Apasserine (/ˈpæsəraɪn/) is anybird of theorderPasseriformes (/ˈpæsərɪfɔːrmiːz/; fromLatinpasser 'sparrow' andformis '-shaped') which includes more than half of all bird species. Sometimes known asperching birds, passerines generally have ananisodactyl arrangement of their toes (three pointing forward and one back), which facilitates perching.
The terms "passerine" and "Passeriformes" are derived from thescientific name of thehouse sparrow,Passer domesticus, whose genus is theLatin word for sparrow.[7] Formerly this meant the songbirds of Europe. Now it also includes perching, non-singing birds from the Americas.[8]
The order is divided into three suborders,Tyranni (non-singing, Americas),Passeri (songbirds), and thebasalNew Zealand wrens.[9] Oscines have the best control of theirsyrinx muscles among birds, producing a wide range ofsongs and other vocalizations, though some of them, such as thecrows, do not sound musical to human beings. Some, such as thelyrebird, are accomplished mimics.[10] TheNew Zealand wrens are tiny birds restricted toNew Zealand, at least in modern times; they were long placed in Passeri.
Pterylosis or the feather tracts in a typical passerine
Most passerines are smaller than typical members of other avian orders. The heaviest and altogether largest passerines are thethick-billed raven[11] and the larger races ofcommon raven, each exceeding 1.5 kg (3.3 lb) and 70 cm (28 in). Thesuperb lyrebird and somebirds-of-paradise, due to very long tails or tail coverts, are longer overall. The smallest passerine is theshort-tailed pygmy tyrant, at 6.5 cm (2.6 in) and 4.2 g (0.15 oz).[12]
The foot of a passerine has three toes directed forward and one toe directed backward, calledanisodactyl arrangement. The hind toe (hallux) is long and joins the leg at approximately the same level as the front toes. This arrangement enables passerine birds to easily perch upright on branches. The toes have no webbing or joining, but in somecotingas, the second and third toes are united at their basal third.
The leg of passerine birds contains an additional special adaptation for perching. A tendon in the rear of the leg running from the underside of the toes to the muscle behind thetibiotarsus will automatically be pulled and tighten when the leg bends, causing the foot to curl and become stiff when the bird lands on a branch. This enables passerines to sleep while perching without falling off.[13][14]
Most passerine birds have 12 tail feathers but thesuperb lyrebird has 16,[15] and several spinetails in the familyFurnariidae have 10, 8, or even 6, as is the case ofDes Murs's wiretail. Species adapted to tree trunk climbing such astreecreepers andwoodcreeper have stiff tail feathers that are used as props during climbing.[16] Extremely long tails used as sexual ornaments are shown by species in different families. A well-known example is thelong-tailed widowbird.
The chicks of passerines arealtricial: blind, featherless, and helpless when hatched from their eggs. Hence, the chicks require extensive parental care. Most passerines lay colored eggs, in contrast with nonpasserines, most of whose eggs are white except in some ground-nesting groups such asCharadriiformes andnightjars, where camouflage is necessary, and in someparasiticcuckoos, which match the passerine host's egg. Thevinous-throated parrotbill has two egg colors, white and blue, to deter the brood parasiticcommon cuckoo.[citation needed]
Clutches vary considerably in size: some larger passerines of Australia such as lyrebirds and scrub-robins lay only a single egg, most smaller passerines in warmer climates lay between two and five, while in the higher latitudes of the Northern Hemisphere, hole-nesting species liketits can lay up to a dozen and other species around five or six.The familyViduidae do not build their own nests, instead, they lay eggs in other birds' nests.[citation needed]
The evolutionary history of the passerine families and the relationships among them remained rather mysterious until the late 20th century. In many cases, passerine families were grouped together on the basis of morphological similarities that, it is now believed, are the result ofconvergent evolution, not a close genetic relationship. For example, thewrens of the Americas and Eurasia,those of Australia, andthose of New Zealand look superficially similar and behave in similar ways, 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.[a]
Advances inmolecular biology and improvedpaleobiogeographical data gradually are revealing a clearer picture of passerine origins and evolution that reconciles molecular affinities, the constraints of morphology, and the specifics of the fossil record.[18] The first passerines are now thought to have evolved in theSouthern Hemisphere in the latePaleocene or earlyEocene, around 50 million years ago.[5][6]
The initial diversification of passerines coincides with the separation of the southern continents in theearly Eocene. TheNew Zealand wrens are the first to become isolated inZealandia, and the second split involved the origin of theTyranni inSouth America and thePasseri in theAustralian continent.[2][6][3] The Passeri experienced a great radiation of forms in Australia. A major branch of the Passeri, theparvorderPasserida, dispersed into Eurasia and Africa about 40 million years ago, where they experienced further radiation of new lineages.[6] This eventually led to three majorPasserida lineages comprising about 4,000 species, which in addition to theCorvida and numerous minor lineages make up songbird diversity today. Extensivebiogeographical mixing happens, with northern forms returning to the south, southern forms moving north, and so on.[3]
Perching birdosteology, especially of the limb bones, is rather diagnostic.[19][20][21] However, the early fossil record is poor because passerines are relatively small, and their delicate bones do not preserve well.Queensland Museum specimens F20688 (carpometacarpus) and F24685 (tibiotarsus) fromMurgon, Queensland, are fossil bone fragments initially assigned toPasseriformes.[19] However, the material is too fragmentary and their affinities have been questioned.[22] Several more recent fossils from theOligocene of Europe, such asWieslochia,Jamna,Resoviaornis, andCrosnoornis,[23] are more complete and definitely represent early passeriforms, and have been found to belong to a variety of modern and extinct lineages.[24]
That suboscines expanded much beyond their region of origin is proven by several fossils from Germany such as a presumed broadbill (Eurylaimidae) humerus fragment from the EarlyMiocene (roughly 20 mya) ofWintershof, Germany, the Late Oligocenecarpometacarpus from France listed above, andWieslochia, among others.[20][6] Extant Passeri super-families 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 onward and into thePliocene (about 10–2 mya).Pleistocene and earlyHolocenelagerstätten (<1.8 mya) yield numerous extant species, and many yield almost nothing but extant species or theirchronospecies and paleosubspecies.
In theAmericas, the fossil record is more scant before the Pleistocene, from which several still-existing families are documented. Apart from the indeterminableMACN-SC-1411 (Pinturas Early/Middle Miocene of Santa Cruz Province, Argentina),[f] an extinct lineage of perching birds has been described from the Late Miocene of California, United States: thePalaeoscinidae with the single genusPalaeoscinis."Palaeostruthus" eurius (Pliocene of Florida) probably belongs to an extant family, most likelypasseroidean.
Phylogenetic relationship of the suborders within the Passeriformes. The numbers are from the list published by theInternational Ornithologists' Union in January 2020.[1][33]
The Passeriformes is currently divided into three suborders:Acanthisitti (New Zealand wrens),Tyranni, (suboscines) andPasseri (oscines or songbirds). The Passeri is now subdivided into two major groups recognized now asCorvides andPasserida respectively containing the largesuperfamiliesCorvoidea andMeliphagoidea, as well as minor lineages, and the superfamiliesSylvioidea,Muscicapoidea, andPasseroidea but this arrangement has been found to be oversimplified. Since the mid-2000s, studies have investigated thephylogeny of the Passeriformes and found that many families fromAustralasia traditionally included in the Corvoidea actually represent more basal lineages within oscines. Likewise, the traditional three-superfamily arrangement within the Passeri has turned out to be far more complex and will require changes in classification.[citation needed]
Major "wastebin" families such as theOld World warblers andOld World babblers have turned out to beparaphyletic and are being rearranged. Several taxa turned out to represent highly distinct lineages, so new families had to be established, some of these – like thestitchbird of New Zealand and theEurasianbearded reedling –monotypic with only one living species.[34] In the Passeri alone, a number of minor lineages will eventually be recognized as distinct superfamilies. For example, thekinglets 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 theirs have been found among comprehensive studies of the living Passeri, though they might be fairly close to some little-studied tropical Asian groups.Nuthatches,wrens, and their closest relatives are currently grouped in a distinct super-familyCerthioidea.
This list is in taxonomic order, placing related families next to one another. The families listed are those recognised by theInternational Ornithologists' Union (IOC).[1] The order and the division into infraorders, parvorders, and superfamilies follows the phylogenetic analysis published by Carl Oliveros and colleagues in 2019.[33][g] The relationships between the families in the suborder Tyranni (suboscines) were all well determined but some of the nodes in Passeri (oscines or songbirds) were unclear owing to the rapid splitting of the lineages.[33]
Relationships between living Passeriformes families based on the phylogenetic analysis of Oliveros et al (2019).[33] Some terminals have been renamed to reflect families recognised by the IOC but not in that study.[1] The IOC familiesAlcippeidae andTeretistridae were not sampled in this study.
^The namewren has been applied to other, unrelated birds in Australia and New Zealand. The 27Australasian "wren" species in the familyMaluridae are unrelated, as are theNew Zealand wrens in the family Acanthisittidae; the antwrens in the familyThamnophilidae; and the wren-babblers of the familiesTimaliidae,Pellorneidae, andPnoepygidae. For the monophyly of the "true wrens", Troglodytidae, see Barker 2004.[17]
^SpecimenSMF Av 504. A flattened right hand of a passerine perhaps 10 cm long overall. If suboscine, perhaps closer toCotingidae than toEurylaimides.[26][21]
^SpecimensSMF Av 487–496;SMNS 86822, 86825-86826;MNHN SA 1259–1263:tibiotarsus remains of small, possibly basal Passeriformes.[20]
^Oliveros et al (2019) use the list of families published by Dickinson and Christidis in 2014.[33][35] Oliveros et al include 8 families that are not included on the IOC list. These are not shown here. By contrast, the IOC list includes 15 families that are not present in Dickinson and Christidis. In 13 of these cases, the position of the additional family in the taxonomic order can be determined from the species included by Oliveros and colleagues in their analysis. No species in the familiesAlcippeidae andTeretistridae were sampled by Oliveros et al so their position is uncertain.[1][33]
^The order of the families within the superfamily Orioloidea is uncertain.[33]
^The order of the families within the superfamily Malaconotoidea is uncertain.[33]
^The order of the families within the superfamily Corvoidea is uncertain.[33]
^The taxonomic sequence of the superfamilies Locustelloidea, Sylvioidea and Aegithaloidea is uncertain, although the order of the families within each of the superfamilies is well determined.[33]
^The order of some of the families within the superfamily Emberizoidea is uncertain.[33]
^The familyTeretistridae (Cuban warblers) is tentatively placed here. The family was not included in the analysis published by Oliveros et al (2019).[33] Dickinson and Christidis (2014) considered the genusTeretistrisIncertae sedis.[38] Barker et al (2013) found that Teretistridae is closely related toZeledoniidae.[37]
^Chatterjee, Sankar (2015).The Rise of Birds: 225 Million Years of Evolution. JHU Press. pp. 206–207.ISBN9781421415901.
^Winkler, D. W., S. M. Billerman, and I.J. Lovette (2020). Lyrebirds (Menuridae), version 1.0. In Birds of the World (S. M. Billerman, B. K. Keeney, P. G. Rodewald, and T. S. Schulenberg, Editors). Cornell Lab of Ornithology, Ithaca, New York, US.https://doi.org/10.2173/bow.menuri1.01
^Madge, S. (2020). Thick-billed Raven (Corvus crassirostris), version 1.0. In Birds of the World (J. del Hoyo, A. Elliott, J. Sargatal, D. A. Christie, and E. de Juana, Editors). Cornell Lab of Ornithology, Ithaca, New York, US.https://doi.org/10.2173/bow.thbrav1.01
^Clock, B. (2020). Short-tailed Pygmy-Tyrant (Myiornis ecaudatus), version 1.0. In Birds of the World (J. del Hoyo, A. Elliott, J. Sargatal, D. A. Christie, and E. de Juana, Editors). Cornell Lab of Ornithology, Ithaca, New York, US.https://doi.org/10.2173/bow.stptyr1.01
^Barker, F.K. (2004). "Monophyly and relationships of wrens (Aves: Troglodytidae): a congruence analysis of heterogeneous mitochondrial and nuclear DNA sequence data".Molecular Phylogenetics and Evolution.31 (2):486–504.Bibcode:2004MolPE..31..486B.doi:10.1016/j.ympev.2003.08.005.PMID15062790.
^Hugueney, Marguerite; Berthet, Didier; Bodergat, Anne-Marie; Escuillié, François; Mourer-Chauviré, Cécile & Wattinne, Aurélia (2003). "La limite Oligocène-Miocène en Limagne: changements fauniques chez les mammifères, oiseaux et ostracodes des différents niveaux de Billy-Créchy (Allier, France)" [The Oligocene-Miocene boundary in Limagne: faunal changes in the mammals, birds and ostracods from the different levels of Billy-Créchy (Allier, France)].Geobios.36 (6):719–731.Bibcode:2003Geobi..36..719H.doi:10.1016/j.geobios.2003.01.002.
^Gál, Erika; Hír, János; Kessler, Eugén & Kókay, József (1998–99). "Középsõ-miocén õsmaradványok, a Mátraszõlõs, Rákóczi-kápolna alatti útbevágásból. I. A Mátraszõlõs 1. lelõhely" [Middle Miocene fossils from the sections at the Rákóczi chapel at Mátraszőlős. Locality Mátraszõlõs I.].Folia Historico Naturalia Musei Matraensis.23:33–78.
^Gál, Erika; Hír, János; Kessler, Eugén; Kókay, József & Márton, Venczel (2000). "Középsõ-miocén õsmaradványok a Mátraszõlõs, Rákóczi-kápolna alatti útbevágásból II. A Mátraszõlõs 2. lelõhely" [Middle Miocene fossils from the section of the road at the Rákóczi Chapel, Mátraszõlõs. II. Locality Mátraszõlõs 2].Folia Historico Naturalia Musei Matraensis.24:39–75.
^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 examplehereArchived 16 October 2007 at theWayback Machine andhereArchived 31 July 2008 at theWayback Machine.
^abcdeCracraft, Joel (2014). "Avian higher-level relationships and classification: passeriformes". InDickinson, E.C.;Christidis, L. (eds.).The Howard & Moore Complete Checklist of the Birds of the World. Vol. 2: Passerines (4th ed.). Eastbourne, UK: Aves Press. pp. xvii-xxxiii [xxiv].ISBN978-0-9568611-2-2.
Mayr, Gerald (2016).Avian evolution: the fossil record of birds and its paleobiological significance. Chichester, West Sussex: John Wiley & Sons.ISBN978-1-119-02076-9.
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