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| subdivision_ranks = Genera | | subdivision_ranks = Genera | ||
| subdivision = See text | | subdivision = See text | ||
| synonyms = * Brachypodidae <small>[[William | | synonyms = * Brachypodidae <small>[[William Swainson|Swainson]], 1831</small> | ||
* Trichophoridae <small>Swainson, 1831</small> | * Trichophoridae <small>Swainson, 1831</small> | ||
* Ixosidae <small>[[Charles Lucien Bonaparte|Bonaparte]], 1838</small> | * Ixosidae <small>[[Charles Lucien Bonaparte|Bonaparte]], 1838</small> | ||
| Line 20: | Line 20: | ||
==Taxonomy== | ==Taxonomy== | ||
The family Pycnonotidae was introduced by the English zoologist [[George Robert Gray]] in 1840 as a subfamily Pycnonotinae of the thrush family [[Turdidae]].<ref>{{ cite book | last=Gray | first=George Robert | author-link=George Robert Gray | year=1840 | title=A List of the Genera of Birds : with an Indication of the Typical Species of Each Genus | location=London | publisher=R. and J.E. Taylor | page=28 | url=https://www.biodiversitylibrary.org/page/13668922 }}</ref><ref>{{cite book | last=Bock | first=Walter J. | year=1994 | title=History and Nomenclature of Avian Family-Group Names | series=Bulletin of the American Museum of Natural History | volume= | The family Pycnonotidae was introduced by the English zoologist [[George Robert Gray]] in 1840 as a subfamily Pycnonotinae of the thrush family [[Turdidae]].<ref>{{ cite book | last=Gray | first=George Robert | author-link=George Robert Gray | year=1840 | title=A List of the Genera of Birds : with an Indication of the Typical Species of Each Genus | location=London | publisher=R. and J.E. Taylor | page=28 | url=https://www.biodiversitylibrary.org/page/13668922 }}</ref><ref>{{cite book | last=Bock | first=Walter J. | year=1994 | title=History and Nomenclature of Avian Family-Group Names | series=Bulletin of the American Museum of Natural History | volume= 222 | publisher=American Museum of Natural History | location=New York | pages=201, 237 | hdl=2246/830 | url=http://digitallibrary.amnh.org/handle/2246/830 }}<!--Linked page allows download of the 48MB pdf--></ref> | ||
The | The Persian word ''bulbul'' (بلبل) is sometimes used to refer to the "[[nightingale]]" as well as the bulbul, but the English word ''bulbul'' refers to the birds discussed in this article.{{sfn|Fishpool|Tobias|2005|pp=[https://archive.org/details/handbookofbirdso0010unse/page/161/mode/1up 161–162]}} | ||
A few species that were previously considered to be members of the Pycnonotidae have been moved to other families. Several Malagasy species that were formerly placed in the genus ''[[Phyllastrephus]]'' are now placed in the family [[Bernieridae]].<ref>{{ cite journal | last1=Cibois | first1=Alice | last2=Slikas | first2=Beth | last3=Shulenberg | first3=Thomas S. | last4=Pasquet | first4=Eric | year=2001 | title=An endemic radiation of Malagasy songbirds is revealed by mitochondrial DNA sequence data | journal=Evolution | volume=55 | issue=6 | pages=1198–1206 | doi=10.1554/0014-3820(2001)055[1198:AEROMS]2.0.CO;2 | pmid=11475055 | doi-access= }}</ref><ref>{{cite journal | last1 = Cibois | first1 = Alice | last2 = David | first2 = Normand | last3 = Gregory | first3 = Steven M. S. | last4 = Pasquet | first4 = Eric | year = 2010 | title = Bernieridae (Aves: Passeriformes): a family-group name for the Malagasy sylvioid radiation | journal = Zootaxa | volume = 2554 | pages = 65–68 | doi = 10.11646/zootaxa.2554.1.6 }}</ref> In addition, the genus ''[[Nicator]]'' containing three African species is now placed in a separate family [[Nicatoridae]].<ref>{{cite journal |last1=Beresford |first1=P. |last2=Barker |first2=F.K. |last3=Ryan |first3=P.G. |last4=Crowe |first4=T.M. |year=2005 |title=African endemics span the tree of songbirds (Passeri): molecular systematics of several evolutionary 'enigmas' |journal=Proceedings of the Royal Society B |volume=272 |issue=1565 |pages=849–858 |doi=10.1098/rspb.2004.2997 |pmid=15888418 |pmc=1599865 }}</ref><ref>{{Cite journal | last1=Johansson | first1=U.S. | last2=Fjeldså | first2=J. | last3=Bowie | first3=R.C.K. | date=2008 | title=Phylogenetic relationships within Passerida (Aves: Passeriformes): A review and a new molecular phylogeny based on three nuclear intron markers | journal=Molecular Phylogenetics and Evolution | volume=48 | issue=3 | pages=858–876 | doi=10.1016/j.ympev.2008.05.029| pmid=18619860 }}</ref> | A few species that were previously considered to be members of the Pycnonotidae have been moved to other families. Several Malagasy species that were formerly placed in the genus ''[[Phyllastrephus]]'' are now placed in the family [[Bernieridae]].<ref>{{ cite journal | last1=Cibois | first1=Alice | last2=Slikas | first2=Beth | last3=Shulenberg | first3=Thomas S. | last4=Pasquet | first4=Eric | year=2001 | title=An endemic radiation of Malagasy songbirds is revealed by mitochondrial DNA sequence data | journal=Evolution | volume=55 | issue=6 | pages=1198–1206 | doi=10.1554/0014-3820(2001)055[1198:AEROMS]2.0.CO;2 | pmid=11475055 | doi-access= }}</ref><ref>{{cite journal | last1 = Cibois | first1 = Alice | last2 = David | first2 = Normand | last3 = Gregory | first3 = Steven M. S. | last4 = Pasquet | first4 = Eric | year = 2010 | title = Bernieridae (Aves: Passeriformes): a family-group name for the Malagasy sylvioid radiation | journal = Zootaxa | volume = 2554 | pages = 65–68 | doi = 10.11646/zootaxa.2554.1.6 }}</ref> In addition, the genus ''[[Nicator]]'' containing three African species is now placed in a separate family [[Nicatoridae]].<ref>{{cite journal |last1=Beresford |first1=P. |last2=Barker |first2=F.K. |last3=Ryan |first3=P.G. |last4=Crowe |first4=T.M. |year=2005 |title=African endemics span the tree of songbirds (Passeri): molecular systematics of several evolutionary 'enigmas' |journal=Proceedings of the Royal Society B |volume=272 |issue=1565 |pages=849–858 |doi=10.1098/rspb.2004.2997 |pmid=15888418 |pmc=1599865 }}</ref><ref>{{Cite journal | last1=Johansson | first1=U.S. | last2=Fjeldså | first2=J. | last3=Bowie | first3=R.C.K. | date=2008 | title=Phylogenetic relationships within Passerida (Aves: Passeriformes): A review and a new molecular phylogeny based on three nuclear intron markers | journal=Molecular Phylogenetics and Evolution | volume=48 | issue=3 | pages=858–876 | doi=10.1016/j.ympev.2008.05.029| pmid=18619860 }}</ref> | ||
A study published in 2007 by Ulf Johansson and colleagues using three nuclear markers found that the genus ''[[Andropadus]]'' was [[monophyletic|non-monophyletic]]. In the subsequent revision, species were moved to three resurrected genera: ''[[Arizelocichla]]'', ''[[Stelgidillas]]'' and ''[[Eurillas]]''. Only the [[sombre greenbul]] (''Andropadus importunus''), was retained in ''Andropadus''.<ref name= | A study published in 2007 by Ulf Johansson and colleagues using three nuclear markers found that the genus ''[[Andropadus]]'' was [[monophyletic|non-monophyletic]]. In the subsequent revision, species were moved to three resurrected genera: ''[[Arizelocichla]]'', ''[[Stelgidillas]]'' and ''[[Eurillas]]''. Only the [[sombre greenbul]] (''Andropadus importunus''), was retained in ''Andropadus''.<ref name=avilist/><ref>{{Cite journal | last1=Johansson | first1=U.S. | last2=Fjeldså | first2=J. | last3=Lokugalappatti | first3=L.G.S. | last4=Bowie | first4=R.C.K. | date=2007 | title=A nuclear DNA phylogeny and proposed taxonomic revision of African greenbuls (Aves, Passeriformes, Pycnonotidae) | journal=Zoologica Scripta | volume=36 | issue=5 | pages=417–427 | doi=10.1111/j.1463-6409.2007.00290.x| s2cid=84799480 }}</ref> A study by Subir Shakya and Frederick Shelden published in 2017 found that species in the large genus ''[[ Pycnonotus]]'' formed several deeply divergent clades. The genus was split and genera were resurrected to accommodate these clades.<ref name=avilist/><ref name=shakya>{{Cite journal | last1=Shakya | first1=Subir B. | last2=Sheldon | first2=Frederick H. | date=2017 | title=The phylogeny of the world's bulbuls (Pycnonotidae) inferred using a supermatrix approach | journal=Ibis | volume=159 | issue=3 | pages=498–509 | doi=10.1111/ibi.12464| doi-access=free }}</ref> | ||
The family forms two main clades. One clade contains species that are only found in Africa; many of these have greenbul in the common name. The second clade contains mostly Asian species but includes a few species that are found in Africa.<ref name=shakya/> | The family forms two main clades. One clade contains species that are only found in Africa; many of these have greenbul in the common name. The second clade contains mostly Asian species but includes a few species that are found in Africa.<ref name=shakya/> | ||
{{Cladogram | {{Cladogram | ||
|caption=Phylogeny based on a study of the babblers by Cai and colleagues published in 2019.<ref name= | |caption=Phylogeny based on a study of the babblers by Cai and colleagues published in 2019.<ref name=avilist/><ref>{{Cite journal | last1=Cai | first1=T. | last2=Cibois | first2=A. | last3=Alström | first3=P. | last4=Moyle | first4=R.G. | last5=Kennedy | first5=J.D. | last6=Shao | first6=S. | last7=Zhang | first7=R. | last8=Irestedt | first8=M. | last9=Ericson | first9=P.G.P. | last10=Gelang | first10=M. | last11=Qu | first11=Y. | last12=Lei | first12=F. | last13=Fjeldså | first13=J. | date=2019 | title=Near-complete phylogeny and taxonomic revision of the world's babblers (Aves: Passeriformes) | journal=Molecular Phylogenetics and Evolution | volume=130 | pages=346–356 | doi=10.1016/j.ympev.2018.10.010 | pmid=30321696 | doi-access=free }}</ref> | ||
|align=centre | |align=centre | ||
|cladogram={{ | |cladogram={{Clade| style=font-size:100%;line-height:100%;width:500px; | ||
|1={{clade | |1={{clade | ||
|1={{clade | |1={{clade | ||
|1='''Pycnonotidae''' – bulbuls ( | |1='''Pycnonotidae''' – bulbuls (163 species) | ||
|2={{clade | |2={{clade | ||
|1={{clade | |1={{clade | ||
|1=[[ | |1=[[Paradoxornithidae]] – parrotbills and myzornis (38 species) | ||
|2=[[ | |2=[[Sylviidae]] – sylviid warblers (32 species) | ||
}} | }} | ||
|2={{clade | |2={{clade | ||
|1=[[Zosteropidae]] – white-eyes ( | |1=[[Zosteropidae]] – white-eyes (152 species) | ||
|2={{clade | |2={{clade | ||
|1=[[Timaliidae]] – tree babblers (58 species) | |1=[[Timaliidae]] – tree babblers (58 species) | ||
|2={{clade | |2={{clade | ||
|1=[[Pellorneidae]] – ground babblers ( | |1=[[Pellorneidae]] – ground babblers (68 species) | ||
|2={{clade | |2={{clade | ||
|1=[[Alcippeidae]] – Alcippe fulvettas (10 species) | |1=[[Alcippeidae]] – Alcippe fulvettas (10 species) | ||
| Line 65: | Line 65: | ||
[[File:Red-whiskered Bulbul-web.jpg|thumb|252px|[[Red-whiskered bulbul]]]] | [[File:Red-whiskered Bulbul-web.jpg|thumb|252px|[[Red-whiskered bulbul]]]] | ||
Currently, there are | Currently, there are 161 recognized species in 28 genera:<ref name=avilist>{{ cite web | author=AviList Core Team | date=2025 | title=AviList: The Global Avian Checklist, v2025 | doi=10.2173/avilist.v2025 | doi-access=free | url=http://www.avilist.org/checklist/v2025/ | access-date=17 November 2025 }}</ref> | ||
* Genus ''[[Andropadus]]'' – [[sombre greenbul]] (formerly contained many species) | * Genus ''[[Andropadus]]'' – [[sombre greenbul]] (formerly contained many species) | ||
| Line 72: | Line 72: | ||
* Genus ''[[Neolestes]]'' – [[black-collared bulbul]] | * Genus ''[[Neolestes]]'' – [[black-collared bulbul]] | ||
* Genus ''[[Bleda (bird)|Bleda]]'' – bristlebills (5 species) | * Genus ''[[Bleda (bird)|Bleda]]'' – bristlebills (5 species) | ||
* Genus ''[[Atimastillas]]'' – | * Genus ''[[Atimastillas]]'' – greenbuls (2 species) | ||
* Genus ''[[Ixonotus]]'' – [[spotted greenbul]] | * Genus ''[[Ixonotus]]'' – [[spotted greenbul]] | ||
* Genus ''[[Thescelocichla]]'' – [[swamp palm bulbul]] | * Genus ''[[Thescelocichla]]'' – [[swamp palm bulbul]] | ||
* Genus ''[[Chlorocichla]]'' – greenbuls (5 species) | * Genus ''[[Chlorocichla]]'' – greenbuls (5 species) | ||
* Genus ''[[Baeopogon]]'' – greenbuls (2 species) | * Genus ''[[Baeopogon]]'' – greenbuls (2 species) | ||
* Genus ''[[Arizelocichla]]'' – greenbuls ( | * Genus ''[[Arizelocichla]]'' – greenbuls (11 species) (formerly in ''Andropadus'') | ||
* Genus ''[[Criniger]]'' – greenbuls (5 species) | * Genus ''[[Criniger]]'' – greenbuls (5 species) | ||
* Genus ''[[Eurillas]]'' – greenbuls (5 species) (formerly in ''Andropadus'') | * Genus ''[[Eurillas]]'' – greenbuls (5 species) (formerly in ''Andropadus'') | ||
| Line 86: | Line 86: | ||
* Genus ''[[Alcurus]]'' – [[striated bulbul]] | * Genus ''[[Alcurus]]'' – [[striated bulbul]] | ||
* Genus ''[[Iole (genus)|Iole]]'' – 7 species | * Genus ''[[Iole (genus)|Iole]]'' – 7 species | ||
* Genus ''[[Hemixos]]'' – | * Genus ''[[Hemixos]]'' – 3 species | ||
* Genus ''[[Acritillas]]'' – [[yellow-browed bulbul]] | * Genus ''[[Acritillas]]'' – [[yellow-browed bulbul]] | ||
* Genus ''[[Ixos]]'' – 5 species | * Genus ''[[Ixos]]'' – 5 species | ||
* Genus ''[[Hypsipetes]]'' – | * Genus ''[[Hypsipetes]]'' – 26 species (includes 3 species formerly in ''Thapsinillas'', one formerly in ''[[Cerasophila]]'' and one formerly in ''Microscelis'') | ||
* Genus ''[[Microtarsus]]'' – 7 species | |||
* Genus ''[[Microtarsus]]'' – | * Genus ''[[Rubigula]]'' – 8 species (formerly in ''Pycnonotus'') | ||
* Genus ''[[Rubigula]]'' – | |||
* Genus ''[[Bare-faced bulbul|Nok]]'' – [[bare-faced bulbul]] (genus introduced in 2017) | * Genus ''[[Bare-faced bulbul|Nok]]'' – [[bare-faced bulbul]] (genus introduced in 2017) | ||
* Genus ''[[Spizixos]]'' – finchbills (2 species) | * Genus ''[[Spizixos]]'' – finchbills (2 species) | ||
* Genus ''[[Pycnonotus]]'' – | * Genus ''[[Pycnonotus]]'' – 31 species (substantially reduced from earlier classification) | ||
===Cladogram=== | ===Cladogram=== | ||
Phylogeny based on a study by Subir Shakya and Frederick Shelden published in 2017 with the revised genera as defined in the [[AviList]] checklist.<ref name=avilist/><ref name=shakya/> The positions of the [[bare-faced bulbul]] (''Nok hualon'') and the [[yellow-browed bulbul]] (''Acritillas indica'') are based on a study by Jérôme Fuchs and colleagues published in 2018.<ref>{{Cite journal | last1=Fuchs | first1=J. | last2=Pasquet | first2=E. | last3=Stuart | first3=B.L. | last4=Woxvold | first4=I.A. | last5=Duckworth | first5=J.W. | last6=Bowie | first6=R.C.K. | date=2018 | title=Phylogenetic affinities of the enigmatic Bare-faced Bulbul ''Pycnonotus hualon'' with description of a new genus | journal=Ibis | volume=160 | issue=3 | pages=659–665 | doi=10.1111/ibi.12580}}</ref> As currently defined the genera ''Chlorocichla'' and ''Arizelocichla'' are not [[monophyletic]]. | |||
{{clade| style=font-size:90%;line-height:90% | |||
|1={{clade | |1={{clade | ||
|label1='''Pycnonotidae''' | |||
|1={{clade | |1={{clade | ||
|label1=Africa clade | |label1=Africa clade | ||
| Line 112: | Line 107: | ||
|1=''[[Andropadus]]'' – sombre greenbul | |1=''[[Andropadus]]'' – sombre greenbul | ||
|2={{clade | |2={{clade | ||
|1=''[[ | |1=''[[Calyptocichla]]'' – golden greenbul | ||
|2=''[[ | |2=''[[Stelgidillas]]'' – slender-billed greenbul | ||
}} | }} | ||
}} | }} | ||
| Line 120: | Line 115: | ||
|2={{clade | |2={{clade | ||
|1={{clade | |1={{clade | ||
|1=''[[Bleda]]'' – bristlebills (5 species) | |1=''[[Phyllastrephus]]'' – greenbuls, brownbuls, leaflove (20 species) | ||
|2={{clade | |||
|1=''[[Criniger]]'' – greenbuls (5 species) | |||
|2=''[[Eurillas]]'' – greenbuls (5 species) | |||
}} | |||
}} | |||
|2={{clade | |||
|1=''[[Bleda_(bird)|Bleda]] - [[Bristlebill]]'' – bristlebills (5 species) | |||
|2={{clade | |2={{clade | ||
|1={{clade | |1=''[[Arizelocichla]]'' – greenbuls (12 species) | ||
|2={{clade | |||
|1={{clade | |1={{clade | ||
|1=''[[Atimastillas]]'' – yellow-throated leaflove | |1=''[[Atimastillas]]'' – yellow-throated leaflove | ||
| Line 135: | Line 138: | ||
|1=''[[Chlorocichla]]'' – greenbuls (5 species) | |1=''[[Chlorocichla]]'' – greenbuls (5 species) | ||
|2={{clade | |2={{clade | ||
|1=''[[ | |1=''[[Baeopogon]]'' – greenbuls (2 species) | ||
|2=''[[ | |2=''[[Chlorocichla simplex]]'' – simple greenbul | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
| Line 157: | Line 152: | ||
|2={{clade | |2={{clade | ||
|1={{clade | |1={{clade | ||
|1=''[[Microtarsus]]'' – (7 species) | |||
|2={{clade | |||
|1=''[[Rubigula]]'' – (8 species) | |||
|2={{clade | |||
|1=''[[Pycnonotus]]'' – (34 species) | |||
|2={{clade | |||
|1=''[[Bare-faced_bulbul|Nok]]'' – bare-faced bulbul | |||
|2=''[[Spizixos]]'' – finchbills (2 species) | |||
}} | |||
}} | |||
}} | |||
}} | |||
|2={{clade | |||
|1=''[[Tricholestes]]'' – hairy-backed bulbul | |1=''[[Tricholestes]]'' – hairy-backed bulbul | ||
|2={{clade | |2={{clade | ||
|1={{clade | |1={{clade | ||
|1=''[[ | |1=''[[Alophoixus]]'' – (8 species) | ||
|2=''[[ | |2=''[[Setornis]]'' – hook-billed bulbul | ||
}} | }} | ||
|2={{clade | |2={{clade | ||
| Line 169: | Line 177: | ||
|2={{clade | |2={{clade | ||
|1={{clade | |1={{clade | ||
|1=''[[ | |1=''[[Acritillas]]'' – yellow-browed bulbul | ||
|2=''[[ | |2=''[[Hemixos]]'' – (4 species) | ||
}} | }} | ||
|2={{clade | |2={{clade | ||
|1=''[[ | |1=''[[Hypsipetes]]'' – (25 species) | ||
|2=''[[ | |2=''[[Ixos]]'' – (5 species) | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
| Line 209: | Line 192: | ||
}} | }} | ||
}} | }} | ||
}} | }} | ||
| Line 217: | Line 199: | ||
==Behaviour and ecology== | ==Behaviour and ecology== | ||
===Breeding=== | ===Breeding=== | ||
The bulbuls are generally [[monogamy|monogamous]]. One unusual exception is the [[yellow-whiskered greenbul]] which at least over part of its range appears to be polygamous and engage in a [[Lek (biology)|lekking]] system. Some species also have [[alloparenting]] arrangements, where non-breeders, usually the young from earlier clutches, help raise the young of a dominant breeding pair.{{sfn|Fishpool|Tobias|2005|p=[https://archive.org/details/handbookofbirdso0010unse/page/151/mode/1up 151]}} Up to five speckled eggs are laid in open tree nests and [[avian incubation|incubated]] by the female. Incubation usually lasts between 11 and 14 days, and chicks [[fledge]] after 12–16 days.{{sfn|Fishpool|Tobias|2005|pp=[https://archive.org/details/handbookofbirdso0010unse/page/154/mode/1up 154–155]}} | The bulbuls are generally [[monogamy in animals|monogamous]]. One unusual exception is the [[yellow-whiskered greenbul]] which at least over part of its range appears to be polygamous and engage in a [[Lek (biology)|lekking]] system. Some species also have [[alloparenting]] arrangements, where non-breeders, usually the young from earlier clutches, help raise the young of a dominant breeding pair.{{sfn|Fishpool|Tobias|2005|p=[https://archive.org/details/handbookofbirdso0010unse/page/151/mode/1up 151]}} Up to five speckled eggs are laid in open tree nests and [[avian incubation|incubated]] by the female. Incubation usually lasts between 11 and 14 days, and chicks [[fledge]] after 12–16 days.{{sfn|Fishpool|Tobias|2005|pp=[https://archive.org/details/handbookofbirdso0010unse/page/154/mode/1up 154–155]}} | ||
===Feeding=== | ===Feeding=== | ||
Bulbuls eat a wide range of foods, ranging from fruit to seeds, nectar, small insects and other arthropods and even small vertebrates. The majority of species are frugivorous and supplement their diet with some insects, although there is a significant minority of specialists, particularly in Africa. Open country species in particular are generalists. Bulbuls in the genus ''[[Criniger]]'' and bristlebills in the genus ''[[ Bleda (bird)|Bleda]]'' will join [[mixed-species feeding flock]]s. | Bulbuls eat a wide range of foods, ranging from fruit to seeds, nectar, small insects and other arthropods and even small vertebrates. The majority of species are frugivorous and supplement their diet with some insects, although there is a significant minority of specialists, particularly in Africa. Open country species in particular are generalists. Bulbuls in the genus ''[[Criniger]]'' and bristlebills in the genus ''[[Bleda (bird)|Bleda]]'' will join [[mixed-species feeding flock]]s. | ||
==Relationship to humans== | ==Relationship to humans== | ||
| Line 234: | Line 216: | ||
==External links== | ==External links== | ||
{{Commons category|Pycnonotidae}} | |||
*''[http://ibc.lynxeds.com/family/bulbuls-pycnonotidae Bulbul photos and videos] on the Internet Bird Collection'' | *''[http://ibc.lynxeds.com/family/bulbuls-pycnonotidae Bulbul photos and videos] on the Internet Bird Collection'' | ||
*''[https://web.archive.org/web/20090605205229/http://www.eeb.cornell.edu/winkler/botw/pycnonotidae.html Family characters]'' | *''[https://web.archive.org/web/20090605205229/http://www.eeb.cornell.edu/winkler/botw/pycnonotidae.html Family characters]'' | ||
Latest revision as of 13:58, 7 February 2026
| Bulbul | |
|---|---|
| |
| Brown-eared bulbul (Hypsipetes amaurotis) | |
Scientific classification 
| |
| : | [[Template:Taxonomy/Pycnonotidae]] Gray, GR, 1840 |
| Genera | |
|
See text | |
| Synonyms | |
| |
The bulbuls are members of a family, Pycnonotidae, of medium-sized passerine songbirds, which also includes greenbuls, brownbuls, leafloves, and bristlebills. The family is distributed across most of Africa and into the Middle East, tropical Asia to Indonesia, and north as far as Japan. A few insular species occur on the tropical islands of the Indian Ocean. There are 166 species in 32 genera. While different species are found in a wide range of habitats, the African species are predominantly found in rainforest, whereas Asian bulbuls are predominantly found in more open areas.
Taxonomy[edit | edit source]
The family Pycnonotidae was introduced by the English zoologist George Robert Gray in 1840 as a subfamily Pycnonotinae of the thrush family Turdidae.[1][2]
The Persian word bulbul (بلبل) is sometimes used to refer to the "nightingale" as well as the bulbul, but the English word bulbul refers to the birds discussed in this article.[3]
A few species that were previously considered to be members of the Pycnonotidae have been moved to other families. Several Malagasy species that were formerly placed in the genus Phyllastrephus are now placed in the family Bernieridae.[4][5] In addition, the genus Nicator containing three African species is now placed in a separate family Nicatoridae.[6][7]
A study published in 2007 by Ulf Johansson and colleagues using three nuclear markers found that the genus Andropadus was non-monophyletic. In the subsequent revision, species were moved to three resurrected genera: Arizelocichla, Stelgidillas and Eurillas. Only the sombre greenbul (Andropadus importunus), was retained in Andropadus.[8][9] A study by Subir Shakya and Frederick Shelden published in 2017 found that species in the large genus Pycnonotus formed several deeply divergent clades. The genus was split and genera were resurrected to accommodate these clades.[8][10]
The family forms two main clades. One clade contains species that are only found in Africa; many of these have greenbul in the common name. The second clade contains mostly Asian species but includes a few species that are found in Africa.[10]
List of genera[edit | edit source]
Currently, there are 161 recognized species in 28 genera:[8]
- Genus Andropadus – sombre greenbul (formerly contained many species)
- Genus Stelgidillas – slender-billed greenbul (formerly in Andropadus)
- Genus Calyptocichla – golden greenbul
- Genus Neolestes – black-collared bulbul
- Genus Bleda – bristlebills (5 species)
- Genus Atimastillas – greenbuls (2 species)
- Genus Ixonotus – spotted greenbul
- Genus Thescelocichla – swamp palm bulbul
- Genus Chlorocichla – greenbuls (5 species)
- Genus Baeopogon – greenbuls (2 species)
- Genus Arizelocichla – greenbuls (11 species) (formerly in Andropadus)
- Genus Criniger – greenbuls (5 species)
- Genus Eurillas – greenbuls (5 species) (formerly in Andropadus)
- Genus Phyllastrephus – greenbuls and brownbuls (20 species)
- Genus Tricholestes – hairy-backed bulbul
- Genus Setornis – hook-billed bulbul
- Genus Alophoixus – 8 species (formerly in Criniger)
- Genus Alcurus – striated bulbul
- Genus Iole – 7 species
- Genus Hemixos – 3 species
- Genus Acritillas – yellow-browed bulbul
- Genus Ixos – 5 species
- Genus Hypsipetes – 26 species (includes 3 species formerly in Thapsinillas, one formerly in Cerasophila and one formerly in Microscelis)
- Genus Microtarsus – 7 species
- Genus Rubigula – 8 species (formerly in Pycnonotus)
- Genus Nok – bare-faced bulbul (genus introduced in 2017)
- Genus Spizixos – finchbills (2 species)
- Genus Pycnonotus – 31 species (substantially reduced from earlier classification)
Cladogram[edit | edit source]
Phylogeny based on a study by Subir Shakya and Frederick Shelden published in 2017 with the revised genera as defined in the AviList checklist.[8][10] The positions of the bare-faced bulbul (Nok hualon) and the yellow-browed bulbul (Acritillas indica) are based on a study by Jérôme Fuchs and colleagues published in 2018.[11] As currently defined the genera Chlorocichla and Arizelocichla are not monophyletic. Template:Clade
Description[edit | edit source]
Bulbuls are short-necked slender passerines. The tails are long and the wings short and rounded. In almost all species the bill is slightly elongated and slightly hooked at the end. They vary in length from 13 cm and 13.3 g (0.47 oz) for the tiny greenbul to 29 cm and 93 g (3.3 oz) in the straw-headed bulbul.[12] Overall the sexes are alike, although the females tend to be slightly smaller. In a few species the differences are so great that they have been described as functionally different species. The soft plumage of some species is colorful with yellow, red or orange vents, cheeks, throat or supercilia, but most are drab, with uniform olive-brown to black plumage. Species with dull coloured eyes often sport contrasting eyerings. Some have very distinct crests. Bulbuls are highly vocal, with the calls of most species being described as nasal or gravelly. One author described the song of the brown-eared bulbul as "one of the most unattractive noises made by any bird".[13]
Behaviour and ecology[edit | edit source]
Breeding[edit | edit source]
The bulbuls are generally monogamous. One unusual exception is the yellow-whiskered greenbul which at least over part of its range appears to be polygamous and engage in a lekking system. Some species also have alloparenting arrangements, where non-breeders, usually the young from earlier clutches, help raise the young of a dominant breeding pair.[14] Up to five speckled eggs are laid in open tree nests and incubated by the female. Incubation usually lasts between 11 and 14 days, and chicks fledge after 12–16 days.[15]
Feeding[edit | edit source]
Bulbuls eat a wide range of foods, ranging from fruit to seeds, nectar, small insects and other arthropods and even small vertebrates. The majority of species are frugivorous and supplement their diet with some insects, although there is a significant minority of specialists, particularly in Africa. Open country species in particular are generalists. Bulbuls in the genus Criniger and bristlebills in the genus Bleda will join mixed-species feeding flocks.
Relationship to humans[edit | edit source]
The red-whiskered bulbuls and red-vented bulbuls have been captured for the pet trade in great numbers and have been widely introduced to tropical and subtropical areas, for example, southern Florida, Fiji, Australia and Hawaii. Some species[which?] are regarded as crop pests, particularly in orchards.[citation needed]
In general, bulbuls and greenbuls are resistant to human pressures on the environment and are tolerant of disturbed habitat. Around 13 species are considered threatened by human activities, mostly specialised forest species that are threatened by habitat loss.
References[edit | edit source]
- ↑ Gray, George Robert (1840). A List of the Genera of Birds : with an Indication of the Typical Species of Each Genus. London: R. and J.E. Taylor. p. 28.
- ↑ Bock, Walter J. (1994). History and Nomenclature of Avian Family-Group Names. Bulletin of the American Museum of Natural History. Vol. 222. New York: American Museum of Natural History. pp. 201, 237. hdl:2246/830.
- ↑ Fishpool & Tobias 2005, pp. 161–162.
- ↑ 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. PMID 11475055.
- ↑ Cibois, Alice; David, Normand; Gregory, Steven M. S.; Pasquet, Eric (2010). "Bernieridae (Aves: Passeriformes): a family-group name for the Malagasy sylvioid radiation". Zootaxa. 2554: 65–68. doi:10.11646/zootaxa.2554.1.6.
- ↑ 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'". Proceedings of the Royal Society B. 272 (1565): 849–858. doi:10.1098/rspb.2004.2997. PMC 1599865. PMID 15888418.
- ↑ 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". Molecular Phylogenetics and Evolution. 48 (3): 858–876. doi:10.1016/j.ympev.2008.05.029. PMID 18619860.
- ↑ 8.0 8.1 8.2 8.3 AviList Core Team (2025). "AviList: The Global Avian Checklist, v2025". doi:10.2173/avilist.v2025. Retrieved 17 November 2025.
- ↑ Johansson, U.S.; Fjeldså, J.; Lokugalappatti, L.G.S.; Bowie, R.C.K. (2007). "A nuclear DNA phylogeny and proposed taxonomic revision of African greenbuls (Aves, Passeriformes, Pycnonotidae)". Zoologica Scripta. 36 (5): 417–427. doi:10.1111/j.1463-6409.2007.00290.x. S2CID 84799480.
- ↑ 10.0 10.1 10.2 Shakya, Subir B.; Sheldon, Frederick H. (2017). "The phylogeny of the world's bulbuls (Pycnonotidae) inferred using a supermatrix approach". Ibis. 159 (3): 498–509. doi:10.1111/ibi.12464.
- ↑ Fuchs, J.; Pasquet, E.; Stuart, B.L.; Woxvold, I.A.; Duckworth, J.W.; Bowie, R.C.K. (2018). "Phylogenetic affinities of the enigmatic Bare-faced Bulbul Pycnonotus hualon with description of a new genus". Ibis. 160 (3): 659–665. doi:10.1111/ibi.12580.
- ↑ Dunning, John B. Jr., ed. (2008). CRC Handbook of Avian Body Masses (2nd ed.). CRC Press. ISBN 978-1-4200-6444-5.
- ↑ Fishpool & Tobias 2005, p. 146.
- ↑ Fishpool & Tobias 2005, p. 151.
- ↑ Fishpool & Tobias 2005, pp. 154–155.
Sources[edit | edit source]
- Fishpool, L.D.C.; Tobias, J.A. (2005). "Family Pycnonotidae (Bulbuls)". In del Hoyo, J.; Elliott, A.; Christie, D.A. (eds.). Handbook of the Birds of the World. Vol. 10: Cuckoo-shrikes to Thrushes. Barcelona, Spain: Lynx Edicions. pp. 124–250. ISBN 978-84-87334-72-6.
External links[edit | edit source]
- Bulbul photos and videos on the Internet Bird Collection
- Family characters