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The Moult of The Bullfinch PDF
The Moult of The Bullfinch PDF
The Moult of The Bullfinch PDF
I. NEWTON
INTRODUCTION
Few detailed field studies have yet been carried out on the moult in any bird species.
This paper describes some observations on the moult of the Bullfinch Pyrrhula pyrrhula
nesa (Carduelinae), and its timing in relation to the breeding season and food-supply
in the woodland of Wytham Estate, near Oxford (58" 80'N.). The Bullfinch is a common
resident in the area, mainly in the woodland, but also in gardens, orchards, and hedge-
rows. It nests in dense shrubs and hedges, and feeds for most of the year mainly on
various seeds, and in spring on buds, including those of various fruit trees (Newton
1964, 1964 a).
Details of moult and weight were obtained from 133 individuals in 1962, from 283
in 1963, and from 247 in 1964; many of these were handled more than once during a
single moult.* They were caught in mist nets placed more or less at random in the
woodland or at feeding places such as patches of Meadowsweet Filipendula ulmaria and
various docks Rumex spp. Netting was restricted to two areas, the Great Wood and
Marley, which have been described by Southern (1954) and Gibb (1954) respectively.
The Great Wood comprises some 400 acres of deciduous trees, dominated by Oak
Quercus robur, Ash Fraxinus excelsior and Sycamore Acer pseudo-platanus, and about
150 acres of conifers. Marley is an open scrubby area of 66 acres, which is dominated
by Oak and Hazel Corylus avellana, with large patches of hawthorns Crataegus spp. and
Elder Sambucus niger.
Wytham Bullfinches started laying each year in early May, soon after fresh seeds
first became available. The laying season lasted until mid-July in 1963 and 1964, but
was prolonged to early September in 1962 when food was especially plentiful in late
summer. After breeding the adults underwent a complete moult, and at the same time
the juveniles underwent a partial moult. The term " moult )' is here taken to cover
both the shedding and the renewal of feathers.
* Full details of moult were not obtained from every bird, and if time was short details only of
wing-moult (or sometimes of wing- and tail-moult) were taken from the adults. This accounts
for the different totals in some of the tables.
42 I. NEWTON : THE MOULT OF THE BULLFINCH P Y R R H U L A P Y R R H U L A IBIS108
are retained until the next moult about a year later. Birds in first-year plumage can be
distinguished by some of the retained juvenile feathers, until at 11-15 months old, the
next moult is so advanced that their plumage is indistinguishable from that of older
birds. All subsequent moults are complete post-nuptial moults and occur at approxi-
mately annual intervals. There are no pre-nuptial moults in the Bullfinch.
In an adult Bullfinch, just prior to moult, the feathers are usually worn, faded and
soiled, and there are often gaps in the plumage where lost feathers have not been replaced.
The fading of the black flight and tail feathers to various shades of brown is presumably
caused by the disintegration of some of the cornified cell-layers of the feather (Miller
1928). Fading was marked in birds moulting late, and was most pronounced in birds
in first-year plumage in which the flight and tail feathers, acquired as nestlings, could
have been up to 15 months old before they were replaced.
The adult moult in the Bullfinch lasts 10-12 weeks and the post-juvenile moult
7-9 weeks. In adults replacement of the body plumage rarely spans the entire moult
period, but averages about as long as the post-juvenile moult. The moult begins in the
adults with the primaries and in the juveniles with the lesser-coverts. The latter are
often the first body feathers to be moulted in adult Bullfinches. In this respect the
Bullfinch differs from most other passerines in which body moult begins first on the
breast, back, scapulars, and head, and only then in the lesser-coverts (Dwight 1900).
Each primary-covert is shed with its corresponding primary, and has usually reached full
length by the time the primary is half grown. The greater-coverts are shed more or less simul-
taneously with each other, or in quick succession from the innermost outwards; sometimes
however, the inner ones are half grown before the outer four or five are dropped. Replacement of
the lesser-coverts begins near the body and spreads first towards the carpal joint and then to the
median-coverts, which are moulted all more or less together; at the same time the carpal-covert
and the feathers of the alula are replaced. On the underside of the wing the coverts are softer and
appear more sparsely distributed; the lesser- and median-coverts are moulted more or less simul-
taneously, and the under greater- and under primary-coverts are moulted in rapid succession
from the carpal joint outwards.
shorter-and are replaced from the central pair outwards. The ten upper coverts were often shed
from the outermost inwards, but this pattern was sometimes modified by the temporary retention
of any of them, and in some individuals no orderly sequence of replacement was discerned. The
under tail-coverts also tend to be replaced centripetally, though variation was even more frequent
than in the upper coverts. The feathers circling the anus are all moulted together.
secondary score
60. -:
3=
50- e
0 .
*d
40- C.
4-
30-
:**
20-
0 .
**
10- *. I*
** a****
:**-**
**
~ - o o o o o * w a * * w * mo e ***:*
1
1. The moult of the primary and secondary feathers in the Bullfinch Pyrrhula pyrrhula.
TABLE
Mean Mean Mean Me
No. of birds with the followin8 feathers in growth:- no: total no. n
growing no: filll- fu
No. o f Primaries Secondaries second- growing Fowp gr
aria renuges primma sec
z e x z e d 1 2 3 4 5 6 7 8 9 1 2 3 4 5 ar
1-10 44 4420 J 1.6
1-20 25 22 25 20 I 3 .O 0.08
1-30 21 9 17 25 19 4 2.8 1.1
1 4 11 1 3 13 11 16 1 4 0.2 3.2 2.4
1-50 14 4 12 14 10 12 1 1*o 3.8 2.9
1-60 24 3 1 1 6 2 4 9 1 2 2 4 1.4 4.2 4.2 0
1-70 12 1 9 1 1 9 2 6 7 2 2 2 1*8 4.6 5.2 0
1-80 8 7 8 8 7 7 4 1 2.5 5.4 6.4 1
1-89 31 2 15 31 IS 17 23 24 4.5 7.3 3
Tail score
50-
. :
. . . 0 .
Oi 0
. :*:
%
30-
20-
10-
10 20 30 40 50 60 70 80 90
Primary score
2. Tail score in relation to primary score in moulting adult Bullfinches Pyrrhula pyrrhula.
FIGURE
Tail score (0-60) and primary score (0-90), which are calculated from the growth stage of
individual feathers (see text), indicate the state of moult in the rectrices and primaries
respectively.
1966 I. NEWTON : THE MOULT OF THE BULLFINCH P Y R R E U L A PYRRHULA 47
The growth rate of individual rectrices could be worked out for only three birds.
In the first, two feathers had each gained one point in seven days (2 mm. per day); in
the second, four feathers had each gained one point in eight days (1.8 mm. per day);
and in the third, five feathers had each gained one point in four days (3.5 mm. per day).
These figures show considerable individual variation but that the growth rate of the
rectrices can be as fast as that of the remiges, which averaged 3-4 mm. per day (see later).
TABLE
3. The moult of the body feathers, and of the small feathers of the alar and caudal
tracts, of the Bullfinch Pyrrhula pyrrhula in relation to primary moult. The jigures
show the number of b i d examined at each stage of moult which were moulting
particular feather tracts. For explanalion of stages of moult see text.
Stage of primary moult
1 2 3 4 5 6 7 8 9
Total number of birds
examined : 40 24 20 14 13 20 10 8 31
Feather group
Upper greater-coverts 4 14 14 8 1
median-coverts 13 9 8 10 5
lesser-coverts 9 14 17 11 7 11 4
Under greater-converts 4 5 6 13
median-coverts 4 8 7 6 2
lesser-coverts 6 7 13 8 5 2
Tertials 12 9 12 16 9 2
Alula 1 2 1 1 2 5 5
Scapulars (shoulder tract) 6 15 16 11 8 10 6 2 13
Axillaries 2 10 14 6 2
Ventral tract 8 16 16 14 13 20 10 8 21
Dorsal tract 9 13 17 14 13 20 10 5 15
Thigh tract 3 9 8 11 17 9 8 27
Leg 5 10 9 13 17 4 4 6
Head 4 9 13 i3 20 8 5 6
Upper tail-coverts 2 12 11 17 10 5 12
Under tail-coverts 6 13 13 17 9 4 7
moult. However, there was considerable variation and after stage 6 most birds were
clothed almost entirely with new and growing feathers, and appeared fully moulted
when seen in the field. Thus the most intense renewal of body feathers occurred at
stages 3-6, when almost every tract and group of feathers was in heavy moult. But full
replacement of the plumage of the dorsal and ventral tracts, two of the largest tracts on
the body, lasted almost the entire moult period. Feathers on the belly and head were
the last to be moulted.
1964
Samnle she* 24 27 20 17
Re-&&Gequation P=1.22+1.095D P= -12.33+1.18D P= -4.29+1.123D P- - 10.11+ 1.133D
Rate of increase in primary
score (points per day) 1.095*0.19 1.18 & 0.059 1.123 f0.088 1.133j~0.14
Duration of moult (days) 82.17 76.26 80.15 79.43
Start of moult
Linearity test
Aug. 1 i14.9
Fi =243 =:
Aug. 10f7.3
F 1 0.589
Aug. 9;tg.l
F1:=2.24
Aug. 9 k 8 . 4
P1g=0.88
Note. Includes retraps, but unusually early or late individuals were omitted from analysis since these can bias the estimates
obtained, especially if the samples are small and the data are not collected regularly throughout the moult period.
by plotting the primary scores of many individuals against date, show little trace of a
sigmoid shape, probably because in the Bullfinch the period when fewer than average
feathers are growing is short compared with the spread in the date of onset of moult in
the whole population.
It is worth mentioning that since the different age and sex groups began moult, on
average, on different dates, if the data for all birds are combined, the spread in the start
of moult is then so great that a regression analysis gives a value for the overall rate of
moult which is too small. Such an analysis can therefore be used to estimate the rate
of moult satisfactorily only when the time of moult in a population is fairly well
synchronised-otherwise the regression analysis gives niisleading results. (For further
discussion of the use of regression analyses on moult data see Evans, in press.)
The mean rate of increase in primary score determined by the regression analyses
varied from about 1.10 to 1.23 points per day, and the duration of moult from 73 to 82
days (Table 5). In both years the yearling males examined started moult on average a
few days earlier than the other groups, probably because some of them were non-breeding
birds (Newton 1964).
Information on the rate of moult in individual birds is available for 21 birds which
were handled more than once during a moult, and at intervals up to 68 days. The rate
of increase in primary score in some of these birds is shown in Fig. 4 in which each line
refers to one bird, and its slope is the rate of moult in points per day. The mean rate
of increase in primary score was 1.30 & 0.2 points per day, which is consistent with the
estimates obtained from the regression analyses. Recorded over shorter periods, the
slowest rate of progress was 0.4 points per day over 14 days, and the fastest six times as
great, 2.5 points per day over 15 days.
Variations in the rate of primary moult in three individuals caught more than twice
during moult are shown in Table 5. The fastest rate of primary moult recorded in
bird B1, was almost three times faster than the slowest. In B2, the rate at which the
primary score increased varied from 1.33 to 2.25 points per day; while B3 maintained
about the same rate throughout. The greatest deviations from the mean rate of moult
D VOL. 108
50 I. NEWI'ON : THE MOULT OF THE BULLFINCH P Y R R H O L A P Y R R H U L A IBIS108
Primary score
I30
0I
. .
O .
0
0
1962
10- 0
30
0
.. . .. . . 0
&
. O
1963
-
8.. to
10- 0 m p
OO r:. 00
0 0.
Primary score
.:!
0
..
0
. 0
30-
0 0
0
. O .
*:.
. . ..
0
0 . 0 0 1 9 6 4
. *
10-
:. 0 .
8
0
0
00
.
1966 I. NEWTON : THE MOULT OF THE BULLFINCH PYRRHULA PYRRHULA 51
appeared in birds in which the interval between successive observations was short.
Thus the longest lines in Fig. 4 run nearly parallel, indicating that the overall rate of
moult was about the same in all these birds, even though the rate may have varied
widely at different stages of moult.
Primary score
TABLE
5. Fluctuations in the rate of moult in three adult BullJinches Pyrrhula pyrrhula.
Rate of increase in
Date of moult score in points
Bird capture Score per day
B1 1 Aug. 63 4
1-46 (11 days)
12 Aug. 63 20
1.77 (26 days)
7 Sept. 63 66
0.67 (3 days)
10 Sept. 63 68
0.89 (18 days)
28 Sept. 63 84
B2. 15Aug. 63 8
1.43 (7 days)
22 Aug. 63 18
2.25 (16 days)
7 Sept. 63 54
1.33 (9 days)
16 Sept. 63 66
B3 10Aug. 63 8
1.50 (10 days)
20 Aug. 63 23
1.43 (14 days)
3 Sept. 63 43
1.48 (21 days)
24 Sept. 63 74
Note. The number of days between successive observations is given in parenthesis in column 3.
52 I. NEWTON : THE MOULT OF THE BULLFINCH PYR R H U LA P Y R R H U ~ L A IBIS108
Assuming a mean rate of 1.3 & 0.2 points per day, the time taken to moult would be 69
days (+-9 days), plus up to about seven more days in those individuals in which moult of
the secondaries ended after moult of the primaries. Three individuals were caught soon
after the start and again just before the end of moult, and assuming that each primary
grows at a rate of 0.4 points per day (see later), the time taken for the complete moult
in these birds would be 68, 70, and 74 days respectively, estimates which are again
consistent with those obtained from the regression analyses (Table 4).
The duration of moult in adult Bullfinches is similar to that of some other passerines
of temperate regions, such as Steller’s Jay Cyanocitta stelleri (Pitelka 1958). On the
other hand, Lesser Redpolls Curduelis$ummeu cabaret moult in only eight weeks (Evans,
in press) and Chaffinches in only six (Marler 1956). In the House Finch of California
moult lasts about 15 weeks (Michener & Michener 1940), and in the Boat-tailed Grackle
13 weeks in the female, compared with 15 in the male (Selander 1958).
next was dropped; while birds apparently free of young, but with the same moult score,
had up to five primaries growing in each wing and never less than two. Birds at earlier
stages of moult showed no such difference.
TABLE
6. The pattern of wing-moult in adult BullJinches Pyrrhula pyrrhula with dependent
young compared to that in birds apparently free of young.
Individual Score of urimaries:- Total number of
growing primaries
1 2 3 4 5 6 per wing
Primary score 20-22
With young 1 (male) 5 5 - - - - a}l.5
2 (female) 5 4 I - - -
Without young 3 (male) 5 4 2 - - -
4 (male)
5 (female)
3
5
3
3
3
2
2
1
-
-
-
-
:>,
3
Primary score 28-30
With young 6 (male) 5 5 5 - - -
7 (male) 5 5 5 - - - ;}*.3
8 (female) 5 5 4 1 - - 2
Without young 9 (male) 5 5 3 I - -
10 (male) 5 5 2 2 - -
11 (male) 4 3 3 2 2 -
12 (female) 5 4 3 2 1 -
13 (female) 5 5 4 I - -
14 (female) 5 4 3 2 1 -
Note. For explanation of primary score see text.
This apparent slowing in the shedding of primaries in birds with young could have
been directly due to " shortage " of food imposed by the dual strain of moult and
parental care. Although there are insufficient data to compare the growth rates of
individual feathers in birds of each category, " fault bars " were never seen on the
feathers of any Bullfinches examined. These are areas of weakness produced on the
feathers of some species when food is short and appear as pale bars across the vane where
the barbs are deficient in barbules and melanin (Riddle 1908).
To sum up, in the Bullfinch different primaries appear to take about the same time
to complete their growth, though the data on this point are limited. Differences in the
rate of moult between individuals, and at different stages of moult in the same individual,
are due mainly to variations in the numbers of growing feathers involved. In addition,
there is some slight evidence that moult in the Bullfinch may be influenced by other
factors, since birds which were feeding young dropped their primaries more slowly than
those which were not.
25-
15-
10-
5-
I
_ I a
caught at the end of September. For instance, on 24 September 1962, six adults were
caught, one of which had just started to moult (score 6), and another (score 84) must
have started nine or ten weeks earlier. T h e mean date of onset of moult in 1962 was
31 August (S.D. 23 days), compared with 9 August (S.D. 11 days) in 1963 and 11 August
(S.D. 12 days) in 1964. I n all years, the adults which moulted earliest finished in early
October; in 1962 there were moulting adults in the population almost until the end of
December, but only until mid-November in the two following years.
Percentage
in moult 1962 1963
Percentage
in mouIt 1964
100
50
FIGURE
6 . T h e moulting season of Bullfinches Pyrrhulu pyrrhtilu near Oxford in three successive
years,
Continuous line=adults; broken line =juveniles.
1966 I. NEWTON : THE MOULT OF THE BULLFINCH PYRREULA PYRREULA 55
Ten birds were caught during moult in two different years, and most of them moulted
at a different time (with respect to the mean date of moult) in each year (Table 7). These
differences presumably result from variations in the time they finished breeding in
different years (see below).
THE RELATIONSHIP BETWEEN THE END OF BREEDING AND THE ONSET OF MOULT
Breeding and moult are processes of high energy demand, and in almost all species
of temperate regions occur in summer when conditions are favourable (Pitelka 1958).
In many species, including the Bullfinch, the moult of the adults begins as their last
young become independent. In all three years Bullfinches started nesting in early May,
but this initial synchrony was soon lost, mainly through the action of predators (Newton
1964). Presumably it was the subsequent lack of synchrony in breeding activity which
caused the spread in the time of onset of moult in the population, since each pair started
to moult as their last young left the nest. The greater spread in the time of onsetof
moult in 1962 than in the two following years was due to a proportion of pairs having
additional broods begun after the end of July (Newton 1964). Such late breeding
occurred in response to a comparative abundance of food in late summer (see later and
Newton 1964), and the onset of moult was thereby delayed.
TABLE
8. The number of adult BullJinches Pyrrhula pyrrhula at different stages of moult
which were known to have dependent young.
Stage of moult Moult not Stage 1 Stage 2 Stage 3 Stage 4
begun
Total number of Bullfinches caught 44 44 25 27 17
Total number feeding young 16* 15* 5* 5* 0
*Note. Birds were known to have dependent young only if they were seen feeding them, or
if they had food in the throat pouch. The figures, which are based on all adults caught August-
October all years, are therefore minimum figures.
In most Bullfinches parental care and moult tend to be mutually exclusive, but in
all three years many of the adults caught in the early stages of moult still had dependent
young, some of which were seen being fed; while some adults had food in the throat
56 I. NEWTON : THE MOULT OF THE BULLFINCH P Y R R H U L A P Y R R E U L A IBIS 108
pouch when caught. Young Bullfinches are normally fed by their parents for 1-2 weeks
after fledging. The number of adults at different stages of moult which were known to
have dependent young is shown in Table 8, which is based on the data for August to
October in all three years. At least a third of all the birds caught at stage 1 of moult,
and a fifth of those at stages 2 and 3 had dependent young, but none at any later stage.
I n those at stage 3 the moult presumably began before their young had left the nest,
unless the young were dependent for an unusually long period. The overlap between
moult and dependence of young occurred not only in those birds in which the onset
of moult was delayed by late breeding, but also in those starting to moult comparatively
early. The earliest breeding Bullfinch in moult was a male caught on 6 August 1963
(score 4). On five occasions both members of known pairs (caught earlier at their nests)
were caught, and in each case the male’s moult was slightly ahead of the female’s.
Similar slight overlap between parental care and moult has been noted in some
individuals of at least four other passerines of temperate regions, the Northern Water-
thrush Seiurus noveboracensis (Eaton 1957), the Plain Tit Parus inornutus (Dixon 1962),
the Lesser Redpoll Carduelis fiammeu cabaret (Evans, in press) and the Chaffinch
FringiZZu coelebs (Marler 1956). Experimental evidence is available for several species,
such as the Red-winged Blackbird (Wright & Wright 1944) and the White-crowned
Sparrow Zonotrichia Zeucophris (Blanchard 1941), that the moult in the males begins
after the testes have regressed so far that no free sperms are present. In the male Andean
Sparrow, however, the correlation between testis regression and moult is only partial,
though in the female of this species breeding and moult are mutually exclusive (Miller
1961). Some larger species of birds, on the other hand, show a protracted replacement
of the remiges which begins during the breeding season; this applies for instance to the
Accipitridae, which depend on full wing-efficiency for their livelihood (Heinroth 1931).
In addition, many species, such as the Glaucous Gull Lams hyperboreus (Johnston 1961),
breed and moult concurrently during the short arctic summer.
THE POST-JUVENILE MOULT
T h e post-juvenile moult was limited to the body feathers and the small feathers of
the alar and caudal tracts; these feathers comprise about three-quarters of the weight
of plumage replaced by the adults (see later). The sequence of post-juvenile moult was
in most respects similar to the body moult of adults which had no dependent young.
Table 9 shows some of the feathers retained by juvenile male and female Bullfinches
according to the date at which they started moulting. The figures are based on all
juveniles caught in 1962 and 1963 after completing a moult of known date. In addition
to the primaries, secondaries, two outer tertials, rectrices, primary-coverts and alula, all
juveniles retained the alula-covert, the carpal-covert, or both. Many birds moulting
late in the season retained up to five greater-coverts; only those birds which moulted
earliest replaced them all. Late-moulting juveniles were never noticed to have the under
wing-coverts in moult. They were among the last feathers acquired by the juvenile
after leaving the nest, and they too were probably retained by birds moulting late in the
season. Exceptionally a few body feathers were also retained, usually in areas that are
normally late to moult, such as the belly. Table 9 shows that juvenile males always
replaced more plumage than juvenile females moulting at the same time, but the reasons
for this are not known. The retention of the juvenile alula-covert or carpal-covert,
which are edged with brown, distinguished first-year birds from adults, in which they
are edged with grey.
Since only certain feathers are replaced at the post-juvenile moult, presumably such
replacements are necessary to reinforce the most vulnerable parts of the plumage;
although not seriously damaged by the time of post-juvenile moult, they might, if not
replaced, become dangerously worn by the next moult. The strongest and stiffest
1966 I. NEWTON : THE MOULT OF THE BULLFINCH P Y R R H U L A P Y R R H U L A 57
TABLE9. The number of juvenile feathers retained by juvenile Bullfinches Pyrrhula
pyrrhula moulting at different times.
Number of birds which retained the following
feathers :-
Moult began Sex No. of birds alula carpal greater-coverts*
examined coverts coverts 1 2 3 4 5
July Male 3 2 3 0 0 0 0 0
Female 3 3 3 1 1 0 0 0
August Male 34 34 33 4 4 2 0 0
Female 23 22 23 14 14 6 4 1
September Male 25 25 25 18 18 14 8 0
Female 20 20 20 20 20 16 12 4
October Male 8 8 8 7 7 5 3
Female 6 6 6 6 6 6 6 6
* Numbered from the outside inwards,
feathers, the flight and tail feathers and primary-coverts, were always retained; the less
resilient greater- and alula-coverts were renewed in all but late-hatched young; while the
lax body-feathers were replaced by all birds. T h e retention of those feathers most
resistant to wear appears to be fairly general in passerines (Dwight 1900), but there are
exceptions such as the House Sparrow Passer domesticus in which the juveniles replace
all their feathers at the post-juvenile moult.
Bullfinches hatched late in the season generally began to moult within a fortnight
of leaving the nest (see later), when they had only just acquired juvenile plumage, or
even while parts of it, such as the rectrices and under wing-coverts, were still growing.
They would presumably gain nothing by replacing feathers only recently acquired.
The energy saved by the late-hatched young in retaining feathers that are replaced by
older young is, however, negligible, since the feathers involved weigh less than a
hundredth of those that are moulted (see later).
A tendency of late-hatched young to replace fewer juvenile feathers than those
hatched earlier has been noted also in the House Finch (Jlichener & Michener 1940),
Boat-tailed Grackle Cassidix mexicanus prosopidicoZa (Selander 1958), Phainopepla
Phainopepla nitens (Miller 1933), Brown-headed Cowbird Molothrus ater and Red-winged
Blackbird AgeZaius phoeniceus (Selander & Giller 1960).
TABLE
10. The percentage of juvenile Bulljinches Pyrrhula pyrrhula starting to moult in
each ten-day period over three successive years.
Ten-day periods
Total
Year caught July August September October November
3 1 2 3 1 2 3 1 2 3 1 2 3
1962 111 3 9 12 17 15 13 5 10 13 2 1 0 0
1963 187 13 2 9 1 6 1 9 1 6 6 1 0 0 0 0 0 0
1964 164 12 2 6 1 3 2 3 1 3 7 6 0 0 0 0 0 0
In 1962, the onset of moult among the juveniles was spread fairly evenly over the
14 weeks from the end of July to the beginning of November (Table 10) and some were
moulting to nearly the end of December (Fig. 6). In the following two years, the onset
was spread over only nine weeks, from the end of July to the end of September, and
there were no moulting juveniles in the population after about mid-November (Fig. 6).
1966 I . NEWTON : THE MOULT OF THE BULLFINCH PPRRHDLA PYRRHULA 59
In all years there was a period when juveniles that had completed their moult occurred
in the population simultaneously with juveniles in which the moult had not begun.
In 1962 the overlap lasted 6-7 weeks, and in the two later years only about one week.
TABLE
11. The mean wetght and moult periods of dayerent groups of feathers in the
Bulljinch Pyrrhula pyrrhula.
Primary score Mean length Total weight Mean weight
at average of moult (by of feathers of feathers
start and end primary score) replaced replaced per
of moult (mgm.)*
, - unit increase
in primary
score
(mgd
Primaries 1-90 90 220 2.44
Secondaries 46-90 45 96 2.13
Tertials 27-69 43 26 0-58
Upper primary-coverts 1-90 90 20 0.22
Upper greater-coverts 13-32 20 32 1.50
Upper lesser- and median-coverts 8-48 41 38 0.93
Alula 65-78 14 7 0.50
Underwing-coverts 37-76 40 54 1.35
Shoulder tract 18-68 51 108 2.12
Ventral tract 13-85 73 736 10.08
Dorsal tract 13-80 68 490 7.21
Thigh tract 30-86 57 80 1*40
Legs 26-68 43 20 0.47
Head 19-70 52 86 1.65
Rectrices 27-79 53 136 2-59
Upper tail-coverts 34-80 47 28 0.59
Lower tail-coverts 30-77 48 24 0.50
* The figures are the averages of five birds; the range of values obtained was never more than
8% of the mean.
During moult, an average of 2.2 gm. of new feathers were produced by adult Bull-
finches, and about 1.7 gm. by juveniles; this is equivalent to lo;/, and 8% respectively
of their live-weights at the start of moult (see later), and 40% and 33% respectively of
their dry-weights (Newton & Evans, in prep.). Since adult moult lasts 10-11 weeks and
post-juvenile moult 7-9 weeks, the mean rate of feather synthesis by weight is about
the same in both.
Fig. 7 shows the rate at which new feathers were synthesised during adult and post-
juvenile moults; this was calculated for the primaries, secondaries and rectrices of adults
from Figs. 3, 1 and 2 respectively; in all other tracts (in the absence of other information)
it was assumed to be constant throughout (see Table 11). Given this assumption, the
production of feathers was slowest at the start and end of moult, but for most of the time
averaged nearly 40mgm. per day (equivalent to 0 6 % of the dry-weight of the bird)
in both adults and juveniles. There was, however, considerable variation and in many
birds feather production must, at times, have far exceeded this.
1.7
1 -5 1 A 5
1 .o 1.0
0.5 0.5
0 0
Stage of moult Stage of moult
FIGURE7. The rate of feather synthesis during adult and post-juvenile moults in the Bullfinch
Pyrrhula pyrrhula. (For explanation of moult stages see text.)
protein, and their constituent amino acids may be needed in proportions very different
from those in which they are present in the proteins of natural foods. It is possible
therefore that a larger weight of protein is required during moult than might be expected
simply to supply enough of the less common amino acids. No relevant information is
available for Bullfinches but it seems probable that extra food is required throughout the
moult, especially since food-shortage on only one or two days is liable to weaken the
structure of growing feathers.
ACTIVITY DURING MOULT
Apart from eating more during moult, birds might conserve energy by resting for
part of each day, especially if food is easy to obtain at this time. Captive Bullfinches,
which normally sing or fly around for most of the day, when in moult were active only
while feeding; the rest of the day was spent mainly in sitting or sleeping in a shaded
position with feathers fluffed. They also bathed less and preened more than at other
times.
TABLE
12. The number of Bullfinches Pyrrhula pyrrhula caught at different stages of moult.
ADULTS
Stage of moult 1 2 3 4 5 6 7 8 9
Number caught 43 27 22 14 9 22 9 7 29
Percentage of total caught 24 15 12 7 5 12 5 4 16
JUVENILES
Stage of moult 1 2 3 4 5 6*
Number caught 140 48 66 43 49 72
Percentage of total caught 31 11 16 10 12 18
Note. The table is based only on the data for 1963 and 1964 when trapping was carried out
with equal intensity throughout the moult period. For explanation of stages see text.
* Birds which had just completed the moult, as indicated by the presence of sheaths on sonie
of the new feathers.
62 I. NEWTON : THE MOULT OF THE BULLFINCH P F R R H U L A l’J*RRHL;Ld IBISio8
Table 12 shows the total number of adult and juvenile Bullfinches caught at different
stages of moult, irrespective of the dates their moult occurred. Although each stage of
moult took about the same time to complete, it may be seen that more birds were caught
in the first and last stages of moult than in the middle stages when more feathers were
being replaced. T h e difference is more striking in the adults than in the juveniles.
This is consistent with a reduction of activity for most of the moult. Moreover, during
any given period, birds in the first and last stages of moult could be caught at most
hours of the day, but birds in heavy moult mainly in the early mornings and in the
evenings. These data, however, indicate an average trend and refer mainly to birds
moulting in the most favourable part of the season. Some individuals, especially those
moulting late, may have to become even more active than normal in order to find enough
food (see later).
It was noticed also that relatively more moulting adults than juveniles were caught
in nets placed in thick cover compared with those placed in more open situations, such
as at patches of Meadowsweet. Such skulking behaviour, which was not apparent among
moulting juveniles, would assist adults in heavy wing-moult to avoid avian predators.
A similar difference in behaviour during moult between adult and juvenile Chaffinches
was noted by Marler (1956).
TABLE
13. The mean weights (with standard deviations) of adult Bulljinches Pyrrhula
pyrrhula according to stage of moult (all years).
August September October
Stage Male Female
Moult no: started (33) 21.7&1-5 (40) 23.2k1.6
1 (22) 22-3 2 1.2 (11) 23.1 f 1.1
2-3 (24) 22-92 1.2 (15) 22.821.0 (4) 23.0
4-5 (8) 23.7 (13) 23.8+ 1.0
6-7 (17) 23.7k1.0 (4) 23-8
8-9 (15) 23-82 1.3
Moult just finished* (18) 24.22 1.2
Notes. The figures in parentheses show the numbers weighed. Standard deviations are given
only when more than ten birds in each class were weighed.
* As indicated by the presence of sheaths on some of the new feathers.
I966 I . NEWTON : THE MOULT OF THE BULLFINCH P Y R R H U L A I’YRRIfUL4 63
TABLE
14. The mean weights (with standard deviations) of juvenile Bulenches Pyrrhula
pyrrhula according to stage of moult.
1962 1963
September October August September October
Stage
Moult not started (4) 21.5 (3) 20.7 (26) 21.5 k 1-0 (3) 21.9
1 (5) 22-9 (13) 22.4k1.2 (30) 21-8k1.1 (18) 21*3+1.2
2 (2) 22.8 (8) 22.5 (9) 22.5 (9) 22.6
3 (3) 22.9 (7) 22-8 (8) 22.8 (5) 23.3
4 (6) 23.8 (11) 24-3 1.2 (6) 23.8 (8) 24.0 (5) 23.7
5 (1) 24.2 (10) 24.3 (1) 24.1
6* (5) 24.8 (22) 24.0 &- 0.9 (11) 23.9$11 .3
1964
August September October November
Stage
Moult not started (30) 21.4&1.0 (7) 21.1
1 ,.
L
(38) 21.8,tl.O
(4) 22.1
(30) 21-611.1
(8) 22.5 (2) 21.2
3 (10) 22.8 (16) 22.611.2(9) 22.4
4 (3) 24.7 (8) 23.5
5 (8) 23.8 (15) 24.3&1-1 (3) 22.8
6* (14) 23.4,tl.O (24) 23.811.0
Notes. The figures in parentheses show the numbers weighed. Standard deviations are given
only when more than ten birds in each class were weighed.
* Moult just finished as indicated by the presence of sheaths on some of the new feathers.
of fat, but in late-fledged juveniles, partly to general body growth as well. In both adults
and juveniles the same range of weight changes occurred in different years and at different
dates in any one year (Tables 13 and 14), as well as in particular individuals caught more
than once during moult.
There is therefore no evidence from these mean weight changes that the moult
imposes a physiological strain on Bullfinches; but as mentioned above, the figures are
almost certainly based on the weights of a sample of healthy birds. If any succumbed,
the chances that they would be caught just beforehand are very small.
TABLE
15. The percentage composition of the food of the BullJinch Pyrrhula pyrrhula in
Wytham Woods, Oxford, from August to November in three successive years.
1962 1963 1964
Aug. Sep. Oct. Nov. Aug. Sep. Oct. Nov. Aug. Sep. Oct. Nov.
No. of obs. 101 162 215 164 102 134 152 131 117 153 201 102
Birch 45
13 36 29 34 20 16 28 14 25 20 15 7
Grasses 1 0 0 16 2 0 0 14 2 0 0
Docks 6 17
4 4 6 15 17 4 16 14 10 12 16
Bramble 9 17 21 12 18 20 25 10 11 14 18
Meadowsweet 17 20 20 3 19 24 26 2 24 26 13 2
Wood avens 4 3 1 0 4 5 1 0 3 5 1 0
Nettle 2 3 8 7 3 8 17 39 3 12 40 55
Elder 1 1 15
0 0 1 1 0 0 1 2 1 0
Privet 0 14 6 0 4 0 0 0 8 2 0
Ash 0 0 1 20 0 0 0 0 0 0 0 1
Othem 3 1 5 5 10 5 4 4 6 4 2 1
64 I. NEWTON : THE MOULT OF THE BULLFINCH P ' Y R H H I J L AP Y R R H U L A IBIS108
In Wytham Woods, the seeds of various herbaceous plants were abundant and
continually replenished from May to September each year. It is unlikely therefore that
food for Bullfinches was short at this time, though no measurements were made. From
October however, there was no fresh growth, and Bullfinches fed mainly on seeds which
were to last them into the winter, namely those of bramble, docks, nettle, birch, and
privet. The seeds of ash were also important, but these were not taken in any quantity
until they had ripened after the end of the moult period; they were plentiful in 1962
and 1964. The crops of birch and privet seeds were largest in 1962 and smallest in 1963.
These seeds formed a higher proportion of the diets of Bullfinches in 1962, when they
were most plentiful, than in the following two years (Table 15).
(1) Adults
It is unfortunate that in 1962 netting was not started until mid-September since in
this year there was prolonged breeding by some Bullfinches and a considerable spread
in the date on which moult started. In this year, the difference in survival between adults
moulting early and late might well have been less than expected, since birds presumably
continue breeding and moult late only in exceptionally favourable years when the seed-
crops, produced in late summer, are large enough to remain abundant into the autumn
1961 I. NEWTON : THE MOULT OF THE BULLFINCH I J Y R R H U L A PYRRIIC'LA 65
(see later). Thus in 1962 almost all the late-moulting adults caught were recovered after
the end of the moult season, which showed that they had not suffered appreciable mortality
as a result of their late moult.
I n 1963, data for analysing survival in the adults are insufficient. I n 1964, however,
five out of eight birds that started to moult in the first half of the season were subsequently
recovered, compared with one out of five birds that started to moult in the latter half.
This suggests that in 1964 the adults moulting later survived less well than those moulting
earlier, though more data are required to substantiate this. T h e alternative explanation,
that more of the late-moulting adults had left the wood, is unlikely, since as mentioned
above, there are no records of Wytham adults moving very far.
( 2 ) Juveniles
Table 16 shows the recoveries of juveniles beginning to moult in different months
1962-64. Only birds caught in the first stage of moult are included. Further, to exclude
birds which might still have been dependent on their parents when they started moult,
all those with under-developed rectrices when caught were omitted from analysis.
Many juveniles were later recovered in Marley, but three were reported from outside
the wood (within two miles) by members of the public and are listed separately. It may
be seen that in all years more of the juveniles moulting early were recovered after
completing their moult than of those moulting late. Though the data are few, they
suggest that late moulting carries a lower survival (=recovery) rate for juveniles than
early moulting. T h e alternative explanation, that more of the late moulting birds had
left the area, is again unlikely since all those recovered outside Marley were birds which
had moulted early.
TABLE
16. Survival of juvenile BultJSnclaesPyrrhula pyrrhula in relation to time of moult.
Start Number Recaught after November
moult examined
in Marley elsewhere Total Percentage
1962
September 9 4 1 5 56
October 14 4 0 4 28
1963
July 4 3 1 4 (1 55
August 18 7 1 8
September 6 2 0 2 33
1964
- . ..
July 5 1
August 47 18
September 39 7
It is not surprising that the early moulting birds survived better, since, in addition
to moulting at the most favourable time, they were by then more experienced at finding
food, for they had already been out of the nest 4-8 weeks compared with only 2-3 wecks
for the late-moultingjuveniles. It is of interest also that in 1962, when the late-moulting
adults survived well, the late-moulting juveniles apparently suffered heavy losses. This
is again presumably because these juveniles were younger, less experienced at finding
food and began moult 2-3 weeks later than their parents.
Finally, it is worth stressing that in all years most birds moulted while food was still
plentiful; only a few (mainly juveniles) were still in heavy moult when food became scarce.
These observations therefore support the view that the moult is timed to occur while
food is still plentiful. Since most of the seeds eaten by Bullfinches during moult are
already available and are eaten to some extent earlier in the summer, the birds have
presumably evolved the ability, as in 1962, to prolong breeding and simultaneously
postpone moult in those years when food is exceptionally plentiful. But e w n in 1962, a
proportion of birds finished breeding and moulted at the normal date. It is possible
E VOL. 108
66 I. NEWTON : THE MOULT OF THE BULLFINCH PYRRHC'LA PYRRHC'LA IBIS108
therefore that selection to avoid a late moult preserves some birds which have a hereditary
tendency to moult early, irrespective of conditions.
DISCUSSION
In normal years in England, a period beginning at the end of July, during which
food is abundant, is probably the most favourable time for moult in the Bullfinch (the
food-supply and the timing of the breeding season being the " ultimate " factors to
which the timing of moult is adapted). Juveniles which left the nest after about mid-
August began to moult within a fortnight, suggesting that the advantage of moulting
then outweighed that of waiting until they were well grown and more experienced at
finding food. Doubtless, therefore, juveniles which left the nest at the end of May could
have begun moult before the end of July if it had been advantageous for them. Further-
more, even the non-breeding males did not begin moult until towards the end of July.
Presumably there is some external regulator, perhaps daylength, which acts as the main
proximate factor initiating the moult at this date. But the effect of daylength is modified
in the adults by the date at which breeding activity ends, and in most of the juveniles
by their age.
The abundance of food and warm weather may not be the only factors making the
end of July the best time for all Bullfinches, adults and juveniles, to begin moult. Many
of the adults examined just before moulting late in 1962 had very worn plumage (see
above), and these were birds which cannot have moulted for more than a year. Even
longer periods might elapse between successive moults unless selection kept the timing
of moult fairly constant from year to year; the plumage of any bird, adult or juvenile,
which moulted early in the summer of one year (before the end of July) might be so
worn by the end of a long breeding season the following year as to impair flight and
insulation.
Natural selection will presumably favour those individuals which raise most broods
during a season. If moult occurs at a time of year when breeding is also possible (but
this has not been proved for the Bullfinch), selection will tend to compress the moult
to within as short a time as possible in the latest part of the period during which food is
abundant, a period which varies in duration from year to year. If food were continuously
plentiful, the latest date at which a moult could be safely accomplished in a resident
species, such as the Bullfinch, would probably be set by the colder and shorter days of
winter when insulation by the plumage is most needed.
In all three study years, most Bullfinches finished moult just before food became
scarce, even though this occurred later in one year than in the other two. Although in
some species moult is markedly slower when food is scarce (Ashmole 1962), even with
abundant food there will obviously be a limit to the rate of moult; it must not proceed
so fast as to hinder flight or serioudy reduce insulation.
ACKNOWLEDGMENTS
My sincere thanks are due to Dr. P. R. Evans and Mr. R. E. Moreau for criticising the manuscript
and for their many excellent suggestions. P. R. Evans gave particular help with the statistical
analyses. I am also indebted to Dr. D. Lack and Dr. C. M. Perrins for useful comments on the
manuscript, and have benefited greatly from discussion with Professor A. J. Cain and Dr. D. W.
Snow. This work was carried out while I was holding grants first from the Department of Scientific
and Industrial Research, and then from the Agricultural Research Council.
SUMMARY
The distribution of feather tracts and their sequence of moult in the Bullfinch is described.
The adult post-nuptial moult, which is complete, lasted 10-12 weeks, and the post-juvenile moult,
which is partial, 7-9 weeks. Adult moult began with the shedding of the first (innermost) primary
and ended with the replacement of the last. Variations in the rate of moult in the flight feathers were
mainly achieved, not by changes in the growth rates of individual feathers, but in the number of
feathers growing concurrently. The primaries were shed more slowly, and the onset of body
moult delayed, in birds which were still feeding late young.
1966 I. NEWTON : THE MOULT OF THE BULLFINCH PYRRHL'LA I'l'RRRULA 67
In 1962, the onset of moult in the adults was spread over 11 weeks from thc end of July to the
beginning of October, and in the two following years over the six weeks, from the end of July to
the beginning of September. The onset of moult was delayed by late breeding, which itself
occurred in response to a comparative abundance of food in late summer, markedly in 1962. I n all
years, the first juveniles to moult started at the end of July, and the last, three weeks after the latest
adults. Juveniles moulting late in the season retained more juvenile feathers than those moulting
earlier.
During moult, adult and juvenile Bullfinches produce feathers equivalent to 40':: and 33%
respectively of their dry weights. In both, for much of the moult, an average of nearly 40 mgm. of
feather material-some 0.6"; of their dry-weight-is laid down each day. The remiges of the
adult comprise only a seventh of the weight of the entire plumage, and it is suggested that their
protracted moult results not so much from their energy requirements, as from the need to maintain
efficient flight. Variation in the rate of moult in the remiges was much less pronounced than in the
body feathers. Bullfinches were less active during moult than at other times of the year. T h e
weights of both adults and juveniles increased during moult.
The food during the moult period is described. I n all years, most Bullfinches finished moulting
just before food became scarce, even though this occurred at different times in different years.
In one year, adults moulting latest in the season probably survived less well than those moulting
earlier; the same was apparently true of the juveniles in all years. T h e timing of moult in the
Bullfinch, and the factors initiating it, are discussed in relation to the breeding season and food-
supply near Oxford.
REFERENCES
ASHMOLE, N. P. 1962. The Black Noddy Anous tenuirostvis on Ascension Island. Ibis 103b:
235-273.
BLANCHARD, B. D. 1941. The White-crowned Sparrows (Zonotrichia leucophrys) of the Pacific
seaboard: environment and annual cycle. Univ. Calif. Publ. Zool. 46: 1-178.
DIXON,K. L. 1962. Notes on the molt schedule of the Plain Titmouse. Condor 64: 134-139.
DWIGHT,K. L. 1900. T h e sequence of plumages and moults of the passerine birds of New York.
Ann. hTewYork Acad. Sci. 13: 73-360.
EATON, S. W. 1957. A life history of Seiuvus noveboracensis. Sci. Studies, St. Bonaventure Univ.
19: 7-36.
EVANS,P. R. I n press. Autumn movements, moult and measurements of the Lesser Redpoll
Carduelis jlammea cabaret. Ibis 108.
GIBB,J. A. 1954. The feeding ecology of tits with notes on Treecreeper and Goldcrest. Ibis 96:
513-543.
JOHNSTON, D. W. 1961. Timing of annual molt in the Glaucous Gulls of northern Alaska.
Condor 63 : 474-478.
KING,J. R. & FARNER, D. S. 1961. Energy metabolism, thermoregulation, and body temperature.
I n ' Biology and Comparative Physiology of Birds '. (Ed. J. A. Marshall.) Vol. 2: 215-288.
London & New York: Academic Press.
KOCH,H. J. & DE BONT,A. F. 1944. Influence de la mue sur l'intensite de mktabolism chez le
Pinson Fringilla coelebs coelebs L. Ann. SOC.Zool. Belg. 75: 81-86.
MARLER, P. 1956. Behaviour of the Chaffinch Fringilla coelebs. Behav., suppl. 5: 147-150.
MICHENER, H. & MICHENER, J. R. 1940. The molt of House Finches of the Pasadena region,
California. Condor 42: 140-153.
MILLER,A. H. 1928. The molts of the Loggerhead Shrike Lanius ludovicianus Linnaeus. Univ.
Calif. Publ. Zool. 30: 393417.
MILLER,A. H. 1933. Postjuvenal molt and the appearance of sexual characters of plumage in
Phainopepla nitens. Univ. Calif. Publ. Zool. 38.
MILLER,A. H. 1961. Molt cycles in equatorial Andean Sparrows. Condor 63: 143-161.
NEWTON, I. 1964. T h e ecology and moult of the Bullfinch. D. Phil. thesis, Oxford University.
NEWTON,I. 1964 a. Bud-eating by Bullfinches in relation to the natural food-supply. J. Appl.
Ecol. 1: 265-279.
NEWTON, I. I n press. Seasonal fluctuations in the weights of Bullfinches. Brit. Birds.
PEREK,M. & SULMAN, F. 1945. The basal metabolic rate in molting and laying hens. Endo-
cinology 36: 240-243.
PITELKA, F. A. 1958. Timing of molt in the Steller Jays of the Queen Charlotte Islands, British
Columbia. Condor 60: 3 8 4 9 .
RIDDLE,0. 1908. The genesis of fault bars in feathers and the cause of alternation of light and
dark fundamental bars. Biol. Bull. Woods Hole 14: 328-370.
SELANDER, R. K. 1958. Age determination and molt in the Boat-tailed Grackle. Condor 60:
355-376.
SELANDER, R. K. & GILLER, D. R. 1960. First-year plumages of the Brown-headed Cowbird and
Red-winged Blackbird. Condor 62: 202-214.
SNOW,D. W. 1958. A Study of Blackbirds. London: Allen & Unwin Ltd.
SOUTHERN, H. N. 1954. Tawny Owls and their prey. Ibis 96: 384-410.
WALLGREN, H. 1954. Energy metabolism of two species of the genus Emberiza as correlated with
distribution and migration. Acta Zool. Fenn. 84: 1-110.
WRIGHT,P. L. & WRIGHT,M. H. 1944. T h e reproductive cycle of the Red-winged Blackbird.
Condor 46: 46-59.
Dr. I . Newton, Edward Grey Institute, Botanic Garden, Oxjord.