|Red-billed Leiothrix, Leiothrix lutea|
Vigors & Horsfield, 1827
Sylviidae (as a tribe, Timaliini), sensu Sibley and Monroe
The babblers, 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 and the Indian Subcontinent. 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).
Morphological diversity is rather high; most species resemble "warblers", jays or thrushes. Along with Sylviidae, Leiothrichidae, Pellorneidae and others; this group is among those Old World bird families with the highest number of species still being discovered.
Timaliids are small to medium birds, ranging in size from the Pygmy Wren-babbler at 9 centimetres in length, to the Giant Laughingthrush at 36 cm. They have strong legs, and many are quite terrestrial. They typically have generalised bills, similar to those of a thrush or warbler, except for the scimitar babblers which, as their name implies, have strongly decurved bills. Most have predominantly brown plumage, with minimal difference between the sexes, but many more brightly coloured species also exist.
This group is not strongly migratory, and most species have short rounded wings, and a weak flight. They live in lightly wooded or scrubland environments, ranging from swamp to near-desert. They are primarily insectivorous, although many will also take berries, and the larger species will even eat small lizards and other vertebrates.
Typical babblers live in communities of around a dozen birds, jointly defending a territory. Many even breed communally, with a dominant pair building a nest, and the remainder helping to defend and rear their young. Young males remain with the group, while females move away to find a new group, and thus avoid inbreeding. They make nests from twigs, and hide them in dense vegetation.
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)Template:CitethisquoteThe 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 tit-like birds that in the past were moved about between the timaliids, the tits, and distinct family status (under the telling name Paradoxornithidae — literally, "puzzling birds"). They are likely 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.
One somewhat controversial approach in the "Taxonomy in Flux" world birdlist proposes splitting the group into four families.
- Sylviidae - Sylvias, Fulvettas, Parrotbills.
- Zosteropidae - White-eyes, Yuhinas etc.
- Timaliidae - Babblers.
- Garrulacidae - Laughingthrushes, Liocichlas, Sibias etc.
This creates a tidier, more manageable arrangement, much as the creation of several smaller families does for the traditional Emberizidae. It remains to be seen though if the innovative split of Garrulacidae from Timaliidae will gain wider acceptance.
List of Species
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).
- Genus Mixornis
- Genus Cyanoderma
- Golden Babbler, Cyanoderma chrysaea
- Chestnut-winged Babbler, Cyanoderma erythroptera
- Crescent-chested Babbler, Cyanoderma melanothorax
- Black-chinned Babbler, Cyanoderma pyrrhops
- Rufous-capped Babbler, Cyanoderma ruficeps
- Buff-chested Babbler, Cyanoderma ambigua
- Rufous-fronted Babbler, Cyanoderma rufifrons
- Genus Spelaeornis
- Genus Pomatorhinus
- Coral-billed Scimitar-Babbler, Pomatorhinus ferruginosus
- Red-billed Scimitar-Babbler, Pomatorhinus ochraceiceps
- Slender-billed Scimitar-Babbler, Pomatorhinus superciliaris
- Streak-breasted Scimitar-Babbler, Pomatorhinus ruficollis
- Taiwan Scimitar-Babbler, Pomatorhinus musicus
- Chestnut-backed Scimitar-Babbler, Pomatorhinus montanus
- White-browed Scimitar-Babbler, Pomatorhinus schisticeps
- Indian Scimitar-Babbler, Pomatorhinus horsfieldii
- Sri Lanka Scimitar-Babbler, Pomatorhinus melanurus
- Genus Megapomatorhinus
- Large Scimitar-Babbler, Megapomatorhinus hypoleucos
- Rusty-cheeked Scimitar-Babbler, Megapomatorhinus erythrogenys
- Spot-breasted Scimitar-Babbler, Megapomatorhinus mcclellandi
- Black-streaked Scimitar-Babbler, Megapomatorhinus gravivox
- Grey-sided Scimitar-Babbler, Megapomatorhinus swinhoei
- Black-necklaced Scimitar-Babbler, Megapomatorhinus erythrocnemis
- Genus Stachyris
- Nonggang Babbler, Stachyris nonggangensis
- Sooty Babbler, Stachyris herberti
- Black-throated Babbler, Stachyris nigricollis
- White-breasted Babbler, Stachyris grammiceps
- Chestnut-rumped Babbler, Stachyris maculata
- Grey-throated Babbler, Stachyris nigriceps
- Grey-headed Babbler, Stachyris poliocephala
- Sikkim Wedge-billed Babbler, Stachyris humei
- Cachar Wedge-billed Babbler, Stachyris roberti
- Snowy-throated Babbler, Stachyris oglei
- Spot-necked Babbler, Stachyris strialata
- White-necked Babbler, Stachyris leucotis
- White-bibbed Babbler, Stachyris thoracica
White-eye cladeIf 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)
- Genus Staphida
- Genus Dasycrotapha – formerly in Stachyris; tentatively placed here
- Flame-templed Babbler, Dasycrotapha speciosa
- Genus Sterrhoptilus – formerly in Stachyris; tentatively placed here
- Genus Zosterornis – formerly in Stachyris; tentatively placed here
- Genus Fulvetta – 7 species of typical fulvettas, formerly in Alcippe
- Spectacled Fulvetta, Fulvetta ruficapilla
- Chinese Fulvetta, Fulvetta striaticollis
- White-browed Fulvetta, Fulvetta vinipectus
- Grey-hooded Fulvetta, Fulvetta cinereiceps
- Taiwan Fulvetta, Fulvetta formosana - formerly in F. cinereiceps
- Streak-throated Fulvetta, Fulvetta manipurensis - formerly in F. cinereiceps
- Ludlow's Fulvetta, Fulvetta ludlowi - tentatively placed here
- Genus Chrysomma – 3 species
- Genus Chamaea – Wrentit
Genera incertae sedis
Relationships unresolved; may or may not be Timaliidae sensu stricto
- Genus Micromacronus - 2 species of miniature-babblers or tit-babblers (timaliine clade?)
- Genus Trichastoma – 7 species, Pellorneum – Napothera assemblage?
- Genus Rimator - Long-billed Wren-babbler
- Genus Ptilocichla – 3 species of wren-babblers
- Genus Pnoepyga – 3 species of wren-babblers
- Genus Dumetia – Tawny-bellied Babbler
- Genus Rhopocichla – Dark-fronted Babbler
- Genus Myzornis - Fire-tailed Myzornis
- Genus Malia – Malia
- Genus Lioptilus - Bush Blackcap (may be related to Sylvia)
- Genus Parophasma – Abyssinian Catbird (may be related to Sylvia)
- Genus Ptyrticus – Thrushbabbler (may be related to Illadopsis)
- Genus Horizorhinus – Dohrn's Thrush-babbler or Principe Flycatcher-babbler (relationships uncertain)
Formerly placed here
Genera whose relationships are now known to lie entirely outside the Timaliidae, no matter how these are delimited:
- Genus Neomixis – jeries
Several lineages of Malagasy warblers, Bernieridae
- Genus Hartertula – formerly in Neomixis
- Wedge-tailed Jery, Hartertula flavoviridis
- Genus Crossleyia
- Yellow-browed Oxylabes, Crossleyia xanthophrys
- Genus Oxylabes
- White-throated Oxylabes, Oxylabes madagascariensis
Belong to the sugarbirds, Promeropidae (Beresford et al. 2005):
- Genus Arcanator – sometimes included in Modulatrix
- Dapple-throat, Arcanator orostruthus
- Genus Modulatrix
- Spot-throat, Modulatrix stictigula
Apparently related to the vireos, Vireonidae (Barker et al. 2004):
- Genus Erpornis
- White-bellied Erpornis or White-bellied "Yuhina", Erpornis zantholeuca – formerly in Yuhina (Cibois et al. 2002)
Belongs to the vireos (Reddy & Cracraft, in press):
- Genus Pteruthius – shrike-babblers
Belongs to the vangas, Vangidae (Schulenberg 2003):
- Genus Kakamega (Cibois 2003a) – uncertain relationships. May be related to thrushes or sugarbirds
- Grey-chested Kakamega or Grey-chested "Illadopsis", Kakamega poliothorax
- 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 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. Proc. R. Soc. B 269(1488): 295-308. doi:10.1098/rspb.2001.1883 PDF fulltext
- Barker, F. Keith; Cibois, Alice; Schikler, Peter A.; Feinstein, Julie & Cracraft, Joel (2004): Phylogeny and diversification of the largest avian radiation. PNAS 101(30): 11040-11045. doi:10.1073/pnas.0401892101 PMID 15263073 PDF fulltext Supporting information
- Beresford, P.; Barker, F.K.; Ryan, P.G. & Crowe, T.M. (2005): African endemics span the tree of songbirds (Passeri): molecular systematics of several evolutionary 'enigmas'. Proc. Roy. Soc. Lond. B 272(1565): 849–858. doi:10.1098/rspb.2004.2997 PDF fulltext Electronic appendix
- 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
- Jønsson, Knud A. & Fjeldså, Jon (2006): A phylogenetic supertree of oscine passerine birds (Aves: Passeri). Zool. Scripta 35(2): 149–186. doi:10.1111/j.1463-6409.2006.00221.x (HTML abstract)
- 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)
- Pasquet, Eric; Bourdon, Estelle; Kalyakin, Mikhail V. & Cibois, Alice (2006). The fulvettas (Alcippe), Timaliidae, Aves): a polyphyletic group. Zoologica Scripta 35, 559–566. doi:10.1111/j.1463-6409.2006.00253.x (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
- Sibley, Charles Gald & Ahlquist, Jon Edward (1990): Phylogeny and classification of birds. Yale University Press, New Haven, Conn.
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