia was also formerly (e.g., Ridgway 1914, Cory 1919, Haffer 1967) considered a separate species, but apparent intergradation with nominate ruficauda (Wetmore 1968) has led to them being considered conspecific (e.g., Peters 1948, Meyer de Schauensee 1970, Tobias et al. 2002).
10a. Galbula cyanescens has been considered a subspecies of G. tombacea by some authors (e.g., REFS?,
Pinto 1937), but <>.
11. Galbula chalcothorax and G. leucogastra were formerly considered conspecific (e.g., Cory 1919, Peters 1948, Meyer de Schauensee 1970, Haffer 1974), but there is no indication of hybridization between the two, and they differ as much or more in plumage than most parapatric jacamars ranked at the species level (Parker & Remsen 1987); they constitute a superspecies (Sibley & Monroe 1990, Tobias et al. 2002).
11a. Galbula dea was formerly (e.g., Cory 1919,
Pinto 1937) placed in the monotypic genus Urogalba.
12. Jacamerops is masculine, so the correct spelling of the species name is aureus (David & Gosselin 2002b).


BUCCONIDAE (PUFFBIRDS) 1
Notharchus hyperrhynchus
White-necked Puffbird 1a, 1b
Notharchus macrorhynchos
Guianan Puffbird 1b
Notharchus swainsoni
Buff-bellied Puffbird 1b
Notharchus pectoralis
Black-breasted Puffbird 2
Notharchus ordii
Brown-banded Puffbird 2
Notharchus tectus
Pied Puffbird 3
Bucco macrodactylus
Chestnut-capped Puffbird 4
Bucco tamatia
Spotted Puffbird 5, 5a
Bucco noanamae
Sooty-capped Puffbird 5, 5a
Bucco capensis
Collared Puffbird
Nystalus radiatus Barred Puffbird 6, 6a
Nystalus striolatus
Striolated Puffbird 6, 6a
Nystalus chacuru
White-eared Puffbird 6a
Nystalus maculatus
Spot-backed Puffbird 7, 7a
Hypnelus ruficollis
Russet-throated Puffbird 8, 9
Malacoptila fusca
White-chested Puffbird 10
Malacoptila semicincta
Semicollared Puffbird 10
Malacoptila striata
Crescent-chested Puffbird 10, 10a
Malacoptila rufa
Rufous-necked Puffbird
Malacoptila panamensis White-whiskered Puffbird 11
Malacoptila fulvogularis
Black-streaked Puffbird 11, 11a
Malacoptila mystacalis
Moustached Puffbird 11
Micromonacha lanceolata
Lanceolated Monklet
Nonnula rubecula Rusty-breasted Nunlet 12
Nonnula sclateri
Fulvous-chinned Nunlet 13
Nonnula brunnea
Brown Nunlet 13
Nonnula frontalis
Gray-cheeked Nunlet 14
Nonnula ruficapilla
Rufous-capped Nunlet 14
Nonnula amaurocephala
Chestnut-headed Nunlet 15
Hapaloptila castanea
White-faced Nunbird 15a
Monasa atra
Black Nunbird 16
Monasa nigrifrons
Black-fronted Nunbird 16
Monasa morphoeus
White-fronted Nunbird 16, 16a
Monasa flavirostris
Yellow-billed Nunbird
Chelidoptera tenebrosa Swallow-winged Puffbird 17




1. The monophyly of the Bucconidae has never been seriously questioned. Within-family relationships in the Bucconidae have not been subjected to any modern analyses; see Rasmussen & Collar (2002) for a summary of literature that supports the traditional linear sequence of genera used here.
1a. Notharchus was merged into Bucco by Cottrell (1968), and this was followed by the AOU (1983), but not by other classifications; see Monroe et al. (1993).
1b. The taxon swainsoni of the Atlantic forest region was formerly (e.g., Cory 1919,
Pinto 1937) considered a separate species from Notharchus macrorhynchos, but Peters (1948) treated them as conspecific; this was followed by most subsequent classifications. Rasmussen & Collar (2002) elevated swainsoni to species rank, and Alvarenga et al. (2002) provided rationale in support of that treatment. SACC proposal passed to elevate swainsoni to species rank. The hyperrhynchus subspecies group was also formerly (e.g., Ridgway 1914, Cory 1919, Pinto 1937) considered a separate species, but it was also treated as conspecific with N. macrorhynchos by Peters (1948). Rasmussen & Collar (2002) also suggested that the hyperrhynchus group might also warrant species rank. SACC proposal passed to elevate hyperrhynchus to species rank; SACC proposals passed to apply English name "Guianan Puffbird" to narrowly distributed macrorhynchos and to retain "White-necked" for widely distributed hyperrhynchus.
2. Some authors (e.g., REFS, Rasmussen & Collar 2002) consider Notharchus pectoralis and N. ordii to form a superspecies.
3. Trans-Andean subspecies subtectus was formerly (e.g., REFS) considered a separate species from Notharchus tectus. <
delete if REF can't be found>
4. Bucco macrodactylus was formerly (e.g.,
Ridgway 1914, Cory 1919, Pinto 1937) placed in the monotypic genus Argicus, but this was merged into Bucco by Peters (1948), and this has been followed by most subsequent classifications, except for Rasmussen & Collar (2002), who resurrected Argicus.
5. Bucco tamatia and B. noanamae were formerly (e.g.,
Ridgway 1914, Cory 1919, Pinto 1937) placed in the genus Nystactes, but this was merged into Bucco by Peters (1948), and this has been followed by most subsequent classifications, except for Rasmussen & Collar (2002), who resurrected Nystactes.
5a. Bucco tamatia and B. noanamae form a superspecies (REFS, Rasmussen & Collar (2002).
6. Nystalus radiatus and N. striolatus may form superspecies (Rasmussen & Collar (2002).
6a. Nystalus radiatus, N. striolatus, and N. chacuru were formerly
(e.g., Ridgway 1914, Cory 1919) treated in a separate genus, Ecchaunornis, but Peters (1948) merged this into Nystalus.
7. Silva (1991) considered striatipectus a separate species from Nystalus maculatus; not followed by Rasmussen & Collar (2002).
Proposal needed.
7a. Called "Spot-bellied Puffbird" in Rasmussen & Collar (2002).
8. Although Cory (1919) and Peters (1948) considered the subspecies bicinctus to be a separate species from Hypnelus ruficollis, Meyer de Schauensee (1966, 1970) considered them conspecific, evidently <
check> on the basis that the subspecies described by Phelps & Phelps (1958) was intermediate between the two, and because another population was also considered intermediate. Rasmussen & Collar (2002) considered bicinctus (with stoicus) as a separate species from H. ruficollis; they reported that hybridization in area of contact was infrequent and certainly not indicative of free interbreeding, as is often stated or implied (e.g., Sibley & Monroe 1990). Proposal needed.
9. Hypnelus was merged into Bucco by Cottrell (1968), but this has not been followed by most subsequent authors.
10. Malacoptila fusca and M. semicincta form a superspecies (Haffer 1987, Rasmussen & Collar 2002); they were once considered conspecific (e.g., Peters 1948), but see Traylor (1951, 1956<?>). Sibley & Monroe (1990) also included M. striata in the superspecies.
10a. Malacoptila striata
was formerly (e.g., Cory 1919) known as M. torquata, but see Peters (1948).
11. Sibley & Monroe (1990) considered Malacoptila panamensis and M. mystacalis to form a superspecies; Rasmussen & Collar (2002) also included M. fulvogularis.
11a. The Colombian subspecies substriata was formerly
(e.g., Cory 1919) considered a separate species from Malacoptila fulvogularis, but Peters (1948) treated them as conspecific; Rasmussen & Collar (2002) treated substriata as a synonym of fulvogularis, regarded by them as a monotypic species.
12. Nonnula rubecula may consist of
more than one species (Rasmussen & Collar 2002).
13. Nonnula sclateri and N. brunnea form a superspecies (REFS); they have been considered conspecific by some authors (e.g., REFS). Also, N. rubecula is sometimes (e.g., (REFs, Rasmussen & Collar 2002) included in this superspecies, but it may be sympatric with N. brunnea w. Amazonia.
14. Many authors (e.g., Meyer de Schauensee 1970) have treated Nonnula frontalis as subspecies of N. ruficapilla, following Meyer de Schauensee (1946b); published evidence for considering them separate species is weak; Sibley & Monroe (1990) and Rasmussen & Collar (2002) considered them to form a superspecies.
15. Nonnula amaurocephala was considered to form a superspecies with N. frontalis and N. ruficapilla by REFS, Rasmussen & Collar (2002).
15a. Called "White-faced Puffbird" in Fjeldså & Krabbe (1990).
16. Rasmussen & Collar (2002) considered Monasa morphoeus and M. nigrifrons to form a superspecies with M. atra, but the first two are broadly sympatric and cannot be considered allospecies.
16a. The subspecies grandior, fidelis, similis, pallescens (with sclateri and minor), and rikeri were formerly
(e.g., Ridgway 1914, Cory 1919) each considered separate species from Monasa morphoeus, but Peters (1948) treated them all as conspecific (and similis as a synonym of fidelis, and rikeri as a synonym of nominate morphoeus).
17. Called "Swallow-wing" by Meyer de Schauensee (1970), Snyder (1966), Haverschmidt (1968), Meyer de Schauensee & Phelps (1978), Sibley & Monroe (1990), Haverschmidt & Mees (1994), and elsewhere; Hilty & Brown (1986) evidently were the first to use "Swallow-winged Puffbird," and this has been followed by Ridgely & Greenfield (2001), Rasmussen & Collar (2002), Hilty (2003).
Proposal?



PICIFORMES 1

CAPITONIDAE (NEW WORLD BARBETS) 2
Capito aurovirens Scarlet-crowned Barbet
Capito dayi Black-girdled Barbet 3
Capito maculicoronatus
Spot-crowned Barbet 4
Capito squamatus
Orange-fronted Barbet 4
Capito hypoleucus
White-mantled Barbet 4a
Capito wallacei
Scarlet-banded Barbet 5
Capito quinticolor
Five-colored Barbet 4a
Capito brunneipectus
Brown-chested Barbet 6, 7
Capito niger
Black-spotted Barbet 6
Capito auratus
Gilded Barbet 6, 6b
Eubucco richardsoni
Lemon-throated Barbet 8, 8a
Eubucco tucinkae
Scarlet-hooded Barbet 9
Eubucco bourcierii
Red-headed Barbet 10
Eubucco versicolor
Versicolored Barbet 8a, 10, 11

SEMNORNITHIDAE (TOUCAN-BARBETS) 2
Semnornis ramphastinus Toucan Barbet


1. Genetic data (Sibley and Ahlquist 1985, 1986, 1990, Johansson et al. 2001, Prychitko & Moore 2003, Fain & Houde 2004, Hackett et al. 2008) support traditional morphological data (e.g., Simpson and Cracraft 1981, Swierczewski and Raikow 1981) that the Piciformes, as constituted here, are a monophyletic group. Most classifications also include jacamars and puffbirds in this order (e.g., Ridgway 1914, Wetmore 1960), but see notes under Galbuliformes.
2. The families Capitonidae, Semnornithidae, and Ramphastidae are each other's closest relatives with respect to Old World barbets (Burton 1984, Prum 1988, Sibley and Ahlquist 1990, Lanyon & Hall 1994, Barker & Lanyon 2000, Johansson et al. 2001, Johannson & Ericson 2003, Moyle 2004). [Old Word barbets are here tangentially but implicitly treated as separate families, Asian Megalaimidae and African Lybiidae; recent genetic data (Moyle 2004) support the monophyly of the barbet radiations within each region.] To emphasize the close relationships among New World taxa, these three families were treated as subfamilies of a single family, Ramphastidae, by AOU (1998).
SACC proposal passed to treat these taxa at family rank. Semnornis is treated as separate family until affinities resolved. Swierczewski and Raikow's (1981) analysis of characters of the hindlimb musculature supported the traditional inclusion (e.g., Meyer de Schauensee 1970) of Semnornis in the barbets, but Prum's (1988a) analysis of morphological data indicated that Semnornis is the sister taxon to the Ramphastidae, not the Capitonidae. Genetic data indicate that Semnornis may be basal to both families (Barker and Lanyon 2000); Moyle (2004) found weak support for that relationship, but also weak support for a sister relationship to Ramphastidae.
3. Capito dayi was considered conspecific with a broadly defined C. niger by Ripley (1945), but see Ripley (1946) and Haffer (1997). Genetic data (Armenta et al. 2005) indicate that C. dayi is definitely not part of the C. niger group, but more closely related to other species of Capito, especially C. quinticolor.
4. Sibley & Monroe (1990) considered Capito maculicoronatus and C. squamatus to form a superspecies; they were considered conspecific by Ripley (1945). Genetic data (Armenta et al. 2005) indicate that they are sister taxa.
4a. Capito hypoleucos was considered conspecific with C. quinticolor by Ripley (1945).
5. Recently described: O'Neill et al. (2000).
6. Capito brunneipectus and C. auratus were formerly (e.g., Peters 1948, Meyer de Schauensee 1970) considered conspecific with C. niger. Haffer (1997) split brunneipectus and auratus from niger because: (1) no good evidence was ever presented for the treatment of the three as conspecific; (2) no evidence exists for gene flow between niger and auratus where they are in contact; and (3) brunneipectus differs dramatically in coloration from auratus and niger. Genetic data (Armenta et al. 2005) support this treatment. Sibley & Monroe (1990), Short & Horne (2001), and Short & Horne (2002a) treated brunneipectus as separate species but not auratus. Capito niger, C. auratus, and C. brunneipectus presumably form a superspecies (Haffer 1997), but Short & Horne (2001) were not confident that brunneipectus belongs in that group; C. niger and C. brunneipectus are monotypic, with all subspecies-level taxa in the group included under C. auratus. Ridgway (1914), Cory (1919),
Pinto (1937), and Chapman (1928) treated auratus as a separate species, but Bond & Meyer de Schauensee (1943), Ripley (1945), and Peters (1948) considered them conspecific.
6b. "Capito aurantiiventris," known from the "Upper Amazon Valley" and formerly (e.g., Cory 1919) treated as a species, was subsequently (e.g., Peters 1948) treated as a synonym of C. auratus amazonicus. "Capito peruvianus," known from eastern Peru and Ecuador, and formerly (e.g., Cory 1919) treated as a species, was subsequently treated as a synonym of C. auratus auratus (<> Chapman 1928).
7. Called "Cinnamon-breasted Barbet" in Sibley & Monroe (1990), Short & Horne (2001), and Short & Horne (2002a).
8. The subspecies aurantiicollis was formerly
(e.g., Ridgway 1914, Cory 1919, Pinto 1937) considered a separate species from Eubucco richardsoni, but Berlioz (1938), Ripley (1945), and Peters (1948) treated them as conspecific.
8a. Eubucco richardsoni and E. versicolor were considered conspecific by Ripley (1945), but see, for example, Traylor (1951b).
9. Eubucco tucinkae was formerly considered to form superspecies with (Sibley & Monroe 1990), the sister taxon of (Haffer 1987), or even conspecific with (e.g., Peters 1948), E. bourcierii, but lowland distribution and habitat makes it unlikely that they are even sister species; Short & Horne (2001) also made this point, but based it on morphology.; see also Traylor (1951b).
10. Eubucco bourcierii and E. versicolor were considered to form a probable superspecies by Parker et al. (1985); cf. Short & Horne (2002a).
11. The subspecies steerii and glaucogularis were formerly
(e.g., Ridgway 1914, Cory 1919) each considered a separate species from Eubucco versicolor, but Berlioz (1938), Ripley (1945), and Peters (1948) treated them all as conspecific.


RAMPHASTIDAE (TOUCANS) 1
Ramphastos toco Toco Toucan
Ramphastos ambiguus Black-mandibled Toucan 23, 24, 25
Ramphastos tucanus
White-throated Toucan 22, 22a, 22b, 23, 23b
Ramphastos sulfuratus
Keel-billed Toucan 18, 21
Ramphastos brevis
Choco Toucan 18, 20
Ramphastos vitellinus
Channel-billed Toucan 18, 19, 19a
Ramphastos dicolorus
Red-breasted Toucan 18, 18a, 18b
Aulacorhynchus prasinus
Emerald Toucanet 1a
Aulacorhynchus sulcatus
Groove-billed Toucanet 2, 3
Aulacorhynchus derbianus
Chestnut-tipped Toucanet 3, 3a
Aulacorhynchus haematopygus
Crimson-rumped Toucanet 4
Aulacorhynchus huallagae
Yellow-browed Toucanet 4
Aulacorhynchus coeruleicinctis
Blue-banded Toucanet 4, 5
Andigena hypoglauca
Gray-breasted Mountain-Toucan 17
Andigena laminirostris
Plate-billed Mountain-Toucan 17
Andigena cucullata
Hooded Mountain-Toucan 17
Andigena nigrirostris
Black-billed Mountain-Toucan
Selenidera spectabilis Yellow-eared Toucanet 14
Selenidera culik
Guianan Toucanet 14, 14a
Selenidera reinwardtii
Golden-collared Toucanet 14, 15
Selenidera nattereri
Tawny-tufted Toucanet 14
Selenidera gouldii
Gould's Toucanet 14, 16
Selenidera maculirostris
Spot-billed Toucanet 14
Pteroglossus viridis
Green Aracari 6
Pteroglossus inscriptus
Lettered Aracari 6, 7, 10b
Pteroglossus bitorquatus
Red-necked Aracari 8
Pteroglossus azara
Ivory-billed Aracari 8, 9, 9a, 10, 10b
Pteroglossus aracari
Black-necked Aracari 10, 10c
Pteroglossus castanotis
Chestnut-eared Aracari 10
Pteroglossus pluricinctus
Many-banded Aracari 10a
Pteroglossus torquatus
Collared Aracari 10a, 11, 11a
Pteroglossus beauharnaesii
Curl-crested Aracari 12
Pteroglossus bailloni
Saffron Toucanet 13



1. Multiple independent data sets identify the Capitonidae/Semnornithidae as the sister to the Ramphastidae (see notes under those families above). The Ramphastidae have a number of unusual characters that distinguish them from all barbets, New World and Old World, including a unique arrangement of the caudal vertebrae and sleeping posture (see Short & Horne 2001) and a unique cranial morphology (Höfling 1991, 1998); the genus Aulacorhynchus shares these characters and is firmly embedded in the Ramphastidae, despite Sibley & Ahlquist's (1990) suggestion that it was intermediate in some respects between toucans and Capitonidae. Genetic data are consistent with the monophyly of the Ramphastidae (Moyle 2004). Genetic data (Barker & Lanyon 2000, Moyle 2004) indicate that Ramphastos is basal to all other toucan genera, and other genetic data sets are consistent with this (Sibley & Ahlquist 1990, Nahum et al. 2003). Moyle (2004) and Weckstein (2004) found that Andigena and Selenidera were sister genera, and that Aulacorhynchus was the sister of Andigena + Selenidera.
SACC proposal passed to change linear sequence of genera to the one used in this classification. Genetic data (Hackett & Lehn 1997, Barker & Lanyon 2000, Moyle 2004, Weckstein 2004) support for the traditional close relationship of Baillonius and Pteroglossus.
1a. Ridgely & Greenfield (2001) suggested that Aulacorhynchus prasinus may consist of more than one species-level taxon, but see Short & Horne (2001), who pointed out that the allopatric taxa are no more distinctive than those known to intergrade. The subspecies lautus, albivitta, cyanolaemus, dimidiatus, and atrogularis, as well as Middle American wagleri and caeruleogularis, were formerly
(e.g., Ridgway 1914, Cory 1919) each considered separate species from (and in some cases not particularly closely related to) Aulacorhynchus prasinus, but Peters (1948) and Haffer (1974) treated them all as conspecific. <add synopsis of Navarro et al. (2001)>. Puebla-Olivares et al. (2008) identified three clades in South America based on mtDNA and proposed species rank for each. Proposal badly needed.
2. The taxon calorhynchus was formerly (e.g., Cory 1919, Peters 1948, Phelps & Phelps 1958a, Meyer de Schauensee 1970) treated as a separate species ("Yellow-billed Toucanet") from Aulacorhynchus sulcatus, but in their area of contact in Venezuela, only individuals with intermediate bill characters are found (Schwartz 1972b); still treated as separate species by Hilty (2003).
3. Aulacorhynchus sulcatus and A. derbianus form a superspecies (Haffer1974, Short & Horne 2001, Short & Horne 2002b); they are treated as separate species because of differences in voice and bill shape.
3a. The whitelianus subspecies group of the Tepui region was formerly
(e.g., Cory 1919) considered a separate species from Aulacorhynchus derbianus, but they were treated as conspecific by Peters (1948).
4. Aulacorhynchus haematopygus, A. huallagae, and A. coeruleicinctis form a superspecies (Haffer1974, Fjeldså & Krabbe 1990, Short & Horne 2001, Short & Horne 2002b).
5. Species name often given incorrectly as "coeruleicinctus."
6. Pteroglossus viridis and P. inscriptus form a superspecies (Haffer 1974, Sibley & Monroe 1990, Short & Horne 2001). <
incorp. Cracraft & Prum 1988>
7. The subspecies humboldti was formerly (e.g., Peters 1948) considered a subspecies of Pteroglossus viridis, but Haffer (1974) included it in P. inscriptus; intermediate specimens from their area of contact led Haffer (1974) to treat humboldti as conspecific with and a subspecies of P. inscriptus. Cory (1919) treated humboldti as a separate species from P. viridis and P. inscriptus. "Pteroglossus didymus," known from eastern Peru and treated as a valid species by Cory (1919), is now considered a synonym of Pteroglossus inscriptus humboldti (Traylor 1958, Friedmann 1958, Borrero 1959, Haffer 1974, Short & Horne 2002b). See Hybrids and Dubious Taxa.
8. Pteroglossus bitorquatus and P. azara are considered to form a superspecies by Haffer (1974), but Short & Horne (2002b) considered P. bitorquatus to be probably more closely related to P. viridis/P. inscriptus. <
incorp. Cracraft & Prum 1988>
9. Haffer (1974) showed that Pteroglossus mariae, formerly (e.g., Peters 1948, Meyer de Schauensee 1970) considered a separate species ("Brown-mandibled Aracari"), forms hybrid zones with subspecies flavirostris wherever they are in contact and is thus best treated as a subspecies, as suspected by Peters (1948) and treated by Cory (1919) and
Pinto (1937); this treatment has been followed by most subsequent authors except Ridgely & Greenfield (2001); previous reports of sympatry (Todd 1943, Meyer de Schauensee 1966) are now considered erroneous (Haffer 1974). Analysis of plumage characters () supports their treatment as sister taxa, but preliminary genetic data (Hackett & Lehn 1997) does not.
9a. Sibley & Monroe (1990) pointed out that Pteroglossus azara is the correct name used for this species, not P. flavirostris, the latter used since Peters (1948).
10. Pteroglossus torquatus (including sanguineus, erythropygius, and Middle American P. frantzii), P. pluricinctus, P. aracari, and P. castanotis were considered to form a superspecies by Haffer (1974); however, P. pluricinctus is widely sympatric with P. castanotis in western Amazonia. Sibley & Monroe (1990) considered Pteroglossus torquatus and P. pluricinctus to form a superspecies, but preliminary genetic data (Hackett & Lehn 1997) provide not support for that relationship. Pteroglossus aracari and P. castanotis are generally considered to be sister species based on plumage similarities (e.g., Prum 1988b); Sibley & Monroe (1990), Short & Horne (2001), and Short & Horne (2002b) considered P. aracari and P. castanotis to form a superspecies.
10b. "Pteroglossus olallae," known only from the type specimen from the Rio Jurua, Brazil, was treated as a species by Peters (1948) and Meyer de Schauensee (1966) but is generally considered a hybrid or aberrant individual. See Hybrids and Dubious Taxa.
10c. "Pteroglossus formosus," known from an uncertain locality and treated as a valid species by Cory (1919), is considered a synonym of Pteroglossus a. aracari by Short & Horne (2002b). See Hybrids and Dubious Taxa.
11. Subspecies sanguineus ("Stripe-billed Aracari") and erythropygius ("Pale-mandibled Aracari") were formerly (e.g., Ridgway 1914, Cory 1919, Peters 1948, Meyer de Schauensee 1970, Dickinson 2003) considered separate species from Pteroglossus torquatus, but hybridization in areas of contact with each other and with nominate torquatus (e.g., Haffer 1967) has led some subsequent authors (e.g., Haffer 1974, Short and Horne 2002b) to consider them as subspecies of P. torquatus. Haffer (1967) interpreted specimen data as indicating free interbreeding between torquatus and sanguineus in northwestern Colombia; these two differ no more from each other than do sanguineus and erythropygius from each other, and so Haffer (1974) considered the latter also as a subspecies of torquatus. Short & Horne (2001) also reported signs of extensive intergradation between sanguineus and erythropygius and between torquatus and sanguineus. Preliminary genetic data (Hackett & Lehn 1997) are consistent with a close relationship among these four taxa, as is the traditional treatment based on plumage characters (e.g., Prum 1988b). Sibley & Monroe (1990) and Ridgely & Greenfield (2001) continued to rank them all as species.
SACC proposal to recognize sanguineus and erythropygius as separate species did not pass.
11a. Called "Spot-breasted Aracari" in Haffer (1974) and
Short & Horne (2001).
12. Unusual crown feathers and face pattern led to former placement of Pteroglossus beauharnaesii
in monotypic genus Bauharnaisius by some authors (e.g., Ridgway 1914, Cory 1919, Pinto 1937). Preliminary genetic data (Hackett & Lehn 1997) indicate that beauharnaesii is nested within Pteroglossus, and perhaps closest to P. bitorquatus.
13. Pteroglossus bailloni has previously been placed nearly universally in a monotypic genus, Baillonius. Peters (1948) placed Baillonius bailloni in Andigena, but genetic data (Hackett & Lehn 1997, Barker & Lanyon 2000, Nahum et al. Moyle 2004, Weckstein 2004) support the widespread view (e.g., Haffer 1974, Short & Horne 2001) that Baillonius and Pteroglossus are sister genera. Recent genetic data (Kimura et al. 2004) further indicate that Baillonius is embedded within Pteroglossus and thus should be merged into that genus.
SACC proposal passed to merge Baillonius into Pteroglossus.
14. The species in the genus Selenidera form a superspecies (Haffer 1974; cf. Short & Horne 2001, 2002b).
14a. The species name for Selenidera culik was formerly (e.g., <check Cory 1919> Pinto 1937) piperivora, but see Peters (1948).
15. The subspecies langsdorffii was formerly (e.g., Cory 1919, Peters 1948) treated as separate species from Selenidera reinwardtii, but they were considered conspecific by Meyer de Schauensee (1966) and subsequent authors.
16. Selenidera gouldii was formerly (e.g., Cory 1919,
Pinto 1937, Peters 1948, Meyer de Schauensee 1970) considered a subspecies of S. maculirostris, but they were treated as separate species by Haffer (1974); they are presumably sister species.
17. Andigena hypoglauca, A. laminirostris, and A. cucullata form a superspecies (Haffer 1974, Sibley & Monroe 1990, Short & Horne 2001, Short & Horne 2002b).
18. Haffer (1974) and Sibley & Monroe (1990) considered Ramphastos sulfuratus, R. brevis, R. vitellinus, and R. dicolorus to form a superspecies.
18a. Genetic data (Weckstein 2004) indicate thatRamphastos toco is the basal species in the genus, and that Haffer's (1974) "croaker" group (R. vitellinus, R. brevis, R. sulfuratus) and "yelper" group (R. tucanus, R. ambiguus/swainsonii) were monophyletic; support for the placement of R. sulfuratus, however, in the "croaker" group was weak.
SACC proposal passed to change linear sequence of species to the one used in this classification.
18b. Called "Green-billed Toucan" in Mazar Barnett & Pearman (2001).
19. Cory (1919) and Meyer de Schauensee (1966, 1970) considered R. culminatus ("Yellow-ridged Toucan") and R. citrolaemus ("Citron-throated Toucan") as separate species from Ramphastos vitellinus. Haffer (1974) treated these as a subspecies of R. vitellinus, and this treatment, actually a partial return to the classification of
Pinto (1937) and Peters (1948), has been followed by most subsequent authors (but not Sibley & Monroe 1990, Hilty 2003). Haffer identified broad hybrid zones between vitellinus and culminatus wherever they meet; see Short & Horne (2001) for additional information. "Ramphastos osculans," known from northern Brazil, Venezuela, and Guyana, and treated as a valid species by Cory (1918), is a population of intergrades between R. culminatus and R. vitellinus (Hellmayr 1933, Haffer 1974). See Hybrids and Dubious Taxa.
19a. The subspecies ariel of eastern and southeastern Brazil was formerly
(e.g., Cory 1919) considered a separate species from Ramphastos vitellinus, but Pinto (1937) and Peters (1948) treated them as conspecific. Weckstein (2004) found that Amazonian ariel was more closely related to culminatus than either was to nominate vitellinus (opposite the relationship suggested by plumage characters; Prum 1988b); also, Weckstein (2004) also found that ariel from the Atlantic forest region was only distantly related to Amazonina ariel.
20. Ramphastos brevis was formerly (e.g., Peters 1948) considered a subspecies of R. ambiguus, but the two differ in vocalizations [REF]; plumage characters suggest that R. brevis and R. sulfuratus are sister species (Prum 1988b), but this is not supported by genetic data (Weckstein 2004).
21. Called "Rainbow-billed Toucan" by Short & Horne (2001) and Short & Horne (2002b).
22. Haffer (1974) showed that R. cuvieri ("Cuvier's Toucan") and R. tucanus form a broad hybrid zone in northern and eastern Amazonia, and, therefore, treated cuvieri (with inca, also considered a separate species by Peters 1948) as a subspecies of Ramphastos tucanus; see also Short & Horne (2001); this treatment has been followed by most subsequent authors, but Sibley & Monroe (1990) continued to treat cuvieri as a species, following earlier classifications (e.g., Cory 1919, Peters 1948, Meyer de Schauensee 1970).
22b. Also known as "White-breasted Toucan" (Haffer 1974, Hilty 2003) and "Red-billed Toucan" (Meyer de Schauensee 1970).
23. Haffer (1974) considered Ramphastos tucanus and R. ambiguus to form a superspecies.
23b. "Ramphastos aurantiirostris," formerly (e.g., Peters 1948, Phelps & Phelps 1958a, Meyer de Schauensee 1966, 1970) treated as a species, it is only a color variant of nominate R. tucanus (Pinto 1938, Haffer 1974). See Hybrids and Dubious Taxa.
24. Haffer (1974) considered the taxon swainsonii to be a subspecies of R. ambiguus, and this treatment has been followed by Short & Horne (2001, 2002b); they have very similar if not identical voices and differ only in color of facial skin and mandible (e.g., Prum 1988b). Others continue to treat them as component species in a superspecies (e.g., AOU 1998, Ridgely & Greenfield 2001). Stiles et al. (1999) noted vocal and biometric differences between abbreviatus and ambiguus and noted that abbreviatus of the Magdalena valley should be treated as conspecific with swainsonii (not ambiguus as treated by some authors) if the species is split. The R. a. swainsonii group and R. a. ambiguus apparently replace one another on opposite slopes of the East Andes and show a 1.4% difference in mtDNA sequences (Donegan et al. 2007).
Proposal needed. Treatment of swainsonii as a species (e.g., Ridgway 1914, Cory 1919, Peters 1948, Meyer de Schauensee 1970) may have persisted because of the sympatry between it and R. brevis, which was described as, and long thought to be, a subspecies of ambiguus (e.g., Peters 1948, Meyer de Schauensee 1966; see Short & Horne 2001, Short & Horne 2002b).
25. Called "Yellow-throated Toucan" by Haffer (1974), Short & Horne (2001), and Short & Horne (2002b).
Proposal needed?



PICIDAE (WOODPECKERS) 1
Picumnus aurifrons Bar-breasted Piculet 2, 3, 4
Picumnus pumilus
Orinoco Piculet 5, 5a
Picumnus lafresnayi
Lafresnaye's Piculet 4, 5
Picumnus exilis
Golden-spangled Piculet 5b, 6, 7
Picumnus sclateri
Ecuadorian Piculet
Picumnus squamulatus Scaled Piculet
Picumnus spilogaster White-bellied Piculet 8, 9
Picumnus minutissimus
Arrowhead Piculet 8, 9, 10, 16
Picumnus pygmaeus
Spotted Piculet 11, 17b
Picumnus steindachneri
Speckle-chested Piculet
Picumnus varzeae Varzea Piculet 11, 12
Picumnus cirratus
White-barred Piculet 12, 13, 14, 16
Picumnus dorbignyanus
Ocellated Piculet 13, 15, 16
Picumnus temminckii
Ochre-collared Piculet 13, 16
Picumnus albosquamatus
White-wedged Piculet 16, 17, 17b
Picumnus fuscus
Rusty-necked Piculet 18
Picumnus rufiventris
Rufous-breasted Piculet
Picumnus fulvescens Tawny Piculet 19, 20
Picumnus limae
Ochraceous Piculet 19, 20
Picumnus nebulosus
Mottled Piculet 20, 20a
Picumnus castelnau
Plain-breasted Piculet 21
Picumnus subtilis
Fine-barred Piculet 21
Picumnus olivaceus
Olivaceous Piculet 22
Picumnus granadensis
Grayish Piculet 22
Picumnus cinnamomeus
Chestnut Piculet
Melanerpes candidus White Woodpecker 23
Melanerpes formicivorus
Acorn Woodpecker 24, 25
Melanerpes cruentatus
Yellow-tufted Woodpecker 26, 27, 28, 29
Melanerpes flavifrons
Yellow-fronted Woodpecker 26, 27
Melanerpes pulcher
Beautiful Woodpecker 26, 30, 31
Melanerpes pucherani
Black-cheeked Woodpecker 26, 30
Melanerpes cactorum
White-fronted Woodpecker 32
Melanerpes rubricapillus
Red-crowned Woodpecker 33, 34, 35
Picoides fumigatus
Smoky-brown Woodpecker 41
Veniliornis kirkii
Red-rumped Woodpecker 44
Veniliornis cassini
Golden-collared Woodpecker 45
Veniliornis spilogaster
White-spotted Woodpecker 42
Veniliornis mixtus
Checkered Woodpecker 36, 37, 38
Veniliornis lignarius
Striped Woodpecker 36, 37
Veniliornis sanguineus
Blood-colored Woodpecker
Veniliornis passerinus Little Woodpecker 42, 43
Veniliornis frontalis
Dot-fronted Woodpecker 42
Veniliornis callonotus
Scarlet-backed Woodpecker
Veniliornis dignus Yellow-vented Woodpecker 39, 40
Veniliornis nigriceps
Bar-bellied Woodpecker 40
Veniliornis affinis
Red-stained Woodpecker 44, 45, 46
Veniliornis chocoensis
Choco Woodpecker 45
Veniliornis maculifrons
Yellow-eared Woodpecker
Piculus leucolaemus White-throated Woodpecker 47, 47a, 48, 49, 35
Piculus litae
Lita Woodpecker 48
Piculus flavigula
Yellow-throated Woodpecker 50
Piculus chrysochloros
Golden-green Woodpecker 51, 52
Piculus aurulentus
White-browed Woodpecker 51
Colaptes rubiginosus
Golden-olive Woodpecker 47, 53, 54, 55
Colaptes rivolii
Crimson-mantled Woodpecker 47, 55, 56, 57
Colaptes atricollis
Black-necked Woodpecker 58, 59, 35
Colaptes punctigula
Spot-breasted Woodpecker 58, 59, 61
Colaptes melanochloros
Green-barred Woodpecker 58, 59, 60, 61
Colaptes pitius
Chilean Flicker 62, 63
Colaptes rupicola
Andean Flicker 62, 63, 64
Colaptes campestris
Campo Flicker 62, 65, 66
Celeus loricatus
Cinnamon Woodpecker 66
Celeus undatus
Waved Woodpecker 66
Celeus grammicus
Scale-breasted Woodpecker 66, 67, 68
Celeus elegans
Chestnut Woodpecker 69, 70, 70a
Celeus lugubris
Pale-crested Woodpecker 69, 70a, 71
Celeus flavescens
Blond-crested Woodpecker 69
Celeus flavus
Cream-colored Woodpecker 72
Celeus spectabilis
Rufous-headed Woodpecker 73
Celeus obrieni
Kaempfer's Woodpecker 73, 73a
Celeus torquatus
Ringed Woodpecker 74, 75
Dryocopus galeatus
Helmeted Woodpecker 76
Dryocopus lineatus
Lineated Woodpecker 76, 77, 78, 79
Dryocopus schulzi
Black-bodied Woodpecker 77, 80, 80a
Campephilus pollens
Powerful Woodpecker 81, 82
Campephilus haematogaster
Crimson-bellied Woodpecker 82, 83, 84
Campephilus rubricollis
Red-necked Woodpecker 81, 85, 85a
Campephilus robustus
Robust Woodpecker
Campephilus melanoleucos Crimson-crested Woodpecker 81, 86, 87
Campephilus gayaquilensis
Guayaquil Woodpecker 81, 87
Campephilus leucopogon
Cream-backed Woodpecker 81
Campephilus magellanicus
Magellanic Woodpecker 88


1. The monophyly of the Picidae has never been seriously questioned. Within the Piciformes, evidence supports a sister relationship to the Old World Indicatoridae (<REFS>, Prychitko & Moore 2003, Cracraft et al. 2004, Fain & Houde 2004, Webb & Moore 2005, Benz et al. 2006, Ericson et al. 2006, Hackett et al. 2008). The linear arrangement and composition of genera below in general follows that of Short (1982), who placed the piculets in a separate subfamily, Picumninae, and divided the typical woodpeckers, Picinae, into six tribes, four of which have representatives in South America: Melanerpini for a broadly defined Melanerpes and Sphyrapicus; Campetherini for a broadly defined Picoides and Veniliornis; Colaptini for Piculus, Colaptes, and Celeus; and Campephilini for Dryocopus and Campephilus. In general, Short's classification, culminating in a monographic treatment of the family (Short 1982), merged many previously recognized genera into many fewer, broadly defined genera. <incorp. Goodge 1972>. Genetic data (Webb & Moore 2005, Benz et al. 2006) that most of these groups are not monophyletic. Webb and Moore (2005), generally supported by Benz et al. (2006), recommended a classification with three tribes for the three major groups in the Picinae: (1) Malarpicini for Colaptes, Piculus, Celeus, Dryocopus, and several Old World genera; (2) Dendropicini for Picoides, Veniliornis, Melanerpes, Sphyrapicus, and several Old World genera; and (3) Campephilus, Chrysocolaptes, and two Old World genera. Proposal needed for change in linear sequence. Genetic data (Benz et al. 2006) support the monophyly and distinctiveness of the Picuminae (Picumnus and Old World Sasia, but not Caribbean Nesoctites) as the sister taxon to all other woodpeckers. Proposal badly needed for subfamily recognition.
2. Species-level taxonomy in the genus Picumnus is in need of major re-evaluation; interbreeding, to varying degrees, between various pairs of parapatric and partially sympatric species is inordinately high; see Short (1982).
3. Following Short (1982), Sibley & Monroe (1990), and Winkler & Christie (2002), Picumnus aurifrons here includes P. borbae (with juruanus), treated as a separate species (as "Bar-breasted Piculet," with aurifrons called "Gold-fronted Piculet") by Meyer de Schauensee (1970) and others. The subspecies wallacii was also formerly
(e.g., Cory 1919) considered a separate species from Picumnus aurifrons, but Peters (1948) treated them as conspecific. The subspecies pusillus was described and treated as a separate species (Pinto 1937), but Peters (1948) treated it as conspecific with P. aurifrons.
4. Picumnus lafresnayi was formerly (e.g., Peters 1948, Meyer de Schauensee 1970) considered a subspecies of P. aurifrons, but see Short (1982) for rationale for treating as a separate species, representing a return to the classification of Cory (1919). The subspecies punctifrons was also formerly
(e.g., Cory 1919) considered a separate species, but Peters (1948) treated them as conspecific. Short (1982) proposed that the sister species of P. lafresnayi could be either P. aurifrons or P. exilis.
5. Picumnus pumilis was formerly (e.g., Short 1982) treated as a subspecies of P. lafresnayi, and they are presumed sister species that form a superspecies (Sibley & Monroe 1990); they overlap slightly in se. Colombia with no sign of interbreeding (Hilty & Brown 1986, Winkler & Christie 2002).
5a. "Picumnus stellae," known from the Río Orinoco, Venezuela, and treated as a valid species by Cory (1919), is now considered a synonym of P. pumilus (Peters 1948). See Hybrids and Dubious Taxa.
5b. Picumnus exilis was considered by Short (1982) to be most closely related to the P. aurifrons group (of Note 3 above).
6. The taxon nigropunctatus was formerly (e.g., Phelps & Phelps 1958a, Meyer de Schauensee 1970) considered a separate species ("Black-spotted Piculet") from Picumnus exilis, but Short (1982) stated that it was a synonym of P. exilis, and this treatment was followed by Sibley & Monroe (1990). It continues to be ranked as a species by Rodner et al. (2000) and Winkler & Christie (2002), based in part on unpublished data of M. Lentino, which is summarized in Winkler & Christie (2002).
proposal needed.
7. The subspecies undulatus, buffoni, and salvini were formerly
(e.g., Cory 1919) each considered a separate species from Picumnus exilis, but Peters (1948) treated them all as conspecific.
8. The subspecies pallidus was formerly (e.g.,
Pinto 1937, Peters 1948) considered a separate species from Picumnus spilogaster, or was considered as a subspecies of Picumnus minutissimus (Meyer de Schauensee 1966); plumage pattern, however, favors treatment as a subspecies of P. spilogaster (Short 1982, Winkler & Christie 2002).
9. Picumnus
spilogaster was formerly (e.g., Cory 1919, Pinto 1937, Peters 1948) known as P. leucogaster, but see Zimmer & Phelps (1950) and Meyer de Schauensee (1966); Peters (1948) considered it a synonym of P. minutissimus, but see <REF>.
10. Called "Guianan Piculet" in Sibley & Monroe (1990) and Dickinson (2003).
proposal needed?
11. Meyer de Schauensee (1966) suggested that Picumnus varzeae might be a subspecies of P. pygmaeus, but see Short (1982).
12. Picumnus varzeae and P. cirratus hybridize to an uncertain extent along the Amazon River (Short 1982).
13. Picumnus cirratus, P. dorbignyanus, and P. temminckii are considered to form a superspecies (Sibley & Monroe 1990, Winkler & Christie 2002); they interbreed to varying and uncertain degrees where parapatric (Short 1982, Winkler & Christie 2002), and thus have all been considered conspecific by some (e.g., Short 1982). Relationships among these three and also P. albosquamatus (see Note 16) are badly in need of detailed study.
14. The Peruvian subspecies jelskii was formerly
(e.g., Cory 1919) considered a separate species from Picumnus cirratus, but Peters (1948) treated them as conspecific, and this has been followed by Short (1982) and most subsequent classifications.
15. Winkler & Christie (2002) pointed out that an error has been perpetuated in the spelling of the species name, usually given incorrectly as "dorbygnianus" (e.g., Meyer de Schauensee 1970).
16. Picumnus albosquamatus interbreeds to varying uncertain degrees with P. dorbignyanus, P. temminckii, and P. cirratus (Short 1982, Winkler & Christie 2002), and may be part of that superspecies (Short 1982); it was considered conspecific with P. minutissimus by Meyer de Schauensee (1966), following Gyldenstolpe (1945), but see Short (1982).
17. The southern Brazilian subspecies guttifer was formerly
(e.g., Cory 1919, Pinto 1937, Peters 1948) considered a separate species from Picumnus albosquamatus/minutissimus, but Meyer de Schauensee (1966) and Short (1982) treated them as conspecific, following Bond and Meyer de Schauensee (1943).
17b. "Picumnus asterias," known only from the type specimen from "Brazil" and treated as a valid species by Cory (1919),
Pinto (1937), and Peters (1948), and as tentatively valid by Meyer de Schauensee (1966), is possibly a variant of P. pygmaeus (Meyer de Schauensee 1966, 1970) or P. albosquamatus guttifer (Short 1982, Sibley & Monroe 1990). "Picumnus arileucus," described from Mato Grosso and treated as a valid species by Pinto (1937), is now considered a synonym of P. albosquamatus corumbanus (Peters 1948). See Hybrids and Dubious Taxa.
18. Picumnus fuscus was considered a doubtful species by Peters (1948) and Meyer de Schauensee (1966), and was not recognized as a species by Meyer de Schauensee (1970); Short (1982) recognized it as a valid species, and this has been followed by subsequent authors.
19. Picumnus fulvescens was formerly (e.g., Meyer de Schauensee 1970) treated as a subspecies of P. limae ; they are presumably sister species (Winkler & Christie 2002); see Short (1982) for rationale for treating fulvescens as a species, and for the possibility that P. l. saturatus is a synonym of P. fulvescens.
20. Short (1982) suspected that Picumnus nebulosus might be closely related to P. fulvescens and P. limae.
20a. "Picumnus iheringi," known from southeastern Brazil; and treated as a valid species by Pinto (1937), is now considered a synonym of P. nebulosus (Gyldenstolpe 1945, Peters 1948).
21. Picumnus castelnau and P. subtilis were considered to be sister species by Short (1982) and Winkler & Christie (2002); they may occasionally hybridize (Short 1982).
22. Picumnus olivaceus and P. granadensis were considered to form a superspecies by Short (1982), Sibley & Monroe (1990), and Winkler & Christie (2002), and evidence for treatment as separate species is weak; they were formerly (e.g., Ridgway 1914) considered conspecific.
23. Melanerpes candidus was formerly (e.g., Cory 1919,
Pinto 1937, Peters 1948, Meyer de Schauensee 1970) placed in the monotypic genus Leuconerpes, but most authors have followed Short (1982) in merging this into Melanerpes.
24. Melanerpes formicivorus was formerly (e.g., Ridgway 1914, Cory 1919) placed in a separate genus Balanosphyra, but most authors have followed Peters (1948) in merging this into Melanerpes.
25. The Colombian subspecies flavigula was formerly
(e.g., Cory 1919) considered a separate species from Middle American Melanerpes formicivorus, but Peters (1948) and Short (1982) treated them as conspecific.
26. Melanerpes cruentatus, M. flavifrons, M. chrysauchen, and M. pucherani were formerly
(e.g., Ridgway 1914, Cory 1919, Pinto 1937) treated in a separate genus, Tripsurus, but Peters (1948) merged this into Melanerpes; these four species were considered by Short (1982) to form a superspecies.
27. Melanerpes cruentatus and M. flavifrons form a superspecies (Short 1982, Sibley & Monroe 1990, Winkler & Christie 2002).
28. Called "Red-fronted Woodpecker" by Short (1982).
29. The form "rubrifrons" was formerly (e.g.,
Pinto 1937, Phelps & Phelps 1958a, Meyer de Schauensee 1966, 1970) considered a separate species, but most recent authors have followed Griscom & Greenway (1941) and Short (1982) in treating it as a regional color morph of M. cruentatus.
30. Melanerpes chrysauchen and M. pucherani form a superspecies (Sibley & Monroe 1990, Winkler & Christie 2002). Short (1982) also included M. cruentatus and M. flavifrons in this superspecies; in head and back pattern, M. flavifrons is more similar to distant M. chrysauchen and M. pucherani than it is to nearby M. cruentatus.

31.
The Colombian taxon pulcher was considered a separate species from Central American Melanerpes chrysauchen by Cory (1919), Eisenmann (1955), and Stiles & Skutch (1989); however, Peters (1948) treated them as conspecific, and that treatement has been followed by most subsequent authors (e.g., Meyer de Schauensee 1979, Short 1982, Hilty & Brown 1986, Winkler et al. 1995, AOU 1998, Winkler & Christie 2002, Dickinson 2003). Wetmore (1968) provided rationale for treating pulcher as a separate species, as noted by Meyer de Schauensee (1966), but this has not been followed by most subsequent authors. SACC proposal passed to recognize pulcher as separate species.
32. Melanerpes cactorum was formerly (e.g., Cory 1919, Peters 1948, Meyer de Schauensee 1970) placed in a separate monotypic genus Trichopicus, but recent authors have followed Short (1982) in merging this into Melanerpes, as suggested long ago by Wetmore (1926).
33. Melanerpes rubricapillus
was formerly (e.g., Ridgway 1914, Cory 1919, Phelps & Phelps 1958a) placed in the genus Centurus, along with many North and Middle American species; Peters (1948) merged Centurus into Melanerpes, and this has been followed by most recent authors.
34. Melanerpes rubricapillus was considered conspecific with Middle American M. pygmaeus by Peters (1948) and Short (1982); they were treated as members of a superspecies by Sibley & Monroe (1990), and Winkler & Christie (2002).
35. Melanerpes rubricapillus
was formerly (e.g., Ridgway 1914, Cory 1919) known as M. subelegans, but see Peters (1948).
36.
Veniliornis lignarius and V. mixtus form a superspecies (Short 1982, Fjeldså & Krabbe 1990, Sibley & Monroe 1990, Winkler & Christie 2002); justification is weak for their treatment as separate species (Short 1970, 1971, 1982); genetic data (Weibel & Moore 2002a, b) confirm their relationship as sister species.
37.
Veniliornis lignarius and V. mixtus were formerly (e.g., Cory 1919, Pinto 1937) treated in a separate genus, Dyctiopicus, but Peters (1948) merged this into Dendrocopos, which was then merged into Picoides by Short (1970, 1971, 1982); see Goodwin (1968) and Ouellet (1978) for differing view. Recent genetic data (Weibel & Moore 2002a, 2002b, Webb & Moore 2005), however, indicate that the widespread genus Picoides is polyphyletic unless Veniliornis and Dendropicos are included. In particular, the two South American species formerly treated in Picoides are more closely related to Veniliornis (as represented by V. nigriceps and V. callonotus) than they are to Northern Hemisphere Picoides; see also Moore et al. (2006). This result is exceptionally robust with respect to analytical techniques, and it includes both mitochondrial and nuclear genes. However, it might be best to wait for additional taxon-sampling before proposing a merger (and to wait for broader rearrangement of Picoides, which consists of at least five lineages worthy of generic recognition, including restoration of Dendrocopos and Dryobates). Date from Moore et al. (2006), however, require removal of lignarius and mixtus from Picoides. SACC proposal passed to transfer to Veniliornis. SACC proposal passed to change linear sequence within Veniliornis.
38. The northeastern subspecies cancellatus was formerly
(e.g., Cory 1919, Pinto 1937) considered a separate species from Veniliornis mixtus, but Peters (1948) treated them as conspecific; they intergrade where their ranges meet in northeastern Paraguay (Short 1982).
39. The Peruvian subspecies valdizani was formerly
(e.g., Cory 1919) considered a separate species from Veniliornis dignus, but Peters (1948) and Short (1982) treated them as conspecific.
40. Short (1982) noted that plumage similarities suggest that Veniliornis dignus and V. nigriceps are sister species.
41.
Veniliornis fumigatus was formerly (e.g., Ridgway 1914, Cory 1919) known as V. oleaginus, but see Peters (1948).
41a. Genetic data (Moore et al. 2006) indicate that Veniliornis fumigatus is embedded within Picoides and is not closely related to Veniliornis sense stricto.
SACC passed to transfer to Picoides.
42. Veniliornis passerinus and V. frontalis are sister taxa (Zimmer 1942a) that form a superspecies (Short 1982); they may hybridize to a limited extent (Short 1982, Winkler & Christie 2002). Short (1982) also noted that plumage similarities suggest that V. spilogaster might be the sister species to V. dignus + V. nigriceps
43. The subspecies taenionotus (with "cearae") of eastern Brazil was formerly
(e.g., Cory 1919) considered a separate species from Veniliornis passerinus, but Zimmer (1942a), Peters (1948), and Short (1982) treated them as conspecific. The subspecies fidelis, agilis, and olvinus were also formerly (e.g., Cory 1919) each considered separate species from V. passerinus, but Zimmer (1942a), Peters (1948), and Short (1982) also treated them all as conspecific.
44. Veniliornis kirkii, V. affinis, V. cassini, and V. maculifrons form a superspecies (Short 1982, Haffer 1987, Sibley & Monroe 1990, Winkler & Christie 2002). However, the apparent broad geographic overlap between V. kirkii and V. a. chocoensis would invalidate the superspecies designation.
45. The taxon chocoensis was formerly (e.g., Meyer de Schauensee 1970) regarded as a subspecies of Veniliornis cassini, but it was transferred to V. affinis by Short (1974, 1982). It was treated as a separate species ("Choco Woodpecker") by Sibley & Monroe (1990), Winkler et al. (1995), Ridgely & Greenfield (2001), and Winkler & Christie (2002), but little evidence is published to support this; chocoensis differs from V. affinis and V. cassini only in minor plumage details.
SACC proposal passed to elevate chocoensis to species rank.
46. The subspecies ruficeps (with "haematostygma" = hilaris; see Zimmer 1942a) and orenocensis were formerly
(e.g., Cory 1919) both considered separate species from Veniliornis affinis, but they were all treated as conspecific by Zimmer (1942a), Peters (1948), and Short (1982).
47. Genetic data (Webb & Moore 2005, Benz et al. 2006) indicate that the genus Piculus is paraphyletic with respect to Colaptes: P. rubiginosus and P. rivolii are embedded within Colaptes.
SACC passed to transfer the latter two to Colaptes.
47a. The name formerly (e.g., Ridgway 1914, Cory 1919) used for the genus Piculus was Chloronerpes, but see Peters (1948).
48. The subspecies litae of the Chocó was formerly
(e.g., Cory 1919) considered a separate species from Piculus leucolaemus; Peters (1948) treated them as conspecific, and this was followed by many subsequent authors (e.g., Meyer de Schauensee 1970, Short 1982). Recently, litae was treated as a species ("Lita Woodpecker") by Sibley & Monroe (1990), Ridgely & Greenfield (2001), and Winkler & Christie (2002) and [REFS], but published evidence to support this is weak. On the other hand, litae resembles P. flavigula in some aspects of its plumage as much as it does P. leucolaemus; in fact, specimens of litae from southwestern Colombia have been misidentified as P. flavigula (REF, Winker & Christie 2002). SACC proposal passed to elevate litae to species rank.
49. Sibley & Monroe (1990) considered Piculus leucolaemus to form a superspecies with Middle American P. simplex and P. callopterus; some authors (e.g., Short 1982, AOU 1983) have considered them all as conspecific, and Peters (1948) considered P. callopterus to be a subspecies of P. leucolaemus. See Wetmore (1968) and Stiles & Skutch (1989) for rationale for treating them as separate species.
50. The subspecies erythropis of eastern and southeastern Brazil was formerly
(e.g., Cory 1919, Pinto 1937) considered a separate species from Piculus flavigula, but Peters (1948) and Short (1982) treated them as conspecific.
51. Piculus chrysolochloros and P. aurulentus form a superspecies (Short 1982, Sibley & Monroe 1990, Winkler & Christie 2002).
52. The subspecies xanthochlorus was formerly
(e.g., Cory 1919) considered a separate species from Piculus chrysochloros, but Peters (1948) and Short (1982) treated them as conspecific.
53. Piculus rubiginosus and Middle American P. auricularis form a superspecies (Short 1982, Sibley & Monroe 1990, Winker & Christie 2002).
54. The Peruvian subspecies chrysogaster was formerly
(e.g., Cory 1919) considered a separate species from Piculus rubiginosus, but Peters (1948) and Short (1982) treated them as conspecific.
55. Short (1982) considered Piculus rivolii to be the sister species to P. rubiginosus/auricularis based on plumage similarities.
56. Piculus rivolii was formerly
(e.g., Cory 1919) treated in a separate genus, Hypoxanthus, but Peters (1948) merged this into Piculus.
57.
The southern subspecies atriceps was formerly (e.g., Cory 1919) considered a separate species from Piculus rivolii, but Peters (1948) and Short (1982) treated them as conspecific.
58. Colaptes atricollis, C. punctigula, and C. melanochloros were formerly
(e.g., Cory 1919, Pinto 1937, Peters 1948, Phelps & Phelps 1958a, Meyer de Schauensee 1970) treated in a separate genus, Chrysoptilus, but Short (1965, 1972a, 1982) merged this into Colaptes. However, plumage similarities of these three species to Piculus suggests that further study may reveal a closer relationship to that genus ; in fact, recent genetic data with limited taxon-sampling suggest that Piculus and South American Colaptes are more closely related to each other than either is to North American Colaptes (Prychitko & Moore 2000, Weibel & Moore 2002a, b; see also Webb & Moore 2005). Ridgely & Greenfield (2001) and Hilty (2003) retained Chrysoptilus for punctigula only.
59.
Colaptes atricollis, C. punctigula, and C. melanochloros were called "Flickers" by Short (1982).
60. The subspecies melanolaimus (with nigroviridis and leucofrenatus) was formerly (e.g., Cory 1919, Traylor 1951c, Meyer de Schauensee 1970; but not Laubmann 1934, Peters 1948) considered a separate species ("Golden-breasted Woodpecker") from Colaptes melanochloros, but they intergrade where in contact (Short 1972a, Winkler & Christie 2002). The subspecies nigroviridis and "mariae" were also formerly
(e.g., Cory 1919) each considered separate species from Colaptes melanochloros, but Peters (1948) treated them all as conspecific; "mariae" is not currently recognized as a valid taxon at any level (Short 1972a, 1982, Winkler & Christie 2002).
61. Short (1972a, 1982) and Sibley & Monroe (1990) considered Colaptes punctigula and C. melanochloros to form a superspecies.
62. Colaptes campestris and C. rupicola were formerly
(e.g., Cory 1919) treated in a separate genus, Soroplex, but Peters (1948) merged this into Colaptes. Colaptes pitius was also formerly (e.g., Cory 1919) treated in a separate monotypic genus, Pituipicus, but Peters (1948) also merged this into Colaptes. Short (1982) considered Colaptes campestris to be the sister species to Colaptes pitius + C. rupicola. Although these South American flickers have been considered congeneric with North American Colaptes since Peters (1948), their distribution and plumage similarities to Piculus and "Chrysoptilus" suggest that their morphological similarities to North American flickers may be due to convergence. Short (1972a) proposed that the broadly defined Colaptes was the sister genus to Piculus, and he suspected that they could be merged into a single genus.
63.
Plumage similarities and somewhat complementary distributions suggest that Colaptes pitius and C. rupicola are sister species (Short 1982).
64. The northern subspecies cinereicapillus and puna were formerly
(e.g., Cory 1919) both considered separate species from Colaptes rupicola, but Peters (1948) and Short (1982) treated them all as conspecific; Short (1972a) interpreted patterns of geographic variation in cinereicapillus, puna, and nominate rupicola to suggest intergradation among the three forms. Short (1982) reported differences in vocalizations between cinereicapillus and the other taxa, but it hybridizes to some extent with C. r. puna where in contact in central Peru.
65. The subspecies campestroides was formerly (e.g., Cory 1919, Meyer de Schauensee 1970; but not
Pinto 1937 or Peters 1948) considered a separate species ("Field Flicker") from Colaptes campestris, but they evidently freely interbreed where in contact (Short 1972a, 1982, Winkler & Christie 2002).
66. Haffer (1974) considered Celeus loricatus, C. grammicus, and C. undatus to form a superspecies; however, Short (1982), Sibley & Monroe (1990), and Winkler & Christie (2002) excluded loricatus from the superspecies.
67. Winkler & Christie (2002) suggested that the subspecies erythropis of southeastern Brazil might deserve recognition as a separate species from Celeus grammicus.
68. Called "Scaly-breasted Woodpecker" by Winkler & Christie (2002).
69. Short (1972b, 1982) and Sibley & Monroe (1990) considered Celeus elegans, C. lugubris, C. flavescens, and Middle American C. castaneus to form a superspecies; Peters (1948) considered lugubris to be conspecific with C. flavescens. Celeus elegans and C. lugubris occasionally hybridize where in contact in Brazil (Short 1972b).
70. The subspecies citreopygius and jumana were formerly
(e.g., Cory 1919, Pinto 1937) both considered separate species from Celeus elegans; Peters (1948) treated citreopygius and jumana as conspecific, but still considered jumana to be a separate species from C. elegans . Meyer de Schauensee (1966) and Short (1972b, 1982) considered the jumana group to be conspecific with C. elegans, and this has been followed by subsequent authors; they evidently intergrade in eastern Venezuela (Short 1972b).
70a. "Celeus roosevelti," described from so