A classification of the bird species of South America

17b. Genetic data (Burns et al. 2003, Burns & Naoki 2004) suggest that Thraupis and Pipraeidea are closely related and presumably sister genera. However, Thraupis itself is polyphyletic (Klicka et al. 2007, Sedano & Burns 2010), so a more complete Thraupidae phylogeny is needed before changes can be made. Sedano & Burns (2010) found that T. bonariensis is the one species of Thraupis that is sister to Pipraeidea. SACC proposal passed to transfer Thraupis bonariensis into Pipraeidea. This was followed by Dickinson & Christidis (2014).
17c. The species name formerly (e.g., REFS, Phelps & Phelps 1950a) for Thraupis episcopus was virens, but see .
18. Zimmer (1944a) suggested that Thraupis sayaca is only a subspecies of T. episcopus; the population in their area of contact, boliviana, is somewhat intermediate in plumage; it was described as a subspecies of T. episcopus but is closer in plumage to T. sayaca (Gyldenstolpe 1945). However, Gyldenstolpe (1945) found the two species sympatric in northern Bolivia; also, the two evidently differ in vocalizations (Ridgely & Tudor 1989). Also, T. glaucocolpa, roughly as distinct phenotypically from T. episcopus as T. sayaca is, is broadly sympatric with T. episcopus (Ridgely & Tudor 1989). Thraupis episcopus, T. sayaca, and T. glaucocolpa were considered to form a superspecies by Sibley & Monroe (1990), but T. glaucocolpa cannot be included because of it is almost completely overlapping in distribution with T. episcopus. Although Middle American and trans-Andean populations, the cana group, were treated as separate species by Ridgway (19##; "Blue Tanager"), they have been treated as conspecific since at least Hellmayr (1936). However, the cana group is superficially similar to T. sayaca, more so than it is to the pale-shouldered Amazonian episcopus group. Therefore, if sayaca is treated as a separate species, perhaps the cana group should also be returned to species rank.
19. Thraupis glaucocolpa was formerly (e.g., Hellmayr 1936, Meyer de Schauensee 1970, Storer 1970a) treated as a subspecies of T. sayaca, but most recent classifications have treated it as a separate species (e.g., Meyer de Schauensee & Phelps 1978, Ridgely & Tudor 1989, Sibley & Monroe 1990).
19a. Sedano & Burns (2011) found that Thraupis cyanocephala was not a member of Thraupis but was close to Anisognathus. Burns et al. (2016) also recommended resurrecting Sporathraupis, as in Dickinson & Christidis (2014). SACC proposal pending to resurrect Sporathraupis.
19aa. The subspecies olivicyanea of the Coastal Range of Venezuela was formerly (e.g., REF Thraupis cyanocephala, but they evidently intergrade where in contact (Meyer de Schauensee 1966).
19b. The Andean subspecies darwinii was formerly (e.g., Chapman 1926, Zimmer 1930) considered a separate species from Thraupis bonariensis.
20. Storer (1970a) merged Bangsia into Buthraupis, but this has not been followed by most subsequent authors; see Monroe et al. (1993). They are not sister genera (Sedano & Burns 2010).
20a. Recently recorded at Cerro Tacarcuna, Colombia (Ruiz-Ovalle & Hurtado-Guerra (2014).
21. Storer (1970a) merged Wetmorethraupis into Buthraupis, but this has not been followed by most subsequent authors; see Isler & Isler (1987). They are not sister genera (Sedano & Burns 2010).
21a. Sibley & Monroe (1990) considered Buthraupis eximia and B. aureodorsalis to form a superspecies. Sedano & Burns (2010) confirmed that they are sister species but also found that they are sister to Chlorornis riefferii. SACC proposal passed to remove from Buthraupis and to resurrect the genus Cnemathraupis for them.
22. Described since Meyer de Schauensee (1970): Blake & Hocking (1974).
22a. Buthraupis wetmorei was formerly (e.g., Hellmayr 1936) placed in a monotypic genus, Tephrophilus. Sedano & Burns (2010) showed that it was not a member of Buthraupis. Burns et al. (2016) recommended resurrecting Tephrophilus, as in Dickinson & Christidis (2014). SACC proposal pending to resurrect Tephrospilus.
23. Called "Santa Marta Mountain-Tanager" by Ridgely & Tudor (1989). SACC proposal to change English name did not pass.
23a. Anisognathus melanogenys and A. lacrymosus form a superspecies (Sibley & Monroe 1990); they were considered conspecific by Hellmayr (1936) and Storer (1970a). See Donegan and Avendaño (2010) for evidence supporting species rank for melanogenys. SACC proposal did not pass to treat melanogenys as a subspecies of lacrymosus.
23b. Anisognathus melanogenys, A. lacrymosus, and A. igniventris were formerly (e.g., Hellmayr 1936) placed in the genus Poecilothraupis, but see Meyer de Schauensee (1966).
23c. SACC proposal to change spelling of English name to "Lachrymose" did not pass.
24. Anisognathus somptuosus was formerly (e.g., Hellmayr 1936, Meyer de Schauensee 1970, Storer 1970a, Ridgely & Tudor 1989) known as A. flavinuchus (or flavinucha), but see Sibley & Monroe (1990). Hilty (2011) elevated the subspecies flavinucha to species rank, with the English name “Bolivian Mountain-Tanager.”
24a. Anisognathus somptuosus and A. notabilis were formerly (e.g., Hellmayr 1936, Zimmer 1944a, Phelps & Phelps 1950a) placed in the genus Compsocoma, but see Meyer de Schauensee (1966). Sedano & Burns (2010) found that their inclusion in Anisognathus was problematic. SACC proposals pending.
24b. Contrary to traditional linear sequences, genetic data (Burns et al. 2003, Burns & Naoki 2004, Sedano & Burns 2010) indicate that Chlorornis is a member of a group of Andean tanager genera that consists of Buthraupis, Anisognathus, Delothraupis, Dubusia, and Calochaetes (and presumably also Bangsia and Wetmorethraupis).
24c. The hyphenated English group-name “Mountain-Tanager” no longer refers to a monophyletic group (e.g. Burns et al. 2016), so the hyphens must be removed. SACC proposal needed. <>
25. The subspecies carrikeri of the Santa Marta Mountains, was described as a separate species from Dubusia taeniata, but was treated as a subspecies of the latter by Meyer de Schauensee (1966) and subsequent authors. Described as a separate species and treated as such by Chapman (1926), the southern subspecies stictocephala was treated as a subspecies of D. taeniata by Hellmayr (1936) and subsequent authors. Vocal differences between it and nominate taeniata are pronounced (Robbins et al., unpubl. data). SACC proposal to elevate stictocephala to species rank did not pass.
25a. Delothraupis was merged into Dubusia by Meyer de Schauensee (1966) and . Genetic data (Burns et al. 2003, Burns & Naoki 2004, Klicka et al. 2007, Sedano & Burns 2010) indicate that Delothraupis and Dubusia are sister taxa. SACC proposal passed to merge these two genera.
25b. Hellack and Schnell (1977, REFs) previously noted that “Saltator” rufiventris was an unusual saltator based on plumage and morphology. Genetic data (Klicka et al. 2007, Chaves et al. 2013) have revealed that “Saltator” rufiventris is definitely not a saltator but a tanager, closely related to Delothraupis and Dubusia. SACC proposal passed to move to Thraupidae. Because there is no other genus name available, the species is maintained here provisionally, as indicated by quotation marks, pending a proposal to move to Dubusia or naming of a new genus. Dickinson & Christidis (2014) transferred it to Dubusia and called it Rufous-bellied Mountain-Tanager. Burns et al. (2016) named a new genus for the species: Pseudosaltator. SACC proposal passed to recognize Pseudosaltator and to change English name to Rufous-bellied Mountain-Tanager.
26. Iridosornis porphyrocephalus and I. analis form a superspecies (Sibley & Monroe 1990); evidence for treating them as separate species is weak (Ridgely & Tudor 1989); they were considered conspecific by Hellmayr (1936). Orcés (1944) reported them to be sympatric in southern Ecuador, but the locality for I. porphyrocephalus is widely regarded as dubious (Ridgely & Tudor 1989, Paynter 1993, Ridgely & Greenfield 2001).
26b. Iridosornis is masculine, so the correct spelling of the species name is porphyrocephalus (David & Gosselin 2002b).
27. Iridosornis reinhardti was considered a subspecies of I. rufivertex by Hellmayr (1936), Zimmer (1944a), and Storer (1970a); they form a superspecies (Parker et al. 1985, Sibley & Monroe 1990).
27a. Although Chlorochrysa is traditionally placed next to Tangara in linear sequences (e.g., Meyer de Schauensee 1970), genetic data (Burns & Naoki 2004) indicate that they are not closely related.
28. Chlorochrysa phoenicotis and C. calliparaea form a superspecies (REF).
28a. Despite being one of the largest genera in the world in terms of species richness, genetic data (Burns and Naoki 2004) strongly support Tangara as a monophyletic group. The genus Tangara was formerly called Calospiza (e.g., Hellmayr 1936), and in literature from the 1800s, Calliste; see Zimmer (1943b) and Meyer de Schauensee (1966). Although Burns and Naoki (2004) found support for the monophyly of the genus, relationships within the genus were found to differ from those implied by traditional linear sequences. SACC proposal passed to change linear sequence. Sedano & Burns (2010), however, found that Tangara was monophyletic only if Thraupis was included. SACC proposal did not pass to modify generic limits. Burns et al. (2014) confirmed this, and Burns et al. (2016) recommended resurrection of several genera and described two new ones to partition Tangara into monophyletic groups; see various Notes 28ab, 28cc, and 36.
28aa. Isler & Isler (1987) proposed that Tangara inornata and T. mexicana were sister species based on similarities in sociality, feeding behavior, habitat, and voice. Genetic data (Burns & Naoki 2004) corroborate this relationship. SACC proposal passed to change linear sequence.
28ab. Isler & Isler (1987) proposed that Tangara palmeri and Middle American T. cabanisi were sister species based on similarities in plumage, habitat, and voice. Burns et al. (2014) confirmed this with genetic data. Burns et al. (2016) described a new genus, Poecilostreptus, for these two species. SACC proposal pending to recognize Poecilostreptus.
28b. Storer (1970a) considered Tangara cyanocephala and T. desmaresti to form a superspecies. Isler & Isler (1987) considered these two to form a species group with T. cyanoventris. Genetic data (Burns & Naoki 2004) indicate that Tangara cyanocephala and T. desmaresti are likely sister species (T. cyanoventris not sampled), and that they form a monophyletic group with T. seledon and T. fastuosa, which are also sister species.
28bb. Isler & Isler (1987) proposed that Tangara chilensis, T. seledon, and T. fastuosa were sister species based on similarities in plumage, behavior, habitat, biogeography, and voice. Genetic data (Burns & Naoki 2004), however, do not corroborate this relationship; see Notes 28b and 37b.
28bbb. Isler & Isler (1987) proposed that Tangara johannae, T. schrankii, T. florida, T. arthus, T. icterocephala, T. xanthocephala, T. chrysotis, and T. parzudakii form a species group based on similarities in plumage and foraging behavior. Genetic data (Burns & Naoki 2004) confirm this as a monophyletic group.
28bbbb. "Tangara gouldi," known only from the type specimens from southeastern Brazil and considered a species by Hellmayr (1936) and Pinto (1944), is considered a hybrid (T. cyanoventris X T. desmaresti) (Bond 1947, Meyer de Schauensee 1966, Storer 1970a). See Hybrids and Dubious Taxa.
28c. Hilty (2003) suspected that the subspecies phelpsi of the Tepuis might deserve recognition as a separate species from Tangara xanthogastra.
28cc. Isler & Isler (1987) proposed that Tangara xanthogastra, T. punctata, T. guttata, T. varia, and T. rufigula form a species group based on plumage and vocal similarities. Genetic data (Burns & Naoki 2004, Burns et al. 2014) confirm this as a monophyletic group. Burns et al. (2016) recommended restoration of Ixothraupis Bonaparte for these five species. SACC proposal pending to recognize Ixothraupis.
28d. Storer (1970a) considered Tangara gyrola and T. lavinia to form a superspecies. Genetic data (Burns & Naoki 2004) confirm this relationship. Isler & Isler (1987) suggested that T. rufigenis might also be part of this group based on plumage and habitat similarities.
28dd. As pointed out by Meyer de Schauensee (1966), the differences among subspecies groups currently treated included in T. gyrola seem to be at least of the same magnitude as those between T. gyrola and T. lavinia; indeed, the viridissima subspecies group and the bangsi subspecies group were formerly (e.g., Ridgway 1902, Chapman 1925) each treated as separate species from T. gyrola.
28e. The species named used for Tangara guttata was formerly (e.g., Hellmayr 1936, Phelps & Phelps 1950a) chrysophrys, but see Meyer de Schauensee (1966) and Storer (1970a).
28f. The subspecies brasiliensis was formerly (e.g., Hellmayr 1936) treated as a separate species from Tangara mexicana, but most classifications have followed Zimmer (1943c) in treated them as conspecific.
29. Some authors (e.g., Hellmayr 1936, Sick 1997) have suspected that Tangara peruviana and T. preciosa are actually color morphs of the same species, but they are largely parapatric with little or no evidence for interbreeding (Firme et al. 2007).
29a. The species named used for Tangara preciosa was formerly (e.g., Hellmayr 1936, Pinto 1944) castanonota, but see Meyer de Schauensee (1966) and Storer (1970a).
29b. The species named used for Tangara vitriolina was formerly (e.g., Hellmayr 1936) ruficapilla, but see Meyer de Schauensee (1966) and Storer (1970a).
30. Described since Meyer de Schauensee (1970): Schulenberg & Binford (1985).
30a. Isler & Isler (1987) proposed that Tangara cayana, T. peruviana, T. preciosa, T. meyerdeschauenseei, T. vitriolina, and West Indian T. cucullata form a species group based on similarities in plumage and habitat. Genetic data (Burns & Naoki 2004) corroborate the close relationship among Tangara cayana, T. vitriolina, T. cucullata, and T. meyerdeschauenseei (T. peruviana and T. preciosa not sampled).
30aa. "Tangara arnaulti," known from one aviary specimen and reluctantly considered a valid species by Hellmayr (1936) and Meyer de Schauensee (1966); presumably a hybrid (T. preciosa X T. cayana) (Hellmayr 1936, Bond 1951a, Meyer de Schauensee 1966, 1970, Storer 1970a, Ingels 1971). See Hybrids and Dubious Taxa.
30b. The southern subspecies fulvicervix was formerly (e.g., Berlepsch 1912) considered a separate species from Tangara ruficervix, a treatment that will almost certainly be restored; they were treated as conspecific by Hellmayr (1936) and subsequent authors; they differ in more plumage features than do many pairs of Tangara treated as separate species, yet they show limited genetic divergence (Burns & Naoki 2004).
30c. Isler & Isler (1987) proposed that Tangara ruficervix, T. labradorides, and T. cyanotis formed a species group based on plumage similarities. Genetic data (Burns & Naoki 2004) do not confirm this relationship; these three species do not cluster within any of the other species groups in the genus.
31. Tangara larvata was formerly (e.g., Hellmayr 1936, Zimmer 1943b, Meyer de Schauensee 1966, 1970) considered a subspecies of T. nigrocincta, but most classifications have generally followed Eisenmann (1957) in treating it as a separate species; see Wetmore et al. (1984) for rationale. Sibley & Monroe (1990) considered them to form a superspecies. Genetic data (Burns & Naoki 2004) show that T. larvata is actually more closely related to T. cyanicollis than either is to T. nigrocincta, thus confirming treatment of T. nigrocincta as a separate species from T. larvata and negating treatment of T. larvata and T. nigrocincta as members of superspecies.
31a. Formerly known as "Golden-masked Tanager" (e.g., AOU 1983), but see .
32. Described since Meyer de Schauensee (1970): Graves & Weske (1987).
32a. Isler & Isler (1987) proposed that Tangara vassorii and T. nigroviridis formed a species group with Central American T. dowii and T. fucosa. Genetic data (Burns & Naoki 2004) confirm their close relationship, with T. vassorii basal to the other three,
33. Graves & Weske (1987) proposed that Tangara heinei, T. phillipsi, T. argyrofenges, and T. viridicollis formed a monophyletic group. Genetic data (Burns & Naoki 2004) corroborate the close relationship of Tangara heinei, T. argyrofenges, and T. viridicollis. Sibley & Monroe (1990) proposed that T. heinei and T. phillipsi formed a superspecies.
34. Called "Silver-backed Tanager" by Isler & Isler (1987), Ridgely & Tudor (1989), Sibley & Monroe (1990), and Ridgely & Greenfield (2001). SACC proposal to change English name did not pass.
35. Called "Straw-backed Tanager" by Isler & Isler (1987), Ridgely & Tudor (1989), Sibley & Monroe (1990), Ridgely & Greenfield (2001), and Hilty (2011). SACC proposal to change English name did not pass.
36. Genetic data (Burns & Naoki 2004) indicate that Tangara cyanoptera is the sister to the T. heinei species complex (see Note 33), thus corroborating its traditional position in linear sequences. Burns et al. (2014) found that these two, plus T. larvata, T. nigrocincta, T. cyanicollis, T. preciosa, T. peruana, T. meyerdeschauenseei, T. vitriolina, T. cucullatus, T. cayana, T. viridicollis, T. phillipsi, and T. argyrofenges, formed a monophyletic group not included in Tangara s.s.; Burns et al. (2016) described a new genus, Stilpnia, for this group. SACC proposal pending to recognize Stilpnia.
36a. Ridgely & Tudor (1989) and Hilty (2003) suspected that the subspecies whitelyi of the Tepui region might deserve treatment as a separate species from Tangara cyanoptera.
37. Ridgely & Tudor (1989) suspected that the subspecies cyanomelas of the Atlantic forest region of Brazil deserves consideration as a separate species status from Tangara velia.
37a. Tangara velia and T. callophrys were formerly (e.g., Hellmayr 1936, Zimmer 1943a, Pinto 1944, Phelps & Phelps 1950a) placed in a separate genus, Tanagrella; similarities in plumage and bill shape have suggested to most authors (e.g., Isler & Isler 1987) that T. velia and T. callophrys are sister species. Genetic data (Burns & Naoki 2004) confirm this relationship and also indicate that T. chilensis is the sister species to these two.
38. Tersina was formerly placed in its own monotypic family, Tersinidae (e.g., Hellmayr 1936, Tordoff 1954a, <>George 1962, Meyer de Schauensee 1970), or subfamily or tribe with the tanagers (e.g., Tersininae as in Storer 1970a, or Tersinini as in ); in all these treatments it was considered the sister taxon to all other tanagers. Genetic data (Bledsoe 1988, Burns 1997, Burns et al. 2002, 2003) as well as morphological data (Raikow 1978), however, indicate that it is embedded within the tanagers, and most closely related to Dacnis and Cyanerpes (Burns et al. 2003).
39. The genera Dacnis and Cyanerpes were formerly (e.g., Meyer de Schauensee 1970, Fjeldså & Krabbe 1990) considered members of a separate family, the Coerebidae, but they were considered to be tanagers on the basis of skull structure by Tordoff (1954a) and were placed in the Thraupidae by Storer (1969, 1970a). This has been followed in most subsequent classifications. Molecular data indicate that they are sister genera, but embedded within the tanagers, forming a group with Tersina (Burns et al. 2002, 2003).
39a. Dacnis albiventris was formerly (e.g., Hellmayr 1935) placed in a monotypic genus, Hemidacnis, but Zimmer (1942d) merged this into Dacnis.
40. Ridgely & Tudor (1989) pointed out the trans-Andean egregia group may deserve species rank. Ridgely et al. (2001) considered egregia a species separate from lineata based on plumage differences and disjunct range. SACC proposal to recognize Dacnis egregia as a separate species did not pass because of insufficient published data.
41. Dacnis hartlaubi was formerly (e.g., Hellmayr 1936, Meyer de Schauensee 1966, 1970, Sibley & Monroe 1990) placed in a monotypic genus, Pseudodacnis and included in the Thraupidae (as "Turquoise Dacnis-Tanager") when other Dacnis were placed in the Coerebidae. Storer (1969, 1970a) merged this genus into Dacnis, and subsequent classifications have followed this treatment.
41a. Cyanerpes lucidus and C. caeruleus form a superspecies (AOU 1983, Sibley & Monroe 1990); they were considered conspecific by Hellmayr (1935), but they are sympatric in northwestern Colombia.
42. The genus Chlorophanes was formerly (e.g., Meyer de Schauensee 1970) considered a member of a separate family, the Coerebidae, but Tordoff (1954a) considered it a tanager on the basis of skull morphology and Storer (1969, 1970a) placed it in the Thraupidae. Genetic data indicate that it is embedded within the tanagers; molecular data do not support a close relationship to Dacnis or Cyanerpes, but its closest relatives remain uncertain (Burns et al. 2002, 2003).