Old World babbler

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Babblers
The White-naped Yuhina (Yuhina bakeri) would be moved to the Zosteropidae if these are retained
The White-naped Yuhina (Yuhina bakeri) would be moved to the Zosteropidae if these are retained
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Suborder: Passeri
Family: Timaliidae
Vigors & Horsfield, 1827
Genera

Dozens; see article text

The Old World babblers or timaliids are a large family of mostly Old World passerine birds. They are rather diverse in size and coloration, but are characterised by soft fluffy plumage. These are birds of tropical areas, with the greatest variety in southeast Asia. The American Wrentit is an enigmatic species that was in recent times placed with the Old World babblers but may, in fact, not belong here. The timaliids are one of two unrelated groups of birds known as babblers, the other being the Australasian Babblers of the family Pomatostomidae (also known as pseudo-babblers).

These birds have strong legs, and many are quite terrestrial. This group is not strongly migratory, and most species have short rounded wings, and a weak flight. Morphological diversity is rather high; most species resemble "warblers", jays or thrushes. This group is among those Old World bird families with the highest number of species still being discovered.

Contents

[edit] Systematics

The systematics of Old World babblers have long been contended. During much of the 20th century, the family was used as a "wastebin taxon" for numerous hard-to-place Old World songbirds (such as Picathartidae or the Wrentit). Ernst Hartert was only half-joking when he summarized this attitude with the statement that, in the passerines,

"Was man nicht unterbringen kann, sieht man als Timalien an." (What one can't place systematically is considered an Old World babbler)

The most obviously misplaced taxa were removed piecemeal towards the end of the last century.

Since then, with the aid of DNA sequence data, it has been confirmed that even the remaining group is not monophyletic. Analysis of mtDNA cytochrome b and 12S/16S rRNA data (Cibois 2003a) spread the Timaliidae that were studied across what essentially was a badly resolved polytomy with Old World warblers and white-eyes. As the typical warblers (genus Sylvia) grouped with some presumed timaliids (such as the fulvettas), it was suggested that some Sylviidae should be moved to the Timaliidae.

As this would include the type genus of the latter, this would lead to a nomenclatorial problem requiring ICZN intervention (Cibois 2003b) and was, at that time, not sensible in any case as the phylogeny of the remaining Old World warblers had not been fully resolved either. The problem with such an approach would be — as many Old World warblers have not been studied with the new results in mind and neither have a number of timaliids — to risk creating a huge, ill-defined family-level clade; consequently, this approach seems to have been put on hold for the time being in favor of a general resorting of the Sylvioidea.

Alström et al. (2006) supported the taxonomic proposal of Cibois (2003b), "if the Timaliidae and several groups of warblers are recognized at the same family level" but of course it is not necessary to unite them to achieve monophyly in both. Notably, one of the few conclusions beyond genus level which received quite robust support in Cibois (2003a) was the distinctness of Sylvia and the related "babblers" from the Timaliidae sensu stricto. Thus, for the time being, it seems wisest to maintain the Sylviidae and Timaliidae as distinct and just split off or move about genera as needed to achieve monophyly.

The parrotbills are somewhat titmouse-like birds that in the past were moved about between the timaliids, the titmice, and own family status (under the telling name Paradoxornithidae — literally, "puzzling birds"). They are certainly not a distinct family; rather, they belong into the Sylvia clade (Cibois 2003a, Alström et al. 2006).

The relationships of the white-eyes (presently Zosteropidae) are not resolved at present. Based on nDNA RAG-1 and c-mos sequence data, Barker et al. (2002) found them likelier to group closer to the timaliids proper than to Sylvia and allies, as did Cibois (2003a). Combining data from nDNA c-myc exon 3, RAG-1 and myoglobin intron 2 sequences with that of mtDNA cytochrome b (Ericson & Johansson 2003) supports their scenario as does a restudy using the myoglobin intron 2 and cytochrome b sequences of a wider (though not denser) range of taxa (Alström et al. 2006)

On the other hand, DNA-DNA hybridization (Sibley & Ahlquist 1990) placed the white-eyes closer to Sylvia. This method is nowadays considered inferior to comparison of long and various DNA sequences, however. Still, it should be noted that no molecular study thus far could resolve the white-eyes' relationships with sufficient confidence beyond the mere fact that they form a clade with "core" Sylviidae and "core" Timaliidae. In this assemblage, they most likely form a monophyletic lineage with the yuhinas (and possibly other "babblers"). Consequently, were the Zosteropidae to be retained as a family, these would be moved there.

In addition, the new studies have shown that several genera (such as Garrulax and Fulvetta) are not monophyletic and need to be split up (Cibois 2003, Pasquet et al. 2006).

[edit] Timaliidae sensu stricto

These genera would be retained in the timaliids in any case; one has been moved here from the sylviids. They make up a few reasonably well-supported clades and a lot of genera with quite unresolved relationships (termed "assemblages" here):

Red-billed Leiothrix, Leiothrix lutea
Red-billed Leiothrix, Leiothrix lutea

Liocichlas and allies clade

Laughingthrush assemblage

White-crested Laughingthrush, Garrulax leucolophus
White-crested Laughingthrush, Garrulax leucolophus
  • Genus Cutia - cuitas (2 species)
  • Genus Garrulax – laughingthrushes (formerly c.50 species). Polyphyletic, consisting of around 11 genera for which see Garrulax.

Pellorneum – Napothera assemblage

Black-capped Babbler, Pellorneum capistratum
Black-capped Babbler, Pellorneum capistratum
  • Genus Pellorneum (7 species) - possibly polyphyletic (Jønsson & Fjeldså 2006)
  • Genus Pseudominla – 4 species of atypical alcippes or "fulvettas", formerly in Alcippe
The Black-lored Babbler (Turdoides melanops) of Africa may comprise more than one species, in which case this bird is T. sharpei.
The Black-lored Babbler (Turdoides melanops) of Africa may comprise more than one species, in which case this bird is T. sharpei.
  • Genus Jabouilleia – atypical scimitar-babblers (2 species)

Timaliine clade

The Mountain Fulvetta (Alcippe peracensis) belongs to the typical alcippes
The Mountain Fulvetta (Alcippe peracensis) belongs to the typical alcippes
  • Genus Spelaeornis (9 species) – typical wren-babblers
  • Genus Macronous – tit-babblers (5 species)

[edit] White-eye clade

If the white-eyes are maintained as a separate family Zosteropidae, this group would have to be included there:

  • Genus Yuhina – yuhinas – 8 species, (found to be polyphyletic and now split into five genera)

[edit] Sylviid clade

The Yellow-eyed Babbler (Chrysomma sinense) is closer to the typical warblers
The Yellow-eyed Babbler (Chrysomma sinense) is closer to the typical warblers

Also includes the parrotbills. If the Sylviidae are maintained as a separate family, this group would have to be included there:

[edit] Genera incertae sedis

Relationships unresolved; may or may not be Timaliidae sensu stricto

  • Genus Crocias – 2 species of crociases (laughingthrush assemblage?)
  • Genus Micromacronus - 2 species of miniature-babblers or tit-babblers (timaliine clade?)
  • Genus Trichastoma – 7 species, Pellorneum – Napothera assemblage?
The Wrentit's relationships have long been elusive.It turned out to related to the typical warblers, too
The Wrentit's relationships have long been elusive.
It turned out to related to the typical warblers, too

[edit] Formerly placed here

Genera whose relationships are now known to lie entirely outside the Timaliidae, no matter how these are delimited

[edit] Other Sylvioidea

Basal lineage of cisticolid warblers, Cisticolidae (Nguembock et al. 2007):

Belong to an as yet unnamed Malagasy warbler clade ("Bernieridae"); see Cibois et al. 2001.

[edit] Other Passerida

Belong to the sugarbirds, Promeropidae (Beresford et al. 2005):

[edit] "Corvida"

Apparently related to the vireos, Vireonidae (Barker et al. 2004):

Belongs to the vireos (Reddy & Cracraft, in press):

Belongs to the vangas, Vangidae (Schulenberg 2003):

[edit] Incertae sedis

[edit] References

  • 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. doi:10.1016/j.ympev.2005.05.015
  • 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. Proc. R. Soc. B 269(1488): 295-308. doi:10.1098/rspb.2001.1883 PDF fulltext
  • Cibois, Alice (2003a): Mitochondrial DNA Phylogeny of Babblers (Timaliidae). Auk 120(1): 1-20. DOI: 10.1642/0004-8038(2003)120[0035:MDPOBT]2.0.CO;2 HTML fulltext without images
  • Cibois, Alice (2003b): Sylvia is a babbler: taxonomic implications for the families Sylviidae and Timaliidae.Bull. B. O. C. 123: 257-261.
  • Cibois, Alice; Slikas, Beth; Schulenberg, Thomas S. & Pasquet, Eric (2001): An endemic radiation of Malagasy songbirds is revealed by mitochondrial DNA sequence data. Evolution 55(6): 1198-1206. DOI:10.1554/0014-3820(2001)055[1198:AEROMS]2.0.CO;2 PDF fulltext
  • Cibois, Alice; Kalyakin, Mikhail V.; Lian-Xian, Han & Pasquet, Eric (2002): Molecular phylogenetics of babblers (Timaliidae): revaluation of the genera Yuhina and Stachyris. J. Avian Biol. 33: 380–390. doi:10.1034/j.1600-048X.2002.02882.x (HTML abstract)
  • Collar, N. J. & Robson, C. 2007. Family Timaliidae (Babblers) pp. 70–291 in; del Hoyo, J., Elliott, A. & Christie, D.A. eds. Handbook of the Birds of the World, Vol. 12. Picathartes to Tits and Chickadees. Lynx Edicions, Barcelona.
  • Ericson, Per G.P. & Johansson, Ulf S. (2003): Phylogeny of Passerida (Aves: Passeriformes) based on nuclear and mitochondrial sequence data. Molecular Phylogenetics and Evolution 29(1): 126–138 doi:10.1016/S1055-7903(03)00067-8 PDF fulltext
  • Nguembock, Billy; Fjeldså, Jon; Tillier, Annie & Pasquet, Eric (2007): A phylogeny for the Cisticolidae (Aves: Passeriformes) based on nuclear and mitochondrial DNA sequence data, and a re-interpretation of a unique nest-building specialization. Molecular Phylogenetics and Evolution 42: 272-286. doi:10.1016/j.ympev.2006.07.008 (HTML abstract)
  • Reddy, Sushma & Cracraft, Joel (in press): Old World Shrike-babblers (Pteruthius) belong with New World Vireos (Vireonidae). Molecular Phylogenetics and Evolution, In Press, Accepted Manuscript, Available online 28 February 2007. doi:10.1016/j.ympev.2007.02.023 (HTML abstract)
  • Schulenberg, T.S. (2003): The Radiations of Passerine Birds on Madagascar. In: Goodman, Steven M. & Benstead, Jonathan P. (eds.): The Natural History of Madagascar: 1130-1134. University of Chicago Press. ISBN 0-226-30306-3

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