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The common chiffchaff (Phylloscopus collybita), or simply the chiffchaff, is a common and widespread leaf warbler which breeds in open woodlands throughout northern and temperate Europe and the Palearctic.
It is a migratory passerine which winters in southern and western Europe, southern Asia and north Africa. Greenish-brown above and off-white below, it is named onomatopoeically for its simple chiff-chaff song. It has a number of subspecies, some of which are now treated as full species. The female builds a domed nest on or near the ground, and assumes most of the responsibility for brooding and feeding the chicks, whilst the male has little involvement in nesting, but defends his territory against rivals, and attacks potential predators.
A small insectivorous bird, it is subject to predation by mammals, such as cats and mustelids, and birds, particularly hawks of the genus Accipiter. Its large range and population mean that its status is secure, although one subspecies is probably extinct.
TaxonomyEdit
The British naturalist Gilbert White was one of the first people to separate the similar-looking common chiffchaff, willow warbler and wood warbler by their songs, as detailed in 1789 in The Natural History and Antiquities of Selborne,<ref name= white>Template:Cite book</ref> but the common chiffchaff was first formally described as Sylvia collybita by French ornithologist Louis Pierre Vieillot in 1817 in his Nouveau Dictionnaire d'Histoire Naturelle.<ref name= viellot>Template:Cite book</ref> The type locality is the French region of Normandy.<ref>Template:Cite book</ref>
Described by German zoologist Friedrich Boie in 1826,<ref>Template:Cite journal</ref> the genus Phylloscopus contains about 80 species of small insectivorous Old World woodland warblers which are either greenish or brown above and yellowish, white or buff below. The genus was formerly part of the Old World warbler family Sylvidae, but has now been split off as a separate family Phylloscopidae.<ref name=alstrom>Template:Cite journal</ref> The chiffchaff's closest relatives, other than former subspecies, are a group of leaf warblers which similarly lack crown stripes, a yellow rump or obvious wing bars; they include the willow, Bonelli's, wood and plain leaf warblers.<ref name= Baker/>
An old synonym, used for the chiffchaff was Phylloscopus rufus (Bechstein).<ref name=rufus>For instance in: Template:Cite encyclopedia</ref>
The common 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.<ref name= ClementHelbig1998>Template:Cite journal</ref><ref name = Sangster2002>Template:Cite journal</ref>
SubspeciesEdit
- P. c. collybita, the nominate form, breeds in Europe east to Poland and Bulgaria, and is described below. It mainly winters in the south of its breeding range around the Mediterranean and in North Africa.<ref name= Baker>Template:Cite book</ref> It has been expanding its range northwards into Scandinavia since 1970 and close to the southern edge of the range of P. c. abietinus.<ref name=Hanson>Template:Cite journal</ref>
- 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 P. c. tristis and P. c. collybita, being grey-washed olive-green above with a pale yellow supercilium, and underparts whiter than in P. c. collybita,Template:Sfn but it has very similar vocalisations to the nominate subspecies.<ref name= Baker/> Due to individual variation, it can be difficult to reliably separate P. c. abietinus and P. c. collybita outside their main breeding and wintering ranges.Template:Sfn Some common chiffchaffs in the Middle East are browner and have a more disyllabic swee-hu call than P. c. abietinus, and may belong to a poorly known taxon "brevirostris";<ref name= dubois2008>Template:Cite journal</ref> further research is needed to clarify the affinities of this form.<ref name= dubois2010>Template:Cite journal</ref>
- P. (c.) tristis, the Siberian chiffchaff, breeds in Siberia east of the Pechora River and winters in the lower Himalayas.<ref name=Baker/> It is also regularly recorded in western Europe in winter, and it is likely that the numbers involved have been underestimated due to uncertainties over identification criteria, lack of good data and recording policies (Sweden and Finland only accept trapped birds).<ref name= dean8>Template:Cite journal</ref> It is a dull subspecies, grey or brownish above and whitish below, with little yellow in the plumage, and the buff-white supercilium is often longer than in the western subspecies. It has a higher pitched suitsistsuisit song and a short high-pitched cheet call.Template:Sfn 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.<ref name = Martens1983/><ref name= Helbig1996>Template:Cite journal</ref> Nominate P. c. collybita and P. c. tristis do not recognize each other's songs.<ref name=Schubert>Template:Cite journal</ref><ref name = MartensMeincke1989>Template:Cite journal</ref> Pending resolution of the status of P. (c.) fulvescens, which is found where the ranges of P. c. abietinus and P. c. tristis connect and may<ref name= Marova>Template:Cite journal</ref> or may not<ref name = MartensMeincke1989/> be a hybrid between these, tristis is maintained in P. collybita.<ref name = Sangster2002/>
Former subspeciesEdit
- P. ibericus, the Iberian chiffchaff is brighter, greener on the rump, and yellower below than P. collybita,<ref name= Baker/> and has a Template:Not a typo song. It was initially named P. brehmii, but the type specimen of that taxon is not an Iberian chiffchaff.<ref name= Iberian>Template:Cite journal</ref> This species is found in Portugal and Spain, west of a line stretching roughly from the western Pyrenees<ref name =Salomon1989>Template:Cite journal</ref> via the mountains of central Spain to the Mediterranean; the Iberian and common chiffchaffs co-occur in a narrow band along this line.<ref name=Balmori>Template:Cite journal</ref> 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. c. collybita in vocalisations,<ref name=Helbig1996/><ref name =Salomon1989/><ref name = SalomonHemim>Template:Cite journal</ref> external morphology,<ref name=Salomon1997>Template:Cite journal</ref> and mtDNA sequences.<ref name=Helbig1996/><ref name = Helbig2001>Template:Cite journal</ref> There is hybridization in the contact zone,<ref name =Salomon1989/><ref name = SalomonHemim/><ref name= BB101/> almost always between male P. ibericus and female P. c. collybita,<ref name= BB101>Template:Cite journal</ref> and hybrids apparently show much decreased fitness;<ref name = Helbig2001/> hybrid females appear to be sterile according to Haldane's Rule.<ref name = Helbig1993>Template:Cite journal</ref> Regarding the latter aspect, the Iberian chiffchaff apparently is the oldest lineage of chiffchaffs and quite distinct from the common chiffchaff.<ref name=Helbig1996/>
- P. canariensis, the Canary Islands chiffchaff is a non-migratory species formerly occurring on the major Canary Islands, which is differentiated from P. collybita by morphology, vocalisations and genetic characteristics, and, of course, is not sympatric with any other chiffchaffs. The nominate western subspecies P. c. canariensis of El Hierro, La Palma, La Gomera, Tenerife, and Gran Canaria is smaller than common chiffchaff, and has shorter, rounder wings.<ref name=Helbig1996/> It is olive-brown above and has a buff breast and flanks;<ref name= Baker/> it has a rich deep Template:Not a typo song, and a call similar to the nominate race.Template:Sfn The eastern P. c. exsul of Lanzarote and possibly Fuerteventura is paler above and less rufous below than its western relative,<ref name= Baker/> and had a harsher call;Template:Sfn it might have been a distinct species,<ref name = Sangster2002/> but it became extinct in 1986 at latest, probably much earlier. The reasons for its extinction are unclear, but it appears always to have been scarce and localised, occurring only in the Haria Valley of Lanzarote.<ref name= Simms >Template:Cite book</ref>
- 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 P. c. tristis, but with a finer darker bill, browner upperparts and buff flanks; its song is almost identical to P. collybita, but the call is a weak Template:Not a typo. P. s. lorenzii is warmer and darker brown than the nominate race;<ref name=Baker/> it is sympatric with common chiffchaff in a small area in the Western Caucasus, but interbreeding occurs rarely, if ever.<ref name=Martens1983/> The mountain chiffchaff differs from tristis in vocalisations,<ref name=Martens1983>Template:Cite journal</ref><ref name="MartensHänel">Template:Cite journal</ref> external morphology,<ref name=Cramp1992>Template:Cite book</ref> and mtDNA sequences.<ref name=Helbig1996/> Its two subspecies appear to be distinct vocally,<ref name=Martens1983/> and also show some difference in mtDNA sequences;<ref name=Helbig1996/> they are maintained at subspecies rank pending further research.<ref name = Sangster2002/>
EtymologyEdit
The common chiffchaff's English name is onomatopoeic, referring to the repetitive Template:Not a typo song of the European subspecies.<ref name= Cocker>Template:Cite book</ref> There are similar names in some other European languages, such as the Dutch {{#invoke:Lang|lang}}, the German {{#invoke:Lang|lang}}, Welsh {{#invoke:Lang|lang}} and Finnish {{#invoke:Lang|lang}}.<ref>Tiltaltti in Glosbe.</ref> The binomial name is of Greek origin; Phylloscopus comes from Template:Transliteration/{{#invoke:Lang|lang}} "leaf", and Template:Transliteration/{{#invoke:Lang|lang}} "to look at" or "to see",<ref name= Terres >Template:Cite book</ref> since this genus comprises species that spend much of their time feeding in trees, while collybita is a corruption of κολλυβιστής (kollubistḗs) "money changer", the song being likened to the jingling of coins.<ref name= Cocker/> In some languages their tree-dwelling habit is hinted in the vernacular name. For example, in Swedish the common chiffchaff is called gransångare, a compound of gran (i.e. "spruce") and sångare, meaning both "singer" and Old World warbler.
DescriptionEdit
The common chiffchaff is a small, dumpy, Template:Convert long leaf warbler. The male weighs 7–8 grammes (0.28–0.31 oz), and the female 6–7 grammes (0.25–0.28 oz).Template:Sfn The spring adult of the western nominate subspecies P. c. 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 (extension of the flight feathers beyond the folded wing).Template:Sfn 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 before migration. 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.Template:Sfn
P. c. collybita in Newcastle upon Tyne
This warbler gets its name from its simple distinctive song, a repetitive cheerful Template:Not a typo. This song is one of the first avian signs that spring has returned. Its call is a Template:Not a typo, less disyllabic than the Template:Not a typo of the willow warbler or Template:Not a typo of the western Bonelli's warbler.<ref name= Collins>Mullarney, Killian; Svensson, Lars, Zetterstrom, Dan; Grant, Peter. (1999). Birds of Europe. London. HarperCollins. pp. 304–306 Template:ISBN</ref>
The song differs from that of the Iberian chiffchaff, which has a shorter Template:Not a typo. However, mixed singers occur in the hybridisation zone and elsewhere, and can be difficult to allocate to species.<ref name = BB101/>
When not singing, the common chiffchaff can be difficult to distinguish from other leaf warblers with greenish upperparts and whitish underparts, particularly the willow warbler. However, that species has a longer primary projection, a sleeker, brighter appearance and generally pale legs. Bonelli's warbler (P. bonelli) might be confused with the common chiffchaff subspecies tristis, but it has a plain face and green in the wings.Template:Sfn The common chiffchaff also has rounded wings in flight, and a diagnostic tail movement consisting of a dip, then sidewards wag, that distinguishes it from other Phylloscopus warblers<ref name= BWP/> and gives rise to the name "tailwagger" in India.<ref name=Simms/>
Perhaps the greatest challenge is distinguishing non-singing birds of the nominate subspecies from Iberian chiffchaff in the field. In Great Britain and the Netherlands, all accepted records of vagrant Iberian chiffchaffs relate to singing males.<ref name = BB101/>
Distribution and habitatEdit
The common chiffchaff breeds across Europe and Asia east to eastern Siberia and north to about 70°N, with isolated populations in northwest Africa, northern and western Turkey and northwestern Iran.Template:Sfn It is migratory, but it is one of the first passerine birds to return to its breeding areas in the spring and among the last to leave in late autumn.<ref name = Collins/><ref name =BWP/> When breeding, it is a bird of open woodlands with some taller trees and ground cover for nesting purposes. These trees are typically at least Template:Convert 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 (P. trochilus) prefers younger trees, while the wood warbler (P. sibilatrix) prefers less undergrowth.Template:Sfn In winter, the common chiffchaff uses a wider range of habitats including scrub, and is not so dependent on trees. It is often found near water, unlike the willow warbler which tolerates drier habitats.Template:Sfn There is an increasing tendency to winter in western Europe well north of the traditional areas, especially in coastal southern England and the mild urban microclimate of London.Template:Sfn These overwintering common chiffchaffs include some visitors of the eastern subspecies abietinus and tristis, so they are certainly not all birds which have bred locally, although some undoubtedly are.<ref name =BWP/>
BehaviourEdit
TerritoryEdit
The male common chiffchaff is highly territorial during the breeding season, with a core territory typically Template:Convert across, which is fiercely defended against other males. Other small birds may also be attacked.Template:Sfn 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.Template:Sfn His 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.<ref name= Rodrigues >Template:Cite journal</ref>
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.Template:Sfn After breeding has finished, this species abandons its territory, and may join small flocks including other warblers prior to migration.<ref name=BWP/>
BreedingEdit
The male common chiffchaff returns to its breeding territory two or three weeks before the female and immediately starts singing to establish ownership and attract a female. When a female is located, the male will use a slow butterfly-like flight as part of the courtship ritual, but once a pair-bond has been established, other females will be driven from the territory. The male has little involvement in the nesting process other than defending the territory.Template:Sfn The female's nest is built on or near the ground in a concealed site in brambles, nettles or other dense low vegetation. The domed nest has a side entrance, and is constructed from coarse plant material such as dead leaves and grass, with finer material used on the interior before the addition of a lining of feathers. The typical nest is Template:Convert high and Template:Convert across.Template:Sfn
The clutch is two to seven (normally five or six) cream-coloured eggs which have tiny ruddy, purple or blackish spots and are about Template:Convert long and Template:Convert across. They are incubated by the female for 13–14 days before hatching as naked, blind altricial chicks.Template:Sfn The female broods and feeds the chicks for another 14–15 days until they fledge. The male rarely participates in feeding, although this sometimes occurs, especially when bad weather limits insect supplies or if the female disappears. After fledging, the young stay in the vicinity of the nest for three to four weeks, and are fed by and roost with the female, although these interactions reduce after approximately the first 14 days.Template:Sfn In the north of the range there is only time to raise one brood, due to the short summer, but a second brood is common in central and southern areas.Template:Sfn
Although pairs stay together during the breeding season and polygamy is uncommon, even if the male and female return to the same site in the following year there is no apparent recognition or fidelity. Interbreeding with other species, other than those formerly considered as subspecies of P. collybita, is rare, but a few examples are known of hybridisation with the willow warbler. Such hybrids give mixed songs, but the latter alone is not proof of interspecific breeding.Template:Sfn
FeedingEdit
Like most Old World warblers, this small species is insectivorous, moving restlessly through 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.Template:Sfn The chiffchaff has been estimated to require about one-third of its weight in insects daily, and it feeds almost continuously in the autumn to put on extra fat as fuel for the long migration flight.Template:Sfn
Predators and threatsEdit
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. 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.Template:Sfn
The common chiffchaff is occasionally a host of brood parasitic cuckoos, including the common and Horsfield's cuckoos,<ref name= Johnsgard >Template:Cite book</ref> but it recognises and rejects non-mimetic eggs and is therefore only rarely successfully brood-parasitised.<ref name= Moksne>Template:Cite journal</ref> Like other passerine birds, the common chiffchaff can also acquire intestinal nematode parasites and external ticks.<ref name = Cork>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name= Jaenson >Template:Cite journal</ref>
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 P. c. abietinus and P. c. tristis, and woodland conservation policies in the range of P. c. collybita, the chiffchaff's future seems assured.Template:Sfn
StatusEdit
The common 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".<ref name=IUCN/>
None of the major subspecies is under threat, but exsul, as noted above, is probably extinct. There is a slow population increase of common chiffchaff in the Czech Republic.<ref>Template:In lang Budníček menší (Phylloscopus collybita). Česká společnost ornitologická (Czech Society for Ornithology), accessed 20 April 2009</ref> The range of at least P. c. collybita seems to be expanding, with northward advances in Scotland, Norway and Sweden and a large population increase in Denmark.<ref name=BWP>Template:Cite book</ref>
ReferencesEdit
SourcesEdit
External linksEdit
- Template:Wikispecies-inline
- Template:Commons-inline
- Template:Wiktionary-inline
- BBC Science and Nature: BBC chiffchaff site
- BBC Science and Nature: Common chiffchaff song (Real Audio streaming)
- Common chiffchaff videos, photos & sounds on the Internet Bird Collection
- RSPB: Chiffchaff, Phylloscopus collybita
- Ageing and sexing (PDF; 3.5 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze