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Cladus: Eukaryota
Supergroup: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Aves
Subclassis: Carinatae
Infraclassis: Neornithes
Parvclassis: Neognathae
Ordo: Passeriformes
Subordo: Passeri
Parvordo: Passerida
Superfamilia: Passeroidea
Familia: Thraupidae
Genera: Anisognathus - Bangsia - Buthraupis - Calochaetes - Calyptophilus - Catamblyrhynchus - Chlorochrysa - Chlorophanes - Chlorophonia - Chlorornis - Chlorospingus - Chlorothraupis - Chrysothlypis - Cissopis - Cnemoscopus - Compsothraupis - Conirostrum - Conothraupis - Creurgops - Cyanerpes - Cyanicterus - Cypsnagra - Dacnis - Delothraupis - Dubusia - Eucometis - Euphonia - Habia - Hemispingus - Hemithraupis - Heterospingus - Iridophanes - Iridosornis - Lamprospiza - Lanio - Mitrospingus - Nemosia - Neothraupis - Nephelornis - Nesospingus - Orchesticus - Oreomanes - Oreothraupis - Orthogonys - Phaenicophilus - Pipraeidea - Piranga - Pseudodacnis - Pyrrhocoma - Ramphocelus - Rhodinocichla - Schistochlamys - Sericossypha - Spindalis - Stephanophorus - Tachyphonus - Tangara - Tersina - Thlypopsis - Thraupis - Trichothraupis - Urothraupis - Wetmorethraupis - Xenodacnis

Name

Thraupidae

The tanagers (sg. pronounced /ˈtænədʒər/) comprise the bird family Thraupidae, 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 species in the genera Euphonia and Chlorophonia, 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 24 cm (9.5 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)

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

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)


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 (5 species)
* Genus Platyspiza - Vegetarian Finch
* 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

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]

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:


* 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)
* Genus Spindalis – spindalises (4 species).

Fringillidae, subfamily Euphoniinae:

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

it] Footnotes

1. ^ Yuri & Mindell (2002)
2. ^ Fjeldså & Rahbek (2006) & Klicka et al. (2007)
3. ^ See http://www.museum.lsu.edu/~Remsen/SACCBaseline.html
4. ^ Burns et al. (2003) & Klicka et al. (2007)
5. ^ See Webster & Webster (1999). If the presence of a free lacrimal bone as found in Haplospiza, Acanthidops, and two of the three Catamenias has any phylogenetic significance then this clade may also include several other "tanager-finches" that share this feature
6. ^ Klicka (2007)
7. ^ Webster & Webster (1999) & Klicka et al. (2007). Probably polyphyletic
8. ^ a b Klicka et al. (2007)
9. ^ Clark (1986)
10. ^ See Lijtmaer et al. (2004) & Robbins et al. (2005). Polyphyletic. Members of this genus are paraphyletic with various members of Sporophila
11. ^ See Robbins et al. (2005). This species is nested within a group containing both Sporophila and Oryzoborus
12. ^ Burns et al. (2003)
13. ^ (See below: Group 1f)
14. ^ Klicka et al. (2007). This species formerly placed near Passerina in the Cardinalidae is related to Phrygilus alaudinus a tanager-finch
15. ^ http://www.zoonomen.net/cit/RI/Genera/RIGenAZ.html
16. ^ Klicka et al. (2007). This genus is very likely polyphyletic within its clade
17. ^ Ridgely & Tudor (1989) p.472
18. ^ Klicka et al. (2007). Some members of this genus paraphyletic with respect to certain Tangara
19. ^ See Burns et al. (2002) for the circumscription of this group the "domed nest clade" or "Tholospiza".
20. ^ See Burns et al. (2002). Exact affinities uncertain but probably sister species to Tiaris olivacea in the "Tholospiza"
21. ^ See http://www.museum.lsu.edu/~Remsen/SACCBaseline10.html Apparently close to mountain-tanagers Dubusia and Delothraupis
22. ^ See Burns et al. (2003) for close relationship of these species
23. ^ See Burns et al. (2003), Klicka et al. (2007) - may be closer to group 1
24. ^ a b Klicka & Spellman (2007)
25. ^ See http://www.museum.lsu.edu/~Remsen/SACCBaseline10.html
26. ^ Klicka et al. (2007). Apparently closest to Saltator atricollis and this species may require moving to Saltatricula
27. ^ See http://www.museum.lsu.edu/~Remsen/SACCBaseline10.html. May be related to the emberizine genus Atlapetes
28. ^ See http://www.museum.lsu.edu/~Remsen/SACCBaseline11.html


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.
* Klicka, J., K. Burns, & G. M. Spellman. 2007. Defining a monophyletic Cardinalini: A molecular perspective. Molecular Phylogenetics and Evolution 45: 1014-1032
* 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.
* Lougheed, S. C., J. R. Freeland, P. Handford, & I. T. Boag. 2000. A molecular phylogeny of warbling-finches (Poospiza): paraphyly in a Neotropical emberizid genus. Molecular Phylogenetics and Evolution 17: 367-378.
* Montereybay. 2000 July. 6-11. Tanagers: Thraupidae Accessed 2006 March 4.
* Naoki, K. 2003. Evolution of Ecological Diversity in the Neotropical Tanagers of the Genus Tangara (Aves: Thraupidae). Dissertation available online, given to Louisiana State University.
* Ridgely, R. S., & G. Tudor. 1989. The Birds of South America, vol. 1. Univ. Texas Press, Austin.
* Robbins, M. B., M. J. Braun, C. J. Huddleston, D. W. Finch, & C. M. Milensky (2005). First Guyana records, natural history, and systematics of the White-winged Seedeater (Dolospingus fringilloides). Ibis 147:334-341.
* 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.
* Yuri, T., and D. P. Mindell. 2002. Molecular phylogenetic analysis of Fringillidae, "New World nine-primaried oscines" (Aves: Passeriformes) Mol. Phylogen. Evol. 23:229-243.

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