Darwin's finches (also known as the Galápagos Finches or as Geospizinae) are a group of 14 or 15 species of Passerine birds, now placed in the tanager family rather than the true finch family. They were first collected by Charles Darwin on the Galápagos Islands during the second voyage of the Beagle. Thirteen are found on the Galápagos Islands and one on Cocos Island. The term Darwin's Finches was first applied by Percy Lowe in 1936, and popularised in 1947 by David Lack in his book Darwin's Finches.^[1][2]

The birds are all about the same size (10–20 cm). The most important differences between species are in the size and shape of their beaks, and the beaks are highly adapted to different food sources. The birds are all dull-colored.

Darwin's theory

During the survey voyage of HMS Beagle, Darwin had no idea of the significance of the birds of the Galápagos. He had learned how to preserve bird specimens while at the University of Edinburgh and had been keen on shooting, but he had no expertise in ornithology and by this stage of the voyage concentrated mainly on geology and mostly left bird shooting to his servant Syms Covington.^[3] Nonetheless, these birds were to play an important part in the inception of Darwin's theory of evolution by natural selection.

On the Galápagos Islands and afterward, Darwin thought in terms of "centres of creation" and rejected ideas of transmutation of species.^[4] From Henslow's teaching he was interested in geographical distribution of species, particularly links between species on oceanic islands and on nearby continents. On Chatham Island he recorded that a mockingbird was similar to those he had seen in Chile, and after finding a different one on Charles Island he carefully noted where mockingbirds had been caught, but paid little attention to the finches. When writing up his notes on the way to Tahiti Darwin was astonished to find that all the mockingbirds caught on Charles Island were of one species, those from Albemarle of another, and those from James and Chatham Islands of a third species. As they sailed home about nine months later this, together with other facts including what he'd heard about Galápagos tortoises, made him wonder about the stability of species.^[5][6]

Following his return from the voyage, Darwin presented the finches to the Geological Society of London at their meeting on 4 January 1837, along with other mammal and bird specimens he had collected. The bird specimens, including the finches, were given to John Gould, the famous English ornithologist, for identification. Gould set aside his paying work and at the next meeting on 10 January reported that birds from the Galápagos Islands which Darwin had thought were blackbirds, "gross-beaks" and finches were in fact "a series of ground Finches which are so peculiar [as to form] an entirely new group, containing 12 species." This story made the newspapers.^[7][8]

Darwin had been in Cambridge at that time. In early March he met Gould again and for the first time got a full report on the findings, including the point that his Galápagos "wren" was another closely allied species of finch. The mockingbirds Darwin had labelled by island were separate species rather than just varieties. Gould found more species than Darwin had anticipated,^[9] and concluded that 25 of the 26 land birds were new and distinct forms, found nowhere else in the world but closely allied to those found on the South American continent.^[8] Darwin now saw that if the finch species were confined to individual islands, like the mockingbirds, this would help to account for the number of species on the islands, and he sought information from others on the expedition. Specimens had also been collected by Captain Robert FitzRoy, FitzRoy’s steward Harry Fuller and Darwin's servant Covington, who had labelled them by island.^[10] From these, Darwin tried to reconstruct the locations where he had collected his own specimens. The conclusions led shortly afterwards to his conversion to the idea of transmutation of species.^[8]

Text from the Voyage of the Beagle

At the time he rewrote his diary for publication as Journal and Remarks (later The Voyage of the Beagle), he described Gould's findings on the number of birds, noting that "Although the species are thus peculiar to the archipelago, yet nearly all in their general structure, habits, colour of feathers, and even tone of voice, are strictly American".^[11] In the first edition of The Voyage of the Beagle Darwin said that "It is very remarkable that a nearly perfect gradation of structure in this one group can be traced in the form of the beak, from one exceeding in dimensions that of the largest gros-beak, to another differing but little from that of a warbler".^[12]

In 1839 Darwin conceived of his theory of natural selection, and by the time of the second edition in 1845 Darwin had brought together his theory. He now added two closing sentences: "Seeing this gradation and diversity of structure in one small, intimately related group of birds, one might really fancy that from an original paucity of birds in this archipelago, one species had been taken and modified for different ends".^[13][14]

The remaining land-birds form a most singular group of finches, related to each other in the structure of their beaks, short tails, form of body and plumage: there are thirteen species, which Mr. Gould has divided into four subgroups. All these species are peculiar to this archipelago; and so is the whole group, with the exception of one species of the sub-group Cactornis, lately brought from Bow Island, in the Low Archipelago. Of Cactornis, the two species may be often seen climbing about the flowers of the great cactus-trees; but all the other species of this group of finches, mingled together in flocks, feed on the dry and sterile ground of the lower districts. The males of all, or certainly of the greater number, are jet black; and the females (with perhaps one or two exceptions) are brown. The most curious fact is the perfect gradation in the size of the beaks in the different species of Geospiza, from one as large as that of a hawfinch to that of a chaffinch, and (if Mr. Gould is right in including his sub-group, Certhidea, in the main group) even to that of a warbler. The largest beak in the genus Geospiza is shown in Fig. 1, and the smallest in Fig. 3; but instead of there being only one intermediate species, with a beak of the size shown in Fig. 2, there are no less than six species with insensibly graduated beaks. The beak of the sub-group Certhidea, is shown in Fig. 4. The beak of Cactornis is somewhat like that of a starling, and that of the fourth subgroup, Camarhynchus, is slightly parrot-shaped. Seeing this gradation and diversity of structure in one small, intimately related group of birds, one might really fancy that from an original paucity of birds in this archipelago, one species had been taken and modified for different ends. In a like manner it might be fancied that a bird originally a buzzard, had been induced here to undertake the office of the carrion-feeding Polybori of the American continent.^[15]

Text from the Origin of Species

Darwin discussed the divergence of species of birds in the Galápagos more explicitly in his chapter on geographical distribution in On the Origin of Species:

The most striking and important fact for us in regard to the inhabitants of islands, is their affinity to those of the nearest mainland, without being actually the same species. [In] the Galapagos Archipelago... almost every product of the land and water bears the unmistakable stamp of the American continent. There are twenty-six land birds, and twenty-five of these are ranked by Mr. Gould as distinct species, supposed to have been created here; yet the close affinity of most of these birds to American species in every character, in their habits, gestures, and tones of voice, was manifest.... The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, yet feels that he is standing on American land. Why should this be so? why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plain a stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which resembles closely the conditions of the South American coast: in fact there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in climate, height, and size of the islands, between the Galapagos and Cape de Verde Archipelagos: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape de Verde Islands are related to those of Africa, like those of the Galapagos to America. I believe this grand fact can receive no sort of explanation on the ordinary view of independent creation; whereas on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists, whether by occasional means of transport or by formerly continuous land, from America; and the Cape de Verde Islands from Africa; and that such colonists would be liable to modification;—the principle of inheritance still betraying their original birthplace.^[16]

Polymorphism in Darwin's finches

Whereas Darwin spent just five weeks in the Galápagos, and David Lack spent three months, Peter and Rosemary Grant and their colleagues have made research trips to the Galápagos for about thirty years, particularly studying Darwin's finches. Here we look briefly at the case of the large cactus finch Geospiza conirostris on Isla Genovesa (formerly Tower Island) which is formed from a shield volcano, and is home to a variety of birds. These birds, like all well-studied groups,^[17] show various kinds of morphism.

Males are dimorphic in song type: songs A and B are quite distinct. Also, males with song A have shorter bills than B males. This is also a clear difference. With these beaks males are able to feed differently on their favourite cactus, the prickly pear Opuntia. Those with long beaks are able to punch holes in the cactus fruit and eat the fleshy aril pulp which surrounds the seeds, whereas those with shorter beaks tear apart the cactus base and eat the pulp and any insect larvae and pupae (both groups eat flowers and buds). This dimorphism clearly maximises their feeding opportunities during the non-breeding season when food is scarce.

If the population is panmixic,^[18][19] then Geospiza conirostris exhibits a balanced genetic polymorphism and not, as originally supposed, a case of nascent sympatric speciation. The selection maintaining the polymorphism maximises the species' niche by expanded its feeding opportunity. The genetics of this situation cannot be clarified in the absence of a detailed breeding program, but two loci with linkage disequilibrium^[20] is a possibility.

Another interesting dimorphism is for the bills of young finches, which are either 'pink' or 'yellow'. All species of Darwin's finches exhibit this morphism, which lasts for two months. No interpretation of this phenomenon is known.^[21]

Taxonomy

Family

For some decades taxonomists have placed these birds in the family Emberizidae with the New World sparrows and Old World buntings (Sulloway 1982). However, the Sibley-Ahlquist taxonomy puts Darwin's finches with the tanagers (Monroe and Sibley 1993), and at least one recent work follows that example (Burns and Skutch 2003). The American Ornithologists' Union, in its North American check-list, places the Cocos Island Finch in the Emberizidae but with an asterisk indicating that the placement is probably wrong (AOU 1998–2006); in its tentative South American check-list, the Galápagos species are incertae sedis, of uncertain place (Remsen et al. 2007).

Species

• Genus Geospiza
  • Large Cactus-finch, Geospiza conirostris
  • Sharp-beaked Ground-finch, Geospiza difficilis
    • Vampire Finch, Geospiza difficilis septentrionalis
  • Medium Ground-finch, Geospiza fortis
  • Small Ground-finch, Geospiza fuliginosa
  • Large Ground-finch, Geospiza magnirostris
    • Darwin's Large Ground-finch, Geospiza magnirostris magnirostris - possibly extinct (1957?)
  • Common Cactus-finch, Geospiza scandens
• Genus Camarhynchus
  • Vegetarian Finch, Camarhynchus crassirostris - sometimes separated in Platyspiza
  • Large Tree-finch, Camarhynchus psittacula
  • Medium Tree-finch, Camarhynchus pauper
  • Small Tree-finch, Camarhynchus parvulus
  • Woodpecker Finch, Camarhynchus pallidus - sometimes separated in Cactospiza
  • Mangrove Finch, Camarhynchus heliobates
• Genus Certhidea
  • Warbler Finch, Certhidea olivacea
• Genus Pinaroloxias
  • Cocos Island Finch, Pinaroloxias inornata

Molecular basis of beak evolution

Developmental research in 2004 found that bone morphogenetic protein 4 (BMP4), and its differential expression during development, resulted in variation of beak size and shape among finches. BMP4 acts in the developing embryo to lay down skeletal features, including the beak.^[22] The same group showed that the different beak shapes of Darwin's finches develop are also influenced by slightly different timing and spatial expression of a gene called calmodulin (CaM).^[23] Calmodulin acts in a similar way to BMP4, affecting some of the features of beak growth. The authors suggest that changes in the temporal and spatial expression of these two factors are possible developmental controls of beak morphology.

Notes

References

• Darwin, Charles (1839), Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Journal and remarks. 1832-1836, III, London: Henry Colburn .
• Darwin, Charles (1845), Journal of researches into the natural history and geology of the countries visited during the voyage of H.M.S. Beagle round the world, under the Command of Capt. Fitz Roy, R.N. 2d edition, London: John Murray .
• Darwin, Charles (1859), On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (1st ed.), London: John Murray, http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text&pageseq=1 
• Darwin, Francis (1887), "Chapter 1, The Foundations of the 'Origin of Species'", in Darwin, Francis, The life and letters of Charles Darwin, including an autobiographical chapter, 2, London: John Murray, http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F1452.2&pageseq=21 
• Desmond, Adrian; Moore, James (1991), Darwin, London: Michael Joseph, Penguin Group, ISBN 0-7181-3430-3, OCLC 185764721 
• Eldredge, Niles (2006), "Confessions of a Darwinist", The Virginia Quarterly Review (Spring 2006): 32–53, http://www.vqronline.org/articles/2006/spring/eldredge-confessions-darwinist/, retrieved 2008-11-04 
• Lack, David (1940), "Evolution of the Galapagos Finches" (^{[dead link]}), Nature 146 (146): 324–327, 07 September 1940, doi:10.1038/146324a0, http://www.nature.com/nature/journal/v146/n3697/abs/146324a0.html, retrieved 2008-12-09 
• Check-list of North American Birds, American Ornithologists' Union, 1998–2006, http://www.aou.org/checklist/index.php3, retrieved 2007-04-09 
• Kevin J. Burns and Alexander F. Skutch (2003), "Tanagers and Tanager-Finches", in Christopher Perrins, ed., The Firefly Encyclopedia of Birds, Firefly Books, pp. 629–631, ISBN 1-55297-777-3, http://www.amazon.com/Firefly-Encyclopedia-Birds-Christopher-Perrins/dp/1552977773/ref=si3_rdr_bb_product/103-5704731-0011011, retrieved 2007-04-09  It is not clear whether this placement was made by Burns and Skutch or by Perrins.
• Keynes, Richard (2000), Charles Darwin’s zoology notes & specimen lists from H.M.S. Beagle., Cambridge University Press, http://darwin-online.org.uk/content/frameset?itemID=F1840&viewtype=text&pageseq=1, retrieved 2008-12-08 
• Burt L. Monroe and Charles G. Sibley, A World Checklist of Birds. New Haven, Conn.: Yale University Press (1993). ISBN 0-300-07083-7. Accessed 2007-04-09. Monroe and Sibley consider the tanagers to be a tribe (Thraupini) of a big family Fringillidae rather than a family of their own (Thraupidae).
• J. V. Remsen, Jr., C. D. Cadena, A. Jaramillo, M. Nores, J. F. Pacheco, M. B. Robbins, T. S. Schulenberg, F. G. Stiles, D. F. Stotz, and K. J. Zimmer. [Version 2007-04-05.] A classification of the bird species of South America. American Ornithologists' Union. Accessed 2007-04-09.
• Steinheimer, F. D. (2004), "Charles Darwin's bird collection and ornithological knowledge during the voyage of H.M.S. Beagle, 1831-1836", Journal of Ornithology 145 (145(4)): 300–320, doi:10.1007/s10336-004-0043-8, http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=A161&pageseq=1, retrieved 2008-12-08 
• Sulloway, Frank J. (1982), "The Beagle collections of Darwin’s finches (Geospizinae)", Bulletin of the British Museum (Natural History), Historical Series (43, No. 2): 49–94, http://darwin-online.org.uk/content/frameset?itemID=A86&viewtype=image&pageseq=1, retrieved 2008-12-08 
• Sulloway, Frank J. (Spring 1982), "Darwin and His Finches: The Evolution of a Legend" (PDF), Journal of the History of Biology 15 (1): 1–53, doi:10.1007/BF00132004, http://www.sulloway.org/Finches.pdf, retrieved 2008-12-09 
• Sulloway, Frank J. (2006), "Why Darwin Rejected Intelligent Design", in Brockman, John, Intelligent Thought: Science versus the Intelligent Design Movement, New York: Vintage, pp. 107–126, http://www.sulloway.org/Why%20Darwin%20Rejected%20Intelligent%20Design%20(2006).pdf, retrieved 2008-12-08 

The tanagers (sg. pronounced /ˈtænədʒər/) are a family, Thraupidae, of birds in the order Passeriformes. The family has an American distribution.

There were traditionally about 240 species of tanagers, but the taxonomic treatment of this family's members is currently in a state of flux. As more of these birds are studied using modern molecular techniques it is expected that some genera may be relocated elsewhere. Already the Euphonias and chlorophonias, which were once considered part of the tanager family, are now treated as members of Fringillidae, in their own subfamily (Euphoniinae). Likewise the genera Piranga (which includes the Scarlet Tanager, Summer Tanager, and Western Tanager), Chlorothraupis, and Habia appear to be members of the Cardinal family^[1], and have been reassigned to that family by the AOU.

Description

Tanagers are small to medium-sized birds. The shortest-bodied species, the White-eared Conebill, is 9 cm (3.8 in) long and weighs 7 grams, barely smaller than the Short-billed Honeycreeper. The longest, the Magpie Tanager is 28 cm (11 in) and weighs 76 grams (2.7 oz). The heaviest is the White-capped Tanager which weighs 114 grams (4 oz) and measures about 23 cm (8.7 in). Both sexes are usually the same size and weight. Tanagers are often brightly colored, but some species are black and white. Birds in their first year are often duller or a different color altogether. Males are typically more brightly coloured than females.

Most tanagers have short, rounded wings. The shape of the bill seems to be linked to the species' foraging habits.

Distribution

Tanagers are restricted to the New World and mainly to the tropics. About 60% of tanagers live in South America, and 30% of these species live in the Andes. Most species are endemic to a relatively small area.

Behaviour

Most tanagers live in pairs or in small groups of 3-5 individuals. These groups may consist simply of parents and their offspring. Birds may also be seen in single species or mixed flocks. Many tanagers are thought to have dull songs, though some are elaborate.

Diet

Tanagers are omnivorous, and their diet varies from genus to genus. They have been seen eating fruits, seeds, nectar, flower parts and insects. Many pick insects off branches. Other species look for insects on the underside of leaves. Yet others wait on branches until they see a flying insect and catch it in the air. Many of these particular species inhabit the same areas, but these specializations alleviate competition.

Reproduction

The breeding season begin in March through until June in temperate areas and in September through October in South America. Some species are territorial while others build their nests closer together. There is little information on tanager breeding behavior or whether they are monogamous or polygamous. Males show off their brightest feathers to potential mates and rival males. Some species' courtship rituals involve bowing and tail lifting.

Most tanagers build cup nests on branches in trees. Some nests are almost globular. Entrances are usually built on the side of the nest. The nests can be shallow or deep. The species of the tree they choose to build their nest in and the nest's position varies among genera. Most species nest in an area hidden by very dense vegetation. There is still no information on the nests of some species.

The clutch size is 3–5 eggs. The female incubates the eggs and builds the nest, but the male may feed the female while she incubates. Both sexes feed the young. Five species have helpers assist in feeding the young. These helpers are thought to be the previous year's nestlings.

Systematics

Phylogenetic studies suggest the true tanagers form three main groups two of which consist of several smaller, well-supported clades.^[2] The list below is an attempt using information gleaned from the latest studies to organise them into coherent related groups, and as such may contain groupings not yet accepted by or are under review by the various ornithological taxonomy authorities.^[3]

Group 1

Mainly dull-coloured forms

Slaty Finch, Haplospiza rustica

a) Conebill and flowerpiercer group (Also contains Haplospiza, Catamenia, Acanthidops, Diglossa, Diglossopis, Phrygilus and Sicalis^[4] traditionally in the Emberizidae)^[5] This group despite having a rather varied bill morphology shows marked plumage similarities. Most are largely grey, blue, or black, and numerous have rufous on the underparts:

• Genus Conirostrum – typical conebills (10 species)
• Genus Oreomanes – Giant Conebill
• Genus Xenodacnis – Tit-like Dacnis
• Genus Catamenia (3 species)
• Genus Diglossa – typical flowerpiercers (14 species)
• Genus Diglossopis – blue flowerpiercers (4 species)
• Genus Haplospiza (2 species). Paraphyletic with 2 species of sierra-finch Phrygilus^[6]
• Genus Acanthidops – Peg-billed Finch
• Genus Phrygilus - sierra-finches (11 species)^[7]
• Genus Sicalis – yellow-finches (12 species). Paraphyletic with Phrygilus^[8]

b) True seedeaters. Traditionally placed in Emberizidae. These genera share a particular foot-scute pattern which suggests that they may form a monophyletic group^[9]:

Male Variable Seedeater, Sporophila corvina

• Genus Sporophila – typical seedeaters (some 55 species)
• Genus Oryzoborus (6 species)^[10]
• Genus Dolospingus – White-naped Seedeater^[11]
• Genus Charitospiza – Coal-crested Finch

c) "Yellow-rumped" clade^[12]:

• Genus Heterospingus (2 species)
• Genus Chrysothlypis (2 species)
• Genus Hemithraupis (3 species)

Brazilian Tanager, Ramphocelus bresilius

d) "Crested" clade (Also contains Coryphospingus & Volatinia traditionally placed in the Emberizidae):

• Genus Ramphocelus – silver-billed tanagers (9 species)
• Genus Lanio – shrike-tanagers (4 species)
• Genus Eucometis – Gray-headed Tanager
• Genus Tachyphonus (8 species)
• Genus Trichothraupis – Black-goggled Tanager
• Genus Stephanophorus – Diademed Tanager
• Genus Coryphospingus (2 species)
• Genus Volatinia – Blue-black Grassquit

e) "Blue Finch" clade. Relationships within Thraupidae uncertain but may be related to "Poospiza" clade^[13]:

• Genus Porphyrospiza - Blue Finch^[14]
• Genus? Phrygilus alaudinus^[15] - Band-tailed Sierra-finch

Orange-headed Tanager, Thlypopsis sordida

f) The "Poospiza" clade - a diverse but close-knit group containing both warbler and finch-like forms:

• Genus Poospiza – Warbling-finches (15 species)^[16]
• Genus Compsospiza - Mountain-finches (2 species)
• Genus Cnemoscopus – Gray-hooded Bush Tanager
• Genus Hemispingus – hemispinguses (12 species)
• Genus Thlypopsis (6 species)
• Genus Pyrrhocoma – Chestnut-headed Tanager
• Genus Cypsnagra – White-rumped Tanager
• Genus Nephelornis – Pardusco

g) Grass & Pampa-finches. Relationships within Thraupidae uncertain but together form a well-supported clade^[8]:

• Genus Emberizoides (3 species)
• Genus Embernagra (2 species)

Male Yellow-bridled Finch, Melanodera xanthogramma

h) A miscellaneous and likely polyphyletic group of unplaced "tanager-finches" (which may or may not include the species called Tanager-finch) whose members when studied will no doubt be relocated to other clades:

• Genus Melanodera (2 species)
• Genus Rowettia – Gough Island Finch
• Genus Nesospiza (2 species)
• Genus Gubernatrix – Yellow Cardinal
• Genus Idiopsar – Short-tailed Finch
• Genus Piezorhina – Cinereous Finch
• Genus Xenospingus – Slender-billed Finch
• Genus Incaspiza – inca-finches (5 species)
• Genus Coryphaspiza – Black-masked Finch
• Genus Rhodospingus – Crimson-breasted Finch
• Genus Donacospiza – Long-tailed Reed-finch (may be related to Poospiza^[17])

i) Basal forms in group 1:

• Genus Conothraupis (2 species)
• Genus Orchesticus – Brown Tanager
• Genus Creurgops (2 species)

Group 2

"Typical" colourful Tanagers

Diversity of Darwin's finches

a) Tropical canopy tanagers:

• Genus Thraupis - T. abbas & episcopus at least^[18]
• Genus Tangara (about 50 species)

b) The "Tholospiza" - Darwin's finches, grassquits, atypical honeycreepers and some seedeaters.^[19] The finch-like forms in this clade were formerly classified in the Emberizidae:

• Genus Geospiza – ground finches (6 species)
• Genus Camarhynchus – tree finches (6 species)
• Genus Certhidea – Warbler Finch
• Genus Pinaroloxias – Cocos Island Finch
• Genus Melopyrrha – Cuban Bullfinch
• Genus Coereba – Bananaquit. Formerly placed in own family Coerebidae^[20]
• Genus Tiaris – grassquits (5 species) - polyphyletic
• Genus Loxipasser – Yellow-shouldered Grassquit
• Genus Euneornis – Orangequit
• Genus Melanospiza – St. Lucia Black Finch
• Genus Loxigilla – Antillean bullfinches (3 species) - polyphyletic

Green-and-gold Tanager, Tangara schrankii

c) Mountain tanagers:

• Genus Cyanicterus – Blue-backed Tanager
• Genus Bangsia – (5 species)
• Genus Buthraupis – (4 species)
• Genus Chlorornis – Grass-green Tanager
• Genus Wetmorethraupis – Orange-throated Tanager
• Genus Anisognathus – (5 species)
• Genus Dubusia – Buff-breasted Mountain-tanager
• Genus Delothraupis – Chestnut-bellied Mountain-tanager
• Genus? Saltator rufiventris - Rufous-bellied 'Saltator'^[21]

Blue-gray Tanager, Thraupis episcopus

d) Typical tanagers:

• Genus Thraupis - Thraupis bonariensis at least belongs here
• Genus Pipraeidea – Fawn-breasted Tanager
• Genus Iridosornis (5 species)

e) Typical multicoloured tanagers (includes Paroaria traditionally placed in either Emberizidae or Cardinalidae):

• Genus Diuca (2 species)
• Genus Lophospingus (2 species)
• Genus Neothraupis – White-banded Tanager
• Genus Cissopis – Magpie Tanager
• Genus Paroaria (5–6 species)
• Genus Schistochlamys (2 species)

f) Green & Golden-collared Honeycreepers^[22]:

• Genus Chlorophanes – Green Honeycreeper
• Genus Iridophanes – Golden-collared Honeycreeper

g) Typical honeycreepers and relatives^[23]:

• Genus Tersina – Swallow Tanager
• Genus Cyanerpes, the typical honeycreepers (4 species)
• Genus Pseudodacnis – Turquoise Dacnis-tanager
• Genus Dacnis, the dacnises (8 species)

h) Basal lineages within group 2:

Green Honeycreeper, Chlorophanes spiza

• Genus Chlorochrysa (3 species)
• Genus Parkerthraustes – Yellow-shouldered Grosbeak (traditionally in Cardinalidae, but biochemical evidence^[24] suggests it is a tanager)
• Genus Nemosia – (2 species)
• Genus Compsothraupis – Scarlet-throated Tanager
• Genus Sericossypha – White-capped Tanager

Group 3

Saltators

• Genus Saltator (16 species; traditionally placed in Cardinalidae, but biochemical evidence suggests they may be tanagers or a sister group^[24])
• Genus Saltatricula – Many-colored Chaco-finch. Traditionally placed in the Emberizidae but may be related to one of the Saltators^[25][26]

Thraupidae incertae sedis

• Genus Mitrospingus (2 species)
• Genus Orthogonys – Olive-green Tanager
• Genus Calochaetes – Vermilion Tanager
• Genus Catamblyrhynchus – Plushcap or Plush-capped Finch
• Genus Oreothraupis – Tanager-finch^[27]
• Genus Urothraupis – Black-backed Bush-tanager
• Genus Rhodinocichla – Rosy Thrush-tanager
• Genus Lamprospiza – Red-billed Pied Tanager
• Genus Phaenicophilus – palm-tanagers (2 species)
• Genus Xenoligea – White-winged warbler
• Genus Microligea – Green-tailed warbler
• Genus Calyptophilus – chat-tanagers (2 species)
• Genus Nesospingus – Puerto Rican Tanager

The Western Tanager (Piranga ludoviciana) seems to be closer to cardinals

Recently split from Thraupidae

Related to Arremonops and other American sparrows in Emberizidae:

• Genus Chlorospingus – bush-tanagers (around 10 species)

Related to the cardinals in Cardinalidae^[28]:

• Genus Piranga – northern tanagers (9 species)
• Genus Habia – ant-tanagers or habias (5 species)
• Genus Chlorothraupis (3 species)
• Genus Amaurospiza (4 species; apparently very close to Cyanocompsa)

Fringillinae, subfamily Euphoniinae:

• Genus Euphonia (over 25 species)
• Genus Chlorophonia (5 species)

Exact affinities uncertain, but lie outside the tanagers:

• Genus Spindalis – spindalises (4 species).

Footnotes

References

• Bent, A. Life Histories of Blackbirds, Orioles, Tanagers, and Allies. New York:Dover Publications:1965. 549 p.
• Burns, K. J., S. J. Hackett, and N. K. Klein. 2002. Phylogenetic relationships and morphological diversity in Darwin's finches and their relatives. Evolution 56: 1240-1252.
• Burns, K. J., S. J. Hackett, and N. K. Klein. 2003. Phylogenetic relationships of Neotropical honeycreepers and the evolution of feeding morphology. J. Avian Biology 34: 360-370.
• Clark, G. A., JR. 1986. Systematic interpretations of foot-scute patterns of Neotropical finches. Wilson Bull. 98: 594-597.
• Fjeldså J. and Rahbek C. (2006). Diversification of tanagers, a species rich bird group, largely follows lowlands to montane regions of South America. Integrative and Comparative Biology 46(1):72-81. Download - http://intl-icb.oxfordjournals.org/cgi/reprint/46/1/72.
• Greeney, H. 2005. Nest and eggs of the Yellow-whiskered Bush Tanager in Eastern Ecuador. Ornitologia Neotropical 16: 437- 438.
• Hellmayr, C. E. 1935. Catalogue of birds of the Americas and the adjacent islands in Field Museum of Natural History. Fieldiana Zoology v.13, pt.8. - for "Coerebidae". (Download available at http://www.archive.org/details/catalogueofbirds138hell)
• Hellmayr, C. E. 1936. Catalogue of birds of the Americas and the adjacent islands in Field Museum of Natural History. Fieldiana Zoology v.13, pt.9. Tersinidae - Thraupidae. (Download available at http://www.archive.org/details/catalogueofbirds139hell)
• Hellmayr, C. E. 1938. Catalogue of birds of the Americas and the adjacent islands in Field Museum of Natural History. Fieldiana Zoology v.13, pt.11. Ploceidae - Catamblyrhynchidae - Fringillidae. (Download available at http://www.archive.org/details/catalogueofbirdso1311hell)
• Infonatura. 2005 June. Birds, mammals, and amphibians of Latin America Accessed 2006 March 4.
• Isler M. Isler P. The Tanagers a Natural History, Distribution, and Identification. Washington D.C.: Smithsonian Institution Press: 1987. 404 p.
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• Latta, S. 2006. et al. Aves de la República Dominicana y Haití. Princeton University Press.

• Lijtmaer, D. A., N. M. Sharpe, P. L. Tubaro & S. C. Lougheed. 2004. Molecular phylogenetics and diversification of the genus Sporophila (Aves: Passeriformes). Mol. Philo. Evol. 33:562-579.
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• Sato, A., C. O'Huigin, F. Figueroa, P. R. Grant, B. R. Grant, H. Tichy, and J. Klein. 1999. Phylogeny of Darwin's finches as revealed by mtDNA sequences. Proc. Nat. Acad. Sci. 96: 5101-5106.
• Webster, J.D. & Webster, J.R. 1999. Skeletons and the genera. of sparrows (Emberizinae). Auk 116: 1054–1074.
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