Perninae

European honey buzzard Pernis apivorus, copyright Aurelie Laurent.

Belongs within: Accipitridae.

The Perninae are an assemblage of lightly built raptors lacking a bony shield above the eye (Lerner & Mindell 2005).

Honey, it’s a buzzard
Published 27 July 2024

The long-winged, relatively lightly built birds of prey known as ‘kites’ have long been thought of as one of the more primitive types of raptors, owing to their often generalist habits. Nevertheless, it has generally been agreed (at least in the last century) that they are not a uniform lineage, being divided between multiple subfamilies. Among these is the Perninae, including the honey buzzards and their relatives.

Black baza Aviceda leuphotes, copyright Vijay Anand Ismavel.

As defined by recent phylogenetic analyses, the Perninae include nine genera of medium-sized raptors (Catanach et al. 2025) with 19 species currently recognised. Though diverse in appearance and habits, pernines are generally united by the absence of the projecting bony shield above the eye found in many other accipitrids (Lerner & Mindell 2005). As a result, they tend to have more of a ‘wide-eyed’ look than other birds of prey. Most feed on a mixture of insects and small vertebrates which they may find by sallying forth from arboreal perches or swooping low over vegetation. The Pacific baza Aviceda subcristata may even eat small fruits. The honey buzzards of the genus Pernis get their name from a particular fondness for social insects and their larvae, often following foraging bees and wasps to find their nests. The hook-billed kites of the genus Chondrohierax are specialists in arboreal snails that they often find by clambering and hopping between branches in a manner more reminiscent of a parrot than a hawk. Many populations of the more widespread C. uncinatus show a strong dimorphism in beak size with large-beaked individuals taking different snail species from their smaller-beaked compatriots (Thiollay 1994).

Hook-billed kite Chondrohierax uncinatus uncinatus, copyright Nick Athanas.

Pernines are most diverse in the Indo-Australian region and South America. Their only representative in Europe is the European honey buzzard Pernis apivorus that migrates between the western Palaearctic and sub-Saharan Africa. Other species of Pernis are found in eastern Asia. The Madagascar serpent-eagle Eutriorchis astur is perhaps the most divergent member of the clade, having previously been associated with other ‘serpent-eagles’ in the subfamily Circaetinae. Eutriorchis astur is extremely endangered and was even feared to be extinct for much of the 20th Century. The cuckoo-hawks and bazas of the genus Aviceda are widespread in much of the Old World tropics; the name ‘cuckoo-hawk’ applied to African species presumably refers to their appearance and hunting style rather than their breeding habits.

Swallow-tailed kite Elanoides forficatus, copyright Joe Nicholson.

In South America, pernines are represented by the Chondrohierax hook-billed kites, the pale kites of the genus Leptodon, and the swallow-tailed kite Elanoides forficatus. The ranges of more widespread species extend into southern North America, especially the swallow-tailed kite that migrates as far north as the southern United States. Two Neotropical pernine species are also considered endangered: the Cuban kite Chondrohierax wilsonii and the white-collared kite Leptodon forbesii. The white-collared kite, restricted to a small area around Pernambuco in Brazil, was until recently mistaken for an aberrant form of the more widespread grey-headed kite Leptodon cayanensis.

Black-breasted buzzard Hamirostra melanosternon, copyright Marie Tarrant.

As well as the Aviceda bazas, the Australasian region is home to species of the genera Henicopernis, Lophoictinia and Hamirostra. Henicopernis includes the long-tailed honey buzzards of New Guinea and surrounding islands. As their name suggests, these are similar in appearance to Pernis though phylogenetic analyses place them closer to other Australasian species. The square-tailed kite Lophoictinia isura and black-breasted buzzard Hamirostra melanosternon are both found on the Australian continent. These two species are similar in appearance to kites of another subfamily, the Milvinae, and have historically been placed therein. The black-breasted buzzard is a relatively large species with a more vertebrate-focused diet than other pernines, and will also scavenge on carrion. The square-tailed kite has more delicate tastes, eschewing carrion and often preferring small birds. It has, we are told, a particular fondness for honeyeaters.

Systematics of Perninae
<==PerninaeLM05
|--Eutriorchis astur Sharpe 1875CHP25
`--+--+--Leptodon Sundevall 1836CHP25, B94 (see below for synonymy)
| | |--L. cayanensis (Latham 1790)CHP25 [=Cymindis cayanensisSS66]
| | |--L. forbesiJT12
| | `--L. uncinatusS18
| `--ChondrohieraxML23
| |--C. uncinatus (Temminck 1822)CHP25
| | |--C. u. uncinatusUSDI77
| | `--C. u. mirusUSDI77
| `--C. wilsoniiJT12
`--+--AvicedaCHP25
| | i. s.: A. caferS66
| | A. jerdoniJT12
| | A. madagascariensisJT12
| |--A. cuculoides Swainson 1837CHP25
| | |--A. c. cuculoidesM-PG70
| | |--A. c. batesi (Swann 1920) [=Baza batesi]M-PG70
| | |--A. c. emini (Reichenow 1894) [=Baza emini]M-PG70
| | `--A. c. verreauxi Lafresnaye 1846M-PG70
| `--+--A. leuphotesCHP25
| `--A. subcristata (Gould 1838)CHP25, WS48 [=Lepidogenys subcristatusWS48]
| |--A. s. subcristatusJ06
| |--A. s. reinwardtiiJ06
| `--A. s. stenozonaJ06
`--+--Pernis Cuvier 1817CHP25, M02
| |--P. apivorus (Linnaeus 1758)CHP25, M02 [=Falco apivorusM02, Circus apivorusF17]
| | |--P. a. apivorusI92
| | `--P. a. japonicus Kuroda 1925I92
| `--+--P. ptilorhynchusCHP25 [=Falco (Pernis) ptilorhynchusHH55, F. (P.) ptilonorhynchus (l. c.)HH55]
| | |--P. p. ptilorhynchusVP89
| | `--P. p. orientalisVP89
| `--+--P. celebensisCHP25
| `--P. steereiCHP25
`--+--Elanoides Vieillot 1818CHP25, B94 [Elanoidinae]
| `--E. forficatus (Linnaeus 1758)CHP25
| |--E. f. forficatusE52
| `--E. f. yetapaE52
`--+--Hamirostra Brown 1846CHP25, B94 [Hamirostrinae]
| |--H. isuraS90
| `--H. melanosternon (Gould 1840) (see below for synonymy)WS48
`--+--Lophoictinia isura (Gould 1838)JT12, WS48 (see below for synonymy)
`--HenicopernisJT12
|--H. infuscatusJT12
`--H. longicaudaJT12

Hamirostra melanosternon (Gould 1840) [=Buteo melanosternon; incl. Gypoictinia melanosterna decepta Mathews 1912]WS48

Leptodon Sundevall 1836 non Mohr 1803 (ICBN)CHP25, B94 [incl. Cymindis Cuvier 1816 non Latreille 1806B94; Cymindidinae, Leptodontinae]

Lophoictinia isura (Gould 1838)JT12, WS48 [=Milvus isurusWS48; incl. M. isurus westraliensis Mathews 1912WS48]

*Type species of generic name indicated

References

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[CHP25] Catanach, T. A., M. R. Halley & S. Pirro. 2025. Enigmas no longer: using ultraconserved elements to place several unusual hawk taxa and address the non-monophyly of the genus Accipiter (Accipitriformes: Accipitridae). Biological Journal of the Linnean Society 144 (2): blae028.

[E52] Eisenmann, E. 1952. Annotated list of birds of Barro Colorado Island, Panama Canal Zone. Smithsonian Miscellaneous Collections 117 (5): 1–62.

[F17] Forster, T. 1817. A synoptical catalogue of British birds; intended to identify the species mentioned by different names in several catalogues already extant. Forming a book of reference to observations on British ornithology. Nichols, Son, and Bentley: London.

[HH55] Husson, A. M., & L. B. Holthuis. 1955. The dates of publication of “Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche Bezittingen” edited by C. J. Temminck. Zoologische Mededelingen 34 (2): 17–24.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[J06] Johnstone, R. E. 2006. The birds of Gag Island, Western Papuan islands, Indonesia. Records of the Western Australian Museum 23 (2): 115–132.

[LM05] Lerner, H. R. L., & D. P. Mindell. 2005. Phylogeny of eagles, Old World vultures, and other Accipitridae based on nuclear and mitochondrial DNA. Molecular Phylogenetics and Evolution 37: 327–346.

[M-PG70] Mackworth-Praed, C. W., & C. H. B. Grant. 1970. African Handbook of Birds ser. 3 vol. 1. Birds of West Central and Western Africa. Longman: London.

[ML23] Mather, E. K., M. S. Y. Lee, A. B. Camens & T. H. Worthy. 2023. A giant raptor (Aves: Accipitridae) from the Pleistocene of southern Australia. Journal of Ornithology 164: 499–526.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[S66] Schlegel, H. 1866. Communication from, on mammals and birds collected in Madagascar. Proceedings of the Zoological Society of London 1866: 419–426.

[SS66] Sclater, P. L., & O. Salvin. 1866. Catalogue of birds collected by Mr. E. Bartlett on the River Uyacali, Eastern Peru, with notes and descriptions of new species. Proceedings of the Zoological Society of London 1866: 175–201.

[S18] Stone, W. 1918. Birds of the Panama Canal Zone, with special reference to a collection made by Mr. Lindsey L. Jewel. Proceedings of the Academy of Natural Sciences of Philadelphia 70: 239–280.

[S90] Storr, G. M. 1990. Birds of Shark Bay area, Western Australia. In: Berry, P. F., S. D. Bradshaw & B. R. Wilson (eds) Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee pp. 299–311. Western Australian Museum.

Thiollay, J. M. 1994. Family Accipitridae (hawks and eagles). In: Hoyo, J. del, A. Elliott & J. Sargatal (eds) Handbook of the Birds of the World vol. 2. New World Vultures to Guineafowl pp. 52–205. Lynx Edicions: Barcelona.

[USDI77] United States Department of the Interior. 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

[VP89] Viney, C., & K. Phillipps. 1989. Birds of Hong Kong 5th ed. Government Printer: Hong Kong.

[WS48] Whittell, H. M., & D. L. Serventy. 1948. A systematic list of the birds of Western Australia. Public Library, Museum and Art Gallery of Western Australia, Special Publication 1: 1–126.

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