Common Chaffinch Fringilla coelebs Scientific name definitions

Common Chaffinch is a medium-sized finch (14–18 cm, 17–29 g) with a conical bill and a peaked hindcrown. Most plumages have two broad white wingbars, a greenish rump, and white tail sides. Adult females are generally dull grayish brown above and buffy grayish below. Adult males vary by population (see Subspecies), but the widespread nominate subspecies has a blue-gray crown and nape, reddish-brown back, and rust-colored face and underparts.

Common Chaffinch has 9 full-length primaries (numbered distally, from innermost p1 to outermost p9), 9 secondaries (numbered proximally from outermost s1 to innermost s9 and including 3 tertials, s7–s9 in passerines), and 12 rectrices (numbered distally, innermost r1 to outermost r6 on each side of the tail). Geographic variation in appearance is moderate to extensive (see Systematics History). The following is based primarily on plumage descriptions of the widespread nominate subspecies in Cramp and Perrins (1 Cramp, S., and C. M. Perrins (Editors) (1994). The Birds of the Western Palearctic, Volume 8: Crows to Finches. Oxford University Press, Oxford, United Kingdom. Close ), Clement et al. (2 Clement, P., A. Harris, and J. Davis (1993). Finches and Sparrows: An Identification Guide. Christopher Helm, London, UK. Close ) and Collar et al. (3 Collar, N. J., I. Newton, and P. Clement (2010). Family Fringillidae (finches). In Handbook of the Birds of the World. Volume 15. Weavers to New World Warblers (J. del Hoyo, A. Elliott and D. A. Christie, Editors), Lynx Editions, Barcelona, Spain. pp. 440–617. Close ), along with examination of Macaulay Library images; see Jenni and Winkler (4 Jenni, L., and R. Winkler (2020). Moult and Ageing of European Passerines. Second edition. Bloomsbury, London, UK. Close ) and Ottenby Bird Observatory (5 Ottenby Bird Observatory (2023). Ringers' Digiguide. https://ringersdigiguide.ottenby.se/ Close ) for illustrated ageing and sexing criteria. See Molts for the use of Humphrey-Parkes molt and plumage terminology (6 Humphrey, P. S., and K. C. Parkes (1959). An approach to the study of molts and plumages. Auk 76(1):1–31. Close ) used by Birds of the World. Plumage appearance rather similar between sexes in juveniles (although some males and females may be separable by flight-feather color) but differs markedly in later plumages; definitive appearance is assumed at the Second Basic Plumage.

Natal Down

Retained natal down in nestlings can be dingy brownish. It can remain on the head and stiffen in fledglings, at which time it can bleach to whitish.

Juvenile (First Basic) Plumage

Present primarily July-August. Both sexes similar to definitive basic female but duller. Upperparts uniformly olive-brown except for buffish-white patch on the nape and duller green rump. Underparts dingy whitish, sometimes with a pale buff wash to the breast and diffuse streaks on the sides. Juvenile primaries and rectrices are thinner and more tapered or pointed at the tips than basic feathers and juvenile body feathers are weaker and more filamentous (barb density sparser) than in later plumages, especially undertail coverts. Juvenile males may average a warmer tinge to the back, auriculars, and breast, and duskier or blacker flight feathers than juvenile females, but too much overlap may exist for reliable sexing of most individuals. It may also be possible that males average more white exposed at the base of the primaries (beyond the primary coverts) than females but study is needed.

Formative and First Alternate Plumages

Present primarily September (when fresh) to July (when worn); see Molts concerning alternate plumages. Body feathering similar in appearance to respective sexes of Definitive Basic Plumage, although averaging duller, especially in males. Formative/First Alternate Plumages are best identified by molt limits among upperwing coverts: a few to most median and inner upperwing greater coverts newer, fuller, and blacker with whiter tips, especially in males, contrasting with retained juvenile outer greater coverts with duller tips and browner juvenile primary coverts without pale edging. One to three tertials (often s8, s8–s9, or s7–s9) or central rectrices (r1) rarely replaced, contrasting with retained browner inner secondaries and outer rectrices. Secondaries brownish with thinner or no dull buff to yellow edging. Retained juvenile outer primaries and rectrices are narrower, more tapered or pointed (less truncate) at the tips, and relatively abraded, especially in May‒July. It may also be possible that juvenile primaries average more white exposed at the base of the primaries (beyond the primary coverts), sex for sex, than basic primaries but study is needed. One to 4 inner grater coverts and possibly some body feathering may be replaced during a First Alternate Molt; look for these feathers to be contrastingly fresher, the coverts browner with yellowish edging in April-July. These birds may show three generations of greater coverts, first alternate, formative, and juvenile.

Definitive Basic and Alternate Plumages

Present primarily September (when fresh) to July (when worn); see below and Molts regarding the presence of alternate plumages in this species.

Female. Head and most of upperparts dull earth-brown, tinged dark grayish, with diffuse paler grayer sides of crown and nape, forming indistinct hind collar (more prominent in fresh plumage in autumn); mantle olive-brown to earth-brown; lower back and rump yellowish green; uppertail coverts duller yellowish brown to olive. Central rectrices dusky brown; remaining rectrices blackish, the outermost two feathers (r6) extensively white with dark tips to the outer webs and the next pair (r5) often with a white at the tip of the inner web. Sides of head primarily olive, sometimes with pale buff to whitish arc above eye and gray of nape extending below auriculars to malar region. Lesser upperwing coverts and scapulars brown, as mantle; median coverts white with dark bases often visible; tertials and inner two greater coverts brown with broad pale buff to yellowish buff margins; remaining greater coverts dark brown to blackish brown, margined buff and tipped white; alula, primary coverts and remiges dusky, finely edged buff to yellowish (forming pronounced panel on secondaries when fresh) the bases of all except outermost three primaries white to yellowish white forming small white patch on closed wing, visible just beyond primary coverts. Underparts dull buffish brown, often tinged grayer (sometimes pinkish) on breast and upper belly; flanks sometimes tinged browner; ventral region whitish becoming paler on undertail coverts; underwing coverts largely dull grayish (cf. ML468243901).

Male. Forehead black; crown, nape, sides of neck, and upper back bright slaty gray-blue; back reddish-brown or chestnut; scapulars bright grayish to bluish-green; lower back yellow-olive; uppertail coverts and central rectrices gray to bluish-slate, the rectrices with black shaft streaks and narrowly edged green; remainder of tail as in female except dark portions black. Lores, ocular region, and auriculars pinkish chestnut to bright vinous-pink tinged orange. Wing feathers as in female except blacker and more boldly patterned overall; inner lesser upperwing coverts grayish to bluish-green, often with white fringes and median coverts white (without visible dark bases) resulting in whiter wing patches in flight; bases to inner primaries may average more extensively white, forming a broader visible bar beyond primary coverts. Chin throat, and most of underparts similar in color to sides of head, usually becoming paler on lower breast and flanks; ventral region and undertail coverts whitish; outer underwing coverts bright gray contrasting bright white inner coverts and axillars (cf. ML207898451). In fresh plumage (October–January), forehead feathering can be fringed dark gray, crown to upper mantle feathers fringed browner, the tips of the greater coverts tinged yellowish, and the pink on face can be washed brownish, creating duller or more camouflaged look overall; feather fringing wears off by spring and plumage becomes brighter.

In both sexes, Definitive Basic Plumage separated from Formative Plumage by having upperwing coverts and secondaries uniform in wear and quality, without molt limits; alula and primary coverts dusky to black; secondaries darker with more extensive yellowish fringing; basic outer primaries and rectrices broad, more truncate (less pointed), dusky to blackish (female) or black (male), and relatively fresh, especially in May‒July. Body feathering also averages brighter than that of Formative Plumage, especially in males.

During April‒July, some birds appear to show 1‒4 newly replaced inner greater coverts, which may indicate the presence of alternate plumages (see Definitive Basic and Altermnate Plumages fo rthat species) These feathers appear contrastingly fresh and more filamentous than basic feathers, and can show brown centers with more yellowish fringing than found on the inner two basic greater coverts. It may be possible that other body feathers can be replaced but appearance of body feathering seems to be effected more by wear than by molt (see above). New inner greater coverts can occur during both the first and definitive cycle, which suggests the presence of both First and Definitive Alternate Plumages (see Molts). Study is needed.

General

Molt and plumage terminology follows Humphrey and Parkes (7 Humphrey, P. S., and K. C. Parkes (1959). An approach to the study of molts and plumages. Auk 76(1):1–31. Close ) as modified by Howell et al. (8 Howell, S. N. G., C. Corben, P. Pyle, and D. I. Rogers (2003). The first basic problem: a review of molt and plumage homologies. Condor 105:635–653. Close ). Under this nomenclature, terminology is based on evolution of molts along ancestral lineages of birds from ecdysis (molts) of reptiles, rather than on molts relative to breeding season, location, or time of the year, the latter generally referred to as “life-cycle” molt terminology (4 Jenni, L., and R. Winkler (2020). Moult and Ageing of European Passerines. Second edition. Bloomsbury, London, UK. Close ). In north-temperate latitudes and among passerines, the Humphrey-Parkes (H-P) and life-cycle nomenclatures correspond to some extent but terms are not synonyms due to the differing bases of definition. Prebasic molts often correspond to “post-breeding“ or “post-nuptial“ molts, preformative molts often correspond to “post-juvenile“ molts, and prealternate molts often correspond with “pre-breeding“ molts of life-cycle terminology. However, for species that suspend prebasic or preformative molts for migration or undergo extensive molts on winter grounds, there is often a lack of correspondence between H-P and life-cycle terms (9 Pyle, P. (2022). Defining moults in migratory birds: a sequence-based approach. Journal of Avian Biology 2022(5):e02958. Close ). The terms prejuvenile molt and juvenile plumage are preserved under H-P terminology (considered synonyms of first prebasic molt and first basic plumage, respectively) and the former terms do correspond with those in life-cycle terminology.

Common Chaffinch is widely reported to undergo a Complex Basic Strategy (cf. 8 Howell, S. N. G., C. Corben, P. Pyle, and D. I. Rogers (2003). The first basic problem: a review of molt and plumage homologies. Condor 105:635–653. Close , 10 Howell, S. N. G. (2010). Peterson Reference Guide to Molt in North American Birds. Houghton Mifflin Harcourt, Boston, Massachusetts, USA. Close ), including complete prebasic molts and a partial preformative molt but no prealternate molts (1 Cramp, S., and C. M. Perrins (Editors) (1994). The Birds of the Western Palearctic, Volume 8: Crows to Finches. Oxford University Press, Oxford, United Kingdom. Close , 4 Jenni, L., and R. Winkler (2020). Moult and Ageing of European Passerines. Second edition. Bloomsbury, London, UK. Close ). However, examination of Macaulay Library images of birds in April‒June shows some evidence of both First and Definitive Prealternate molts (see below), in which case Common Chaffinch undergoes a Complex Alternate Strategy.

Prejuvenile (First Prebasic) Molt

This molt is complete in Fringillid finches, in the nest. Little information available on this molt in Common Chaffinch but it occurs during the nesting season, primarily in May‒July.

Preformative Molt

Partial, primarily in June‒October, mostly or entirely prior to autumn migration but possibly or likely may suspend molt for migration and resume it on wintering grounds; study needed. In Europe, starts at age ca. wk 5 or 20‒30 d after Prejuvenile Molt completes, and lasts ca. 45‒50 days (references in 1 Cramp, S., and C. M. Perrins (Editors) (1994). The Birds of the Western Palearctic, Volume 8: Crows to Finches. Oxford University Press, Oxford, United Kingdom. Close ). Includes body feathers, most to all lesser coverts, some to all inner median coverts, and no to most inner greater coverts, occasionally 1‒3 tertials and rarely 1‒2 central rectrices but no other flight feathers. In a sample of > 6,000 birds from Europe, 3‒10 coverts usually replaced (mean 8.5 and all 10 coverts replaced in 14% of birds), tertials replaced in 4.6% (with only 0.3% replacing all three tertials), and rectrices replaced in 0.8% of birds (4 Jenni, L., and R. Winkler (2020). Moult and Ageing of European Passerines. Second edition. Bloomsbury, London, UK. Close ; see also 5 Ottenby Bird Observatory (2023). Ringers' Digiguide. https://ringersdigiguide.ottenby.se/ Close ). Photoperiodic control of the Preformative Molt was studied by Dolnik and Gavrilov (11 Dolnik, V. R., and V. M. Gavrilov (1980). Photoperiodic control of the molt cycle in the Chaffinch (Fringilla coelebs). Auk 97(1):50–62. Close ). The Preformative Molt of birds from late broods molt more rapidly than those from early broods and photoperiodic control systems appear to synchronize the termination of the molt in birds from early and late broods.

First and Definitive Prealternate Molts

These molts have not been documented for Common Chaffinch but examination of Macaulay Library images of birds in April-June (see those and reference to others at the bottom of the Plumages page) appears to indicate that some individuals in both the first and definitive cycles may replace the innermost 1‒4 greater coverts, suggesting a limited First and Definitive Prealternate Molt, respectively, as can be found in other finches and passerines; these feathers are often replaced during prealternate molts in passerines due to there increased solar exposure (4 Jenni, L., and R. Winkler (2020). Moult and Ageing of European Passerines. Second edition. Bloomsbury, London, UK. Close , 12 Pyle, P. (2022). Identification Guide to North American Birds. Part I, Second Edition. Slate Creek Press, Forest Knolls, CA, USA. Close ). This molt, if present, may occur most often on winter grounds in highly migratory populations, where it may have been missed (cf. 9 Pyle, P. (2022). Defining moults in migratory birds: a sequence-based approach. Journal of Avian Biology 2022(5):e02958. Close ); study needed.

Definitive Prebasic Molt

Complete, primarily mid-June to early October, with duration at the individual level reported at 72‒80 d (see 1 Cramp, S., and C. M. Perrins (Editors) (1994). The Birds of the Western Palearctic, Volume 8: Crows to Finches. Oxford University Press, Oxford, United Kingdom. Close for details). Primaries (and corresponding primary coverts) are replaced distally (p1 to p9), secondaries are replaced proximally from s1 and proximally and distally from the central or innermost tertial (s8 or s9), and rectrices are generally replaced distally (r1 to r6) on each side of the tail, though variation in sequence of rectrix molt may occur.

Bioenergetics and photoperiodic control of the Definitive Prebasic Molt in Common Chaffinch were studied by Dolnik and Gavrilov (13 Dolnik, V. R., and V. M. Gavrilov (1979). Bioenergetics of molt in the Chaffinch (Fringilla coelebs). Auk 96:253–264. Close , 11 Dolnik, V. R., and V. M. Gavrilov (1980). Photoperiodic control of the molt cycle in the Chaffinch (Fringilla coelebs). Auk 97(1):50–62. Close ). Metabolized energy increased during the first part of molt and then decreased following molt completion. The caloric equivalent of weight loss at night was lower in molting than in non-molting birds and lower at high than at low ambient temperatures, due largely to the greater amounts of water loss involved. The timing of onset and completion of the Definitive Prebasic Molt in Chaffinches is a remote expression of vernal photostimulation. Under natural conditions, day length immediately before and during the molt does not control either the time of onset or of termination of molt but an additional control system increases the rate of the molt in its early phase and decreases the rate of molt in its late phase, which may play an adaptive role in synchronizing the end of the molt among birds that begin molt at different times.

The following is based on the literature (1 Cramp, S., and C. M. Perrins (Editors) (1994). The Birds of the Western Palearctic, Volume 8: Crows to Finches. Oxford University Press, Oxford, United Kingdom. Close , 2 Clement, P., A. Harris, and J. Davis (1993). Finches and Sparrows: An Identification Guide. Christopher Helm, London, UK. Close , 3 Collar, N. J., I. Newton, and P. Clement (2010). Family Fringillidae (finches). In Handbook of the Birds of the World. Volume 15. Weavers to New World Warblers (J. del Hoyo, A. Elliott and D. A. Christie, Editors), Lynx Editions, Barcelona, Spain. pp. 440–617. Close ), along with examination of Macaulay Library images.

Bill and Gape

The bill of juveniles is dusky brownish and that of adult females is brownish, darkest at tip, palest at base of lower mandible. The bill of adult males is pale bluish gray with dark tip for most of the year, often becoming yellowish or buffish at base during molting periods and in early winter. The gape is covered by head feathering in adults; in nestlings and fledglings it is swollen and yellowish.

Iris

In all ages and sexes, iris is dark brown.

Tarsi and Toes

In all ages and sexes, tarsi and toes vary from pale pinkish-brown to dark gray.

Linear Measurements

Total Length: 140–180 mm.

Mass

17–29 g.

Common Chaffinch has often been considered conspecific with African Chaffinch (Fringilla spodiogenys), Canary Islands Chaffinch (Fringilla canariensis), Madeira Chaffinch (Fringilla maderensis), and Azores Chaffinch (Fringilla moreletti) (14 Hartert, E. J. O. (1921). Die Vögel der paläarktischen Fauna. Systematische Übersicht der in Europa, Nord-Asien und der Mittelmeerregion vorkommenden Vögel. Band 3(17). R. Friedländer und Sohn, Berlin, Germany. Close , 15 Mayr, E. (1968). Subfamily Fringillinae - chaffinches and Brambling. Pp. 202-206 in: Paynter (1968). Close , 16 Cramp, S., and C. M. Perrins (Editors) (1994). Handbook of the Birds of Europe, the Middle East and North Africa. The Birds of the Western Palearctic. Volume 9. Buntings and New World Wood Warblers. Oxford University Press, Oxford, United Kingdom. Close , 17 Dickinson, E. C., and L. Christidis, Editors (2014). The Howard and Moore Complete Checklist of the Birds of the World. Fourth Edition. Volume 2. Aves Press, Eastbourne, UK. Close , 18 Clements, J. F., T. S. Schulenberg, M. J. Iliff, T. A. Fredericks, J. A. Gerbracht, D. Lepage, S. M. Billerman, B. L. Sullivan, and C. L. Wood (2022). The eBird/Clements checklist of Birds of the World: v2022. Cornell Laboratory of Ornithology, Ithaca, NY, USA. Close ), but is here split on the basis of genetic, morphological, plumage and vocal differences (19 Lynch, A., and A. J. Baker (1994). A population memetics approach to cultural evolution in Chaffinch song: differentiation among populations. Evolution 48(2):351–359. Close , 20 Lachlan, R. F., M. N. Verzijden, C. S. Bernard, P. P. Jonker, B. Koese, S. Jaarsma, W. Spoor, P. J. B. Slater, and C. ten Cate (2013). The progressive loss of syntactical structure in bird song along and island colonization chain. Current Biology 23:1896-1901 Close , 21 Illera, J. C., L. G. Spurgin, E. Rodriguez-Exposito, M. Nogales, and J. C. Rando (2016). What are we learning about speciation and extinction from the Canary Islands? Ardeola 63(1):15–33. Close , 22 Recuerda, M., J. C. Illera, G. Blanco, R. Zardoya, and B. Milá (2021). Sequential colonization of oceanic archipelagos led to a species-level radiation in the common chaffinch complex (Aves: Fringilla coelebs). Molecular Phylogenetics and Evolution 164:107291 Close ). In plumage, Common Chaffinch is the most distinct of the five, with the adult male characterized by rufous-brown, cinnamon-brown, or pinkish-brown cheeks, auriculars, which continues on the the throat, breast, and belly, becoming only slightly paler on the belly; gray crown and nape; and brownish to cinnamon-brown back. This contrasts with all other chaffinches, which either have a yellowish-green mantle and blue-gray back as in African Chaffinch, or largely blue-gray upperparts (with some taxa having olive-brown or greenish mantle) in Canary Islands Chaffinch, Madeira Chaffinch, and Azores Chaffinch. African Chaffinch further differs from Common Chaffinch in its blue-gray to slate-colored auriculars and cheek with contrasting white eye-arcs, and pale pinkish-brown throat and breast, which fades to whitish on the belly. The island species further differ from Common Chaffinch in their pale orange to ochre face and upper-breast coloration, which fades to whitish on the belly (16 Cramp, S., and C. M. Perrins (Editors) (1994). Handbook of the Birds of Europe, the Middle East and North Africa. The Birds of the Western Palearctic. Volume 9. Buntings and New World Wood Warblers. Oxford University Press, Oxford, United Kingdom. Close , 22 Recuerda, M., J. C. Illera, G. Blanco, R. Zardoya, and B. Milá (2021). Sequential colonization of oceanic archipelagos led to a species-level radiation in the common chaffinch complex (Aves: Fringilla coelebs). Molecular Phylogenetics and Evolution 164:107291 Close ). In song, the island taxa are distinct in having reduced syntactical structure, with structure declining from Azores Chaffinch to Madeira Chaffinch to Canary Islands Chaffinch (20 Lachlan, R. F., M. N. Verzijden, C. S. Bernard, P. P. Jonker, B. Koese, S. Jaarsma, W. Spoor, P. J. B. Slater, and C. ten Cate (2013). The progressive loss of syntactical structure in bird song along and island colonization chain. Current Biology 23:1896-1901 Close ). Syntax structure may be an important aspect of song that is selected for by females and in territorial bouts, such that changes in syntax structure may play an important role in speciation (20 Lachlan, R. F., M. N. Verzijden, C. S. Bernard, P. P. Jonker, B. Koese, S. Jaarsma, W. Spoor, P. J. B. Slater, and C. ten Cate (2013). The progressive loss of syntactical structure in bird song along and island colonization chain. Current Biology 23:1896-1901 Close ). Lastly, the five taxa of chaffinch show significant genetic divergence, with Common Chaffinch and African Chaffinch appearing to be sister taxa that are in turn sister to the three Atlantic island species (22 Recuerda, M., J. C. Illera, G. Blanco, R. Zardoya, and B. Milá (2021). Sequential colonization of oceanic archipelagos led to a species-level radiation in the common chaffinch complex (Aves: Fringilla coelebs). Molecular Phylogenetics and Evolution 164:107291 Close ).

16 subspecies in 6 groups recognized (23 Clements, J. F., P. C. Rasmussen, T. S. Schulenberg, M. J. Iliff, T. A. Fredericks, J. A. Gerbracht, D. Lepage, S. M. Billerman, B. L. Sullivan, and C. L. Wood (2023). The eBird/Clements checklist of Birds of the World: v2023. Cornell Laboratory of Ornithology, Ithaca, NY, USA. Close ). Nominate subspecies varies clinally; subspecies sarda, syriaca, solomkoi, and alexandrovi possibly represent extreme limit of variation of nominate, but further research required; proposed subspecies hortensis (from Anhalt, in central Germany), balearica (Mallorca), tyrrhenica (Corsica), schiebeli (Crete), caucasica (southern Caucasus region), and wolfgangi (Tomsk, in western Siberia) all considered to fall within range of variation of nominate. Additional proposed subspecies scotica (described from Carmunnock, in southwestern Scotland) and hibernicus (Glengariff, in southwestern Ireland) are both synonymized with gengleri.

In a comprehensive molecular phylogenetic study of the Fringilla coelebs complex that used thousands of genetic markers from across the genome, Common Chaffinch appeared to be sister to African Chaffinch (Fringilla spodiogenys). Together, these two species appeared to be sister to the three Atlantic islands taxa, Canary Islands Chaffinch (Fringilla canariensis), Madeira Chaffinch (Fringilla maderensis), and Azores Chaffinch (Fringilla moreletti) (22 Recuerda, M., J. C. Illera, G. Blanco, R. Zardoya, and B. Milá (2021). Sequential colonization of oceanic archipelagos led to a species-level radiation in the common chaffinch complex (Aves: Fringilla coelebs). Molecular Phylogenetics and Evolution 164:107291 Close ).

Has hybridized with Brambling (Fringilla montifringilla) (43 McCarthy, E. M. (2006). Handbook of Avian Hybrids of the World. Oxford University Press, Oxford, UK. Close ).

• Common x African Chaffinch (hybrid) Fringilla coelebs x spodiogenys
• Common Chaffinch x Brambling (hybrid) Fringilla coelebs x montifringilla

British Isles, Continental Europe, Balearic Islands, Sardinia (Italy), Crete (Greece), Cyprus, Levant, Iraq, Iran, Turkmenistan, and Siberia. Some populations winter to Africa.

Introduced Range

Introduced (nominate subspecies or gengleri) in southern South Africa (Cape Town area) and New Zealand.

Extralimital Records

Vagrant to Saudi Arabia (recently more regular in eastern provinces), Kuwait, United Arab Emirates, and east to northwest Thailand, eastern China (some regular overwintering), Korea, and Japan; also eastern North America.

Lowland and lower montane deciduous, mixed and conifer woods with slight preference for beech (Fagus), hornbeam (Carpinus), mature oak (Quercus), spruce (Picea) and pine (Pinus), forest edges and glades; also copses, heaths, edges of tundra and agricultural areas, hedgerows, orchards, parks, and gardens. In non-breeding season more widespread in similar habitat and open agricultural areas, particularly weedy fallow and stubble fields, olive (Olea) groves, palms, wadis, and desert oases. Sea-level to 2,500 m.

Resident, partially migratory, and migratory in different parts of the range. Breeders in the north and northeast of the range move south and southwest between mid-September and the end of November, to wintering areas within breeding range in central and southern Europe, around eastern Mediterranean, also northwest Africa, and farther south in central Asia, onward or further movements within range usually triggered by onset of severe weather; return north from late February to early May, adult males making earliest movements, ahead of adult females and first-year individuals. Evidence from ringing shows that juveniles and adult females move farther than males; wintering population in Sweden, Britain, Belgium, and Netherlands mostly males, in Ireland predominantly females. Some evidence of winter site-fidelity among migrants wintering in Britain in subsequent years. Most movement diurnal, in small to medium-sized flocks, often visibly along lines of hills, river valleys, and coasts (particularly at headlands) and through high mountain passes; in some areas, particularly Baltic, North Sea, and eastern Mediterranean coasts, passage often continuous for several days when weather conditions favorable; sometimes huge numbers involved, e.g., daily peaks of hundreds of thousands of individuals at Falsterbo (southern Sweden), and 60,000 in a single day in Schwäbische Alb (southern Germany). Western breeders move farthest west and those from more eastern populations winter progressively farther east. Those from Scandinavia head west or south through Denmark and Netherlands (seldom directly across North Sea) to non-breeding grounds mainly in central and southern British Isles, Netherlands to western France and western Spain, peak arrival in southeast England second half October, subsequent movements north and west to northern Wales and Ireland; return route reverse of autumn one, but more frequently crossing North Sea on broad front in less than a day. Breeders from Finland, eastern Baltic region, Poland, and northwest Russia move between south and southwest to winter areas mainly in southwest France and western Spain, with smaller numbers Britain, southern Germany and Switzerland, where local breeders make relatively short flight to southern France, northern Italy, eastern Spain and Balearic Islands; Polish breeding birds migrate before main passage from farther north; those from Czech Republic head southwest to southern France, smaller number reaching Spain and Italy, and birds from Slovakia winter mainly northern Italy, joining others from western Russia, Hungary, Switzerland, Germany, France, and smaller numbers from Finland and Poland; in western Alps passage into Switzerland of first-year females reaches peak 3–4 days ahead of adult females, first-year males up to a week later, and adult males peaking up to 12 days later. Numerous in winter in northwest Africa, where most arrive via Strait of Gibraltar, mostly of central European origin but small numbers also from western Russia. In Russia, northern breeders form large post-breeding flocks from early August and move south and southwest from early September to mid-October, wintering within southern area of breeding range and in central and southern Europe; in north-central Siberia breeding areas along Yenisey River deserted by mid-September, return passage through Caucasus late February or early March, and farther north from late March or early April, continues to early May, males arriving back on breeding territory on average one week ahead of females; regular on passage in western and central Mongolia, western China, and throughout Kazakhstan, with main direction in autumn southwest through Ural Valley to northern Caspian region and Tien Shan foothills, also some birds wintering in eastern China (Hebei, Liaoning); in northern Iran common visitor early October to early April in areas south of Caspian Sea; winter visitor also in northwestern and southeastern Afghanistan, northern and western Pakistan (regular Baluchistan, scarce in north), Kashmir, occasional or sporadic northwest India and Nepal. Breeding birds in Britain (subspecies gengleri) largely sedentary, with 90% of ringing recoveries up to 5 km from natal area (remainder moving less than 50 km), and French breeders largely sedentary, but small numbers move short distance west or southwest (occasionally reaching Spain); proportion of population migratory increases farther east and north, e.g. 60% of breeders in Belgium sedentary, and small numbers wintering southern Scandinavia (mostly Sweden and southern Norway); irregular in winter in western Russia, but small numbers in Saint Petersburg region; annual in varying numbers in Iceland, mostly in spring and autumn, involving individuals drifted off passage to/from Scandinavia. Wintering birds (females often predominating) arrive late September to mid-November and outnumber local breeding populations in Greece, Türkiye, and Cyprus, and a common passage migrant and winter visitor in Lebanon, Syria, Israel, northern Jordan, and Iraq; small numbers of nominate subspecies winter northern Egypt and coastal Libya (mostly Tripolitania) between late October and early April.

Site-fidelity variable, strong in males in England and in pairs southern Finland; evidence from Russian study that ca. 90% of year-old individuals return to breed within 1 km of natal site.

Food Capture and Consumption

Actively forages on the ground throughout year, also at all levels in bushes and trees in spring and summer; perches briefly on near-vertical tree trunks (particularly during outbreaks of bark beetles), and occasionally pursues and catches flying insects, also hovers briefly to take invertebrates from leaves or beside suspended fat or peanuts at garden feeders . Wades in shallow water; puts head beneath surface to collect caddis fly larvae, and removes outer case as if it were a seed husk. On ground , it has a distinctive shuffling or jerky, forward-hopping or walking motion, simultaneously nodding head.

In the nonbreeding season, the nominate subspecies often forages in flocks consisting solely of males or solely of females and immatures (this behavior less frequently recorded for other subspecies) and in association with Brambling (Fringilla montifringilla), European Greenfinch (Chloris chloris), and sparrows (Passer).

Major Food Items

Diet the most varied in the family, mostly small invertebrates and their larvae, also seeds and buds.

Animal items include springtails (Collembola), mayflies (Ephemeroptera), small dragonflies (Odonata), stoneflies (Plecoptera), grasshoppers (Orthoptera), earwigs (Dermaptera), cockroaches (Blattodea), bugs (Hemiptera), lacewings (Neuroptera), moths (Lepidoptera), flies (Diptera), caddis flies (Trichoptera), ants and wasps (Hymenoptera), beetles (Coleoptera), harvestmen (Opiliones), millipedes (Diplopoda), centipedes (Chilopoda); also snails (Pulmonata) and small earthworms (Lumbricidae).

Seeds, buds (mostly of trees in spring), and flowers include those of yew (Taxus), cypress (Chamaecyparis), juniper (Juniperus), larch (Larix), spruce (Picea), pine (Pinus), fir (Abies), Douglas-fir (Pseudotsuga), willow (Salix), poplar (Populus), birch (Betula), alder (Alnus), elder (Sambucus), hornbeam (Carpinus), beech (Fagus), chestnut (Castanea), oak (Quercus), elm (Ulmus), mulberry (Morus), tulip-tree (Liriodendron), sycamore (Acer), lime (Tilia), ash (Fraxinus), olive (Olea), mistletoe (of genus Viscum), nettle (Urtica), knotgrass (Polygonum), dock (Rumex), goosefoot (Chenopodium), beet (Beta), orache (Atriplex), glasswort (Salicornia), amarant (Amaranthus), chickweed (Stellaria), mouse-ear (Cerastium), corn spurrey (Spergula), buttercup (Ranunculus), meadowsweet (Filipendula), rose (Rosa), bramble (Rubus), strawberry (Fragaria), currant (Ribes), cinquefoil (Potentilla), apple (Malus), pear (Pyrus), cherry (Prunus), rowan (Sorbus), hawthorn (Crataegus), cotoneaster (Cotoneaster), clover (Trifolium), wood-sorrel (Oxalis), cranesbill (Geranium), spurge (Euphorbia), sea-buckthorn (Hippophae), willowherb (Epilobium), evening primrose (Oenothera), dwarf alpenrose (Rhodothamnus), bilberry (Vaccinium), crowberry (Empetrum), grape (Vitis), primrose (Primula), bedstraw (Galium), thrift (Armeria), comfrey (Symphytum), forget-me-not (Mysotis), plantain (Plantago), sunflower (Helianthus), nightshade (Solanum), figwort (Scrophularia), snowberry (Symphoricarpos), teasel (Dipsacus), sedges (Carex), rushes (Juncus), and various grasses and cereals.

Nestlings fed with insect larvae, mainly aphids (Aphidoidea), and caterpillars (Lepidoptera).

Song , by male only, mainly March to mid-July throughout most of range, a descending musical rattle of several notes followed by accelerated final flourish, "chip chip chip, tell tell tell, cherry-erry-erry tissi cheweeeoo," usually repeated from prominent perch in tree or bush; subsong, mostly from ground or from dense cover, occasionally given also by female, softer or more subdued than full song and mostly series of chirps, warbles and rattling notes. Slight variations in song throughout range, e.g., birds in northern Europe give short, sharp "chip" or "chink" similar to call of Great Spotted Woodpecker (Dendrocopos major) immediately following end of song. Calls include sharp, distinctive "pink" or "chink," singly or as double-note (latter seemingly unknown in central and southern Europe), nominate subspecies also has variations of "twit" or "fit," also a loud "wheet," "whit" or "tsip" most often given in spring; high-pitched wheezing "eeese" and a soft "chup" or "tsup" flight call; male also gives loud and drawn-out "hooeed," regionally variable, often given on overcast or wet days (hence known as "rain-call"). In flight, frequently gives soft "tsup," "tap" or "chup" note; female usually silent, but sometimes gives soft "chip" or "chirrip" or, in spring, may respond to male song with soft "zi" or wheezing "si-si-si."

Territorial Behavior

Solitary nester, territorial.

Mating System and Operational Sex Ratio

Monogamous, exceptionally bigamous.

Courtship, Copulation, and Pair Bond

Territorial male, with sleeked plumage (highlighting wingbars), displays to female by singing and by making low, short, undulating moth-like flight with wings extended and beaten rapidly while head and tail held low, and on landing crouches, turns sideways to female, becomes motionless, lifting or flicking wings, and then tilts body on side closest to female and raises wing for several seconds, revealing flanks and belly (may also alternately switch wings depending on location of female, each time raising wing closest to her), before relaxing and moving off in moth-like flight to repeat display, or female flies in rapid zigzag chase with male in pursuit; mating usually takes place towards start of nest-building, may be initiated by either sex, but successful only if female solicits with wings lowered and shivered, tail partly raised and breast feathers fluffed while giving loud "seep" calls. Pair bond frequently maintained into subsequent years (sedentary subspecies gengleri).

Season mid-March to mid-July; one brood, exceptionally two.

Nest placed up to 35 m above ground on branch, against trunk or in fork of tree or bush.

Nest, built entirely by female, a deep cup of plant fibers, grass, fine roots, lichens, moss, bark strips, animal hair and feathers.

Clutch 4–5 eggs, pale bluish-green to light reddish, variably marked with purple-brown blotches, spots or fine streaks.

Incubation by female, period 10–16 days.

Nestling period 11–18 days.

In early courtship, male initially dominant, but female gradually assumes dominance during egg-laying and incubation. Chicks fed and cared for mostly by female, male contributes little or up to one-third of food for young. Young fed by both parents for up to 21 days after fledging.

Breeding success: of 10,967 eggs in Britain, 59% hatched and 41% of these produced fledged young; in 1950–60 proportion of successful clutches increased from 38% of those begun in early Apr to ca. 50% in late May (owing largely to reduced predation as vegetation cover increased), 2.6–4.5 young fledged from successful nests depending on month and clutch size (clutches of 5–6 most successful). In a western Russia study, 63% of 3,859 eggs hatched and 52% fledged young, averaging 3.1/successful nest; of 20 nests in a German study, average of 3.7 young/successful nest. Breeds in first year.

Longevity at least 14 years.

Losses of eggs and young due to predation by crows (Corvidae), red squirrels (Sciurus vulgaris) and gray squirrels (Sciurus carolinensis), cats, also stoats and weasels (Mustela), nest failures caused also by human predation or disturbance, bad weather and desertion following loss of parent.

Territory Size

Size of breeding territory (used for courtship and nesting) variable, average in southern England 0.7 ha, at start of breeding season territory larger, size decreasing as areas forfeited to later-arriving settlers.

Not globally threatened; evaluated as Least Concern by BirdLife International under IUCN Red List Criteria. Common to locally abundant. Estimated European breeding population (including Russia west of the Urals) 230,000,000 pairs, most of which in Britain, Fennoscandia, Germany, Croatia, Belarus, and Russia. Breeding densities are variable: in Britain highest densities are 631 birds/km² in Scots pine (Pinus sylvestris) woods, average 58 pairs/km² in woodland and 21 pairs/km² on farmland; in Fennoscandia more numerous in deciduous woods (49–145 pairs/km²) than in conifers (12–102 pairs), and in Bialowieza Forest, in eastern Poland, up to 157 pairs/km² in deciduous forest and up to 100 pairs/km² in coniferous forest; in Switzerland, more numerous (up to 190 pairs/km²) in subalpine spruce than in mixed oak-hornbeam forest (max 110 pairs). Numbers in much of range largely stable; slight increase in Spain, Britain, Denmark, Croatia, and Ukraine during 20th century as a result of changes in forest structure and management.

Chuenchom Hansasuta received a Doctor of Dental Surgery from Chulalongkorn University. During her long and distinguished career in dentistry, she studied and practiced in places such as Thammasat University (Thailand), State University of New York at Buffalo (USA), and University of Connecticut Health Center (USA) and retired in 2020. Chuenchom always had an intense curiosity for birds that over time, evolved into an acute interest in plumages. She has long been active in education and volunteering, becoming chairwoman of The Flyway Foundation and actively engaging and educating the public in the study of birds and their plumages.