Timaliidae

Life-forms

Superregnum: Eukaryota
Cladus: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Aves
Subclassis: Carinatae
Infraclassis: Neornithes
Parvclassis: Neognathae
Ordo: Passeriformes
Subordo: Passeri
Parvordo: Passerida
Superfamilia: Sylvioidea
Familia: Timaliidae
Subfamiliae: Leiothrichinae - Pellorneinae - Timaliinae - Zosteropinae

Overview of genera

Actinodura - Alcippe - Arcanator - Babax - Chamaea - Chrysomma - Conostoma - Crocias - Crossleyia - Cutia - Dumetia - Gampsorhynchus - Garrulax - Hartertula - Heterophasia - Illadopsis - Jabouilleia - Kakamega - Kenopia - Kupeornis - Leiothrix - Liocichla - Lioptilus - Macronous - Malacocincla - Malacopteron - Micromacronus - Minla - Modulatrix - Mystacornis - Myzornis - Napothera - Neomixis - Oxylabes - Panurus - Paradoxornis - Parophasma - Pellorneum - Phyllanthus - Pomatorhinus - Pteruthius - Ptilocichla - Ptyrticus - Rhopocichla - Rimator - Spelaeornis - Sphenocichla - Stachyris - Timalia - Trichastoma - Turdoides - Xiphirhynchus - Yuhina

Name

Timaliidae

References

* Gelang, M.; Cibois, A.; Pasquet, E.; Olsson, U.; Alström, P.; Ericson, P.G.P. 2009: Phylogeny of babblers (Aves, Passeriformes): major lineages, family limits and classification. Zoologica scripta, 38: 225-236.

Vernacular names
English: Old World babblers
Türkçe: Timalyagiller

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).

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.

Characteristics

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.[1]

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.[1]

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.[1]

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)[cite this quote]

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 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.[2]

* 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.

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).

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).

Liocichlas and allies clade

* Genus Liocichla – liocichlas (4 species)

* Genus Actinodura – barwings (7 species)

* Genus Minla
o Red-tailed Minla, Minla ignotincta

* Genus Chrysominla
o Chestnut-tailed Minla, Chrysominla strigula

* Genus Siva
o Blue-winged Minla or Siva, Siva cyanouroptera

* Genus Leiothrix (2 species)

* Genus Heterophasia – sibias (8 species)

* Genus Alcippe – "fulvettas" or typical alcippes (tentatively placed here)
o Grey-cheeked Fulvetta, Alcippe morrisonia (polyphyletic?)
o Mountain Fulvetta, Alcippe peracensis (paraphyletic?)
o Brown Fulvetta, Alcippe brunneicauda
o Black-browed Fulvetta, Alcippe grotei – formerly in peracensis
o Brown-cheeked Fulvetta, Alcippe poioicephala
o Javan Fulvetta, Alcippe pyrrhoptera – tentatively placed here
o Nepal Fulvetta, Alcippe nipalensis – tentatively placed here

* Genus Cutia - cuitas (2 species)

* Genus Turdoides (27 species)

* Genus Garrulax – laughingthrushes (formerly c.50 species). Polyphyletic, consisting of around 11 genera for which see Garrulax.

* Genus Babax – babaxes (3 species)

* Genus Pellorneum (7 species) - possibly polyphyletic (Jønsson & Fjeldså 2006)

* Genus Kenopia
o Striped Wren-babbler, Kenopia striata

* Genus Malacopteron (6 species)

* Genus Schoeniparus – "fulvettas" or atypical alcippes, formerly in Alcippe
o Rufous-throated Fulvetta, Schoeniparus rufogularis
o Dusky Fulvetta, Schoeniparus brunnea
o Rusty-capped Fulvetta, Schoeniparus dubia

* Genus Pseudominla – 4 species of atypical alcippes or "fulvettas", formerly in Alcippe

* Genus Gampsorhynchus
o White-hooded Babbler, Gampsorhynchus rufulus

* Genus Malacocincla (5 extant species)
o Vanderbilt's Babbler, Malacocincla sepiarium vanderbilti – extinct? (late 20th century?)

* Genus Napothera – atypical wren-babblers (paraphyletic). Sometimes split into several genera.
o Large Wren-babbler, Napothera (Turdinus) macrodactyla
o Rusty-breasted Wren-babbler, Napothera (Turdinus) rufipectus
o Black-throated Wren-babbler, Napothera (Turdinus) atrigularis
o Marbled Wren-babbler, Napothera (Turdinus) marmorata
o Limestone Wren-babbler, Napothera (Gypsophila) crispifrons
o Streaked Wren-babbler, Napothera brevicaudata
o Mountain Wren-babbler, Napothera crassa
o Eyebrowed Wren-babbler, Napothera epilepidota
o Luzon Wren-babbler, Napothera (Robsonius) rabori

* Genus Jabouilleia – atypical scimitar-babblers (2 species)

* Genus Graminicola – formerly in Sylviidae, tentatively placed here
o Rufous-rumped Grassbird (or "-babbler"), Graminicola bengalensis

Timaliine clade (putative subfamily Timaliinae)

* Genus Pomatorhinus – scimitar-babblers (9 species)

* Genus Xiphirhynchus – Slender-billed Scimitar-babbler

* Genus Stachyris (polyphyletic)
* Stachyris group
o Grey-throated Babbler, Stachyris nigriceps

* Unresolved
o Buff-chested Babbler, Stachyris ambigua
o White-breasted Babbler, Stachyris grammiceps
o Sooty Babbler, Stachyris herberti
o Nonggang Babbler, Stachyris nonggangensis
o Grey-headed Babbler, Stachyris poliocephala
o Snowy-throated Babbler, Stachyris oglei
o Spot-necked Babbler, Stachyris striolata
o White-necked Babbler, Stachyris leucotis
o Black-throated Babbler, Stachyris nigricollis
o White-bibbed Babbler, Stachyris thoracica
o Chestnut-rumped Babbler, Stachyris maculata
o Chestnut-winged Babbler, Stachyris erythroptera
o Crescent-chested Babbler, Stachyris melanothorax

* Genus Stachyridopsis
o Rufous-fronted Babbler, Stachyridopsis rufifrons
+ Deignan's Babbler, Stachyridopsis (rufifrons) rodolphei
o Rufous-capped Babbler, Stachyridopsis ruficeps
o Golden Babbler, Stachyridopsis chrysaea
o Black-chinned Babbler, Stachyridopsis pyrrhops

* Genus Spelaeornis (9 species) – typical wren-babblers

* Genus Sphenocichla
o Wedge-billed Wren-babbler, Sphenocichla humei

* Genus Macronus – tit-babblers (5 species)

* Genus Timalia
o Chestnut-capped Babbler, Timalia pileata

White-eye clade

If the white-eyes are maintained as a separate family Zosteropidae, this group would have to be included there:
The White-collared Yuhina (Yuhina diademata) would be moved to the Zosteropidae if these are retained

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

* Genus Staphida
o Striated Yuhina, Staphida castaniceps
o Chestnut-collared Yuhina, Staphida torqueola
o Chestnut-crested Yuhina, Staphida everetti

* Genus Dasycrotapha – formerly in Stachyris; tentatively placed here
o Flame-templed Babbler, Dasycrotapha speciosa

* Genus Sterrhoptilus – formerly in Stachyris; tentatively placed here
o Mindanao Pygmy Babbler (or Yuhina), Sterrhoptilus plateni
o Visayan Pygmy Babbler (or Yuhina), Sterrhoptilus pygmaeus
o Golden-crowned Babbler (or Yuhina), Sterrhoptilus dennistouni
o Black-crowned Babbler (or Yuhina), Sterrhoptilus nigrocapitata
o Rusty-crowned Babbler (or Yuhina), Sterrhoptilus capitalis

* Genus Zosterornis – formerly in Stachyris; tentatively placed here
o Chestnut-faced Babbler (or Yuhina), Zosterornis whiteheadi
o Luzon Striped Babbler (or Yuhina), Zosterornis striatus
o Panay Striped Babbler (or Yuhina), Zosterornis latistriatus
o Negros Striped Babbler (or Yuhina), Zosterornis nigrorum
o Palawan Striped Babbler (or Yuhina), Zosterornis hypogrammicus

Sylviid clade
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:

* Genus Pseudoalcippe
o African Hillbabbler, Pseudoalcippe abyssinica – formerly in Illadopsis

* Genus Lioparus – formerly in Alcippe
o Golden-breasted Fulvetta, Lioparus chrysotis

* Genus Fulvetta – 7 species of typical fulvettas, formerly in Alcippe
o Spectacled Fulvetta, Fulvetta ruficapilla
o Chinese Fulvetta, Fulvetta striaticollis
o White-browed Fulvetta, Fulvetta vinipectus
o Grey-hooded Fulvetta, Fulvetta cinereiceps
o Taiwan Fulvetta, Fulvetta formosana - formerly in F. cinereiceps
o Streak-throated Fulvetta, Fulvetta manipurensis - formerly in F. cinereiceps
o 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 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?
* 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

The Wrentit's relationships have long been elusive, but it is probably a relative of the typical warblers.

* Genus Malia – Malia
* Genus Lioptilus - Bush Blackcap (may be related to Sylvia)
* Genus Parophasma – Abyssinian Catbird (may be related to Sylvia)
* Genus Kupeornis – 3 species of mountain-babblers. Perhaps related to Phyllanthus
* Genus Phyllanthus - Capuchin Babbler
* 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:

Other Sylvioidea

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

* Genus Neomixis – jeries
o Common Jery, Neomixis tenella
o Green Jery, Neomixis viridis
o Stripe-throated Jery, Neomixis striatigula

Several lineages of Malagasy warblers, Bernieridae

* Genus Hartertula – formerly in Neomixis
o Wedge-tailed Jery, Hartertula flavoviridis
* Genus Crossleyia
o Yellow-browed Oxylabes, Crossleyia xanthophrys
* Genus Oxylabes
o White-throated Oxylabes, Oxylabes madagascariensis

Other Passerida

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

* Genus Arcanator – sometimes included in Modulatrix
o Dapple-throat, Arcanator orostruthus
* Genus Modulatrix
o Spot-throat, Modulatrix stictigula

"Corvida"

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

* Genus Erpornis
o 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
o Black-headed Shrike-babbler, Pteruthius rufiventer
o White-browed Shrike-babbler, Pteruthius flaviscapis
o Green Shrike-babbler, Pteruthius xanthochlorus
o Black-eared Shrike-babbler, Pteruthius melanotis
o Chestnut-fronted Shrike-babbler, Pteruthius aenobarbus

Belongs to the vangas, Vangidae (Schulenberg 2003):

* Genus Mystacornis
o Crossley's Babbler-vanga, Mystacornis crossleyi

Incertae sedis

* Genus Kakamega (Cibois 2003a) – uncertain relationships. May be related to thrushes or sugarbirds
o Grey-chested Kakamega or Grey-chested "Illadopsis", Kakamega poliothorax

References

1. ^ a b c Perrins, C. (1991). Forshaw, Joseph. ed. Encyclopaedia of Animals: Birds. London: Merehurst Press. pp. 188–190. ISBN 1-85391-186-0.
2. ^ Sylvioidea III (http://jboyd.net/Taxo/List23.html#timaliidae)

* 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

* 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 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.