name = Passerines
Early Eocene(Wangerripian) to Recent
image_width = 250px
House Sparrow("Passer domesticus")
ordo = Passeriformes
ordo_authority = Linnaeus, 1758
diversity_link = #Taxonomic list of Passeriformes families
diversity = Roughly 100 families, around 5,400 species
type_species = "Fringilla domestica"
type_species_authority = Linnaeus, 1758
*Passeriand see text
A passerine is a
birdof the order Passeriformes, which includes more than half of all bird species. Sometimes known as perching birds or, less accurately, as songbirds, the passerines form one of the most diverse terrestrial vertebrateorders: with around 5,093 species[ [http://elibrary.unm.edu/sora/Auk/v063n01/p0064-p0069.pdf Ernst Mayr, "The Number of Species of Birds", "The Auk," Volume 63, Number 1 (January, 1946), p.67] ] , it is roughly twice as diverse as the largest of the mammalorders, the Rodentia.
The names "passerines" and "Passeriformes" are derived from "Passer domesticus", the
scientific nameof the type species– the House Sparrow– and ultimately from the Latinterm "passer" for true sparrows and similar small birds.
Origin and evolution
The evolutionary history of and relationships among the passerine families remained rather mysterious until around the end of the 20th century. Many passerine families were grouped together on the basis of morphological similarities that, it is now believed, are the result of
convergent evolution, not a close genetic relationship. For example, the " wrens" of the northern hemisphere, those of Australia, and those of New Zealandlook very similar and behave in similar ways, and yet belong to three far-flung branches of the passerine family tree; they are as unrelated as it is possible to be while remaining Passeriformes.
Much research remains to be done, but advances in
molecular biologyand improved paleobiogeographical data are gradually revealing a clearer picture of passerine origins and evolution. It is now thought that the first passerines evolved in Gondwanaat some time in the Paleogene, maybe around the Late Paleocenesome 60–55 million years ago(mya)Fact|date=April 2008. The initial split was between the Tyranni, the songbirds, the Eurylaimidesand the New Zealand "wrens", which must have diverged during a short period of time (some million years at most). The Passeriformes apparently evolved out of a fairly close-knit cladeof " near passerines" which contains such birds as the Piciformes, Coraciiformes, and Cuculiformes. [Johansson & Ericson (2003)]
A little later, a great radiation of forms took place out of
Australia-New Guinea: the Passerior songbirds. A major branch of the Passeri, "Parvorder Passerida", emerged either as the sister group to the basal lineages and corvoids ("Parvorder Corvida"), or more likely as a subgroup of it, and expanded deep into Eurasiaand Africa, where there was a further explosive radiation of new lineages. This eventually led to three major passeridan lineages comprising about 4,000 species, which in addition to the corvoidan clade and numerous minor lineages make up songbird diversity today. There has been extensive biogeographicalmixing, with northern forms returning to the south, southern forms moving north, and so on.
Earliest passerines Perching bird
osteology, especially of the limb bones, is rather diagnostic. [See e.g. Boles (1997), Manegold "et al." (2004), Mayr & Manegold (2006)] However, the early fossil record is poor because the first Passeriformes were apparently on the small side of the present size range, and their delicate bones did not preserve well. QM specimens F20688 ( carpometacarpus) and F24685 ( tibiotarsus) from Murgon, Queenslandare fossil bone fragments clearly recognizable as passeriform; they represent two species of approximately some 10 and some 20 cm in overall length and prove that some 55 mya, barely into the Early Eocene, early perching birds were recognizably distinct. [Boles (1997)] A quite similar group, the Zygodactylidae(named for their zygodactylous approach to perching) independently arose at much the same time – and possibly from closely related ancestors – in the landmasses bordering the North Atlantic, which at that time was only some two-thirds of its present width.
Until the discovery of the Australian fossils, it was believed for some time that "
Palaeospiza bella" from the Priabonian Florissant Fossil Beds( Late Eocene, around 35 mya) was the oldest known passeriform. However, it is now considered a non-passeriform near passerine.
Bathans Formationat the Manuherikia Riverin Otago, New Zealand, MNZS42815 (a distalright tarsometatarsusof a Tui-sized bird) and several bones of at least one species of Saddleback-sized bird have recently been described. These date from the Early to Middle Miocene( Awamoanto Lillburnian, 19-16 million years ago). [Worthy "et al." (2007)]
Modern knowledge about the living passerines' interrelationships (see the list of families below) suggests that the last common ancestor of all living Passeriformes was a small forest bird, probably with a stubby tail [The last common ancestor of all songbirds most likely had a decidedly longer tail. See del Hoyo "et al." (2003, 2004).] and an overall drab coloration, but possibly with marked
sexual dimorphism. The latter trait seems to have been lost and re-evolved multiple times in songbird evolution alone, judging from its distribution among the extant lineages: the common ancestor of Passerida for example was almost certainly not markedly dimorphic considering the trait is very rare among the basal lineages of these, but very common among the youngest passerid clade, the Passeroidea; on the other hand among the basalmost Passeri there are a considerable number of strongly dimorphic lineages such as the very ancient Menuridaeas well as many Meliphagoideaand Corvoidea. Sexual dimorphism is also not uncommon in the Acanthisittidaeand prominent in some suboscines such as the Pipridaeand Cotingidae.
Early European passerines
In Europe, perching birds are not too uncommon in the fossil record from the
Oligoceneonwards, but most are too fragmentary for a more definite placement:
Wieslochia" (Early Oligocene of Frauenweiler, Germany)
*Passeriformes gen. et sp. indet. (Early Oligocene of Luberon, France) – suboscine or basal [Specimen SMF Av 504. A flattened right hand of a passerine perhaps 10 cm long overall. If suboscine, perhaps closer to
Cotingidaethan to Eurylaimides: Roux (2002), Mayr & Manegold (2006)]
*Passeriformes gen. et spp. indet. (Late Oligocene of France) – several suboscine and oscine taxa [Huguenet "et al." (2003), Mayr & Manegold (2006)]
*Passeriformes gen. et spp. indet. (Middle Miocene of France and Germany) – basal? [Specimens SMF Av 487-496;
SMNS86822, 86825-86826; MNHNSA 1259–1263: tibiotarsusremains of small, possibly basal Passeriformes: Manegold "et al." (2004)]
*Passeriformes gen. et spp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary) – at least 2 taxa, possibly 3; at least one probably Oscines [A partial
coracoidof a probably Muscicapoidea, possibly Turdidae; distal tibiotarsusand tarsometatarsusof a smallish to mid-sized passerine which may be the same as the preceding; proximal ulnaand tarsometatarsus of a Paridae-sized passerine: Gál "et al." (1998-1999, 2000)]
"Wieslochia" was possibly not a member of any extant suborder. That not only the Passeri expanded much beyond their region of origin is proven by an undetermined
broadbill(Eurylaimidae) from the Early Miocene(roughly 20 mya) of Wintershof, Germany, and the indeterminate Late Oligocene suboscine from France listed above. Even very basal Passeriformes might have been common in Europe until the Middle Miocene, some 12 mya. [Manegold "et al." (2004)] Extant Passeri superfamilies were quite distinct by that time and are known since about 12–13 mya when modern genera were present in the corvoidean and basal songbirds. The modern diversity of Passerida genera is known mostly from the Late Miocene onwards and into the Pliocene(about 10–2 mya). Pleistoceneand early Holocene lagerstätten(<1.8 mya) yield numerous extant species, and many yield almost nothing but extant species or their chronospeciesand paleosubspecies.
Americas, the fossil record is more scant before the Pleistocene, from which several still-existing suboscine families are documented. Apart from the indeterminable MACN-SC-1411 (Pinturas Early/Middle Miocene of Santa Cruz Province, Argentina), [Distal right humerus, possibly suboscine: Noriega & Chiappe (1991, 1993)] an extinct lineage of perching birds has been described from the Late Miocene of California, USA: the Palaeoscinidaewith the single genus " Paleoscinis". "Palaeostruthus" eurius" (Pliocene of Florida) probably belongs to an extant family, most likely passeroidean.
Late Quaternary prehistoric birds."
ystematics and taxonomy
InitiallyFact|date=February 2007 , the Corvida and Passerida were classified as "
parvorders" in the suborder Passeri; in accord with the usual taxonomicpractice, they would probably be ranked as infraorders. As originally envisioned in the Sibley-Ahlquist taxonomy, they contained, respectively, the large superfamilies Corvoidea and Meliphagoidea as well as minor lineages, and the superfamilies Sylvioidea, Muscicapoidea and Passeroidea.
This arrangement has been found to be overly simplified by more recent research. Since the mid 2000s, literally dozens of studies are being published which try rather successfully to resolve the
phylogenyof the passeriform radiation. For example, the Corvida in the traditional sense were a rather arbitrary assemblage of early and/or minor lineages of passeriform birds of Old World origin, generally from the region of Australia, New Zealand, and Wallacea. The Passeri on the other hand can be made monophyletic by moving some families about, but the "clean" three-superfamily-arrangement has turned out to be far more complex and it is uncertain whether future authors will stick to it.
Major "wastebin" families such as the
Old World warblers and Old World babblers have turned out to be paraphyleticand are being rearranged. Several taxa turned out to represent highly distinct species-poor lineages and consequently new families had to be established, some of them – like the Stitchbirdof New Zealandand the Eurasian Bearded Reedling– monotypicwith only one living species. [The former does not even have recognized subspecies, while the latter is one of the most singular birds alive today. Good photos of a Bearded Reedling are for example [http://montereybay.com/creagrus/Reedling_bearded-WEH.jpghere] and [http://www.naturephoto-cz.com/photos/mraz/bearded-reedling-05a04012.jpghere] .] . It seems likely that in the Passeri alone, a number of minor lineages will eventually be recognized as distinct superfamilies. For example, the kinglets constitute a single genus with less than 10 species today, but seem to have been among the first perching bird lineages to diverge as the group spread across Eurasia. No particularly close relatives of them have been found among comprehensive studies of the living Passeri, though it is suspected that they might be fairly close to some little-studied tropical Asian groups. Treatment of the nuthatches, wrens, and their closest relatives as a distinct subfamily Certioideais increasingly considered justified; the same might eventually apply to the titmiceand their closest relatives.
This process is still continuing. Therefore, the arrangement as presented here is subject to change. However, it should take precedence over unreferenced conflicting treatments in family, genus and species articles here; see the next section for default sources.
Taxonomic list of Passeriformes families
This list is in taxonomic order, placing related species/groups next to each other. The Passerida subdivisions are updated as needed from the default sequence of the "
Handbook of the Birds of the World", [del Hoyo "et al." (2003-)] based on the most modern and comprehensive studies. [Lovette & Bermingham (2000), Cibois "et al." (2001), Barker "et al." (2002, 2004), Ericson & Johansson (2003), Beresford "et al." (2005), Alström "et al." (2006), Jønsson & Fjeldså (2006)] Updates are added as necessary.
Regarding arrangement of families
The families are sorted into a somewhat unusual sequence. This is because so many reallocations have taken place since about 2005 that a definite arrangement has not been established yet. The present sequence is an attempt to preserve as much of the traditional sequence while giving priority to adequately addressing the relationships between the families.
Acanthisittidae: New Zealand "wrens"
Eurylaimides– Old World suboscines (or Broad-billed suboscines). Probably a separate suborder.
Eurylaimoidea– broadbills and allies
Sapayoidae: Broad-billed Sapayoa
** Superfamily Pittoidea
Tyrannides- New World suboscines
** Superfamily N.N. – "bronchophones"
Tyrannidae: tyrant flycatchers
Tityridae: tityras and allies.
Furnariidae: ovenbirds and woodcreepers
Formicariidae: antpittas, antthrushes and typical tapaculos. Possibly polyphyletic.
Conopophagidae: gnateaters and gnatpittas
*** N.N.: atypical "
tapaculos" (crescent-chests and allies)
Songbirds or oscines
*Basal Passeri – the most ancient true songbirds, endemic to Australia. Sometimes considered a superfamily "Menuroidea"Verify source|date=January 2008 .
Meliphagoidea– mainly insectivores and nectarivores, distribution centered on Australo-Melanesian region extending into surroundings, notably the Pacific.
Maluridae: fairy-wrens, emu-wrens and grasswrens
Dasyornithidae: bristlebirds. Formerly in Acanthizidae.
Acanthizidae: scrubwrens, thornbills, and gerygones
** Meliphagoidea "incertae sedis"
Pardalotidae: pardalotes. Formerly in Acanthizidae, might be included in Meliphagidae.
Acanthorhynchus": spinebills. Usually included in Meliphagidae; might be considered a monotypicfamily if Pardalotidae are considered valid too.
Corvoidea– a highly diverse group of global distribution, but most plentiful in the Australasian region and surroundings. The oldest truly globally successful group of passerines, they include among them what may well be the most intelligent and the most spectacular of the order.
Melanocharitidae: berrypeckers and longbills. Tentatively placed here.
Callaeidae: New Zealand wattlebirds. Tentatively placed here.
** Family N.N.:
Stitchbird. Tentatively placed here.
Cnemophilidae: satinbirds. Tentatively placed here.
Campephagidae: cuckoo-shrikes and trillers
Pachycephalidae: whistlers and allies. Delimitation with regards to several proposed families and subfamilies requires thorough study.
Oriolidae: orioles and Figbird
** Paramythiidae: Tit Berrypecker and Crested Berrypecker. Formerly in Passerida.
Artamidae: woodswallows, butcherbirds, currawongs and Australian Magpie
Malaconotidae: puffback shrikes, bush shrikes, tchagras and boubous
Platysteiridae: wattle-eyes. Formerly in Passerida. Probably paraphyletic.
Pityriaseidae: Bornean Bristlehead. Tentatively placed here.
Prionopidae: helmetshrikes and woodshrikes
Monarchidae: monarch flycatchers
Paradisaeidae: birds of paradise
Corcoracidae: White-winged Chough and Apostlebird
Corvidae: crows, ravens and jays
** Corvoidea "incertae sedis"
Vireolanius": shrike-vireos. Usually included in Vireonidae, possibly a monotypic family,
Erpornis": White-bellied Erpornis. Formerly in " Yuhina" (Passerida: Timaliidae); possibly a monotypic family, possibly in Vireonidae
Colluricinclidae: shrike-thrushes. Often included in Pachycephalidae but perhaps recognizable as a subfamily at least.
Cinclosomatidae: whipbirds and allies. Contains Psophodidaebut that might make it paraphyletic. At least some species belong in Pachycephalidae if Falcunculinae are not considered a distinct family.
Falcunculidae: Shrike-tit and allies. Usually included in Pachycephalidae; might be distinct family or merged in Cinclosomatidae or PsophodidaeVerify source|date=January 2008 .
pitohuis. Usually included in Pachycephalidae but seem closer to Oriolidae and best considered a distinct family including " Oreoica" and possibly other Pachycephalidae "sensu lato".
* Passeri (mainly "
Corvida") " incertae sedis"
** Possible superfamily "Ptilonorhynchoidea"Verify source|date=January 2008 – bowerbirds and Australian treecreepers
Climacteridae: Australian treecreepers
Turnagridae: Piopio ( extinct)
** Possible superfamily N.N. - logrunners and pseudo-babblers
Petroicidae: Australian robins
** Possible superfamily N.N.
Chaetopidae: rock-jumpers. Recently split from Turdidae.
Eupetidae: Malaysian Rail-babbler. Recently split from Cinclosomatidae.
** Possible monotypic superfamily Reguloidea – kinglets
** Possible monotypic superfamily N.N.
** Family N.N.:
Hyliotas. Recently split from Sylviidae.
Irenidae: fairy-bluebirds. Reguloidea? Basal to/in Passeroidea?
Chloropseidae: leafbirds. Reguloidea? Basal to/in Passeroidea?
Sylvioidea– mostly insectivores, distribution centered on the Indo-Pacificregion. Few occur in the Australian region and fewer still in the Americas. Usually sleek and drab birds, few have pronounced sexual dimorphism.
Hirundinidae: swallows and martins
Phylloscopidae: leaf-warblers and allies. Recently split from Sylviidae.
Aegithalidae: long-tailed tits or bushtits [Gill, F., Wright, M. & Donsker, D. (2008). IOC World Bird Names (version 1.6). Available at http://www.worldbirdnames.org/]
Cettiidae: ground-warblers and allies. Recently split from Sylviidae.
Megaluridae: grass-warblers and allies. Recently split from Sylviidae.
Malagasy warblers. A newly assembled family.
Acrocephalidae: marsh- and tree-warblers. Recently split from Sylviidae.
Cisticolidae: cisticolas and allies
Sylviidae: "true/sylviid warblers" and parrotbills. Might be merged in Timaliidae. Monophyly needs confirmation.
Zosteropidae: white-eyes. Probably belongs in Timaliidae.
Timaliidae: (Old World) babblers. Monophyly needs confirmation.
** Sylvioidea "incertae sedis"
African warblers": A proposed clade, but monophyly needs confirmation. Formerly in Sylviidae.
Donacobius": Black-capped Donacobius. Monotypicfamily? Tentatively placed here; possibly closest to Megaluridae. Formerly in Troglodytidae and Mimidae.
Nicator": Relationships unresolved, monotypic family? Tentatively placed here; formerly in Pycnonotidae.
Muscicapoidea– mostly insectivores, near-global distribution centered on Old World tropics. One family endemic to Americas. Nearly absent (except introductions) from the Australian region. Usually rather stocky for their size, most are quite dark and dull though Sturnidaeare commonly iridescent and/or colorful. Sexual dimorphism often absent, sometimes pronounced.
Muscicapidae: Old World flycatchers and chats. Monophyly needs confirmation.
Turdidae: thrushes and allies. Monophyly needs confirmation.
Buphagidae: oxpeckers. Formerly usually included in Sturnidae.
Sturnidae: starlings and possibly Philippine creepers. Placement of latter in Muscicapoidea seems good, but inclusion in Sturnidae requires confirmation; possibly distinct family Rhabdornithidae.
Mimidae: mockingbirds and thrashers
Certhioidea- wrens and allies. Sometimes included in Muscicapoidea.
Tichodromadidae: Wallcreeper: Traditionally placed as a subfamily of the nuthatches and more rarely of the treecreepers, no study has been able to verify either placement this far. Thus it is better considered a monotypic family, at least for the time being.
Salpornithidae: Spotted Creeper. Tentatively placed here; often considered a subfamily of the Certhidae.
Passeroidea– mostly herbivores including many seed-eaters, near-global distribution centered on Palearcticand Americas. Includes the Nine-primaried oscines (probably a subclade). A very high proportion of colorful and highly sexually dimorphic forms.
Passeridae: true sparrows
Motacillidae: wagtails and pipits
Urocynchramidae: Przewalski's Finch. Recently split from Fringillidae; tentatively placed here.
Peucedramidae: Olive Warbler
Estrildidae: estrildid finches (waxbills, munias, etc)
Viduidae: indigobirds and whydahs
Fringillidae: true finches and Hawaiian honeycreepers. Possibly polyphyletic.
Icteridae: grackles, New World blackbirds, and New World orioles
Parulidae: New World warblers
Thraupidae: tanagers and allies
Emberizidae: buntings and American sparrows
** Passeroidea "incertae sedis"
Coerebidae: Bananaquit. Family invalid or not monotypic; reallocation pending.
* Passerida "incertae sedis" - Rather basal Passerida, most of which seem to constitute several small but distinct lineages that could be considered superfamilies. Most occur in Asia, Africa and North America.
Panurus": Bearded Reedling (Bearded "Tit"). Relationships enigmatic. Formerly in " Paradoxornithidae", might be included in Sylvioidea as monotypic family Panuridae or even constitute the smallest passerine superfamily.
** Possible superfamily Paroidea – titmice and allies. Might be included in Sylvioidea.
Paridae: tits, chickadees and titmice
Remizidae: penduline tits. Sometimes included in Paridae.
Stenostiridae: stenostirids ("flycatcher-tits"). A newly assembled family; sometimes included in Paridae.
** Possible superfamily Bombycilloidea – waxwings and allies. Included in Muscicapoidea if Sittoidea/Certhioidea are not considered a distinct superfamily.
Dulidae: Palmchat. Tentatively placed here.
Ptilogonatidae: silky flycatchers. Tentatively placed here.
Hypocoliidae: Hypocolius. Tentatively placed here.
** Possible superfamily "Dicaeoidea" – sunbirds and flowerpeckers. Might be included in Passeroidea.
** Possible monotypic superfamily N.N.
Promeropidae: sugarbirds. Might be included in Passeroidea.
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