Bulbul

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iBulbuls

Light-vented Bulbul, Pycnonotus sinensis

Scientific classification

Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Order:	Passeriformes
Family:	Pycnonotidae

Genera

See text.

Bulbuls (Pycnonotidae) are a family of medium-sized passerine songbirds resident in Africa and tropical Asia. There are about 130 species.

These are mostly frugivorous birds. Some are colorful with yellow, red or orange vents, cheeks, throat or supercilia, but most are drab, with uniform olive brown to black plumage. Some have very distinct crests.

These are noisy and gregarious birds with often beautiful striking songs.

Many of these species inhabit tree tops, while some are restricted to the undergrowth. Up to five purple-pink eggs are laid in an open tree nests and incubated by the female.

The Red-whiskered Bulbul, Pycnonotus jocosus, has been widely introduced to tropical and subtropical areas, for example southern Florida, USA.

[edit] Systematics

The traditional layout was to divide the bulbuls into 4 groups, named Pycnonotus, Phyllastrephus, Criniger, and Chlorocichla groups after characteristic genera (Delacour, 1943). However, more recent analyses demonstrated that this arrangement was probably based on erroneous interpretation of characters:

Studies of the mitochondrial cytochrome b sequence found that five species of Phyllastrephus did not belong to the bulbuls, but to an enigmatic group of songbirds from Madagascar instead (Cibois et al., 2001; see below for the species in question). Similarly, analysis of DNA sequences of the RAG1 and RAG2 genes suggests that the genus Nicator was not a bulbul either (Beresford et al., 2005). That the previous arrangement had failed to take into account biogeography was indicated by the study of Pasquet et al. (2001) who demonstrated the genus Criniger must be divided into an African and an Asian (Alophoixus) lineage. Using analysis of 2 mitochondrial and one nuclear DNA sequences, Moyle & Marks (2006) found one largely Asian lineage and one African group of genera; the Golden Greenbul seemed to be very distinct and form a group of its own. Some taxa are not monophyletic, and more research is necessary to determine relationships within the larger genera.

Family Pycnonotidae

Genus Pycnonotus (paraphyletic)
- "Ancient" Asian bulbuls
- Black-headed Bulbul, Pycnonotus atriceps
- Puff-backed Bulbul, Pycnonotus eutilotus
- Black-and-white Bulbul, Pycnonotus melanoleucus
- Pycnonotus proper
- Black-crested Bulbul, Pycnonotus melanicterus
- Grey-bellied Bulbul, Pycnonotus cyaniventris
- Spectacled Bulbul, Pycnonotus erythropthalmos
- Straw-headed Bulbul, Pycnonotus zeylanicus
- Red-eyed Bulbul, Pycnonotus brunneus
- Olive-winged Bulbul, Pycnonotus plumosus
- Yellow-vented Bulbul, Pycnonotus goiavier
- Common Bulbul, Pycnonotus barbatus
- Black-fronted Bulbul, Pycnonotus nigricans
- White-cheeked Bulbul, Pycnonotus leucogenys
- Unassigned
- Striated Bulbul, Pycnonotus striatus
- Cream-striped Bulbul, Pycnonotus leucogrammicus
- Spot-necked Bulbul, Pycnonotus tympanistrigus
- Grey-headed Bulbul, Pycnonotus priocephalus
- Styan's Bulbul, Pycnonotus taivanus
- Scaly-breasted Bulbul, Pycnonotus squamatus
- Red-whiskered Bulbul, Pycnonotus jocosus
- Brown-breasted Bulbul, Pycnonotus xanthorrhous
- Light-vented Bulbul, Pycnonotus sinensis
- Cape Bulbul, Pycnonotus capensis
- White-spectacled Bulbul, Pycnonotus xanthopygos
- White-eared Bulbul, Pycnonotus leucotis
- Red-vented Bulbul, Pycnonotus cafer
- Sooty-headed Bulbul, Pycnonotus aurigaster
- Blue-wattled Bulbul, Pycnonotus nieuwenhuisii (disputed)
- Yellow-wattled Bulbul, Pycnonotus urostictus
- Orange-spotted Bulbul, Pycnonotus bimaculatus
- Stripe-throated Bulbul, Pycnonotus finlaysoni
- Yellow-throated Bulbul, Pycnonotus xantholaemus
- Yellow-eared Bulbul, Pycnonotus penicillatus
- Flavescent Bulbul, Pycnonotus flavescens
- White-browed Bulbul, Pycnonotus luteolus
- Streak-eared Bulbul, Pycnonotus blanfordi
- Cream-vented Bulbul, Pycnonotus simplex

Genus Spizixos
- Crested Finchbill, Spizixos canifrons
- Collared Finchbill, Spizixos semitorques

Genus Tricholestes
- Hairy-backed Bulbul, Tricholestes criniger

Genus Setornis
- Hook-billed Bulbul, Setornis criniger

Genus Alophoixus (possibly polyphyletic)
- Finsch's Bulbul, Alophoixus finschii
- White-throated Bulbul, Alophoixus flaveolus
- Puff-throated Bulbul, Alophoixus pallidus
- Ochraceous Bulbul, Alophoixus ochraceus
- Gray-cheeked Bulbul, Alophoixus bres
- Yellow-bellied Bulbul, Alophoixus phaeocephalus
- Golden Bulbul, Alophoixus affinis

Genus Iole
- Olive Bulbul, Iole virescens
- Grey-eyed Bulbul, Iole propinqua
- Buff-vented Bulbul, Iole olivacea
- Yellow-browed Bulbul, Iole indica

Genus Hemixos
- Ashy Bulbul, Hemixos flavala
- Chestnut Bulbul, Hemixos castanonotus

Genus Ixos (paraphyletic)
- Close to Hemixos
- Streaked Bulbul, Ixos malaccensis
- Close to Hypsipetes
- Philippine Bulbul, Ixos philippinus
- Unassigned
- Sulphur-bellied Bulbul, Ixos palawanensis
- Streak-breasted Bulbul, Ixos siquijorensis
- Yellowish Bulbul, Ixos everetti
- Zamboanga Bulbul, Ixos rufigularis
- Mountain Bulbul, Ixos mcclellandii
- Sunda Bulbul, Ixos virescens

Genus Microscelis
- Brown-eared Bulbul, Microscelis amaurotis (sometimes included in Ixos)

Genus Hypsipetes
- Madagascar Bulbul, Hypsipetes madagascariensis
- Black Bulbul, Hypsipetes leucocephalus
- Seychelles Bulbul, Hypsipetes crassirostris
- Comoro Bulbul, Hypsipetes parvirostris
- Reunion Bulbul, Hypsipetes borbonicus
- Mauritius Bulbul, Hypsipetes olivaceus
- Nicobar Bulbul, Hypsipetes virescens
- White-headed Bulbul, Hypsipetes thompsoni

Genus Calyptocichla
- Golden Greenbul, Calyptocichla serina

Genus Phyllastrephus
- Leaf-love Greenbul, Phyllastrephus scandens
- Cabanis' Greenbul, Phyllastrephus cabanisi
- Fischer's Greenbul, Phyllastrephus fischeri
- Placid Greenbul, Phyllastrephus placidus
- Terrestrial Brownbul, Phyllastrephus terrestris
- Northern Brownbul, Phyllastrephus strepitans
- Pale-olive Greenbul, Phyllastrephus fulviventris
- Gray-olive Greenbul, Phyllastrephus cerviniventris
- Baumann's Greenbul, Phyllastrephus baumanni
- Toro Olive Greenbul, Phyllastrephus hypochloris
- Cameroon Olive Greenbul, Phyllastrephus poensis
- Sassi's Greenbul, Phyllastrephus lorenzi
- Yellow-streaked Bulbul, Phyllastrephus flavostriatus
- Grey-headed Greenbul, Phyllastrephus poliocephalus
- Tiny Greenbul, Phyllastrephus debilis
- White-throated Greenbul, Phyllastrephus albigularis
- Icterine Greenbul, Phyllastrephus icterinus
- Liberian Greenbul, Phyllastrephus leucolepis
- Xavier's Greenbul, Phyllastrephus xavieri

Genus Andropadus (possibly polyphyletic)
- Cameroon Mountain Greenbul, Andropadus montanus
- Shelley's Greenbul, Andropadus masukuensis
- Little Greenbul, Andropadus virens
- Grey Greenbul, Andropadus gracilis
- Ansorge's Greenbul, Andropadus ansorgei
- Plain Greenbul, Andropadus curvirostris
- Slender-billed Greenbul, Andropadus gracilirostris
- Sombre Greenbul, Andropadus importunus
- Yellow-whiskered Bulbul, Andropadus latirostris
- Western Mountain Greenbul, Andropadus tephrolaemus
- Eastern Mountain Greenbul, Andropadus nigriceps
- Stripe-cheeked Bulbul, Andropadus milanjensis

Genus Criniger
- Red-tailed Greenbul, Criniger calurus
- Western Bearded Greenbul, Criniger barbatus
- Eastern Bearded Greenbul, Criniger chloronotus
- Yellow-bearded Greenbul, Criniger olivaceus
- White-bearded Greenbul, Criniger ndussumensis

Genus Bleda
- Common Bristlebill, Bleda syndactyla
- Green-tailed Bristlebill, Bleda eximia
- Grey-headed Bristlebill, Bleda canicapilla

Genus Thescelocichla
- Swamp Greenbul, Thescelocichla leucopleura

Genus Chlorocichla
- Simple Greenbul, Chlorocichla simplex
- Yellow-throated Greenbul, Chlorocichla flavicollis
- Yellow-necked Greenbul, Chlorocichla falkensteini
- Yellow-bellied Greenbul, Chlorocichla flaviventris
- Joyful Greenbul, Chlorocichla laetissima
- Prigogine's Greenbul, Chlorocichla prigoginei

Genus Ixonotus (pending confirmation of placement)
- Spotted Greenbul, Ixonotus guttatus

Genus Baeopogon
- Honeyguide Greenbul, Baeopogon indicator
- Sjostedt's Greenbul, Baeopogon clamans

Genus Neolestes
- Black-collared Bulbul, Neolestes torquatus

The last genus might be allied to Calyptocichla or not be a bulbul at all.

Taxa until recently included in the Pycnonotidae are:

Genus Bernieria
- Long-billed Greenbul, Bernieria madagascariensis

Genus Xanthomixis (possibly polyphyletic)
- Spectacled Greenbul, Xanthomixis zosterops
- Appert's Greenbul, Xanthomixis apperti
- Dusky Greenbul, Xanthomixis tenebrosus
- Gray-crowned Greenbul, Xanthomixis cinereiceps

Genus Nicator
- Yellow-spotted Nicator, Nicator chloris
- Eastern Nicator, Nicator gularis
- Yellow-throated Nicator, Nicator vireo

The first two belong to the "Malagasy warblers"; the affiliations of Nicator are unknown at present.

[edit] References

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; Slikas, Beth; Shulenberg, 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

Delacour, J. (1943): A revision of the genera and species of the family Pycnonotidae (bulbuls). Zoologica 28(1): 17-28.

Moyle, Robert G. & Marks, Ben D. (2006): Phylogenetic relationships of the bulbuls (Aves: Pycnonotidae) based on mitochondrial and nuclear DNA sequence data. Molecular Phylogenetics and Evolution 40(3): Pages 687-695. DOI:10.1016/j.ympev.2006.04.015 (HTML abstract)