Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Cladus: Avemetatarsalia
Cladus: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauriformes
Cladus: Dracohors
Cladus: Dinosauria
Cladus: Saurischia
Cladus: Eusaurischia
Subordo: Theropoda
Cladus: Neotheropoda
Cladus: Averostra
Cladus: Tetanurae
Cladus: Avetheropoda
Cladus: Coelurosauria
Cladus: Tyrannoraptora
Cladus: Maniraptoromorpha
Cladus: Maniraptoriformes
Cladus: Maniraptora
Cladus: Pennaraptora
Cladus: Paraves
Cladus: Eumaniraptora
Cladus: Avialae
Infraclassis: Aves
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Ornithuromorpha
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Cladus: Neoaves
Cladus: Telluraves
Cladus: Australaves
Ordo: Passeriformes
Subordo: Passeri
Infraordo: Corvida
Superfamilia: Meliphagoidea
Familia: Meliphagidae
Genus: Ptilotula
Species: P. flavescens – P. fusca – P. keartlandi – P. ornata – P. penicillata – P. plumula
Name
Ptilotula Mathews, 1912
Typus: Ptilotis flavescens Gould, 1840 = Ptilotula flavescens
References
Primary references
Mathews, G.M. 1912. A Reference-List to the Birds of Australia. Novitates Zoologicae, a journal of zoology in connection with the Tring Museum 18(3): 171–455 BHL DOI: 10.5962/bhl.part.1694 Reference page. p. 414
Additional references
Nyári, A.S. & Joseph, L. 2011. Systematic dismantlement of Lichenostomus improves the basis for understanding relationships within the honeyeaters (Meliphagidae) and the historical development of Australo-Papuan communities. Emu 111: 202–211. DOI: 10.1071/MU10047 ResearchGate Reference page.
Ptilotula is a genus of honeyeater consisting of species occurring in Australia and Papua New Guinea. The genus consists of six former members of Lichenostomus, and was created after a molecular analysis showed the genus was polyphyletic.[2] The International Ornithologists' Union accepted this change and officially included the genus in reference lists from 2013.[3] The type species is the yellow-tinted honeyeater (Ptilotula flavescens).[2] Birds in this genus typically occupy dry open forest and woodland habitats, and can be found in arid and semi-arid environments.[2][4]
Species
The genus includes six species:[3]
Yellow-tinted honeyeater (Ptilotula flavescens)
Grey-headed honeyeater (P. keartlandi)
Yellow-plumed honeyeater (P. ornata)
Grey-fronted honeyeater (P. plumula)
Fuscous honeyeater (P. fusca)
White-plumed honeyeater (P. penicillata)
Description
Ptilotula species are medium-sized honeyeaters ranging from 13 cm to 18.5 cm in size, with the yellow-tinted honeyeater (P. flavescens) smallest at 13–15.5 cm and the yellow-plumed honeyeater (P. ornatus) largest at 14–18.5 cm.[5] The characteristic features all species share are a plain face with a black proximal stripe and either yellow or white distal plumes across the sides of the neck.[4]
Systematics and Taxonomy
Until recently, the members of Ptilotula were considered part of Lichenostomus.[2] They were recognised as a clade within this genus, displaying similarities in morphological characters and habitat preferences.[2][4] Extensive molecular phylogenetic analyses of the honeyeater family Meliphagidae occurred during the first decade of the 21st century, resulting in Lichenostomus being split into seven genera.
The name Ptilotula was first proposed by the Australian ornithologist Gregory Mathews in 1912.[6][7] Prior to 1912 most honeyeaters were placed in either Meliphaga or Melithreptus.[6] He attempted to resolve this by placing 14 species into Ptilotis, a genus originally described by John Gould; but Mathews recognised his treatment was polyphyletic. He rectified this by creating several new genera from Ptilotis, placing the white-plumed honeyeater (P. penicillatus) and the yellow-tinted honeyeater (P. flavescens) in Ptilotula.[8] However, when contributing to the Second Official Checklist of the Birds of Australia, the Royal Australasian Ornithologists Union rejected Mathews' phylogenic treatment as they did not agree with the formation of so many new genera. Mathews compromised by allowing these species to remain in Meliphaga and noting Ptilotula as a sub-genus.[9][10]
In subsequent works, Mathews continued to list Ptilotula as a genus rather than sub-genus.[10][11] In addition to the two species listed above, he included the grey-headed honeyeater (P. keartlandi), the yellow-plumed honeyeater (P. ornatus) and the grey-fronted honeyeater (P. plumulus) in the genus. Nevertheless, other authors continued to follow the RAOU standard using Meliphaga for all these species.[12][13][14]
In 1975 the Australian ornithologist Richard Schodde argued that the criteria used to determine membership in Meliphaga was too broad and that if applied consistently, more than half of the family would be placed in this genus.[15] He split Meliphaga into three genera, placing the Ptilotula clade in Lichenostomus. The development of molecular analyses resulted in later research which agreed with Schodde's assessment, but while identifying Ptilotula as a clade early techniques were unable to give sufficient weight to warrant a split from Lichenostomus.[4] By 2010 newer techniques clearly showed that Lichenostomus was polyphyletic and needed to be reviewed.[16]
In 2011, Nyari and Joseph were finally able to show that the Ptilotula clade warranted promotion to genus.[2] Their assessment confirmed the evolutionary relationship Mathews had proposed in 1931, albeit with the inclusion of the fuscous honeyeater (P. fuscus) which he had placed in the monotypic genus Paraptilotis.[11] They also showed that three other Lichenostomus species, varied honeyeater (Gavicalis versicolor), singing honeyeater (G. virescens) and mangrove honeyeater (G. fasciogularis) could also be placed in Ptilotula due to being closely related. These three species are much larger (16–24 cm) and all share a black band from the bill through the eye to the neck, a character absent in the Ptilotula complex.[5] For this reason, they decided not to include them in Ptilotula and instead placed them in their own genus Gavicalis.
Evolutionary history
Ptilotula are derived from an unknown ancestor which occupied dry woodland in central Australia.[4] Walter Boles described a fossil leg bone found in Riversleigh, Queensland from an as yet unnamed Pliocene species as characteristic of tibias from the Lichenostomus-Meliphaga complex.[17] It is most similar in size to those of P.keartlandi and P.plumulus, two species which are resident in the region [17] and closely related sister species.[2] The environment has been dry woodland since the late Miocene to early Pliocene[18] hence the similarities to extant species indicate this fossil may represent an ancestor to these species.[17]
References
"Melaphagidae". aviansystematics.org. The Trust for Avian Systematics. Retrieved 2023-07-16.
Nyári, Árpád S.; Joseph, Leo (2011). "Systematic dismantlement of Lichenostomus improves the basis for understanding relationships within the honeyeaters (Meliphagidae) and historical development of Australo–Papuan bird communities". Emu. 111 (3): 202–211. doi:10.1071/mu10047. S2CID 85333285.
Gill, Frank; Donsker, David (eds.). "Honeyeaters". World Bird List Version 6.1. International Ornithologists' Union. Retrieved 28 January 2016.
Christidis, Len; Schodde, R. (1993). "Relationships and radiations in the Meliphagine Honeyeaters, Meliphaga, Lichenostomus and Xanthotis (Aves, Meliphagidae) - Protein evidence and its integration with morphology and ecogeography". Australian Journal of Zoology. 41 (3): 293–316. doi:10.1071/ZO9930293.
Higgins, P.J.; Peter, J.M.; Steele, W.K. (2001). Handbook of Australian, New Zealand and Antarctic Birds: Tyrant-flycatchers to Chats. Oxford, England.: Oxford University Press.
Mathews G.M. (1912). "A reference-list to the birds of Australia". Novitates Zoologicae. 18: 171–455 [414]. doi:10.5962/bhl.part.1694.
Salomonsen, F. (1967). "Family Maliphagidae, Honeyeaters". In Paynter, R.A. Jnr. (ed.). Check-list of birds of the world (Volume 12). Vol. 12. Cambridge, Mass.: Museum of Comparative Zoology. p. 365.
Mathews, Gregory (1913). A list of the birds of Australia. London: Witherby & Co. pp. 280–281.
Royal Australasian Ornithologists' Union Checklist Committee (1926). Official checklist of the birds of Australia : supplements 1 to 9 (2nd ed.). Melbourne: H. J. Green.
Mathews, Gregory (1946). A working list of Australian birds including the Australian quadrant and New Zealand. Sydney: Shepherd & Newman.
Mathews, Gregory (1931). A list of the birds of Australasia (including New Zealand, Lord Howe and Norfolk Islands and the Australasian Antarctic quadrant). London: Taylor and Francis for the author.
Terril, S.E.; Rix, C.E. (March 1, 1950). ". The birds of South Australia- their Distribution and Habitat". The South Australian Ornithologist. 19 (6–8): 53–100.
Officer, Hugh R. (Brigadier) (1964). Australian Honeyeaters. Melbourne: Bird Observers Club. pp. 39–63.
An Index of Australian Bird Names. Division of Wildlife Research Technical paper No. 20. CSIRO Division of Wildlife Research. 1969. pp. 54–56.
Scodde, R. (1975). Interim List of Australian Songbirds. Passerines. Melbourne: Royal Australasian Ornithologists Union.
Gardner, Janet L.; Trueman, John W.H.; Ebert, Daniel; Joseph, Leo; Magrath, Robert D. (2010). "Phylogeny and evolution of the Meliphagoidea, the largest radiation of Australasian songbirds". Molecular Phylogenetics and Evolution. 55 (3): 1087–1102. doi:10.1016/j.ympev.2010.02.005. PMID 20152917.
Boles, W. (2005). "Fossil honeyeaters (Meliphagidae) from the Late Tertiary of Riversleigh, north-western Queensland". Emu. 105 (1): 21–26. doi:10.1071/MU03024. S2CID 82659390.
Archer, Michael; Godthelp, Henk; Hand, Suzanne J.; Megirian, Dirk (1989). "Fossil Mammals of Riversleigh, North-western Queensland: Preliminary Overview of Biostratigraphy, Correlation and Environmental Change". Australian Zoologist. 25 (2): 29–66. doi:10.7882/AZ.1989.001.
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