Old World warblers are a large group of birds formerly grouped together in the bird family Sylviidae. They are not closely related to the
New World warblers. 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 number 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.[1]
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.[1]
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.[citation needed] 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.[2]
Sibley & Ahlquist (1990) united the "Old World warblers" with the babblers and other
taxa in a
superfamilySylvioidea 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
mtDNAcytochrome 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.
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)
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)
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.
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
Leaf warblers or phylloscopes. 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.
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.
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 Stenostira – Together with some "odd flycatchers", they form the new family
Stenostiridae. They are closely related to
Paridae (Beresford et al. 2005)
^Sefc, K. M.; Payne, R. B.; Sorenson, M. D. (2003). "Phylogenetic relationships of African sunbird-like warblers: Moho Hypergerus atriceps, Green Hylia Hylia prasina and Tit-hylia Pholidornis rushiae". Ostrich. 74: 8–17.
doi:
10.2989/00306520309485365.
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^Johansson, U.S.; Fjeldså, J.; Bowie, R.C.K. (2008). "Phylogenetic relationships within Passerida (Aves: Passeriformes): A review and a new molecular phylogeny based on three nuclear intron markers". Mol. Phylogenet. Evol. 48 (3): 858–876.
doi:
10.1016/j.ympev.2008.05.029.
PMID18619860.
^Fuchs, J.; Fjeldså, J.; Bowie, R. C. K.; Voelker, G.; Pasquet, E. (2006). "The African warbler genus Hyliota as a lost lineage in the oscine songbird tree: Molecular support for an African origin of the Passerida". Mol. Phylogenet. Evol. 39 (1): 186–197.
doi:
10.1016/j.ympev.2005.07.020.
PMID16182572.
References
Alström, P.; Ericson, P. G. P.; Olsson, U.; Sundberg, P. (2006). "Phylogeny and classification of the avian superfamily Sylvioidea". Molecular Phylogenetics and Evolution. 38 (2): 381–397.
doi:
10.1016/j.ympev.2005.05.015.
PMID16054402.
Baker, K. (1997). Warblers of Europe, Asia, and North Africa. Helm
ISBN0-7136-3971-7.
Fuchs, J.; Fjeldsa, J.; Bowie, R. C. K.; Voelker, G.; Pasquet, E. (2006). "The African warbler genus Hyliota as a lost lineage in the Oscine songbird tree: Molecular support for an African origin of the Passerida". Molecular Phylogenetics and Evolution. 39 (1): 186–197.
doi:
10.1016/j.ympev.2005.07.020.
PMID16182572.
Shirihai, H., Gargallo, G., & Helbig, A. J. (2001). Sylvia Warblers. Helm
ISBN0-7136-3984-9.
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.
ISBN0-00-219404-X.