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{{Short description|Family of birds}}
{{other uses}}
{{Other uses}}
{{Automatic taxobox
{{Automatic taxobox
| name = White-eyes
| name = White-eyes
Line 5: Line 6:
| image_caption = [[Silvereye]] (''Zosterops lateralis''), adult (right) and juveniles
| image_caption = [[Silvereye]] (''Zosterops lateralis''), adult (right) and juveniles
| taxon = Zosteropidae
| taxon = Zosteropidae
| authority = <small>(disputed)</small>[[Charles Lucien Bonaparte|Bonaparte]], 1853
| authority = [[Charles Lucien Bonaparte|Bonaparte]], 1853
| subdivision_ranks = Genera
| subdivision_ranks = Genera
| subdivision =
| subdivision = See text
''[[Apalopteron]]''<br/>
''[[Cleptornis]]''<br/>
''[[Heleia]]''<br/>
''[[Hypocryptadius]]''<br/>
''[[Lophozosterops]]''<br/>
''[[Megazosterops]]''<br/>
''[[Oculocincta]]''<br/>
''[[Rukia (bird)|Rukia]]''<br/>
''[[Speirops]]''<br/>
''[[Tephrozosterops]]''<br/>
''[[Woodfordia]]''<br/>
''[[Yuhina]]''<br/>
''[[Zosterops]]''
}}
}}


[[Image:Zosterops palpebrosus1.jpg|thumb|242px|[[Oriental white-eye]], ''Zosterops palpebrosus'']]
[[Image:Zosterops palpebrosus1.jpg|thumb|242px|[[Indian white-eye]], ''Zosterops palpebrosus'']]
The '''white-eyes''' are small [[passerine]] [[bird]]s native to tropical, subtropical and temperate [[Sub-Saharan Africa]], southern and eastern [[Asia]], and [[Australasia]]. White-eyes inhabit most tropical islands in the [[Indian Ocean]], the western [[Pacific Ocean]], and the [[Gulf of Guinea]]. Discounting some widespread members of the genus ''[[Zosterops]]'', most species are [[Endemism in birds|endemic]] to single islands or [[archipelagos]]. The [[silvereye]], ''Zosterops lateralis'', naturally colonised [[New Zealand]], where it is known as the "wax-eye" or ''tauhou'' ("stranger"), from 1855. The silvereye has also been [[Introduced species|introduced]] to the [[Society Islands]] in [[French Polynesia]], while the [[Japanese white-eye]] has been introduced to [[Hawaii]].
The '''white-eyes''' are a [[family (biology)|family]], '''Zosteropidae''', of small [[passerine]] [[bird]]s native to tropical, subtropical and temperate [[Sub-Saharan Africa]], southern and eastern [[Asia]], and [[Australasia]]. White-eyes inhabit most tropical islands in the [[Indian Ocean]], the western [[Pacific Ocean]], and the [[Gulf of Guinea]]. Discounting some widespread members of the genus ''[[Zosterops]]'', most species are [[Endemism in birds|endemic]] to single islands or [[archipelagos]]. The [[silvereye]], ''Zosterops lateralis'', naturally colonised [[New Zealand]], where it is known as the "wax-eye" or ''tauhou'' ("stranger"), from 1855. The silvereye has also been [[Introduced species|introduced]] to the [[Society Islands]] in [[French Polynesia]], while the [[Japanese white-eye]] has been introduced to [[Hawaii]].


==Characteristics==
==Characteristics==
White-eyes are mostly of undistinguished appearance, the plumage being generally greenish olive above, and pale grey below. Some species have a white or bright yellow throat, breast, or lower parts, and several have buff flanks. As their common name implies, many species have a conspicuous ring of tiny white feathers around their eyes.<ref name=EoB>{{cite book |editor=Forshaw, Joseph|author= Lindsey, Terence|year=1991|title=Encyclopaedia of Animals: Birds|publisher= Merehurst Press|location=London|pages= 207|isbn= 1-85391-186-0}}</ref> The scientific name of the group also reflects this latter feature, being derived from the [[Ancient Greek]] for "girdle-eye". They have rounded wings and strong legs. Like many other nectarivorous birds, they have slender, pointed bills, and brush-tipped tongues.<ref name=EoB/>{{sfn|van Balen|2008|p=[https://archive.org/details/handbookofbirdso0013unse/page/413/mode/1up 413]}} The size ranges up to {{convert|15|cm|in|abbr=on}} in length.


All the species of white-eyes are sociable, forming large flocks that only separate on the approach of the breeding season. They build trees [[bird nest|nest]]s and lay two to four eggs which are usually pale blue.{{sfn|van Balen|2008|p=[https://archive.org/details/handbookofbirdso0013unse/page/432/mode/1up 432]}} Though mainly insectivorous, they eat [[nectar]] and fruits of various kinds. The silvereye can be a problem in [[Australia]]n vineyards, by piercing the [[grape]] allowing infection or insect damage to follow.{{sfn|van Balen|2008|p=[https://archive.org/details/handbookofbirdso0013unse/page/429/mode/1up 429]}}
White-eyes are mostly of undistinguished appearance, the plumage being generally greenish olive above, and pale grey below. Some species have a white or bright yellow throat, breast or lower parts, and several have buff flanks. As their common name implies, many species have a conspicuous ring of tiny white feathers around their eyes.<ref name=EoB>{{cite book |editor=Forshaw, Joseph|author= Lindsey, Terence|year=1991|title=Encyclopaedia of Animals: Birds|publisher= Merehurst Press|location=London|pages= 207|isbn= 1-85391-186-0}}</ref> The scientific name of the group also reflects this latter feature, being derived from the [[Ancient Greek]] for "girdle-eye". They have rounded wings and strong legs. Like many other nectivorous birds, they have slender, pointed bills, and brush-tipped tongues.<ref name=EoB/> The size ranges up to {{convert|15|cm|in|abbr=on}} in length.

All the species of white-eyes are sociable, forming large flocks which only separate on the approach of the breeding season. They build tree [[bird nest|nest]]s and lay two to four unspotted pale blue eggs.{{Citation needed|date=June 2008}} Though mainly insectivorous, they eat [[nectar]] and fruits of various kinds. The silvereye can be a problem in [[Australia]]n vineyards, through piercing the [[grape]] allowing infection or insect damage to follow.{{Citation needed|date=June 2008}}

White-eyes are the city bird of [[Kurayoshi, Tottori|Kurayoshi City]], in Tottori, Japan.{{citation needed|date=March 2017}}


==Systematics==
==Systematics==
The white-eyes were long considered a distinct [[family (biology)|family]] '''Zosteropidae''' because they are rather homogeneous in [[morphology (biology)|morphology]] and [[ecology]], leading to little [[adaptive radiation]] and [[divergence (biology)|divergence]].
The family Zosteropidae was introduced (as a subfamily Zosteropinae) in 1853 by the French naturalist [[Charles Lucien Bonaparte]].<ref>{{ cite journal | last=Bonaparte | first=Charles Lucien | author-link=Charles Lucien Bonaparte | year=1853 | title=Classification ornithologique par series | language=French | journal=Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences | volume=37 | pages=641–647 [644, No. 83] | url=https://www.biodiversitylibrary.org/page/1216804 }}</ref><ref>{{cite book | last=Bock | first=Walter J. | year=1994 | title=History and Nomenclature of Avian Family-Group Names | series=Bulletin of the American Museum of Natural History | volume= 222 | publisher=American Museum of Natural History | location=New York | pages=154, 229 | url=http://digitallibrary.amnh.org/handle/2246/830 }} Linked page allows download of the 48MB pdf</ref> The white-eyes were long considered a distinct [[family (biology)|family]] Zosteropidae because they are rather homogeneous in [[morphology (biology)|morphology]] and [[ecology]], leading to little [[adaptive radiation]] and [[divergence (biology)|divergence]].


The genus ''[[Apalopteron]]'', formerly placed in the [[Meliphagidae]], was transferred to the white-eyes on genetic evidence{{Citation needed|date=April 2007}}. It differs much in appearance from the typical white-eyes, ''[[Zosterops]]'', but is approached by some [[Micronesia]]n [[taxa]]; its colour pattern is fairly unusual save the imperfect white-eye-ring.
The genus ''[[Apalopteron]]'', formerly placed in the [[Meliphagidae]], was transferred to the white eyes in 1995 on genetic and behavioral evidence. It differs much in appearance from the typical white-eyes, ''[[Zosterops]]'', but is approached by some [[Micronesia]]n [[taxa]]; its color pattern is fairly unusual save the imperfect white eye-ring.<ref name="Springer 1995">{{cite journal |last1=Springer |first1=Mark S. |last2=Higuchi |first2=Hiroyoshi |last3=Ueda |first3=Keisuke |last4=Minton |first4=Jason |last5=Sibley |first5=Charles G. |title=Molecular Evidence That the Bonin Islands "Honeyeater" Is a White-eye |journal=Journal of the Yamashina Institute for Ornithology |date=1995 |volume=27 |issue=2 |pages=66–77_1 |doi=10.3312/jyio1952.27.66 |url=https://www.jstage.jst.go.jp/article/jyio1952/27/2/27_2_66/_pdf|doi-access=free }}</ref>


In 2003, Alice Cibois published the results of her study of [[mtDNA]] [[cytochrome b|cytochrome ''b'']] and [[12S rRNA|12S]]/[[MT-RNR2|16S rRNA]] [[DNA sequence|sequence]] data. According to her results, the white-eyes were likely to form a [[clade]] also containing the [[yuhina]]s, which were until then placed with the [[Old World babbler]]s, a large "[[wastebin taxon|wastebin]]" family. Previous molecular studies (e.g. Sibley & Ahlquist 1990, Barker ''et al.'' 2002) had together with the morphological evidence tentatively placed white-eyes as the Timaliidae's closest relatives already. But some questions remained, mainly because the white-eyes are all very similar birds in habitus and habits, while the Old World babblers are very diverse (because, as we now know, the group as formerly defined was [[polyphyletic]]).
In 2003, Alice Cibois published the results of her study of [[mtDNA]] [[cytochrome b|cytochrome ''b'']] and [[12S rRNA|12S]]/[[MT-RNR2|16S rRNA]] [[DNA sequence|sequence]] data. According to her results, the white-eyes were likely to form a [[clade]] also containing the [[yuhina]]s, which were until then placed with the [[Old World babbler]]s, a large "[[wastebin taxon|wastebin]]" family.<ref>{{Cite journal | last=Cibois | first=Alice | date=2003 | title=Mitochondrial DNA phylogeny of Babblers (Timaliidae) | journal=The Auk | volume=120 | issue=1 | pages=35–54 | doi=10.1093/auk/120.1.35 | doi-access=free}}</ref> Previous molecular studies (e.g. Sibley & Ahlquist 1990, Barker ''et al.'' 2002) had together with the morphological evidence tentatively placed white-eyes as the Timaliidae's closest relatives already. But some questions remained, mainly because the white-eyes are all very similar birds in habitus and habits, while the Old World babblers are very diverse (because, as we now know, the group as formerly defined was [[polyphyletic]]).


Combined with the yuhinas (and possibly other Timaliidae), the limits of the white-eye clade to the "true" Old World babblers becomes indistinct. Therefore, the current (early 2007) opinion weighs towards merging the group into the Timaliidae, perhaps as a [[subfamily]] ("Zosteropinae"). Few white-eyes have been thoroughly studied with the new results in mind, however, and almost all of these are from ''Zosterops'' which even at this point appears [[lumpers and splitters|over-lumped]]. Also, many "Old World babblers" remain of unresolved relationships. Whether there can be a clear delimitation of a white-eye subfamily or even a young or emerging family is a question that requires a more comprehensive study of both this group and Timaliidae to resolve (Jønsson & Fjeldså 2006).
Combined with the yuhinas (and possibly other Timaliidae), the limits of the white-eye clade to the "true" Old World babblers becomes indistinct. Therefore, the current (early 2007) opinion weighs towards merging the group into the Timaliidae, perhaps as a [[subfamily]] ("Zosteropinae"). Few white-eyes have been thoroughly studied with the new results in mind, however, and almost all of these are from ''Zosterops'' which even at this point appears [[lumpers and splitters|over-lumped]]. Also, many "Old World babblers" remain in unresolved relationships. Whether there can be a clear delimitation of a white-eye subfamily or even a young or emerging family is a question that requires a more comprehensive study of both this group and Timaliidae to resolve (Jønsson & Fjeldså 2006).


For example, a revision of the yuhinas and the genus ''[[Stachyris]]'' (Cibois ''et al.'' 2002), based on the same genes as Cibois (2003), revealed that the Philippine species placed in the latter genus by some were actually yuhinas. However, when the review by Jønsson & Fjeldså (2006) was published, no study had tried to propose a phylogeny for the newly defined yuhinas including the white-eyes. Therefore, Jønsson & Fjeldså (2006) give a rather misleading phylogeny for the group. It appears as if the yuhinas are polyphyletic, with the [[white-collared yuhina]] being closer to the ancestor of the ''Zosterops'' white-eyes than to other yuhinas including the species moved from ''Stachyris'' (Cibois ''et al.'' 2002).
For example, a revision of the yuhinas and the genus ''[[Stachyris]]'' (Cibois ''et al.'' 2002), based on the same genes as Cibois (2003), revealed that the Philippine species placed in the latter genus by some were actually yuhinas. However, when the review by Jønsson & Fjeldså (2006) was published, no study had tried to propose a phylogeny for the newly defined yuhinas including the white eyes. Therefore, Jønsson & Fjeldså (2006) give a rather misleading phylogeny for the group. It appears as if the yuhinas are polyphyletic, with the [[white-collared yuhina]] being closer to the ancestor of the ''Zosterops'' white-eyes than to other yuhinas including the species moved from ''Stachyris'' (Cibois ''et al.'' 2002).


In the past, the [[Madanga]] (''Madanga ruficollis'') was included in this family but studies now place it as an atypical member of the [[Motacillidae]].<ref>{{cite journal|doi=10.1098/rsos.140364|pmid=26064613|title=Dramatic niche shifts and morphological change in two insular bird species|journal=Royal Society Open Science|volume=2|issue=3|pages=140364|year=2015|last1=Alstrom|first1=P.|last2=Jonsson|first2=K. A.|last3=Fjeldsa|first3=J.|last4=Odeen|first4=A.|last5=Ericson|first5=P. G. P.|last6=Irestedt|first6=M.|pmc=4448822}}</ref>
In the past, the [[Madanga]] (''Madanga ruficollis'') was included in this family but studies now place it as an atypical member of the [[Motacillidae]].<ref>{{cite journal|doi=10.1098/rsos.140364|pmid=26064613|title=Dramatic niche shifts and morphological change in two insular bird species|journal=Royal Society Open Science|volume=2|issue=3|pages=140364|year=2015|last1=Alstrom|first1=P.|last2=Jonsson|first2=K. A.|last3=Fjeldsa|first3=J.|last4=Odeen|first4=A.|last5=Ericson|first5=P. G. P.|last6=Irestedt|first6=M.|pmc=4448822|bibcode=2015RSOS....240364A}}</ref>


The cladogram below showing the relationships between families is based on a study of babblers by Tianlong Cai and collaborators published in 2019.<ref name=cai>{{Cite journal | last1=Cai | first1=T. | last2=Cibois | first2=A. | last3=Alström | first3=P. | last4=Moyle | first4=R.G. | last5=Kennedy | first5=J.D. | last6=Shao | first6=S. | last7=Zhang | first7=R. | last8=Irestedt | first8=M. | last9=Ericson | first9=P.G.P. | last10=Gelang | first10=M. | last11=Qu | first11=Y. | last12=Lei | first12=F. | last13=Fjeldså | first13=J. | date=2019 | title=Near-complete phylogeny and taxonomic revision of the world's babblers (Aves: Passeriformes) | journal=Molecular Phylogenetics and Evolution | volume=130 | pages=346–356 | doi=10.1016/j.ympev.2018.10.010 | pmid=30321696 | doi-access=free }}</ref><ref name=ioc/>
===Diversity===
{{clade | style=font-size:90%;line-height:90%
'''Family: ZOSTEROPIDAE'''
|1={{clade
* '''Genus ''[[Speirops]]'''''
|1={{clade
** [[Fernando Po speirops]], ''Speirops brunneus''
|1=[[Pycnonotidae]] – bulbuls (167 species)
** [[Principe speirops]], ''Speirops leucophoeus''
|2={{clade
** [[Black-capped speirops]], ''Speirops lugubris''
|1={{clade
** [[Cameroon speirops]], ''Speirops melanocephalus''
|1=[[Sylviidae]] – sylviid babblers (32 species)
* '''Genus ''[[Zosterops]]''''' – typical white-eyes (some 75 species, 1–3 recently [[extinct]]; polyphyletic)
|2=[[Paradoxornithidae]] – parrotbills and myzornis (38 species)
* '''Genus ''[[Rukia (bird)|Rukia]]''''' – Eastern Carolines white-eyes
}}
** [[Long-billed white-eye]], ''Rukia longirostra''<!-- BirdConserv.Res.Contrib.01:001 -->
|2={{clade
** [[Truk white-eye]], ''Rukia ruki''<!-- BirdConserv.Res.Contrib.01:001 -->
|1='''Zosteropidae''' – white-eyes (152 species)
* '''Genus ''[[Cleptornis]]'''''
|2={{clade
** [[Golden white-eye]], ''Cleptornis marchei''<!-- BirdConserv.Res.Contrib.01:001 -->
|1=[[Timaliidae]] – tree babblers (58 species)
* '''Genus ''[[Tephrozosterops]]'''''
|2={{clade
** [[Rufescent white-eye]], ''Tephrozosterops stalkeri''
|1=[[Pellorneidae]] – ground babblers (68 species)
[[Image:Zosterops borbonicus borbonicus p1180876.jpg|thumb|242px|Mascarene white-eye<br/>''Zosterops b. borbonicus'' ("Réunion grey white-eye")]]
|2={{clade
* '''Genus ''[[Lophozosterops]]'''''
|1=[[Alcippeidae]] – Alcippe fulvettas (10 species)
** [[Crested white-eye]], ''Lophozosterops dohertyi''
|2=[[Leiothrichidae]] – laughingthrushes and allies (133 species)
** [[Black-masked white-eye]], ''Lophozosterops goodfellowi''
}}
** [[Mees's white-eye]], ''Lophozosterops javanicus''
}}
** [[Grey-hooded white-eye]], ''Lophozosterops pinaiae''
}}
** [[Streaky-headed white-eye]], ''Lophozosterops squamiceps''
}}
** [[Cream-browed white-eye]], ''Lophozosterops superciliaris''
}}
* '''Genus ''[[Oculocincta]]'''''
}}
** [[Pygmy white-eye]], ''Oculocincta squamifrons''
}}
* '''Genus ''[[Heleia]]'''''
}}
** [[Thick-billed white-eye]], ''Heleia crassirostris''
** [[Spot-breasted white-eye]], ''Heleia muelleri''
* '''Genus ''[[Chlorocharis]]'''''
** [[Mountain blackeye]], ''Chlorocharis emiliae''
* '''Genus ''[[Woodfordia]]'''''
** [[Sanford's white-eye]], ''Woodfordia lacertosa''
** [[Bare-eyed white-eye]], ''Woodfordia superciliosa''
* '''Genus ''[[Megazosterops]]''''' – sometimes placed in ''Rukia''
** [[Giant white-eye]], ''Megazosterops palauensis''<!-- BirdConserv.Res.Contrib.01:001 -->
* '''Genus ''[[Hypocryptadius]]'''''
** [[Cinnamon ibon]], ''Hypocryptadius cinnamomeus''
* '''Genus ''[[Apalopteron]]'''''
**[[Bonin white-eye]], ''Apalopteron familiare'' (formerly Bonin honeyeater)


The cladogram below showing the relationships between the genera is based on the study by Carl Oliveros and collaborators that was published in 2021. The genera ''[[Apalopteron]]'', ''[[Tephrozosterops]]'' and ''[[Rukia]]'' were not sampled in this study. The genus ''[[Megazosterops]]'' was found to be nested in ''[[Heleia]]''.<ref>{{Cite journal | last1=Oliveros | first1=C.H. | last2=Andersen | first2=M.J. | last3=Moyle | first3=R.G. | date=2021 | title=A phylogeny of white-eyes based on ultraconserved elements | journal=Molecular Phylogenetics and Evolution | volume=164 | pages=107273 | doi=10.1016/j.ympev.2021.107273 | doi-access=free }}</ref> The earlier study by Cai and collaborators found a generally similar phylogeny but with ''[[Cleptornis]]'' as [[sister taxon|sister]] to ''Heleia''. Cai's study found that ''Apalopteron'' was nested within ''Heleia'' with weak support and that ''[[Tephrozosterops]]'' was sister to ''[[Zosterops]]''.<ref name=cai/>
[[Image:Yuhina diademata - 20060115.jpg|thumb|242px|[[White-collared yuhina]] (''Yuhina diademata''), a close relative of the white-eyes]]
{{clade|style=font-size:90%;line-height:90%
If the white-eyes are maintained as a separate family or subfamily, this yuhinas would have to be included there too.
|label1='''Zosteropidae'''
|1={{clade
|1=''[[Parayuhina]]'' – white-collared yuhina
|2={{clade
|1=''[[Staphida]]'' – yuhinas (3 species)
|2={{clade
|1=''[[Yuhina]]'' – yuhinas (7 species)
|2={{clade
|1={{clade
|1=''[[Cleptornis]]'' – golden white-eye
|2={{clade
|1=''[[Dasycrotapha]]'' – babblers (3 species)
|2=''[[Sterrhoptilus]]'' – babblers (4 species)
}}
}}
|2={{clade
|1=''[[Zosterornis]]'' – babblers (5 species)
|2={{clade
|1=''[[Heleia]]'' and ''[[Megazosterops]]'' – white-eyes (11 species altogether)
|2=''[[Zosterops]]'' – white-eyes, speirops and the [[silvereye]] (110 species)
}}
}}
}}
}}
}}
}}
}}


==List of genera==
* '''Genus ''[[Yuhina]]''''' – yuhinas
The family contains 149 species divided into 13 genera:<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela Rasmussen | date=January 2021 | title=ylviid babblers, parrotbills, white-eyes | work=IOC World Bird List Version 11.1 | url=https://www.worldbirdnames.org/bow/sylvias/ | publisher=International Ornithologists' Union | access-date=17 June 2021 }}</ref>
** [[White-collared yuhina]], ''Yuhina diademata''
[[Image:Yuhina diademata - 20060115.jpg|thumb|242px|[[White-collared yuhina]] (''Parayuhina diademata''), a close relative of the white-eyes]]
** [[White-naped yuhina]], ''Yuhina bakeri''
* ''[[Parayuhina]]'' – white-collared yuhina
** [[Whiskered yuhina]], ''Yuhina flavicollis''
* ''[[Staphida]]'' – (3 species)
** [[Burmese yuhina]], ''Yuhina humilis''
* ''[[Yuhina]]'' – yuhinas (7 species)
** [[Stripe-throated yuhina]], ''Yuhina gularis''
* ''[[Dasycrotapha]]'' – babbler and pygmy babblers (3 species)
** [[Rufous-vented yuhina]], ''Yuhina occipitalis''
* ''[[Sterrhoptilus]]'' – babblers (4 species)
** [[Taiwan yuhina]], ''Yuhina brunneiceps''
* ''[[Zosterornis]]'' – striped babblers (5 species)
** [[Black-chinned yuhina]], ''Yuhina nigrimenta''
* ''[[Cleptornis]]'' – golden white-eye
** [[Striated yuhina]], ''Yuhina castaniceps''
* ''[[Rukia]]'' – white-eyes (2 species)
** [[Indochinese yuhina]], ''Yuhina torqueola''
* ''[[Megazosterops]]'' – giant white-eye
** [[Chestnut-crested yuhina]], ''Yuhina everetti''
* ''[[Heleia]]'' – heleias (10 species)
* '''Genus ''[[Dasycrotapha]]''''' – formerly in ''Stachyris''; tentatively placed here
* ''[[Apalopteron]]'' – Bonin white-eye
** [[Flame-templed babbler]], ''Dasycrotapha speciosa''
* ''[[Tephrozosterops]]'' – rufescent darkeye
** [[Mindanao pygmy babbler]], ''Dasycrotapha plateni''
* ''[[Zosterops]]'' – (110 species including 3 recently [[extinct]])
** [[Visayan pygmy babbler]], ''Dasycrotapha pygmaea''
* '''Genus ''[[Sterrhoptilus]]''''' – formerly in ''Stachyris''; tentatively placed here
** [[Golden-crowned babbler]] (or yuhina), ''Sterrhoptilus dennistouni''
** [[Black-crowned babbler]] (or yuhina), ''Sterrhoptilus nigrocapitata''
** [[Rusty-crowned babbler]] (or yuhina), ''Sterrhoptilus capitalis''
* '''Genus ''[[Zosterornis]]''''' – formerly in ''Stachyris''; tentatively placed here
** [[Chestnut-faced babbler]] (or yuhina), ''Zosterornis whiteheadi''
** [[Luzon striped babbler]] (or yuhina), ''Zosterornis striatus''
** [[Panay striped babbler]] (or yuhina), ''Zosterornis latistriatus''
** [[Negros striped babbler]] (or yuhina), ''Zosterornis nigrorum''
** [[Palawan striped babbler]] (or yuhina), ''Zosterornis hypogrammicus''


==References==
==References==
{{Reflist}}
{{Reflist}}

==Sources==
*{{ cite book | last=van Balen | first=S. | year=2008 | chapter=Family Zosteropidae (White-eyes) | editor1-last=del Hoyo | editor1-first=J. | editor2-last=Elliott | editor2-first=A. | editor3-last=Christie | editor3-first=D.A. | title=Handbook of the Birds of the World | volume=13: Penduline-tits to Shrikes | location=Barcelona, Spain | publisher=Lynx Edicions | isbn=978-84-96553-45-3 | pages=402–485 | chapter-url=https://archive.org/details/handbookofbirdso0013unse/page/402/mode/1up | chapter-url-access=registration }}


==Further reading==
==Further reading==
*Alström, Per; Ericson, Per G.P.; Olsson, Urban & Sundberg, Per (2006): Phylogeny and classification of the avian superfamily Sylvioidea. ''Molecular Phylogenetics and Evolution'' '''38'''(2): 381–397. <small>{{doi|10.1016/j.ympev.2005.05.015}} {{PMID|16054402}}</small>
*{{cite journal | last1 = Alström | first1 = Per | last2 = Ericson | first2 = Per G.P. | last3 = Olsson | first3 = Urban | last4 = Sundberg | first4 = Per | year = 2006 | title = Phylogeny and classification of the avian superfamily Sylvioidea | journal = Molecular Phylogenetics and Evolution | volume = 38 | issue = 2| pages = 381–397 | doi = 10.1016/j.ympev.2005.05.015 | pmid = 16054402 }}
*Barker, F. Keith; Barrowclough, George F. & Groth, Jeff G. (2002): A phylogenetic hypothesis for passerine birds: taxonomic and biogeographic implications of an analysis of nuclear DNA sequence data. ''[[Proceedings of the Royal Society#Proceedings of the Royal Society B|Proc. R. Soc. B]]'' '''269'''(1488): 295–308. <small>{{doi|10.1098/rspb.2001.1883}}</small> [http://www.tc.umn.edu/~barke042/pdfs/Barker.et.al02.pdf PDF fulltext]
*{{cite journal | last1 = Barker | first1 = F. Keith | last2 = Barrowclough | first2 = George F. | last3 = Groth | first3 = Jeff G. | year = 2002 | title = A phylogenetic hypothesis for passerine birds: taxonomic and biogeographic implications of an analysis of nuclear DNA sequence data | journal = [[Proceedings of the Royal Society#Proceedings of the Royal Society B|Proc. R. Soc. B]] | volume = 269 | issue = 1488| pages = 295–308 | doi = 10.1098/rspb.2001.1883 | pmc = 1690884 | pmid = 11839199 }}
*{{cite journal | last1 = Cibois | first1 = Alice | last2 = Kalyakin | first2 = Mikhail V. | last3 = Lian-Xian | first3 = Han | last4 = Pasquet | first4 = Eric | year = 2002 | title = Molecular phylogenetics of babblers (Timaliidae): revaluation of the genera ''Yuhina'' and ''Stachyris'' | journal = [[Journal of Avian Biology|J. Avian Biol.]] | volume = 33 | issue = 4 | pages = 380–390 | doi = 10.1034/j.1600-048X.2002.02882.x }}
*Cibois, Alice (2003): Mitochondrial DNA Phylogeny of Babblers (Timaliidae). ''[[Auk (journal)|Auk]]'' '''120'''(1): 1–20. <small>[[Digital Object Identifier|DOI]]: 10.1642/0004-8038(2003)120[0035:MDPOBT]2.0.CO;2</small> [http://findarticles.com/p/articles/mi_qa3793/is_200301/ai_n9225604 HTML fulltext without images] {{dead link|date=October 2013}}
*Cibois, Alice; Kalyakin, Mikhail V.; Lian-Xian, Han & Pasquet, Eric (2002): Molecular phylogenetics of babblers (Timaliidae): revaluation of the genera ''Yuhina'' and ''Stachyris''. ''[[Journal of Avian Biology|J. Avian Biol.]]'' '''33''': 380–390. <small>{{doi|10.1034/j.1600-048X.2002.02882.x}}</small> (HTML abstract)
* del Hoyo, J.; Elliot, A. & Christie D. (editors). (2006). Handbook of the Birds of the World. Volume 12: Picathartes to Tits and Chickadees. Lynx Edicions. {{ISBN|978-84-96553-42-2}}
* del Hoyo, J.; Elliot, A. & Christie D. (editors). (2006). Handbook of the Birds of the World. Volume 12: Picathartes to Tits and Chickadees. Lynx Edicions. {{ISBN|978-84-96553-42-2}}
*Jønsson, Knud A. & Fjeldså, Jon (2006): A phylogenetic supertree of oscine passerine birds (Aves: Passeri). ''[[Zoologica Scripta|Zool. Scripta]]'' '''35'''(2): 149–186. <small>{{doi|10.1111/j.1463-6409.2006.00221.x}}</small> (HTML abstract)
*{{cite journal | last1 = Jønsson | first1 = Knud A. | last2 = Fjeldså | first2 = Jon | year = 2006 | title = A phylogenetic supertree of oscine passerine birds (Aves: Passeri) | journal = [[Zoologica Scripta|Zool. Scripta]] | volume = 35 | issue = 2| pages = 149–186 | doi = 10.1111/j.1463-6409.2006.00221.x | s2cid = 85317440 }}
*{{ cite journal | last1=Lim | first1=B.T.M. | last2=Sadanandan | first2=K.R. | last3=Dingle | first3=C. | last4=Leung | first4=Y.Y. | last5=Prawiradilaga | first5=D.M. | last6=Irham | first6=M. | last7=Ashari | first7=H. | last8=Lee | first8=J.G.H. | last9=Rheindt | first9=F.E. | year=2018 | title=Molecular evidence suggests radical revision of species limits in the great speciator white‑eye genus ''Zosterops'' | journal=Journal of Ornithology | volume= | issue= | pages= | doi=10.1007/s10336-018-1583-7 }}
*{{ cite journal | last1=Lim | first1=B.T.M. | last2=Sadanandan | first2=K.R. | last3=Dingle | first3=C. | last4=Leung | first4=Y.Y. | last5=Prawiradilaga | first5=D.M. | last6=Irham | first6=M. | last7=Ashari | first7=H. | last8=Lee | first8=J.G.H. | last9=Rheindt | first9=F.E. | year=2018 | title=Molecular evidence suggests radical revision of species limits in the great speciator white‑eye genus ''Zosterops'' | journal=Journal of Ornithology | volume= 160| pages= 1–16| doi=10.1007/s10336-018-1583-7 | s2cid=51890116 }}
* Mees, G. F. (1957): A Systematic Review of the Indo-Australian Zosteropidae Parts I. Zoologische Verhandelingen 35:1–204 [http://www.repository.naturalis.nl/document/148848 PDF]
* {{cite journal | last1 = Mees | first1 = G. F. | year = 1957 | title = A Systematic Review of the Indo-Australian Zosteropidae Parts I. | url = http://www.repository.naturalis.nl/document/148848 | journal = Zoologische Verhandelingen | volume = 35 | pages = 1–204 }}
* Mees, G. F. (1961): A Systematic Review of the Indo-Australian Zosteropidae Parts II. Zoologische Verhandelingen 50:1–168 [http://www.repository.naturalis.nl/document/148911 PDF]
* {{cite journal | last1 = Mees | first1 = G. F. | year = 1961 | title = A Systematic Review of the Indo-Australian Zosteropidae Parts II. | url = http://www.repository.naturalis.nl/document/148911 | journal = Zoologische Verhandelingen | volume = 50 | pages = 1–168 }}
* Mees, G. F. (1969): A Systematic Review of the Indo-Australian Zosteropidae Parts III. Zoologische Verhandelingen 102:1–390 [http://www.repository.naturalis.nl/document/148979 PDF]
* {{cite journal | last1 = Mees | first1 = G. F. | year = 1969 | title = A Systematic Review of the Indo-Australian Zosteropidae Parts III | url = http://www.repository.naturalis.nl/document/148979 | journal = Zoologische Verhandelingen | volume = 102 | pages = 1–390 }}
* Mees, G F (1953): Article: An attempt at a natural classification of certain Zosteropidae of the Indo-Australian Archipelago. Zoologische Mededelingen 32:57–68 [http://www.repository.naturalis.nl/document/150367 PDF]
* {{cite journal | last1 = Mees | first1 = G F | year = 1953 | title = An attempt at a natural classification of certain Zosteropidae of the Indo-Australian Archipelago | url = http://www.repository.naturalis.nl/document/150367 | journal = Zoologische Mededelingen | volume = 32 | pages = 57–68 }}
* [[Charles Sibley|Sibley, Charles Gald]] & [[Jon Edward Ahlquist|Ahlquist, Jon Edward]] (1990): ''Phylogeny and classification of birds''. Yale University Press, New Haven, Conn.
* [[Charles Sibley|Sibley, Charles Gald]] & [[Jon Edward Ahlquist|Ahlquist, Jon Edward]] (1990): ''Phylogeny and classification of birds''. Yale University Press, New Haven, Conn.
*{{ cite journal | last1=Wells | first1=D.R. | year=2017a | title=''Zosterops'' white-eyes in continental South-East Asia. 1: proposed refinements to the regional definition of Oriental White-eye ''Z. palpebrosus'' | journal=Bulletin of the British Ornithologists' Club | volume=137 | issue=2 | pages=100-109 | doi=10.25226/bboc.v137i2.2017.a12 }}
*{{ cite journal | last1=Wells | first1=D.R. | year=2017a | title=''Zosterops'' white-eyes in continental South-East Asia. 1: proposed refinements to the regional definition of Oriental White-eye ''Z. palpebrosus'' | journal=Bulletin of the British Ornithologists' Club | volume=137 | issue=2 | pages=100–109 | doi=10.25226/bboc.v137i2.2017.a12 | doi-access=free }}
*{{ cite journal | last1=Wells | first1=D.R. | year=2017b | title=''Zosterops'' white-eyes in continental South-East Asia. 2: what is ''Zosterops auriventer'' Hume? | journal=Bulletin of the British Ornithologists' Club | volume=137 | issue=2 | pages=110-117 | doi=10.25226/bboc.v137i2.2017.a13 }}
*{{ cite journal | last1=Wells | first1=D.R. | year=2017b | title=''Zosterops'' white-eyes in continental South-East Asia. 2: what is ''Zosterops auriventer'' Hume? | journal=Bulletin of the British Ornithologists' Club | volume=137 | issue=2 | pages=110–117 | doi=10.25226/bboc.v137i2.2017.a13 | doi-access=free }}
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[[Category:Zosteropidae|*]]
[[Category:Zosteropidae|*]]

Latest revision as of 08:00, 7 September 2024

White-eyes
Silvereye (Zosterops lateralis), adult (right) and juveniles
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Superfamily: Sylvioidea
Family: Zosteropidae
Bonaparte, 1853
Genera

See text

Indian white-eye, Zosterops palpebrosus

The white-eyes are a family, Zosteropidae, of small passerine birds native to tropical, subtropical and temperate Sub-Saharan Africa, southern and eastern Asia, and Australasia. White-eyes inhabit most tropical islands in the Indian Ocean, the western Pacific Ocean, and the Gulf of Guinea. Discounting some widespread members of the genus Zosterops, most species are endemic to single islands or archipelagos. The silvereye, Zosterops lateralis, naturally colonised New Zealand, where it is known as the "wax-eye" or tauhou ("stranger"), from 1855. The silvereye has also been introduced to the Society Islands in French Polynesia, while the Japanese white-eye has been introduced to Hawaii.

Characteristics

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White-eyes are mostly of undistinguished appearance, the plumage being generally greenish olive above, and pale grey below. Some species have a white or bright yellow throat, breast, or lower parts, and several have buff flanks. As their common name implies, many species have a conspicuous ring of tiny white feathers around their eyes.[1] The scientific name of the group also reflects this latter feature, being derived from the Ancient Greek for "girdle-eye". They have rounded wings and strong legs. Like many other nectarivorous birds, they have slender, pointed bills, and brush-tipped tongues.[1][2] The size ranges up to 15 cm (5.9 in) in length.

All the species of white-eyes are sociable, forming large flocks that only separate on the approach of the breeding season. They build trees nests and lay two to four eggs which are usually pale blue.[3] Though mainly insectivorous, they eat nectar and fruits of various kinds. The silvereye can be a problem in Australian vineyards, by piercing the grape allowing infection or insect damage to follow.[4]

Systematics

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The family Zosteropidae was introduced (as a subfamily Zosteropinae) in 1853 by the French naturalist Charles Lucien Bonaparte.[5][6] The white-eyes were long considered a distinct family Zosteropidae because they are rather homogeneous in morphology and ecology, leading to little adaptive radiation and divergence.

The genus Apalopteron, formerly placed in the Meliphagidae, was transferred to the white eyes in 1995 on genetic and behavioral evidence. It differs much in appearance from the typical white-eyes, Zosterops, but is approached by some Micronesian taxa; its color pattern is fairly unusual save the imperfect white eye-ring.[7]

In 2003, Alice Cibois published the results of her study of mtDNA cytochrome b and 12S/16S rRNA sequence data. According to her results, the white-eyes were likely to form a clade also containing the yuhinas, which were until then placed with the Old World babblers, a large "wastebin" family.[8] Previous molecular studies (e.g. Sibley & Ahlquist 1990, Barker et al. 2002) had together with the morphological evidence tentatively placed white-eyes as the Timaliidae's closest relatives already. But some questions remained, mainly because the white-eyes are all very similar birds in habitus and habits, while the Old World babblers are very diverse (because, as we now know, the group as formerly defined was polyphyletic).

Combined with the yuhinas (and possibly other Timaliidae), the limits of the white-eye clade to the "true" Old World babblers becomes indistinct. Therefore, the current (early 2007) opinion weighs towards merging the group into the Timaliidae, perhaps as a subfamily ("Zosteropinae"). Few white-eyes have been thoroughly studied with the new results in mind, however, and almost all of these are from Zosterops which even at this point appears over-lumped. Also, many "Old World babblers" remain in unresolved relationships. Whether there can be a clear delimitation of a white-eye subfamily or even a young or emerging family is a question that requires a more comprehensive study of both this group and Timaliidae to resolve (Jønsson & Fjeldså 2006).

For example, a revision of the yuhinas and the genus Stachyris (Cibois et al. 2002), based on the same genes as Cibois (2003), revealed that the Philippine species placed in the latter genus by some were actually yuhinas. However, when the review by Jønsson & Fjeldså (2006) was published, no study had tried to propose a phylogeny for the newly defined yuhinas including the white eyes. Therefore, Jønsson & Fjeldså (2006) give a rather misleading phylogeny for the group. It appears as if the yuhinas are polyphyletic, with the white-collared yuhina being closer to the ancestor of the Zosterops white-eyes than to other yuhinas including the species moved from Stachyris (Cibois et al. 2002).

In the past, the Madanga (Madanga ruficollis) was included in this family but studies now place it as an atypical member of the Motacillidae.[9]

The cladogram below showing the relationships between families is based on a study of babblers by Tianlong Cai and collaborators published in 2019.[10][11]

Pycnonotidae – bulbuls (167 species)

Sylviidae – sylviid babblers (32 species)

Paradoxornithidae – parrotbills and myzornis (38 species)

Zosteropidae – white-eyes (152 species)

Timaliidae – tree babblers (58 species)

Pellorneidae – ground babblers (68 species)

Alcippeidae – Alcippe fulvettas (10 species)

Leiothrichidae – laughingthrushes and allies (133 species)

The cladogram below showing the relationships between the genera is based on the study by Carl Oliveros and collaborators that was published in 2021. The genera Apalopteron, Tephrozosterops and Rukia were not sampled in this study. The genus Megazosterops was found to be nested in Heleia.[12] The earlier study by Cai and collaborators found a generally similar phylogeny but with Cleptornis as sister to Heleia. Cai's study found that Apalopteron was nested within Heleia with weak support and that Tephrozosterops was sister to Zosterops.[10]

Zosteropidae

Parayuhina – white-collared yuhina

Staphida – yuhinas (3 species)

Yuhina – yuhinas (7 species)

Cleptornis – golden white-eye

Dasycrotapha – babblers (3 species)

Sterrhoptilus – babblers (4 species)

Zosterornis – babblers (5 species)

Heleia and Megazosterops – white-eyes (11 species altogether)

Zosterops – white-eyes, speirops and the silvereye (110 species)

List of genera

[edit]

The family contains 149 species divided into 13 genera:[11]

White-collared yuhina (Parayuhina diademata), a close relative of the white-eyes

References

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  1. ^ a b Lindsey, Terence (1991). Forshaw, Joseph (ed.). Encyclopaedia of Animals: Birds. London: Merehurst Press. p. 207. ISBN 1-85391-186-0.
  2. ^ van Balen 2008, p. 413.
  3. ^ van Balen 2008, p. 432.
  4. ^ van Balen 2008, p. 429.
  5. ^ Bonaparte, Charles Lucien (1853). "Classification ornithologique par series". Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences (in French). 37: 641–647 [644, No. 83].
  6. ^ Bock, Walter J. (1994). History and Nomenclature of Avian Family-Group Names. Bulletin of the American Museum of Natural History. Vol. 222. New York: American Museum of Natural History. pp. 154, 229. Linked page allows download of the 48MB pdf
  7. ^ Springer, Mark S.; Higuchi, Hiroyoshi; Ueda, Keisuke; Minton, Jason; Sibley, Charles G. (1995). "Molecular Evidence That the Bonin Islands "Honeyeater" Is a White-eye". Journal of the Yamashina Institute for Ornithology. 27 (2): 66–77_1. doi:10.3312/jyio1952.27.66.
  8. ^ Cibois, Alice (2003). "Mitochondrial DNA phylogeny of Babblers (Timaliidae)". The Auk. 120 (1): 35–54. doi:10.1093/auk/120.1.35.
  9. ^ Alstrom, P.; Jonsson, K. A.; Fjeldsa, J.; Odeen, A.; Ericson, P. G. P.; Irestedt, M. (2015). "Dramatic niche shifts and morphological change in two insular bird species". Royal Society Open Science. 2 (3): 140364. Bibcode:2015RSOS....240364A. doi:10.1098/rsos.140364. PMC 4448822. PMID 26064613.
  10. ^ a b Cai, T.; Cibois, A.; Alström, P.; Moyle, R.G.; Kennedy, J.D.; Shao, S.; Zhang, R.; Irestedt, M.; Ericson, P.G.P.; Gelang, M.; Qu, Y.; Lei, F.; Fjeldså, J. (2019). "Near-complete phylogeny and taxonomic revision of the world's babblers (Aves: Passeriformes)". Molecular Phylogenetics and Evolution. 130: 346–356. doi:10.1016/j.ympev.2018.10.010. PMID 30321696.
  11. ^ a b Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (January 2021). "ylviid babblers, parrotbills, white-eyes". IOC World Bird List Version 11.1. International Ornithologists' Union. Retrieved 17 June 2021.
  12. ^ Oliveros, C.H.; Andersen, M.J.; Moyle, R.G. (2021). "A phylogeny of white-eyes based on ultraconserved elements". Molecular Phylogenetics and Evolution. 164: 107273. doi:10.1016/j.ympev.2021.107273.

Sources

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Further reading

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