• 378 species in the Neotropics
• DR personal total: 217 species (57%), 38 photo'd

These aren't your father's Tanagers any more.

The Thraupidae page that was here in 2000 began: "No one is quite sure what makes a 'tanager' right now, but as a Supreme Court justice once said about pornography: 'I may not be able to define it, but I know it when I see it.' Tanagers may be hard to define, but usually you know one when you see one. They are just drop-dead gorgeous birds of the Neotropics."

Well . . . wrong. Sort-of. Some of the "drop-dead gorgeous birds" in the Neotropics are Tanagers, including those in the speciose genus Tangara, like Silver-throated Tanager (above). And a good many are traditional tanagers, such as White-lined Tanager (female, left). But a whole lot of the birds that appear genetically to be tanagers go by different names. The full story may surprise you. It is based on genetic evidence published within the decade. [The most recent information is in Burns et al. (2014). That has not yet been widely incorporated into this page. An update is needed.]

What about the Western Tanager, which I featured on my previous version of this family? Not a tanager — it belongs, to the Cardinalidae [Cardinals, Grosbeaks & allies]. Same goes for Scarlet Tanager, Summer Tanager, and Hepatic Tanager. Not tanagers. The sought-after ant-tanagers of the tropical undergrowth? Not tanagers. The bright and colorful chlorophonias and euphonias? Not tanagers.

So what is a tanager? It's still hard to define, but the genetic evidence is trickling in, and a lot of plain-colored birds appear to tanagers. Darwin's finches on the Galapagos — such as Small Ground-Finch (right) — that may be a tanager. A seedeater on a tropic fenceline — like the male Variable Seedeater (below) — that may be a tanager. The grassquits, Bananaquit, and the saltators — they are all tanagers. These aren't your father's Tanagers any more.

The genetic evidence is impressive; e.g., Burns et al. (2002, 2003), Burns & Naoki (2004), Yuri & Mindell (2002), Klicka et al. (2000, 2007). Workers in the past had paid too much attention to bill shape, and not enough to biology, and convergent evolution masked many relationships (Isler & Isler 1987). Thraupidae is now an entirely Neotropical family whose members are essentially non-migratory. Although little has yet been made official in new checklists, tomorrow's Thraupidae should be a huge family of very diverse Neotropical birds. With ~378 species, it is among the largest bird families on earth. The "old" Thraupidae had ~ 242 species (Isler & Isler 1987). Included now are many birds bearing the name "tanager," but also many called finch, conebill, seedeater, dacnis, or honeycreeper. Purple Honeyeater (left) is a widespread example of the latter.

The genetic evidence divides the true tanagers into two groups. One group includes the original genus Thraupis, and its 9 species, including the very widespread and abundant Blue-gray Tanager (right).

This is also the group that includes the honeycreepers, dacnises, grassquits, seedeaters, Caribbean bullfinches, and Galapagos finches. Among the surprises is that the various "cardinals" of South America, in the genus Paroaria, fit into this set of tanagers. One of those is the widespread Red-capped Cardinal (below, a cool shot by Greg Lasley).

Among this set of tanagers is the genus Tangara (~50 species) are generally bright, gaudy, and travel in mixed-species flocks. Combining great loveliness and rarity is Azure-rumped Tanager (left in a fantastic shot by Lou Jost). It is limited to a narrow elevational range in the mountains of Chiapas, Mexico, and adjacent western Guatemala.

It is the Andes where the Tangara tanagers really shine. One encounters mixed-species flocks which include a dozen or more species, some with such alliterative names as Beryl-spangled Tanager T. nigroviridis, Spangle-cheeked Tanager T. dowii, or Flame-faced Tanager T. parzudakii. The foothills have birds like Bay-headed Tanager (below left). The Amazonian lowlands have fewer species, but the widespread Paradise Tanager T. chilensis is as gaudy as they come.

Some genetic data indicate that the Galapagos finches — represented here by Woodpecker Finch (right) — are embedded within the Thraupidae as part of a group that includes Yellow-faced Grassquit (above left), Bananaquit (above right), and several Caribbean genera (Burns et al. 2002, 2003, Yuri and Mindell 2002, Klicka et al. 2000, 2007).

The relationships of all these species have long been controversial. The grassquit is in genus Tiaris, but Tiaris itself is paraphyletic, with Yellow-faced Grassquit, the type species of the genus, not being not particularly close to other "Tiaris." Grassquits had once been placed with the cardinalines, and then with the emberizines based on skeletal morphology (e.g., AOU 1957). Now they are all tanagers; Burns et al. (2014).

The Galapagos finches were once placed in a separate subfamily [Geospizinae] from other sparrows and finches in the family Emberizidae. Their origins have been a point of contention back to the days of Darwin. They are often called "Darwin's Finches" because of their mythologized influence on his theory of evolution (in truth Darwin did not pay much attention to the finches; he was more impressed with variation in mockingbirds and tortoises). They have themselves been the subject of extensive study, including the amazing work of Peter & Rosemary Grant (Weiner 1994).

Until recently, the relationships of Bananaquit remained unresolved; some thought it a parulid warbler but many have considered it a monotypic family (e.g., AOU 1998) or subfamily (Ridgely & Tudor 1989). Bledsoe (1988) included it within the Thraupidae; Sibley & Monroe (1990) also thought it was a tanager (which they considered to be just a tribe Thraupini of a huge Fringillidae family). [It is possible that someday scientists will lump all the families near the end of the passerines — tanagers, emberizids, finches, New World warblers, icterids, cardinals & grosbeaks — into a single humongous family on the grounds that they all evolved comparatively recently, in contrast to longer-evolved families, but let's hope it doesn't go that far!] The SACC even considered Bananaquit a family until 2007. It is now clearly a tanager.

There is a second major group in the "new" Thraupidae. A basal set in this other group are 9 species of brightly-patterned Ramphocelus tanagers. I've heard that, spectrographically, the intense red on male Ramphocelus is the brightest red in the bird world.

Passerini's Tanager (left) is one of them, and there is strong sexual dimorphism (male above, female below). This used to be called 'Scarlet-rumped Tanager.' While males look alike throughout its range, the females in western Costa Rica are decidedly different. 'Scarlet-rumped' has recently been split into Passerini's (the widespread bird of Atlantic lowlands from Mexico to Panama) and Cherrie's (on the Pacific slope of Costa Rica).

Another Ramphocelus is the gorgeous burgundy-velvet Silver-beaked Tanager (male; below left).

A number of undergrowth skulkers are in this set of tanagers, including the "crested" tanagers in genus Tachyphonus. These are birds like Flame-crested Tanager T. cristatus and Fulvous-crested Tanager T. surinamus, and non-crested species such as White-lined Tanager (male, above right; the female was shown at the top of this family page). The "white line" is actually a bit of the underwing coverts visible in normal views (above right).

Gray-headed Tanager (right) is in its own genus; it joins flocks of other species in the lowland undergrowth. Shrike-tanagers, genus Lanio, are placed here. So are the honeycreepers — represented here by Shining Honeycreeper (below left) — plus conebills (genus Conirostrum), Giant Conebill (Oreomanes), and Tit-like Dacnis (Xenodacnis).

This second set of tanagers includes numerous anomalous species. Slender-billed Finch (above center) is has a thin, bright yellow, finch-like bill; it is restricted to arid scrub in coastal south Peru and north Chile. The Plushcap (above right) is restricted to bamboo thickets high in the Andes. It is so atypical that for many years it was considered to be in its own family ["Catamblyrhynchidae"]. The Pardusco Nephelornis onelli is an oddity found only in isolated elfin forests in central Peru. Tanager-Finch Oreothraupis arremonops likes lush Pacific slope cloud forests. Hemispingus is a genus of tanagers [but Chlorospingus bush-tanagers are emberizids].

Also here are some 14 species of Diglossa flowerpiercers, represented here by Slaty Flowerpiercer (female right; males are black). The strange upturned, hook-tipped bill is used to pierce the base of flowers to get at the nectar there.

Orange-throated Tanager (left © Dan Singer, among the few to photo this rarity) was only discovered to science in 1964 (Lowery & O'Neill 1964); the fascinating story of this "bird without a name" is told by Stap (1991). Wetmorethraupis is a monotypic genus and remains rare and hard to find near the Rio Marañón, and restricted to a narrow altitudinal range (600-800m) in mature forest in northern Peru and southernmost Ecuador. The list of local and endemic species among the tanagers is quite long.

Thraupidae is such a huge family that we can barely skim the surface here. I've gotten this far without mentioning the enigmatic Swallow-Tanager Tersina viridis, once thought to be in its own monotypic family ["Tersinidae"]. There are also many special tanagers that are exceptionally rare and elusive. Cherry-throated Tanager Nemosia rourei had previously been known only from a specimen in southeast Brazil from 1870 and one report 47 years ago, but was just recently rediscovered (you can read about this find, and see photos, in Pacheco 1998).

For a time there was mystery as to the placement of saltators. They have long been considered with cardinals and grosbeaks, but genetic evidence shows this was wrong. Burns et al. (2014) determined they were solidly "tanagers." They found that 15 of the 16 species of Saltator formed a distinctive clade, worthy of subfamily status, and named the clade Saltatorinae. This subfamily includes the very widespread Buff-throated Saltator (below). The exception was Rufous-bellied Saltator "Saltator" rufiventris of the eastern Andes of southern Bolivia and extreme northern Argentina. It is apparently close to mountain-tanagers, but needs a new genera. [As of this update, the paper proposing generic changes is still unpublished.]

It is my hope that the ornithologists don't tinker too much with the name "tanager." We have birds in many different families that are called "chat," or "finch," or "redstart." Just because Slender-billed Finch is now a tanager does not mean it is wise to revise the English name and cause literary confusion of years. I can live with some small changes here or there — "Stripe-headed Tanager" became Stripe-headed Spindalis without much problem — and, in fact, the genus Spindalis is now in different family, the Phoenicophilidae — but my plea would be to avoid wholesale English name changes in the quest for the "perfect" name. Yet, in the end, no matter how a "tanager" eventually comes to be defined or named, these are fascinating and often colorful birds of the Neotropics. They, with hummingbirds, parrots, and toucans, are a core component of any Neotropical adventure.

I have another page focused mostly on tanagers endemic to coastal Brazil. Someday some of these photos should be moved over to this discusion.

Photos: The Silver-throated Tanager Tangara icterocepala was at Mirador Cinchona, Costa Rica, on 18 Dec 2007. The female White-lined Tanager Tachyphonus rufus was at Asa Wright Nature Reserve, Trinidad, on 24 Dec 2006, as was the male shown farther down the page. The female Small Ground-Finch Geospiza fuliginosa was on Santa Cruz I., Galapagos, on 25 Sep 1989. The male Variable Seedeater Sporophila aurita was near Puerto Viejo, Costa Rica, on 17 Dec 2007. The male Purple Honeyeater Cyanerpes caeruleus was at Asa Wright, Trinidad, on 25 Dec 2006. The Blue-gray Tanager Thraupis episcopus was at Puerto Viejo, Costa Rica, on 17 Dec 2007. Greg Lasley photographed the bathing Red-capped Cardinal Paroaria gularis at Hato el Cedros, Venezuela, on 25 Apr 1992. The wonderful berry-eating Azure-rumped Tanager Tangara cabanisi was photographed by Lou Jost at Canyon Hondo, below El Triunfo, Chiapas, Mexico, in 1986. The acrobatic Bay-headed Tanager Tangara gyrola was on the Osa Peninsula, Costa Rica, on 26 Dec 2007. The Blue-winged Mountain-Tanager Anisognathus somptuosus was foraging with a canopy flock below a ridge on the Nono-Mindo Road, Ecuador, in April 1992. The Yellow-faced Grassquit Tiaris olivacea was on Cozumel Is., Mexico, on Sep . The Bananaquit Coereba flaveola was at Asa Wright, Trinidad, on 24 Dec 2006. The Woodpecker Finch Camarhynchus pallidus was on Santa Cruz I., Galapagos, on 25 Sep 1989. The pair of Passerini's Tanager Ramphocelus passerinii was at Puerto Viejo, Costa Rica, on 19 Dec 2007. The male Silver-beaked Tanager R. carbo was at Asa Wright, Trinidad, on 25 Dec 2006. The Gray-headed Tanager Eucometis penicillata was in a mixed flock at Bosque del Rio Tigre, Osa Peninsula, Costa Rica, on 25 Dec 2007. The Shining Honeycreeper Cyanerpes lucidus was at Puerto Viejo, Costa Rica, on 19 Dec 2007. The Plushcap Catamblyrhynchus diadema was in bamboo below Abra Malaga, Dept. Cuzco, Peru, on 17 June 1987. The Slaty Flowerpiercer Diglossa plumbea was at Savegre, Costa Rica, on 23 Dec 2007. Dan Singer photographed the Orange-throated Tanager Wetmorethraupis sterrhopteron on 23 Sep 2005 at Nuevo Salem, Peru. The Buff-throated Saltator Saltator maximus was at Puerto Viejo, Costa Rica, on 20 Dec 2007. All photos © Don Roberson, except those credited © Lou Jost, © Greg W. Lasley, and © Dan Singer, and used with permission; all rights reserved.

Bibliographic essay
Family Book:
Isler, M. L., and P. R. Isler. 1987. The Tanagers: Natural History, Distribution, and Identification. Smithsonian Institution Press, Washington, D. C.

Writing in 1987, when the DNA-DNA hybridization studies by Sibley & Ahlquist (1990) were just advance rumors, Isler & Isler wrote this about tanagers, which they had already reduced them to a subfamily, the Thraupinae:

"Tanagers belong to a very large group of birds known as nine-primaried oscines (or songbirds)... Characteristics have never been found that distinguish clearly among the subgroups of New World nine-primaried oscines commonly known as wood-warblers, tanagers, cardinals, and buntings (citations omitted). Despite the unclear boundaries, bill shape and presumed feeding behavior have been used to define families or subfamilies. Wood-warblers generally have slender pointed bills and feed mostly on insects, tanagers have thicker bills (some with a notch on the upper mandible) and eat mostly fruit, and New World finches have the stoutest bills and take mostly seeds. . . . Bill form and feeding habits appear less and less to be a good reflection of phylogenetic relationships among the New World nine-primaried songbirds. Recent studies have shown that bill form can change quite rapidly in evolutionary time [and] feeding behaviors within each of the three groups . . . have been found to be highly variable. . . . Despite the increasing certainty that tanagers are not presently definable as a monophyletic group we employ the rather traditional taxonomic arrangement . . . . It will take many years to reassess the relationships among the New World nine-primaried songbirds, and in the meantime, it is convenient and appropriate to use a well-known existing taxonomy. Even within the confines of this taxonomy, tanagers exhibit a remarkable variety of morphology and behavior."

The Islers were prescient. Things would change, and yet the book remains solid. It is not in any of the "regular" series of family books, which makes a refreshing change. It is more thorough and scholarly than most family volumes in the past two decades. Indeed, reading the acknowledgements is like reading the "all-star" line-up for Neotropical ornithology: Steve Hilty, the late & great Ted Parker, and Bob Ridgely "were the principal reviewers," and Gary Graves, John O'Neill, Van Remsen & Tom Schulenberg "reviewed large sections of the book." This is possibly as authoritative as any recent similar effort for any family in the world, and for that the reader is intensively grateful.

I especially like the distribution maps which show all the major rivers of the Neotropics so one can easily get a feel for the range of the species (e.g., either within a watershed or many, or restricted to mountains between watersheds). Maps focus on either South or Central America, depending on the species involved. Besides the maps, the discussions of subspecific variation is good; details of vocalizations are excellent; and the habitat, behavior, and natural history information is outstanding. The plates — in field guide style by Morton Isler — are accurate, even if not particularly evocative.

All that having been said, there are a few criticisms. First, the book should probably have been called "The Tanagers of the Neotropics" because the three primarily North American breeding species (Western, Scarlet, Summer) are treated as an afterthought. This proved accurate in the long run — these would prove to be in the Cardinalidae — but here are no breeding range maps for these, and the map for Hepatic Tanager cuts off at mid-Mexico with the phrase "extends to southeastern U.S." [sic]. This typographical mistake (obviously, southwestern U.S. was intended) is illustrative of the [lack of] care given to birds north of the border. The primary sources cited tend to be the Bent series (from 1956!) and earlier. Secondly, the plates, albeit accurate enough for i.d. purposes, do not capture the breath-taking beauty of these birds. The Glistening-green Tanager Chorochrysa phoenicotis, for example, just knocks your socks off in the field! [See spectacular photos in Dunning 1970]. Here, it is a dull green bird.

Yet these drawbacks detract little from the outstanding text. For the real scoop on the 'traditional' tanagers this is the book. Also see the recently-published chapter in the Handbook of Birds of the World project.

Literature cited:

American Ornithologists' Union. 1957. Check-List of North American Birds. 5th ed. A.O.U., Washington, D. C.

American Ornithologists' Union. 1998. Check-List of North American Birds. 7th ed. A.O.U., Washington, D. C.

Barker, F.K., K.J. Burns, J. Klicka, S.M. Lanyon, and I.J. Lovette. 2013. Going to extremes: contrasting rates of diversification in a recent radiation of New World passerine birds. Syst. Biol. 62: 298-320.

Bledsoe, A. H. 1988. Nuclear DNA evolution and phylogeny of the New World nine-primaried oscines. Auk 105: 504-515.

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.

Burns, K.J., and K. Naoki. 2004. Molecular phylogenetics and biogeography of Neotropical tanagers in the genus Tangara. Molec. Phylog. Evol. 32: 838-854.

Burns, K.J., A.J. Schultz, P.O. Title, N.A. Mason, F.K. Barker, J. Klicka, S.M. Lanyon, and I.J. Lovette. 2014. Phylogenetics and diversification of tanagers (Passeriformes: Thraupidae), the largest radiation of Neotropical songbirds. Molec. Phylo. Evol. 75: 41-77.

Dunning, J. S. 1970. Portraits of Tropical Birds. Livingston Publ. Co., Wynnewood, PA.

Klicka, J., K. Burns, and G.M. Spellman. 2007. Defining a monophyletic Cardinalini: A molecular perspective. Molec. Phylog. Evol. 45: 1014 1032.

Klicka, J., K. P. Johnson, and S. M. Lanyon. 2000. New World nine-primaried oscine relationships: constructing a mitochondrial DNA framework. Auk 117: 321-336.

Lowery, G. H., Jr., and J. P. O'Neill. 1964. A new genus and species of tanager from Peru. Auk 81: 125-131.

Pacheco, J. F. 1998. Cherry-throated Tanager Nemosia rourei rediscovered. Cotinga 9: 41.

Ridgely, R. S., and G. Tudor. 1989. The Birds of South America. Vol. 1: The Oscine Passerines. Univ of Texas, Austin.

Sibley, C. G., and J. E. Ahlquist. 1990. Phylogeny and Classification of Birds: A Study in Molecular Evolution. Yale Univ. Press, New Haven, CT.

Sibley, C. G., and B. L. Monroe. 1990. Distribution and Taxonomy of Birds of the World. Yale Univ. Press, New Haven, CT.

Stap, D. 1991. A Parrot without a Name: The Search for the Last Unknown Birds on Earth. Alfred A. Knopf, New York.

Weiner, Jonathan. 1994. The Beak of the Finch: A Story of Evolution in Our Time. Knopf, New York.

Yuri, T., and D.P. Mindell. 2002. Molecular phylogenetic analysis of Fringillidae, "New World nine-primaried oscines" (Aves: Passeriformes). Molec. Phylog. Evol. 23: 229-243.