Passerine
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 Fringilla coelebs (chaffinch), male
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A passerine is a bird of the giant order Passeriformes. More than half of all species of bird are passerines. Sometimes known as perching birds or, less accurately, as songbirds, the passerines form one of the most spectacularly diverse terrestrial vertebrate orders: with around 5,400[verification needed] species, it is roughly twice as diverse as the largest of the mammal orders, the Rodentia.
The group gets its name from the Latin name for the House Sparrow (Passer domesticus).
Characteristics
Many passerines are songbirds and have complex muscles to control their syrinx; many gape in the nest as infants to beg for food.
The order is divided into two suborders, Tyranni, and Passeri (oscines). Oscines have the most control of their syrinx muscles among birds, producing a wide range of songs and other vocalizations (though some of them, such as the crows, do not sound like it, while others like the Lyrebird are accomplished imitators).
Most passerines are smaller than typical members of other avian orders. The heaviest and altogether largest passerine is the Thick-billed Raven; some Common Ravens come close and the two species of Lyrebird are longer overall.[verification needed]
The foot of a passerine has three toes directed forward without any webbing or joining, and one toe directed backward. The hind toe joins the leg at the same level as the front toes. In other orders of birds the toe arrangement is different.
Most passerines lay coloured eggs, in contrast to non-passerines, where the colour is white except in some ground nesting groups such as Charadriiformes and nightjars, where camouflage is necessary, and some parasitic cuckoos which have to match the passerine host's egg.
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, of Australia, and of New Zealand all look 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 yet remaining Passeriformes.
Much research remains to be done, but advances in molecular biology and improved paleobiogeographical data are gradually revealing a clearer picture of passerine origins and evolution. It is now thought that the early passerines evolved in Gondwana at some time in the Paleogene, maybe around the Late Paleocene (c. 60-55 mya). The initial split was between the Tyranni, the ancestors of the songbirds, the Eurylaimides and the New Zealand "wrens" which must have diverged during a short period of time (some million years at best). The Passeriformes apparently evolved out of a fairly close-knit clade of "near passerines" which contains such birds as the Piciformes, Coraciiformes, and Cuculiformes (Johnasson & Ericson 2003).
A little later, a great radiation of forms took place out of Australia-New Guinea: the Passeri or songbirds. A major branch of the passerine tree, the 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 Eurasia and Africa, where there was a further explosive radiation of new lineages. This eventually led to the corvoidan clade, three major passeridan lineages, and numerous minor lineages making up songbird diversity today. There has been extensive biogeographical mixing, with northern forms returning to the south, southern forms moving north, and so on.
Fossil record
Perching bird osteology, especially of the leg bones, is rather diagnostic. Still, the first Passeriformes were apparently forms 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, Queensland are fossil bone fragments clearly recognizable as passeriform; they prove that some 55 mya, early perching birds were recognizably distinct. The Murgon fossils represent two species of approximately some 10 and some 20 cm overall length. 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.
In Europe, perching birds are not too uncommon in the fossil record from the Oligocene onwards, but most are too fragmentary for a more definite placement:
- Wieslochia (Early Oligocene of Frauenweiler, Germany)
- Passeriformes gen. et sp. indet. SMF Av 504 (Late Oligocene of Luberon, France) - suboscine
- Passeriformes gen. et sp. indet. (Late Oligocene of France)
- Passeriformes gen. et sp. indet. SMN Av 487-496 (Middle Miocene of Petersbuch, Germany)
- Passeriformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary) - 3 spp.: Passeroidea, Muscicapoidea, Paridae?
- Passeriformes gen. et sp. indet. SMNS 86822,86825-86826
A set of Early/Middle Miocene (approx. 15 mya) fossils from Sansan, France (MNHN SA 1259-1263) is doubtfully passeriform. 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. Wieslochia was apparently not a member of any extant suborder either. Extant Passeri superfamilies were well distinct by about 12-13 mya and modern genera were present in the corvoidean and basal songbirds. The modern diversity of Passerida genera evolved mostly from the Late Miocene onwards and into the Pliocene (about 10-2 mya). Pleistocene lagerstätten (<1.8 mya) yield numerous to near-exclusively extant species, or their chronospecies and paleosubspecies.
In the Americas, the fossil record is more scant. Apart from the indeterminable MACN-SC-1411 (Pinturas Early/Middle Miocene of Santa Cruz Province, Argentina), an entirely extinct lineage of perching birds has been described from the USA:
- Palaeoscinidae
- Paleoscinis (Late Miocene of California)
"Palaeostruthus" eurius (Pliocene of Florida) probably belongs to an extant family, most likely passerioidean.
Systematics and taxonomy
This list is in taxonomic order, placing related species/groups next to each other. The subdivisions follow Lovette & Bermingham's 2000 study of the c-mos proto-oncogene nDNA sequence, which has been validated by subsequent research as regards its general findings.
Initially[citation needed], the Corvida and Passerida were classified as "parvorders" in the suborder Passeri; in accord with the usual taxonomic prectice, they would probably be ranked as infraorders. As originally envisioned (see also Sibley-Ahlquist taxonomy), they contained the superfamilies Corvoidea and minor lineages, and the superfamilies Sylvioidea, Muscicapoidea and Passeroidea, respectively.
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 phylogeny of 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 the Wallacea. The Passeri on the other hand can be made monophyletic by moving some families about, but the "clean" 3-superfamily-arrangement has turned out to be far more complex.
Several taxa turned out to be isolated species-poor lineages and consequently new families had to be established; 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, 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. Likewise, major "wastebin" families such as the Old World warblers and Old World babblers have turned out to be paraphyletic.
This process is still continuing. Therefore, the arrangement as presented here is not definite and may be subject to change without previous notice until the relevant literature has been reviewed.
However, it should take precedence over unreferenced and conflicting treatments in family, genus and species articles.
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. It was attempted to preserve as much of the former arrangement, while giving priority to adequately address the relationships between the families (see also Jønsson & Fjeldså 2006).
Taxonomic list of Passeriformes families
Suborder Acanthisitti
- Acanthisittidae: New Zealand "wrens"
Suborder Tyranni
Suboscines
- Infraorder Eurylaimides - Old World suboscines (or Broad-billed suboscines; probably a separate suborder)
- Superfamily Eurylaimoidea - broadbills and allies
- Eurylaimidae: broadbills
- Philepittidae: asities
- Sapayoidae: Broad-billed Sapayoa
- Superfamily Pittoidea
- Pittidae: pittas
- Superfamily Eurylaimoidea - broadbills and allies
- Infraorder Tyrannides - New World suboscines
- Superfamily N.N. - "bronchophones"
- Tyrannidae: tyrant flycatchers
- Cotingidae: cotingas
- Pipridae: manakins
- Superfamily Furnarioidea - tracheophones
- Furnariidae: ovenbirds and woodcreepers
- Thamnophilidae: antbirds
- Formicariidae: antpittas, antthrushes and typical tapaculos (possibly polyphyletic)
- Conopophagidae: gnateaters and gnatpittas
- N.N.: atypical "tapaculos" (crescent-chests and allies)
- Superfamily N.N. - "bronchophones"
Suborder Passeri
Songbirds or oscines
- "Parvorder" Corvida (paraphyletic)
- Menuridae: lyrebirds
- Atrichornithidae: scrub birds
- Climacteridae: Australian treecreepers
- Maluridae: fairy-wrens, emu-wrens and grasswrens
- Meliphagidae: honeyeaters and chats
- Promeropidae: sugarbirds
- Pardalotidae: pardalotes, scrubwrens, thornbills, and gerygones
- Petroicidae: Australian robins
- Orthonychidae: logrunners
- Pomatostomidae: Australasian babblers
- Cinclosomatidae: whipbirds and allies
- Neosittidae: sittellas
- Pachycephalidae: whistlers, shrike-thrushes, pitohuis and allies
- Dicruridae: monarch flycatchers and allies
- Campephagidae: cuckoo shrikes and trillers
- Oriolidae: orioles and Figbird
- Paramythiidae: tit berrypecker and crested berrypecker - formerly in Passerida
- Melanocharitidae: berrypeckers and longbills - formerly in Passerida
- Artamidae: wood swallows, butcherbirds, currawongs and Australian Magpie
- Paradisaeidae: birds of paradise
- Corvidae: crows, ravens and jays
- Corcoracidae: White-winged Chough and Apostlebird
- Irenidae: fairy-bluebirds
- Chloropseidae: leafbirds - tentatively placed here
- Laniidae: shrikes
- Prionopidae: helmetshrikes
- Malaconotidae: puffback shrikes, bush shrikes, tchagras and boubous
- Pityriaseidae: Bornean Bristlehead
- Platysteiridae: wattle-eyes or puffback flycatchers - formerly in Passerida
- Aegithinidae: ioras - tentatively placed here
- Vireonidae: vireos
- Vangidae: vangas
- Ptilonorhynchidae: bowerbirds
- Turnagridae: Piopio
- Callaeidae: New Zealand wattlebirds
- Family N.N.: Stitchbird
- Picathartidae: rockfowl. Tentatively placed here.
- Chaetopidae: rock-jumpers. Recently split from Turdidae. Tentatively placed here.
- Infraorder Passerida
- Superfamily Sylvioidea - mostly insectivores, distribution centered on the Indo-Pacific region. Few occur in the Americas.
- Alaudidae: larks
- Hirundinidae: swallows and martins
- Phylloscopidae: leaf-warblers and allies. Recently split from Sylviidae.
- Aegithalidae: long-tailed tits
- Cettiidae: ground-warblers and allies. Recently split from Sylviidae.
- Megaluridae: grass-warblers and allies. Recently split from Sylviidae.
- "Bernieridae": Malagasy warblers. A newly assembled family.
- Acrocephalidae: marsh- and tree-warblers. Recently split from Sylviidae.
- Pycnonotidae: bulbuls
- Cisticolidae: cisticolas and allies
- Sylviidae: "true/sylviid warblers" and parrotbills. Might be merged into Timaliidae. Monophyly needs confirmation.
- Zosteropidae: white-eyes. Probably belongs into Timaliidae.
- Timaliidae: babblers. Monophyly needs confirmation.
- Sylvioidea incertae sedis
- "African warblers": A proposed clade, but monophyly needs confirmation. Formerly in Sylviidae.
- Donacobius: Black-capped Donacobius. Monotypic family? Tentatively placed here; possibly closest to Megaluridae. Formerly in Troglodytidae.
- Nicator: Relationships unresolved, monotypic family? Tentatively placed here; formerly in Pycnonotidae.
- Panurus: Bearded Reedling (Bearded "Tit"). Relationships enigmatic, monotypic family Panuridae? Tentatively placed here; formerly in "Paradoxornithidae".
- Superfamily Muscicapoidea - mostly insectivores, near-global distribution centered on Old World tropics. One family endemic to Americas.
- Cinclidae: dippers
- 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
- Superfamily Passeroidea - mostly herbivores, near-global distribution centered on Palearctic and Americas. Includes the Nine-primaried oscines (probably a subclade).
- Passeridae: true sparrows
- Prunellidae: accentors
- Motacillidae: wagtails and pipits
- Urocynchramidae: Przewalski's Finch. Recently split from Fringillidae; tentatively placed here.
- Peucedramidae: Olive Warbler
- Estrildidae: estrildid finches (waxbills, munias, etc)
- Ploceidae: weavers
- Viduidae: indigobirds and whydahs
- Fringillidae: true finches. Possibly polyphyletic.
- Drepanididae: Hawaiian honeycreepers. Might be merged into Fringillidae.
- Icteridae: grackles, New World blackbirds, and New World orioles
- Parulidae: New World warblers
- Thraupidae: tanagers and allies
- Cardinalidae: cardinals
- 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 superfamilies. Most occur in Asia, Africa and North America
- Possible superfamily "Dicaeoidea" - sunbirds and flowerpeckers
- Nectariniidae: sunbirds
- Dicaeidae: flowerpeckers
- Possible superfamily Bombycilloidea - waxwings and allies
- Bombycillidae: waxwings
- Dulidae: Palmchat. Tentatively placed here.
- Ptilogonatidae: silky flycatchers. Tentatively placed here.
- Hypocoliidae: Hypocolius. Tentatively placed here.
- Possible superfamily Paroidea - titmice and allies
- 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 Sittoidea or Certhioidea - wrens and allies.
- Sittidae: nuthatches
- Tichodromadidae: Wallcreeper. Tentatively placed here.
- Certhiidae: treecreepers
- Salpornithidae: Spotted Creeper. Tentatively placed here; might belong into Certhidae.
- Troglodytidae: wrens
- Polioptilidae: gnatcatchers
- Possible monotypic superfamily N.N.
- Promeropidae: sugarbirds
- Possible monotypic superfamily N.N.
- Family N.N.: Hyliotas. Recently split from Sylviidae.
- Possible superfamily Reguloidea - kinglets. Tentatively placed here.
- Regulidae: kinglets
- Possible superfamily "Dicaeoidea" - sunbirds and flowerpeckers
- Superfamily Sylvioidea - mostly insectivores, distribution centered on the Indo-Pacific region. Few occur in the Americas.
See also
ReferencesISBN links support NWE through referral fees
- Johansson, Ulf S. & Ericson, Per G. P. (2003): Molecular support for a sister group relationship between Pici and Galbulae (Piciformes sensu Wetmore 1960). J. Avian Biol. 34(2): 185–197. Digital object identifier (DOI): 10.1034/j.1600-048X.2003.03103.x PDF fulltext
- Jønsson, Knud A. & Fjeldså, Jon (2006): A phylogenetic supertree of oscine passerine birds (Aves: Passeri). Zool. Scripta 35(2): 149–186. Digital object identifier (DOI): :10.1111/j.1463-6409.2006.00221.x (HTML abstract)
- Lovette, Irby J.& Bermingham, Eldredge (2000): c-mos Variation in Songbirds: Molecular Evolution, Phylogenetic Implications, and Comparisons with Mitochondrial Differentiation. Molecular Biology and Evolution 17(10): 1569–1577. PDF fulltext
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