Until recently considered conspecific with T. hiemalis and T. pacificus; degree of differentiation from these taxa still in need of study and clarification, as the forms are seemingly undiagnosable except by song (although degree of vocal difference across full ranges of present species and T. pacificus has not been researched). Recent DNA work suggests that species as here constituted consists of four distinct clades, in Europe, Caucasus, Nepal and E Asia, respectively, which has been interpreted as cryptic speciation. Many races doubtfully valid, e.g. zagrossiensis sometimes subsumed within hyrcanus; proposed race †orii (Daito Is, in Japan) is included here in fumigatus. Twenty-nine subspecies recognized.
9–10 cm; 6–12 g. Nominate race has pale buff supercilium, brown lores and ear coverts spotted off-white; crown dark brown, back warmer brown, rump more rufous-brown; shoulders and upperwing coverts rufous-brown with darker barring; primaries and secondaries brown, barred darker; rectrices warm brown with dark transverse bars; chin and throat pale buff-brown, centre of throat sometimes whiter; chest and belly warmer rufous-brown, central belly uniform buff-white, flanks darker rufous-brown, blackish barring on lower flanks; eye brown; bill brown, pale base; legs light brown. Sexes similar. Juvenile has indistinctly mottled breast, more obscure barring on flanks. Races differ mainly in size, general coloration, and extent of barring: islandicus is substantially larger than nominate, bill heavier, generally darker brown, barring more pronounced, especially below; indigenus is darker, duller and less rufous than nominate; hirtensis is similar to previous but larger, more grey-brown above, pale and well barred below; zetlandicus is much darker above and below than previous, more rufous-brown; fridariensis resembles last but paler above and below; borealis is also paler, and more heavily barred; hebridensis is more buffy, less heavily barred and with less stout bill; koenigi is darker, more earth-brown and less rufous, than nominate; kabylorum is paler above and below than nominate; juniperi is like previous, but bill longer, feet dark-coloured; cypriotes is more extensively barred below than last; hyrcanus is more greyish brown than nominate, with pale throat, more extensively barred underparts; zagrossiensis resembles previous but paler; tianschanicus is also paler and greyer, with fainter barring; pallescens is dull and grey, with long bill; kurilensis is darker than last; fumigatus has shorter bill than previous; mosukei is similar to previous, but darker and more reddish brown; ogawae is darker, duller and more sooty; taivanus is greyer and less rufous than last; magrathi has dense barring extending over mantle and breast; neglectus is similar to previous but darker, with smoky-brown throat; subpallidus differs from last in paler coloration; nipalensis differs in having still darker plumage; talifuensis differs in paler plumage with reduced barring; szetschuanus is more olive than previous, with heavier barring; idius is smaller and greyer than previous; and dauricus is similar to last but darker.
Male has remarkable song, long and complex, a series of tinkling trills one after the other for seconds on end; female not known to sing. Races in Old World sing differently from the two recently split species in America. Calls include various sharp “tac” notes and loud churrs; in America, sharp, high-pitched note in W (i.e. T. pacificus) but lower-pitched, richer note in E (i.e. T. hiemalis). Territorial males exposed to song playback shortly after dawn increased their song output before sunrise one day after the simulated intrusion, which indicates that dawn singing functions in territory defence#R. Young begin singing in Jul of the year they hatch#R. Two instances of adult singing vigorously in response to appearance of an adder (Vipera berus)#R.
Highly varied. Breeds in moist forest with extensive understorey; in W Europe, commonest in deciduous and mixed woodland, well-vegetated suburbs, urban parks and gardens, moorland scrub, and offshore islands with very scant cover. In mixed deciduous-coniferous woodland along a human-density gradient in Sweden, numbers significantly depressed as human numbers increased#R. Habitat use more restricted in some parts of range; e.g. in N Africa found mostly in forested stream valleys at elevations of 1200–1800 m. From sea-level to above treeline; highest-known occurrence at 4575 m in Himalayas.
Food and feeding
Bulk of food invertebrates; arthropods, including spiders (Araneae), beetles (Coleoptera), earwigs (Dermaptera), orthopterans, etc. Also small vertebrates, such as small fish, tadpoles and young frogs. Some vegetable matter, including various berries and some seeds#R; sometimes seaweed. Usually forages low in vegetation, singly or in pairs; will also flycatch, and has been seen to wade in shallow water, immersing head in search of prey; occasionally forages under snow#R. In Norway, routinely begins foraging several hours before sunrise during winter#R. Recorded also as associating with foraging badgers (Meles meles), capturing prey disturbed by them.
Egg-laying from late Mar or early Apr in W Europe, a week or more later in C Europe, early Apr to May on Cyprus, and second half of May in European Russia. Usually double-brooded in Britain and Israel. Polygyny quite frequent in some W European populations. Nest domed, with side entrance hole, constructed of grass and fibres (often wet when placed in position, giving very tight structure on drying), mean dimensions c. 11·3 cm high, 13 cm wide and 14·5 cm deep, internal chamber 6·2 cm in diameter and 5·6 cm high; male builds surplus nests, female selects that to be used and adds lining herself; located in variety of sites, frequently in dense vegetation but also in cavity or crevice, often artificial, as well as odd sites such as pocket of a hanging coat, in rocky locations often in crevice in rock face (e.g. Fair Isle race fridariensis#R), on St Kilda (race hirtensis) often in crevice in ruin; usually low down. Outside of breeding season, up to nine birds may roost communally in nest boxes on cold winter nights#R. Nest may be reused in subsequent years, sometimes by individuals other than original builders. Clutch 3–9 eggs, usually 5–8, larger clutches (of up to 17) probably the result of two females laying in same nest, island races tend to lay smaller clutches; eggs clear white, spotted with pale brown or reddish brown, especially at blunt end, more rarely almost immaculate; incubation by female alone, period c. 16 days; young fed by both parents, brooded only by female, nestling period 14–19 days, mean c. 17 days; fledglings cared for another 9–18 days before becoming totally independent. Adults have been observed feeding young of Great Tits (Parus major) and, in turn, young of present species have been repeatedly fed by a Common Chiffchaff (Phylloscopus collybita)#R. Nesting success in old-growth forest in Poland was 40%; 91% of nest losses resulted from predation#R. Nests parasitized only rarely, by cuckoos (Cuculus spp.)#R#R.
Resident, migratory and partially migratory. Movements nocturnal. In Britain and Ireland largely sedentary, with most ringing recoveries showing movements of 50 km or less, although several recoveries from S coast into C & S France; races on offshore islands in Britain largely sedentary, but one record of zetlandicus from mainland Scotland (Aberdeenshire). N populations in continental Europe highly migratory; individuals ringed in Norway, in Sweden and on E Baltic coast recovered in, respectively, Belgium and Germany, S Spain, and SW France; largely absent from Scandinavia and N European Russia in winter. As for many migrant passerines in W Europe, timing of spring migration has become progressively earlier over last 40 years, probably because of climate change#R. Further S in Europe, movements seem to be mostly altitudinal in nature. In E Asia, breeding areas in E Siberia, Kamchatka and Sakhalin vacated in winter, but island populations appear to be sedentary; in C Asia most movements altitudinal, although detailed data lacking.
Status and conservation
Not globally threatened (Least Concern). Common or abundant over much of range. European population estimated at 20,500,000–26,500,000 pairs, of which at least 7,000,000 in UK. Some island populations may be very small; for example, totals of singing males of fridariensis on Fair Isle (7·6 km²) oscillated between ten and 50 between 1950 and 2015#R#R. Designation of St Kilda race hirtensis as a separate species in 1884 led to a frenzy of collecting activity, although this did not, in all likelihood, put the race in serious danger; it currently numbers 200–300 pairs. Estimates of borealis (Faeroe Is) are in the region of 250–500 pairs. In Japan, race mosukei (Izu Is) is listed as Endangered, and orii (Daito Is) has recently become extinct. Generally, a highly successful species. Has great ability, especially in Europe, to adapt to modified habitat, and at peaks of abundance may be one of the three commonest species in Britain. Free-ranging house cats (Felis catus) a significant source of mortality in some UK cities#R. Hard winters with prolonged snow cover can cause catastrophic declines in numbers in non-migratory populations, but these normally very temporary, and recovery typically very swift. Severe winter weather especially hard on juveniles, which may suffer declines in survival rates of 25% as a result#R. In Kazakhstan, singing males of C Asian race tianschanicus recently (Jun 2012) found in Altai foothills near Buktharma, a considerable N extension of previously known range#R.
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