PLUMIK;PS AND MILT

Intanbcticm

Since mult of feathers is an integral of

annual physiological cycle of birds, a detailed study of the

process and its relation to the other major events of the cycle

was undertaken. Moult and breeding are considered to be major

energy demanding events in the annual cycle of birds. A study of

the temporal relationship between these two activities may give an

insight into the ecological pressure acting on a particular given

powlation. The plumages and mult cycle in the Whitebreasted

Kingfisher are described in this chapter.

Materials & MetMs

Adult specimens of the Whitebreasted Kingfisher

were collected by shooting, and after rcmving the gut, gonads and

thyroid they were preserved in 143% formalin. Later these

preserved specimens were dried in the laboratory and their mult

was studied. ?'he data obtained over four years from 1988 to 1991

were pooled. For the study of the post-juvenal u t two

nestlings in the late stage collected from two nests were kept in

captivity. Of them, one died just after the mult has started,

and the other was maintained in capticity till the end of the

study period. ChanEfes taking place in the bill and p l w of

nestling were also noted. &tails of the post juvenal moult were

also recorded.

In the given adult speci- the extent of mxlt

was assessed by assigning numerical score to each individual using

a system followed by Miller (1961), Ashmle (1962) and Newton

(1966). Individual primary feathers of moulting birds were given

a score from '0. (old) to 5 (new) according to their state of

growth. The sum of the scores of individual primaries was taken

as an index of the general. state of mult. The total score for

both wings is the sum of the individual values for the ten

primaries (l0i3). In recording the moult of individual feathers,

the following categories and scores were used.

Old feather - 0

Feather miss.ing or in small pin - 1

Feather in large pin or brush stage - 2 Feather brush to half-grown - 3

Feather half to three- quarters grown - 4

Feather three-quarters to full mwn - 5

Based on the numerical scoring system given above, the m l t in

the Whitebreasted Kingfisher is divided into 1-10 stages (Table

23). The moult of the secondaries and rectrics was also recorded

using the same method, and the relation of their moulting with the

primary stages was noted. Stage of moult on the body was scored

in seven regions, viz, capital, spinal, ventral, humeral, femoral,

marginal and c m a l from 0 to 3; '0' for no active feather

replacement, '1' for relatively small numbers of feather papillae,

-2, for 50% of feather growth and "3' for relatively heavy feather

growth. The above scoring system is similar to the one proposed

by Niles (1972). The mean date and duration for the start of

moult were estimated by regression analysis of Julian date of

moult score ( P h , 1976). The moult of primary, secondary and

tail coverts and alula was also carried out.

The appearance of plumage and moult in different

stages of the Whitebreasted Kingfisher are as follows : -

The newly hatched chick of the Whitebreasted

Kingfisher is fully naked and flesh c o l d . No natal down

feather coat is noted before the growth of the juvenal plumage.

At the tims of nest leaving the fledgling appears to be fully

feathed. The juvenal plumage is very mch similar to that of

he adult kingfisher, but duller.

The tips of the primaries are round and the

feathers of white breast region have short black stripes on their

margins. The length of the primaries and secondaries of juvenile

is less than those of adult. The average measurements of juvenile

flighL feathers (N-6) are given in Table 10

Pl- of first year bird

The pl- of the first year bid is very similar

to that of adult and becomes brighter after the post-juvenal

moult. The tips of the primaries a m pointed. The size of the

feathers is similar to that of adult feathers.

There is no sexual dimorphism in the plumage of the

Whitebreasted Kingfisher. The plumage of both male and female is

identical in colour. The feathers of the head, dorsal and lateral

surface of neck, underparts of abdomx and wings are deep brown.

A conspicuous white 'shirt frontr extends f m the bill to the

breast on the ventral side. The secondaries, rectrices and the

feathers on the back are brilliant turquoise blue in colour. The

prkies have white tips. The coverts of secondaries are black

in colour. The tips of the primaries of the adult are pointed.

The size of the feathers is larger i n comparison to those of

juveniles. The aver- measuremnts of f l ight feathers of adult

(10 cases) are given in Table 10).

Tha fea ibr tracts and their pattern of m3ult

The distr i tut ion of the feather t racts (pterylae)

is best seen in a nestling when the feathers are i n sheaths or

still small. There are eight t racts of which 3 are feathered and

arranged as i n most paserines (Dwight, 19PM). In a l l these

t racts , mlt begins a t a number of points ( loci ) from which it

spreads i n a characteristic manner.

I . Capital tract. This t rac t covers the entire head and

passes into the spinal t rac t .

11. Spinal tract (dorsal tract). The dorsal t rac t includes the

feathers of nape, back and nuop. I t is continuous with the

capital t rac t and ends p t e r i o r l y i n front of the preen gland.

111. Ventral tract. The ventral t ract starts under the b i l l ,

runs up to the end of neck and then divides into two main branches

down thrwgh the belly to end in front of anal c i rc le t .

I V . lbrmural tract (douldor t rac t ) . Each half of tlw humera1

t rac t connects with the ventral t rac t and then curves dorsally to

form a band of feathers across the humerus.

V.F-Dral tract (thigh t rad) . I t comprises a pair of narrow

oblique bands of feathers on each flank.

VI. CNsal tract. C m a l t rac t comprises the feathers of the

legs.

The body feathers begin their m u l t a l m s t a t the

same time in a l l the t racts mmtioned above. There is no

particular locus in any of the t racts where the replacement of

feathers starts. The replacement of body feathers is compleki

before the completion of the primary moul t .

VII. Ths alar tract. The f l ight feathers, thei r various coverts

and the alula quil ls constitute the alar t ract . The Whitebreasted

Kingfisher has 10 primaries and 15 secondaries. (Plate - 11)

Following the system used by Sbssmann and S t r e s s m (1966) the

primaries are referred to by the symbol H (Handwing) and

secondaries by the symbol A (Arming). The primaries here are

numbered f m the carpal joint and spreads outwards. The upper

greater pr- covert is shed with corresponding primary and

becomes f u l l grown when the primary feather is half grown. Upper

W a n primary covertsare also replaced f r v m f i r s t to the l a s t .

There are no upper lesser coverts for the primaries. Under

pla te 11

Showing alar t r a c t (dorsal view) of the Whitebreasted Kingfisher

primary coverts do not have any strict pattern. The primary

coverts except the upper greater primary coverts are replaced

together se r i a l ly .

The secondaries are numbered from the carpal joitlt

inwards (Plate 11). The secondary q u i l l s moult from two loci i .e.

from A1 and All. Moulting progresses from both s ides and meets at

A4 which is the last secondary to moult. Another sequence of

moult of the secondaries starts from All and proceeds to the last

feather A15.

The upper greater secondary coverts f a l l from the

first feather and then gradually proceed ascendantly to the last.

The upper median and lesser secondary coverts and a l so a l l the

under secondary coverts do not follow a clear pattern of moult.

They are replaced almost together.

There are three a lu l a r q u i l l s numbered from 1-3,

inside outward. They moult from 1 tr~ 3.

VIII. The caudal trect. The caudal tract includes 12 -trices

and the i r upper and under coverts. The mlt of rectrices is from

the middle pair to outwards. The upper and lower coverts moult i n

advance to rectrices moult. The moul t of these coverts is

centrifugous.

In the adult specim=ns (N-79) examined by me, mult

was noted in all the feather tracts. At the powlation level

moult started in July and completed in December.

Marlt of p M e 9 , and seadades. The shedding of h e first

primary (HI) signified the onset of moult which usually extended

from July to December (Table 2%; Fig.6). The second primary was

shed when the previous feather was partly grown. The old one was

dropped out before the new one was mhed out. Corresponding

feathers in both wings were usually shed simltaneously showing

the same pattern of growth. Replacement of the secondaries

started a little late extending from August to December (Table

21). (Eleventh Secondary( All) was shed when the primary moult was

in the stage 3 ) , which was then followed by A12 and AllZl

simltaneously. First secondary usually was shed when the primary

moult was in the stage 6. The last moulting secondary was

A or A4. 3

Moult in the Whitebreasted Kingfisher was divided

into ten successive stages based upon the primary score (Table

23). Monthwise occurmce of different stages of primary moult is

given in Table 24. Table 25 gives details of moult on the

individual primary feathers together with the mean number of

Table 20. Monthly record of primary moult in the Whitebreasted Kingfisher, Halcyon snymtv~is fusca

No. of birds Primaries Monthly

Month collect-Not tion m l t - mlting completed 1 2 3 4 5 6 7 8 9 10

ing

June 11 10

July 11 4

Ausust 10 1

September 9 -

October 10 -

November 10 - kxmber 10 -

January 8 -

Flg.6 Moult of the prlmarlee In the Whltebmaeted Klnglleher In

dltbrent month8

Table 21. Monthly record of secondary moult in the Whitebreasted Kingfisher, Halcyun snlyrnensis fusca

Month Jun Jul Aug. Sep. k t . Nov. Dec. Jan.

Monthly collection 11 11 10 9 10 10 10 8

No. of birds

Not moulting 11 10 3 - - - - -

Moulting 11 1 7 9 8 7 3 1

Completed - - - - 2 3 7 7

Secondaries No. of birds moultins

Table 22. Monthly record of retrix moult in the Whitebreasted Kingfisher, Hdcyon sqvznensis fusm

No. of birds Rectrices Monthly

Month collection not mxllt- mxllting completed 1 2 3 4 5 6 ing

June 11 11 - - old feathers

July 11 11 - - old feathers

August 10 6 4 - 3 1 2 1 2 1

September 9 - 9 - 6 5 5 5 3

October 10 - 7 3 2 4 4 6

November 10 - 5 5 1 2 2 5

Member 10 - 2 8 1 1 1 1 2 1

J=ww 8 - - 8 New feathers

growing primaries in differnt stages of primary moult. At the

timg of the beginning and the end of the mult there was only a

single feather growing but the number of feathers growing in

between was generally two to three.

Table 23. Scheme f o r calculati.ng m u l t stages i n the Whitebreasted Kingfisher, Halcyo;~ w1~1(311.s.f s fusca

No. of birds in e8ch Moult stage Primary score stage

Table 24. Data showing nwult stages i n the Whitebreasted Kingfisher, Halcyon wrnensis fusca

No. of b i d s Moult stages

Month Monthly m l t i n g ccinpleted 1 2 3 4 5 6 7 8 9 10 col lect ion

June 11 1 - 1

July 11 7 - 3 2 1 1

September 9 9 - 1 3 4 1

October 10 8 2 1 3 6

December 10 3 7 2 8

January 8 1 7 8

Total 6 2 1 3 2 2 5 6 6 3 1

Table 25. Moult of primaries in the Whitebreasted Kingfist~er, Halcyvxn sqvznmnsis fusca

No. of birds Primaries Mean no. of Stage in different 1 2 3 4 5 6 7 8 9 10 growing ~rimaries

s'azes

1 6 5 2 1.16

2 2 2 2 2 3.m

3 1 1 1 1 3 .m 4 3 1 2 3 2 2.66

5 2 1 2 2 2.50

6 2 2 1 1.50

7 5 1 5 1 1.40

8 6 6 2 1.33

9 6 4 6 1.67

10 31 3 10 0.41

Moult& in the secondaries began when the H4 was

moulting or the moult was in the stage 3 (Table 26). Secondary

d t started with the shedding of a single feather, and later the

rate of moulting soon got accelerated with three to five feathers

growing at a t b in each wing. The rate of increase in the

secondary score was slow in the first half of the primary moult

and then increased in the second half F i g 7 Moult of the

secondaries completed at the same tim as the primary moult (N =

1'7). In two cases secondary moult proceeded after the completion

of the primary d t .

Table 26. Moult of secondaries in relation to primaries in the . Whitebreasted Kingfisher, Halcyon sqnnensis fuscd

No. of birds Secondaries Mean No. Stages in different 1 2 3 4 5 6 7 8 9 la 11 12 13 14 15 of growing

stages secondaries

Mt of Rectrices. In the Whitebreasted Kingfisher, moult of

-trices took place between August and December(Tab1e 22). In

most cases (N = 25) rectrices moulted centrifugally and

smetrically. Some irregularity was noticed in two cases studied

so far. The tail m>ult started in the middle pair of feathers

when the sixth primary (H6) was growing (Table 27). All the birds

in the stage 6 examined had the tail feathers moulting. The rate

of moulting was very fast in the stage 7, and generally 2 to 4

tail feathers were found growing at a time (Fig. 8). The

Flg.7 Secondary ecore in relatlon to Primary ecore in the Whltebreaeted

Klngfleher

PRlUpslV SCORE (Ll - 100 )

termination of mult was restricted to the last feather. The tail

mult became complete before the completion of the primary moult.

Table 2'7. Moult of rectrices in relation to primaries in the Whitebreasted Kingfisher, Halcyon snlyn~er~sis fUsca

No. of birds Rectrices Mean no. of growing stase in different 1 2 3 4 5 6 rectrices

stages.

wt inwingcwerts, tail coverts. and all other body tracts.

The upper greater primary coverts moulted along with the primary

feathers. The upper greater secondary coverts started mlting

when the moult was in the stage 4 and ended before the completion

of secondary mult. The moulting of the upper greater secondary

coverts started from the first feather and gradually p d e d

ascendantly to the last. Moult of the upper median primary

coverts, &an and lesser secondary coverts and the under wing

coverts did not follow a clear pattern of mult. All these

feathers moulted along with the body feathers. The u p p r and

Flg.8 Tall aeore In telatlon to Prlrnary score In the Whltebreasted Klngflsher

lower coverts of t i e tail moulted i n advance to the moult of

rectr ices, but in few cases the upper coverts followed the m u l t

of rectr ices. The pattern of m u l t i n tail coverts was

centrifugoUs.

The m u l t sequence in the body tracts was assessed

simply on the basis of new or pin feathers and proper recording

was mde according to the percentage of m u l t . The moult of the

bodv feathers started almst a t the same time i n a l l the t r a c t s i n

the primary stage 5 and ended before the tenth primary became full

grown. Percentage of number of birds moulting its body feathers

is given i n Fig. 8.

Moult of the a lu la se r i a l ly s tar ted from the f i r s t

to the third i n the primary stage 5.

Table 28 gives the timing and duration of moult i n

a l l the body tracts, wing coverts, tail coverts and a lu la shown i n

relation to the primary moult. Fig. 9 gives monthly record of the

percentage of birds showing the nwult of body feathers.

Table 28. Moult of Wing and tail coverts, body feathers and alula in relation to primary stages i n the Whitebmsted Kingfisher Halcyon ~ z n e n s i s fusca.

Stages 1 2 3 4 5 6 1 8 9 1 0

Birds i n stages 6 2 1 3 2 2 5 6 6 3 1

No. of birds nwulting

Primary Upper greater

Upper mxlian

Lower greater

Lower median

Secondary Um?er greater

Upper median

Lower greater

Lower median

Tail Upper

Lower

Capital t rac t

Spinal t rac t

Humera1 t rac t

F w r a l t rac t

Marginal t ract

C m a l t r ac t

Ventral t rac t

Alula

Jun

Fig.0 Monthly record ot moult in the body tract8 of the Whlbbrea8bd

Klngtl8her

Jul Oct NO\ Dec Jan

Post-juvenal m l t in the Whitebreasted Kingfisher

is described on the basis of a study on two fledglings reared in

captivity from 1989 to 1990. Cne bird r-ved from nest on 12th

May 1989 started multing on July 18, beginning from the first

primary. This bird later died before the completion of the mult

or when the second primy was moulting. The second one was taken

out from the nest hole on 14th Hay 1990. Moult of this bird began

on 16th July and ended on 20th January 1991.

The moult in the juvenile was quite similar to that

of the adult in its duration and pattern. The onset of mult in

juveniles was marked by tfle shedding of the first primary

gradually proceeded towards the last. When the third and fourth

primaries were growing, secondaries started moulting from All to

AI5 and AIQl to Ag and soon the first feather A1 moulted when the

fifteenth secondary was multing. The last moulting feather was

A4. The =trices showed an irregular pattern of mult which

occurred when the eighth primary was in the growing stage. The

upper greater prFmary coverts showed no sign of replacement along

with their respective primaries. Upper lesser primary coverts and

lower primary coverts were not replaced. The upper greater

secondary, median secondary and lesser secondary coverts were shed

and replaced. The under secondary coverts did not show any sign

of mlt. The tail coverts moulted and were replaced. The

moulting of wing and tail coverts w a s similar to t h a t of adul t

moult. The body moult f i r s t noted on the h m r a l tract and l a t e r

spread to a l l the other t r a c t s . The total duraLion of the

post-juvenal moult w a s noted to be of 190 days. One of the

peculiar features of the moult i n juvenile was that the secondary

moult continued after the completion of primary moult. Large

samples are needed to ar r ive a t a def in i te conclusion about the

post-juvenal mwlt i n the Whitebreasted Kingfisher.

Timing and duration of moult

The t im and duration of the adul t m n d t was

calculated by using regression analysis, and date as dependent

variable and primary score a s independent variable (Pimn, 1976).

The regression (Fig. 10) indicated that progression of m x l t was

l inear and the start of d t (primary) was s m where arvund late

June and ended in mid-November. The duration of m u l t was

dculated appmximately to be 140 days and the average increase

of primary score per day was 1.39.

In the Whitebreasted KCngfisher, the juvenal

plumge does not show mch variation i n colour from that of the

adul t plumage. The colour of plumage of the juvenile is du l l when

Fig. 10 Regression of primary moult score on date

in the Whitebreasted Kingfisher

53 103

DAYS ( Day 1 . July 1

compared with the plumage of adult. Milstein (1962) noted the

dirty apFearance of plumage in the juvenile of the Angola

P~%c€isher. k i n g the post-juvenal moult all the flight

feathers, rectrices and almst all the b d y feathers are replaced.

The juvenile kingfishers when moulting during the year they are

hatched seem to retain some feathers especially few wing averts.

I have not made a detailed study of the percentage of feathers

retained during the post-juvenal mwlt. S a m available data on

the passerine species indicate that the percentage of feathers

retained during the post-juvenal moult may be influenced by

several fact~rs. Pitelka (1945) noted the retention of some

coverts through the post-juvenal moult in northen races of

A p I ~ l a z m cccm1escen.s and stated that the more extensive

retention of coverts in the Southern races might be attributed

either to greater wear or to the early breeding. In Moloffuus

ater and Agelains p I n ? ~ l i m s , a wrelation between the total

number of retained juvenal feathers and the degree of cranial

ossification (indicating age) at the time of post-juvenal mult

has been established (Selander and Giller, 1960). The adaptive

significance of the retained feathers as pointed out by Miller

(1928) was thoroughly discussed by Selander and Giller (1960).

Certain late developing feathers, like the under wing coverts,

h u s e of their protected position do not wear before they mult

during first annual moult. Thus, the metabolic energy is

conserved by not moulting them during post-juvenal m l t .

W i n g the post juvenal moult the Whitebreasted

Kingfisher renews the remiges and rectrices but some passerines

retain them. In this kingfisher, difference in size between the

flight feathers of the juveniles and those of the adults alone

would favour a removal of wing feathers during the post-juvenal

moult. Since the juvenile remiges and rectrices are shorter than

those of the adult birds, it would be advantageous to the bird to

remove them before an increase in body weight seriously interfers

with its power of flight.

The timing and duration of moult in the various

species of birds is related to the ecology and other events in the

annual cycle (Newton, 1968). In the Whitebreasted Kingfisher, the

moult proceeds without interruption from July to hcember. In

this bird, the duration of the annual moult is about six mntlls

which is a lengthy period in comparison to that in other birds.

The duration of moult is recorded as about 56 days for Catduelis

f l a m m (Evans, 19661, 60 dqvs for Zonotddda capensis (Miller,

1961), 60-70 days for Qanocl tta stellei (Pitelka, 1958) 70-84

days for fjcz'z'11ula py~r11ula (Newton, 1966), 77 days in Tawny Owl

(Hirons et a1 . , 1984) 85-110 days for &ssi d i x mexi caicls (Selander

1958) and 105 days for the House Finch (Michener and Michener,

1940). Longer duration of moult was also noted in the birds of

tropical warm clhte (Naik and Andrews, 1966; Naik and

Shivanarayan 1969a; Naik and Naik, 1969; Mathew, 1975). Waders

breeding in tropical latitude tend ta taka longer time than those

which mult in temprate areas (Pearson, 1981, 1984). buthwaite

(1971b) recorded that in Pied Kingfisher, k y l e zudis the captive

birds took six mnths for the completion of moult. The duration

of moult in the Whitebreasted Kingfisher is therefore closely

comparable to the duration of moult of other birds residing in

tropical climate.

The sequence of mult in the remiges and rectrices

of the Whitebreasted Kinsfisher is similar to that found amng the

passerines. It is known that the mult sequence of primaries of

birds in general follows certain patterns which m y be

characteristic of certain natural groups. According to Stresemann

(1963b), in a small sized bird, descending sequence of primry

moult is of the saw advantage as an ascending sequence, but even

then the primaries usually nwult in the same order among the

phylogenetically related birds. In the Whitebreasted Kingfisher,

d g e s and rectrices mult symnetrically as in nwst other birds.

Nonoally there is some w-ordination between mult of primaries,

secondaries and tail feathers. The sequence in the moult of

d g e s of the Kingfisher has some advantage as noted in

passerines . The inner secondaries ) usually start moulting

when the second primary (H2) is already replaced and the first

secondary (Al)mults a little later as the primary mult is half

way through. The advantage of this sequence of moult is that the

rmulting feathers are separated by the intact ones. The mult of

rectrices starts when a few primaries are renewed. Usually

Lail coverts are replaced before the moulting of the tail feathers

which is of advantage to the bird. In the absence of rectrices,

tail coverts are of importance to the bird in landing (Fisher,

1959; Evans, 1966). Co-ordination between the primary moult and

the secondary tail moult has been reported in the Jungle Babbler

(Naik and Andrews, 1966).

The body moult of the Whitebreasted Kingfisher

occurs in the latter half of the entire period of nwult. One of

the pecular features of the body moult is that there are no

particular loci on any of the tracts from where the moult starts.

In most of the cases I have studied, the body uwult occurs in all

the tracts simltaneously. Similar case of body moult was

observed in Goldfinch (Middleton, 1977).

Breeding and mult are two physiological events

which involve mch mtabolic strain in the life of birds. Of the

two, breeding, which is the more important activity occurring at

the most favourable part of the year, could be ultimate factor

influencing the timing and tempo of moult. The breeding and moult

are mtually exclusive in a majority of birds, and this is

generally considered as an adaptive feature from the stand mint

of budgeting the metabolic energy. In the Whitebreasted

Kingfisher, there is a temporal separation of breeding and nwult

as the feather moult starts only after the tednation of breeding

activities. The separation of breeding and moult in m a n y bird

species is regarded as an adaptation for spreading the energy

demands of their activities (Payne, 1972; Murton and Westwood,

1977). In the Whitebreasted Kingfisher, the breeding activity,

which is noted from January to June, is followed by the moult

extending from July to December. The breeding is cited as an

important adaptive strategy in the annual cycle (Heitmyer, 1985,

1987; Him115 et al., 1984; Miller, 1986). Moult is a costly

process and is reported to increase basal metabolism by 9-46%

(Loworn and Brazen, 1988). King and Farner (1961) tentatively

estimated a mean increase in energy intake of 7.6 per cent in the

House Sparrows during the post-nuptial moult assuming no change in

other energy demanding functions. According to King and Murphy

(1985), moult will time the events if the birds are unable to meet

the cost of moults simltaneously with other events. In the House

Sparrow, Passer dumesticus breeding and moult were not possible

simltantwusly l x r x a u a of energy constraints (Kendeigh, 1973) and

tlis is probably also the case for many passerines. The breeding

and moult are temporally separated in Finches and htings

(Newton, 1968), Lesser Stripped Swallow (Farle, 19881, Purple

Martin (Niles, 1972) and Towny Owl (Hirons et al., 1984). Payne

(1972) stated that withdrawal of reproductive hormones for birds

with non-overlapping schedules of moulting and breeding showed

initiation of moult, while birds with overlapping schedules of

moult and breeding showed no initiation which is independent of

reproduction.

The primary nwult in the Whitebreasted Kingfisher

starts in July, progresses gradually and is completed in December.

The moult of other feathers including body feathers occurs within

this period tut it is confined to a shorter duration. The growth

of new feathers places additional nutritional demands and in nwst

birds of temperate region moult frequently occurs soon after

breeding (Payne, 1972; Ginn and Melville, 1983). In Towny Owls,

the parents continue to feed their chicks for 2 or 3 months after

fledging and moult appears to start soon after the young leave the

nest ( H i m et al. 1984). In the Whitebreasted Kingfisher, the . nestlings fledge by June and no feeding of young is seen in July

when the primary nwult starts. So it can be concluded that the

breeding and nwult are completely separated in this bird which may

be an adaptive feature of the annual cycle.