7/25/2019 Sakai 1996

1/4

F e c u n d i t y o f fe m a l e a n g e l fi sh

Cen t r opyge e r r uga t us

i n d e p e n d e n t o f b o d y s iz e : f ie l d c o ll e c ti o n o f s p a w n e d e g g s

Y o i c h i S a k a i

Laboratory o f Animal Sociology Faculty of Science Osaka City University Sugimoto 3-3-138 Sumiyoshi-ku

Osaka 558 Japan

Present address: Biological Laboratory Faculty o f Liberal Arts Chukyo University Yagoto Nagoya 466 Japan

Received 27 November 1995; in revised form 8 March 1996; accep t ed 11 Apr i l 1996

I ch thyo log ica l

Research

9 The IchthyologicalSociety of Japan 1996

Sakai, Y. 199 6. Fecundity o f female angelfish,

Centropyge errugatus

indepen-

dent of body siz e: field col lect ion o f spawned eggs. Ichthyol . Res. , 43 2):

186-189.

K e y w o r d s . -

Pelagic-egg collection; batch fecundity; body size; spawning fre-

quency; protogyny.

any r e e f f is hes , s uch as w ras s es , pa r ro t -

f i s hes , g rouper s and ange l f i s hes , s pawn

p e l a g i c e g g s i n to t h e w a t e r c o l u m n r e v ie w i n

Thres he r , 1984 ) . The f ecu nd i t y r ep roduc t i ve e f -

fo r t) o f t hes e f i s hes has been es t i m a t ed m o s t l y

f r o m g o n a d w e i g h t o r t h e n u m b e r o f m a t u r e e g g s

in the ovary Fedd ern , 1965; Blaxter , 1969;

R o ber t s on and W arner , 1978 ; W arner and

R ober t s on , 1978 ; W oo t t on , 1990 ) . I t i s known ,

however , t ha t f em al es do no t u s ua l l y r e l eas e a l l

r i pen i ng eggs i n t he ova r i e s i n each s pawn i ng

even t F eddern , 1965 ). T here fo re , i n o rde r to e s -

t i m a t e f ecund i t y accu ra t e l y , t he eggs r e l eas ed

d u r i n g a s p a w n i n g e v e n t s h o u l d b e c o l l e c t e d i m -

m e d i a t e l y the rea f t e r. How ever , th i s has bee n

c o n d u c t e d o n l y f o r a f e w p a i r - s p a w n i n g f is h e s i n

t he i r na t u ra l hab i t a t s R e i nbo t h , 1973 ; B au er and

Bau er, 1981; S ha piro et al . , 1994).

T h e m a t i n g b e h a v i o r o f p r o t o g y n o u s a n g e l -

f i shes P om acan t h i da e ) has been we l l docu -

m en t e d Th res he r , 1984; Moyer , 1990 ), bu t t he

ac t ua l r ep roduc t i ve ou t pu t f rom each pa i r -

s p a w n i n g a c t i vi t y re m a i n s u n k n o w n . T h e f e c u n -

d i t y o f i nd i v i dua l f em al es o f the s m a l l ange l f i s h

Centropyge fer ruga tus

w a s s t u d i e d b y e g g c o l -

l ec t i on i n t he f i e l d , t he m e t hod o f egg co l l ec t i on

b e i n g d e s c r i b e d a n d r e l a t i o n s h i p s b e t w e e n f e -

cun d i t y and body s i ze a s we l l a s o t he r eco l og i ca l

f ac t o r s d i s cus s ed he re i n .

a t e r ia l s a n d e t h o d s

T h e p r e s e n t s t u d y w a s c a r r i e d o u t o n c o r a l

r e e f s i n f ro n t o f t h e S e s o k o M a r i n e S c i e n c e C e n -

t e r o f t h e U n i v e r s i t y o f t h e R y u k y u s , S e s o k o I s -

l and 26~ 127~ Ok i naw a , J apan .

H a r e m s o f

Centropyge ferrugatus

were d i s t r i b -

u t e d a l o n g t h e r e e f e d g e a t d e p t h s o f 1 - 5 m

S aka i , 1996) . Each i nd i v i dua l was d i s t i ngu i s hed

b y v a r i a t io n s i n t h e p a t t e rn o f s p o ts o n t h e b o d y

and t he to t a l l eng t h TL) v i s ua l l y e s t i m a t ed to

t h e n e a r e s t 3 m m . T h e m e a n d i f fe r e n c e b e t w e e n

e s ti m a t e d a n d m e a s u re d T L s w a s 1 .5 m m S D =

1 .0 , n= 14 ) fo r f i sh co l l ec t ed ou t s i de t he s t udy

a r e a . T h e b r e e d i n g s e a s o n w a s f r o m e a r l y M a y

t o m i d N o v e m b e r , m a l e - f e m a l e p a i r s r e l e a s i n g

pe l ag i c gam et es a f t e r a s ho r t a s cen t i n t o t he

w a t e r c o l u m n a r o u n d s u n s e t S a k a i a n d K o h d a ,

1995 ). F e m al es s pawn ed a l m os t da i l y t h rou gho u t

t he b reed i ng s eas on Y . S aka i , unpub l , da ta ; s ee

Resul t s ) .

T h e s p a w n e d e g g s f r o m 1 2 f e m a l e s i n 4

h a r e m s w e r e c o l l e c t e d i n J u n e a n d J u l y 1 9 9 1

w a t e r t e m p e r a t u re : 2 5 - 2 9 ~ a n d f r o m 7 f e -

m a l e s i n 3 o f t h e 4 h a r e m s i n S e p t e m b e r 1 99 1

2 6 - 2 9 ~ T h e g a m e t e c l o ud , v i s ib l e f o r a b o u t

10 s ec , was s coo ped up t o t he wa t e r s u r face

m a x a s c e n t s p e e d : 2 0 c m p e r s e c ) u s i n g a h a n d -

h e l d p l a n k t o n n e t o f 0 .4 m m m e s h e g g s i ze : c a .

0 .7 m m i n d i am et e r ; H i ok i e t a l. , 1990 ) , 47 cm i n

m o u t h d i a m e t e r a n d 5 5 c m i n d e p th . E a c h e g g -

c l u t ch was s ecu red i n a 100 cc bo t t l e a t t he w a t e r

s u r face , t he en t i r e ope ra t i on t ak i ng 3 0 -6 0 s ec .

E g g s o f 2 f e m a l e s w e r e t h u s c o l l e c t e d o n 1 4 s u c -

c e s s i v e e v e n i n g s i n M a y . B e c a u s e t h e m e a n

n u m b e r o f r e l e a se d e g g s p e r s p a w n i n g o n t h e

f i rs t 7 even i ngs 1488 eggs_+878 S D an d 1060

eggs _+433) d i d no t d i f f e r s i gn i f i can t l y f rom

t h o s e c o l l e c t e d o n t h e s u b s e q u e n t 7 e v e n i n g s

7/25/2019 Sakai 1996

2/4

F e c u n d i t y o f A n g e l f is h 187

1 4

10 3`

102

n o sp I

104

0 3 1 0 3

0

: .~ 1 0 2

C } n o s p

0 1 4

t -

O

~ ~ 10 3

1 2

9

104

1 0 3

1 2 i

n o sp I

H a r e m 1

H a r e m 3 i

i Ju n e / Ju l y

i H a r e m 2

J u n e / J u l y

H a r e m 4 ( I q a i r - g r o u p )

I

S e p t e m b e r

I

S e p t e m b e r

9

Fig. 1 . Relationship betw een lunar cycle and batch

fecund i ty o f female

Centropyge errugatus.

Twelve

individuals from 4 harems were surveyed (7 in both

June /July and September). Th e largest female, secon d

largest and third largest in each harem are shown by

circles, triangles and squares, respe ctively (the fourth

female in harem 2 is shown by squares and broken

lines). The th ird 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 fe-

males in harem 3 al l d isappeared before the survey in

September. D ue to a typhoon, the las t day of egg-col-

lect ion for harem 2 in Septem ber was dis turbed, no

sp: no spawning; ?: no egg-collection; @: new m oon ;

9 full moon.

( 1 3 9 2 a n d 1 4 4 0 M a n n - W h i t n ey U -

t e st , z - - 0 . 2 a n d 0 . 7, p - - 0 . 9 a n d 0 . 5 ) , e g g - c o l l e c -

t i o n s f o r o t h e r f e m a l e s w e r e d o n e o n 7 s u c c e s -

1 4

>,

0

c-

--1

0

c-

o

m

1 3

1 2

5 6

i i . . . . . . . . . i . . . . . . . . . . . . . . . . . . i

8 9 1 11

Body size of fema les TL cm)

Fig. 2 . Relationship betw een batch fecundity and

body s ize o f female

Centropyge ferrugatus

in

June/July 19 91. Average and S D are show n for each

fema le that spawned every 7 evenings ( 9 and those

which did not spawn on at least one evening (@:

num ber o f spawnings indicated on figure).

s i v e e v e n i n g s . T o c h e c k a l l s p a wn i n g s i n a h a re m

o n e a c h e v e n i n g , a m a l e w a s f o l l o w e d c o n ti n u -

o u s l y f r o m 1 h r b e f o r e s u n s e t u n ti l i t b e c a m e i n-

a c t i v e . E g g c o l l e c t i o n f a i l e d i n 6 (5 % ) o u t o f 1 1 3

s p a w n i n g s o b s e r v e d . F o r t h o s e f e m a l e s i n v o l v ed ,

" t o t a l n u m b e r o f e g g s p e r w e e k " w e r e e s t i m a t e d

a s f o l lo w s : to t a l n u m b e r o f e g g s d u r in g n d a y s

7/n .

Af t e r l a n d i n g , e a c h e g g c l u t c h wa s f i x e d i n

2 % f o r m a l in , a n d t h e n u m b e r o f e g g s c o u n t e d

u n d e r a b i n o c u l a r m i c ro s c o p e . I n t h i s p a p e r , t h e

n u m b e r o f r e l e a s e d e g g s p e r s p a w n i n g i s re -

f e r r e d t o a s " b a t c h f e c u n d i t y " (W o o t t o n , 1 9 9 0 ) .

R e s u l t s a n d i s c u s s i o n

B a t c h f e c u n d i t y o f e a c h f e m a l e

Centropyge

ferrugatus

v a r i e d d a y b y d a y f r o m 1 1 0 t o 2 8 0 0

(F i g . 1 ) , t h e a v e ra g e fo r e a c h f e m a l e r a n g i n g b e -

t w e e n 7 4 0 - 2 0 8 0 ( F ig . 2 ) . B a u e r a n d B a u e r

( 1 9 8 1 ) r e p o r t e d t h e b a t c h f e c u n d i t y o f

Cen

tropyge s p p . i n a n a q u a r i u m t o b e c a . 1 0 0 -2 0 0 ,

w h i c h s e e m s t o b e a c o n s i d e r a b l e u n d e r e s t i m a -

t i o n , p ro b a b l y d u e t o a n i n a c c u ra c y i n t h e i r c o l -

l e c t io n m e t h o d - - t h e r e c i r c u la t i o n o f t h e a qu a r -

i u m wa t e r , i n c l u d i n g e g g s , t h ro u g h a p l a n k t o n

n e t f o r 6 - 8 h r . T h e b a t c h f e c u n d i t y o f C. ferru

gatus d i d n o t d i f f e r b e t w e e n J u n e / J u l y a n d S e p -

t e m b e r ( W i l c o x o n s i g n e d - r a n k s te s t, n = 7 , z =

7/25/2019 Sakai 1996

3/4

188 Y . S a k a i

0 . 3 4 , p = 0 . 7 4 ; F i g . 1 ). F e m a l e b o d y s i ze w a s n o t

s i g n i f i c a n t l y c o r r e l a t e d w i t h a v e r a g e b a t c h f e -

c u n d i t y ( S p e a r m a n ' s c o r r e l a t i o n c o e f f i c i e n t ,

J u n e / Ju l y , G = 0 . 1 3 , n = 12 , z = 0 . 4 4 , p = 0 . 6 6 , F i g .

2 ; S e p t e m b e r, G = 0 . 2 3 , n - - 7 , z - - 0 . 5 7 , p = 0 . 5 7 ) .

M a l e s p e r f o r m e d c o u r t s h i p b e h a v i o r t o w a r d

a l l f e m a l e s o n e a c h e v e n i n g . H o w e v e r , f e m a l e s

s o m e t i m e s r e je c t e d t h e c o u r t s h i p a n d h i d i n s h e l-

t e r s w i t h o u t s p a w n i n g ( F i g . 1 ) ; 5 o f 1 2 f e m a l e s

d i d n o t s p a w n o n e v e n i n g s 1 - 4 o f 7 e v e n in g s ( 1 2

o f 8 4 f e m a l e - e v e n i n g s , 1 4 % ) i n J u n e /J u l y , a n d i n

S e p t e m b e r 2 o f 7 f em a l e s f a i le d t o s p a w n o n 2 - 3

e v e n i n g s (5 o f 4 6 f e m a l e - e v e n i n g s , 1 1 % ) . T h e

s p a w n i n g f r e q u e n c y w a s n o t c o r r e l a t e d w i t h f e -

m a l e b o d y s i z e ( S p e a r m a n ' s c o r r e l a t i o n c o ef fi -

c ie n t, J u ne /J u ly , G = - 0 . 2 , n = 1 2 , z = 0 . 6 7 , p =

0 .5 1 ; S e p te m b e r , G = - 0 . 1 6 , n = 7 , z = 0 .3 9 , p =

0 . 7 0 ), n o r d i d i t d i f fe r b e t w e e n J u n e / J u l y a n d

S e p t e m b e r ( W i l c o x o n s i g n e d -r a n k s te s t, n = 7 ,

z = 0 . 8 2 , p = 0 . 4 1 ) . T h e t o ta l n u m b e r o f e g g s

s p a w n e d b y a f e m a l e o v e r o n e w e e k d i d n o t s i g -

n i f ic a n t l y d if f e r b e t w e e n t h e h a r e m s o b s e r v e d i n

d i f f e r e n t l u n a r p h a s e s ( K r u s k a l - W a l l i s t e s t :

J u n e / J u l y , nl=3 n2=4 n3--2 n4=3 H = 6 . 3 ,

p = 0 . 1 0 ; S e p t e m b e r , nl=3 n2=3 n3--1

H = 0 . 6 , p = 0 . 7 5 ; F i g. 1 ). A l t h o u g h lu n a r p e r io d -

i c i t y o f s p a w n i n g i s k n o w n i n Centropyge s p p .

( L o b e l , 1 9 7 8 ; T h r e s h e r , 1 9 8 2 ) , t h i s d o e s n o t a p -

p e a r t o b e c o m m o n a m o n g a n g e l f i s h e s

( T h r e s h e r , 1 9 8 4 ) .

B e c a u s e t h e t o t a l n u m b e r o f e g g s re l e a se d

o v e r o n e w e e k w a s n o t s i g n i f i c a n t l y c o r r e l a t e d

w i t h f e m a l e b o d y s i z e ( S p e a r m a n ' s c o r r e l a t i o n

c o e f fi c i en t , J u n e / J u ly , G = 0 . 0 3 , n = 1 2 , z = 0 . 0 9 ,

p = 0 . 9 3 , F i g. 3 ; S e p t e m b e r , G = 0 . 1 1 , n = 7 ,

z = 0 . 2 7 , p = 0 . 7 9 ) , i n a d d i t io n to th e a b s e n c e o f

s e a s o n a l c h a n g e s i n s p a w n i n g f r e q u e n c y a n d

b a t c h f e c u n d i t y , a s m e n t i o n e d a b o v e , i t a p p e a r e d

t h a t f e m a l e b o d y s i z e m a y n o t b e r e l a t e d t o

s p a w n i n g s u c c e s s t h r o u g h o u t t h e b r e e d i n g s ea -

s o n . T h i s is c o n t r a r y t o th e w e l l - k n o w n te n d e n c y

f o r f e m a l e r e p r o d u c t i v e a b i l i t y t o g e n e r a l l y i n -

c r e a s e w i t h b o d y s i z e i n f i s h e s ( r e v i e w e d b y

W o o t t o n , 1 9 9 0 ) , a n d m a y b e r e l a t e d t o th e o c c u r -

r e n c e o f p r o t o g y n o u s s e x c h a n g e i n C. ferruga-

tus ( M o y e r , 1 9 9 0 ; S a k a i , 1 9 9 6 ) . I n g o n o c h o r i s t i c

s p e c i e s , f e m a l e s h a v e t o i n v e s t m u c h e n e r g y a s

p o s s i b l e i n e g g p r o d u c t i o n , s o a s t o m a x i m i z e

t h e i r l i f e t i m e r e p r o d u c t i v e s u c c e s s ; t h u s l a r g e r

f e m a l e s h a v e h i g h e r f e c u n d i t y . I n p r o t o g y n o u s

s p e c i e s , o n t h e c o n t r a r y , t h i s w o u l d n o t a l w a y s

I0s

o

v

N 10'

..Q

E

f-

0

~ 10: ~

7

0

9 0

0 0

9

0

0

0 9

i , , , ~ . . . . . i . . . . . . . r l l l l l l l l l l

8 9 10 11

Body size of females TL cm)

F i g . 3 . R e l a t i o n s h i p b e t w e e n to t a l n u m b e r o f e g g s

r e l e a s e d o v e r 7 d a y s a n d b o d y s i z e o f f e m a l e

Cen-

tropyge ferrugatus i n J u n e / J u ly 1 9 9 1 . 9 f e m a l e s

w h i c h s p a w n e d e v e r y e v e n i n g ; @ : t h o s e w h i c h d i d

n o t s p a w n o n a t l e a s t o n e e v e n i n g .

b e t h e b e s t s tr a te g y . L a r g e f e m a l e s m a y i n c r e a s e

t h e i r l i f e t i m e r e p r o d u c t i v e s u c c e s s b y i n v e s t i n g

e n e r g y i n g r o w t h r a th e r t h a n e g g p r o d u c t i o n b e -

c a u s e l a r g e r b o d y s i z e i s a d v a n t a g e o u s f o r l a t e r

b e c o m i n g a d o m i n a n t m a le .

Acknowledgments.-

I a m g r a t e fu l to M . K o h d a a n d

T . K u w a m u r a f o r s u p p o r t a n d a d v i c e t h r o u g h o u t t h is

s tudy . Cr i t i ca l r ead ing by M. K ohda , T . K uw amur a ,

M . S a n o , G . S . H a r d y a n d a n a n o n y m o u s re v i e w e r le d

t h e m a n u s c r i p t t o s i g n if ic a n t i m p r o v e m e n t . I a l s o

t h a n k c o l l e a g u e s o f t h e A n i m a l S o c i o l o g y L a b o r a -

tory , O saka Ci ty U niver s i ty , f o r f r u i t f u l d i scuss ion . I

a p p r e c i a t e d th e h e l p g i v e n b y K . S a k ai a n d t h e s t a f f

o f t h e S e s o k o M a r i n e S c i e n c e C e n t e r ( S M S C ) , U n i -

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