The sapayoa or broad-billed sapayoa (''Sapayoa aenigma'') is a

suboscine The Tyranni (suboscines) are a suborder of passerine birds that includes more than 1,000 species, the large majority of which are South American. It is named after the type genus '' Tyrannus''. These have a different anatomy of the syrinx mus ...

passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...

found in lowland rainforests in Panama and north-western South America. As the epithet ''aenigma'' ("the

enigma Enigma may refer to: *Riddle, someone or something that is mysterious or puzzling Biology *ENIGMA, a class of gene in the LIM domain Computing and technology * Enigma (company), a New York-based data-technology startup * Enigma machine, a family ...

") implies, its relationships have long been elusive. It is easy to overlook, but appears to be common in a wide range and is not considered threatened by the IUCN.

Taxonomy and systematics

The sapayoa was formally described by the German ornithologist Ernst Hartert in 1903 under the present binomial name ''Sapayoa aenigma''. It has always been considered a

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...

genus, ''Sapayoa'', and historically regarded as a New World

; in particular, it was assigned to the manakin family (

Pipridae The manakins are a family, Pipridae, of small suboscine passerine birds. The group contains some 54 species distributed through the American tropics. The name is from Middle Dutch ''mannekijn'' "little man" (also the source of the different bird ...

). However, the

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

was listed as ''

incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...

'' (position uncertain) in the Sibley-Ahlquist taxonomy, because

"preliminary DNA-DNA hybridization comparisons ... indicate that this species is either a relative of the Old World
Eurylaimidae The Eurylaimidae are a family of suboscine passerine birds that occur from the eastern Himalayas to Indonesia and the Philippines. The family previously included the sapayoa from the Neotropics, the asities from Madagascar, and the Calyptomeni ...
or a sister group of all other Tyrannida, as suggested by earlier biochemical studies .... In any event, it is not a close relative of manakins or any other recent tyrannoid."

More recent research suggests that it is not a New World suboscine at all, but an Old World suboscine. In 2004, it was shown that the sapayoa is an outlier to the New World suboscines. In an earlier analysis based on

nDNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. I ...

myoglobin intron 2 and

GAPDH Glyceraldehyde 3-phosphate dehydrogenase (abbreviated GAPDH) () is an enzyme of about 37kDa that catalyzes the sixth step of glycolysis and thus serves to break down glucose for energy and carbon molecules. In addition to this long establishe ...

intron 11

sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is calle ...

data, the authors found the sapayoa

"as a deep branch in the group of
broadbill Broadbill may refer to the bird families: *the Eurylaimidae, a family of birds known as the ''Asian and Grauer’s broadbills'' *the Calyptomenidae, a family of birds known as the ''African and green broadbills'' Broadbill may also refer to: *Bro ...
s and
pitta Pittas are a family, Pittidae, of passerine birds found in Asia, Australasia and Africa. There are thought to be 40 to 42 species of pittas, all similar in general appearance and habits. The pittas are Old World suboscines, and their closest ...
s of the Old World tropics."Fjeldså et al. (2003)

Accordingly, the sapayoa would be the last surviving New World species of a lineage that evolved in

Australia-New Guinea The continent of Australia, sometimes known in technical contexts by the names Sahul (), Australia-New Guinea, Australinea, Meganesia, or Papualand to distinguish it from the country of Australia, is located within the Southern and East ...

when Gondwana was in the process of splitting apart. The sapayoa's ancestors are hypothesized to have reached South America via the

Western Antarctica West Antarctica, or Lesser Antarctica, one of the two major regions of Antarctica, is the part of that continent that lies within the Western Hemisphere, and includes the Antarctic Peninsula. It is separated from East Antarctica by the Transa ...

Peninsula. Nowadays, the sapoyoa is sometimes placed in the family

Eurylaimidae The Eurylaimidae are a family of suboscine passerine birds that occur from the eastern Himalayas to Indonesia and the Philippines. The family previously included the sapayoa from the Neotropics, the asities from Madagascar, and the Calyptomeni ...

with the broadbills. Others tentatively place the sapayoa in the

asity The asities are a family of birds, Philepittidae, that are endemic to Madagascar. The asities consist of four species in two genera. The ''Neodrepanis'' species are known as sunbird-asities and were formerly known as false sunbirds.del Hoyo, J. ...

family Philepittidae otherwise found only in Madagascar and sometimes included in the broadbill family. However, the divergence between the broadbills and the sapayoa found in the 2003 study is only slightly less deep than that between the sapayoa and the pittas. It is even possible, though unlikely, that the present species is actually closer to the pittas than to the broadbills. Consequently, it is now placed in its own monotypic

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

, Sapayoidae.

Description

The sapayoa is a small, olive-colored bird, somewhat paler below and with a yellowish throat. Its habitus resembles a bigger, longer-tailed, broader-billed female

manakin The manakins are a family, Pipridae, of small suboscine passerine birds. The group contains some 54 species distributed through the American tropics. The name is from Middle Dutch ''mannekijn'' "little man" (also the source of the different bird ...

. It is rare to uncommon in the

forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...

understory, favoring ravines and small streams.Ridgely & Tudor (1994) p.689, plate 46. It is usually seen in pairs or mixed-species flocks. It spends long periods perching, then sallies up to pick fruit or catch insects, on foliage or in mid air, with its flat, wide bill in a way reminiscent of flatbills. The sapayoa builds a nest suspended from a branch usually above a stream. It is a pear-shaped structure with the larger end at the top and fibres hanging beneath. The entrance is at the side.

Breeding behaviour

Sapayoa was previously believed to be monogamous. However, a study observed sapayoas engaging in cooperative breeding.

Footnotes

References

* Banks, Richard C.; Chesser, R. Terry; Cicero, Carla; Dunn, Jon L.; Kratter, Andrew W.; Lovette, Irby J.; Rasmussen, Pamela C.; Remsen, J. V. Jr.; Rising, James D.; Stotz, Douglas F. & Winker, Kevin (2008). Forty-ninth supplement to the American Ornithologists' Union ''Check-list of North American Birds''. ''The Auk'' 125(3): 758–768. PDF fulltext
* Chesser, R. Terry (2004). Molecular systematics of New World suboscine birds. '' Mol. Phylogenet. Evol.'' 32(1): 11–24. PDF fulltext
* Fjeldså, Jon; Zuccon, Dario; Irestedt, Martin; Johansson, Ulf S. & Ericson, Per G.P. (2003). ''Sapayoa aenigma'': a New World representative of 'Old World suboscines'. '' Proc. R. Soc. B'' 270(Supplement 2): 238–241. PDF fulltextElectronic supplement
* Irestedt, M.; Ohlson, J.I.; Zuccon, Dario; Källersjö, M. & Ericson, Per G.P. (2006). Nuclear DNA from old collections of avian study skins reveals the evolutionary history of the Old World suboscines (Aves: Passeriformes). '' Zool. Scripta'' 35(6): 567–580. PDF fulltext
* Kemp, Alan & Sherley, Greg H. (2003). Asities. ''In:'' Perrins, Christopher (ed.): ''Firefly Encyclopedia of Birds'' p. 421. Firefly Books. * Remsen, J. V., Jr.; Cadena, C. D.; Jaramillo, A.; Nores, M.; Pacheco, J. F. Robbins, M. B.; Schulenberg, T. S., Stiles, F. G.; Stotz, D. F.; & Zimmer, K. J. Version (2009). A classification of the bird species of South America. American Ornithologists' Union
HTML full text
* Ridgely, Robert S. & Tudor, Guy (1994). ''The Birds of South America'' (Volume 2: The suboscine passerines). University of Texas Press, Austin. * Sibley, Charles Gald & Monroe, Burt L. Jr. (1990). ''Distribution and taxonomy of the birds of the world: A Study in Molecular Evolution''. Yale University Press, New Haven, CT.

External links

Don Roberson's Bird Families of the World

Avibase The World Bird Database
{{Taxonbar, from=Q1090032 Birds of Colombia Birds of the Tumbes-Chocó-Magdalena Birds described in 1903