Editing Passerine (section)

==Origin and evolution==
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 of [[convergent evolution]], not a close genetic relationship. For example, the [[Troglodytidae|wrens of the Americas and Eurasia]], [[Maluridae|those of Australia]], and [[Acanthisittidae|those 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.{{efn|The name [[wren]] has been applied to other, unrelated birds in Australia and New Zealand. The 27 [[Australasian realm|Australasian]] "wren" species in the family [[Maluridae]] are unrelated, as are the [[New Zealand wren]]s in the family Acanthisittidae; the antwrens in the family [[Thamnophilidae]]; and the wren-babblers of the families [[Timaliidae]], [[Pellorneidae]], and [[Pnoepygidae]]. For the monophyly of the "true wrens", Troglodytidae, see Barker 2004.<ref name=barker04>{{cite journal | last=Barker | first=F.K. | date=2004 | title=Monophyly and relationships of wrens (Aves: Troglodytidae): a congruence analysis of heterogeneous mitochondrial and nuclear DNA sequence data |journal=Molecular Phylogenetics and Evolution |volume=31 |issue=2 | pages=486–504 | doi=10.1016/j.ympev.2003.08.005 | pmid=15062790| bibcode=2004MolPE..31..486B }}</ref> }}

Advances in [[molecular biology]] and improved [[paleobiogeography|paleobiogeographical]] 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.<ref>{{cite journal|doi=10.1111/j.1096-3642.2004.00118.x|title=The evolutionary radiation of modern birds (Neornithes): Reconciling molecules, morphology and the fossil record|date=June 2004|last1=Dyke|first1=Gareth J.|last2=Van Tuinen|first2=Marcel|journal=Zoological Journal of the Linnean Society|volume=141|issue=2|pages=153–177|doi-access=free}}</ref> The first passerines are now thought to have evolved in the [[Southern Hemisphere]] in the late [[Paleocene]] or early [[Eocene]], around 50&nbsp;million years ago.<ref name="ericson2002">{{cite journal|pmc=1690883|pmid=11839192|date=7 February 2002|last1=Ericson|first1=P.G.|last2=Christidis|first2=L.|last3=Cooper|first3=A.|last4=Irestedt|first4=M.|last5=Jackson|first5=J.|last6=Johansson|first6=U.S.|last7=Norman|first7=J.A.|title=A Gondwanan origin of passerine birds supported by DNA sequences of the endemic New Zealand wrens|volume=269|issue=1488|pages=235–241|doi=10.1098/rspb.2001.1877|journal=[[Proceedings of the Royal Society B]]}}</ref><ref name=ClaramuntCracraft2015/>

The initial diversification of passerines coincides with the separation of the southern continents in the [[early Eocene]]. The [[New Zealand wren]]s are the first to become isolated in [[Zealandia]], and the second split involved the origin of the [[Tyranni]] in [[South America]] and the [[Passeri]] in the [[Australia (continent)|Australian continent]].<ref name=ericson2003/><ref name=ClaramuntCracraft2015/><ref name=selvatti2015/> The Passeri experienced a great radiation of forms in Australia. A major branch of the Passeri, the [[parvorder]] [[Passerida]], dispersed into Eurasia and Africa about 40 million years ago, where they experienced further radiation of new lineages.<ref name=ClaramuntCracraft2015/> This eventually led to three major [[Passerida]] lineages comprising about 4,000 species, which in addition to the [[Corvida]] and numerous minor lineages make up songbird diversity today. Extensive [[biogeographical]] mixing happens, with northern forms returning to the south, southern forms moving north, and so on.<ref name=selvatti2015/>

===Fossil record===
====Earliest passerines====
[[File:Superb lyrbird in scrub.jpg|thumb|Male [[superb lyrebird]] (''Menura novaehollandiae''): This unique [[songbird]] shows strong [[sexual dimorphism]], with a peculiarly [[apomorph]]ic display of plumage in males.]]
Perching bird [[osteology]], especially of the limb bones, is rather diagnostic.<ref name=boles97>{{cite journal|author= Boles, Walter E. |year=1997|title= Fossil songbirds (Passeriformes) from the Early Eocene of Australia|journal=[[Emu (journal)|Emu]] |volume=97|issue=1|pages= 43–50|doi=10.1071/MU97004|bibcode=1997EmuAO..97...43B }}</ref><ref name=manegold04>{{cite journal|author1=Manegold, Albrecht |author2=Mayr, Gerald |author3=Mourer-Chauviré, Cécile |name-list-style=amp |year=2004|title= Miocene Songbirds and the Composition of the European Passeriform Avifauna|journal=[[Auk (journal)|Auk]] |volume=121|issue=4|pages= 1155–1160|doi=10.1642/0004-8038(2004)121[1155:MSATCO]2.0.CO;2|s2cid=73668280 |doi-access=free}}
</ref><ref name=mayr06/> 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]]) from [[Murgon, Queensland]], are fossil bone fragments initially assigned to [[Passeriformes]].<ref name=boles97/> However, the material is too fragmentary and their affinities have been questioned.<ref>{{cite journal |last1=Mayr |first1=G |title=The age of the crown group of passerine birds and its evolutionary significance–molecular calibrations versus the fossil record |journal=Systematics and Biodiversity |date=2013 |volume=11 |issue=1 |pages=7–13 |doi=10.1080/14772000.2013.765521|bibcode=2013SyBio..11....7M |s2cid=85167051 }}</ref> Several more recent fossils from the [[Oligocene]] of Europe, such as ''[[Wieslochia]]'', ''[[Jamna (bird)|Jamna]]'', ''[[Resoviaornis]],'' and ''[[Crosnoornis]]'',<ref>Bochenski, Z.M., Tomek, T., Bujoczek, M. (2021) [https://link.springer.com/article/10.1007/s10336-021-01858-0#citeas A new passeriform (Aves: Passeriformes) from the early Oligocene of Poland sheds light on the beginnings of Suboscines]. ''J Ornithol''. https://doi.org/10.1007/s10336-021-01858-0</ref> are more complete and definitely represent early passeriforms, and have been found to belong to a variety of modern and extinct lineages.<ref name=":0">{{Cite journal |last1=Lowi-Merri |first1=Talia M. |last2=Gjevori |first2=Martina |last3=Bochenski |first3=Zbigniew M. |last4=Wertz |first4=Krzysztof |last5=Claramunt |first5=Santiago |date=2024-12-31 |title=Total-evidence dating and the phylogenetic affinities of early fossil passerines |url=https://www.tandfonline.com/doi/full/10.1080/14772019.2024.2356086 |journal=Journal of Systematic Palaeontology |language=en |volume=22 |issue=1 |doi=10.1080/14772019.2024.2356086 |bibcode=2024JSPal..2256086L |issn=1477-2019}}</ref>

From the [[Saint Bathans#Geology and palaeontology|Bathans Formation]] at the [[Manuherikia River]] in [[Otago]], New Zealand, [[Museum of New Zealand Te Papa Tongarewa|MNZ]] S42815 (a [[Anatomical terms of location#Proximal and distal|distal]] right [[tarsometatarsus]] of a [[tūī|tui]]-sized bird) and several bones of at least one species of [[Saddleback (bird)|saddleback]]-sized bird have recently been described. These date from the Early to [[Middle Miocene]] ([https://books.google.com/books?id=gto4AAAAIAAJ&pg=PA584 Awamoan] to [[Lillburnian]], 19–16&nbsp;mya).<ref>{{cite journal|author1=Worthy, Trevor H. |author2=Tennyson, A.J.D. |author3=Jones, C. |author4=McNamara, J.A. |author5=Douglas, B.J. |year=2007|title= Miocene waterfowl and other birds from central Otago, New Zealand|journal=[[Journal of Systematic Palaeontology]] |volume=5|issue=1|pages= 1–39|doi=10.1017/S1477201906001957|bibcode=2007JSPal...5....1W |author1-link=Trevor H. Worthy |hdl=2440/43360 |s2cid=85230857 |hdl-access=free }}</ref>

====Early European passerines====
[[File:Wieslochia.jpg|thumb|upright|''[[Wieslochia]]'' fossil]]

In Europe, perching birds are not too uncommon in the fossil record from the [[Oligocene]] onward, belonging to several lineages:
* ''[[Wieslochia]]'' (Early Oligocene of Frauenweiler, Germany) – suboscine<ref name=":0" /><!--*Naturwissensch91:173; ActPalPol51:315 -->
* ''[[Resoviaornis]]'' (Early Oligocene of Wola Rafałowska, Poland) – oscine<ref name=":0" />
* ''[[Jamna (bird)|Jamna]]'' (Early Oligocene of Jamna Dolna, Poland) – basal<ref name=":0" />
* ''[[Winnicavis]]'' (Early Oligocene of Lower Silesian Voivodeship, Poland)
* ''[[Crosnoornis]]'' (Early Oligocene of Poland) - suboscine<ref name=":0" />
* Passeriformes gen. et sp. indet. (Early Oligocene of Luberon, France) – suboscine<ref name=":0" /> or basal{{efn|Specimen [[Senckenberg Museum|SMF]] Av 504. A flattened right hand of a passerine perhaps 10&nbsp;cm long overall. If suboscine, perhaps closer to [[Cotingidae]] than to [[Eurylaimides]].<ref>{{cite journal|last= Roux | first=T. |year=2002|title= Deux fossiles d'oiseaux de l'Oligocène inférieur du Luberon |trans-title=Two bird fossils from the Lower Oligocene of Luberon|journal=Courrier Scientifique du Parc Naturel Régional du Luberon |volume=6|pages=38–57  | url=http://documents.irevues.inist.fr/bitstream/handle/2042/58077/CS_2002_6_38.pdf?sequence=1 }}</ref><ref name=mayr06/> }}
* Passeriformes gen. et spp. indet. (Late Oligocene of France) – several suboscine and oscine taxa<ref>{{cite journal|author1=Hugueney, Marguerite |author2=Berthet, Didier |author3=Bodergat, Anne-Marie |author4=Escuillié, François |author5=Mourer-Chauviré, Cécile |author6=Wattinne, Aurélia |name-list-style=amp |year=2003|title= La limite Oligocène-Miocène en Limagne: changements fauniques chez les mammifères, oiseaux et ostracodes des différents niveaux de Billy-Créchy (Allier, France) |trans-title=The Oligocene-Miocene boundary in Limagne: faunal changes in the mammals, birds and ostracods from the different levels of Billy-Créchy (Allier, France)|journal=[[Geobios]]|volume=36|issue=6|pages=719–731|doi=10.1016/j.geobios.2003.01.002|bibcode=2003Geobi..36..719H }}</ref><ref name=mayr06>{{cite journal|last1=Mayr | first1=Gerald | last2=Manegold | first2=Albrecht |name-list-style=amp |year=2006|title= A Small Suboscine-like Passeriform Bird from the Early Oligocene of France|journal=[[Condor (journal)|Condor]] |volume=108|issue=3|pages= 717–720|doi=10.1650/0010-5422(2006)108[717:ASSPBF]2.0.CO;2 | doi-access=free }}</ref>
* Passeriformes gen. et spp. indet. (Middle Miocene of France and Germany) – basal?{{efn|Specimens [[Senckenberg Museum|SMF]] Av 487–496; [[SMNS]] 86822, 86825-86826; [[MNHN]] SA 1259–1263: [[tibiotarsus]] remains of small, possibly basal Passeriformes.<ref name=manegold04/>}}
* Passeriformes gen. et spp. indet. (Sajóvölgyi Middle Miocene of Mátraszőlős, Hungary) – at least 2 taxa, possibly 3; at least one probably Oscines.{{efn|A partial [[coracoid]] of a probable Muscicapoidea, possibly [[Turdidae]]; [[Anatomical terms of location#Proximal and distal|distal]] [[tibiotarsus]] and [[tarsometatarsus]] of a smallish to mid-sized passerine that may be the same as the preceding; [[Anatomical terms of location#Proximal and distal|proximal]] [[ulna]] and tarsometatarsus of a [[Paridae]]-sized passerine.<ref>{{cite journal |author1=Gál, Erika |author2=Hír, János |author3=Kessler, Eugén |author4=Kókay, József |name-list-style=amp |year=1998–99 |title=Középsõ-miocén õsmaradványok, a Mátraszõlõs, Rákóczi-kápolna alatti útbevágásból. I. A Mátraszõlõs 1. lelõhely |trans-title=Middle Miocene fossils from the sections at the Rákóczi chapel at Mátraszőlős. Locality Mátraszõlõs I. |journal=Folia Historico Naturalia Musei Matraensis |volume=23 |pages=33–78 }}</ref><ref>
{{cite journal |author1=Gál, Erika |author2=Hír, János |author3=Kessler, Eugén |author4=Kókay, József |author5=Márton, Venczel |name-list-style=amp |year=2000 |title=Középsõ-miocén õsmaradványok a Mátraszõlõs, Rákóczi-kápolna alatti útbevágásból II. A Mátraszõlõs 2. lelõhely |trans-title=Middle Miocene fossils from the section of the road at the Rákóczi Chapel, Mátraszõlõs. II. Locality Mátraszõlõs 2 |journal=Folia Historico Naturalia Musei Matraensis |volume=24 |pages=39–75 }}</ref> }}
* Passeriformes gen. et sp. indet. (Middle Miocene of Felsőtárkány, Hungary) – oscine?{{efn|A [[humerus]] [[diaphysis]] piece of a swallow-sized passerine.<ref name=hir01>{{cite journal |last1=Hír | first1=János | last2=Kókay | first2=József | last3=Venczel | first3=Márton |last4=Gál | first4=Erika |author5=GKessler, Eugén |name-list-style=amp |year=2001 |title=Elõzetes beszámoló a felsõtárkányi "Güdör-kert" n. õslénytani lelõhelykomplex újravizsgálatáról |trans-title=A preliminary report on the revised investigation of the paleontological locality-complex "Güdör-kert" at Felsõtárkány, Northern Hungary |journal=Folia Historico Naturalia Musei Matraensis |volume=25 |pages=41–64 | url=http://adatbank.transindex.ro/vendeg/htmlk/pdf6928.pdf |archive-url=https://web.archive.org/web/20210316054055/http://adatbank.transindex.ro/vendeg/htmlk/pdf6928.pdf |archive-date=2021-03-16 |url-status=live  }}</ref> }}
* Passeriformes gen. et sp. indet. (Late Miocene of Polgárdi, Hungary) – [[Sylvioidea]] ([[Sylviidae]]? [[Cettiidae]]?)<ref name=hir01/><!-- this should be moved to Sylvioidea -->
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 Early [[Miocene]] (roughly 20&nbsp;mya) of [[:de:Wintershof (Eichstätt)|Wintershof]], Germany, the Late Oligocene [[carpometacarpus]] from France listed above, and ''[[Wieslochia]]'', among others.<ref name=manegold04/><ref name="ClaramuntCracraft2015"/> 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 the [[Pliocene]] (about 10–2&nbsp;mya). [[Pleistocene]] and early [[Holocene]] [[lagerstätten]] (<1.8&nbsp;mya) yield numerous extant species, and many yield almost nothing but extant species or their [[chronospecies]] and paleosubspecies.

====American fossils====
{{See also|Late Quaternary prehistoric birds}}
In the [[Americas]], the fossil record is more scant before the Pleistocene, from which several still-existing families are documented. Apart from the indeterminable [[MACN]]-SC-1411 (Pinturas Early/Middle Miocene of Santa Cruz Province, Argentina),{{efn|Distal right [[humerus]], possibly suboscine.<ref>{{cite journal|author1=Noriega, Jorge I. |author2=Chiappe, Luis M. |name-list-style=amp |year=1991|title= El más antiguo Passeriformes de America del Sur. Presentation at VIII Journadas Argentinas de Paleontologia de Vertebrados |trans-title=The most ancient passerine from South America|journal=Ameghiniana |volume=28|issue=3–4|pages=410|url=https://books.google.com/books?id=hqDVeF8W8OkC}}</ref><ref>{{cite journal|author1=Noriega, Jorge I. |author2=Chiappe, Luis M. |name-list-style=amp |year=1993|title= An Early Miocene Passeriform from Argentina|journal=[[Auk (journal)|Auk]] |volume=110|issue=4|pages= 936–938|url=http://sora.unm.edu/sites/default/files/journals/auk/v110n04/p0936-p0938.pdf |archive-url=https://web.archive.org/web/20140313222943/http://sora.unm.edu/sites/default/files/journals/auk/v110n04/p0936-p0938.pdf |archive-date=2014-03-13 |url-status=live|doi= 10.2307/4088653 |jstor=4088653}}</ref>}} an extinct lineage of perching birds has been described from the Late Miocene of California, United States: the [[Palaeoscinidae]] with the single genus ''[[Palaeoscinis]]''. ''"Palaeostruthus" eurius'' (Pliocene of Florida)<!-- Condor84:240 --> probably belongs to an extant family, most likely [[passeroidea]]n.