Fecundity of female angelfish, Centropyge ferrugatus,

independent of body size: field collection of spawned eggs

Yoichi Sakai

Laboratory of Animal Sociology, Faculty of Science, Osaka City University, Sugimoto 3-3-138, Sumiyoshi-ku,
Osaka 558, Japan
Present address: Biological Laboratory, Faculty of Liberal Arts, Chukyo University, Yagoto, Nagoya 466, Japan
(Received 27 November 1995; in revised form 8 March 1996; accepted 11 April 1996)

Ichthyological
Research
9 The IchthyologicalSociety of Japan 1996

Sakai, Y. 1996. Fecundity of female angelfish, Centropygeferrugatus, independent of body size: field collection of spawned eggs. Ichthyol. Res., 43 (2):
186-189.
Key words.- Pelagic-egg collection; batch fecundity; body size; spawning frequency; protogyny.

any reef fishes, such as wrasses, parrotfishes, groupers and angelfishes, spawn
pelagic eggs into the water column (review in
Thresher, 1984). The fecundity (reproductive effort) of these fishes has been estimated mostly
from gonad weight or the number of mature eggs
in the ovary (Feddern, 1965; Blaxter, 1969;
Robertson and Warner, 1978; Warner and
Robertson, 1978; Wootton, 1990). It is known,
however, that females do not usually release all
ripening eggs in the ovaries in each spawning
event (Feddern, 1965). Therefore, in order to estimate fecundity accurately, the eggs released
during a spawning event should be collected immediately thereafter. However, this has been
conducted only for a few pair-spawning fishes in
their natural habitats (Reinboth, 1973; Bauer and
Bauer, 1981; Shapiro et al., 1994).
The mating behavior of protogynous angelfishes (Pomacanthidae) has been well documented (Thresher, 1984; Moyer, 1990), but the
actual reproductive output from each pairspawning activity remains unknown. The fecundity of individual females of the small angelfish
Centropyge ferrugatus was studied by egg collection in the field, the method of egg collection
being described and relationships between fecundity and body size as well as other ecological
factors discussed herein.

Materials and Methods

The present study was carried out on coral

reefs in front of the Sesoko Marine Science Center of the University of the Ryukyus, Sesoko Is-

land (26~
127~
Okinawa, Japan.
Harems of Centropyge ferrugatus were distributed along the reef edge at depths of 1-5 m
(Sakai, 1996). Each individual was distinguished
by variations in the pattern of spots on the body
and the total length (TL) visually estimated to
the nearest 3 mm. The mean difference between
estimated and measured TLs was 1.5 mm (SD=
1.0, n= 14) for fish collected outside the study
area. The breeding season was from early May
to mid November, male-female pairs releasing
pelagic gametes after a short ascent into the
water column around sunset (Sakai and Kohda,
1995). Females spawned almost daily throughout
the breeding season (Y. Sakai, unpubl, data; see
Results).
The spawned eggs from 12 females in 4
harems were collected in June and July 1991
(water temperature: 25-29~
and from 7 females in 3 of the 4 harems in September 1991
(26-29~
The gamete cloud, visible for about
10 sec, was scooped up to the water surface
(max ascent speed: 20 cm per sec) using a handheld plankton net of 0.4 mm mesh (egg size: ca.
0.7 mm in diameter; Hioki et al., 1990), 47 cm in
mouth diameter and 55 cm in depth. Each eggclutch was secured in a 100 cc bottle at the water
surface, the entire operation taking 30-60 sec.
Eggs of 2 females were thus collected on 14 successive evenings in May. Because the mean
number of released eggs per spawning on the
first 7 evenings (1488 eggs_+878 SD and 1060
eggs_+433) did not differ significantly from
those collected on the subsequent 7 evenings

187

F e c u n d i t y of Angelfish

10 4

i
iHarem 2

Harem 1

10 4

June/July

10 3`

>,
"0
c--1
0

102

co

no sp I
104

June/July
Harem 3

03
0

103

:.~

102

10 3

i
10 2
,

, i

, i

10

11

Body size of f e m a l e s (TL cm)

Fig. 2. Relationship between batch fecundity and

body size of female Centropyge ferrugatus in
June/July 1991. Average and SD are shown for each
female that spawned every 7 evenings (9 and those
which did not spawn on at least one evening (@:
number of spawnings indicated on figure).

C} no sp
0

tO
~'~

10 4

September
Harem 4 (Iqair-group)

10 3

102

104

September

103

102 i

no sp I

ID

Fig. 1. Relationship between lunar cycle and batch

fecundity of female Centropygeferrugatus. Twelve
individuals from 4 harems were surveyed (7 in both
June/July and September). The largest female, second
largest and third largest in each harem are shown by
circles, triangles and squares, respectively (the fourth
female in harem 2 is shown by squares and broken
lines). The third female in harem 4 disappeared on
the fifth evening in June/July. The largest and third
largest in harem 4, the largest in harem 2, and all females in harem 3 all disappeared before the survey in
September. Due to a typhoon, the last day of egg-collection for harem 2 in September was disturbed, no
sp: no spawning; ?: no egg-collection; @: new moon;
9 full moon.

(1392
and 1440
Mann-Whitney Utest, z--0.2 and 0.7, p--0.9 and 0.5), egg-collections for other females were done on 7 succes-

sive evenings. To check all spawnings in a harem

on each evening, a male was followed continuously from 1 hr before sunset until it became inactive. Egg collection failed in 6 (5%) out of 113
spawnings observed. For those females involved,
"total number of eggs per week" were estimated
as follows: total number of eggs during n days
7/n.
After landing, each egg clutch was fixed in
2% formalin, and the number of eggs counted
under a binocular microscope. In this paper, the
number of released eggs per spawning is referred to as "batch fecundity" (Wootton, 1990).

Results and Discussion

Batch fecundity of each female Centropyge
ferrugatus varied day by day from 110 to 2800
(Fig. 1), the average for each female ranging between 740-2080 (Fig. 2). Bauer and Bauer
(1981) reported the batch fecundity of Centropyge spp. in an aquarium to be ca. 100-200,
which seems to be a considerable underestimation, probably due to an inaccuracy in their collection method--the recirculation of the aquarium water, including eggs, through a plankton
net for 6-8 hr. The batch fecundity of C. ferrugatus did not differ between June/July and September (Wilcoxon signed-ranks test, n=7, z=

188

Y. Sakai

0.34, p=0.74; Fig. 1). Female body size was not

significantly correlated with average batch fecundity (Spearman's correlation coefficient,
June/July, G=0.13, n = 12, z=0.44, p=0.66, Fig.
2; September, G=0.23, n--7, z--0.57,p=0.57).
Males performed courtship behavior toward
all females on each evening. However, females
sometimes rejected the courtship and hid in shelters without spawning (Fig. 1); 5 of 12 females
did not spawn on evenings 1-4 of 7 evenings (12
of 84 female-evenings, 14%) in June/July, and in
September 2 of 7 females failed to spawn on 2-3
evenings (5 of 46 female-evenings, 11%). The
spawning frequency was not correlated with female body size (Spearman's correlation coefficient, June/July, G = - 0 . 2 , n=12, z=0.67, p =
0.51; September, G = - 0 . 1 6 , n=7, z=0.39, p =
0.70), nor did it differ between June/July and
September (Wilcoxon signed-ranks test, n = 7 ,
z=0.82, p=0.41). The total number of eggs
spawned by a female over one week did not significantly differ between the harems observed in
different lunar phases (Kruskal-Wallis test:
June/July, nl=3, n2=4, n3--2, n4=3, H=6.3,
p=0.10; September, nl=3, n2=3, n3--1,
H=0.6, p=0.75; Fig. 1). Although lunar periodicity of spawning is known in Centropyge spp.
(Lobel, 1978; Thresher, 1982), this does not appear to be common among angelfishes
(Thresher, 1984).
Because the total number of eggs released
over one week was not significantly correlated
with female body size (Spearman's correlation
coefficient, June/July, G=0.03, n=12, z=0.09,
p=0.93, Fig. 3; September, G=0.11, n=7,
z=0.27, p=0.79), in addition to the absence of
seasonal changes in spawning frequency and
batch fecundity, as mentioned above, it appeared
that female body size may not be related to
spawning success throughout the breeding season. This is contrary to the well-known tendency
for female reproductive ability to generally increase with body size in fishes (reviewed by
Wootton, 1990), and may be related to the occurrence of protogynous sex change in C. ferrugatus (Moyer, 1990; Sakai, 1996). In gonochoristic
species, females have to invest much energy as
possible in egg production, so as to maximize
their lifetime reproductive success; thus larger
females have higher fecundity. In protogynous
species, on the contrary, this would not always

I0s
o
v

N 10'!

0
0

0
0

..Q

E
f0

~--

10: ~
7

i , , , ~

.....

. . . . . . .

i i r l l l l l l l l l l

10

11

Body size of females (TL cm)

Fig. 3. Relationship between total number of eggs

released over 7 days and body size of female Centropyge ferrugatus in June/July 1991. 9 females
which spawned every evening; @: those which did
not spawn on at least one evening.

be the best strategy. Large females may increase

their lifetime reproductive success by investing
energy in growth rather than egg production because larger body size is advantageous for later
becoming a dominant male.

Acknowledgments.- I am grateful to M. Kohda and

T. Kuwamura for support and advice throughout this
study. Critical reading by M. Kohda, T. Kuwamura,
M. Sano, G. S. Hardy and an anonymous reviewer led
the manuscript to significant improvement. I also
thank colleagues of the Animal Sociology Laboratory, Osaka City University, for fruitful discussion. I
appreciated the help given by K. Sakai and the staff
of the Sesoko Marine Science Center (SMSC), University of the Ryukyus, during the field work. This is
Contribution #285 from SMSC.
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189

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