P. Fossum, H. Bj~rke og R. Sætre Fiskeridirektoratets Havforskningsinstitutt
Postboks 1870, 5024 Bergen
SAMMENDRAG
Denne rapporten gir noen forelopige resultater fra sildelarveunder- s~kelsene på Møre i april 1986. Den behandler horisontal- og verti- kalfordeling av larvene i relasjon til det fysiske miljoet samt deres vekst og næringsopptak.
Gytefeltene kommer klart frem i fordelingen av de minste larvene.
Feltene var de samme som ble funnet i 1985. I tillegg fant det sted en mindre gyting ute
påeggakanten. Hovedtrekkene i sirkulasjons- monsteret synes
åvære det samme som i 1985. Ved
åkombinere hydro- grafiske data, larvefordeling og drivboyer ser det ut for at man kan få frem både larvenes viktigste driftsruter samt områder hvor larvene holdes tilbake i 10-15 dager.
Larvepopulasjonen i 1986 var totalt dominert av plommesekklarver.
Antall larver større enn 12
mmvar omkring 10% av det som ble funnet
isamme periode i 1985.
11986 ble det også observert lavere vekst og næringsopptak hos larvene enn i 1985. Alt dette tyder på at rekrutteringen til sildebestanden i 1986 var dårligere enn i 1985.
Storstedelen av larvene ble funnet dypere enn
60 m.Innslaget av ny-
klekkede larver Øket med dypet mens eldre larver ble hovedsaklig
funnet i de Øvre 40
m.Forste næringsopptak ble funnet i 3-6 dagers
gamle larver. Dietten var dominert av rauåte larver.
STUDIES ON HERRING M V A E OFF CdESTERN FIORWAY E61e
1986
by
P. Fossum, H . Bjarke and R . Sætre I n s t i t u t e of Marine Research P. O . Box 1870, N-5024 Bergen
Norway
ABSTRACT
T h i s r e p o r t g i v e s some p r e l i m i n a r y r e s u l t s from a s t u d y on t h e spawning grounds of t h e Norwegian spring-spawning h e r r i n g i n A p r i l 1986. I t d e a l s with t h e h o r i z o n t a l and v e r t i c a l d i s t r i b u t i o n of h e r r i n g l a r v a e i n r e l a t i o n t o t h e p h y s i c a l environment a s w e l l a s growth and d i e t .
The spawning a r e a can c l e a r l y be seen i n t h e d i s t r i b u t i o n of l a r v a e
<
9 mm and t h e s e were t h e same a s i n 1985. I n a d d i t i o n , a s m a l l e r spawning a r e a o f f s h o r e a t t h e c o n t i n e n t a l s l o p e was i d e n t i f i e d .The main f e a t u r e s of t h e c i r c u l a t i o n p a t t e r n of t h e a r e a seems t o be t h e same a s i n
1985.
The combined u s e of hydrography, l a r v a e d i s t r i - b u t i o n and Argos d r i f t e r s seem a b l e t o r e v e a l both t h e l a r v a l d r i f t a s w e l l a s r e t e n t i o n a r e a s with a r e s i d e n c e time of 10-15 d a y s ,The l a r v a l population i n 1986 was t o t a l l y dominated by yolksac l a r v a e . The number of l a r v a e
>
12 mm was about 10% of t h a t found d u r i n g t h e same p e r i o d i n 1985, I n 1986 a slower growth and a lower f e e d i n g r a t i o i n a l l l a r v a l s t a g e s was observed compared t o 1985, T h i s i n d i - c a t e poorer recruitment i n 1986 than i n 1985.The m a j o r i t y of t h e l a r v a e were found deeper than 60 m opposed t o 1985. The percentage of newly hatched l a r v a e i n c r e a s e d with depth.
Older l a r v a e were found mainly i n t h e upper 40 m. Patchy d i s t r i b u t i o n made a comparison between l a r v a e caught a t day and a t n i g h t d i f f i c u l t . F i r s t f e e d i n g was observed i n 3-6 days o l d l a r v a e . The d i e t of t h e l a r v a e was dominated by copepods n a u p l i i . A s h r i n k a g e of 40-50% i n dryweight and
3-7%
i n l e n g t h due t o p r e s e r v a t i o n was observed.Sampling of l a r v a e of t h e Norwegian spring-spawning h e r r i n g has been c a r r i e d o u t f o r a long period ( e . g . WLBORG 1960, DRAGESUND 1970, SELI- VERSTOV 1974 and BJØRKE 1 9 8 1 ) . The main o b j e c t i v e of t h e s e i n v e s t i g a - t i o n s has been t o l o c a t e spawning a r e a s and t o o b t a i n t h e f i r s t i n d i - c a t i o n of t h e r e c r u i t m e n t . The i n c r e a s e of t h e h e r r i n g s t o c k i n more r e c e n t y e a r s and development of new g e a r s and methods, a c t u a l i z e t h e formulation of a p r o j e c t t o s t u d y t h e r e c r u i t m e n t mechanisms.
I n
1985
a p i l o t study was c a r r i e d o u t (BJØRKE, FOSSUM and SÆTRE, 1 9 8 6 ) . The p r o j e c t was l a t e r included i n a n a t i o n a l program t o s t u d y t h e p o s s i b l e consequences on f i s h eggs and l a r v a e of o i l explo- r a t i o n on t h e Norwegian c o n t i n e n t a l s h e l f n o r t h of 6 2 ' ~ (FØYN and BJØRKE, 1986). T h i s program i s given t h e acronym HELP (Havforsknings- i n s t i t u t t e t s Egg- og Larveprogram) and i s supposed t o l a s t f o r t h e p e r i o d e 1986-90.The o b j e c t i v e s of HELP i s t o :
-
o b t a i n d e t a i l e d knowlegde of t h e d i s t r i b u t i o n i n s p a c e and time of t h e important commercial f i s h s p e c i e s d u r i n g t h e i r e a r l y l i f e s t a g e along t h e Norwegian c o a s t ,-
s t u d y t h e r e p r o d u c t i v e biology of t h e same s p e c i e s .-
s t u d y t h e r e c r u i t m e n t mechanisms of t h e Norwegian spring-spawning h e r r i n g and t h e Arcto-Norwegian cod.-
s t u d y t h e p h y s i c a l f a c t o r s a f f e c t i n g both t h e t r a n s p o r t - d i s p e r s i o n and t h e l i v i n g c o n d i t i o n of t h e e a r l y l i f e s t a g e s of f i s h .The p r e s e n t r e p o r t g i v e s some p r e l i m i n a r y r e s u l t s from t h e i n v e s t i - g a t i o n on t h e l a r v a e of t h e Norwegian spring-spawning h e r r i n g i n 1986.
MATERIALS AND METRODS
The s t u d y was c a r r i e d o u t during t h e p e r i o d s 29 March
- 7
A p r i l and9 - 18
A p r i l . The southern p a r t of t h e a r e a was covered d u r i n g both t h e s e p e r i o d s ( F i g . l ) . Herring l a r v a e were sampled with a modified c o n i c a l n e t of 0.5 m* opening and375
p m mesh s i z e (ELLERTSEN e t a l .1984)
from 150 m ( o r5
m above t h e bottom) t o t h e s u r f a c e . The v e r t i - c a 1 d i s t r i b u t i o n s of temperature, s a l i n i t y , n u t r i e n t s and c h l o r o p h y l l c o n t e n t s were observed by Nansen h a u l s . Five Argos s a t e l l i t t e - t r a c k e d , d r i f t i n g buoys were deployd. These were equipped with a 10 m 2 window b l i n d drogue a t t a c h e d t o t h e buoys v i a a 30 m t e t h e r l i n e .The m a t e r i a l f o r t h e v e r t i c a l s t u d i e s was d e r i v e d from two experiments on Buagrunnen; one from 5-7 A p r i l with a Mocness l m 2 sampler(mesh 3 7 5 ~ ) (WIEBE e t a l . ) ,and one from A p r i l
5
and 6 with a opening/closing Juday n e t (mesh 375p and opening O.5m2
).
Dif f i c u l t i e s i n changing of n e t s a t p r e c i s e depths arosed d u r i n g t h e Mocness experiment and t h i s caused a s l i g h t overlapping of depth i n t e r v a l s (Table 1.) About 50 m 3 was f i l t e r e d with t h e Mocness sampler w i t h i n each depth i n t e r v a l and about 1 0 m 3 with t h e v e r t i c a l n e t . To confirm abundance of l a r v a e a v e r t i c a l plankton hau1 was made between each Mocness h a u l . Bottom depth v a r i e d between 120 and 90 m d u r i n g t h e Mocness experiment.During t h e sampling with t h e Juday n e t t h e s h i p was anchored a t a depth of 120 m .
The h e r r i n g l a r v a e used t o morphometric measurements and g u t c o n t e n t a n a l y s i s were preserved i n
4%
formalin i n 10%.
s e a water.
Only t h e food organisms t h a t could be recognized through t h e e p i t h e l i u m of t h e g u t were examined, because of g u t c o n t e n t voidance i n h e r r i n g l a r v a e d u r i n g c a t c h i n g and f i x a t i o n (HAU1981).
No q u a n t i t a t i v e a n a l y s e s of t h e d i e t of t h e h e r r i n g l a r v a e was performed due t o t h e same reason.The same procedure was followed i n 1986 a s w i t h t h e 1985 m a t e r i a l (BJØRKE, FOSSUM & SAXRE 1986) and a comparative a n a l y s i s between t h e s e two sets of d a t a could t h u s be made.
The h e r r i n g l a r v a e used t o d e s c r i b e t h e h o r i z o n t a l d i s t r i b u t i o n were measured on board. Standard l e n g t h (SL) measurements of 50 h e r r i n g l a r v a e p e r hau1 ( i f p r e s e n t ) were taken t o n e a r e s t mm below. For morp- hometric measurements i n t h e l a b o r a t o r y a m a t e r i a l of 647 h e r r i n g
l a r v a e were analysed
.
From each s t a t i o n 20 l a r v a e ( i E p r e s e n t ) were c l a s s i f i e d according t o DOYLE (1977) and ØIESTAD (1983). and measured t o n e a r e s t 0 . 1 mm below. V i s i b l e prey organisms were d i s s e c t e d o u t of t h e g u t and c l a s s i f i e d i n t o one of t h e following two groups; copepod eggs o r copepod n a u p l i i . The l a r v a e were then r i n s e d i n f r e s h w a t e r , d r i e d t o c o n s t a n t weight and weighed on a Cahn e l e c t r o b a l a n c e t o t h e n e a r e s t pg.Because of shrinkage due t o p r e s e r v a t i o n (THEILACKER & DORSEY 1 9 8 0 ) , a r e p r e s e n t a t i v sample of h e r r i n g l a r v a e ( N = 47) from t h e whole a r e a of d i s t r i b u t i o n were s e p e r a t e d i n t o d i f f e r e n t s t a g e s when f r e s h onboard.
They were measured t o n e a r e s t O . l m m below, r i n s e d i n f r e s h water and brought t o t h e l a b o r a t o r y where they were d r i e d t o c o n s t a n t weight and weighed on a Cahn e l e c t r o b a l a n c e t o t h e n e a r e s t pg. This sample r e p r e - s e n t an e s t i m a t e of t h e t r u e s t a n d a r d l e n g t h and d r y weight i n t h e l a r v a l p o p u l a t i o n , and a comparison between t h e s e l a r v a e and t h e pre- served ones g i v e s information of t h e shrinkage d u r i n g f i x a t i o n .
RESULTS AND DISCUSSION
Hydrography
Surface temperatures ( F i g . 2 ) and s a l i n i t i e s ( F i g . 3 ) were
4.5
O - 6 . 5 ' ~ and 33-35%. r e s p e c t i v e l y . The n o r t h e r n p a r t of t h e a r e a was c l e a r l y i n f l u e n c e d by A t l a n t i c Water masses while i n t h e s o u t h e r n p a r t C o a s t a l Water was dominant. The same p a t t e r n could be s e e n i n t h e s u r f a c e d i - s t r i b u t i o n of n i t r a t e and s i l i c a t e ( F i g s . 4-5) with high v a l u e s i n t h e n o r t h e r n and low v a l u e s i n t h e s o u t h e r n p a r t . The s u r f a c e d i s t r i b u t i o n of c h l o r o p h y l l g ( F i g .6 )
showed very low v a l u e s i n t h e n o r t h e r n a r e a . The h i g h e s t c o n c e n t r a t i o n s were found along t h e s h e l f break i n t h e c e n t r a l a r e a .Fig.
7
shows t h e t y p i c a l v e r t i c a l hydrographic s t r u c t u r e from t h e s o u t h e r n (SECTION A ) , t h e c e n t r a l (SECTION B ) and t h e n o r t h e r n (SECTION C ) p a r t of t h e i n v e s t i g a t e d a r e a . The l o c a t i o n of t h e s e c t i - ons appear i n Fig. 1. C o a s t a l Water ( S<
3 5 % . ) covers t h e upper 100 m i n t h e s o u t h e r n and c e n t r a l s h e l f a r e a . I n t h e northernmost s e c t i o n(SECTION C ) water of c o a s t a l o r i g i n i s seen a t t h e outermost s t a t i o n s . T h i s i s probably a r e s u l t of t h e s p l i t t i n g of t h e Norwegian C o a s t a l
Current f u r t h e r south a l s o i n d i c a t e d i n t h e d i s t r i b u t i o n of s u s f a c e s a l i n i t y ( F i g . 3 ) . According t o LJØEN and NAli(KEN
(1969)
t h e Norwegianr C o a s t a l Current s p l i t s i n t o two branches between63' and 6 4 ' ~ ~
The main branch run p a r a l l e l and c l o s e t o t h e c o a s t on t h e e o a s t a l s i d e of Haltenbanken (Area F, F i g .1 7 ) .
The o t h e r branch follows t h e edge oft h e c o n t i n e n t a l s l o p e on top of t h e A t l a n t i c water.
SECTION A was c a r r i e d o u t twice;
31
March and16
A p r i l ( F i g , 71, The a n g l e of t h e s l o p i n g i n t e r f a c e between t h e C o a s t a l and t h e A t l a n t i c Water have decreased d u r i n g t h e f i r s t p a r t of A p r i l r e s u l t i n g i n a more seaward e x t e n t i o n of t h e C o a s t a l Water. There i s a pronounced r e - d u c t i o n of t h e n i t r a t e c o n t e n t i n t h e upper l a y e r s from t h e end of March t o mid-April i n d i c a t i n g t h a t a phytoplankton blsomhas
occurred between t h e o b s e r v a t i o n s . During t h e same p e r i o d e t h e c o n t e n t s of c h l o r o p h y l l g i n t h e upper 30 m of t h e s e c t i o n i n c r e a s e d from 0 - 1-
0 . 3 t o
1 . 5 - 4.5
mg/m3 ( F i g .8 ) .
Though t h e s u r f a c e c h l o r o p h y l l v a l u e s9-18
A p r i l i n t h e n o r t h e r n a r e a i s low ( F i g .G),
reminiseences of a precedent phytoplankton bloom a r e seen i n SECTION C ( F i g ,8 ) .
F i g .
9
shows t h e d r i f t i n g t r a c k s from t h e Argos kuoys. The bottom topography of t h e a r e a i s r a t h e r complica.ted and t h i s i n f l u e n c e s t h e c i r c u l a t i o n p a t t e r n . There a r e f o u r major shallow banks i n t h e a r e a : Buagrunnen ( C ) , F r ~ y a b a n k e n (D ) , Haltenbanken ( F ) and Sklinnabanken( E ) . The l o c a t i o n of t h e s e a r e seen i n F i g .
17.
Around t h e s e banks t h e topographic s t e e r i n g of t h e c u r r e n t favour an a n t i - c y c l o n i c e i r c u - l a t i o n . North of Buagrunnen and of F r ~ y a b a n k e n th e water mainly flows eastwards and merge i n t o t h e channel between Haltenbankena d
t h e c o a s t . This seems t o be t h e main r o u t e f o r t h e northward d r i f t i n g l a r v a e . The c u r r e n k speed of t h e upper l a y e r of t h i c a r e a i s between 30 and 60 cm/s which means7-15
n a u t i c a l miles/day, OE t h e two Argos buoys which were d r i f t i n g f u r t h e r northward. one ended up a t66'~ 26
May and t h e o t h e r one c l o s e t o 6 9 ' 3 0 ' ~ a t
18
June ( F i g , L O )H o r i z o n t a l l a r v a e d i s t r i b u t i o n
The h a t c h i n g of h e r r i n g l a r v a e s t a r t e d i n t h e southernmost a r e a around
15
March and a t Buagrunnen (Area C , F i g .17)
around L A p r i l , A t both l o c a t i o n s t h e maximum h a t c h i n g occurred i n mid-April (BJØRKE, HANSEN and MELLE,1987) .
The h o r i z o n t a l d i s t r i b u t i o n of h e r r i n g l a r v a e of t h r e e d i f f e r e n t l e n g t h groups from t h e f i r s t coverage 29 March
- 7
A p r i l i s shown i n F i g s .11-13.
The d i s t r i b u t i o n of t h e youngest l a r v a e ( F i g . l i ) i n d i - c a t e t h r e e s e p a r a t e d spawning a r e a s ; Buagrunnen, c l o s e t o t h e s h o r e i n t h e southernmost a r e a and a t t h e s h e l f break around 6 3 ' ~ . A s can be seen t h e r e a r e very few l a r v a eL
12 mm ( F i g .1 3 ) .
During t h e second coverage (9-18 A p r i l ) t h e t o t a l number of l a r v a e
< 9
mm i s h i g h e r ( F i g .1 4 ) .
Two of t h e apparent spawning a r e a s a r e t h e same a s on t h e previous coverage. A d d i t i o n a l l y , t h e r e seems t o be a minor spawning n e a r FrØyabanken (Area D, Fig. 1 7 ) . The number of l a r v a eL
12 mm ( F i g . 1 6 ) i s only about 10% of t h a t found d u r i n g t h e same p e r i o d i n1985
(BJØRKE, FOSSUM and SÆTRE, 1 9 8 6 ) . T h i s may i n d i - c a t e t h a t t h e recruitment success i n 1986 was c o n s i d e r a b l y lower than i n1985.
Off t h e southernmost c o a s t a l spawning a r e a (Area A , Fig.
17)
a s w e l l a s a t Buagrunnen t h e Argos d r i f t e r s i n d i c a t e r e t e n t i o n a r e a s of t h e l a r v a e with a recidence time of t h e water of 10-15 days. Fig.17
i s an attempt t o summarize t h e information on t h e l a r v a l d r i f t . It i s based on l a r v a e d i s t r i b u t i o n , t h e t r a c k s of t h e Argos d r i f t e r s and on hydrography. The d r i f t i n g time from t h e spawning a r e a s t o t h e p a s s i n g of t h e 6 5 ' ~ l a t t i t u d e seems t o be 40-50 days from t h e southernmost spawning a r e a and 20-30 days from Buagrunnen. A s a s i t g n l f i c a n t pso- p o r t i o n of t h e l a r v a e p o p u l a t i o n i s below t h e drogue depth of t h e Argos buoys and a s d i s p e r s i o n mechanisms i s n o t c o n s i d e r e d , t h e c a l - c u l a t e d d r i f t i n g times is probably underestimates.V e r t i c a l l a r v a e d i s t r i b u t i o n .
Table 1 shows t h e number of l a r v a e p e r m 2 s u r f a c e s a p l e d d u r i n g t h e Mocness experiment. Larvae without yolk s a c and without t h e c h a r a c t e r i s t i c s of s t a g e 2a d e s c r i b e d by DOYLE (1977) a r e o m i t t e d from t h e t a b l e . Although t h e Mocness sampling w i t h i n t h e a r e a was l o c a t e d by abundance of l a r v a e i n v e r t i c a l n e t - h a u l s , some of t h e Mocnes tows were without any l a r v a e . This i n d i c a t e patchy d i s t r i b u t i o n of t h e l a r v a e
.
Table 1. Number of l a r v a e p e r m 2 s u r f a c e s m p l e d d u r i n g t h e Mocness experiment.
Date
5
A p r i l 6 A p r i lHour
18
20 21 23 24 02 04 06 07 09 12 14 15Dep t h
t a b l e continued:
Date 6 A p r i l
7
A p r i lHour 16 17
19
20 21 22 23 00 02 03 04 05 07Dep t h
t a b l e continued:
Date 7 A p r i l
Hour 08
O9
10 11 12 1314
15 16 17 20 T o t a lDepth
F i g .
18
shows t h e v e r t i c a l d i s t r i b u t i o n of l a r v a e when a l l depth i n - t e r v a l s above 80 m were sampled. Nearly 60 % of t h e l a r v a e were found i n t h e 80-56 m depth i n t e r v a l . Most of t h e s e l a r v a e were newly h a t - ched, i . e . s t a g e l a . The percentage of t h e s e l a r v a e i n c r e a s e d with depth and only few were found i n t h e upper 20 m . The percentage of t h e o l d e r s t a g e s , however, i n c r e a s e d i n t h e upper l a y e r s and s t a g e 2a were found only i n t h e s u r f a c e l a y e r .F i g .
19
shows t h e d i s t r i b u t i o n d u r i n g broad d a y l i g h t i . e . between 09 and 15 h r s . GMT. More than55%
of t h e l a r v a e were found i n t h e 80-65 m i n t e r v a l . F i g . 20 shows t h e v e r t i c a l d i s t r i b u t i o n of t h e l a r v a e d u r i n g darkness i . e . between 21 and 02 h r s . GMT. S t i l l most of t h e l a r v a e were found i n t h e 80-85 m i n t e r v a l and s t a g e I b was dominating. The number of l a r v a e caught a t n i g h t were much lesser t h a n t h a t caught a t d a y l i g h t . T h i s i s probably caused by a patchy d i s t r i b u t i o n of l a r v a e(Table 1 ) . T h i s i s most probably a l s o t h e reason f o r t h e absence of l a r v a e i n t h e deepest hau1 a t n i g h t . Because of t h e patchy d i s t r i b u t i - on of t h e l a r v a e i t i s d i f f i c u l t t o make any comparison between s t a g e d i s t r i b u t i o n of l a r v a e caught a t d a y l i g h t and i n d a r k n e s s .
Table 2. Number of l a r v a e p e r m 2 s u r f a c e sampled d u r i n g t h e c l o s i n g n e t experiment.
Date
5
A p r i l 6 A p r i lHour
Og
1113 14 17 19
21 23 01 03 05 07 T o t a lDep t h
Table 2 shows t h e number of l a r v a r p e r m 2 s u r f a c e d u r i n g t h e c l o s i n g n e t experiment. Larvae without yolk s a c and w i t h s u t t h e c h a r a c t e r i s t i c s of s t a g e 2 a d e s c r i b e d by DOYLE (1977) a r e omitted from
t h e t a b l e . F i g . 21 shows t h e v e r t i c a l d i s t r i b u t i o n of t h e l a r v a e
Fn
Table 2. Most of t h e l a r v a e were found i n t h e 100-81 m i n t e r v a l and t h e m a j o r i t y of t h e s e were i n s t a g e l b . During t h i s experiment newly hatched were recorded i n a l l depth i n t e r v a l s without any c l e a r p a t - t e r n . Older l a r v a e , however, were mainly found i n t h e upper l a y e r s .During a s i m i l a r experiment i n 1985 i n t h e same a r e a (BJØRKE, FOSSUM and SÆTRE 1986) more than 65% of t h e l a r v a e were caught i n t h e upper 60 m . I n c l u d i n g t h e l a r v a e omitted from Table l and Table 2 t h e per- centage of l a r v a e i n t h e upper 60 m were 12% d u r i n g t h e Mocness ex- periment and
14%
d u r i n g t h e c l o s i n g n e t experiment i n 1986. I n 1985 t h e h i g h e s t abundance of l a r v a e were found i n t h e middle of t h e pyc- n o c l i n e . I n 1986 t h e pycnocline was less pronounced than i n1985
( F i g . 7 S e c t i o n A ) . I n 1985 few newly hatched l a r v a e ( 1 a r v a e<
9mm)were found. These two f a c t s might e x p l a i n t h e h i g h e r abundance of l a r v a e i n t h e deeper i n t e r v a l s observed i n 1986.Lengthlstage d i s t r i b u t i o n of f i x e d l a r v a e .
Table
3
shows t h e l e n g t h d i s t r i b u t i o n of f i x e d h e r r i n g l a r v a e caught i n 1986 c l a s s i f i e d according t o DOYLE (1977) and ØIESTAD (1983). Fig.24 shows t h e frequency d i s t r i b u t i o n of t h e s e l a r v a e .
93%
of t h e l a r v a e were measured t o t h e mearest mm below, while t h e rest were measured t o t h e n e a r e s t 1/10 mm (Table4 ) .
36%
of t h e l a r v a e< 9
mm i n Table3
were i n s t a g e l a i . e . t h r e e days o l d o r younger, and of l a r v a e
< 8
mm, 46%
were i n s t a g e l a . The spawning grounds f o r h e r r i n g can t h u s be more p r e c i s e l y l o c a t e d by mapping d i s t r i b u t i o n of f i x e d l a r v a e s m a l l e r than8
mm. On t h e o t h e r hand only 16%
of t h e l a r v a e i n Table 3 were s m a l l e r than8
mm while43 %
of t h e l a r v a e were s m a l l e r than9
mm, By mapping d i s t r i b u t i o n of f i x e d l a r v a e s m a l l e r than8
mm one might thus l o o s e information of spawning grounds because of low abundance of such l a r v a e . I n a d d i t i o n , 64%
of t h e l a r v a e i n s t a g e l a a r e omitted from t h e mapping because they a r e e q u a l t o o r l a r g e r than8
mm. F i g . 11 and F i g .1 4
which shows t h e d i s t r i b u t i o n of f r e s h l a r v a e< 9
mm g i v e s t h u s a reasonable good l o c a t i o n of t h e spawning grounds of h e r r i n g when only l e n g t h d a t a of t h e l a r v a e a r e a v a i l i a b l e . Mapping of l a r v a e i n s t a g e l a w i l l , however, g i v e a b e t t e r l o c a t i o n of t h e spawning grounds of Norwegian spring-spawning h e r r i n g .Table
3.
Length/stage d i s t r i b u t i o n of f i x e d h e r r i n g l a r v a e caught i n1986.
Length (mm)
la
l b 1c 1 d 2a TOT...
005
l 1006 167 38 8 70 283
007 515 353 71 271
1210008 749 1049 158 515 2471
009 400 1547 474 617 3038
010
57 761 712 397
219-29
011
24 6
187 25 197
012 1
7 31 9 48
013 4 3 7
014
2 2015
1 1...
TOT
1888 3774 1491 1995 39 9187
Shrinkage of h e r r i n g l a r v a e
I n Table
4
t h e d a t a of shrinkage due t o formalin f i x a t i o n i s given.Table
4.
P e r c e n t shrinkage d u r i n g formalin f i x a t i o nS t a g e Nos of l a r v a e Shrinkage i n Shrinkage i n Fixed Unfixed l e n g t h
(g)
d r y weight%)The t a b l e shows t h a t t h e s h r i n k a g e both i n l e n g t h and weight i s r e h - t i v e l y c o n s t a n t between t h e d i f f e r e n t s t a g e s . The shrinkage i n l e n g t h i s r e l a t i v e l y moderate, while t h e shrinkage i n weight i s very l a r g e and t h e weight i s almost halved d u r i n g t h e f i x a t i o n p e r i o d . The d i f - f e r e n c e s i n mean l e n g t h of t h e same s t a g e s between t h e l a r v a e i n Table
4
and t h e ones r e p r e s e n t e d i n Fig. 24 might be due t o more a c c u r a t e measurements of t h e l a r v a e i n Table4.
Condition of h e r r i n g l a r v a e
The m a t e r i a l i n t h i s examination was sampled d u r i n g t h e second covera- ge and c o n s i s t s of 647 h e r r i n g l a r v a e of s t a n d a r d l e n g t h 6.2-14.5 mm.
The bulk of t h e l a r v a e belonged t o t h e yolk s a c s t a g e s l a - l d . Few l a r v a e had s t a r t e d t o develope t h e d o r s a l f i n , s t a g e 2 a , and t h e deve- lopment of t h e l a r v a l population was delayed compared t o t h e 1985 s e a s o n , when most of t h e l a r v a e sampled i n t h e same area and a t t h e same time were i n t h e 2a s t a g e (BJØRKE
,
FOSSUM & SÆTRE 1 9 8 6 ) .The mean s t a n d a r d l e n g t h , d r y weight and number of l a r v a e i n each s t a g e a r e shown i n Table
5.
Table
5.
The l a r v a l m a t e r i a l sampled i n 1986.Substage Nos. of Mean s t a n d a r d Mean d r y
l a r v a e l e n g t h (mm) SD weight (ligl SD
The mean h a t c h i n g weight cmd t h e mean weight of t h e Larvae I n
the
dif- f e r e n t y o l k s a c s t a g e s was unchanged compared t o t h e 1985 m a t e r i a l(BJØRXE
,
FOSSUM & SÆTRE 1986).
The delayed development of t h e l a r v a l population i n 1986 compared t o 1985, seemes t o be due t o t h e l a t e h a t c h i n g t h i s y e a r . Mastirnum bat- ching was observed i n mid A p r i l
,
(BJØRKE, HANSEM and MELLE 1987) a d l a r v a e hatched e a r l i e r i n t h e season a r e probably vanished.A s t a n d a r d l e n g t h l d r y weight p l o t of t h e p r e s e n t m a t e r i a l i s shown i n F i g . 22. There i s no s t r i c t r e l a t i o n s h i p between t h e s t a n d a r d l e n g t k and t h e dry weight, i n d i c a t e d by a e x p o n e n t i a l c o r r e l a t i o n c o e f f e s c i - e n t r 2 = 0 . 4 0 , i n c o n t r a s t t o t h e E985 m a t e r i a l with a corresponding c o r r e l a t i o n c o e f f i s c i e n t r 2 =0.81.
There seemes t o be s i g n i f i e a n t l e s s growth of t h e h e r r i n g l a r v a e i n i986 than i n 1985, with a d e c r e a s e of t h e growth parameter ( s l o p e ) from 0 . 1 8 t o 0.11. The same p r e s e r v a t i o n procedure was followed both i n 1985 and 1986, and t h i s i n d i c a t e t h a t t h e l a r v a l population i n 1986 was exposed t o more marginale food c o n d i t i o n s than i n t h e previous y e a r .
Some of t h e v a r i a t i o n i n t h e d a t a may be introduced through t h e pre- s e r v a t i o n , a s t h e r e i s a s t r o n g e r l e n g t h l weight r e l a t i o n s h i p i n t h e p l o t of t h e l a r v a e n o t exposed t o Eormalin ( F i g . 2 3 ) . Therefore t h e n e c e s s i t y of a comparable p r e s e r v a t i o n procedure and t o s t u d y t h e l e n g t h
/
weight r e l a t i o n s h i p of unpreserved and presesved l a r v a e muct be s t r e s s e d .F i g . 24 shows t h e l e n g t h frequency d i s t r i b u t i o n of t h e l a r v a e i n t h e d i f f e r e n t s t a g e s . The jump i n s t a n d a r d l e n g t h from s t a g e La t o l b Is probably due t o t h e s e n s i t i v e n e s s of t h e newly hatched l a r v a e t o band- l i n g . They a r e e a s i l y exposed t o shrinkage d u r i n g c a t c h i n g . Exeept f o r t h i s t h e l e n g t h frequency d i s t r i b u t i o n of t h e y o l k s a c l a r v a e seemes t o be comparable t o t h e one from t h e previous y e a r .
The d i e t of t h e l a r v a e d u r i n g S t a g e s l a - 2 a , r e p r e s e n t i n g a t i m e span e s t i m a t e d t o be 23 days (BJØRKE, FOSSUM & SÆTRE 1986) i s shown i n Fig. 25. The dominating food organism through t h i s p e r i o d was copepod
a a u p l i i which contributec!
80 %
t o t h e g t t c c a n t e - a t , Tkie only o t h e r food orgmism found was copepod e g g s (20%j, The youngest l a r v a e found with g u t c o n t e n t were3-6
days o l d . I n t h i s period copepod eggs and copepod n a u p l i i were of e q u a l importmce,
S u t e r on t h e d i e t was dominated o*copepod n a u p l i i . No copepod e g g s were found i n Larvae o l d e r t h m
11
days. There i s a pronounced reduction i n t h e f e e d i n g r a t i o (number of food organisms p e r l a r v a l lut) i n a l l ctagec from 298"o 1986,The reduced f e e d i n g r a t i o and growth parameter t o g e t h e r with t h e t o t a l dominance of yolksac Larvae a s Late a s mid A p r i l , i n d i c a t e
1986
a s a y e a r with poor r e c r u i t m e n t of t h e Norweglan cpring- spawning h e r r i n g , L n v e s t i g a t i o n s c a r r i e d o u t i n May gave a d d i t i o n a l information of t h l s (Nedseaas p e r s , comm,), a d t h e O-group Index e s t a b l i s h e d l a t e i n t h e autumn confirmed t h a t t h i s y e a r c l a c s was a poor one ( R a t t i n g e n p e r s * comm. ) ,CONCEUCIONS
The spawning a r e a can c l e a r l y be seen i n t h e d i s t r i b u t i o n of l a r v a e
<
9
mm and t h e s e were t h e same a s i n2985.
I n a d d i t i o n a s m a l l e r spaw- n i n g a r e a o f f s h o r e a t t h e c o n t i n e n t a l s l o p e was i d e n t i f i e d ,The main f e a t u r e s of t h e c i r c u l a t i o n p a t t e r n of t h e a r e a seems t o be t h e same a s i n
198s.
The combined use of bydrography, l a r v a e d i s t r i b u - t i o n and Argos d r i f t e r s ceem a b l e t o r e v e a l b o t h t h e Larval d r i f t as welI a s r e t e n t i o n a r e a s with a r e s i d e n c e t i m e of10-15
days,The l a r v a l population i n l986 was t o t a l l y dominuted by yolksac l a r v a e . The number of l a r v a e
>
L2 mm woc about 18% of t h a t found d u r i n g t h e same p e r i o d i n1985, In
1986 a slower growth a d a lower f e e d i n g r a t i o i n a l l l a r v a l s t a g e s was observed compared t o 2985, T h i s i n d i c a t e poorer r e c r u i t m e n t i n 1986 t k m i n1985,
The m a j o r i t y of t h e l a r v a e weie found deeper than 60 m a s opposed t o
1985.
The percentage of newly hatched Earvae I.ncreased with depth.Older l a r v a e were found mainly i n t h e upper 40 m , Patchy d i s t r i b u t i o n made a comparison between l a r v a e caugbt a t day and a t n i g h t d i f f i c u l t .
Mapping of t h e e a r l i e s t s t a g e of l a r v a l development i n s t e a d of tlie s m a l l e s t l a r v a e groups g i v e s a b e t l e r Xocation of the spawning
grounds ,
F i r s t f e e d i n g was observed i n 3-6 days o l d l a r v a e . The d i e t of t h e l a r v a e was dominated by copepod n a u p l i i . A shrinkage of 40-45% i n dry- weight and
3-7%
i n l e n g t h due t o p r e s e r v a t i o n was observed.REFERENCES
BJØRKE, H . 1981. D i s t r i b u t i o n of f i s h eggs and l a r v a e from Stad t o Lofoten d u r i n g A p r i l 1976-80. Pp 583-603
& I
Roald Sætre and Martin Mork (Eds.) The Norwegian Coastal Current. U n i v e r s i t y of Bergen1981: 895
p.BJØRKE, H . , P. FOSSUM and R . SÆTRE, 1986. D i s t r i b u t i o n , d r i f t and c o n d i t i o n of h e r r i n g l a r v a e o f f western Norway i n 1985. Coun
.
Meet. i n t . Coun. Explor. Sea, (H:39):
1-15.
BJØRKE, H., K . HANSEN and W. MELLE,
1987.
S i l d e k l e k k i n g og s e i g y t i n g på More i1986.
I n s t i t u t e of Marine ~ e s e a r c h ' s Egg- and Larvae Program (HELP),
Report 1987, No.4.
DOYLE, M . J .
1977.
A morphological s t a g i n g system f o r t h e l a r v a l devel- opment of t h e h e r r i n g , (Clupea harengus L . ) . J . mar. b i o l . A s s . ,51:
859-867.DRAGESUND, 0 . 1970. F a c t o r s i n f l u e n c i n g y e a r - c l a s s s t r e n g t h of Nor- wegian s p r i n g spawning h e r r i n g . FiskDir. Skr. S e r . HavUnders.,l5
:
381-450.
ELLERTSEN, B . , P. FOSSUM, P. SOLEMDAL, S. SUNDBY and S. TILSETH,
1984.
A c a s e s t u d y on t h e d i s t r i b u t i o n of cod l:.rvae and a v a i l a b i l i t y of prey organisms i n r e l a t i o n t o p h y s i c a l p r o c e s s e s i n Lofoten.
I n : E. DAHL, D.S. DANIELSEN, E . MOKSNESS and P . SOLEMDAL ( E d s . ) , The Propagation of Cod Gadus morhua L. F l ~ d e v i g e n r a p p o r t s e r . ,
L:
453 - 477.
FØYN, L , and K . BJØRKE,
1986,
S t r a t e g i e s i n assescment of p o t e n t i a l o i l p o l l u t i o n e f f e c t s on t h e f i s h r e s o u r c e s . Coun. Meet. i n t . Coun. Explor. Sea, (E:34).HAY, D.E.
1981.
E f f e c t s of c a p t u r e and f i x a t i o n on g u t c o n t e n t s and body s i z e of P a s i c i c h e r r i n g l a r v a e . Rapp. P.-v. Reun. Cons.perm. i n t . Explor. Mer,
a:
395-400.LJØEN, R . and 0 . NAKKEN, 1969. On t h e hydrography of t h e s h e l f water o f f Møre and Helgeland. F i s k D i r . Skr. S e r . HavUnders
. , 15:
285-
294.
SELIVERSTOV, A.S.
1974.
V e r t i c a l migrations of l a r v a e of t h e A t 1 a n t ~ - Scandian Herring (Clupea harengus L . ) P. 253-262&
BLAXTER, J . H . S. e d . The e a r l y l i f e h i s t o r y of f i s h . Springer-Verlag. B e r l i n .THEILACKER, G. and DORSEY, K . 1980. Larval f i s h d i v e r s i t y , a summer of l a b o r a t o r y and f i e l d r e s e a r c h . I O C Workshop Report no. 28:
105-142.
WIBORG, K.F. 1960. I n v e s t i g a t i o n s on eggs and l a r v a e of commercial f i s h e s i n Norwegian c o a s t a l and o f f s h o r e waters i n
1957-58.
F i s k . - D i r . Skr. S e r . HavUnders., 12( 7 ) :
1-27.WIEBE, P.H., BURT, K . H . , BOYD, S.H. and MORTON, A.W., 1976. A m u l t i p l e opening/closing n e t and environmental s e n s i n g system f o r sampling zooplankton. J . Mar
.
Res., 34:
313-326.ØIESTAD, V . 1983. Growth and s u r v i v a l of h e r r i n g l a r v a e and f r y (Clupea harengus L . ) exposed t o d i f f e r e n t f e e d i n g regimes i n experimental ecosystems; outdoor b a s i n s and p l a s t i c bags. Ph. d , t h e s i s : l - 2 9 9 . U n i v e r s i t y of Bergen.
F i g . l . Grid o f s t a t i o n s , 29 March - 1 8 A p r i l 1986. B a t h y m e t r i c c o n t o u r s f o r each 100 m a r e i n c l u d e d . I n s e r t e d map shows t h e l o c a t i o n o f t h e s t u d i e d a r e a .
Fig. 3 . S u r f a c e c a l i n i t y 9-18 April 1986.
F i g . 4 . S u r f a c e v a l u e s of n i t r a t e 9-18 A p r i l 1 9 8 6 .
F i g . 5 . S u r f a c e v a l u e s of s i l i c a t e 9-18 A p r i l 1 9 8 6 .
Fig. 6. Surface values of chlorophyll a 9-18 April 1986.
.
l
SECTION A 16 APRIL 1986
14 APRIL 1986
F i g . 7 . Hydrographic s e c t i o n A , B and C . L o c a t i o n of t h e s e i s i n d i c a t e d i n F i g . l .
Ei DEPLOYMENT POSITION A RECOVERY POSITION
F i g . 9. T r a c k s o f t h e d r i f t i n g Argos buoys drogued a t 30 m depth.
Fig. 10. T r a c k s o f two o f t h e Argos buoys f u r t h e r n o r t h .
F i g . 11. D i s t r i b u t i o n o f h e r r i n g larvae < 9 mm ( ~ / m 2 ) , 29 March-7 April 1986.
2 F i g . 1 2 . D i s t r i b u t i o n of h e r r i n g l a r v a e between 9 a n d 11 mm (N/m ) ,
29 March - 7 A p r i l 1 9 8 6 .
F i g . 14- D i s t r i b u t i o n o f h e r r i n g l a r v a e < 9 mm (N/m 2 ) 9-18 A p r i l 1 9 8 6 .
4" 5 O 6" 7 - '8 9 O 10- 11' l z O
I l l
30i 300 200
500 LOG LOC
i : 3 0 8
. . .
6 5'-
300
. . . . . .
6 1 0
.
e . .
HERRING LARVAE -
9 m m 5 (Nlrn2) < 12mm 9 - 18 APRIL 1986
F i g . 1 5 . D i s t r i b u t i o n o f h e r r i n g l a r v a e between 9 and 11 mm ( ~ / m 2 )
,
9-18 A p r i l 1986.
F i g . 1 6 . D i s t r i b u t i o n o f h e r r i n g l a r v a e > 11 mm (N/m 2 ) , 9-18 A p r i l 1986.
F i g . 1 7 . Observed spawning a r e a s , ( c i r c l e s ) and t e n t a t i v e l a r v a l d r i f t r o u t e s . The i n d i c a t e d d r i f t i n g time i s u n t i l p a s s i n g t h e 65 o N l a t i t u d e .
VERTICAL OISTRI3UTION m s s
VERTICAL DISTRIBUTION
m 5 5 FIAGE
r I
IERTiCAL CICTRIEUTION
mNES5
t . . . r I . . n ~ I . 9 5 x 8 . . . . i . r . * l . . - - r
0 1 0 2 0 3 0 w 5 0 m
PER CENT
F i g . 18. V e r t i c a l d i s t r i b u t i o n o f l a r v a e ; number, s t a g e and p e r c e n t a g e f o r t h e Mocness m a t e r i a l .
VERTICAL OISTRIBUTION
W3PeSf DAY VERTICAL DISTRIBUTION
ICEPESS OAY STAGE
VERTICAL OISTRIBUTION
iUSESS DAY
F i g . 19. V e r t i c a l d i s t r i b u t i o n of l a r v a e ; number. s t a g e and percentage f o r the Mocness m a t e r i a l during day time.
YERTIChL DISTRIBUTION
t C M 3 3 N I C F T
VERTICAL DISTRIBUTION
KUES-3 N1W STAGE
VERTICAL DISTRIBUTIOM
FPXXZSS N I W
z I
F i g . 20. V e r t i c a l d i s t r i b u t i o n o f l a r v a e ; number, stage and p e r c e n t a g e f o r t h e Mocness m a t e r i a l d u r i n g n i g h t .
VERTICAL OISTRIBUTION
-ING NET
l
VERTICAL DISTRIBUTION
CLOSINS HET STAGE
VERTICAL DISTRIBUTION
=OS ING NE7 s I
PER CENT
F i g . 2 1 . V e r t i c a l d i s t r i b u t i o n o f l a r v a e ; n u m b e r , s t a g e and.
p e r c e n t a g e f o r t h e c l o s i n g n e t .
STANDARD LENGTH
(M
F i g . 22. The l e n g t h / d r y w e i g h t p l o t o f t h e p r e s e n t h e r r i n g l a r v a e m a t e r i a l .
I
DRY WEIGHT (PLI)
STANDARD LENGTH
(MM)
F i g . 2 3 . The l e n g t h / d r y w e i g h t p l o t o f t h e l a r v a e n o t exposed t o f o r m a l i n f i x a t i o n .
F i g . 2 4 . The p e r c e n t l e n g t h d i s t r i b u t i o n o f t h e d i f f e r e n t s t a g e s .
I
FEEOING RATIO-COP NAUPLII
I
t--*[OF! EGG
0.2
r
I
DAYS POST HATCHINGI
F i g . 25. The d i e t o f t h e l a r v a e i n t h e p e r i o d 3-23 days p o s t h a t c h i n g .