SUMMARY
In the course of a 10-year study (1936-1946) of the Yellow-eyed Penguin, Megadyptes antipodes, weights were taken at 7-day intervals from 56 days before the egg was laid to 48 days after hatching-a period of about 146 days. Males reach their peak weight at the end of the off-season and drop steadily during the pre-egg stage until just before egg-laying. Females reach their peak in weight a week later than the males, and for four weeks are heavier than the males, which again exceed the females in weight as the eggs appear. As incubation progresses, both sexes steadily increase in weight, but another drop occurs during the 48 days after the hatching of the chick. Weight fluctuations in two species of petrel, the Broad-billed Prion, Pachyptila vittata [=forsteri], and the Diving Petrel, Pelecanoides urinatrix, were also examined. The same pattern of fluctuation seemed to occur in these two species as in the penguin Megadyptes. The results of the study were compared with the findings of other workers on other species. They were found to follow closely the deductions of Wolfson, who studied migrant Oregon Juncos, Junco oreganus. They differ, however, from data on non-migratory birds, most of which apparently reach peak weights in the winter. Megadyptes resembles Juncos in that a heavy deposit of fat develops as the pre-egg period approaches. In both species this is coincident with growth of the gonads. The pattern of weight fluctuation indicates that incubation in birds is not the arduous task that it is supposed to be, but that feeding of young is a time of stress. The penguin Megadyptes reaches its peak weight for the year at the beginning of the molt, its minimum weight at the end; the petrel Pachyptila apparently does not lose weight during the molt. Evidence from other workers on other species indicates that in some birds there is a gain of weight during the molt; in others, a loss; in still others, no significant change.
ROM August 1936 to May 1946 I carried out a banding study of
the Yellow-eyed Penguin, Megadyptes antipodes, on the Otago
Peninsula, New Zealand. In all, I paid 973 visits to the breeding areas,
which are 20 miles from the city of Dunedin with the outermost col-
onies 20 miles apart. The observations .concerned 88 male and 96 fe-
male penguins which were found breeding for at least one season. A
total of 292 matings was noted.
Weights were recorded as often as possible throughout the year.
Finally, in the tenth season, weights were taken at 7-day intervals, from
a point 56 days before the deposition of the first egg to the end of the
guard stage, when the chicks are abandoned in the daytime and the
two parents fish simultaneously.
In addition, seasonal weights were taken of two species of petrels--
the Broad-billed Prion, Pachyptila vittata [--- forsteri] and the Diving
Petrel, Pelecanoides urinatrix. These weights were taken with much less
thoroughness, but even so, the results should indicate general tendencies
and serve for comparison. The petrel research was carried out on the
tiny island of Whero off the southeast corner of Stewart Island.
In conclusion, I have compared the weights of the petrels and pen-
guins with the findings of other workers on other species. This paper
owes its inspiration largely to the researches of Wolfson (1945). My
thanks are due to Mr. R. K. Wilson, who drew the graphs.
WEIGHT FLUCTUATION IN THE ]PENGUIN MEGADYPTES
The weekly weights of the penguin Megadyptes are given in Tables
1 and 2; their differences are worked out statistically in Table 3. There
is little difference from week to week; only 7 of the 42 differences are
significant3 Four other types of interval were worked out. In weights
taken at two-weekly intervals (Graph 1), 9 differences out of 20 are
significant; at three-weekly intervals, 5 out of 12; and the division of
each stage into two equal parts gives 5 significant differences out of 10.
It will be observed that as the intervals are widened the differences in
weight tend to become statistically more significant. A further tend-
ency, however, is for significant phases in the breeding cycle to be
obscured.
Before allowing that a difference between means is significant, I have followed
the ruling that it must be at least four times the value of the probable error of the
difference.
L.E. WEIGHT FLUCTUATION 161
Richdale
In an endeavor to obviate this factor, I made a final broad group-
ing, as shown in Graph 2. All but one of the eight differences are
significant, and the one exception is nearly so. One important point,
however, is hidden. This is the fact that the females are heavier than
the males for four weeks preceding the laying of the first egg, a phe-
nomenon best appreciated by the use of weekly intervals as shown in
Tables 1 and 2. Graph 2 shows several points between 56 days before
the first egg and 48 days after hatching where a change in weight oc-
curs that is significant statistically. Moreover, a close study of the be-
havior of the sexes during this period shows a correlation with the sta-
tistics.
The period from 42 days before laying to the laying of the first
egg is best discussed in weekly intervals, since wider intervals fail to
reflect important changes in behavior. In the interval 42-36 days be-
fore egg-laying, the males have reached their heaviest weight, after
which a steady decline occurs till the end of the pre-egg period, when
TABLE 1
MEAN WEEKLY WEIGItTS OF MALE PENGUINS FROlVI 56 DAYS BEFORE EGO
TO END OF GUARD STAGE
Class
Stage interval n Mean r 2 PEru s Range
days lbs. lbs. lbs.
Pre-egg 56-50 4 14.13 .70 .23 13 -14.
49-43 2 13.38 .22 .10 13-13
42-36 9 14.22 .45 .10 131-14,
35-29 10 13.82 .83 ,17 121-15
28-22 7 12.50 .77 .20 11-14,
21-15 13 11.96 .46 .08 11 -12,
14- 8 17 11.57 .72 .11 10 /-13
7- 1 17 11.47 .50 .08 10-12,
Post-egg 0- 6 20 12.04 .61 .09 10a/g-13
7-13 33 11.93 .84 .09 10-14
14-20 16 11.90 .73 .12 11 -13
21-27 18 11.79 .59 .09 10a/g-12
28-34 21 12.33 .91 ,13 11 /-14
35-41 25 12.57 .60 ,08 11-13
42-48 20 12.27 .61 .09 11 -13'
Guard 0- 6 31 12.51 .85 .10 11 -14
7-13 i 26 12.59 .84 .11 11-14
14-20 27 12.12 .91 .11 10-14
21-27 25 11.74 .58 .07 10-13
28-34 22 11.47 .59 .08 10-13
35-41 21 11.62 .97 .14 10-14,
42-48 12 11.91 .83 .16 10/-12
x n ---- aumber of weights taken.
1/2 ---- standard deviation.
a PEru = probable error of mean.
162 THE WILSON BULLETIN eptember 1947
Vol. 59, No. 3
TABLE 2
MEAN WEEKLy WEIGHTS OF FEMALE PENGUINS FROM 56 DAYS BEFORE EGG
TO END OF GuAm) STAGE
Class
Stage interval
days
Pre-egg 56-50
49-43
42-36
35-29
28-22
21-15
14- 8
Post-egg 0- 6
7-13
14-20
21-27
28-34
35-41
42-48
Guard 0- 6
7-13
14-20
21-27
28-34
35-41
42-48
Mean
lbs.
12.58
12.75
11.92
13.81
13.30
13.05
12.20
11.57
10.42
10.40
10.76
10.59
10.94
11.19
11.29
11.46
11.26
10.84
11.88
11.27
10.87
10.43
2
.72
.56
.62
.62
.63
.52
.48
.55
.53
.79
.63
.61
.62
.85
.67
.62
.94
.51
.52
P E m 8
.13
.24
.17
.12
.09
.10
.08
.08
.09
.13
.10
.09
.11
.09
.10
.12
.08
.07
Range
lbs.
12-13
121-12
11 -12
12-14
12-14
12 -14
101-12
9 -11
9-11
10 -12
9-11
10 -12
10M-12
10-12¬
10 -12
9a-13
9a-12
10 -12
9a-13,
10 -12
number of weights taken.
standard deviation.
probable error of mean.
their weight tends to increase. From 40 days before the egg appears, the
males begin to stay ashore in the daytime, and they remain there
for much longer periods than the females. Although some of the fe-
males begin to stay ashore quite early, they spend far more time than
their partners at sea, fishing. Because of this fact, the females con-
tinue to increase in weight till the 35-29 day interval.
The result is that the weights of the two sexes in the 35-29 day
interval are nearly equal. After this interval the males are lighter than
the females till the last week of the pre-egg period, when the weights
are once more equal. In the 21-15 day interval, females are on the
average slightly over one pound heavier than males.
With the appearance of the first egg, the story changes abruptly.
In the 7-1 day interval before egg-laying, the males begin to seek
food more often and the females tend to remain ashore. For some days
after the first egg is laid the females rarely enter the water and the
males seldom stay ashore in the daytime. (There are, of course, ex-
ceptions to this general rule.) Thus the females reach their lowest
L.z. WEIGHT FLUCTUATION 163
Richdale
o=
',
Graph 1. Mean weights of Yellow-eyed Penguins at two-weekly intervals from
56 days before egg-laying to the end of the guard stage.
TABLE 3
D:IFF.P, ENCF. S :IN WEEKLy MEANS TAKEN :FP, O3Z[ TABLES 1 AD 2
Male Female
Intervals
Difference PEa 1 Difference PEal
lbs. lbs.
56-50 and 49-43 .75 .251 .17 .13
49-43 and 42-36 .84 .141 .73 .25
42-36 and 35-29 .40 .197 1.89 .34
35-29 and 28-22 1.32 .262 .51 .29,
28-22 and 21-15 .54 .215 .25 .201
21-15 and 14- 8 .39 .136 .85 .151
14- 8 and 7- 1 .10 .136 .63 .13,
7- 1 and 0- 6 .57' .120 1.15 .1D
0- 6 and 7-13 .11 .127 .02 .10
7-13 and 14-20 .03 .150 .36 .11:
14-20 and 21-27 .11 .150 .17 .12
21-27 and 28-34 .54 .158 .35 .151
28-34 and 35-41 .24 .152 .25 ! .16!
35-41 and 42-48 .30 .120 .10 .13,
42-48 and 0- 6 .24 .134 .21 i .14,
0- 6 and 7-13 .08 .149 .20 .131
7-13 and 14-20 .47 .162 .42 .141
14-20 and 21-27 .38 .130 .04 I .13,
21-27 and 28-34 .27 .106 .39 .151
28-34 and 35-41 .15 .163 .40 .14,
35-41 and 42-48 .29 .213 .34 i .101
PEa:probable error of difference.
Statistically significant differences are given in boldface.
164 THE WILSON BULLETIN September 1947
Vol. 59, No. 3
Graph 2. Mean weights of Yellow-eyed Penguins at wide intervals from 56 days
before egg-laying to the end of the guard stage.
weight level during the first two weeks after the egg is laid, whereas
the males reach theirs before the egg is laid.
From these respective depression poifits the weights of both sexes
gradually increase during incubation until the highest peak in weight
for the post-egg period is reached in both sexes (during the interval
from 42-48 days post-egg to 0-6 days guard). Soon after the parents
begin to feed the chicks, a decline occurs. This seems to indicate that
even though the parents themselves acquire food daily (which they did
not during the incubation stage), the additional task of feeding the
chicks has an adverse influence on their weight. At the end of the
guard stage, weights of the adults have once more reached a low level.
The graph for the guard stage tends to be irregular. This is the
most difficult period for which to obtain an even sampling, since there
is a great disparity in weight between a bird that has been ashore for
nearly 24 hours and one that has just returned from feeding. The true
test would be to weigh the birds at a definite period say five or six
hours after they had returned home. This, however, would be an al-
most impossible task.
WEIGHT FLUCTUATION IN PETRELS
Before attempting to interpret Tables 4, 5, and 6 it will be advisa-
ble to summarize briefly the behavior of the species concerned; more
detailed accounts have already been published (Richdale, 1943, 1944,
1945). Pachyptila will be dealt with first. The period from May.12
Richdale
WEIGHT FLUCTUATION
TABLE 4
SEASONAL WEIGItTS OF TIlE BROAD-BILLED PRION
165
Periods Remarks n I Mean
May 12 to 20 Winter months 13 206.6
Aug. 24 to Sept. 2 Laying begins end
of August 19 189.4
Dec. 6 to 16 End of chick-feed-
ing stage 27 189.8
Feb. 7 to March 7 All in molt 22 197.2
er 2
gms.
14.75
16.65
17.65
13.70
2.73
2.54
2.26
1.92
Range
177 to 227
160 to 220
170 to 234
172 to 225
number of weights taken.
standard deviation.
probable error of mean.
to May 20, as shown in Table 4, is the middle of the off-season. All the
birds handled during this period had completed their molt, but their
breeding status was unknown. In the winter months Pachyptila visits
the shore at irregular intervals.
August 24 to September 2 marks the end of the pre-egg stage and
the beginning of laying. One or two of the 19 birds weighed during this
period were incubating; the breeding status of the others was un-
known.
The adults caught between December 6 and 16 were all feeding
chicks (which were near the end of their stay in the burrows). Some
adults were caught before they had fed {heir young and others after-
wards. Since chicks may receive up to 80 grams of food in a night
(Richdale, 1944:194), the range in weight of feeding adults is very
wide.
Finally, the span between February 7 and March 7 is the time
the molt occurs. On February 7, 1942, I was present when Pachyptila
made its first appearance on the island since the completion of the
breeding season a little more than a month earlier. All individuals in-
spected were molting, and all showed considerable wear on the toes--
TABLE 5
DIFFERENCES IN SEASONAL WEIGIITS OF TIlE BROAD-BILLED PRION
Periods Difference PEd I
Between May and August
" August and December
" December and February
" February and May
" December and May
ms.
17.2
.4
7.4
9.4
16.8
3,73
3.40 '
2,96
3.35
3.54
PE d - probable error of difference.
166 THE WILSON BULLETIN September 1947
Vol. 59, No. 3
indicating that they were old birds. Their breeding status was un-
known.
For Pelecanoides (Table 6) only two periods are available. The
first extends from August 23 to 26, which probably represents the be-
ginning of the pre-egg period. The birds were all caught and weighed
at night just after they had landed. Some were known to be breeding
birds, but others were unbanded. Those handled between December
22 and January 30 were all attending chicks and were caught some-
times before and sometimes after they had fed the chicks. For this
reason a wide range in weights is apparent.
TABLE 6
SEASONAL WEIGtITS OF TtIE DrvrNo PETt
Periods Remarks ' n Mean a PE na a Range
gms. gms. gms.
Aug. 23 to 26 End of winter months 26 136.15 6.25 .83 125 to 15;
Dec. 22 to Chick-feeding stage 100 124.10 10.27 .68 108 to 151
Jan. 30
Difference 12.04 ñ 1.07
x n __-- number of weights taken.
2 a: standard deviation.
PEnt: probable error of mean.
The difference between the December and February weights of
Pachyptila, as shown in Tabl 5, is not significant as it stands. When
it is realized, however, that the December weights include, in some
cases, a quantity of undelivered food for the chicks, it is clear that
the real difference would be much greater and possibly significant.
This suggests that the month at sea during which the molt com-
mences is a period of recovery from the stress of feeding young.
During the period of molt, February 7 to March 7, individuals
were weighed on four occasions. These weights did not show any sig-
nificant difference. If this represents the true state of affairs, Pachyp-
tila does not lose weight while molting.
In Pachyptila the difference between the May and December
weights is significant. May is the middle of the off-season, and the
weights would probably be much higher at the beginning of the pre-
egg stage, about the end of July. This is the case in Pelecanoides,
which also shows a significant difference between the early pre-egg
stage and chick stage weights (Table 6). It would seem, then, that
weights in Pachyptila and Pelecanoides rise during the off-season to
a peak at the beginning of the pre-egg stage and drop to a low point
during the. feeding of the chicks.
Finally, Table 4 also indicates that, as in Megadyptes, there is a
depression point at the end of the pre-egg stage in Pachyptila. This
L.E. WEIGHT FLUCTUATION 167
Richdale
is corroborated by the fact that there is a significant difference between
the May and the late August weights.
DISCUSSION
According to Baldwin and Kendeigh (1938:431, 435) most species
of birds show a maximum weight in winter and early spring and a de-
crease at the approach of the breeding season. Nice's observations
(1937:25-26) on Song Sparrows, Melospiza rn1/2lodia, were similar.
Marples (1942 and 1945) indicates a similar trend in the Little Owl,
Athene noctua, and in the Wax-eye, Zosterops lateralis.
Wolfson (1945:109-121), however, o.bjects that these generaliza-
tions are only partially true. He states that there is in Oregon Juncos,
Junco oreganus, and other passerines a significant correlation between
an increase to a maximum body weight and the beginning of the
spring migration. He produces convincing evidence in support of his
contention. His impressive graph (p. 120) indicates that, in the
spring, migrants increase rapidly in weight, whereas residents continue
to decrease. Further, he points out that the increase in weight of the
migrants is due to the deposition of fat, subcutaneously and intraperi-
toneally (p. 109).
The statements of Wolfson are interesting in view of what happens
in Megadyptes. Not only is there a tendency to follow the same pat-
tern of weight fluctuations as migrant passerines, but also, heavy de-
posits of fat develop as the breeding season approaches.
From the beginning of winter to six weeks before egg-laying, there
is a gradual steady increase in weight and fat deposition in both sexes.
This is true not only of the group of penguins as a whole recorded in
Graph 1, but is also true, in the main, of the individuals that I was
able to weigh several times during the period. Further, although no
birds were collected for autopsy, it was obvious that the males, in par-
ticular, were extremely fat by August 24, the beginning of the pre-egg
stage. Even their ability to walk was impaired.
Side by side with this gradual development there is manifested a
growth in the testes and presumably in the ovary. Dissection of two
males killed by a fall of rock toward the end of the winter period
showed that the testes were much enlarged. The physiological state
of the penguins eventually reached a condition that allowed them to
stay ashore for a time without food.
Doubtless there are external as well as internal factors which help
to bring about these changes culminating in the pre-egg stage. If so,
these external factors, one of which is probably the increase in day-
length, apparently act differently on the two sexes. The females reach
their peak weight and fat deposition later than the males and are later
in beginning to stay ashore in the daytime; they do not, at first, fast
as frequently as the males. Wolfson (p. 110) found a comparable situ~
168 THE WILSON BULLETIN September 1947
Vol. 59, No. 3
ation in migrant passerines, a fact which explains why females arrive
on the breeding grounds later than the males.
To sum up, it will be seen that in Megadyptes a steady increase in
body weight and fat deposition precedes the pre-egg stage and that the
females lag behind the males. The sexes, in this way, are prepared for
the behavior patterns which will be released in the succeeding period.
This preparation occurs at the correct time presumably because of the
response of the endocrine glands to external factors in the environment.
The response of the endocrine system then causes a change in the
metabolism which results in an accumulation of surplus fat.
During the last four weeks of the pre-egg stage the female Mega-
dyptes is heavier than the male (Tables 1 and 2). The reason for this
is noted above and is not caused by the advent of the eggs, which aver-
age about 136 grams and are laid four days apart. In the third week
before laying, occurs the greatest disparity in weight between the
sexes, when the females average a little more than one pound heavier.
This advantage ih female weight in species in which the male is the
heavier during most of the year has been recorded elsewhere. When
occurring just before laving it has been thought to be due to the pres-
ence of the eggs (Nice, 1937:27;1938:7. Marples, 1942:247). An-
other possibility is that the male's weight may tend to fall because of
his greater activity, especially noticeable in species which are highly
territorial. Such a possibility is suggested by Nice (I 937: 25).
In species in which only the female incubates there are records that
she continues to be heavier even after the eggs are laid and during the
subsequent incubation. Nice (1937:26-27; 1938:7) found this tend-
ency in Song Sparrows. She also quotes Riddle and Braucher as stat-
ing that there is an eight per cent rise in weight during incubation in
pigeons and doves. From this it would seem that incubation is not the
arduous task that some people think. As for the male, in species in
which he does not share the task of incubation (so far as records are
available), his weight continues to fall. This is recorded in Song Spar-
rows (Nice, 1937:22) and in the Little Owl (Marples, 1942:247).
In Megadyptes, incubation is shared equally by the sexes. The fe-
male remains lighter than the male throughout, and, in fact, at the be-
ginning of the period she reaches one of her two lowest depression
points. This is due to an increase in fasting toward the end of the pre-
egg stage and particularly during egg-laying. From this low level she
rises steadily till the eggs are hatched. The male's weight also registers
a steady rise. These facts support Nice's view that incubation is a
recuperative period. It would seem that in species in which both sexes
incubate both male and female gain in weight. If the female alone in-
cubates, the female alone becomes heavier. A knowledge of what hap-
pens if only the male incubates, as in the phalaropes, would be of con-
siderable interest.
L.E. WEIGHT FLUCTUATION 169
Richdale
With the hatching of the chicks there seems to be a general drop
in weight for at least those parents which feed the young. This
applies to Megadyptes. (In this penguin there is a rise again, how-
ever, after the end of the guard stage, a condition characteristic of
this species and probably of all penguins.) Nice (1937:27) states that
feeding young is strenuous in the two sexes for both Song Sparrows and
Tree Sparrows, Spizella arborea. The same conditions appear to apply
in the Little Owl (Marpies, 1942:pl. 20).
Finally, weights connected with the molt will be considered. Im-
mediately following the breeding season in autumn, the molt com-
mences for most penguins of both sexes. The greatest amount of fat
is developed (being greater than at the end of the winter months)
and the heaviest weight of the yearly cycle is attained just as the molt
commences, but on completion of the molt the birds register the mini-
mum weight for the year.
As regards other families of birds, Nice (1937:22) indicates that
Song Sparrows are "undoubtedly" at their lowest weight in August
and September, when the molt occurs. Beck found a heavy loss at
this time in four domestic fowls (Nice, 1938:7). In the Little Owl,
Marples (1942:247) thinks that the minor loss in weight he recorded in
the autumn was due to molt.
The contrary view is held by Baldwin and Kendeigh (1938:463),
who say that "molting and renewal of feathers in August and Septem-
ber is not joined with a decrease in weight; rather there is an increase
in weight at that time." In support of this, Laskey (Nice, 1938:7)
reports the "highest weight during the inactive period of molting" in
male Mockingbirds, Mimus polyglottos.
In petrels, it would seem from the meagre evidence available that
they reach peak weight at the beginning of the pre-egg stage and de-
crease steadily till the egg is laid. They are again low in weight at
the end of the chick stage, and up to this point the graph of their
weight closely resembles that of Megadyptes. During the molt Pachyp-
tila, at least, does not lose weight.
In conclusion, it is obvious that not all species of birds reach their
maximum weight in the winter months and then decrease as the breed-
ing season approaches. Some, like the migrant passerines, penguins,
and at least some petrels, reach a peak at the beginning of the pre-egg
stage. The weight of incubating birds tends to rise; if a bird does not
incubate, its weight tends to fall. Parents that feed the young also
lose weight, but no data are available on parents that do not feed the
young. As regards the molt, some birds--particularly penguins--cer-
tainly lose weight during the process. On the other hand, there is evi-
dence that some birds do not and that others even gain weight. It is
abundantly clear that data collected on bird weights are of inestimable
value in clarifying many physiological and psychological problems.
LITERATURE CITED
BALDWIN, S. PRE/TISS, and S. CI-IARLES KENDEIGI-I
1938 Variations in the weights of birds. Auk, 55: 416-467.
MARPLES, B. J.
1942 A study of the Little Owl, Athene noctua, in New Zealand. Trans.
Royal Soc. o! N.Z., 72: 237-252.
1945 Zosterops lateralis at Dunedin, New Zealand. Erau, 44: 277-287.
NICE, /ARGARET /ORSE
1937 Studies in the life history of the Song Sparrow. 1. A population study
of the Song Sparrow. Trans. L{nn. Soc. N.Y., 4.
1938 The biological significance of bird weights. Bird-Banding, 9: 1-11.
L.E. WEIGHT FLUCTUATION 171
Richdale
RICIIDALE, L. E.
1943 The Kuaka or Diving Petrel, Pelecanoides urinatrix (Gmelin). Emu,
43: 24-48, 97-107.
1944 The Parara or Broad-billed Prion, Pachyptila viitara (Gmelin). Emu,
43: 191-217.
1945 Supplementary notes on the Diving Petrel. Trans. Royal Soc. of N.Z.,
75: 42-53.
W OLi*SON ALBERT
1945 The role of the pituitary, fat d. eposition, and body weight in bird
migration. Condor, 47: 95-127.
DEPARTMENT 017 Z00LOGY UNIVERSITY 017 0TAG0 DUNEDIN NEW
ZEALAND