Life History of the Spotfin Killifish, Fundulus luciae ...

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1976

Life History of the Spotfin Killifish, Fundulus luciae (Pisces Life History of the Spotfin Killifish, Fundulus luciae (Pisces

Cyprinodontidae), in Fox Creek Marsh, Virginia Cyprinodontidae), in Fox Creek Marsh, Virginia

Donald M. Byrne College of William and Mary - Virginia Institute of Marine Science

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Recommended Citation Recommended Citation Byrne, Donald M., "Life History of the Spotfin Killifish, Fundulus luciae (Pisces Cyprinodontidae), in Fox Creek Marsh, Virginia" (1976). Dissertations, Theses, and Masters Projects. Paper 1539617475. https://dx.doi.org/doi:10.25773/v5-rs1b-kg72

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LIFE HISTORY OF THE SPOTFIN KILLIFISH,

FUNDULUS LUCIAE (PISCES: CYPRINODONTIDAE),

IN FOX CREEK MARSH, VIRGINIA

A T h e s i s

P r e s e n t e d t o

The F a c u l t y o f t h e S c h o o l o f M a r in e S c i e n c e

The C o l l e g e o f W i l l i a m and Mary i n V i r g i n i a

I n P a r t i a l F u l f i l l m e n t

Of t h e R e q u i r e m e n t s f o r t h e D eg ree o f

M a s t e r o f A r t s

by

D ona ld M i c h a e l Byrne

APPROVAL SHEET

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

t h e r e q u i r e m e n t s f o r t h e d e g r e e o f

M a s t e r o f A r t s

A u t h o r

LIBRARYof

VIRGINIA 1N3TITUTB o f

.M ARINS SCIENCE

A p p r o v e d , J a n u a r y 1976

£ ...Jeftm/A. M u s ic k 7\0h

Jo h n

M a rv in L. Wass

David E . Z w e m e r

B ru c e N e i l s o n

TABLE OF CONTENTS

PageACKNOWLEDGMENTS...................... i v

LIST OF TABLES...................................................................................................... v

LIST OF FIGURES.................................................................................................... v i

ABSTRACT...................................................................................................................... v t i

INTRODUCTION........................................................................................................... 2

STUDY AREA................................................................................................................. 4

METHODS AND MATERIALS..................................................................................... 7

RESULTS................................................................................................................ 11

HABITAT........................................................................................................... 11

SPECIES ASSOCIATIONS........................................................................... 13

MALE REPRODUCTIVE CYCLE.................................................................... 13

FEMALE REPRODUCTIVE CYCLE............................................................... 16

SPAWNING BEHAVIOR.................................................................................. 21

SPAWNING CONDITIONS AND SIT E S .................................................... 23

DEVELOPMENT AND GROWTH...................................................................... 23

DIET................................................................................................................... 32

PARASITES...................................................................................................... 36

HYBRIDIZATION............................................................................................. 41

DISCUSSION................................................................................................................ 42

LITERATURE CITED................................................................................................. 51

ACKNOWLEDGMENTS

I w i s h t o e x p r e s s my a p p r e c i a t i o n t o J o h n M u s ic k ,

my m a j o r p r o f e s s o r , f o r h i s g u i d a n c e d u r i n g t h i s

i n v e s t i g a t i o n , and J o h n M e r r i n e r , D av id Z w e r n e r , M arv in

Wass an d B ru c e N e i l s o n , t h e members o f my t h e s i s c o m m i t t e e ,

f o r t h e i r c r i t i c a l r e v i e w o f t h e m a n u s c r i p t . Dave Z w e rn e r

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

p a r a s i t e s .

I am a l s o i n d e b t e d t o Ken A b l e , who h e l p e d i n i t i a t e

t h e s t u d y , Mike W i l l i a m s , f o r h i s c a r e f u l i l l u s t r a t i o n s ,

a n d D ebb ie S p r i n k l e , f o r t y p i n g t h e m a n u s c r i p t .

LIST OF TABLES

Table Page1. F e c u n d i t y o f F u n d u l u s l u c i a e by t o t a l

l e n g t h i n c r e m e n t s and egg s i z e . ............................. 17

2 . F r e q u e n c y d i s t r i b u t i o n f o r eggs > 1 .6 mmd i a m e t e r i n f e m a le s c o l l e c t e d a tFox C r e e k Marsh f rom A p r i l t h r o u g hA u g u s t and mean num ber o f e gg s p e rf e m a le f o r e a c h c o l l e c t i o n d a t e ........................... 18

3 . V a r i a t i o n i n t h e d i a m e t e r and num ber o fo i l g l o b u l e s p e r egg o f 40 f e r t i l i z e d F u n d u lu s l u c i a e e g g s .....................................................19

4 . M o r p h o l o g i c a l and m e r i s t i c m e a s u r e m e n ts o fl a r v a l a n d j u v e n i l e F u n d u l u s l u c i a e ................ 29

5 . D i e t o f F u n d u lu s l u c i a e ........................... 33

6 . M e tazo an p a r a s i t e s o f F u n d u lu s l u c i a e .......................... 37

7 . L o c a t i o n and f r e q u e n c y o f p a r a s i t e s on341 F u n d u l u s l u c i a e ........................................................ 39

8 . Growth o f F u n d u l u s h e t e r o c l i t u s andF u n d u lu s d i a p h a n u s . . .4 4

v

LIST OF FIGURES

Figure Page1. Map o f t h e York R i v e r sh o w in g l o c a t i o n s

w h e re F u n d u l u s l u c i a e was c o l l e c t e d ............... 6

2 . D i s t r i b u t i o n o f c o n t a c t o r g a n s on ana d u l t m a le F u n d u l u s l u c i a e .......................................15

3 . M a tu r e u n f e r t i l i z e d ovum, 1 .9 2 mm i nd i a m e t e r , o f F u n d u lu s l u c i a e ..................................20

4 . F u n d u l u s l u c i a e l a r v a e a t d i f f e r e n t s t a g e so f d e v e l o p m e n t .....................................................................28

5 . D o r s a l p i g m e n t a t i o n p a t t e r n s o f F u n d u lu sh e t e r o c l i t u s and F . l u c i a e l a r v a e ..................... 31

ABSTRACT

F u n d u lu s l u c i a e , a r a r e and p o o r l y known c y p r i n o d o n t , was c o l l e c t e d from a s i n g l e s i t e t h r o u g h o u t a y e a r a t Fox C re e k Marsh a n d , o c c a s i o n a l l y , f rom o t h e r s a l t m a r s h e s a l o n g t h e York R i v e r , V i r g i n i a . The s p e c i e s o c c u r r e d in s h a l l o w d i t c h e s , m u d h o le s and t i d a l r i v u l e t s l o c a t e d i n d e n s e s t a n d s o f S p a r t i n a a l t e m i f l o r a i n h i g h i n t e r t i d a l a r e a s . T h ese h a b i t a t s w e re b r a c k i s h ( 0 . 2 - 1 2 . 7 / 7 ) , so m e t im e s oxygen d e f i c i e n t ( < 1 m g/1) and e x h i b i t e d s e a s o n a l e x t r e m e s i n t e m p e r a t u r e ( 4 - 3 6 ° C ) .

At Fox C re e k Marsh t h e sp a w n in g s e a s o n e x t e n d e d f rom m i d - A p r i l t o m i d - A u g u s t . F e m a le s c o n t a i n e d ova e x h i b i t i n g c o n t i n u o u s g r a d a t i o n o f s i z e , b u t t h e num ber o f m a t u r e and i n c i p i e n t m a t u r e ova ( > 1 . 6 mm d i a m e t e r ) n e v e r e x c e e d e d 16 p e r f i s h . M ales p o s s e s s e d c o n t a c t o r g a n s on a r e a s f r e q u e n t l y i n c o n t a c t w i t h t h e f e m a l e d u r i n g c o u r t s h i p .

F . l u c i a e l a r v a e w e re o b t a i n e d f rom l a b o r a t o r y f e r t i l i z e d e gg s and t h e Fox C r e e k Marsh c o l l e c t i o n s i t e . M e r i s t i c and m o r p h o l o g i c m e a s u r e m e n t s w e re made on 27 s p e c i m e n s r a n g i n g f rom 5 . 3 t o 1 7 . 0 mm TL (TL = t o t a l l e n g t h ) .

The j u v e n i l e s t a g e was a t t a i n e d a t 11 -13 mm TL.S e x u a l m a t u r i t y o c c u r r e d a t 2 4 -2 7 mm TL f o r m a l e s and 2 8 - 3 0 mm TL f o r f e m a l e s . The maximum s i z e a t t a i n e d by m a l e s was 40 mm TL and by f e m a l e s , 47 mm TL. A g ing w i t h s c a l e s was u n s u c c e s s f u l , b u t t h e l i f e sp a n o f t h e s p e c i e s i s p r o b a b l y a b o u t one y e a r .

A n a l y s i s o f t h e c o n t e n t s o f 303 d i g e s t i v e t r a c t s showed l u c i a e was o m n iv o r o u s . D e t r i t u s , d i a t o m s , o s t r a c o d s , d i p t e r a n s and c o p e p o d s c o n s t i t u t e d t h e m a j o r c o m p o n e n ts o f t h e d i e t .

E v e r y m a j o r m e t a z o a n p a r a s i t e g r o u p , e x c e p t C e s t o d a , was r e p r e s e n t e d on F. l u c i a e . Monogenea w ere t h e m o s t n u m e r o u s , w i t h 5 7 . 2%~i n c i d e n c e .

F . l u c i a e i s a h a r d y e u r y h a l i n e s p e c i e s c a p a b l e o f t o l e r a t i n g r i g o r o u s e n v i r o n m e n t a l c o n d i t i o n s and u t i l i z i n g a v a r i e t y o f fo o d s o u r c e s . I t i s n o t r a r e , b u t a p p a r e n t l y r e s t r i c t e d t o h i g h i n t e r t i d a l m a r s h a r e a s w h e re i t may h a v e b e e n o v e r l o o k e d p r e v i o u s l y . I t s s m a l l s i z e , s h o r t l i f e sp a n and low f e c u n d i t y p r o b a b l y p r e v e n t a more w i d e s p r e a d d i s t r i b u t i o n .

* •v n

LIFE HISTORY OF THE SPOTFIN KILLIFISH,

FUNDULUS LUCIAE (PISCES: CYPRINODONTIDAE),

IN FOX CREEK MARSH, VIRGINIA

I n t r o d u c t i o n

Members o f t h e f a m i l y C y p r i n o d o n t i d a e o c c u r t h r o u g h o u t

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

a l s o i n s a l t and b r a c k i s h w a t e r . I n N o r t h A m e r ic a t h e

f a m i l y i s r e p r e s e n t e d by s e v e r a l g e n e r a , o f w h ic h t h e

l a r g e s t i s F u n d u l u s . A lo ng t h e e a s t c o a s t o f t h e U n i t e d

S t a t e s F u n d u lu s and o t h e r k i l l i f i s h s p e c i e s a r e common

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

s a l t and b r a c k i s h m a r s h e s . I c h t h y o f a u n a l s u r v e y s o f s u c h

a r e a s f r e q u e n t l y show k i l l i f i s h e s t o be among t h e m o s t

a b u n d a n t f i s h e s c o l l e c t e d ( R i c h a r d s and C a s t a g n a 1 9 7 0 ) .

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

a d a p t a b i l i t y , an d h a r d i n e s s o f t h e s e f i s h e s h a v e made them

s u c c e s s f u l l a b o r a t o r y s u b j e c t s f o r a v a r i e t y o f r e s e a r c h

t o p i c s i n e m b r y o lo g y , g e n e t i c s , p h y s i o l o g y , e n d o c r i n o l o g y ,

c y t o l o g y and b e h a v i o r . D e s p i t e s u c h e x t e n s i v e l a b o r a t o r y

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

h e t e r o c l i t u s , t h e r e h a v e b e e n few f i e l d s t u d i e s o f n a t u r a l

p o p u l a t i o n s . L i t t l e i s known a b o u t t h e l e s s n um erous and

c o n s e q u e n t l y l e s s a v a i l a b l e s p e c i e s , su c h a s F u n d u lu s

l u c i a e , t h e s p o t f i n k i l l i f i s h .

The s c i e n t i f i c l i t e r a t u r e p e r t a i n i n g t o F u n d u lu s

l u c i a e c o n s i s t s m a i n l y o f c o l l e c t i o n r e c o r d s and l a b o r a t o r y

s t u d i e s o f t h e g e n u s F u n d u l u s . B a i r d (18 5 5 ) d e s c r i b e d t h e

2

3

s p e c i e s f rom sp e c im e n s c o l l e c t e d a t B e a s e l y P o i n t , New

J e r s e y . I t h a s s u b s e q u e n t l y b e e n r e p o r t e d f rom Long I s l a n d

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

s a l t m a r s h e s (S m i th 1892 , F o w le r 1912 , 1914 , 1927; C ra w f o r d

1 92 0 ; N i c h o l s and B r e d e r 1927; H i l d e b r a n d an d S c h r o e d e r

1928; H i l d e b r a n d 1941; B u t n e r and B r a t t s t r o m 1960; R i c h a r d s

a n d B a i l e y 1967; J o r g e n s o n 1 9 6 9 ) . The s p e c i e s h a s b e e n

i n c l u d e d i n g e n e r i c s t u d i e s o f taxonomy (Brown 1 9 5 4 ) ,

k a r y o l o g y (Chen 1 9 7 1 ) , s a l i n i t y t o l e r a n c e ( G r i f f i t h 1974)

a n d e r y t h r o c y t e m o rp h o lo g y ( S r i v a s t a v a and G r i f f i t h 1 9 7 4 ) .

The m o s t c o m p r e h e n s iv e a c c o u n t o f i t s b i o l o g y , i n c l u d i n g

s to m a c h c o n t e n t s , r e p r o d u c t i v e s e a s o n and egg s i z e was

g i v e n by H i l d e b r a n d and S c h r o e d e r ( 1 9 2 8 ) .

S e v e r a l a u t h o r s r e p o r t e d F u n d u l u s l u c i a e a s r a r e

( H i l d e b r a n d and S c h r o e d e r 1928; R i c h a r d s and B a i l e y 1 9 6 7 ) .

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

R i v e r , G l o u c e s t e r C o u n ty , V i r g i n i a p r o v i d e d t h e i m p e t u s f o r

a l i f e h i s t o r y s t u d y o f t h i s p u r p o r t e d l y r a r e s p e c i e s . A

s u r v e y o f m a r s h e s a l o n g t h e York R i v e r l o c a t e d o t h e r

p o p u l a t i o n s and one o f t h e s e , i n Fox C r e e k M a rsh , was

s e l e c t e d f o r s t u d y . H a b i t a t c h a r a c t e r i s t i c s , f o o d h a b i t s ,

f e c u n d i t y , sp a w n in g s e a s o n , g r o w th and d e v e l o p m e n t ,

b e h a v i o r , s p e c i e s a s s o c i a t i o n s and p a r a s i t e s o f t h i s

k i l l i f i s h a r e d e s c r i b e d h e r e i n .

S tu d y A rea

Fox C r e e k Marsh ( F i g u r e 1) i s l o c a t e d on t h e n o r t h

b a n k o f t h e York R i v e r , a p p r o x i m a t e l y 27 km f rom t h e m o u th .

S m a l l f r e s h w a t e r s t r e a m s d r a i n i n t o t h e m a r s h f rom

s u r r o u n d i n g w oods . B r a c k i s h w a t e r f rom t h e Y ork R i v e r f l o w s

i n w i t h e v e r y f l o o d t i d e and i s d i s t r i b u t e d t h r o u g h o u t t h e

m a r s h v i a a m e a n d e r i n g n e t w o r k o f l a r g e and s m a l l t i d a l

c r e e k s . N e a r t h e r i v e r , s a l t m a r s h c o r d g r a s s ( S p a r t i n a

a l t e r n i f l o r a ) c o v e r s t h e m a rs h s u r f a c e , e x t e n d i n g f rom t h e

m a in t i d a l c r e e k s t o s t e e p wooded b a n k s t h a t d e l i m i t t h e

m a r s h b o u n d a r i e s . I s o l a t e d s t a n d s o f s a l tm e a d o w hay

( S p a r t i n a p a t e n s ) , m a r s h e l d e r ( I v a f r u t e s c e n s ) and

g r o u n d s e l t r e e (B a c c h a r i s h a l i m i f o l i a ) a r e s c a t t e r e d w i t h i n

t h e p r o f u s i o n o f c o r d g r a s s . T h i s v e g e t a t i o n p a t t e r n e x t e n d s

u p s t r e a m f o r a b o u t 1 km u n t i l c o r d g r a s s i s e v e n t u a l l y

r e p l a c e d by n a r r o w - l e a v e d c a t t a i l (Typha a n g u s t i f o l i a ) .

W i t h i n t h e c o r d g r a s s - c a t t a i l t r a n s i t i o n zone a n d a l o n g t h e

m a r s h b o u n d a r i e s o t h e r p l a n t s p e c i e s o c c u r s u c h a s swamp

r o s e m a l lo w (H i b i s c u s m o s c h e u t o s ) , s e a s h o r e m a l lo w

K o s t e l e t z k y a v i r g i n i c a ) , sm a r tw ee d ( Polygonum l a p a t h i f o l i u m ) ,

mock b i s h o p ’s weed (P t i l i m n i u m c a p i l l a c e u m ) and s a l t m a r s h

b u l r u s h ( S c i r p u s r o b u s t u s ) . T r e e s and s h r u b s on t h e b a n k s

i n c l u d e r e d c e d a r ( J u n i p e r u s v i r g i n i a n a ) , h o l l y ( I l e x o p a c a ) ,

m o u n t a i n l a u r e l (K a lm ia l a t i f o l i a ) , r e d m a p le (A c e r ru b r u m ) ,

5

S p a n i s h oak (Q u e rc u s d i g i t a t a ) , swamp m a g n o l i a (M a g n o l i a

v i r g i n i a n a ) , wax m y r t l e (M y r i c a c e r i f e r a ) , t u p e l o (N y ssa

s y l v a t i c a ) and l o b l o l l y p i n e (P in u s t a e d a ) .

A l o c a l i z e d c o n c e n t r a t i o n o f F u n d u lu s l u c i a e was fo u nd

amid t h e d e n s e c o r d g r a s s g ro w th i n a h i g h m a r s h a r e a w h ic h

was n e a r b o t h a wooded b a n k and t h e c o r d g r a s s - c a t t a i l

t r a n s i t i o n z o n e . The l o c a l e p o s s e s s e d num erous p u d d l e s ,

w a t e r - f i l l e d m u d h o le s and t i d a l r i v u l e t s i n w h ic h F u n d u lu s

l u c i a e o c c u r r e d and f rom w h ic h a l l o f t h e Fox C r e e k Marsh

s p e c im e n s w e re o b t a i n e d .

6

F i g u r e 1 . Map o f t h e Y o rk R i v e r sho w in g l o c a t i o n s w h e re

F u n d u l u s l u c i a e was c o l l e c t e d : A, Ware R i v e r ;

B, C a rm in es I s l a n d s ; C, Fox C r e e k M arsh ;

D, Roane M arsh ; E, York tow n C r e e k . The i n s e t

shows t h e l o c a t i o n o f t h e Y o rk R i v e r i n

r e l a t i o n t o t h e C h e sa p e a k e Bay a r e a .

37*

00* i 3 MILES

M ethods and M a t e r i a l s

A d u l t F u n d u l u s l u c i a e w e re c o l l e c t e d a t Fox C r e e k Marsh

w i t h a d i p n e t (38 cm d i a m e t e r , 3 mm m esh) f rom 2 J u n e 1973

t h r o u g h 8 J u l y 1974 . Two a d d i t i o n a l c o l l e c t i o n s w e re made

on 27 J u l y and 12 A u g u s t 1975 t o p r o v i d e m a t e r i a l from a

s p a r s e l y r e p r e s e n t e d t im e p e r i o d . A t o t a l o f 24 s a m p le s

w e re c o l l e c t e d d u r i n g d a y l i g h t h o u r s a t low w a t e r l e v e l s .

Sample s i z e was k e p t s m a l l t o a v o i d d e p l e t i n g t h e p o p u l a t i o n .

A t l e a s t f i v e i n d i v i d u a l s o f e a c h s e x w e re s e l e c t e d t o

o b t a i n e q u a l r e p r e s e n t a t i o n o f t h e d i m o r p h i c s e x e s and

v a r i e t y i n s i z e . D u r in g c o l d w e a t h e r f i s h a p p e a r e d s c a r c e

and f e w e r w e re c o l l e c t e d . S a l i n i t y and d i s s o l v e d oxygen

s a m p le s w ere t a k e n i n c o n j u n c t i o n w i t h e a c h f i s h c o l l e c t i o n .

S a l i n i t y was d e t e r m i n e d w i t h a Beckman RS7B i n d u c t i o n

s a l i n o m e t e r and d i s s o l v e d oxygen by t h e W i n k l e r t i t r a t i o n

m e th o d ( S t r i c k l a n d and P a r s o n s 1 9 6 8 ) . W a te r t e m p e r a t u r e

was m e a s u r e d w i t h a m e r c u r y s t e m t h e r m o m e t e r .

F i s h w e re p r e s e r v e d i m m e d i a t e l y a f t e r c a p t u r e i n 10%

b u f f e r e d f o r m a l i n and s t o r e d f o r l a b o r a t o r y e x a m i n a t i o n .

T o t a l l e n g t h o f e a c h f i s h was r e c o r d e d and a p a t c h o f s c a l e s

f ro m t h e s c a l e rows a b o v e t h e l a t e r a l l i n e , j u s t a n t e r i o r t o

t h e i n s e r t i o n o f t h e d o r s a l f i n , on t h e l e f t s i d e , w e re

removed f o r a g i n g . S c a l e s w e re m oun ted on g l a s s s l i d e s i n

H o y e r ’ s medium (Humason 1972) and e x am in ed w i t h b o t h a

8

m i c r o s c o p e a n d s c a l e p r o j e c t i o n m a c h in e f o r g r o w th m a r k s .

O b s e r v a t i o n s o f r e p r o d u c t i v e s t a t e , d i g e s t i v e t r a c t

c o n t e n t s and p a r a s i t e s w e re r e c o r d e d f rom t h e p r e s e r v e d

s p e c i m e n s . Male r e p r o d u c t i v e r e a d i n e s s was d e t e r m i n e d by

t h e p r e s e n c e o r a b s e n c e o f c o n t a c t o r g a n s (W i ley and

C o l l e t t e 1970) and m a c r o s c o p i c a p p e a r a n c e o f t h e t e s t e s .

F e c u n d i t y was m e a s u r e d by t e a s i n g t h e o v a r y a p a r t and

c o u n t i n g a l l e g g s w i t h a d i a m e t e r ^ 0 .3 2 mm. Eggs w ere

m e a s u r e d w i t h an o c u l a r m i c r o m e t e r and g r o u p e d i n t o t h r e e

s i z e c a t e g o r i e s : 0 . 3 2 - 0 . 9 6 mm, 0 . 9 7 - 1 . 6 mm and > 1 .6 mm.

The d i g e s t i v e t r a c t was e x c i s e d and e m p t i e d upon a g l a s s

s l i d e . C o n t e n t s w e re i d e n t i f i e d and e n u m e r a t e d w i t h

m i c r o s c o p i c e x a m i n a t i o n . The f i n s , body s u r f a c e , g i l l s and

v i s c e r a w e re e x am ined f o r m e t a z o a n p a r a s i t e s . Each p a r a s i t e

e n c o u n t e r e d , e x c e p t t h e m o n o g e n e t i c and d i g e n e t i c t r e m a t o d e s ,

was i d e n t i f i e d t o s p e c i e s i f p o s s i b l e by t h e m e th o d s o f

F e r n a n d o e t a l . (1972) and t h e k e y s o f C r e s s e y ( 1 9 7 2 ) ,

Hoffman ( 1 9 6 7 ) , Meehean ( 1 9 4 0 ) , R o b e r t s ( 1 9 7 0 ) , Y am agu t i

(1 9 6 1 , 1 9 6 3 a , 1963b) and Y orke and M a p l e s to n e ( 1 9 6 2 ) . A

few i n d i v i d u a l s o f e a c h morph o f t h e a b u n d a n t mono- and

d i g e n e t i c t r e m a t o d e s w e re s e l e c t e d a s a su b s a m p le f o r

i d e n t i f i c a t i o n .

S i n g l e c o l l e c t i o n s w e re made a t Ware C r e e k (4 March

1 9 7 3 ) , Roane M arsh (24 J u n e 1973) and Y ork tow n C r e e k (30

J u n e 1 9 7 3 ) . S p ec im en s f rom t h e s e l o c a l i t i e s w e re t r e a t e d i n

t h e same f a s h i o n a s t h o s e f rom Fox C r e e k M arsh and d a t a w ere

com bined f o r t h e r e s p e c t i v e t im e p e r i o d s .

9

L a r v a l and j u v e n i l e F u n d u l u s l u c i a e w e r e c o l l e c t e d

f rom Fox C r e e k M arsh w i t h an a q u a r iu m d i p n e t ( 0 .3 5 mm m e s h ) .

S p e c im en s w e re i m m e d i a t e l y p r e s e r v e d i n 5% b u f f e r e d f o r m a l i n .

A d d i t i o n a l l a r v a e w e re p r o c u r e d f rom e g gs spawned i n a q u a r i a

by c a p t i v e a d u l t s . Newly h a t c h e d l a r v a e w e re o b t a i n e d by

s t r i p p i n g and f e r t i l i z i n g e gg s i n p l a s t i c p e t r i d i s h e s . The

e g g s d e v e l o p e d and h a t c h e d a t room t e m p e r a t u r e i n b r a c k i s h

w a t e r ( 3 . 9 ^ ) .

A t o t a l o f 27 l a r v a l and j u v e n i l e f i s h , 5 . 3 - 1 7 . 0 mm i n

t o t a l l e n g t h , w e re ex am in e d u n d e r a s t e r o s c o p i c d i s s e c t i o n

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

w i t h g r o w th and d e v e l o p m e n t . M e asu rem en ts o f t o t a l l e n g t h ,

s t a n d a r d l e n g t h , d e p t h , h e a d l e n g t h , ey e d i a m e t e r and s n o u t

l e n g t h w e re made on t h e l e f t s i d e o f e a c h s p e c im e n w i t h an

o c u l a r m i c r o m e t e r . C o u n ts w e r e made o f v e r t e b r a e ,

b r a n c h i o s t e g a l r a y s and f i n r a y s .

R e p r o d u c t i v e and f e e d i n g b e h a v i o r w e re o b s e r v e d i n t e n

g a l l o n a q u a r i a f i l l e d w i t h w a t e r o f 0 . 1 , 3 . 9 and 16.94?

s a l i n i t y . The a q u a r i a w e re e s t a b l i s h e d 21 A p r i l 1974 and

m a i n t a i n e d a t a m b i e n t t e m p e r a t u r e and p h o t o p e r i o d u n t i l

9 J u n e 1974 . Each a q u a r i u m c o n t a i n e d 2 - 3 sp a w n in g mops and

u p t o 15 a d u l t f i s h f ro m Fox C r e e k M arsh , o n e - t h i r d o f

w h i c h w e re m a l e . E ach sp a w n in g mop was c o n s t r u c t e d o f

a p p r o x i m a t e l y 120 20 cm l o n g s t r a n d s o f g r e e n a c r y l i c y a r n

bou n d t o g e t h e r and a n c h o r e d w i t h a c l e a n o y s t e r s h e l l i n

t h e s a n d c o v e r i n g t h e t a n k b o t to m . The mops w e re

p e r i o d i c a l l y e x am in e d f o r e g g s . T h o se d i s c o v e r e d w e re

10

rem oved t o p l a s t i c p e t r i d i s h e s f o r d e v e l o p m e n t and h a t c h i n g .

The d i s h e s w e re k e p t a t room t e m p e r a t u r e and c o n t a i n e d w a t e r

f rom t h e t a n k i n w h ic h t h e e g g s had b e e n spaw ned . A d u l t

f i s h w e re f e d b r e a d c ru m b s , c h o p p ed f r o z e n sh r im p and

ch o p ped l i v e e a r t h w o r m s . W a te r was f i l t e r e d and a e r a t e d by

a p l a s t i c f i l t e r b o x w i t h i n t h e t a n k .

R e s u l t s

H a b i t a t

The h a b i t a t o c c u p i e d by F u n d u l u s l u c i a e was s i m i l a r a t

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

d i t c h e s , s m a l l p o o l s and t i d a l r i v u l e t s w i t h s o f t mud-

d e t r i t u s b o t t o m s . T h ese a r e a s w ere u s u a l l y s i t u a t e d among

d e n s e s t a n d s o f S p a r t i n a a l t e m i f l o r a and c o n t a i n e d s h a l l o w

w a t e r , se ld om d e e p e r t h a n 0 . 5 m and f r e q u e n t l y o n ly a few

c e n t i m e t e r s d e e p , e x c e p t d u r i n g h i g h s p r i n g o r s t o r m t i d e s .

F i s h w e re o c c a s i o n a l l y o b s e r v e d p a r t i a l l y e m e rg e d , w r i g g l i n g

f rom one p u d d l e t o a n o t h e r t h r o u g h w a t e r l e s s t h a n 1 cm i n

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

o f f i r m mud f rom w h ic h t a n g l e d m a t s o f S p a r t i n a r o o t s o f t e n

p r o t r u d e d . C a v e - l i k e i n d e n t a t i o n s o c c u r r e d w h e re t h e m a r s h

h a d b e e n u n d e r c u t and p a r t i a l l y subm erged wads o f d e c a y i n g

v e g e t a t i o n w e re p r e s e n t . T h e s e f e a t u r e s and t h e s o f t

b o t t o m p r o v i d e d c o v e r f o r t h e f i s h , w h i c h w ou ld d a r t i n t o

them and h i d e when d i s t u r b e d .

W a te r d e p t h a t t h e Fox C r e e k Marsh c o l l e c t i o n s i t e was

d e p e n d e n t on t i d e h e i g h t . H ig h s p r i n g and s t o r m t i d e s

i n u n d a t e d t h e m a r s h , i n c l u d i n g t h e a r e a w h e re l u c i a e was

c o n c e n t r a t e d . A f u t i l e c o l l e c t i o n a t t e m p t was made on one

s u c h o c c a s i o n and s p e c im e n s w e r e n o t c a p t u r e d i n t h e u s u a l

11

12

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

A p p a r e n t l y t h e f i s h d i s p e r s e d o v e r t h e f l o o d e d m a rs h

s u r f a c e . O p p o r t u n i t i e s t o d i s p e r s e w e re l i m i t e d b e c a u s e n o t

a l l h i g h t i d e s subm erged t h e h i g h m a r s h . Some p r o d u c e d o n l y

s l i g h t i n c r e a s e s i n d e p t h and o t h e r s had no e f f e c t w h a t e v e r .

W a te r c u r r e n t s and s a l i n i t y a t Fox C r e e k M arsh w ere

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

t i d e s , f r e s h w a t e r f l o w e d c o n t i n u o u s l y , b u t w e a k l y , t h r o u g h

t h e l u c i a e h a b i t a t and d i l u t e d t h e b r a c k i s h w a t e r . The

l o w e s t s a l i n i t y r e c o r d e d was 0.24? and v a l u e s l e s s t h a n 1-4

w e re common. When h i g h t i d e s f l o o d e d t h e m a r s h , s t r o n g

c u r r e n t s w ere c r e a t e d i n t h e d i t c h e s and r i v u l e t s by t h e

e n s u i n g e b b . Such t i d e s w e re a l s o a c c o m p a n ie d by i n c r e a s e s

i n s a l i n i t y . The h i g h e s t s a l i n i t y r e c o r d e d was 1 2 . 7C.

W a te r t e m p e r a t u r e and d i s s o l v e d oxygen a t Fox C r e e k

M arsh w e re e x t r e m e l y v a r i a b l e t h r o u g h o u t t h e y e a r . W a te r

t e m p e r a t u r e r a n g e d f rom 4 t o 3 6 °C. I c e fo rm ed d u r i n g w i n t e r

c o l d s p e l l s , b u t was n o t l o n g - l a s t i n g and was se ld o m t h i c k e r

t h a n a few m i l l i m e t e r s . D i s s o l v e d oxygen r a n g e d f rom l e s s

t h a n 1 t o 6 . 8 m g /1 .

F . l u c i a e was p r e s e n t a t t h e Fox C r e e k M arsh c o l l e c t i o n

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

c o n d i t i o n s . I t a p p e a r e d m o s t a b u n d a n t i n s p r i n g and summer,

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

and d a r t i n g t h r o u g h t h e w a t e r . A t su c h t i m e s many f i s h

w e r e e a s i l y c a p t u r e d w i t h a few sweeps o f t h e d i p n e t .

D u r in g c o l d , o v e r c a s t w e a t h e r , h o w e v e r , t h e s e g r o u p s w ere

13

n o t o b s e r v e d . Spec im en s w e re c o l l e c t e d o n l y by s a m p l i n g i n

u n d e r c u t s , d e c a y i n g v e g e t a t i o n and b o t to m s u b s t r a t e , w h ic h

y i e l d e d few i n d i v i d u a l s . D u r in g w i n t e r warm s p e l l s swimming

f i s h r e a p p e a r e d , b u t o n l y i n a f r a c t i o n o f t h e i r s p r i n g and

summer a b u n d a n c e .

S p e c i e s A s s o c i a t i o n s

F o u r o t h e r s p e c i e s o f f i s h e s w e re c a p t u r e d i n a r e a s

w h e re F\_ l u c i a e was p r e s e n t . T h ese w e re t h e mummichog

(F u n d u l u s h e t e r o c l i t u s ) , t h e s h e e p s h e a d minnow (C y p r in o d o n

v a r i e g a t u s ) , t h e r a i n w a t e r k i l l i f i s h (L u c a n i a p a r v a ) and t h e

m o s q u i t o f i s h (Gambusia a f f i n i s ) . a f f i n i s and p a r v a

w e r e b o t h r a r e i n l u c i a e h a b i t a t s . The f o r m e r was

c a p t u r e d o n l y a t Fox G re e k M arsh and t h e l a t t e r o n ly a t

Ware R i v e r and Roane M a rsh . h e t e r o c l i t u s and C .

v a r i e g a t u s w e re r e l a t i v e l y common a t a l l l o c a t i o n s . At Fox

C r e e k M arsh C_ _ v a r i e g a t u s was t a k e n i n f r e q u e n t l y , b u t a d u l t

and j u v e n i l e F\_ h e t e r o c l i t u s w e re a b u n d a n t and f r e q u e n t l y

o u tn u m b e re d F\_ l u c i a e i n s a m p l e s .

Male R e p r o d u c t i v e C y c le

Male s e x u a l m a t u r i t y was a t t a i n e d a t a t o t a l l e n g t h o f

2 4 -2 7 mm and was s i g n a l l e d by a c h an g e i n c o l o r a t i o n .

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

p l a i n g r a y i s h - g r e e n f e m a l e s . At m a t u r i t y , 1 0 -14 d a r k

v e r t i c a l b a r s fo rm ed a l o n g t h e s i d e s and a b l a c k o c e l l u s

a p p e a r e d on t h e p o s t e r i o r r a y s o f t h e d o r s a l f i n . F u r t h e r

14

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

a c t i v i t y : t h e v e r t i c a l b a r s and o c e l l u s became i n t e n s i f i e d

an d t h e b e l l y , p e l v i c and a n a l f i n s , l o w e r c a u d a l p e d u n c l e

and c a u d a l f i n became y e l l o w i s h - o r a n g e .

C o n t a c t o r g a n s w e re p r e s e n t on t h r e e a r e a s o f b r e e d i n g

m a l e s : t h e s i d e s , h e a d and m e d i a l f i n s . They r e s e m b l e d

f l e s h y p a p i l l a e w i t h s i n g l e s h o r t t e r m i n a l b r i s t l e s and

w e r e v i s i b l e o n l y u n d e r m a g n i f i c a t i o n . T hose on t h e s i d e s

w e re r e p r e s e n t e d by a s i n g l e p a p i l l a on t h e p o s t e r i o r t i p

o f e a c h s c a l e . On m o s t m a l e s c o n t a c t o r g a n s e x t e n d e d

l o n g i t u d i n a l l y f rom j u s t b e h i n d t h e p e l v i c f i n b a s e t o t h e

a n t e r i o r edge o f t h e c a u d a l p e d u n c l e and d o r s o - v e n t r a l l y

o v e r t h e e n t i r e s i d e , e x c l u d i n g t h e t o p t h r e e and b o t to m

two s c a l e ro w s . Head p a p i l l a e w e re s m a l l e r t h a n t h o s e on

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

o p e r c l e . They seemed r an d o m ly s c a t t e r e d and w e re n o t

a s s o c i a t e d w i t h s c a l e s . P a p i l l a e on t h e d o r s a l and a n a l

f i n s w e re a s l a r g e a s t h o s e on t h e s i d e s and o r i g i n a t e d

from f i n r a y s . They w ere c l u s t e r e d n e a r t h e m a r g i n o f e a c h

f i n , w i t h few o r n o n e n e a r t h e b a s e . I n d i v i d u a l d i f f e r e n c e s

e x i s t e d i n t h e e x a c t num ber and l o c a t i o n o f p a p i l l a e , b u t i n

g e n e r a l t h e y f o l l o w e d t h e d i s t r i b u t i o n d e s c r i b e d ( F i g u r e 2 ) .

C o n t a c t o r g a n s w e re p r e s e n t on 130 o f 131 a d u l t m a le s

c o l l e c t e d from March t h r o u g h A u g u s t and w e r e a b s e n t on 36

m a l e s c o l l e c t e d f rom S e p t e m b e r t h r o u g h F e b r u a r y ,

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

15

F i g u r e 2 D i s t r i b u t i o n o f c o n t a c t o r g a n s on an a d u l t

m a le F u n d u lu s l u c i a e . Each m ark r e p r e s e n t s

one c o n t a c t o r g a n .

16F em ale R e p r o d u c t i v e C y c le

F e m a le s a t t a i n e d s e x u a l m a t u r i t y a t a t o t a l l e n g t h o f

2 8 -3 0 mm. M a t u r i t y was n o t m arked by any c h a n g e i n

c o l o r a t i o n and s e c o n d a r y s e x c h a r a c t e r i s t i c s w e re n o t e v i d e n t

d u r i n g t h e spa w n in g s e a s o n . B o th a d u l t an d j u v e n i l e f e m a le s

w e re g r a y i s h - g r e e n , w i t h a few p o s s e s s i n g n u m e r o u s , f a i n t ,

v e r t i c a l b a r s a l o n g t h e s i d e s .

Spawning s e a s o n was i n d i c a t e d by t h e p r e s e n c e o f m a t u r e

and i n c i p i e n t m a t u r e ova ( > 1 . 6 mm d i a m e t e r ) i n o v a r i e s o f

a d u l t f e m a le s and e x t e n d e d from m i d - A p r i l t o m id - A u g u s t .

D u r in g t h i s p e r i o d o v a r i e s c o n t a i n e d e g g s e x h i b i t i n g a

c o n t i n u o u s g r a d a t i o n o f s i z e ( T a b le 1 ) . Egg s i z e and egg

f r e q u e n c y d i s p l a y e d an i n v e r s e r e l a t i o n s h i p . The number o f

e ggs > 1 .6 mm v a r i e d s p o r a d i c a l l y among s a m p l e s , s u g g e s t i n g

d i s c o n t i n u o u s and p e r h a p s s y n c h r o n o u s sp a w n in g ( T a b le 2 ) .

T h e i r f r e q u e n c y i n a s a m p l e , h o w e v e r , d i d n o t c o r r e s p o n d

w i t h any l u n a r o r t i d a l p h a s e .

M a tu re e g gs ( a b o u t 2 mm d i a m e t e r ) w e re d e m e r s a l ,

s p h e r i c a l t o s l i g h t l y e l l i p t i c a l and w e re c h a r a c t e r i z e d by

a p o l a r a g g r e g a t e o f o i l g l o b u l e s and p a p i l l a r y

o r n a m e n t a t i o n o f t h e c h o r i o n ( F i g u r e 3 ) . A d h e s iv e

g e l a t i n o u s t h r e a d s w e re e v i d e n t on m o s t f r e s h l y s t r i p p e d

o r spawned e g g s . M e a s u re m e n ts o f egg s i z e and o i l g l o b u l e

num ber w ere made on 40 f e r t i l i z e d e g gs ( T a b l e 3 ) .

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19

T a b l e 3 . V a r i a t i o n i n t h e d i a m e t e r and num ber o f o i l

g l o b u l e s p e r eg g o f 40 f e r t i l i z e d F u n d u l u s l u c i a e e g g s

________________ C h a r a c t e r __________S t a t i s t i c V a lu e ________

Mean 1 .9 6

Egg D i a m e te r (mm) Range 1 . 7 6 - 2 . 1 8

SD 0 . 1 0

Mean 2 1 . 8

O i l G l o b u l e s p e r Egg Range 5 -5 8

SD 1 3 .3 5

20

F i g u r e 3 . M a tu r e u n f e r t i l i z e d ovum, 1 .9 2 mm i n d i a m e t e r ,

o f F u n d u l u s l u c i a e sh o w in g t h e c h a r a c t e r i s t i c*

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

o r n a m e n t a t i o n o f t h e c h o r i o n .

21Spaw ning B e h a v i o r

Spaw ning b e h a v i o r was o b s e r v e d on t h r e e o c c a s i o n s :

t w i c e i n a q u a r i a (29 A p r i l 1974 , 20 May 1974) and once a t Fox

C r e e k M arsh (23 A p r i l 1 9 7 4 ) . The b e h a v i o r o b s e r v e d was

s i m i l a r on a l l t h r e e d a t e s and may be d i v i d e d i n t o t h r e e

s e q u e n c e s : a p p r o a c h , p u r s u i t and sp a w n in g a c t .

A p p ro a c h i n v o l v e d t h e movement o f a m a le t o w a rd a s l o w -

swimming o r s t a t i o n a r y f e m a l e . The m a l e ' s a p p r o a c h was

u s u a l l y made i n t h e same p l a n e a s t h e f e m a le and from e i t h e r

s i d e o r b e h i n d , b u t n e v e r h e a d - o n . The f e m a le u s u a l l y

r e s p o n d e d by swimming a s h o r t d i s t a n c e away, b u t i f s h e d i d

n o t , s u b s e q u e n t c o n t a c t w i t h t h e m a l e , c o n s i s t i n g o f a

s l i g h t b r u s h o r g e n t l e b u t t , a lw a y s p ro m p te d h e r t o do s o .

The m a le t h e n e i t h e r i g n o r e d h e r o r r e p e a t e d h i s a c t i o n s ,

w i t h t h e same r e s u l t . A m a le w o u ld o f t e n make s e v e r a l

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

u s u a l l y e l i c i t e d p u r s u i t b e h a v i o r . S h o r t r e t r e a t s by t h e

f e m a l e became l o n g e r and l o n g e r , w i t h t h e m a le r a p i d l y

f o l l o w i n g , u n t i l e v e n t u a l l y b o t h b e g an swimming s i d e by s i d e

f o r i n t e r v a l s o f a b o u t 13 s e c o n d s .

The s i d e by s i d e swimming c h a r a c t e r i s t i c o f t h e

p u r s u i t s e q u e n c e was p u n c t u a t e d by s h o r t p a u s e s o f 5 - 1 0 s e c .

d u r a t i o n . D u r in g t h e s e t h e m a le o f t e n r u b b e d and b u t t e d h i s

h e a d a n d body on t h e h e a d , s i d e s and b e l l y o f t h e f e m a l e .

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

swimming. W h i le swimming, t h e m a le f o l l o w e d t h e f e m a le v e r y

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

22

h i s e y e o p p o s i t e t h e f e m a l e ' s o p e r c u l u m . O c c a s i o n a l l y ,

w h i l e i n t h i s p o s i t i o n , h i s m e d i a l f i n s w e re m o m e n t a r i l y

b e n t t o w a rd and a r o u n d t h e f e m a l e ' s b o d y . T h i s was

p e r f o r m e d f rom e i t h e r t h e r i g h t o r l e f t s i d e .

P u r s u i t b e h a v i o r e v e n t u a l l y c u l m i n a t e d i n t h e sp a w n in g

a c t . The sp a w n in g a c t was o n l y o b s e r v e d o n c e , i n an

a q u a r i u m . A f t e r 60 m i n u t e s o f p u r s u i t b e h a v i o r a m a le

c o r n e r e d a f e m a le b e tw e e n t h e f i l t e r , w h i c h o b s c u r e d t h e

a c t , and t h e g l a s s w a l l o f t h e t a n k . One egg was p r o d u c e d

a n d was s u s p e n d e d b e tw e e n t h e g l a s s w a l l and f i l t e r by

c l e a r , a d h e r e n t t h r e a d s . A f t e r t h i s t h e m a le c o n t i n u e d t o

a p p r o a c h t h e same f e m a l e , b u t h i s a t t e n t i o n was more o f t e n

d i r e c t e d to w a r d o t h e r f e m a l e s i n t h e a q u a r i u m .

Male t e r r i t o r i a l b e h a v i o r was c e n t e r e d a ro u n d f e m a le s

b e i n g c o u r t e d . A g o n i s t i c b e h a v i o r was som e t im es o b s e r v e d

b e tw e e n two m a l e s p u r s u i n g t h e same f e m a l e . I t c o n s i s t e d o f

movement by one m a le t o w a rd t h e o t h e r , o f t e n w i t h an

a t t e m p t e d n i p , and r e s u l t e d i n t h e f l i g h t o f t h e m a le b e i n g

t h r e a t e n e d . M ales w ere p r o m is c u o u s and f r e q u e n t l y s w i t c h e d

t h e i r a t t e n t i o n f rom on e f e m a le t o a n o t h e r . The i n i t i a t i v e

u s u a l l y r e s i d e d w i t h t h e m a l e . A f e m a le once a p p r o a c h e d

a n d b u t t e d a m a l e , b u t e l i c i t e d no r e s p o n s e . D u r in g

sp a w n in g a c t i v i t y t h e s i d e b a r s , o c e l l u s and y e l l o w i s h -

o r a n g e u n d e r p a r t s o f t h e m a le w e re v i v i d , b u t t h e d ra b

g r a y i s h - g r e e n o f t h e f e m a le r e m a in e d u n c h a n g e d .

23

Spawning C o n d i t i o n s and S i t e s

Spawning o c c u r r e d i n a q u a r i a c o n t a i n i n g w a t e r o f 0 . 1 ,

3 . 9 and 16.9^<? a t t e m p e r a t u r e s o f 2 0 - 2 5 °C and e x p o s e d t o

n a t u r a l p h o t o p e r i o d s o f 1 3 - 1 / 2 t o 1 5 - 1 / 4 h o u r s . Eggs w e re

d e p o s i t e d on sp a w n in g m ops , w h e re t h e y f i r m l y a d h e r e d t o

s t r a n d s o f y a m . Eggs o c c u r r e d t h r o u g h o u t e a c h mop,

i n d i v i d u a l l y and i n c l u s t e r s o f two t o f o u r . Most w ere

s o l i t a r y and l o c a t e d a t o r n e a r t h e b a s e o f t h e mop, w h e re

t h e s t r a n d s w e re d e n s e s t . Eggs w ere n o t fo u n d e l s e w h e r e i n

t h e t a n k s and t h e i n s t a n c e o f one b e i n g spawned b e tw e e n t h e

f i l t e r and t a n k w a l l was e x c e p t i o n a l .

D u r i n g t h e sp a w n in g s e a s o n a t Fox C r e e k Marsh (m id -

A p r i l t o m id - A u g u s t ) w a t e r t e m p e r a t u r e s o f 1 9 -3 6 ° C ,

s a l i n i t i e s o f 0 . 2 - 9 . ! ^ ? , d i s s o l v e d oxyg en s o f < 1 - 6 . 8 mg/1

an d p h o t o p e r i o d s o f 1 3 - 1 / 2 - 1 5 - 1 / 3 h o u r s w e re r e c o r d e d .

S paw ning mops w ere p l a c e d i n a r e a s o c c u p i e d by F u n d u lu s

l u c i a e , b u t no e g gs w e re f o u n d on them. Eggs w e re n o t fo u n d

on n a t u r a l s u b s t r a t e s .

D ev e lo p m en t and Growth

The l e n g t h o f t h e e m b ry o n ic p e r i o d o f F u n d u l u s l u c i a e

seemed t o be i n f l u e n c e d by s a l i n i t y . Eggs k e p t i n f r e s h

( 0 . 1 L%o) and b r a c k i s h (3 .9 ;^) w a t e r a t 2 0 - 2 5 °C u s u a l l y

h a t c h e d i n 1 2 -1 6 d a y s , b u t some, i n c l u d i n g m o s t e g gs i n

16.9^?, h a t c h e d much l a t e r . One egg i n 16 . 9 ^ h a t c h e d o n ly

a f t e r 43 d ays an d 13 a f t e r 33 d a y s . T h e s e l a t e n t embryos

w e r e m ore d e v e l o p e d a t t h e t i m e o f e m e rg en ce t h a n t h o s e

24

e m e r g in g a f t e r a 12 -16 day e m b ry o n ic p e r i o d . The l a t t e r

w e r e 5 . 3 - 6 . 0 mm i n t o t a l l e n g t h , p o s s e s s e d l a r g e y o l k s a c s

and r a y l e s s v e n t r a l f i n f o l d s . In c o n t r a s t , l a t e n t embryos

w e r e l a r g e r ( 6 . 0 - 6 . 3 mm i n t o t a l l e n g t h ) and p o s s e s s e d v e r y

r e d u c e d y o l k s a c s and v e n t r a l f i n f o l d s w i t h p a r t i a l l y

fo rm ed a n a l r a y s .

A y o l k - s a c l a r v a w h ic h h a t c h e d a t 14 d ay s was p r e s e r v e d

a n d i s d e p i c t e d i n F i g u r e 4A. The c a u d a l f i n c o n t a i n e d 12

r a y s and was c o n f l u e n t w i t h t h e u n p i g m e n t e d , r a y l e s s f i n f o l d s .

The d o r s a l f i n f o l d t e r m i n a t e d b e tw e e n t h e 1 7 t h and 1 8 th

n e u r a l s p i n e s ( c o u n t i n g f o r w a r d f rom t h e c a u d a l f i n ) and t h e

v e n t r a l f i n f o l d e x t e n d e d t o t h e p o s t e r i o r edge o f t h e y o l k

s a c . The y o l k s a c was r e l a t i v e l y l a r g e , w i t h a w e l l -

d e v e l o p e d n e t w o r k o f v i t e l l i n e v e s s e l s , and o i l g l o b u l e s

w e r e d i s p e r s e d a b o u t i t s s u r f a c e . The p e c t o r a l f i n s

c o n t a i n e d p a r t i a l l y fo rm ed r a y s . The h e a d p o s s e s s e d a

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

s n o u t ( 1 6 . 5 i n h e a d ) , w e l l - f o r m e d e y e s an d f i v e

b r a n c h i o s t e g a l r a y s . A p a i r o f o t o l i t h s , t h e l a r g e r a b o u t

t h r e e t i m e s t h e s i z e o f t h e s m a l l e r , w e r e l o c a t e d s l i g h t l y

p o s t e r i o r t o e a c h e y e . The o t o l i t h s w e re n o t s t a t i o n a r y and

c h a n g e d p o s i t i o n when t h e sp e c im e n was m a n i p u l a t e d . The

v e r t e b r a l column c o n s i s t e d o f 31 v e r t e b r a e . Numerous s m a l l

m e l a n o p h o r e s w e re p r e s e n t on t h e d o r s a l s u r f a c e o f t h e h e a d

a n d b a c k , a l o n g t h e f i n f o l d b a s e s and o v e r t h e y o l k s a c ,

e s p e c i a l l y t h e d o r s a l p o r t i o n . A few w e re a l s o p r e s e n t on

t h e l a t e r a l a s p e c t s o f t h e h e a d .

25

The f i r s t m o r p h o l o g i c a l c h a n g e s w e re n o t e d i n l a r v a e o f

6 - 7 mm TL (TL = t o t a l l e n g t h ) ( F i g u r e 4B) . S p ec im en s o f t h i s

s i z e n o l o n g e r p o s s e s s e d a y o l k s a c and a S - s h a p e d

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

t h e v i s c e r a l c a v i t y . The m o u th h a d become p r o t r a c t i l e and

s l i g h t l y u p t u r n e d , w i t h t h e l o w e r j aw p r o t r u d i n g s l i g h t l y i n

f r o n t o f t h e u p p e r . V i l l i f o r m t e e t h w e re p r e s e n t a l o n g t h e

f o r w a r d edg e o f e a c h j a w . F i n r a y d e v e l o p m e n t h a d commenced

i n t h e v e n t r a l and d o r s a l f i n f o l d s , w i t h t h e a p p e a r a n c e o f

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

r a y s h a d i n c r e a s e d t o 1 6 - 2 0 , b u t t h e p e c t o r a l f i n r a y s were

s t i l l o n l y p a r t i a l l y f o r m e d . A l s o , t h e l a t e r a l l i n e was now

d i s c e r n i b l e .

S u b s e q u e n t d e v e l o p m e n t c o n s i s t e d p r i m a r i l y o f f i n r a y

f o r m a t i o n , s q u a m a t i o n and p i g m e n t a t i o n ( F i g u r e s 4C and 4 D ) .

F u r t h e r c h a n g e s w e re n o t o b s e r v e d i n t h e m ou th o r a l i m e n t a r y

t r a c t , o t h e r t h a n s i z e i n c r e a s e . O t o l i t h s r e m a in e d v i s i b l e

i n l a r v a e a s l a r g e a s 9 mm TL, b u t i n c r e a s e d o p a c i t y o f t h e

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

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

c o u n t s o f s p e c im e n s l a r g e r t h a n 9 mm TL. V e r t e b r a l c o u n t s

o f s m a l l e r l a r v a e r a n g e d b e tw e e n 31 and 33 , t h e a d u l t

c o m p le m e n t . A l l s p e c i m e n s , r e g a r d l e s s o f s i z e , p o s s e s s e d

a t l e a s t f i v e b r a n c h i o s t e g a l r a y s and a few , l a r g e r t h a n 8

mm TL, p o s s e s s e d a s h o r t , i n n e r m o s t s i x t h r a y .

A l l f i n s w e re c o m p l e t e l y fo rm ed on s p e c im e n s l a r g e r

t h a n 13 mm TL. A l t h o u g h c o n s i d e r a b l e i n d i v i d u a l v a r i a t i o n

26

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

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

f i n s . M e d i a l f i n d e v e l o p m e n t i n v o l v e d t h e e l e v a t i o n o f t h e

f i n f o l d s i n a r e a s o f r a y f o r m a t i o n and t h e i r g r a d u a l d e s c e n t

e l s e w h e r e . F i n f o l d v e s t i g e s w e re p r e s e n t on l a r v a e o f 11 -1 2

mm TL, b u t w e re a b s e n t f rom l a r g e r i n d i v i d u a l s . The num ber

o f r a y s p r e s e n t i n t h e v a r i o u s f i n s a t d i f f e r e n t t o t a l

l e n g t h s i s l i s t e d i n T a b l e 4 .

The p i g m e n t a t i o n p a t t e r n o f t h e y o l k - s a c l a r v a r e m a in e d

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

s q u a m a t i o n . Some m e l a n o p h o r e s a p p e a r e d a d j a c e n t t o f i n r a y s

an d n e a r t h e f i n f o l d b a s e s and d o r s a l p o r t i o n o f t h e v i s c e r a l

c a v i t y . The b e l l y and s i d e s o f t h e b o d y , h o w e v e r , r e m a in e d

u n p i g m e n t e d .

S q u a m a t io n was f i r s t o b s e r v e d on l a r v a e o f 1 0 -1 1 mm TL.

S c a l e s a r o s e on t h e s i d e s f i r s t , t h e n t h e b a c k , h e a d and

f i n a l l y t h e b e l l y . S p ec im en s e x h i b i t i n g s q u a m a t io n a l s o

d i s p l a y e d m e l a n o p h o r e s i n p r e v i o u s l y u n p ig m e n te d a r e a s .

S q u a m a t io n and p i g m e n t a t i o n w e re c o m p l e t e on i n d i v i d u a l s o f

1 1 -1 3 mm TL, e x c e p t f o r t h e b e l l y , w h ic h was c o m p l e t e on

s p e c im e n s o f 17 mm TL and l a r g e r .

Changes i n body p r o p o r t i o n s d u r i n g d e v e l o p m e n t w ere

g r a d u a l and s u b t l e . They a r e sum m ar ized i n T a b l e 4 .

A t Fox C r e e k M arsh , l a r v a e o f F u n d u l u s l u c i a e and

F u n d u l u s h e t e r o c l i t u s o c c u r r e d t o g e t h e r and w ere v e r y

s i m i l a r i n a p p e a r a n c e . The s m a l l e s t s p e c im e n s c o u l d be

d i s t i n g u i s h e d , h o w e v e r , by t h e d o r s a l p i g m e n t a t i o n p a t t e r n

27

o f t h e h e a d and b a c k ( F i g u r e 5 ) . On FL_ l u c i a e t h i s p a t t e r n

c o n s i s t e d o f many s m a l l , d a r k , e v e n l y d i s t r i b u t e d

m e l a n o p h o r e s . The m e l a n o p h o r e s on F ^ h e t e r o c l i t u s , by

c o m p a r i s o n , a p p e a r e d l e s s n u m e ro u s , l a r g e r and b l o t c h y ,

e s p e c i a l l y o v e r t h e h e a d . I n a d d i t i o n , sp e c im e n s o f F .

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

s p o t a t t h e b a s e o f t h e f i r s t d o r s a l r a y . T h i s s p o t , a

d e n s e a g g r e g a t i o n o f m e l a n o p h o r e s , was n o t p r e s e n t i n F .

l u c i a e . The m i d - d o r s a l s t r i p e o f F ^ l u c i a e , a f i e l d mark

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

( R i c h a r d s and B a i l e y 1 9 6 7 ) , was n o t p r e s e n t i n sp e c im e n s

l e s s t h a n 11 mm TL.

The maximum s i z e a t t a i n e d by F u n d u lu s l u c i a e i s

a p p r o x i m a t e l y 40 mm s t a n d a r d l e n g t h , w h ic h makes i t t h e

s m a l l e s t s p e c i e s i n t h e gen u s F u n d u lu s (Rosen 1 9 7 3 ) . I n t h e

p r e s e n t s t u d y t h e l a r g e s t m a le and f e m a le c o l l e c t e d w ere

r e s p e c t i v e l y 40 and 47 mm TL. Most m a l e s , h o w e v e r , r a n g e d

b e tw e e n 28 -3 5 mm TL and m o s t f e m a l e s r a n g e d b e tw e e n 3 0 -4 0

mm TL. A s h o r t l i f e sp a n i s i n d i c a t e d by t h e s m a l l s i z e

a t t a i n e d by t h e s p e c i e s . S c a l e s exam ined f o r age

d e t e r m i n a t i o n i n some i n s t a n c e s p o s s e s s e d a s i n g l e

i n d i s t i n c t a n n u l u s . U s u a l l y a n n u l i c o u l d n o t b e d i s c e r n e d .

I t seems u n l i k e l y t h a t i n d i v i d u a l s s u r v i v e much l o n g e r t h a n

o n e y e a r .

28

F i g u r e 4 F u n d u l u s l u c i a e l a r v a e a t d i f f e r e n t s t a g e s o f

d e v e l o p m e n t : A, 5 . 3 mm TL; B, 6 . 2 mm TL;

C, 9 . 5 mm TL; D, 1 1 . 2 mm TL.

Tabl

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A OA Pd • A C

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r—1 r -l X GG o •rH

r—l X ) GG 3 G 04J G V4 Sno O X 04J

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A Pd Pd XX Pm CO ' '—H CJ CQ PC

mvor—l

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31

F i g u r e 5 . D o r s a l p i g m e n t a t i o n p a t t e r n s o f F u n d u lu s

h e t e r o c l i t u s (A) and F\_ l u c i a e (B) l a r v a e .

F . l u c i a e i s c h a r a c t e r i z e d by a homogeneous

p a t t e r n o f s m a l l d e n s e m e l a n o p h o r e s . F .

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

a n d a s p o t n e a r t h e b a s e o f t h e f i r s t d o r s a l r a y .

i ? ' * . ; v . r *

2 A * ? * V *

* *

***• «* . * « * * .V * r **

32D i e t

A n a l y s i s o f d i g e s t i v e t r a c t c o n t e n t s showed F u n d u lu s

l u c i a e i s om n iv o ro u s ( T a b le 5 ) . The p r i n c i p a l i t e m s

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

( m a i n l y m o s q u i t o e s and c h i r o n o m i d l a r v a e ) and c o p e p o d s . A

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

a r t h r o p o d s , w e re a l s o e a t e n .

The s m a l l s a m p le s i z e s o f l a t e au tum n and w i n t e r p r e v e n t

a c c u r a t e r e p r e s e n t a t i o n o f s e a s o n a l d i e t t r e n d s , b u t s e a s o n a l

p e a k s an d d e p r e s s i o n s a r e d i s c e r n i b l e f o r some fo o d i t e m s .

F o r e x a m p le , d e t r i t u s and d i a t o m s w ere p r e s e n t i n m o s t

s p e c im e n s t h r o u g h o u t t h e y e a r , e x c e p t A u g u s t and S e p te m b e r

( a n d November f o r d i a t o m s ) . H o m o p te ra n s e x h i b i t e d t h e

c o n v e r s e o f t h i s p a t t e r n . Such v a r i a t i o n i s p r o b a b l y t h e

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

p r e f e r e n c e s .

F e e d i n g b e h a v i o r was o b s e r v e d i n a q u a r i a , i n a s h a l l o w

o u t d o o r w a d in g p o o l and a t Fox C re e k M arsh . V i s i o n seemed

i m p o r t a n t i n l o c a t i n g f o o d i t e m s . At Fox C r e e k M a rsh ,

s l o w l y swimming f i s h w e re o b s e r v e d t o s t o p s u d d e n l y , s t a r e

i n t e n t l y a t t h e s u b s t r a t e b e f o r e them a n d , a f t e r a few

m om en ts , d a r t f o r w a r d and n i p a t i t . I n d i v i d u a l s c o n f i n e d

i n an o u t d o o r w a d in g p o o l jum ped o u t o f t h e w a t e r t o c a p t u r e

s m a l l h o m o p te r a n s r e s t i n g on t h e s i d e s , j u s t a b o v e t h e

s u r f a c e . H om o p te ran s n o t i m m e d i a t e l y c a p t u r e d w ere

f r e q u e n t l y k n o c k e d o n t o t h e s u r f a c e and s u b s e q u e n t l y

i n g e s t e d . Jum p in g was e l i c i t e d f rom a q u a r i u m - h e l d f i s h by

•rH CD<D • ^ 4-J j- i

X CO •r4 CD '-N CJ4J e CO JQ CP CD

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•H d d i—1 CJd •H CD CD r-4 O•rl d J4 J-Icd •rl cd cd4-J cd ad 4-J CD P-o d CJ d r-l 0o o d •r-l CM C/3

o CDCO d BI—1 CO •H CDcd d o 4 2 x-m 0£d CD d 4-J M2 dd b •r-l i—i <0

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s t i c k i n g p i e c e s o f e a r th w o rm t o t h e a q u a r i u m g l a s s n e a r t h e

s u r f a c e o f t h e w a t e r . F i s h 3 0 -4 0 mm TL s u c c e s s f u l l y r e a c h e d

worm s e g m e n ts a s h i g h a s 26 mm. A q u a r iu m - h e ld f i s h a l s o

consum ed i t e m s f l o a t i n g on t h e s u r f a c e , s i n k i n g t h r o u g h t h e

w a t e r and r e s t i n g on t h e b o t t o m . P a r t i c l e s t o o l a r g e f o r

c o m p l e t e i n g e s t i o n w e re so m e t im es g r a s p e d b e tw e e n t h e j aw s

and s h a k e n v i g o r o u s l y . A f r a g m e n t s m a l l enough f o r

c o n s u m p t i o n was o c c a s i o n a l l y removed by t h i s a c t i o n .

P a r a s i t e s

R e p r e s e n t a t i v e s o f e v e r y m a j o r m e t a z o a n p a r a s i t e g r o u p ,

e x c e p t C e s t o d a , o c c u r r e d on F u n d u lu s l u c i a e ( T a b le 6 ) . Some

p a r a s i t e s , b e c a u s e o f t h e i r m i n u t e s i z e and t h e l a c k o f l i v e

m a t e r i a l , may n o t h a v e b e e n d e t e c t e d . T re m a to d e a b u n d a n c e

i s , a t b e s t , a minimum e s t i m a t e . Monogenea and D ig e n e a

r e s p e c t i v e l y c o n s t i t u t e d t h e m o s t num erous and m o s t c o n s i s t

e n t l y p r e s e n t g r o u p s .

The o r g a n s m o s t f r e q u e n t l y p a r a s i t i z e d w e re t h e g i l l s

(Monogenea and C o p e p o d a ) , a l i m e n t a r y t r a c t (D ig e n e a ) and

l i v e r ( A c a n t h o c e p h a l a and N e m a t o d a ) ( T a b l e 7 ) . E x c e p t f o r

t h e m o n o g e n e t i c t r e m a t o d e s an d c o p e p o d s , m o s t p a r a s i t e s w e re

im m a tu r e , e n c y s t e d f o r m s . P a r a s i t i z e d f i s h e x h i b i t e d no

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

i n d i v i d u a l s c o n t a i n i n g l a r g e s p e c im e n s o f E u s t r o n g y l o i d e s

s p . T h e s e n e m a to d e s w e re f r e q u e n t l y more t h a n 60 mm l o n g

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

w h e r e t h e y w e re c o i l e d i n c y s t s . F i s h c o n t a i n i n g them

p o s s e s s e d d i s t e n d e d b e l l i e s .

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41

H y b r i d i z a t i o n

T h r e e F u n d u lu s l u c i a e e gg s w e re s t r i p p e d and s u c c e s s f u l l y

f e r t i l i z e d by s p e r m a t o z o a f rom F u n d u lu s h e t e r o c l i t u s . The

r e c i p r o c a l c r o s s , i n v o l v i n g 11 h e t e r o c l i t u s e g g s , was

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

c o n t a i n i n g b r a c k i s h w a t e r ( 3 . 9 fw) a t 20 -25°C an d h a t c h e d i n

1 4 -2 0 d a y s . The l a r v a e w e re t r a n s f e r r e d t o l a r g e r c o n t a i n e r s

and s u r v i v e d f o r t h r e e w e e k s , a t w h ic h t i m e t h e e x p e r i m e n t

was t e r m i n a t e d . L a r v a e r e s e m b l e d t h e s p e c i e s o f t h e m a t e r n a l

p a r e n t .

D i s c u s s i o n

F u n d u l u s l u c i a e i s a h a r d y , e u r y h a l i n e s p e c i e s c a p a b l e

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

oxygen d e f i c i e n c i e s a s s o c i a t e d w i t h t h e h i g h i n t e r t i d a l s a l t

m a r s h . I t was c o l l e c t e d a t Fox C r e e k Marsh f rom w a t e r s o f

4 - 3 6 °C. J o r g e n s o n (1969) c o l l e c t e d s p e c im e n s f rom 3 6 .1 ° C .

F . l u c i a e o c c u r r e d i n f r e s h w a t e r a t Fox C r e e k M arsh and was

t a k e n f rom 0. i4o by R i c h a r d s a n d B a i l e y ( 1 9 6 7 ) . I t h a s b e e n

r e p o r t e d by J o r g e n s o n ( 1 9 6 9 ) , R i c h a r d s and B a i l e y (1967) and

G r i f f i t h (1974) f rom s a l i n i t i e s e x c e e d i n g 2 7 ^ and de S y l v a

e t a l . (1962) c o l l e c t e d t h e s p e c i e s f rom 4 1 G r i f f i t h

(1974) d e t e r m i n e d l u c i a e c o u l d t o l e r a t e s a l i n i t i e s a s

h i g h a s 1 0 0 ^ b u t c o u l d n o t e n d u r e a b r u p t t r a n s f e r f rom

s a l i n e t o f r e s h w a t e r . T o l e r a n c e o f f r e s h w a t e r was

a c h i e v e d by g r a d u a l a c c l i m a t i o n , b u t s u b s t a n t i a l m o r t a l i t i e s

s t i l l o c c u r r e d w i t h i n a few m o n th s . A p p a r e n t l y l u c i a e

c a n e n d u r e im p e rm a n e n t f r e s h w a t e r c o n d i t i o n s , a s e x i s t e d a t

t h e Fox C r e e k M arsh c o l l e c t i o n s i t e , b u t n o t a c o n s t a n t

f r e s h w a t e r e n v i r o n m e n t . A t Fox C r e e k M arsh l u c i a e

o c c u r r e d i n w a t e r c o n t a i n i n g ^ . 1 . 0 m g/1 d i s s o l v e d ox ygen .

M o r p h o l o g i c a l and p e r h a p s p h y s i o l o g i c a l a d a p t a t i o n s e n a b l e

F . l u c i a e and o t h e r c y p r i n o d o n t s t o i n h a b i t oxygen d e f i c i e n t

w a t e r s . S r i v a s t a v a and G r i f f i t h (1 9 74 ) m e a s u r e d e r y t h r o c y t e

s i z e s o f 20 F u n d u l u s s p e c i e s an d c o r r e l a t e d s m a l l b l o o d c e l l

42

43

s i z e , a s p o s s e s s e d by F\_ l u c i a e , w i t h e x i s t e n c e i n oxygen

d e f i c i e n t h a b i t a t s . S m a l l c e l l s may r e p r e s e n t an a d a p t a t i o n

t o s u c h e n v i r o n m e n t s b e c a u s e t h e y h a v e g r e a t e r s u r f a c e /

vo lume r a t i o s , and t h u s p e r m i t more e f f i c i e n t ga s e x c h a n g e .

The u p t u r n e d m ou ths and f l a t t e n e d h e a d s o f c y p r i n o d o n t s

r e p r e s e n t m o r p h o l o g i c a l a d a p t a t i o n s w h ic h p e r m i t u t i l i z a t i o n

o f t h e oxygen r i c h l a y e r o f w a t e r a t t h e a i r - w a t e r i n t e r f a c e

d u r i n g c o n d i t i o n s o f oxygen d e f i c i e n c y (Lewis 1 9 7 0 ) .

F . l u c i a e h a s e r r o n e o u s l y b e e n c o n s i d e r e d r a r e , b e c a u s e

i t s h i g h m a r s h h a b i t a t h a s b e e n o v e r l o o k e d . C o l l e c t i o n

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

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

Such s u r v e y s commonly employ s e i n e n e t s t o c o l l e c t s h a l l o w

w a t e r f i s h e s and s t a t i o n s a r e e s t a b l i s h e d o n l y w h e re u s e o f

t h e n e t s i s f e a s i b l e . The d e n s e l y v e g e t a t e d , s o f t - b o t t o m e d

m a r s h p o o l s an d d i t c h e s w h e re F\_ l u c i a e was fo u n d i n

a b u n d a n c e w ould n a t u r a l l y be e x c l u d e d .

A l t h o u g h n o t r a r e , t h e s p o t f i n k i l l i f i s h was fo u n d t o

b e o n l y l o c a l l y a b u n d a n t . The e n v i r o n m e n t a l s t r e s s w h ic h

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

m ore w i d e s p r e a d a n d n um erous t h a n i s a c t u a l l y t h e c a s e .

T h r e e f a c t o r s may c o n t r i b u t e t o i t s r e s t r i c t e d , s p o r a d i c

d i s t r i b u t i o n : s h o r t l i f e s p a n , s m a l l s i z e a n d low f e c u n d i t y .

D i s t i n c t a n n u l i w e re n o t o b s e r v e d on s c a l e s o f F .

l u c i a e , b u t a l i f e sp a n o f o n l y a b o u t one y e a r i s i m p l i e d by

t h e s m a l l maximum s i z e a t t a i n e d by t h e s p e c i e s . F .

h e t e r o c l i t u s and 5\_ d i a p h a n u s p o s s e s s v i r t u a l l y t h e same

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s i z e e g g s and l e n g t h s a t h a t c h i n g a s l u c i a e . A f t e r one

y e a r o f g r o w th t h e i r s i z e a p p r o x i m a t e s t h e maximum s i z e

a t t a i n e d by t h e s p o t f i n k i l l i f i s h ( T a b le 8 ) . I f t h e g ro w th

r a t e o f l u c i a e i s s i m i l a r t o t h e g r o w th r a t e s o f t h e s e

two s p e c i e s , t h e n m o s t s p o t f i n k i l l i f i s h do n o t s u r v i v e much

l o n g e r t h a n one y e a r . R e p r o d u c t i o n w ou ld be l i m i t e d t o one

s e a s o n and p o p u l a t i o n s w o u ld c o n s i s t o f o n l y two y e a r

c l a s s e s .

The s m a l l s i z e a t t a i n e d by l u c i a e may p r e v e n t i t s

c o l o n i z a t i o n o f a d j a c e n t t i d a l c r e e k s and low m a rs h a r e a s .

T h e s e e n v i r o n m e n t s w o u ld e x p o s e t h e s p e c i e s t o s t r o n g d a i l y

t i d a l c u r r e n t s , w h ic h i t may n o t be a b l e t o w i t h s t a n d , and

p r e d a t o r s , s u c h a s l a r g e f i s h an d c r a b s . The h i g h i n t e r t i d a l

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

S m a l l s i z e c e r t a i n l y l i m i t s f e c u n d i t y . M a tu r e F.

l u c i a e ova a r e a b o u t 2 mm i n d i a m e t e r and t h e o v a r y o f an

a v e r a g e s i z e f e m a l e c a n n o t c o n t a i n many. The g r e a t e s t

n um ber fo u n d w i t h i n one f e m a l e was 16. O v a r i a n s i z e

r e s t r i c t i o n s may b e somewhat c o m p e n s a te d i f s e v e r a l

i n d i v i d u a l s p a w n in g s o c c u r d u r i n g a b r e e d i n g s e a s o n . The

sp a w n in g s e a s o n w ou ld n e c e s s a r i l y be p r o t r a c t e d , w h ic h i s

t h e c a s e . F e m a le s c o n t a i n i n g m a t u r e ova w ere c o l l e c t e d a t

Fox C r e e k M arsh f rom m i d - A p r i l t o m id - A u g u s t and H i l d e b r a n d

a n d S c h r o e d e r (1928) r e p o r t e d l u c i a e spawned f rom A p r i l

t o O c t o b e r a t B e a u f o r t , N. C. T h i s c o m p e n s a to r y

r e p r o d u c t i v e s t r a t e g y i s f u r t h e r s u p p o r t e d by t h e c o n t i n u o u s

g r a d a t i o n i n egg s i z e e x h i b i t e d by f e m a le s d u r i n g t h e

46

b r e e d i n g s e a s o n . H i l d e b r a n d (1917) fo u n d s e v e r a l s i z e s o f

e g g s s i m u l t a n e o u s l y p r e s e n t i n o v a r i e s o f C y p r in o d o n

v a r i e g a t u s and d e m o n s t r a t e d t h a t a s i n g l e f e m a le was c a p a b l e

o f sp a w n in g num erous t im e s d u r i n g t h e r e p r o d u c t i v e s e a s o n .

Boyd and Simmonds (1974) i n d u c e d h e t e r o c l i t u s t o spawn

t h r o u g h o u t t h e y e a r by m a i n t a i n i n g a b r o o d s t o c k i n a s p e c i a l

a q u a r i u m s y s t e m . The r e l e a s e o f e g g s o v e r an e x t e n d e d p e r i o d

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

F o s t e r (1967) b e l i e v e s t h i s r e p r o d u c t i v e s t r a t e g y i s

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

F o s t e r (1967) h a s s t a t e d t h a t many s p e c i e s o f k i l l i f i s h

a p p e a r t o spawn a l m o s t d a i l y i n c a p t i v i t y f o r p e r i o d s o f two

m o n th s o r l o n g e r . Such c o n t i n u i t y was n o t d i s p l a y e d by

F u n d u l u s l u c i a e i n t h e n a t u r a l e n v i r o n m e n t . The mean

num ber o f m a t u r e ova p e r f e m a le seemed d e p e n d e n t on

c o l l e c t i o n d a t e ( T a b l e 2) a n d e x h i b i t e d a b im o d a l

d i s t r i b u t i o n o f e i t h e r ^ 1 . 0 o r _>6.0 e g g s p e r f i s h . Such

d e p e n d e n c e and b i m o d a l i t y w o u ld n o t b e e x p e c t e d i f i n d i v i d u a l

f e m a l e s w e re sp aw n in g d a i l y . Schm elz (1964) fo u n d spa w n in g

a c t i v i t y o f F ^ h e t e r o c l i t u s i n c r e a s e d a r o u n d new and f u l l

moons . Spawning o f l u c i a e a p p e a r e d t o be s y n c h r o n o u s ,

b u t was c o r r e l a t e d w i t h n e i t h e r l u n a r n o r t i d a l c y c l e s .

F o s t e r (1967) d e s c r i b e d s e x u a l b e h a v i o r o f m a le

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

h e a d b o b b i n g and o t h e r a c t i v i t i e s . The b e h a v i o r e x h i b i t e d by

F . l u c i a e m a l e s seems c o n s e r v a t i v e an d s i m p l e by c o m p a r i s o n .

Some b e h a v i o r may h a v e b e e n o v e r l o o k e d , h o w e v e r , b e c a u s e o f

i n s u f f i c i e n t o b s e r v a t i o n .

47

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

o c c u r on many s p e c i e s and a r e p r e s e n t on t h o s e p a r t s o f t h e

body a n d f i n s w h ic h c o n s i s t e n t l y come i n t o c o n t a c t w i t h t h e

f e m a l e d u r i n g c o u r t s h i p and sp a w n in g ( F o s t e r 1 9 6 7 ) . The

c o n t a c t o r g a n s o f l u c i a e m a l e s c o n fo rm t o t h i s

d i s t r i b u t i o n . The s i d e s o f t h e h e a d and body c o n t a c t t h e

f e m a l e d u r i n g s i d e by s i d e swimming, a s do t h e d i s t a l

p o r t i o n s o f t h e m e d i a l f i n s when t h e y a r e b e n t t o w a rd and

a r o u n d t h e f e m a l e . The r e p o r t by F o w le r (1916) t h a t c o n t a c t

o r g a n s do n o t o c c u r on l u c i a e i s i n c o r r e c t .

F u n d u l u s l u c i a e sp a w n in g s i t e s w e re n o t d e t e c t e d i n t h e

n a t u r a l e n v i r o n m e n t , b u t t h e d e p o s i t i o n o f e g g s i n sp a w n in g

mops i n a q u a r i a s u g g e s t s a f i b r o u s s u b s t r a t e may be u t i l i z e d .

The a d h e s i v e f i l a m e n t s p o s s e s s e d by t h e eggs p r o b a b l y s e r v e

t o a n c h o r them t o t h e s u b s t r a t e a n d , by t h e a t t a c h m e n t o f

p a r t i c u l a t e d e t r i t u s , c a m o u f l a g e them f rom p o t e n t i a l

p r e d a t o r s .

The s p a w n in g , e m b r y o n ic d e v e lo p m e n t and h a t c h i n g o f

e g g s i n 0 . 1 , 3 . 9 and 1 6 . 9 ^ a t t e s t s t o t h e e u r y h a l i n i t y o f t h e

s p e c i e s . The a b i l i t y o f a d u l t F ^ l u c i a e t o spawn and p r o d u c e

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

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

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

was n o t m o n i t o r e d . The d e l a y e d h a t c h i n g e x h i b i t e d by eggs

i n 1 6 . 9 ^ w a r r a n t s f u r t h e r i n v e s t i g a t i o n . D e la y e d h a t c h i n g

h a s b e e n r e p o r t e d f o r o t h e r c y p r i n o d o n t s . Milkman (1954)

was a b l e t o d e l a y h a t c h i n g o f F\_ h e t e r o c l i t u s e g g s by p l a c i n g

48

r e c e n t l y f e r t i l i z e d ova i n a f l a s k , b u b b l i n g i n oxygen and

t h e n s e a l i n g t h e f l a s k . He d e m o n s t r a t e d t h a t a l t h o u g h

e x t r a oxygen h a d an i n h i b i t o r y e f f e c t on h a t c h i n g , i t a l o n e

was n o t r e s p o n s i b l e f o r t h e i n h i b i t i o n o b s e r v e d . H a r r i n g t o n

(1959) o b s e r v e d d e l a y e d h a t c h i n g i n c o n f l u e n t e s e g g s

e x p o s e d t o d e s i c c a t i n g c o n d i t i o n s . One egg p r o d u c e d a

v i a b l e l a r v a a f t e r 95 d a y s . I s h i d a ( 1 9 4 4 a , 1944b) exam ined

t h e e f f e c t s o f t o n i c i t y and e l e c t r o l y t e s on h a t c h i n g i n

O r y z i a l a t i p e s . He fo u n d h a t c h i n g was a c c e l e r a t e d by

h y p o t o n i c s o l u t i o n s o f KC1 and NaCl a n d h y p e r t o n i c s o l u t i o n s

o f N aC l , b u t was i n h i b i t e d by h y p e r t o n i c s o l u t i o n s o f KC1

and C aCl2 . H a t c h i n g o f l u c i a e eggs seems i n h i b i t e d by

s a l i n i t y , b u t o t h e r f a c t o r s may be r e s p o n s i b l e and c a n n o t be

d i s c o u n t e d .

The l i f e c y c l e o f t h e s p o t f i n k i l l i f i s h i s a p p a r e n t l y

c o m p l e t e d w i t h i n one l o c a l i t y a n d d oes n o t i n v o l v e m i g r a t i o n s .

A t Fox C r e e k M arsh l a r v a e and j u v e n i l e s w e re c a p t u r e d i n t h e

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

t h r o u g h o u t t h e y e a r . R e c e n t o f f s p r i n g f e e d and grow

t h r o u g h o u t t h e summer, f a l l and w i n t e r , w h ic h i s u s u a l l y

m i l d i n V i r g i n i a , and a r e r e a d y t o spawn i n A p r i l . Gamete

m a t u r a t i o n i s p r o b a b l y i n d u c e d by i n c r e a s i n g t e m p e r a t u r e and

day l e n g t h . F o s t e r (1967) o b s e r v e d t h a t t h e p o s t r e p r o d u c t i v e

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

M ale and f e m a le l u c i a e p r o b a b l y do n o t s u r v i v e t o spawn

a s e c o n d y e a r .

49

The v a r i e t y o f i t e m s fo u n d w i t h i n t h e d i g e s t i v e t r a c t

Ej . l u c i a e i n d i c a t e s t h e s p e c i e s i s n o n - s p e c i f i c i n i t s

f o o d h a b i t s . I t i s p r o b a b l y o p p o r t u n i s t i c a s w e l l , f e e d i n g

on t h o s e i t e m s m o s t o f t e n e n c o u n t e r e d . The s e a s o n a l

a b u n d a n c e o f Hom optera i n t h e d i e t o f E\_ l u c i a e p r o b a b l y

r e f l e c t s t h e i r s e a s o n a l a b u n d a n c e i n n a t u r e and i s an

ex a m p le o f o p p o r t u n i s t i c f e e d i n g . H i l d e b r a n d and S c h r o e d e r

(19 2 8 ) r e p o r t e d c r u s t a c e a n s , m o l l u s k s and a n n te l id worms i n

t h e s to m a c h s o f n i n e F_;_ l u c i a e . M o l l u s k s and a n n e l i d s w e re

r a r e i n sp e c im e n s e x am in ed d u r i n g t h e p r e s e n t s t u d y .

D i f f e r e n c e s i n t h e f a u n a a v a i l a b l e f o r c o n s u m p t io n may

a c c o u n t f o r t h i s d i s c r e p a n c y .

The f e e d i n g b e h a v i o r d i s p l a y e d by F ^ l u c i a e was s i m i l a r

t o t h a t d e s c r i b e d f o r o t h e r f u n d u l i n e k i l l i f i s h e s by F o s t e r

( 1 9 6 7 ) . The a b i l i t y o f l u c i a e t o jump and s u c c e s s f u l l y

c a p t u r e o r g a n i s m s abo v e t h e s u r f a c e o f t h e w a t e r

d e m o n s t r a t e s t h e k e e n v i s i o n i t p o s s e s s e s . Jum p in g h a s a l s o

b e e n r e p o r t e d i n F . h e t e r o c l i t u s ( C h i d e s t e r 1 9 1 6 ) .

D i l l o n (1966) h a s l i s t e d p a r a s i t e s r e p o r t e d from

F u n d u l u s s p e c i e s . A l l o f t h e g r o u p s and m o s t o f t h e g e n e r a

ob s e r v e d on l u c i a e a r e i n c l u d e d i n t h e l i s t , t h o u g h none

a r e r e p o r t e d f rom F ^ l u c i a e . The o n ly p a r a s i t e p r e v i o u s l y

r e p o r t e d f rom t h e s p o t f i n k i l l i f i s h i s Oodinium

c y p r i n o d o n t u r n , a d i n o f l a g e l l a t e fo u n d on t h e g i l l s ( L a w le r

1 9 6 8 ) .

Many o f t h e o r g a n i s m s p a r a s i t i z i n g l u c i a e w e re

i m m a tu r e , i n d i c a t i n g t h e s p e c i e s may be an i m p o r t a n t

50

i n t e r m e d i a t e h o s t . E x c e p t f o r t h e M onogenea , t h e i n c i d e n c e

o f p a r a s i t e s was n o t p a r t i c u l a r l y h i g h . T h e i r p r e s e n c e i s

p r o b a b l y d e t r i m e n t a l , e s p e c i a l l y when l a r g e r s p e c i e s s u c h a s

E u s t r o n g y l o i d e s o c c u r . S u f f i c i e n t num bers o f p a r a s i t e s may

g r e a t l y c o n t r i b u t e t o o r c a u s e m o r t a l i t y .

H y b r i d i z a t i o n b e tw e e n c y p r i n o d o n t s p e c i e s h a s b e en

f o u n d t o o c c u r b o t h i n n a t u r e (Hubbs , W a lk e r and J o h n s o n

1943) and i n t h e l a b o r a t o r y (Hubbs and Drewry 1959 , 1 9 6 2 ) .

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

c o m p a t i b i l i t y and i n d i c a t e s a c l o s e r e l a t i o n s h i p b e tw e e n t h e

p a r e n t a l s p e c i e s . The s u c c e s s f u l r e c i p r o c a l c r o s s e s o f F .

l u c i a e and F\_ h e t e r o c l i t u s i n t h e l a b o r a t o r y t h e r e f o r e

i m p l i e s a c l o s e r e l a t i o n s h i p b e tw e e n them . K a r y o l o g i c a l

c o n f i r m a t i o n was n o t o b t a i n e d , h o w e v e r , so p a r t h e n o g e n e t i c

d e v e lo p m e n t c a n n o t be d i s c o u n t e d . The s u c c e s s f u l c r o s s i n g

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

may o c c u r and s u g g e s t s t h e i m p o r t a n c e o f r e p r o d u c t i v e

b e h a v i o r i n i t s p r e v e n t i o n .

LITERATURE CITED

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Boyd, J . F . and R. C. Simmonds. 1974 . C o n t in u o u sl a b o r a t o r y p r o d u c t i o n o f f e r t i l e F u n d u lu s h e t e r o c l i t u s (Walbaum) e g g s l a c k i n g c h o r i o n i c f i b r i l s . J . F i s h B i o l . 6: 3 8 9 - 3 9 4 .

Brown, J . L. 1954 . A r e v i e w o f t h e c y p r i n o d o n t genusF u n d u l u s o f t h e e a s t e r n U n i t e d S t a t e s . M.S. T h e s i s . C o r n e l l U n i v e r s i t y .

B u t n e r , A. and B. H. B r a t t s t r o m . 1960. L o c a l movement i n M e n id ia and F u n d u l u s . C o p e ia 1 9 6 0 ( 2 ) : 1 3 9 - 1 4 1 .

Chen , T. R. 1971 . A c o m p a r a t i v e chromosome s t u d y o f t w e n ty k i l l i f i s h s p e c i e s o f t h e g e n us F u n d u lu s ( T e l e o s t e i : C y p r i n o d o n t i d a e ) . Chromosoma 3 2 ( 4 ) :4 3 6 - 4 5 3 .

C h i d e s t e r , F . E. 1916 . A b i o l o g i c a l s t u d y o f t h e more i m p o r t a n t o f t h e f i s h e n e m ie s o f t h e s a l t m a r s h m o s q u i t o e s . B u l l . 300 N. J . A gr . Exp. S t a .P a g e s 1 - 1 6 .

C r a w f o r d , D. R. 1920 . N o t e s on F u n d u lu s l u c i a e .A q u a t i c L i f e 5: 7 5 - 7 6 .

C r e s s e y , R. F . 1972. The g e n u s A r g u l u s ( C r u s t a c e a :B r a n c h i u r a ) o f t h e U n i t e d S t a t e s . U. S. E n v i r o n . P r o t e c t . Agency W a te r P o l l u t . C o n t . R e s . S e r .18050 ELD02/72. U. S. Governm ent P r i n t i n g O f f i c e , W a s h i n g t o n , D. C. 13 pp .

de S y l v a , D. P . , F . A. K a l b e r , J r . and C. N. S h u s t e r , J r . 1 962 . F i s h e s and e c o l o g i c a l c o n d i t i o n s i n t h e s h o r e z o n e o f t h e D e law a re R i v e r e s t u a r y , w i t h n o t e s on o t h e r s p e c i e s c o l l e c t e d i n d e e p e r w a t e r . U n iv . D e l . Mar. Lab . I n f o r m . S e r . P u b l i c a t i o n 5 . 164 pp .

D i l l o n , W. A. 1966 . P r o v i s i o n a l l i s t o f p a r a s i t e so c c u r r i n g on F u n d u lu s s p p . Va. J . S c i . 17 ( 1 ) : 2 1 - 3 1 .

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F e r n a n d o , C. H . , J . I . F u r t a d o , A. V. G u s s e v , G. Hanek and S. A. K a k o n g e . 1972. Methods f o r t h e s t u d y o ff r e s h w a t e r f i s h p a r a s i t e s . U n i v e r s i t y o f W a t e r l o o . B i o l . S e r . No. 12. 76 pp .

F o s t e r , N. R. 1967 . C o m p a r a t i v e s t u d i e s on t h e b i o l o g yo f k i l l i f i s h e s ( P i s c e s : C y p r i n o d o n t i d a e ) . Ph .D .T h e s i s . C o r n e l l U n i v e r s i t y . 369 p p .

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56

VITA

D onald M i c h a e l Byrne

B o m i n T e a n e c k , New J e r s e y , O c t o b e r 26 , 1949.

G r a d u a t e d f rom New M i l f o r d H igh S c h o o l i n New M i l f o r d ,

New J e r s e y , J u n e 1967 , B . A . , U n i v e r s i t y o f V i r g i n i a ,

1971 . Employed by V i r g i n i a I n s t i t u t e o f M a r in e S c i e n c e

a s l a b o r a t o r y t e c h n i c i a n , D e p a r t m e n t o f P h y s i c a l O c e a n o g ra p h y ,

S e p te m b e r 1971 - F e b r u a r y 1972 , and l a b o r a t o r y s p e c i a l i s t ,

D e p a r t m e n t o f M i c r o b i o l o g y - P a t h o l o g y , March 1972 -

F e b r u a r y 1974 .

In S e p te m b e r 1972 , t h e a u t h o r e n t e r e d t h e C o l l e g e

o f W i l l i a m and Mary, D e p a r tm e n t o f M ar in e S c i e n c e and

i n May 1974 , became a g r a d u a t e a s s i s t a n t i n t h e D e p a r tm e n t

o f I c h t h y o l o g y .

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