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, 11SEMNORNITHIDAE (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