Old World Warbler

Old World warblers are a large group of birds formerly grouped together in the bird family Sylviidae. The family held over 400 species in over 70 genera, and were the source of much taxonomic confusion. Two families were split out initially, the cisticolas into Cisticolidae and the kinglets into Regulidae. In the past ten years they have been the subject of much research and many species are now placed into other families, including the Acrocephalidae, Cettiidae, Phylloscopidae, and Megaluridae. In addition some species have been moved into existing families or have not yet had their placement fully resolved. A smaller family of warblers, together with some babblers formerly placed in the family Timaliidae and the parrotbills, are retained in a much smaller family Sylviidae.

Characteristics

Most Old World warblers are of generally undistinguished appearance, though some Asian species are boldly marked. The sexes are often identical, but may be clearly distinct, notably in the genus '' Sylvia''. They are of small to medium size, varying from 9 to 16 centimetres in length, with a small, finely pointed bill. Almost all species are primarily insectivorous, although some will also eat fruit, nectar, or tiny seeds.

The majority of species are monogamous and build simple, cup-shaped nests in dense vegetation. They lay between two and six eggs per clutch, depending on species. Both parents typically help in raising the young, which are able to fly at around two weeks of age.

Systematics

In the late 20th century, the Sylviidae were thought to unite nearly 300 small insectivorous bird species in nearly 50 genera. They had themselves been split out of the Muscicapidae. The latter family had for most of its existence served as perhaps the ultimate wastebin taxon in the history of ornithology. By the early 20th century, about every insectivorous Old World "songster" known to science had at one point been placed therein, and most continued to be so.

Only after the mid-20th century did the dismantling of the "pan-Muscicapidae" begin in earnest. However, the Sylviidae remained a huge family, with few clear patterns of relationships recognisable. Though by no means as diverse as the Timaliidae (Old World babblers) (another "wastebin taxon" containing more thrush-like forms), the frontiers between the former "pan-Muscicapidae" were much blurred. The largely southern warbler family Cisticolidae was traditionally included in the Sylviidae. The kinglets, a small genus in a monotypic family Regulidae, were also frequently placed in this family. The American Ornithologists' Union includes the gnatcatchers, as subfamily Polioptilinae, in the Sylviidae.

Sibley & Ahlquist (1990) united the "Old World warblers" with the babblers and other taxa in a superfamily Sylvioidea as a result of DNA–DNA hybridisation studies. This demonstrated that the Muscicapidae as initially defined were a form taxon which collected entirely unrelated songbirds. Consequently, the monophyly of the individual "songster" lineages themselves was increasingly being questioned.

More recently, analysis of DNA sequence data has provided information on the Sylvioidea. Usually, the scope of the clade was vastly underestimated and only one or two specimens were sampled for each presumed "family". Minor or little-known groups such as the parrotbills were left out entirely (e.g. Ericson & Johansson 2003, Barker ''et al.'' 2004). These could only confirm that the Cisticolidae were indeed distinct, and suggested that bulbuls (Pycnonotidae) were apparently the closest relatives of a group containing Sylviidae, Timaliidae, cisticolids and white-eyes.

In 2003, a study of Timaliidae relationships (Cibois 2003a) using mtDNA cytochrome ''b'' and 12S/ 16S rRNA data indicated that the Sylviidae and Old World babblers were not reciprocally monophyletic to each other. Moreover, ''Sylvia'', the type genus of the Sylviidae, turned out to be closer to taxa such as the yellow-eyed babbler (''Chrysomma sinense'') (traditionally held to be an atypical timaliid) and the wrentit (''Chamaea fasciata''), an enigmatic species generally held to be the only American Old World babbler. The parrotbills, formerly considered a family Paradoxornithidae (roughly, "puzzling birds") of unclear affiliations also were part of what apparently was a well distinctive clade.

Cibois suggested that the Sylviidae should officially be suppressed by the ICZN as a taxon and the genus ''Sylvia'' merged into the Timaliidae (Cibois 2003b), but doubts remained. Clearly, the sheer extent of the groups concerned made it necessary to study a wide range of taxa. This was begun by Beresford ''et al.'' (2005) and Alström ''et al.'' (2006). They determined that the late-20th-century Sylviidae united at least four, but probably as many as seven major distinct lineages. The authors propose the creation of several new families ( Phylloscopidae, Cettiidae, Acrocephalidae, Megaluridae) to better reflect the evolutionary history of the sylvioid group.

The Sylviidae, in turn, receive several taxa from other families. Nonetheless, the now-monophyletic family has shrunk by nearly 80% for the time being, now containing 55 species in 10 genera at least. It is entirely likely however that with further research, other taxa from those still ''incertae sedis'' among its former contents, the Timaliidae, the Cisticolinae, or even the Muscicapidae will be moved into this group.

Species

Family Sylviidae ''sensu stricto''

True warblers (or sylviid warblers) and parrotbills. A fairly diverse group of smallish taxa with longish tails. Mostly in Asia, to a lesser extent in Africa. A few range into Europe; one monotypic genus on the west coast of North America.

* Genus '' Sylvia'' – typical warblers (c. 20 species). Paraphyletic or contains ''Parisoma''
** Temperate Eurasian superspecies ("''atricapilla-borin'' group")
*** Eurasian blackcap, ''Sylvia atricapilla''
*** Garden warbler, ''Sylvia borin''
** ''Parisoma'' superspecies
*** Brown parisoma, ''Sylvia lugens''
*** Banded parisoma, ''Sylvia boehmi''
*** Layard's warbler, ''Sylvia layardi''
*** Chestnut-vented warbler, ''Sylvia subcaeruleum''
** ''curruca'' clade
*** Yemen warbler, ''Sylvia buryi'' – sometimes placed in ''Parisoma''
*** Arabian warbler, ''Sylvia leucomelaena''
*** Western Orphean warbler, ''Sylvia hortensis''
*** Eastern Orphean warbler, ''Sylvia crassirostris''
*** Lesser whitethroat, ''Sylvia curruca''
*** Hume's whitethroat, ''Sylvia althaea''
*** Desert whitethroat, ''Sylvia minula''
**** Margelanic whitethroat, ''Sylvia (minula) margelanica''
** ''communis''-''melanocephala'' assemblage
*** Barred warbler, ''Sylvia nisoria'' – tentatively place here
*** Asian desert warbler, ''Sylvia nana''
*** African desert warbler, ''Sylvia deserti''
*** Common whitethroat, ''Sylvia communis''
*** Spectacled warbler, ''Sylvia conspicillata''
*** Tristram's warbler, ''Sylvia deserticola''
*** Dartford warbler, ''Sylvia undata''
*** Marmora's warbler, ''Sylvia sarda''
*** Balearic warbler, ''Sylvia balearica''
*** Rüppell's warbler, ''Sylvia rueppelli''
*** Cyprus warbler, ''Sylvia melanothorax''
*** Western subalpine warbler, ''Sylvia iberiae''
*** Eastern subalpine warbler, ''Sylvia cantillans''
*** Moltoni's warbler, ''Sylvia subalpina''
*** Sardinian warbler, ''Sylvia melanocephala''
**** ''Sylvia (melanocephala) momus''
**** Fayyum warbler, ''Sylvia melanocephala/momus norissae'' – doubtfully distinct, extinct (c. 1940)
*** Menetries's warbler, ''Sylvia mystacea''

* Genus ''Pseudoalcippe'' – African hill babbler. Formerly in ''Illadopsis'' (Timaliidae)
* Genus '' Rhopophilus'' – Chinese hill warbler. Formerly in Cisticolidae
* Genus '' Lioparus'' – golden-breasted fulvetta. Formerly in ''Alcippe'' (Timaliidae)
* Genus '' Paradoxornis'' – typical parrotbills (18 species). Formerly in Paradoxornithidae; polyphyletic
* Genus '' Conostoma'' – great parrotbill. Formerly in Paradoxornithidae; tentatively placed here
* Genus '' Fulvetta'' – typical fulvettas (7 species). Formerly in ''Alcippe'' (Timaliidae)
* Genus '' Chrysomma'' – 3 species. Formerly in Timaliidae
* Genus '' Chamaea'' – wrentit

Moved to family Pellorneidae

* Genus ''Graminicola''
** Rufous-rumped grassbird ("-babbler") ''Graminicola bengalensis''

Moved to family Cisticolidae

* Genus ''Bathmocercus'' – rufous-warblers
** Black-capped rufous-warbler, ''Bathmocercus cerviniventris''
** Black-faced rufous-warbler, ''Bathmocercus rufus''
* Genus ''Sceptomycter'' – sometimes merged into ''Bathmocercus''. Cisticolidae?
** Mrs Moreau's warbler, ''Sceptomycter winifredae''
* Genus '' Poliolais'' – Cisticolidae or more basal like bulbuls?
** White-tailed warbler, ''Poliolais lopezi''
* Two to 14 of the 15 tailorbirds

Moved to family Acrocephalidae

Marsh and tree warblers or acrocephalid warblers. Usually rather large "warblers", most are olivaceous brown above with much yellow to beige below. Usually in open woodland, reed beds or tall grass. Mainly southern Asia to western Europe and surroundings ranging far into Pacific, some in Africa. The genus limits are seriously in need of revision; either most species are moved into ''Acrocephalus'', or the latter is split up though there is presently insufficient knowledge as to how.

* Genus '' Acrocephalus'' – marsh warblers (about 35 species)
* Genus ''Hippolais'' – tree warblers (8 species)
* Genus ''Chloropeta'' – yellow warblers (3 species)
* Genus '' Nesillas'' – brush warblers (4 living species, 1 recently extinct)

Moved to Malagasy warblers

See Cibois ''et al.'' (2001)
* Genus ''Thamnornis''
** Thamnornis, ''Thamnornis chloropetoides''
* Genus '' Cryptosylvicola''
** Cryptic warbler, ''Cryptosylvicola randriansoloi''

Moved to family Locustellidae

Grass warblers and allies. Mid-sized and usually long-tailed species; sometimes strongly patterned but generally very drab in overall colouration. Often forage on the ground. Old World and into Australian region, centred on the Indian Ocean; possibly also one species in South America. A not too robustly supported clade that requires further study.

* Genus '' Bradypterus'' – megalurid bush-warblers (11 species)
* Genus '' Locustella'' – grass warblers (more than 20 species)
* Genus '' Megalurus'' – typical grassbirds (10 species)
* Genus ''Amphilais'' – grey emutail
* Genus '' Elaphrornis'' – Sri Lanka bush warbler
* Genus ''Schoenicola'' – (2 species)
* Genus '' Buettikoferella'' – buff-banded thicketbird
* Genus ''Chaetornis'' – bristled grassbird

The black-capped donacobius, ''Donacobius atricapillus'', which was long considered an aberrant wren or mockingbird is apparently quite closely related, and might possibly be considered the only American species of this family.

Moved to family Cettiidae

Typical bush warblers and relatives or cettiid warblers. Another group of generally very drab species, tend to be smaller and shorter-tailed than Megaluridae. Usually frequent shrubland and undergrowth. Continental Asia, and surrounding regions, ranging into Africa and southern Europe.

* Genus ''Pholidornis'' – formerly in Remizidae; tentatively placed here
** Tit hylia, ''Pholidornis rushiae''
* Genus ''Hylia'' – tentatively placed here
** Green hylia, ''Hylia prasina''
* Genus '' Abroscopus'' – Abroscopus warblers
** Rufous-faced warbler, ''Abroscopus albogularis''
** Yellow-bellied warbler, ''Abroscopus superciliaris''
** Black-faced warbler, ''Abroscopus schisticeps''

* Genus '' Erythrocercus'' – monarch-warblers. Formerly Monarchinae.
** Chestnut-capped flycatcher, ''Erythrocercus mccallii''
** Little yellow flycatcher, ''Erythrocercus holochlorus''
** Livingstone's flycatcher, ''Erythrocercus livingstonei''
* Genus '' Urosphena'' – stubtails
** Timor stubtail, ''Urosphena subulata''
*** Babar stubtail, ''Urosphena subulata advena'' – extinct (mid-20th century)
** Bornean stubtail, ''Urosphena whiteheadi''
** Asian stubtail, ''Urosphena squameiceps''
** Pale-footed bush warbler, ''Urosphena pallidipes''
** Neumann's warbler, ''Urosphena neumanni''
* Genus '' Tesia'' – tesias
** Javan tesia, ''tesia superciliaris''
** Slaty-bellied tesia, ''Tesia olivea''
** Grey-bellied tesia, ''Tesia cyaniventer''
** Russet-capped tesia, ''Tesia everetti''
* Genus '' Horornis'' – bush warblers (some 13 species).
* Genus '' Cettia'' – bush warblers (4 species).
* Genus '' Tickellia''
** Broad-billed warbler, ''Tickellia hodgsoni''
* Genus '' Phyllergates''
** Mountain tailorbird, ''Phyllergates cucullatus''
** Rufous-headed tailorbird, ''Phyllergates heterolaemus''

Moved to family

Aegithalidae

* Genus '' Leptopoecile'' – tit-warblers. Tentatively placed there.
** White-browed tit-warbler, ''Leptopoecile sophiae''
** Crested tit-warbler, ''Leptopoecile elegans''

Moved to family Phylloscopidae

Leaf warblers or phylloscopid warblers. A group very variable in size, often vivid green colouration above and yellow below, or more subdued with greyish-green to greyish-brown plumage. Catch food on the wing fairly often. Eurasia, ranging into Wallacea and Africa.

* Genus '' Phylloscopus'' – leaf warblers (c. 55 species). (includes former genus '' Seicercus'')
** Green-crowned warbler, ''Phylloscopus burkii''
** Grey-crowned warbler, ''Phylloscopus tephrocephalus''
** Whistler's warbler, ''Phylloscopus whistleri''
** Bianchi's warbler, ''Phylloscopus valentini''
** Martens's warbler, ''Phylloscopus omeiensis''
** Alström's warbler, ''Seicercus soror''
** White-spectacled warbler, ''Phylloscopus affinis'' – paraphyletic
*** Bar-winged white-spectacled warbler, ''Seicercus (affinis) intermedius''
** Grey-cheeked warbler, ''Phylloscopus poliogenys''
** Chestnut-crowned warbler, ''Phylloscopus castaniceps''
** Yellow-breasted warbler, ''Phylloscopus montis''
** Sunda warbler, ''Phylloscopus grammiceps''

Moved to family Macrosphenidae

African warblers. Also "''Sphenoeacus'' group". An assemblage of usually species-poor and apparently rather ancient "odd warblers" from Africa. Ecomorphologically quite variable. Monophyly requires confirmation.

* Genus ''Sylvietta'' – crombecs
** Green crombec, ''Sylvietta virens''
** Lemon-bellied crombec, ''Sylvietta denti''
** White-browed crombec, ''Sylvietta leucophrys''
*** Chapin's crombec, ''Sylvietta (leucophrys) chapini'' – possibly extinct (late 20th century?)
** Northern crombec, ''Sylvietta brachyura''
** Philippa's crombec, ''Sylvietta philippae''
** Red-capped crombec, ''Sylvietta ruficapilla''
** Red-faced crombec, ''Sylvietta whytii''
** Somali crombec, ''Sylvietta isabellina''
** Long-billed crombec, ''Sylvietta rufescens''
* Genus '' Melocichla''
** Moustached grass warbler, ''Melocichla mentalis''
* Genus ''Achaetops''
** Rockrunner, ''Achaetops pycnopygius''
* Genus '' Sphenoeacus''
** Cape grassbird, ''Sphenoeacus afer''
* Genus ''Cryptillas''.
** Victorin's warbler, ''Cryptillas victorini''
* Genus ''Macrosphenus'' – longbills
** Kemp's longbill, ''Macrosphenus kempi''
** Yellow longbill, ''Macrosphenus flavicans''
** Grey longbill, ''Macrosphenus concolor''
** Pulitzer's longbill, ''Macrosphenus pulitzeri''
** Kretschmer's longbill, ''Macrosphenus kretschmeri''

"Sylviidae" ''incertae sedis''

Taxa that have not been studied. Most are likely to belong to one of Sylvioidea families listed above. Those in the Australian-Pacific region are probably Megaluridae. These taxa are listed in the sequence used in recent years.

* Genus ''Dromaeocercus'' – emutails. Locustelidae?
** Brown emutail, ''Dromaeocercus brunneus''
** Grey emutail, ''Dromaeocercus seebohmi'' – sometimes separated in ''Amphilais''
* Genus ''Phyllolais'' – Cisticolidae?
** Buff-bellied warbler, ''Phyllolais pulchella''
* Genus ''Graueria''
** Grauer's warbler, ''Graueria vittata''
* Genus '' Eremomela'' – eremomelas. Cettiidae?
** Salvadori's eremomela, ''Eremomela salvadorii''
** Yellow-vented eremomela, ''Eremomela flavicrissalis''
** Yellow-bellied eremomela, ''Eremomela icteropygialis''
** Senegal eremomela, ''Eremomela canescens''
** Green-backed eremomela, ''Eremomela pusilla''
** Green-capped eremomela, ''Eremomela scotops''
** Yellow-rumped eremomela, ''Eremomela gregalis''
** Rufous-crowned eremomela, ''Eremomela badiceps''
** Turner's eremomela, ''Eremomela turneri''
*** Western Turner's eremomela, ''Eremomela turneri kalindei'' – probably extinct (early 1980s?)
** Black-necked eremomela, ''Eremomela atricollis''
** Burnt-neck eremomela, ''Eremomela usticollis''
* Genus '' Randia'' – Malagasy warblers?
** Rand's warbler, ''Randia pseudozosterops''
* Genus '' Bowdleria'' – fernbirds. Sometimes merged into ''Megalurus''. Locustellidae?
** Fernbird, ''Bowdleria punctata''
** Chatham fernbird, ''Bowdleria rufescens'' – extinct (c. 1900)
* Genus ''Chaetornis'' – bristled grassbird. Locustellidae?
* Genus ''Schoenicola'' – grassbirds. Basal Locustellidae?
** Broad-tailed grassbird, ''Schoenicola platyura''
** Fan-tailed grassbird, ''Schoenicola brevirostris''
* Genus ''Cincloramphus'' – songlarks. Basal Locustellidae?
** Brown songlark, ''Cincloramphus cruralis''
** Rufous songlark, ''Cincloramphus mathewsi''
* Genus '' Buettikoferella'' – probably Locustellidae
** Buff-banded bushbird, ''Buettikoferella bivittata''
* Genus ''Megalurulus'' – thicketbirds. Probably Locustellidae
** New Caledonian grassbird, ''Megalurulus mariei''
** Bismarck thicketbird, ''Megalurulus grosvenori''
** Bougainville thicketbird, ''Megalurulus llaneae''
** Santo thicketbird, ''Megalurulus whitneyi''
** Rusty thicketbird, ''Megalurulus rubiginosus''
* Genus '' Trichocichla'' – long-legged warbler

Not in Sylvioidea

Entirely unrelated songbirds hitherto placed in Sylviidae

* Genus '' Amaurocichla'' – Apparently a Passeroidea; very close to, or part of the Motacillidae
** Bocage's longbill or São Tomé short-tail, ''Amaurocichla bocagei''
* Genus ''Stenostira'' – Together with some "odd flycatchers", they form the new family Stenostiridae. They are closely related to Paridae (Beresford ''et al.'' 2005)
** Fairy flycatcher, ''Stenostira scita''
* Genus ''Hyliota'' – hyliotas. Basal Passerida with no known relatives, perhaps somewhat closer to Promeropidae (sugarbirds)
** Yellow-bellied hyliota, ''Hyliota flavigaster''
** Southern hyliota, ''Hyliota australis''
** Usambara hyliota, ''Hyliota usambarae''
** Violet-backed hyliota, ''Hyliota violacea''
* Genus '' Newtonia'' – newtonias. Now in Vangidae (vangas); possibly polyphyletic (Yamagishi ''et al.'' 2001)
** Dark newtonia, ''Newtonia amphichroa''
** Common newtonia, ''Newtonia brunneicauda''
** Archbold's newtonia, ''Newtonia archboldi''
** Red-tailed newtonia, ''Newtonia fanovanae'' – tentatively placed here

See also

* List of extinct birds

Notes

References

*
*Baker, K. (1997). ''Warblers of Europe, Asia, and North Africa''. Helm .

Supporting information

Electronic appendix
*
*
*
*del Hoyo, J.; Elliot, A. & Christie D. (editors). (2006). ''Handbook of the Birds of the World''. Volume 11: Old World Flycatchers to Old World Warblers. Lynx Edicions. .
*
*
*Shirihai, H., Gargallo, G., & Helbig, A. J. (2001). ''Sylvia Warblers''. Helm .
* Sibley, C. G. & Ahlquist, J. E. (1990). ''Phylogeny and classification of birds''. Yale University Press, New Haven, Conn.
*Simms, E. (1985). ''British warblers''. Collins, London. .
*

External links

Old World warblers videos
on the Internet Bird Collection
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{{Interwiki extra, qid=Q187014

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Bird common names