Moult papers:

references, abstracts and comments. Where there is no abstract, an abstract has been written, where abstracts are too long they have been abridged, where the English is poor it has been improved. Abstracts/summaries in languages other than English have been translated into English, some extremely bad abstracts (and even headings) from Ringing & Migration have been corrected when called for. The comment is personal, it points out errors and possible follow-ups, it is begun: CP:

ascending (ascendent, ascendant) primary moult: begins with the outermost primary.
descending (descendent, descendant) primary moult: begins with the innermost primary.
distal primary: the most distant ones from the body
prebasic moult = postnuptial moult
proximal primary: the ones closest to the body
transilient mode: from the last moulted remige there is a forward or backward jump, omitting one or several intermediary remiges.
dorsal: on the back side
ventral: on the belly side

G, H, I, J, K, L, M

Galbraith, H. (1977): The Post-nuptial Moult of a Migratory Population of Pied Wagtails. Ring. & Migr. 1: 184 - 186.

Gargallo, G. (1995): Interrupted moult of some passerine birds in Southern Europe. Ring. & Migr. 16: 117 - 120.

The first records of an interrupted pre-nuptial moult of Bonelli's Warbler Phylloscopus bonelli are described together with more cases of interrupted moult in Blackcap Sylvia atricapilla, Siskin Carduelis spinus, Pied Flycatcher Ficedula hypoleuca and Subalpine Warbler Sylvia cantillans moltonii. It is stated that some birds of this insular race of Subalpine Warbler can undergo a partial instead of complete moult and, consequently, it is suggested that some could undergo a complete pre-nuptial moult.

Gargallo, G. & O. Clarabuch (1995): Exten(t of) moult and ageing in six species of passerines. Ring. & Migr. 16: 178 - 189.

This paper describes the post-juvenile moult of the Chiffchaff Phylloscopus collybita, Serin Serinus serinus, Greenfinch Carduelis chloris, GoldfinchCarduelis carduelis and Linnet Carduelis cannabina and the first pre-nuptial moult of the Woodchat Shrike Lanius senator. These data are used to evaluate correct ageing criteria for each species, and some new and more reliable ageing methods are also presented. Special attention was given to the overall extent and pattern of moult in order to age birds more reliably - particularly when working with populations which show an extensive partial moult. The published cases of complete post-juvenile moult in some cardueline finches are discussed.

Gauci, C. & J. Sultana (1981): The moult of the Fan-tailed Warbler. Bird Study 28: 77 - 86.

Fan-tailed Warblers, both adults and juveniles, have a complete moult in autumn, mainly between June and October. Adult males moult in about 92 days while adult females average 67 days. First-year birds moult in 75 days (females) and 81 days males. From spring to early autumn birds can be aged according to the abrasion of wing and tail-feathers. Adults, as well as some juveniles, can also be sexed. Primaries moult in the normal way, i.e. descendantly, but moult of the secondaries proceeds in both directions and finishes on the 4th. Many birds moult the tertials and some of the greater coverts twice - at the beginning and at the end of moult. This phenomenon has apparently not been encountered previously in other Palearctic passerines studied. Exceptionally, some primaries and secondaries may also be moulted twice. Head feathers take longest to moult. Adult males start moulting about two weeks before females. Many juveniles, especially males, suspend moult and show breeding behaviour. When moult is resumed, the renewed feathers are moulted once again. First-year females from early broods develop brood-patches. Young hatched in the latter part of the season start to moult at an earlier age than those fledging from early broods.

CP:Small materials, fitted by eye; suggested durations are not conclusive.

Ginn, H. B. (1975): The timing and sequence of the complete annual moult in the Dunnock (Prunella modularis) in Britain over an eleven year period. J. Orn. 116: 263 - 280.

Data from the British Trust for Ornithology's Moult Enquiry collected in Britain from 1960 to 1970 were analysed to discover for the adult Dunnock (Prunella modularis), the moulting period, the rate of moult in the individual, the sequence of feather replacement and the relative timing of moult in different tracts, excluding the small contour feathers. The feather scores recorded on the moult cards were used to produce scatter diagrams on which calculations were based. The various ways of using such diagrams are discussed briefly. It is suggested that the speed of moult can often be estimated by visual appraisal, but regression analysis if used should be of dates of capture against individual moult scores, n o t of moult scores against each date....
The sequence of moult is similar to that of most other small passerines but the rate is rather faster than might be expected for this species which is remarkably sedentary in Britain.
The use of moult for ageing at certain times is suggested; Dunnocks in new plumage before mid-August are almost certainly juveniles as are birds in tail and body moult with new remiges.
The timing and rate of moult in different years and parts of Britain were compared and statistically significant differences were found in some cases. Birds caught in moult in different years were found not to be inherently early or late moulters. The probable relationship between the end of the breeding activity and the start of moult is discussed. (...)

Ginn, H. B. & D. S. Melville (1983): Post-nuptial moult in the Garden Warbler. Bird Study 16: 131 - 132.

Gladwin, T. W. (1969): Moult in Birds. BTO Guide 19.

Glutz von Blotzheim, U. N. (1972): Zur Mauser von Charadrius hiaticula, dubius und alexandrinus.. J. Orn. 113: 323 - 333.

Contrary to the information given by WITHERBY (1943) and other handbooks the Ringed, Lesser Ringed and Kentish Plover show quite different moult patterns and plumage sequences.
The Dutch and German Ringed Plovers spending the winter on the shores of western and south-western Europe acquire their first nuptial plumage by a partial moult of the juvenile plumage and the adult nuptial plumage by a lengthy complete moult; the moult of Scandinavian "hiaticula" differs at least in respect of phenology. Amongst Ringed Plovers breeding in the Russian and Siberian Tundra and wintering in south-eastern Africa even the young birds are forced to moult the wing feathers before the beginning of the spring migration (E. and V. STRESEMANN 1963, 1966). The wing-feathers therefore are renewed during the post-juvenile moult. The complete postnuptial moult produces a modestly coloured winter plumage; the black and white markings of head and breast are the results of a restricted prenuptial moult.
Within their first year Little Ringed and Kentish Plover acquire a winter and a first nuptial plumage; the winter plumage is obtained by moult of parts of the juvenile pluamge, the nuptial plumage by partial moult of the winter plumage. The moult and plumage sequences after the first breeding season are the same as in Ch. h. tundrae with one exception concerning the Kentish Plover. In this species, the excessive wear of the nuptial dress, due to the light plumage and the exposed habitat, requires a special "summer moult" already before the beginning of the post-nuptial moult.<(...)

Gratto, C. L. & R. I. G. Morrison (1981): Partial postjuvenile wing moult of the Semipalmated Sandpiper Calidris pusilla.. WSG Bull. 33: 33 - 37.

Green, G. H. & R. W. Summers (1975): Snow Bunting Moult in Northeast Greenland. Bird Study 12: 9 - 17.


Hansen, K. 1985: Afbrudt fældning hos Grå Fluesnapper Muscicapa striata. DOFT 79: 60 - 62, 1985.

In spring 1983, 75 specimens of Spotted Flycatchers were examined on the island of Hjelm in Kattegatt. 21 of these birds showed a state of aberrant wing-moult, concerning the secondaries. Tab. 1 shows the pattern of arrested moult in these birds.
Attention is drawn to the fact, that 12 of these 21 birds still had an old innermost secondary (s6). It is suggested, that these birds must have renewed s6 in autumn, and then arrested the moult before migrating to Africa. One specimen among 65 birds from Africa, examined by Stresemann (J. Orn. 104: 101-111, 1963), supports this theory. The skin was collected on 18 November, and it has s6 new while all other secondaries are still old.

Harris, M. P. & R. F. Yule 1977: The moult of the Puffin Fratercula arctica. Ibis 119: 535 - 541.

(from the discussion) The basic pattern of timing of wing moult appears to be that the majority of adult and immature Puffins replace their primaries in late winter prior to returning to the colonies Many first-year birds moult in mid-summer and these do not visit land that year. The pattern is obscured by two factors (...) firstly, the breeding season varies greatly from colony to colony. (...)Secondly, some birds in adult plumage are still not breeding, as is shown by an influx of fresh-plumaged recently-moulted adult-beaked non-breeders into the colonies a month or two after the breeders have returned.

Harper, D. G. C. 1984: Moult interruption in passerines resident in Britain. Ring. & Migr. 5: 101 - 104.

The note records ten cases of moult interruption in three species of passerines resident in Britain: Robin Erithacus rubecula, House Sparrow Passer domesticus and Greenfinch Carduelis chloris. In addition previous record of moult interruption are reviewed and possible functions discussed. The importance of distinguishing between cases of "arrested" and "suspended" moult is stressed; many published records of moult interruption do not do so.

Hasselquist, D, A. Hedenström, Å. Lindström & S. Bensch (1988): The seasonally divided flight feather moult in the Barred Warbler Sylvia nisoria - a new moult pattern for European passerines. Orn. Scand. 19: 280 - 286.

Wing and tail feather moult in Barred Warblers was studied during three summers in SE Sweden. Birds arriving at their breeding grounds had on average 3.2 - 4.9 fresh secondaries and 3.9 - 9.4 fresh tail feathers, most certainly moulted in winter quarters. A few primaries and tertials had also been renewed. After breeding, normally all primaries and tertials were moulted. Primary moult duration was approximately 40 d for the individual and 60 d for the population. On average, only 0.7 - 1.4 secondaries were shed. No individuals renewed any winter-grown secondaries the following summer. Normally, 3 - 4 (mainly central) tail feathers were moulted. Thus, Barred Warblers moult primaries, tertials and central tail feathers in summer, and secondaries and a varying number of tertials and tail feathers in winter. The juvenile winter moult of secondaries shows that the seasonally divided flight feather moult is actively initiated already in the first winter; this is interpreted as a preparation for an early departure from breeding grounds the following summer. This moult pattern has not previously been documented in European passerines.

Haukioja, E. & J. Reponen (1968): Varpusen Passer domesticus sulkasadosta. Transl.: Moult in the House Sparrow. Porin Lintutiet. Yhd. Vuosik. (Yearbook of Porin Orn. Soc.) 2: 49 - 51.

Haukioja, E. & P. Kalinainen (1969): Transl.: Postnuptial moult of Willow Warbler (Phylloscopus trochilus), Whitethroat (Sylvia communis) and Meadow Pipit (Anthus pratensis). Porin Lintutiet. Yhd. Vuosik. (Yearbook of Porin Orn. Soc.) 2: 75 - 78.

Haukioja, E. (1971): Processing moult card data with reference to the Chaffinch Fringilla coelebs. Orn. Fenn. 48: 25 - 31.

The duration of primary moult in the Chaffinch (in Finland) is a little more than 70 days as calculated by two different methods.(...)

CP: Haukiojas value is based on the regression Moult score versus Time (which approximates the duration of the moult of the population) and retraps of moulting birds.

Haukioja, E. (1971): Flightlessness in some moulting passerines in Northern Europe. Orn. Fenn. 48: 101 - 116.

The effects of wing and tail moult on flying ability have been evaluated for four passerine species (Luscinia svecica, Phylloscopus trochilus, Motacilla flava, M. alba) in Finnish Lappland and Sylvia communis in southern Finland. All species have a short postnuptial moult (between 35 and 50 days) and in L. svecica, Phylloscopus trochilus and Sylvia communis some moulting specimens unable and/or unwilling to fly have been caught. The stage of the moult affects the probability of a bird being caught so that birds which are at the peak of their moult are more difficult to catch with nets. The phenomenon of flightlessness is therefore obviously more common than can be evaluated from moult card data.
Flightlessness depends on the fact that not only are many wingfeathers lost simultaneously but also that in Luscinia svecica, and to a lesser extent in Phylloscopus trochilus, tail feathers are lost simultaneously. The tail moult is most intense during a phase when there are not very large gaps in the wing: this helps in part to maintain as high a degree as possible of flying ability. Wagtails (open field species) can avoid flightlessness, in spite of a short moult, by a prolonged tail moult.

Haukioja, E. & P. Kalinainen (1972): Transl.: The ecology of some passerines during the moulting period. Porin Lintutiet. Yhd. Vuosik. (Yearbook of Porin Orn. Soc.) 3: 5-16.

Some analyses of moulting data collected during the summers of 1968 and 1969 in the delta of the Koskemäki River (61° 30' N, 21° 30' E), in western Finland, are presented.(...) According to a regression equation the postnuptial moult of the Yellow Wagtail (Motacilla flava) lasts about 40 days, which, if anything, is too long an estimate.(...)
Factors affecting on the timing of the moult are discussed. If we accept the idea that temperate zone passerines reproduce as fast as they can, the following course of events is probable. Breeding lasts as long as the numbers of parents lost before the following breeding season due to the attempt to breed are lower than the numbers of young which reach maturity. In the course of the summer the risk of losses rapidly increases because the clutch size is smaller and thus even the potential breding result is lower. Besides, losses of nestlings are generally higher in late broods than in early ones and, after independence, the young from late broods probably survive less well than those from early broods. Taking into account that parents have less time for late summer events (moult, premigratory fat-deposition), the later they try to breed, then also their losses are probably larger and breeding has to come to a stop rather early. Moulting is the following phase in many passerines.(...)
Some possibilities of moult study as a part of an ecological study are discussed. For example, the importance of the moulting period in the timing of breeding and other cycles is stressed. The cessation of breeding is determined to a large degree by the need to moult at a favourable time. The shortage of available food for nestlings is probably also important but the paramount importance of a favourable time for moulting is shown by the fact that some passerines moulting after autumn migration are capable of laying two clutches per season in spite of late arrival (In Finland e.g. the Reed Warbler, Acrocephalus scirpaceus and probably the Grasshopper Warbler Locustella naevia). Besides this group the only passerines laying two clutches per season in Northern Europe seem to be those capable of postponing their moult and migration to late summer or autumn (and probably are often capable of moulting using vegetable food). Species moulting before migration and eating animal food normally only breed once, evidently because the favourable time before autumn migration is too short for another brood and moult.

CP: The mid section brought out with red is not quite to the point, when it comes to the Scanian breeding population of Motacilla f. flava; this population has a tendency to "push" breeding and ignore moult, and there may be something of a deviating moral here. The question is: is this particular Darwinist approach - or part of it - refuted by the Scanian example, or do Scanian flava merely bring out the imperative environmental constraints in full relief? H. & K. were inspired by Cody, M. L. (1966): A general theory of clutch size. Evolution 20: 174 - 184, but most likely by a few general works as well; there is a taste of David Lack to the whole approach.

Hedenström, A., S. Bensch, D. Hasselquist, Å. Lindström, D. Pearson & S. Åkesson (1992): Viewpoint. Split moult: stress or strategy? Ring. & Migr. 13: 179 - 180.

Hedenström, A., Lindström, Å. & J. Pettersson (1995): Interrupted moult in adult Willow Warblers Phylloscopus trochilus during autumn migration through Sweden. Ornis Svecica 5: 69 - 74.

Heinroth, O. (1898): Verlauf der Schwingen- und Schwanzmauser der Vögel. Sitzungsbericht der Ges. naturforsch. Freunde zu Berlin, No. 98

Henriksen, K. (1985): Den postnuptiale fældning af svingfjerene hos Hjejle Pluvialis apricaria. DOFT 79: 141-150, 1985.

Moult of the remiges was examined in 1268 wings from adult Golden Plovers Pluvialis apricaria collected in Denmark. Data on moult of a few museum skins collected in N. Scandinavia were added.
(...)Knowing the relative feather-mass of each primary (Tab. 1), the score of the primaries was converted to mass of new feather material.(...)
Moult of the primaries began with the shedding of the innermost primary (P1) and proceeded outwards towards the wingtip with a maximum of four feathers growing simultaneously. Only 0.8 % of the examined birds deviated from this descendant moult pattern. The inner primaries were shed in a more rapid succession than the outer ones, but the rate of progression during the beginning of the primary moult seemed to be slow in comparison with other waders moulting in temperate Europe.
One hundred and four wings showed evidence of suspended primary moult, and most (82 %) of these suspensions occurred with six or seven new primaries in the wing. Nearly all (92 %) of these suspensions were found in birds collected in the last half of August and the first half of September. In these two periods respectively 13 % and 14 % of the birds had suspended primary moult.
The secondaries were moulted in two groups. The moult of the first eleven, numbered from the wrist (S1-S11), most often started with the shedding of S1 or one of the two feathers S10 or S11. After the termination of the primary moult most birds had new S1 and S11, but the rest of these secondaries, and especially those in the centre of the row, were moulted to a lesser extent. On the average a little more than five of these eleven secondaries were moulted every year.
The sequence of moult in the tertials (S12-S15) was quite irregular and much individual variation concerning the onset of this moult was evident. Some birds initiated the tertial moult with the loss of the innermost primary, others not until the sixth primary was fully grown. Usually one or two tertials were growing at a time and suspensions were common, occurring in 207 of 526 birds with from one to three new tertials.
(...) The mean duration of primary moult in the individual bird was 101 days, and the total moulting period of these feathers covered 158 days.
...a complete separation of primary moult in the Golden Plover from breeding and migration would add to the enhanced difficulties of winter survival, compared to estuarine wintering waders in Europe, this species seems to experience. It is shown that about half of the Golden Plovers would have to complete the primary moult in December as a result of such a postponement. This late completion of moult, involving the long outer primaries, would coincide with a period of building-up of nutritional reserves and with cold-weather movements.

Hereward, A.C. (1979): The Autumn Moult of the Yellow Wagtail. Ringing and Migration 2: 113 - 117.

The Yellow Wagtail Motacilla flava flavissima undergoes a rapid complete post-nuptial moult in Britain. Duration of primary moult is 43-45 days and the mean starting date is 19 July. Tail moult is variable in its pattern. The extent of the post-juvenile moult of greater coverts varies by season and between years. The most peculiar feature of moulting Yellow Wagtails is that they seem to start migrating before completing moult.

Herroelen, P. (1960): De rui van de Boerenzwaluw, Hirundo rustica L.,in Belgisch-Kongo. Gerfaut 50: 87 - 99.

Holmberg, T. (1992): The primary moult of the Brambling Fringilla montifringilla evaluated by four different methods. Orn. Svec. 2: 139 - 146.

The primary moult of the Brambling Fringilla montifringilla was studied between 1988 and 1991 at Ånnsjön (63°15'N; 12°28'E) in the Swedish mountain region. The duration of primary moult was estimated in four different ways: 1) By linear regression of population data; 2) By calculating the mean duration of moult of 34 recaptured birds, assuming a linear relationship between time and moult progression; 3) By using data from the same recaptured birds but correcting for the non-linear progression of moult; 4) By using field data on the rate of shedding of different primaries and the growth rate of different individual feathers to construct a moult score versus time curve. Linear regression of population data gave results varying from 32 to 70 days depending on population and method of analysis. The other three methods all resulted in a primary moult duration of 58 - 60 days. The recaptured birds often showed a weight decrease between captures but this seemed not to affect the moult speed. The mean starting date of moult of the population varied from 8 July to 18 July in the four years. The males tended to start a little earlier than the females, but the difference is not significant. The estimated duration of the primary moult (ca 59 days) is considerably longer than the result (46.5 days) of another recently published study. The difference is probably due to different methods of analysis.

Holmes, R. T. (1966): Molt cycle of the Red-backed Sandpiper (Calidris alpina) in western North America. Auk 83: 517 - 533.

In breeding populations of C. alpina in northern Alaska (71 ° N), the prebasic molt of adults is condensed into the short span of the arctic summer and overlaps with almost the whole of the breeding season. Molt begins early in the summer, and new flight feathers are completely grown before departure for winter quarters. At lower-latitude breeding localities in western Alaska (60-66° N), molt apparently starts later and breeding begins earlier than at higher latitudes, with the result that less overlap occurs. Even in the northern populations, however, the major portion of energy expenditure on molt comes in late summer when breeding is over and young Red-backs are fully grown.
Molt takes place during slow phases of both fall and autumn migrations, energy reserves evidently being sufficient to support both activities. In periods of rapid migration, molt is suspended.
The specialization in the molt schedule of C. alpina relate to the fact that this species, with its winter quarters in north-temperate regions, remains in the arctic through the end of summer, during which time it exploits tundra food sources in the absence of possible competitors.

Holmes, R. T. (1971): Latitudinal differences in the breeding and molt schedules Of Alaskan Red-backed Sandpipers. Condor 73: 93 - 99. Internetversion av denna uppsats

On the Yukon-Kuskokwim Delta in western Alaska, adult C. alpina arrive in mid-May, establish territories, pair, nest, and care for young before starting their prebasic molt. After the young fledge, the red-backs shift from inland marsh tundra to coastal habitats, at which time molt intensifies. Feather replacement is complete prior to arrival in late September and October on their north temperate wintering quarters. The result is that breeding and molt are fitted, in sequence, into the summer schedule of events and do not overlap. In the C. alpinu populations at Barrow, Alaska, 10" latitude farther north, breeding and molt overlap and molt proceeds at a faster rate. Such modifications are results of adaptive responses to the shorter summers at the higher latitudes and, to a lesser extent, to the more severe and unpredictable weather there. Results support the hypothesis that the timing of breeding season events in arctic shorebird populations is strongly affected by the length and character of the summer season.

Hulscher, J. B. (1977): The progress of wing-moult of Oystercatchers Haematopus ostralegus at Drachten, Netherlands. Ibis 119: 507 - 512.

Humphrey, P. S & K. C. Parkes (1959): An approach to the study of moult and plumages. Auk 76: 1 - 31.

[CP]: This paper introduces one American, often used convention, separating between "juvenal", "basic" (winter) and "alternate" (breeding) plumages. The adult plumage is "definitive basic" and "definitive alternate".

Humphrey, P. S & K. C. Parkes (1963): Comments on the study of plumage succession. Auk 80: 496 - 503.

Hyytiä, K. & P. Vikberg (1973): Autumn migration and moult of the Spotted Flycatcher Muscicapa striata and the Pied Flycatcher Ficedula hypoleuca at the Signilskär bird station. Orn. Fenn. 50: 134 - 143.


Imboden, C. (1973): Überzählige Handschwinge am Flügel einer Türkentaube Streptopelia decaocto. Orn. Beob. 70: 19 - 26.

1. The right wing of a Collared Dove held in captivity has 11 functional primaries, whereas the left wing shows the normal number of 10.
2. The additional feather is inserted on the metacarpus where there are normally 6 primaries. All these 7 metacarpal primaries are regularly distributed over the metacarpus and are of normal size.
3. On the abnormal wing there is also one more greater and one more under wing-covert, but the number of the median wing-coverts is the same on both wings.
4. (...)On the left and right wing the beginning and the end of the moult are synchronous, but the right wing moults one more feather than the left one during the same time. (...)

Ingolfsson, A. (1970): The moult of remiges and rectrices of Great Black-backed Gulls Larus marinus and Glaucous Gulls L. hyperboreus in Iceland. Ibis 112: 83 - 92.

The moult of primaries, secondaries, and rectrices in two closely-related gulls, the Great Black-backed Gull, Larus marinus and the Glaucous Gull L. hyperboreus, was studied in Iceland. Both gulls moult their primaries in an extremely regular sequence, starting with the 1st (innermost) and ending with the 10th (outermost) feather. Usually two, less often one or three, primaries are growing per wing during the primary moult, which lasts for about six or seven months. Growing primaries were estimated to lengthen on the average by 8.7 mm per day in marinus and 7.8 mm per day in hyperboreus.
The secondaries, usually 24 in number, are shed in two moult waves, one starting with the innermost feather soon after the start of the primary moult and then progressing slowly outwards, the other beginning with the outermost secondary after the primary moult is about half completed and tyhe progressing rapidly inwards. The moult is completed just before the end of the primary moult as the two moult waves meet at about the 16th secondary. There are no marked differences between the two gulls in the moult of secondaries.
The moult of rectrices shows large variations in both species, some feathers being much more irregular than others in their time of shedding. In both species, indications of an obscured centrifugal pattern of replacement are seen, although the 5th (next to the outermost) rectrix is usually the last one to be shed. Significant differences were observed between the two species in the degree of regularity of shedding of some feathers and in the average position in the moulting sequence of others. The moult of rectrices starts soon after the moult of primaries is half completed. The feathers are then shed in rapid succession, and the moult is completed some time before the end of the primary moult.
The need for good powers of flight at all times is undoubtedly the reason for the protracted primary moult. This in turn causes the moult to start early, in adults sometimes before the eggs are laid; immatures moult even earlier than this. The rectrix moult and the main part of the secondary moult do not begin in adults until the young have fledged, but then progress very rapidly. Presumably, the loss of some of these feathers would impair the flying ability to an extent sufficient to make it difficult for the gulls to care for their young, while the rapid moult is necessary in order for the replacement of these feathers to be completed by the time the primary moult is over.

Insley, H., L. Young & B. Dudley (1980): Primary moult in the Collared Dove. Bird Study 27: 101 - 107.

Analysis of 1,211 moult cards for Collared Doves, collected between 1964 and 1978, showed that all have a complete annual moult and that some young birds may be in moult twice during a single calendar year. Data collected at Twyford, Hampshire, over a 15 month period in 1977 and 1978 showed that adult doves had mean starting and completion dates for primary moult of 16 June and 30 August (=75 day period), thus averaging 0.67 units per day. The BTO data for adults were probably less accurate because of the inclusion of some incorrectly aged juveniles (which had the effect of extending the moult period); these gave mean dates of 31 May and 7 October, a period of 129 days and a rate of 0.39 units per day. The mean dates for juveniles at Twyford and from the BTO data corresponded much better, although Twyford data were imbalanced by a gap from mid-August to early October. Mean starting dates were 5 July and 3 July and for completion were 9 December and 22 February (Twyford and BTO collections respectively). The mean rate of moult for individual juvenile Collared Doves was calculated from 50 birds retrapped at Twyford over periods of up to 97 days apart; these indicated that 128 days were required for the completion of primary moult, at a rate of 0.39 units per day. It was found that some juveniles enter a period of arrested moult between November and March; the proportion increased from 9.8 % in November to apeak of 51.4 % in January, declining to 14.3 % in March. The period of moult for adults coincides with the breeding season, and there is no evidence that moult imposes physiological stress upon either adults or juveniles.


Jepsen, P. U. (1973): Studies of the moult migration and wing-feather moult of the Goldeneye (Bucephala clangula) in Denmark. Danish Review of Game Biology 8 (6): 1 - 23.

Johnson, C. & C. D. T. Minton (1980): The primary moult of the Dunlin Calidris alpina at the Wash. Orn. Scand. 11: 190 - 195.

The primary moult of the Dunlin Calidris alpina at the Wash, East England, is described. A new statistical method for estimating the duration of the moulting period of a population is developed and individual moult duration is also considered, using population parameters and data from retraps. Differences were found in the timing and duration of moult season in different years. Within any one year late moulting individuals moulted more rapidly than early birds. The adaptive significance of these differences within the annual cycle is discussed.

Jukema, J. (1986): De voorjarsrui bij Goudplevieren Pluvialis apricaria in Friesland. Limosa 59: 111 - 113.


Kania, W. (1990): The primary moult of breeding Dunlins Calidris alpina in the central Taimyr in 1989. Wader Study Group Bull. 60: 17 - 19.

Kasparek, M. (1976): Über Populationsunterschiede im Mauserverhalten der Rauchschwalbe (Hirundo rustica). Vogelwelt 97: 121 - 132.

Kasparek, M. (1980): Jahreszeitliche Aspekte der Mauser der Rohrammer Emberiza schoeniclus L. Ökol. Vögel 2: 1 - 36.

Kasparek, M. (1981): Die Mauser der Singvögel Europas - ein Feldführer. Dachverband Deutscher Avifaunisten.

Keast, A. (1968): Moult in birds of the Australian dry country relative to rainfall and breeding. J. Zool. Lond. 155: 185 - 200.

King, J. R. (1972): Postnuptial and postjuvenal moult in Rufous Collared Sparrows in northwestern Argentina. Condor 74: 5 - 16.

The molts of Zonotrichia capensis were studied in populations inhabiting three biotopes ( chaco woodland, transition-zone savannah, and mountain semi-desert) at 27"s latitude in northwestern Argentina. There was no prenuptial molt in any population. Postnuptial molt required about 80 days in individuals in all populations, and 110-120 days in the populations. Molt was begun late in the breeding season in the lowland populations but was then suspended temporarily in about one-quarter of the individuals, apparently during the terminal breeding effort of the year. There was no suspended molt in the mountain population.
Postjuvenal molt required about 50 days in individuals and led to a plumage indistinguishable from that of adults. Replacement of the "tertials" ( S7-9) was common, but replacement of the central rectrices was not. Postjuvenal molt began synchronously in the majority of immature birds, regardless of age. Some individuals were four to five months old at the start of postjuvenal molt, while others were only six to seven weeks old (reckoned from the beginning of incubation). Patterns of molt are summarized and compared for Zonotrichia spp. in Costa Rica, Colombia, Peru, northern Argentina, and North America. Some hypotheses are offered concerning evolutionary trends in molt patterns.

King, J. R. (1994): Initiation of remige moult by first-year Great Reed Warblers Acrocephalus a. arundinaceus in Europe. Ring. & Migr. 15: 123 - 126.

Koch, H. J. & A. F. De Bont (1944): Influence de la mue sur l'intensité de métabolisme cher le Pinson, Fringilla coelebs coelebs L.. Ann. Soc. Zool. Belg. 75: 81-86.

Koopman, K. (1986): Primary moult and weight changes of Ruffs in The Netherlands in relation to migration. Ardea 74: 69 - 77.

This paper gives an analysis of primary moult in relation to body-weight of 823 male and 304 female Ruffs caught in The Netherlands. In late June the first males start primary moult. Females start in early July. Almost all males present in The Netherlands in late summer start primary moult but only part of the females do so. This fits with observations on Ruffs arriving in their African wintering grounds in early autumn. Birds wghich complete the moult of their primaries in The Netherlands have finished by early October (males) or mid October (females) at the latest. The completion of primary moult takes about 72 days in both sexes. However, from early August onwards many Ruffs suspend the moult when the growth of five new primaries has been completed. Ruffs in active moult have lowest weights: 180-190 g in males and 105-115 g in females. When primary moult is likely to be suspended (only one pinion growing), body-weight increases. Birds in suspended moult are heaviest: males weigh 2590 g and females 145 g.
It is argued that Ruffs, which mainly winter in West Africa, reach their winter quarters in a non-stop flight of approximately 4500 km. After early September (females) or mid September (males), Ruffs may be unable to deposit sufficient fat reserves for a non-stop flight to West Africa. As Ruffs would be unable to complete primary moult before these dates, the moult is often suspended. The minority of males that do complete the primary moult in The Netherlands, probably also spends the winter here. They have sufficiently large reserves of fat to move to France in one flight when a sudden period of frost would force them to do so.

Koopman, K. (1996): The partial pre-breeding primary moult in Common Terns Sterna hirundo.. Ring. & Migr. 17: 11 - 14.

In this paper aspects of twice-moulted primaries in Common Terns as described by Craik (1994) for Scotland are compared with those for The Netherlands. Similarity between the two populations is great. However the conclusions drawn are different. In the Scotland study it was concluded that feeding conditions in the winterquarters determine the number of twice-moulted primaries. Basing itself on birds of known age and on retraps in following years the study in the Netherlands states that the variation is a natural one, with a weak tendency of males and older birds to have slightly higher moulting scores. The second moult of the innermost primaries possibly functionally assures a rather fresh wing for the breeding season. It probably does not function as a distinguishing feature with the Arctic Tern, since, the Arctic Tern excepted, all terns breeding in Europe moult their innermost primaries twice.(...)

de Kroon, G. H. J. (1986): Moult and feather growth in the Water Rail Rallus aquaticus on the Waddensea Island of Vlieland. Ring. & Migr. 7: 57 - 59.

From examination of 452 juvenile and 108 adult Water Rails it was established that moult mainly occurs in June, July and August. Flight feathers were shed simultaneously. Contour feather replacement continued until December.


Lehikoinen, E. & M. Laukonen (1977): Frequency of tail-moult in immature Greenfinches. Orn. Fenn. 54: 133 - 134.

Lehikoinen, E. & P. Niemilä (1984): (Transl.) Moult study on passerines. Lintumies 12: 33 - 44.

Leisler, B (1972): Die Mauser des Mariskensängers (Acrocephalus melanopogon) als ökologisches Problem. J. Orn. 113: 191 - 206.

1. The moustached warbler goes through a complete postnuptial moult, a complete postjuvenal moult and possibly a partial prenuptial moult. It is the only Acrocephalus warbler, where juveniles moult completely in the same calendar year in which they are born.
2. Birds of the Nuesiedler See-area renew the entire plumage before migration. As a rule, postjuvenal moult lasts from mid August to the end of September.
3. In caged birds the moult of primaries lasts 46 to 49 days.
4. In general pattern and sequence of wing moult follow the typical passerine scheme.(...)
6. Some data suggest that postjuvenal moult is accelerated in birds born late in the year.
7. The pre-migratory moult of the birds of the Neusiedler See-area is interpreted in terms of an ecological adaptation to their year round distribution. Reasons against a complete moult of these birds in the northernmost parts of the species winterquarters are poor food conditions, relatively short day lengths and unstable climatic conditions. A relatively short duration of moult, the species ecological feeding niche and a departure relatively late in the season favour on the other hand the moult in the breeding area. As a result of the moustached warbler's skulking habit in harsh coppice of Cyperaceae, feather wear is severe during the whole year.
8. In the literature specimens of the moustached warbler undergoing or finishing the complete moult are mentioned only from the end of October and the beginning of November and only on the southernmost part of the species specific range of winterquarters which are situated near to these.
9. (...)As yet it can not be decided as to whether the differences in the onset of moult are due to individual or to geographical variation of the species.

Leisler, B (1977): Observations on the moult of the Great Reed Warbler Acrocephalus arundinaceus. Ibis 119: 204 - 206.

Lindström, Å., Visser, G. H. & S. Daan (1993): The energetic cost of feather synthesis is proportional to basal metabolic rate. Phys. Z. 66: 490-510.

(...)The cost of feather synthesis was estimated at 836 and 683 kJ per (g dry feathers) in the bluethroats and redpolls, respectively. (...) The Cf for a species with known BMRm may be estimated from the equation Cf = 270BMRm. Species with a relatively high BMRm for their size also have a relatively high Cf. The tight association of Cf and BMRm between species, and the low efficiency values of feather synthesis, suggests that feather production costs include more than the costs for keratin synthesis: they mainly consist of costs of maintaining tissues necessary for feather production.

Lundberg, P. & L.-O. Eriksson. (1984): Postjuvenile moult in two northern Scandinavian Starling Sturnus vulgaris populations - evidence for difference in the circannual time-program. Orn. Scand. 15: 105 - 109.

The postjuvenile moult pattern of captive individuals from two Scandinavian (Umeå 63°20'N and Andøya 69°16'N) Starling populations was studied. The birds were hand-raised and kept under constant environmental conditions which made it possible to investigate possible endogenous differences in the circannual time-program of the moult. Birds from Umeå went through a complete moult significantly faster than did birds from Andøya. This was particularly pronounced when considering the remiges. Birds from Andøya also showed a greater temporal segregation of the moult of different parts of the plumage. The results show that there is a difference in the endogenous circannual rhythm of moult between birds from the two populations. This is in good agreement with the different natural annual cycles birds from these two populations show, i.e. birds from Andøya are non-migratory and lay two clutches per year, whereas birds from Umeå are completely migratory and lay only a single clutch per year.


Maragna P. & M. Pesente. (1997): Complete moult confirmed in a Great Reed Warbler Acrocephalus arundinaceus population breeding in northern Italy. Ring. & Migr. 18: 57 - 58.

(from text) At Palude Busatello (45.07 N, 11.51 E), ten adults showed active primary moult. "There was no evidence in first-year birds of active or interrupted moult in any wing or tail feather tract."

Marks, J. S. (1993): Molt of Bristle-thighed Curlews in the Northwestern Hawaiian Islands. Auk 110: 573 - 587.

I studied molt of Bristle-thighed Curlews (Numenius tahitiensis) on Laysan Island in the Northwestern Hawaiian Islands from 1988 to 1991. Adult curlews underwent a complete prebasic molt between August and December. Duration of primary molt was about 92 days, which is rapid compared with other shorebirds that molt in tropical and southern latitudes. Adults replaced large numbers of primaries and secondaries simultaneously, and about 50 % of the birds became flightless during molt. The prealternate molt began in winter and ended in early spring; it involved the body feathers and variable numbers of rectrices but no remiges. Juveniles molted body feathers and some rectrices during the first autumn and winter but did not replace their juvenal primaries until the spring and summer of their second calendar year. Second-year curlews began replacing their new first basic primaries in late summer, in some cases before the outer juvenal primaries had been dropped. The delayed first prebasic primary molt is probably an adaptation allowing inexperienced birds to devote energy expenditure in their first winter to obtaining food rather than to molting remiges. Because second-year birds remain on the wintering grounds throughout the year and do not prepare for migration, there is no selection against replacing new primaries. Unlike most shorebirds, adult Bristle-thighed Curlews gained mass steadily throughout the autumn and winter. Their rapid prebasic molt in autumn may be an adaptation to allow the birds ample time to build up fat reserves during winter. I suggest that the absence of rich intertidal feeding areas and the frequency of winter storms make it difficult for curlews to take on large fat stores in short time periods as do species that winter on continental coasts. The lack of predators and the small size of remote oceanic islands obviate the need for curlews to maintain peak flight efficiency, allowing birds to become flightless during molt in autumn and to carry increasingly large fat stores throughout the winter.

Massey, B. W. & J. L. Atwood (1978): Plumages of the Least Tern. Bird-Banding 49: 360 - 371.

Massi, A. & F. Spina (1996): Wing moult of the Bearded Reedling (Panurus biarmicus) in Northern Italy. Vogelwarte 38: 180 - 187.

Between 1985-1992 a total of 3,153 Bearded Reedling moult cards have been collected in Val Campotto (Ferrara, Northern Italy). Aim of the study was to analyse the duration and dynamics of wing moult, also with respect to age and sex classes. A regular replacement of primaries has been observed, without any marked change in speed during its progression. Primaries are shed starting from P1, ending with P10 (descendant moult); secondaries moult from S1 to S6 (ascendant moult), and the sequence of tertiary moult is T2, T3 and finally T1. The onset of moult for secondaries, tertials and rectrices takes place once P4 has been shed. A total of 186 retraps of moulting birds has been used to calculate the duration of moult, which resulted in an average estimate of 66.8 10.9 days, being longer than that recorded for more northern populations. The mean starting date of primary moult is July 22nd, ranging from May 31st to August 14th. No variations in moult duration and sequence have been detected when taking into account the different age and sex classes.

Mathiasson, S. (1973): A moulting population of non-breeding Mute Swans with special reference to flight-feather moult, feeding ecology and habitat selection. Wildfowl 24: 43 - 53.

McCanch, N. (1998): Swallow Hirundo rustica with unmoulted greater coverts in spring. Ring. & Migr. 19: 30.

Mead, C. J (1975): Juvenile hirundines starting primary moult in Europe. Ring. & Migr. 1: 57.

Mead, C. J (1980): Sand Martins moulting primaries in Britain. Bird Study 27: 51 - 53.

Mead, C. J. & B. R. Watmough (1976): Suspended moult of trans-Saharan migrants in Iberia. Bird Study 23: 187 - 196.

During autumn ringing operations in Iberia all trans-Saharan migrants trapped (c. 6,5000) were examined for evidence of suspended moult of primaries. Only a minority of those caught were adult birds but 94 individuals of 16 species were found to have suspended their moult for the autumn migration. Two species, Savi's Warbler and Spotted Flycatcher, showed a radical departure from the 'normal' sequence, but the other 14 species behaved as follows:
(i) Most individuals migrating with suspended primary moult: Temminck's Stint, Turtle Dove, Scops Owl, Red-necked Nightjar, Bee-eater, House Martin, Woodchat
(ii)Most renewing before migrating, but some retaining old feathers (usually only a few): Orphean Warbler, Whitethroat, Willow Warbler, Pied Flycatcher.
(iii) Most migrating with old plumage, but a few renewing some feathers (usually only a few): Golden Oriole, Garden Warbler, Bonelli's Warbler.
The aberration in the case of the Spotted Flycatcher was that the well-known reversal of the moult sequence could be detected; otherwise this species could be classed as category (iii). Adult Savi's Warblers may migrate with completely old, completely or partly renewed flight feathers. At least six Savi's Warblers had started their primary moult in the middle of the tract rather than with the primary nearest the body.(...)

Mester, H. & W. Prünte (1982): Die "sektorale" postjuvenile Handschwingenmauser der Carduelinen in Südeuropa. J. Orn. 123: 381 - 399.

The postjuvenal moult shown by Goldfinches which inhabit the Balearic Islands is quite exceptional. The fact that these birds acquire some new remiges at an age of only a few months is already outstanding for a member of the family Fringillidae. More remarkable is the mode of this moult, however, as it does not follow the usual passerine pattern at all. Not only the three inner secondaries are changed, these forming a unit, but in most cases a number of primaries as well, namely those which cover and protect the outermost part of the wing when it is folded up.
There is strong evidence that just the same geographical variation of the juvenile moult pattern occurs in the Linnet and Serin, too, and that in some Greenfinches this variation even reaches Central-Europe. The process can start at a focus between P1 and P6. There also exists individual variation insofar as some juvenile birds replace only the feather just mentioned whilst others even replace a group of seven or eight primaries. In spite of this the adaptive significance of the pattern is evident: The flight-feathers shed are exposed to worst mechanical and especially solar wear just as the tertials are. In this respect insolation, reaching its peak on the islands in July, seems to be a very important factor and all in all more harmful to the remiges than rainy weather. Moreover all greater coverts and at least in most cases the bastard wing are changed when first winter plumage is attained. The moult of rectrices is not exactly centrifugal in most instances.
Emphasis is laid on the finding that populations of a species which are exposed to weather-conditions, that are so much destructive to feathers, developed an addittional moult. So the wing tip, normally especially subjected to wear, gets a new shelter against rain and sun light. The function of the primaries, which are replaced by new ones in late summer, is merely to act as a kind of protective screen in the same way as tertials are.
As has been mentioned before, this mode of moulting flight feathers is a unique peculiarity insofar as there exists no constant centre. With great variability it can be shifted in direction of the wing tip until P6, which is part of the process in all cases, whilst on the other hand the outermost primary (P9) is never shed. In this respect the moult remains incomplete. Statistically in both samples, in the Greenfinches and in the Goldfinches as well, the localisation of the focus is approaching a binomial distribution.

Michener, H & J. R. Michener (1940): The moult of House Finches of the Pasadena region, California. Condor 40: 140 - 153.

This work is based on observations over a period of four years on the House Finches handled at a banding station banding from 2000 to 2500 House Finches per year and having many times that many repeat captures.
Subsequent to the postjuvenal molt the annual autumnal or postnuptial molt is the only molt of the species.
The postnuptial molt is complete. It begins with the falling of primary 1 and ends with the completion of secondary 6 ; the time required is approximately 105 days, varying from 90 to 120 for different individuals. This molt has been observed in prog- ress from May 15 to November 10.
There is a general sequence of molt of the feathers in a tract and of the tracts rela- tive to each other, but there are also many variations from this sequence. The extent of the postjuvenal molt was studied by notching all flight feathers on the right side of over 2000 juveniles.
The postjuvenal molt of the early-hatched young was found to be the same as the postnuptial molt of the adults while the latest-hatched young shortened the molt period by beginning the molt soon after leaving the nest, by retaining all juvenal flight feathers, and by molting all the other tracts more nearly at the same time. Those hatched at intermediate times varied the time of beginning the molt relative to age, molted a varying number of the flight feathers, and compressed the molt of the other tracts proportionately to the time available between the date of hatching and early November. Although there are individual differences, it is generally true that the postjuvenal male plumages are red on the head, throat and rump in the earliest-hatched birds and in those hatched successively later, these areas are less and less red and then less and less yellow until the latest hatched show only a dull yellowish tinge.

Mitchell, P. I., Scott, I. & P. R. Evans (1996): Is moult an idicator of population structure in Redshank Tringa totanus during autumn? Ring. & Migr. 17: 93 - 100.

Redshank from Iceland (Tringa totanus robusta) and from Britain (T. t. brittanica) spend the non-breeding season and moult period around the British coast. Only biometric data distinguish the two races. Furness & Baillie (1981) proposed that the proportions of the two races present on the Wash (or any other site) in late summer could be estimated during the period of primary moult, based on the premise that britannica commences moult earlier than robusta. We tested Furness & Baillie's approach on a moulting mixed-race assemblage of Redshank at Teesmouth, NE England. Using a model of speed of primary moult derived from captive Redshank we show that there is no consistent difference in starting date or duration of primary moult between the two races. We conclude that the proportions of two races of Redshank in Britain during the moult period cannot be determined using moult score.

Morrison, R. I. G. (1976): Moult of the Purple Sandpiper Calidris maritima in Iceland. Ibis 118: 237 - 246.

The autumn moult pattern of adult Purple Sandpipers Calidris maritima in Iceland is described. The duration of the moult was estimated to be 5 1/2 - 7 weeks (c. 40 - 50 days). Females generally started moult before males and moult did not appear to overlap breeding. Information from other areas is reviewed. A mechanism by which the duration of moult is shortened amongst various species is by an increase in the number of feathers growing concurrently during the moult. Likely reasons for the placing of the moult in the annual cycle of the Purple Sandpiper are discussed, and appear to be related to the exceptionally northerly winter distribution of the species.

Morton, M. L., J. R. King & D. S. Farner (1969): Postnuptial and post-juvenal moult in White-crowned Sparrows in central Alaska. Condor 71: 376 - 385.

The phenology of the postnuptial and postjuvenal molts in the White-crowned Sparrow was ascertained from examination of 500 free-living birds near Fairbanks, Alaska, during 1962. At the same time, the pattern of postnuptial molt was studied in 17 captive adults exposed to natural temperature and photoperiod. There were no conspicuous differences between free-living and captive birds, and the data from these groups were combined in the general analysis.
Postnuptial molt began in the population in the first week of July and ended by 25 August. In individual birds, the complete postnuptial molt required about 48 days. Renewal of the primary remiges required 37 days in the average individual; all nine primaries grew concurrently during the last 11 days of this period. Renewal of the secondary remiges required 26 days in the average individual, and a maximum of six secondaries grew concurrently. The renewal of the rectrices required about 25 days, all 12 growing concurrently. Molt of the contour plumage usually began with the shedding of the first primary and continued for about one week after the completion of growth in the flight feathers.
Postjuvenal molt began in the population in the last few days of June and was essentially ended by 20 August. The duration of molt in an individual bird was about 34 days.
Midsummer flock-formation began in immature White-crowned Sparrows in the third week of July, when the postjuvenal molt was well advanced. Adults initially remained apart, but began to join the immatures in flocks in the first weel of August during the declining phase of the postnuptial molt. Migration, or premigratory gathering, of the local population began soon after 15 August, and White-crowned Sparrows were absent from the region after 1 September. A few of them must have begun the southward movement before body molt was finished.
The summer molt of White-crowned Sparrows in central Alaska is characterized by its brevity, intensity of flight-feather growth, and conspicuous precision in timing within the population. At present these attributes appear to be quantitatively unique, although future studies of small migrant songbirds at high latitudes may reveal that they are part of a common pattern of adaptation.

Morton, M. L. & D. E. Welton (1973): Postnuptial moult and its relation to reproductive cycle and body-weight in mountain White-crowned Sparrows (Zonotrichia leucophrys oriantha). Condor 75: 184 - 189.

The timing and progress of postnuptial molt was measured in wild and captive Zonotrichia leucophrys oriantha on their central Sierra Nevada breeding ground. Molt began with the loss of the first primary and with contour feathers of the dorsal cervical region. The replacement of new flight feathers proceeded regularly and their total daily increment of growth was nearly constant. The period of feather renewal lasted for an average of 48.8 days.
Evidently the feeding of young and postnuptial molt are incompatible events energetically because they seldom occurred simultaneously in a given adult. The nonoverlap of these events affected the onset of molt in mated pairs in that females fed fledglings for a longer period than males and molt began in them about 8 days later than in males. The chronology of molt for the population was also affected by reproductive activities. Molt began about a week earlier in 1968, a dry year, than in 1969 and 1970, wet years more suitable for a prolonged period of reproduction
Due to fat accumulation, body weight increased by about 25 % between the end of reproduction and migration. Fattening and molt were compatible energetically in that about half of the total weight gain occurred during the period of molt and half immediately following the completion of molt.
Compared to a previously studied migratory conspecific, Z. l. gambelii, feather growth proceeded more slowly in Z. l. oriantha in the major categories of flight feathers, but the total duration of postnuptial molt was the same. different metabolic preparations for autumnal migration are employed by these races in that Z. l. gambeliifattens after molt is completed and Z. l. oriantha fattens both during and after molt.


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