The ischnoceran chewing lice (Phthiraptera: Ischnocera) of bulbuls (Aves: Passeriformes: Pycnonotidae), with descriptions of 18 new species

  • Daniel R. Gustafsson Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Laboratory of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, Guangzhou, China
  • Tomas Najer Department of Veterinary Sciences, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences, Kamycka 129, 165 00 Prague 6, Czech Republic
  • Fasheng Zou Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Laboratory of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, Guangzhou, China
  • Sarah E. Bush School of Biological Sciences, University of Utah, 257 S. 1400 E., Salt Lake City, Utah, 84112, USA
Keywords: Philopteridae, Brueelia, Guimaraesiella, Philopteroides, Pycnonotidae


The ischnoceran chewing lice known from bulbuls are discussed and revised, and 18 new species are described. These are: Brueelia celer sp. nov. from Pycnonotus cafer bengalensis Blyth, 1845 and Pycnonotus cafer primrosei Deignan, 1949; Brueelia colindalei sp. nov. from Hemixos castanonotus canipennis Seebohm, 1890; Brueelia doisuthepensis sp. nov. from Alophoixus ochraceus ochraceus (Moore, 1858); Brueelia galeata sp. nov. from Alophoixus pallidus henrici (Oustalet, 1896); Brueelia hermetica sp. nov. from Pycnonotus barbatus layardi Gurney, 1879; Brueelia leiae sp. nov. from Alophoixus flaveolus burmanicus (Oates, 1899); Brueelia robertrankini sp. nov. from Pycnonotus jocosus jocosus (Linnaeus, 1758) and Pycnonotus jocosus pattani Deignan, 1948; Brueelia yunnanensis sp. nov. from Ixos mcclellandii similis (Rothschild, 1921); Guimaraesiella brunneomarginata sp. nov. from Pycnonotus goiaver samarensis Rand & Rabor, 1960 and Pycnonotus goiaver personatus Hume, 1873; Guimaraesiella caligogularis sp. nov. from Pycnonotus plumosus plumosus Blyth, 1845 and Alophoixus bres tephrogenys (Jardine & Selby, 1833); Guimaraesiella cinnamomea sp. nov. from Iole propinqua propinqua (Oustalet, 1903) and Iole viridescens cinnamomeoventris Baker, 1917; Guimaraesiella ixi sp. nov. from Ixos mcclellandii peracensis (Hartert & Butler, 1898); Guimaraesiella lorica sp. nov. from Hypsipetes leucocephalus nigerrimus Gould, 1863; Guimaraesiella mayoensis sp. nov. from Hypsipetes everetti everetti (Tweeddale, 1877); Guimaraesiella phlaoalopha sp. nov. from Alophoixus pallidus henrici (Oustalet, 1896); Philopteroides holosternus sp. nov. from Pycnonotus goiavier goiavier (Scopoli, 1786); Philopteroides longiclypeatus sp. nov. from Hypsipetes everretti samarensis Rand & Rabor, 1959; Philopteroides haerixos sp. nov. from Ixos mcclellandii holtii (Swinhoe, 1861) and Alophoixus pallidus henrici (Oustalet, 1896). The following new host records are provided: Hemixos castanonotus canipennis Seebohm, 1890, for Guimaraesiella flavala (Najer & Sychra in Najer et al., 2012); Pycnonotus blanfordi conradi (Finsch in Finsch & Conrad, 1873) for Philopteroides cucphuongensis Mey, 2004. Philopterus cucphuongensis is tentatively redescribed and illustrated based on specimens from a non-type host species. The species descriptions of the following species are amended slightly, based on re-examinations of type specimens: Brueelia alophoixi Sychra in Sychra et al., 2009; Guimaraesiella cucphuongensis (Najer & Sychra in Najer et al., 2012); Guimaraesiella flavala (Najer & Sychra in Najer et al., 2012). We propose to move Philopterus hiyodori Uchida, 1949, to the genus Craspedorrhynchus Kéler, 1938. The species Sturnidoecus acutifrons (Uchida, 1949) and Penenirmus guldum (Ansari, 1955) are considered species inquirenda. An updated checklist of ischnoceran lice known from bulbuls is provided, as well as a key to all ischnoceran species known from bulbuls.


Ansari R.A.M. 1947. Mallophaga (Ischnocera) infesting birds in the Punjab (India). Proceedings of the National Institute of Science India 13: 253–303.

Ansari R.A.M. 1955. Studies on the ischnoceron [sic] Mallophaga infesting birds in Pakistan. Proceedings of the Seventh Pakistan Scientific Conference, Biology, Bahawalpur 5–59.

Ansari R.A.M. 1956. Studies in phthirapteran parasites (Mallophaga) infesting birds in the Panjab. Indian Journal of Entomology 17: 394–400.

Ansari R.A.M. 1958a. Studies on Ischnoceran Mallophaga infesting birds in the Panjab [Part I]. Indian Journal of Entomology 20: 46–62.

Ansari R.A.M. 1958b. Studies on Ischnoceran Mallophaga infesting birds in the Panjab [Part II]. Indian Journal of Entomology 20: 77–103.

Blyth E. 1845. Notices and descriptions of various new or little known species of birds. Journal of the Asiatic Society of Bengal 14: 546–602.

Blyth E. 1846. Notices and descriptions of various new and little known species of birds. Journal of the Asiatic Society of Bengal 15: 1–54.

Bush S.E., Weckstein J.D., Gustafsson D.R., Allen J., DiBlasi E., Shreve S.M., Boldt R., Skeen H.R. & Johnson K.P. 2016. Unlocking the black box of feather louse diversity: a molecular phylogeny of the hyper-diverse genus Brueelia. Molecular Phylogenetics and Evolution 94: 737–751.

Chu X., Dik B., Gustafsson D.R., Che X., Zhang Q. & Zou F. 2019. The influence of host body size and food guild on prevalence and mean intensity of chewing lice (Phthiraptera) on birds in Southern China. Journal of Parasitology 105: 334–344.

Clay T. 1951. An introduction to the classification of the avian Ischnocera (Mallophaga): Part I. Transactions of the Royal Entomological Society of London 102: 171–194.

Clements J.F., Schulenberg T.S., Iliff M.J., Roberson D., Fredericks T.A., Sullivan B.L. & Wood C.L. 2019. The eBird/Clements Checklist of Birds of the World: ver. 2019. Available from (accessed 23 Sep. 2019).

Eichler W. 1949. Phthirapterorum nova genera. Bolletino della Società Entomologica Italiana 79: 11–13.

Eichler W. 1951. Die Federlinge der Drosseln. In: Bedeutung der Vogelwelt in Forschung und Praxis – Zusammenstellung der Vorträge gehalten auf der Ersten Ornithologen-Tagung in der Deutschen Demokratischen Republik am 21 und 22 Oktober 1950 in Leipzig: 29–47. Leipzig, Germany.

Eichler W. 1952. Notulae Mallophagologicae. XXVI. Rhombiceps n. g. und andere neue Federlings-gattungen. Zoologische Anzeiger 149: 74–78.

Gajdosova M., Sychra O., Kreisinger J., Sedlacek O., Nana E.D., Albrecht T. & Munclinger P. 2020. Patterns of host-parasite associations in tropical lice and their passerine hosts in Cameroon. Ecology and Evolution 10: 6512–6524.

Gállego J., Martin Mateo M.P. & Aguirre J.M. 1987. Malófagos de rapaces españolas. II. Las especies de género Craspedorrhynchus Kéler, 1938 parásitas de Falconiformes, con descripcíon de tres especies nuevas. Eos 63: 31–66.

Gustafsson D.R. & Bush S.E. 2017. Morphological revision of the hyperdiverse Brueelia-complex (Insecta: Phthiraptera: Ischnocera: Philopteridae) with new taxa, checklists and generic key. Zootaxa 4313: 1–443.

Gustafsson D.R., Malysheva O.D., Tolstenkov O.O. & Bush S.E. 2019a. Five new species of Guimaraesiella (Phthiraptera: Ischnocera) from broadbills (Aves: Passeriformes: Calyptomenidae, Eurylaimidae). Journal of Parasitology 105: 846–857.

Gustafsson D.R., Bush S.E. & Palma R.L. 2019b. The genera and species of the Brueelia-complex (Phthiraptera: Philopteridae) described by Mey (2017). Zootaxa 4615: 252–284.

Gustafsson D.R., Clayton D.H. & Bush S.E. 2019c. Twelve new species of Guimaraesiella (Phthiraptera: Ischnocera: Philopteridae) from “babblers” (Passeriformes: Leiothrichidae, Pellorneidae, Timaliidae) with a description of a new subgenus and a key to its species. Zootaxa 4543: 451–497.

Gustafsson D.R., Zou F., Oslejskova L., Najer T. & Sychra O. 2019d. Four new species of Brueelia Kéler, 1936 (Phthiraptera: Ischnocera) from African hosts, with a redescription of Nirmus bicurvatus Piaget, 1880. European Journal of Taxonomy 507: 1–48.

Hellenthal R.A. & Price R.D. 2003. The genus Myrsidea Waterston (Phthiraptera: Menoponidae) from bulbuls (Passeriformes: Pycnonotidae), with descriptions of 16 new species. Zootaxa 354: 1–20.

Hopkins G.H.E. & Clay T. 1952. A Check List of the Genera and Species of Mallophaga. British Museum (Natural History), London, United Kingdom.

Johnson K.P. & Price R.D. 2006. Five new species of Myrsidea Waterston (Phthiraptera: Menoponidae) from bristlebills and greenbuls (Passeriformes: Pycnonotidae) in Ghana. Zootaxa 1177: 27–37.

Kéler S. von. 1936. Über einige Mallophagen aus Rossitten. Arbeiten in morphologische und taxonomische Entomologie von Berlin-Dahlem 3: 256–264.

Light J.E., Nessner C.E., Gustafsson D.R., Wise S.R. & Voelker G. 2016. Remarkable levels of avian louse (Insects: Phthiraptera) diversity in the Congo Basin. Zoologica Scripta 45: 538–551.

Linnaeus C. von. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. 10th Edition. Launrentii Salvi, Holmiae [= Stockholm, Sweden].

McClure H.E., Ratanaworabhan N., Emerson K.C., Hoogstraal H., Nadchatram N., Kwanyuen P., Atyeo W.T., Maa T.C., Wilson N. & Wayupong L. 1973. Some Ectoparasites of the Birds of Asia. Applied Scientific Research Corporation of Thailand, Bangkok, Thailand

Menkhorst P., Rogers D., Clarke R., Davies J., Marsack P. & Franklin K. 2017. The Australian Bird Guide. Christopher Helm, Bloomsbury, London & New York, United Kingdom and United States.

Mey E. 1994. Beziehungen zwischen Larvenmorphologie und Systematik der Adulti bei den Vogel-Ischnozeren (Insecta, Phthiraptera, Ischnocera). Mitteilungen aus dem Zoologischen Museum in Berlin 70: 3–84.

Mey E. 2001. A new Craspedorrhynchus species (Phthiraptera, Ischnocera) from Australia, with an annotated checklist of this chewing louse genus. Deutsche Entomologische Zeitschrift 48: 117–132.

Mey E. 2004. Zur Taxonomie, Verbreitung und parasitophyletischer Evidenz des Philopterus-Komplexes (Insecta, Phthiraptera, Ischnocera). Ornithologischer Anzeiger 43: 149–203.

Mey E. 2017. Neue Gattungen und Arten aus dem Brueelia-Komplex (Insecta, Phthiraptera, Ischnocera, Philopteridae s. l.). Rudolstädter Naturhistorische Schriften 22: 85–215.

Mey E. & Barker S.C. 2014. Eine neue auf den Feenvögeln (Irenidae) lebende Brueelia-Art (Insecta, Phthiraptera, Ischnocera, Philopteridae), nebst Anmerkungen zur Gattung Brueelia Kéler, 1936 sensu lato. Rudolstädter Naturhistorische Schriften 19: 73–114.

Najer T., Sychra O., Hung N.M., Capek M., Podzemny P. & Literak I. 2012. New species and new records of chewing lice (Phthiraptera: Amblycera and Ischnocera) from bulbuls (Passeriformes: Pycnonotidae) in Vietnam. Zootaxa 3357: 37–48.

Najer T., Papousek I., Adam C., Trnka A., Quach V.T., Nguyen C.N., Figure R., Literak I. & Sychra O. 2020. New records of Philopterus (Ischnocera: Philopteridae) from Acrocephalidae and Locustellidae, with description of one new species from Regulidae. European Journal of Taxonomy 632: 1–37.

Najer T., Papousek I., Sychra O., Sweet A.D. & Johnson K.P. 2021. Combining nuclear and mitochondrial loci provides phylogenetic information in the Philopterus Complex of lice (Psocodea: Ischnocera: Philopteridae). Journal of Medical Entomology 58: 252–260.

Naz S., Najer T. & Gustafsson D.R. 2020. An annotated list of the species of lice (Insecta: Phthiraptera) described by Mohammad A.-R. Ansari. Zootaxa 4809: 401–448.

Neumann L.G. 1906. Notes sur les Mallophages. Bulletin de la Société zoologique de France 31: 54–60.

Nitzsch C.L. 1818. Die Familien und Gattungen der Thierinsekten (Insecta epizoica); als ein Prodromus einer Naturgeschichte derselben. E.F. Germar’s Magazin der Entomologie 3: 261–318.

Paton K. B. 1985. The red-whiskered bulbul in South Australia. South Australian Ornithologist 29: 189–190.

Price R.D., Hellenthal R.A. & Palma R.L. 2003. World checklist of chewing lice with host associations and keys to families and genera. In: Price R.D., Hellenthal R.A., Palma R.L., Johnson K.P. & Clayton D.H. (eds) The Chewing Lice: World Checklist and Biological Overview. Illinois Natural History Survey Special Publication 24. Champaign, Illinois, United States.

Rand A.L. & Rabor D.S. 1959. Three new birds from the Philippine Islands. Fieldiana Zoology 39: 275–277.

Rand A.L. & Rabor D.S. 1960. Birds of the Philippine Islands: Siquijor, Mount Malindang, Bohol, and Samar. Fieldiana Zoology 35: 223–442.

Riley J.H. 1940. Six new forms of birds from Indochina. Proceedings of the Biological Society of Washington 53: 131–134.

Rothschild L.W. 1921. On a collection of birds from west-central and north-western Yunnan. Novitates Zoologicae 28: 14–67.

Shakya S.B. & Sheldon F.H. 2017. The phylogeny of the world’s bulbuls (Pycnonotidae) inferred using a supermatrix approach. Ibis 159: 498–509.

Soler Cruz M.D., Benítez Rodríguez R. & Florido Navio A. 1987. Some Mallophaga (Brueeliinae) from birds of the family Fringillidae. Acta Parasitologica Polonica 31: 241–246.

Sychra O., Literak I., Hung N.M. & Podzemny P. 2009. Chewing lice from wild passerines (Aves, Passeriformes) from Vietnam, with description of a new species of the genus Brueelia (Phthiraptera, Ischnocera, Philopteridae). Acta Parasitologica 54: 154–157.

Takano O.M., Voelker G., Gustafsson D.R. & Light J.E. 2019. Molecular phylogeny and novel host associations of avian chewing lice (Insecta: Phthiraptera) from South Africa. Systematic Entomology 44: 289–304.

Temminck C.J. 1826. Nouveau recueil de planches coloriées d’oiseaux: pour servir de suite et de complément aux planches enluminées de Buffon, édition in-folio et in-4° de l’Imprimerie royale, 1770. Livraison 84. Chez Legras Imbert et Comp., Strasbourg, France.

Uchida S. 1948. Studies on the biting-lice (Mallophaga) of Japan and adjacent territories (Suborder Ischnocera Pt I). The Japanese Medical Journal 1: 303–326.

Uchida S. 1949. Studies on the biting-lice (Mallophaga) of Japan and adjacent territories (Suborder Ischnocera Pt II). The Japanese Medical Journal 1: 535–556.

Valim M.P. & Palma R.L. 2013. Three new species of the genus Philopteroides Mey, 2004 (Phthiraptera, Ischnocera, Philopteridae) from New Zealand. ZooKeys 297: 71–89.

Zheng G. (ed.) 2017. A Checklist on the Classification and Distribution of the Birds of China (Third Edition). Science Press, Beijing, China.

Złotorzycka J. 1964. Mallophaga parasitizing Passeriformes and Pici. II. Brueeliinae. Acta Parasitologica Polonica 12: 239–282.

How to Cite
Gustafsson, D. R., Najer, T., Zou, F., & Bush, S. E. (2022). The ischnoceran chewing lice (Phthiraptera: Ischnocera) of bulbuls (Aves: Passeriformes: Pycnonotidae), with descriptions of 18 new species. European Journal of Taxonomy, 800(1), 1-88.