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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
Ordo: 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: Euavialae
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: Passerida
Superfamilia: Sylvioidea

Familia: Phylloscopidae
Genera: Phylloscopus
Name

Phylloscopidae Alström, Ericson, Olsson, & Sundberg, 2006
References
Primary references

Alström, P.; Ericson, P.G.P.; Olsson, U.; & Sundberg, P. 2006. Phylogeny and classification of the avian superfamily Sylvioidea. Molecular Phylogenetics and Evolution 38 (2): 381–397. DOI: 10.1016/j.ympev.2005.05.015 PDF Reference page.

Additional references

Alström, P., Rheindt, F.E., Zhang, R., Zhao, M., Wang, J., Zhu, X., Gwee, C.Y., Hao, Y., Ohlson, J., Jia, C., Prawiradilaga, D.M., Ericson, P.G.P., Lei, F., Olsson, U. 2018. Complete species-level phylogeny of the leaf warbler (Aves: Phylloscopidae) radiation. Molecular Phylogenetics and Evolution 126: 141–152 DOI: 10.1016/j.ympev.2018.03.031 Reference page.

Vernacular names
Deutsch: Laubsängerartige
English: Leaf Warblers
中文: 柳莺科

Leaf warblers are small insectivorous passerine birds belonging to the genus Phylloscopus.

Leaf warblers were formerly included in the Old World warbler family but are now considered to belong to the family Phylloscopidae, introduced in 2006. The family originally included the genus Seicercus, but all species have been moved to Phylloscopus in the most recent classification. Leaf warblers are active, constantly moving, often flicking their wings as they glean the foliage for insects along the branches of trees and bushes. They forage at various levels within forests, from the top canopy to the understorey. Most of the species are markedly territorial both in their summer and winter quarters. Most are greenish or brownish above and off-white or yellowish below. Compared to some other "warblers", their songs are very simple. Species breeding in temperate regions are usually strongly migratory.
Illustration of Phylloscopus bonelli, Phylloscopus coronatus, Phylloscopus trochilus, Phylloscopus sibilatrix by John Gerrard Keulemans

Description

The species are of various sizes, often green-plumaged above and yellow below, or more subdued with greyish-green to greyish-brown colours, varying little or not at all with the seasons. The tails are not very long and contain 12 feathers (unlike the similar Abroscopus species, which have 10 tail feathers). Many species are more easily identified by their distinctive songs than their dull plumage.[1] These are very small passerines with adult body masses that can vary from 3.5 to 17 g (0.12 to 0.60 oz) and in some cases, such as the Chinese leaf warbler, are among the lightest passerines anywhere. Several of the larger species are similar in size including the large-billed leaf warbler, Radde's warbler and the pale-legged leaf warbler.[2] Total length can vary from 9 to 14.5 cm (3.5 to 5.7 in).[3]
Distribution and habitat

Its members occur in Eurasia, ranging into Wallacea and Africa with one species, the Arctic warbler, breeding as far east as Alaska. Many of the species breed at temperate and high latitudes in Eurasia and migrate substantial distances to winter in southeastern Asia, India, or Africa. One example is Tickell's leaf warbler, which breeds in scrub at high elevation in the Himalayas and on the Tibetan Plateau and then moves down-slope and south to winter in the Himalayan foothills of India and Burma.[4] Most live in forest and scrub and many are canopy or sub-canopy dwellers.
Behavior and ecology

The family Phylloscopidae comprises many small tree-loving warbler species that feed by gleaning insects from leaves or catching food on the wing.[4]
Taxonomy

The genus Phylloscopus was introduced by the German zoologist Friedrich Boie in 1826 to accommodate a single species, the willow warbler, which is therefore considered as the type species.[5][6] The name combines the Ancient Greek phullon meaning "leaf" and skopos meaning "seeker" (from skopeo, "to watch").[7] Phylloscopus is the only genus placed in the family Phylloscopidae that was introduced in 2006 by the Swedish ornithologist Per Alström and coworkers.[8]

Aegithaloidea

Phylloscopidae – leaf warblers (81 species)

Hyliidae – hylias (2 species)

Aegithalidae – bushtits (13 species)

Erythrocercidae – flycatchers (3 species)

Scotocercidae – streaked scrub warbler

Cettiidae – bush warblers and allies (32 species)



Cladogram showing the family relationships based on a study by Carl Oliveros and colleagues published in 2019.[9] The number of species is taken from the bird list maintained by Frank Gill, Pamela Rasmussen and David Donsker on behalf of the International Ornithological Committee (IOC).[10]

The genus contains 81 species:[10] Of these, eleven species were formerly placed in the genus Seicercus, but a 2018 molecular phylogeny study indicated that the genus Seicercus is a synonym of Phylloscopus, leaving the family Phylloscopidae with a single genus, Phylloscopus.[11]

Wood warbler, Phylloscopus sibilatrix
Western Bonelli's warbler, Phylloscopus bonelli
Eastern Bonelli's warbler, Phylloscopus orientalis
Buff-barred warbler, Phylloscopus pulcher
Ashy-throated warbler, Phylloscopus maculipennis
Hume's leaf warbler, Phylloscopus humei
Yellow-browed warbler, Phylloscopus inornatus
Brooks's leaf warbler, Phylloscopus subviridis
Chinese leaf warbler, Phylloscopus yunnanensis
Lemon-rumped warbler, Phylloscopus chloronotus
Sichuan leaf warbler, Phylloscopus forresti
Gansu leaf warbler, Phylloscopus kansuensis
Pallas's leaf warbler, Phylloscopus proregulus
Tytler's leaf warbler, Phylloscopus tytleri
Yellow-streaked warbler, Phylloscopus armandii
Radde's warbler, Phylloscopus schwarzi
Sulphur-bellied warbler, Phylloscopus griseolus
Tickell's leaf warbler, Phylloscopus affinis (includes the Alpine leaf warbler, P. a. occisinensis)
Smoky warbler, Phylloscopus fuligiventer
Dusky warbler, Phylloscopus fuscatus
Plain leaf warbler, Phylloscopus neglectus
Buff-throated warbler, Phylloscopus subaffinis
Willow warbler, Phylloscopus trochilus
Mountain chiffchaff, Phylloscopus sindianus
Canary Islands chiffchaff, Phylloscopus canariensis
Eastern Canary Islands chiffchaff, Phylloscopus canariensis exsul (extinct: 1986?)
Western Canary Islands chiffchaff, Phylloscopus canariensis canariensis
Common chiffchaff, Phylloscopus collybita
Iberian chiffchaff, Phylloscopus brehmii
Eastern crowned warbler, Phylloscopus coronatus
Ijima's leaf warbler, Phylloscopus ijimae
Philippine leaf warbler, Phylloscopus olivaceus
Lemon-throated leaf warbler, Phylloscopus cebuensis
Yellow-throated woodland warbler, Phylloscopus ruficapilla
Brown woodland warbler, Phylloscopus umbrovirens
Red-faced woodland warbler, Phylloscopus laetus
Laura's woodland warbler, Phylloscopus laurae
Black-capped woodland warbler, Phylloscopus herberti
Uganda woodland warbler, Phylloscopus budongoensis
White-spectacled warbler, Phylloscopus intermedius – (previously Seicercus affinis)
Grey-cheeked warbler, Phylloscopus poliogenys – (previously placed in Seicercus)
Green-crowned warbler, Phylloscopus burkii – (previously placed in Seicercus)
Grey-crowned warbler, Phylloscopus tephrocephalus – (previously placed in Seicercus)
Whistler's warbler, Phylloscopus whistleri – (previously placed in Seicercus)
Bianchi's warbler, Phylloscopus valentini – (previously placed in Seicercus)
Alström's warbler, Phylloscopus soror – (first described in 1999; previously placed in Seicercus)
Martens's warbler, Phylloscopus omeiensis – (first described in 1999; previously placed in Seicercus)
Green warbler, Phylloscopus nitidus
Two-barred warbler, Phylloscopus plumbeitarsus
Greenish warbler, Phylloscopus trochiloides
Emei leaf warbler, Phylloscopus emeiensis
Large-billed leaf warbler, Phylloscopus magnirostris
Sakhalin leaf warbler, Phylloscopus borealoides
Pale-legged leaf warbler, Phylloscopus tenellipes
Japanese leaf warbler, Phylloscopus xanthodryas
Kamchatka leaf warbler, Phylloscopus examinandus
Arctic warbler, Phylloscopus borealis
Chestnut-crowned warbler, Phylloscopus castaniceps – (previously placed in Seicercus)
Sunda warbler, Phylloscopus grammiceps – (previously placed in Seicercus)
Yellow-breasted warbler, Phylloscopus montis – (previously placed in Seicercus)
Limestone leaf warbler, Phylloscopus calciatilis – (first described in 2010)
Sulphur-breasted warbler, Phylloscopus ricketti
Yellow-vented warbler, Phylloscopus cantator
Western crowned warbler, Phylloscopus occipitalis
Blyth's leaf warbler, Phylloscopus reguloides
Claudia's leaf warbler, Phylloscopus claudiae
Hartert's leaf warbler, Phylloscopus goodsoni
Kloss's leaf warbler, Phylloscopus ogilviegranti – (formerly considered as a subspecies of Davison's leaf warbler)
Hainan leaf warbler, Phylloscopus hainanus
Davison's leaf warbler, Phylloscopus intensior – (previously white-tailed leaf warbler, Phylloscopus davisoni)
Grey-hooded warbler, Phylloscopus xanthoschistos
Mountain leaf warbler, Phylloscopus trivirgatus
Negros leaf warbler, Phylloscopus nigrorum
Timor leaf warbler, Phylloscopus presbytes
Flores leaf warbler, Phylloscopus floresianus – (formerly considered as a subspecies of the Timor leaf warbler)
Rote leaf warbler, Phylloscopus rotiensis (first described in 2018)[12]
Makira leaf warbler, Phylloscopus makirensis
Sulawesi leaf warbler, Phylloscopus nesophilus – (formerly considered as a subspecies of Lompobattang leaf warbler)[13]
Lompobattang leaf warbler, Phylloscopus sarasinorum[13]
Kolombangara leaf warbler, Phylloscopus amoenus
Island leaf warbler, Phylloscopus poliocephalus
Numfor leaf warbler, Phylloscopus maforensis
Biak leaf warbler, Phylloscopus misoriensis

Two birds were described in 2020 but have not yet been recognised as species by the International Ornithologists' Union.[10][14]

The alpine leaf warbler, Phylloscopus occisinensis, was reclassified as conspecific with Tickell's leaf warbler (P. affinis) by the IOC, but other authorities such as eBird still consider it distinct.[13][15]
References

Baker, Kevin (2010-06-30). Warblers of Europe, Asia and North Africa. A&C Black. p. 17. ISBN 9781408131701.
Dunning, John B. Jr., ed. (2008). CRC Handbook of Avian Body Masses (2nd ed.). CRC Press. ISBN 978-1-4200-6444-5.
Winkler, D. W., S. M. Billerman, and I.J. Lovette (2020). Leaf Warblers (Phylloscopidae), version 1.0. In Birds of the World (S. M. Billerman, B. K. Keeney, P. G. Rodewald, and T. S. Schulenberg, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA.
"Lead-Warblers Phylloscopidae". creagrus.home.montereybay.com. 25 May 2006. Retrieved 2017-09-29.
Mayr, Ernst; Cottrell, G. William, eds. (1986). Check-list of Birds of the World. Volume 11. Cambridge, Massachusetts: Museum of Comparative Zoology. p. 221.
Boie, Friedrich (1826). "Generalübersicht der ornithologischen Ordnungen Familien und Gattugen". Isis von Oken (in German). 19. col. 972.
Jobling, James A. (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. p. 305. ISBN 978-1-4081-2501-4.
Alström, P.; Ericson, P.G.P.; Olsson, U.; Sundberg, P. (2006). "Phylogeny and classification of the avian superfamily Sylvioidea". Molecular Phylogenetics and Evolution. 38 (2): 381–397. doi:10.1016/j.ympev.2005.05.015. PMID 16054402.
Oliveros, C.H.; et al. (2019). "Earth history and the passerine superradiation". Proceedings of the National Academy of Sciences of the United States of America. 116 (16): 7916–7925. doi:10.1073/pnas.1813206116. PMC 6475423. PMID 30936315.
Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (January 2022). "Bushtits, leaf warblers, reed warblers". IOC World Bird List Version 12.1. International Ornithologists' Union. Retrieved 7 June 2022.
Alström, P.; Rheindt, F.E.; Zhang, R.; Zhao, M.; Wang, J.; Zhu, X.; Gwee, C.Y.; Hao, Y.; Ohlson, J.; Jia, C.; Prawiradilaga, D.M.; Ericson, P.G.P.; Lei, F.; Olsson, U. (2018). "Complete species-level phylogeny of the leaf warbler (Aves: Phylloscopidae) radiation" (PDF). Molecular Phylogenetics and Evolution. 126: 141–152. doi:10.1016/j.ympev.2018.03.031.
Ng, Nathaniel. S. R.; Prawiradilaga, Dewi. M.; Ng, Elize. Y. X.; Suparno; Ashari, Hidayat; Trainor, Colin; Verbelen, Philippe; Rheindt, Frank. E. (2018). "A striking new species of leaf warbler from the Lesser Sundas as uncovered through morphology and genomics". Scientific Reports. 8 (1): 15646. doi:10.1038/s41598-018-34101-7.
"Species Updates – IOC World Bird List". Retrieved 2021-05-28.
Rheindt, F.E.; Prawiradilaga, D.M.; Ashari, H.; Suparno; Gwee, C.Y.; Lee, G.W.X.; Wu, M.Y.; Ng, N.S.R. (2020). "A lost world in Wallacea: description of a montane archipelagic avifauna". Science. 367 (6474): 167–170. doi:10.1126/science.aax2146. See supplement.

"Alpine Leaf Warbler - eBird". ebird.org. Retrieved 2021-06-18.

Further reading

Badyaev, Alexander V. & Leaf, Elizabeth S. (1997): "Habitat associations of song characteristics in Phylloscopus and Hippolais warblers". Auk 114(1): 40–46.

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