Chiffchaff
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| Chiffchaff | ||||||||||||||
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Subspecies tristis in India
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| Phylloscopus collybita (Vieillot, 1817) |
The Common Chiffchaff or simply Chiffchaff, (Phylloscopus collybita), is a common and widespread leaf warbler which breeds in open woodlands throughout northern and temperate Europe and Asia. It is probably the best-known of all Old World "warbler"s.
It is a small insectivorous bird, greenish above and off-white below, named for its simple chiff-chaff song. It has a number of subspecies, some of which are now normally treated as full species.
This bird is less strongly migratory and hardier than most leaf warblers; northern breeders winter in southern and western Europe (north to Britain), southern Asia and north Africa. Along with the Sand Martin, it is the first passerine to return in the spring, and one of the last to leave in late autumn.
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The Chiffchaff’s English name is onomatopoeic, referring to the repetitive chiff-chaff song of the European subspecies.[2] The binomial name is of Greek origin; Phylloscopus comes from phyllon, "leaf", and skopeo, "to look at" or "to see",[3] since this genus comprises species that spend much of their time feeding in trees, and collybita is a corruption of kollubistes, "money changer", the song being likened to the jingling of coins.[2]
The Chiffchaff breeds across Europe and Asia east to eastern Siberia and north to about 70oN, with isolated populations in northwest Africa, northern and western Turkey and northwestern Iran.[4]
It is a bird of open woodlands with some taller trees and ground cover for nesting. The trees should be at least 5 metres (16 ft) high, with undergrowth that is an open, poor to medium mix of grasses, bracken, nettles or similar plants. Its breeding habitat is quite specific, and even near relatives do not share it; for example, the Willow Warbler prefers younger trees, and the Wood Warbler less undergrowth.[4]
In winter, the Chiffchaff uses a wider range of habitats including scrub, and is not dependent on for trees. It is often found near water, unlike the Willow Warbler which tolerates drier habitats. There is an increasing tendency in western Europe to winter well north of the traditional areas, especially in coastal southern England, and the mild urban microclimate of London.[4] These birds include some eastern visitors of the subspecies abietinus and tristis, so they are certainly not all locally breeding birds staying to overwinter, although some undoubtedly are.[5]
The Chiffchaff is a small, dumpy, 10-12 centimetres (4 in) long, leaf warbler. The spring adult of the western nominate subspecies collybita has brown-washed dull green upperparts, off-white underparts becoming yellowish on the flanks, and a short whitish supercilium. It has dark legs, a fine dark bill, and short primary projection. As the plumage wears, it gets duller and browner, and the yellow on the flanks tends to be lost, but after the breeding season there is a prolonged complete moult prior to migration.[4] It has rounded wings in flight.[5]
The newly-fledged juvenile is browner above than the adult, with yellow-white underparts, but moults about 10 weeks after acquiring its first plumage. After moulting both the adult and the juvenile have brighter and greener upperparts, and a paler supercilium.[4]
This warbler gets its name from its simple distinctive song, a repetitive cheerful chiff-chaff. This song is one of the first avian signs that spring has returned. Its call is a hweet, less disyllabic than the hooeet of Willow Warbler or hu-it of Western Bonelli's Warbler.[6]
When not singing, the Chiffchaff can be confused with other leaf warblers, which in general have greenish upperparts and whitish underparts. The most likely confusion species are the Willow Warbler, Phylloscopus trochilus and Bonelli's Warbler, P. bonelli.[4] The Willow Warbler has a longer primary projection, usually pale legs, and a sleeker, brighter appearance. Bonelli's Warbler might be confused with tristis Chiffchaff, but has a very plain face and green in the wings.[4]
The Chiffchaff has a diagnostic tail movement consisting of a dip, then sidewards wag, that differentiates it from other Phylloscopus warblers.[5]
The British naturalist Gilbert White was one of the first people to separate the similar-looking Chiffchaff, Willow Warbler and Wood Warbler by their songs, as detailed in The Natural History and Antiquities of Selborne,[7]but the Common Chiffchaff was first formally described as Sylvia collybita by French ornithologist Louis Viellot in 1817 in his Nouvelle Dictionnaire d'Histoire Naturelle.[8]
The Chiffchaff has three still commonly accepted subspecies, together with some from the Iberian Peninsula, the Canary Islands, and the Caucasus which are now more often treated as full species.[9][10]
- P. c. collybita, the nominate form, breeds in Europe east to Poland and Bulgaria, and is described above. It mainly winters in the south of its breeding range around the Mediterranean, and in North Africa.[11]
- P. c. abietinus occurs in Scandinavia and northern Russia, and winters from southeastern Europe and northeastern Africa east to Iraq and western Iran. It is intermediate in appearance between tristis and collybita, but has very similar vocalisations to the nominate subspecies.[11]
- P. (c.) tristis, the Siberian Chiffchaff, breeds in Siberia east of the Pechora River and winters in the lower Himalayas.[11] It is a dull subspecies, grey or brownish above and whitish below, with little yellow in the plumage. It has a higher pitched suitsistsuisit song and a short high-pitched cheet call.[4]
- It is sometimes considered to be a full species due to its distinctive plumage and vocalisations, being similar to P. s. sindianus in these respects.[12][13] Nominate collybita and tristis do not recognize each other's songs.[14][15] Pending resolution of the status of P. (c.) fulvescens which is found where the ranges of abietinus and tristis connect and may [16] or may not[15] be a hybrid between these, tristis is maintained in P. collybita.[10]
- P. ibericus, the Iberian Chiffchaff is brighter, greener on the rump, and yellower below than P. collybita,[11] and has a tit-tit-tit-tswee-tswee song. It was initially named P. brehmii, but the type specimen of that taxon is not an Iberian Chiffchaff.[17] This species is found in Portugal and Spain, west of a line stretching roughly from the western Pyrenees[18] via the mountains of central Spain to the Mediterranean; the Iberian and Common Chiffchaffs co-occur in a narrow band along this line.[19] Apart from the northernmost section, the precise course of the contact zone is not well-documented. A long-distance migrant, this species winters in western Africa.
- It differs from P.collybita in vocalizations,[18][20][13] external morphology,[21] and mtDNA sequences.[22][13] There is little hybridization in the contact zone,[18][20] and hybrids apparently show much decreased fitness;[22] hybrid females appear to be sterile according to Haldane's Rule.[23] Regarding the latter aspect, it is interesting to note that the Iberian Chiffchaff apparently is the oldest lineage of chiffchaffs and quite distinct from the Common Chiffchaff.[13]
- P. canariensis, the Canary Islands Chiffchaff is a non-migratory species formerly occurring on the major Canary Islands. The nominate western subspecies P. c. canariensis of El Hierro, La Palma, La Gomera, Tenerife, and Gran Canaria is olive-brown above and has a buff breast and flanks.[11] The eastern P. c. exsul of Lanzarote and possibly Fuerteventura is paler above and less rufous below than its western relative,[11] and might have been a distinct species,[10] but it became extinct in 1986 at latest, probably much earlier. P. canariensis is differentiated from P. collybita by morphological, bioacoustical and genetical characteristics,[13] and of course is not sympatric with any other chiffchaffs.
- P. sindianus, the Mountain Chiffchaff, is found in the Caucasus (P. s. lorenzii) and Himalayas (P. s. sindianus), and is an altitudinal migrant, moving to lower levels in winter. The nominate subspecies is similar to tristis Chiffchaff, but with a finer darker bill, browner upperparts and buff flanks. P. s. lorentzii is warmer and darker brown than the nominate race.[11]; it is sympatric with Common Chiffchaff in a small area in the Western Caucasus, but interbreeding occurs rarely if ever.[12]
- The Mountain Chiffchaff differs from Chiffchaff in vocalizations,[24][12] external morphology,[25] and mtDNA sequences.[13] Its two subspecies appear to be distinct vocally,[12] and also show some difference in mtDNA sequences;[13] they are maintained at subspecies rank pending further research.[10]
The male Chiffchaff is highly territorial during the breeding season, with a core territory typically 20 metres (66 ft) across, which is fiercely defended against other males. Other small birds may also be attacked. The male is inquisitive and fearless, attacking even dangerous predators like the Stoat if they approach the nest, as well as egg-thieves like the Eurasian Jay.[4]
The male’s song, given from a favoured prominent vantage point, appears to be used to advertise an established territory and contact the female, rather than as a paternity guard strategy.[26]
Beyond the core territory, there is a larger feeding range which is variable in size, but typically ten or more times the area of the breeding territory. it is believed that the female has a larger feeding range than the male.[4] After breeding has finished, the Chiffchaff abandons its territory, and may join small flocks including other warblers prior to migration.[5]
The Chiffchaff's nest is built in low shrub, and the four to seven eggs are incubated by the female for 13-14 days to hatching, with another 14-15 days until the chicks fledge.
Like most Old World warblers, this small species is insectivorous, moving restlessly though foliage or briefly hovering. It has been recorded as taking insects, mainly flies from more than 50 families, along with other small and medium-sized invertebrates. It will take the eggs and larvae of butterflies and moths, particularly those of the Winter Moth.[4]
The male Chiffchaff weighs 7-8 grammes (0.28-0.31 oz), and the female 6-7 grammes (0.25-0.28 oz). it has been estimated that this species needs to consume about one-third of its weight in insects daily. In the autumn, the Chiffchaff feeds almost continuously to put on extra fat as fuel to prepare for the rigours of the long migration flight.[4]
As with most small birds, mortality in the first year of life is high, but adults aged three to four years are regularly recorded, and the record is more than seven years.[4]
Eggs, chicks and fledglings of this ground-nesting species are taken by stoats, weasels and crows such as the European Magpie, and the adults are hunted by birds of prey, particularly the Sparrowhawk. Small birds are also at the mercy of the weather, particularly when migrating, but also on the breeding and wintering grounds.[4]
The Chiffchaff is occasionally a host of brood parasitic cuckoos, including the Common and Horsfield's Cuckoos.[27] The Chiffchaff recognises and rejects non-mimetic eggs and is therefore only rarely successfully parasitised. [28]
The main effect of humans on this species is indirect, through woodland clearance which affects the habitat, predation by cats, and collisions with windows, buildings and cars. Only the first of these has the potential to seriously affect populations, but given the huge geographical spread of abietinus and tristis, and conservation policies in the range of collybita, the Chiffchaff's future seems assured.[4]
The Chiffchaff has an enormous range, with an estimated global extent of 10 million square kilometres (3.8 million square miles) and a population of 60-120 million individuals in Europe alone. Although global population trends have not been quantified, the species is not believed to approach the thresholds for the population decline criterion of the IUCN Red List (that is, declining more than 30 percent in ten years or three generations). For these reasons, the species is evaluated as "least concern".[1]
None of the major subspecies is under threat, but exsul, as noted above, is probably extinct. The range of at least collybita seems to be expanding, with northward advances in Scotland, Norway and Sweden and a large population increase in Denmark.[5]
- ^ a b BirdLife International (2004). Phylloscopus collybita. 2006 IUCN Red List of Threatened Species. IUCN 2006. Retrieved on 12 May 2006. Database entry includes justification for why this species is of least concern
- ^ a b Cocker, Mark; Mabey, Richard (2005). Birds Britannica. London: Chatto & Windus, 378-9. ISBN 0-7011-6907-9.
- ^ Terres, John K. (1980). The Audubon Society encyclopedia of North American birds. New York: Alfred A. Knopf, Inc., 1001. ISBN 0517032880.
- ^ a b c d e f g h i j k l m n o Clement, Peter (1995). The Chiffchaff. London: Hamlyn. ISBN 0600579786.
- ^ a b c d e Snow, David; Perrins, Christopher M (editors) (1998). The Birds of the Western Palearctic concise edition (2 volumes). Oxford: Oxford University Press. ISBN 0-19-854099-X. p1337-1339
- ^ Mullarney, Killian; Svensson, Lars, Zetterstrom, Dan; Grant, Peter. (1999). Birds of Europe. London. HarperCollins. p 304-306 ISBN 0002197286
- ^ White, Gilbert [1789] (1887). The Natural History and Antiquities of Selborne. London: Cassell & Company, pp. 38-39. OCLC 3423785.
- ^ (French) Viellot, Louis Jean Pierre (1817): Nouvelle Dictionnaire d'Histoire Naturelle nouvelle édition, 11, 235.
- ^ Clement, P. & Helbig, Andreas J. (1998): Taxonomy and identification of chiffchaffs in the Western Palearctic. Brit. Birds 91: 361–376.
- ^ a b c d Sangster, George; Knox, Alan G.; Helbig, Andreas J. & Parkin, David T. (2002):Taxonomic recommendations for European birds. Ibis 144(1): 153–159. doi:10.1046/j.0019-1019.2001.00026.x ]
- ^ a b c d e f g Baker, Kevin (1997). Warblers of Europe, Asia and North Africa (Helm Identification Guides). ISBN 0713639717.
- ^ a b c d (German) Martens, Jochen (1982): Ringförmige Arealüberschneidung und Artbildung beim Zilpzalp, Phylloscopus collybita. Das lorenzii-Problem. Zeitschrift für Zoologische Systematik und Evolutionsforschung 20: 82–100.
- ^ a b c d e f g Helbig, Andreas J.; Martens, Jochen; Seibold, I.; Henning, F.; Schottler, B; Wink, Michael (1996): Phylogeny and species limits in the Palearctic Chiffchaff Phylloscopus collybita complex: mitochondrial genetic differentiation and bioacoustic evidence. Ibis 138(4): 650–666
- ^ (German) Schubert, M. (1982): Zur Lautgebung mehrerer zentralasiatischer Laubsänger-Arten (Phylloscopus; Aves, Sylviidae). Mitteilungen aus dem Zoologischen Museum Berlin 58: 109–128.
- ^ a b (German) Martens, Jochen; Meincke, C. (1989): Der sibirische Zilpzalp (Phylloscopus collybita tristis): Gesang und Reaktion einer mitteleuropäischen Population im Freilandversuch. Journal für Ornithologie 130(4): 455–473. [with English abstract] doi:10.1007/BF01918465
- ^ (Russian) Marova, I. M. & Leonovich, V. V. (1993) [Hybridization between Siberian (Phylloscopus collybita tristis) and East European (Ph. collybita abietinus) Chiffchaffs in the area of sympatry.] Sbornik Trudov Zoologicheskogo Muzeya, Moskovskogo Gosudarstvennogo Universiteta 30: 147–163.
- ^ Svensson, Lars (2001). "The correct name of the Iberian Chiffchaff Phylloscopus ibericus Ticehurst 1937, its identification and new evidence of its winter grounds". Bulletin of the British Ornithologists' Club 121: 281-296.
- ^ a b c Salomon, Marc (1989): Song as a possible reproductive isolating mechanism between two parapatric forms. The case of the chiffchaffs Phylloscopus c. collybita and P. c. brehmii in the western Pyrenees. Behaviour 111(1-4): 270–290.
- ^ (Spanish) Balmori, Alfonso; Cuesta, Miguel Ángel; Caballero, José María (2002): Distribución de los mosquiteros ibérico (Phylloscopus brehmii) y europeo (Phylloscopus collybita) en los bosques de ribera de Castilla y León (España). [With English abstract]. Ardeola 49(1): 19-27.
- ^ a b Salomon, Marc; Hemim, Y. (1992):Song variation in the Chiffchaffs (Phylloscopus collybita) of the western Pyrenees – the contact zone between collybita and brehmii forms. Ethology 92(4): 265–282.
- ^ Salomon, Marc; Bried, J.; Helbig, Andreas J.; Riofrio, J. (1997) Morphometric differentiation between male Common Chiffchaffs, Phylloscopus [c.] collybita Vieillot, 1817, and Iberian Chiffchaffs, P. [c.] brehmii Homeyer, 1871, in a secondary contact zone (Aves: Sylviidae). Zoologischer Anzeiger 236: 25–36.
- ^ a b Helbig, Andreas J.; Salomon, Marc; Bensch, S. & Seibold, I. (2001):Male-biased gene flow across an avian hybrid zone: evidence from mitochondrial and microsatellite DNA. Journal of Evolutionary Biology 14: 277–287. doi:10.1046/j.1420-9101.2001.00273.x
- ^ Helbig, Andreas J.; Salomon, Marc; Wink, Michael; Bried, Joël (1993): Absence de flux genique mitochondrial entre le Pouillots "veloces" medio-européen et ibérique (Aves: Phylloscopus collybita, P. (c.) brehmii); implications taxonomiques. Résultats tirés de la PCR et du séquencage d'ADN. C. R. Acad. Sci. III 316: 205-210.
- ^ (German) Martens, Jochen; Hänel, Sabine (1981): Gesangformen und Verwandtschaft der asiatischen Zilpzalpe Phylloscopus collybita abietinus und Ph. c. sindianus. Journal für Ornithologie 122(4): 403–427. [with English abstract]doi:10.1007/BF01652928
- ^ Cramp, Stanley. (ed.) (1992): The Birds of the Western Palearctic, Vol. 6. Oxford University Press, Oxford.
- ^ Rodrigues, Marcos (March 1996). "Song activity in the chiffchaff: territorial defence or mate guarding?". Animal Behaviour 51 (3): 709-716.
- ^ Johnsgard, Paul A. (1997). The Avian Brood Parasites: Deception at the Nest. Oxford University Press, 196. ISBN 0195110420.
- ^ Moksnes, Arne; Roskaft, Eivin (Jan. - Mar 1992). "Responses of Some Rare Cuckoo Hosts to Mimetic Model Cuckoo Eggs and to Foreign Conspecific Eggs". Ornis Scandinavica 23 (1): 17-23. doi:10.2307/3676422.
- BBC Science and Nature: BBC Chiffchaff site
- BBC Science and Nature: Common Chiffchaff song (Real Audio streaming)
- Internet Bird Collection: Common Chiffchaff videos
- RSPB: Chiffchaff, Phylloscopus collybita