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 Cathartiformes (click)
Part 2. Accipitriformes 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 Tityridae) (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,
14
Trogon mesurus Ecuadorian
Trogon 11, 13
Trogon melanurus Black-tailed Trogon 11, 12
Trogon chionurus White-tailed Trogon 1a, 2, 3, 4
Trogon viridis Green-backed
Trogon 1a, 2, 3, 4
Trogon caligatus Gartered Trogon 8,
8a
Trogon violaceus Violaceous Trogon 8,
8a
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 of orders that consists
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, Ornelas et al. 2009) 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 Choc— 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 Amazonian T. viridis. SACC proposal passed
to recognize chionurus as a species.
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 (including chionurus) 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, Ornelas et al. 2009) 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); it 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). SACC proposal passed
to recognize caligatus as a species.
8a. Trogon violaceus (including caligatus)
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,
Ornelas et al. 2009) 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 Amazonian 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, T. mesurus, 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, Ornelas et al. 2009) 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).
13.
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.
SACC proposal passed
to recognize mesurus as a species.
14. Called "White-eyed Trogon" in Sibley & Monroe (1990)
and "Choc— 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;
FjeldsŒ & Krabbe (1990) stated that heliactin may be 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 "Bogot‡," 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 relationship between the 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). FjeldsŒ &
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, Mayr et al. 2003, Manegold 2005) 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, Hšfling & Alvarenga 2001) suggested
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, Mayr et al. 2003, Cracraft et al. 2004,
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). Proposal needed to merge into Piciformes.
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 R’o 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 proposal
passed to apply English name "Guianan Puffbird" to narrowly
distributed macrorhynchos; SACC
proposal passed 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); this has been followed by most subsequent classifications,
except for Rasmussen & Collar (2002), who resurrected Argicus. Penhallurick (2008) noted that Cyphos has priority over 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 FjeldsŒ & 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 bailloni Saffron Toucanet
13
Pteroglossus viridis Green Aracari 6
Pteroglossus inscriptus Lettered Aracari
6, 7, 10b
Pteroglossus torquatus Collared Aracari
10a, 11, 11a
Pteroglossus aracari Black-necked Aracari
10, 10c
Pteroglossus castanotis Chestnut-eared
Aracari 10
Pteroglossus pluricinctus Many-banded
Aracari 10a
Pteroglossus azara Ivory-billed Aracari
8, 9, 9a, 10, 10b
Pteroglossus beauharnaesii Curl-crested
Aracari 12
Pteroglossus bitorquatus Red-necked Aracari
8
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 (Hšfling 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), Weckstein (2004), and Pereira & Wajntal
(2008) found that Andigena and Selenidera were sister genera, and
Moyle (2004) and Weckstein (2004) found that Aulacorhynchus was the
sister of Andigena + Selenidera. SACC proposal passed to change linear sequence of genera
to the one used in this classification. 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, FjeldsŒ & 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. Cory (1919) treated the subspecies humboldti as a separate species
from P. viridis and P. inscriptus. Peters (1948) treated humboldti as 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. Genetic data (Pereira & Wajntal 2008) support the
close relationship of the three taxa and indicate that humboldti and inscriptus
are sister taxa. "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. Genetic data (Pereira & Wajntal
2008) P. pluricinctus, P. castanotis,
and P. aracari form a monophyletic
group. <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 (REF) supports and genetic data (Pereira & Wajntal 2008; cf.
Hackett & Lehn 1997) their treatment as sister taxa.
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 genetic data (Hackett &
Lehn 199, Pereira & Wajntal 20087) provide no 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, but genetic data (Pereira & Wajntal 2008) suggest
that P. azara is more closely related
to P. castanotis + pluricinctus than to P. aracari.
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. Genetic
data (Hackett & Lehn 1997, Pereira & Wajntal 2008) 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). Genetic data (Hackett & Lehn 1997, Pereira & Wajntal
2008) indicate that beauharnaesii is not only nested within Pteroglossus,
but also the sister species to P. bitorquatus. SACC
proposal passed to change linear sequence.
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, Eberhard and Bermingham 2005, Pereira and Wajntal 2008) further indicate
that Baillonius is embedded within Pteroglossus and thus should
be merged into that genus.
Morphological and vocal data (Haffer 1974, Sick 1997, Short and Horne
2001, Hšfling 2004) are also consistent with this merger. SACC proposal passed to merge Baillonius into Pteroglossus.
SACC proposal passed to change linear sequence.
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 that Ramphastos 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
Amazonian 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 Picumninae (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 pumilus 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 R’o 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 treatment 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, FjeldsŒ & 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 proposal 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 proposal 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 Choc— 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.
calopterus; some authors (e.g., Short 1982, AOU 1983) have considered
them all as conspecific, and Peters (1948) considered P. calopterus
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 chrysochloros and P. aurulentus form a
superspecies (Short 1982, Sibley & Monroe 1990, Winkler & Christie
2002).
52. The subspecies xanthophyllous 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, Chrysopterus, 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 southwestern
Brazil and treated as a valid species by Cory (1919),
is now considered a probable hybrid or backcross between C. elegans
jumana and C. lugubris (Short 1972b). See Hybrids and
Dubious Taxa.
71. The subspecies kerri was formerly (e.g.,
Cory 1919) considered a separate species from Celeus lugubris,
but Peters (1948) and Short (1972b, 1982) treated them as conspecific.
72. Celeus flavus was formerly (e.g., Cory
1919, Pinto 1937) treated in a separate monotypic genus, Crocomorphus,
but Peters (1948) merged this into Celeus.
73. Celeus obrieni, known from one specimen from Piau’, Brazil,
is traditionally (e.g., Short 1973, 1982) considered a subspecies of C.
spectabilis, but it differs so dramatically in plumage from C. spectabilis
that this seems unlikely (Whittaker & Oren 1999, Winkler & Christie
2002). SACC proposal
passed to elevate obrieni to species rank.
73a. Formerly known as "Caatinga Woodpecker." SACC proposal passed to change English
name.
74. Celeus torquatus was formerly (e.g., Cory
1919, Pinto 1937) treated in a separate genus, Cerchneipicus, but Peters
(1948) merged this into Celeus.
75. The subspecies tinnunculus (with occidentalis) was
formerly (e.g., Cory 1919, Pinto 1937) considered a
separate species from Celeus torquatus, but Peters (1948) and
Short (1982) treated them as conspecific.
76. Dryocopus lineatus and D. galeatus were formerly (e.g., Ridgway 1914, Cory 1919,
Pinto 1937) treated in a separate genus, Ceophloeus, but Peters (1948)
merged this into Dryocopus. Short
(1982) noted that D. galeatus shares some plumage and structural
features with Celeus, especially C. spectabilis.
77. Dryocopus lineatus and D. schulzi, along with
North American D. pileatus, are considered to form a superspecies
by Mayr and Short (1970), Short (1982), Sibley & Monroe (1990), and Winkler
& Christie (2002); D. lineatus and D. schulzi
hybridize to a limited extent where their ranges meet (Short 1982).
78. The subspecies fuscipennis of western Ecuador and northwestern Peru
was formerly (e.g., Cory 1919) considered a separate
species from Dryocopus lineatus, but Peters (1948) and
Short (1982) treated them as conspecific.
79. The subspecies erythrops was formerly (e.g., Cory 1919, Pinto 1937, Peters 1948) considered a separate
species from Dryocopus lineatus it has been considered a color
morph of D. lineatus (Pinto 1947, Pergolani de Costa 1962), but
see Short (1975, 1982) for treatment as a subspecies of D. lineatus.
80. Dryocopus schulzi was formerly (e.g.,
Cory 1919) treated in a separate genus, Neophloeotomus, but Peters
(1948) merged this into Dryocopus.
80a. "Dryocopus shiptoni," known from Tucum‡n,
Argentina, and treated as a valid species by Cory (1918), is considered a
variant of D. schulzi (Peters 1948, Pergolani de Costa 1962,
Meyer de Schauensee 1966, Short 1982). See Hybrids and
Dubious Taxa.
81. Campephilus pollens, C. rubricollis, C. melanoleucos, C.
leucopogon, and C. gayaquilensis were formerly (e.g., Ridgway 1914, Cory
1919, Pinto 1937) treated in a separate genus, Scapaneus, but Peters
(1948) merged this into Phloeoceastes,
and this was followed by Phelps & Phelps (1958a) and Meyer de
Schauensee (1970). Most recent authors have followed REFS, Short (1982) in
merging Phloeoceastes into Campephilus. <check robustus>
82. Short (1982) considered Campephilus pollens and C. haematogaster
to be sister species based on plumage and morphology.
83. Campephilus haematogaster was formerly (e.g., Ridgway 1914, Cory
1919, Pinto 1937) treated in a separate monotypic genus, Cniparchus, but
Peters (1948) merged this into Phloeoceastes, which was then
merged into Campephilus by REFS?, Short (1982).
84. Ridgely & Greenfield (2001) suggested that the subspecies splendens
of the Western Andes might deserve recognition as a separate species from Campephilus
haematogaster, as it had been previously by (REF).
85. The southwestern subspecies trachelopyrus
was formerly (e.g., Cory 1919, Pinto 1937) considered
a separate species from Campephilus rubricollis, but they
intergrade in western Amazonia (Peters 1948, Short 1982).
85a. Called "Red-headed Woodpecker" in FjeldsŒ & Krabbe (1990),
presumably a lapsus.
86. The northwestern subspecies malherbii was formerly (e.g., Cory 1919) considered a separate species from Campephilus
melanoleucos, but Peters (1948) treated
them as conspecific; they intergrade in eastern Colombia (Short 1982).
87. Campephilus melanoleucos and C. gayaquilensis,
along with Middle American C. guatemalensis, are considered to
form a superspecies by Short (1982), FjeldsŒ & Krabbe (1990), Sibley &
Monroe (1990), and Winkler & Christie (2002); Campephilus melanoleucos
and C. gayaquilensis were considered conspecific by Peters
(1948), but see Meyer de Schauensee (1966).
88. Campephilus magellanicus was formerly (e.g., Cory 1919) treated in a separate monotypic genus, Ipocranter,
but Peters (1948) merged this into Campephilus.
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Part 6. Suboscine Passeriformes, A (Furnariidae) (click)