UBC Theses and Dissertations

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UBC Theses and Dissertations

The development and relative growth of the scales of the Pacific herring (Clupea Pallasii) Gwyn, Agnes Margaret 1939

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•Mi , q % a t i> => T H E D E V E L O P M E N T A N D R E L A T I V E G R O W T H O E T H E S C A L E S O E T H E P A C I F I C H E R R I N G ( C L U P E A P A L L A S I I ) b y A g n e s - M . G w y n A T h e s i s S u b m i t t e d 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 i n t h e D e p a r t m e n t o f Z O O L O G Y T h e U n i v e r s i t y o f B r i t i s h C o l u m b i a O C T O B E R , 1 9 3 9 TABLE OF CONTENTS . ACK2TOWLFJDGMENTS ' INTRODUCTION SURVEY OF THE LITERATURE Theory of Age D e t e r m i n a t i o n P r o o f of the Theory Growth C a l c u l a t i o n s THE PROBLEM MATERIAL METHODS THE STRUCTURE, SHAPE, AND S I Z E OF THE SCALES S t r u c t u r e Shape and S i z e T I M E A N D O R D E R O F A P P E A R A N C E O F S C A L E S R E L A T I V E ' ' G R O W T H O F B O D Y A N D S C A L E R e l a t i v e Growth of Body and S c a l e of F i s h i n T h e i r F i r s t Year Growth i n Length of the A n t e r i o r F i e l d (1) i n R e l a t i o n t o the Body Length (L) i n F i s h More than One Year o f Age. SUMMARY AND CONCLUSIONS BIBLIOGRAPHY TABLES Table 1 Table 2 ( a ) Tabl e 2 (b) Table ? Table 4 Table 5 EXPLANATION OF PLATES AND FIGURES PLATES AND FIGURES ACKNOWLEDGMENTS The present i n v e s t i g a t i o n was c a r r i e d out at the U n i v -e r s i t y of B r i t i s h Columbia .and. the P a c i f i c B i o l o g i c a l S t a -t i o n , . Nanaimo. The work was done under the d i r e c t i o n o f Dr. A.L. T e s t e r , P a c i f i c B i o l o g i c a l S t a t i o n , t o whom the author i s deeply i n d e b t e d f o r s u g g e s t i o n s and c r i t i c i s m and w i l l i n g n e s s t o be of a s s i s t a n c e at a l l t i m e s . Dr. T e s t e r a l s o c o m p i l e d .the f i n a l graphs. M a t e r i a l was o b t a i n e d from c o l l e c t i o n s made by Mr. J.L. McHugh of the B i o l o g i c a l S t a t i o n . G r a t e f u l acknowledgment i s made t o Mr. McHugh f o r h i s c o o p e r a t i o n i n t h i s matter and f o r c r i t i c i s m o f the ma n u s c r i p t . • G r a t e f u l acknowledgment' i s made t o Dr. W.A. Clemens, D i r e c t o r of the P a c i f i c B i o l o g i c a l S t a t i o n , who I s respon-s i b l e f o r f i n a n c i a l a s s i s t a n c e through t h e F i s h e r i e s Re- ' search Board, and a l s o f o r making arrangements f o r the work t o be c a r r i e d on a t t h e S t a t i o n . Dr. C. McLean F r a s e r , Head of the Department of Zoology at t h e U n i v e r s i t y of B r i t i s h Columbia, has a s s i s t e d mater-i a l l y by a l l o w i n g the use of h i s l i b r a r y and by h i s w i l l i n g -ness t o gi v e h e l p f u l a d v i c e . Other members of the s t a f f o f . t h e B i o l o g i c a l S t a t i o n and f e l l o w - s t u d e n t s and members of the s t a f f a t the U n i v e r -s i t y of B r i t i s h Columbia have added t o the p l e a s u r e of t h i s work by t h e i r r e a d y c o o p e r a t i o n . THE :DEVELOPMENT AND RELATIVE -GROWTH OF THE SCALES OF THE PACIFIC HERRING (CLUPBA PALLASH) by Agnes M. Gwyn INTRODUCTION The u s e f u l n e s s of the s c a l e s o f f i s h as a means of s t u d y i n g age, growth, and r a c e has come i n t o r e c o g n i t i o n s i n c e the b e g i n n i n g of the c e n t u r y . A l r e a d y t h e r e has ac-cumulated .a v a s t amount of l i t e r a t u r e d e a l i n g w i t h v a r i o u s phases of s c a l e s t u d y f o r many s p e c i e s of f i s h . I t i s now almost u n i v e r s a l l y r e c o g n i z e d t h a t i n many s p e c i e s i t - i s p o s s i b l e t o determine the age of t h e f i s h from the number of growth checks or " a n n u l i " which appear on t h e s c a l e s . I t has f u r t h e r been r e c o g n i z e d t h a t , from a knowledge of t h e r e l a t i o n s h i p betvreen t h e growth of the s c a l e and the growth of t h e body of the f i s h , i t s h o u l d be p o s s i b l e to t r a c e t h e annual growth of the f i s h throughout i t s l i f e span. The present i n v e s t i g a t i o n i s p r i m a r i l y concerned w i t h t r a c i n g the development of t h e growth of the s c a l e i n r e l a -t i o n t o the growth of the body i n t h e P a c i f i c h e r r i n g (Clupea p a l l a s i i ) , a s p e c i e s of f i s h which i s of c o n s i d e r a b l e commercial importance t o the p r o v i n c e of B r i t i s h Columbia - 2 -a n d u p o n w h i c h n o w o r k h a s b e e n p u b l i s h e d i n t h i s r e g a r d u p t o t h e p r e s e n t . B e f o r e c o n s i d e r i n g t h e p r o b l e m i n m o r e d e t a i l i t i s c o n -s i d e r e d a d v i s a b l e t o p r e s e n t a b r i e f r e v i e w o f t h e l i t e r a t u r e d e a l i n g w i t h s c a l e s t u d i e s t o s h o w t h e p r e s e n t s t a t e o f k n o w l e d g e a n d t h e g e n e r a l i z a t i o n s w h i c h m a y b e m a d e f r o m t h e w o r k o f o t h e r s i n t h i s v e r y i m p o r t a n t p h a s e o f f i s h e r i e s r e -s e a r c h . SURVEY OF THE LITERATURE -Theory of Age D e t e r m i n a t i o n The t h e o r y of s c a l e r e a d i n g i s s i m i l a r - t o t h a t of f i n d i n g the age of t r e e s from t h e number of r i n g s . ' Leeu-wehhoeck (1696) noted t h a t the outer s u r f a c e of the s c a l e o f t h e e e l showed a d e f i n i t e p a t t e r n . Zones of w i d e l y separated r i n g s a l t e r n a t e d w i t h zones of c l o s e l y a p p r o x i -mated r i n g s . He i n t e r p r e t e d the patt-ern a s b e i n g th e r e -s u l t of a p e r i o d of r a p i d growth f o l l o w e d by a. p e r i o d of almost complete c e s s a t i o n of growth, and he presumed t h a t each s e t was formed d u r i n g a summer and w i n t e r o f the a n i -mal's l i f e . S c a l e s of t h e t r o u t , salmon, cod, w h i t i n g , and many o t h e r s p e c i e s o f f i s h have been found t o show these same c h a r a c t e r i s t i c s . I n the h e r r i n g , the s c a l e p a t t e r n i s of a d i f f e r e n t n a t u r e . Whereas i n the m a j o r i t y of the above s c a l e s the p a t t e r n c o n s i s t s of more or l e s s c i r c u l a r bands l a i d down i n a l t e r n a t i n g s e r i e s f i r s t - w i d e a p a r t then c l o s e l y approx-imated, i n t h e h e r r i n g s c a l e the bands have become t r a n s -verse and are l a i d down a t almost equal i n t e r v a l s . The s t r i a e do not appear i n zones showing .each year's growth, but marks on t h e s c a l e s which have been i n t e r p r e t e d as " a n n u l i " c o n s i s t of narrow c l e a r zones conforming t o the shape of the s c a l e . The s t r i a e a r e c r o s s these areas. P r o o f of t h e Theory. S u b s t a n t i a t i o n - o f t h e f a c t t h a t the w i n t e r checks or " a n n u l i " are formed each year has been g i v e n by many workers. 'Hoffbauer (1899) worked w i t h c a r p of known age which he kept i n a q u a r i a and was a b l e t o study t h e growth of the s c a l e w i t h growth of the f i s h . W a l t e r (1-901) worked a l s o w i t h c a r p of known age and found r e s u l t s to agree w i t h ' t h e former, but he a l s o found f a c t o r s such as f a l s e checks and m i s p l a c e d s c a l e s which rendered the s c a l e r e a d i n g d o u b t f u l i n some cases. Dahl (1910.) worked w i t h h a t c h e r y t r o u t and salmon of known age, and found complete agreement between the number of a n n u l i on t h e s c a l e s and t h e number of years of l i f e . Eraser. (191-8-) r e p o r t e d s i m i l a r f i n d i n g s f o r f o u r - y e a r -o l d h a t c h e r y r e a r e d .-sockeye salmon (Oncorhynchus n e r k a ) . Winter checks which agreed w i t h t h e known age were found t o be p r e s e n t on a l l f i s h examined at t h a t t i m e . E r a s e r (1921) a l s o r e p o r t e d t h e capture of marked, salmon which showed s i m i l a r agreement. "Wing-e ( 1 9 1 5 ) found t h a t , i n cod which had been marked and l i b e r a t e d and r e c a p t u r e d a f t e r a p e r i o d of s i x months, one y e a r , and two y e a r s , the s c a l e s i n each case showed a n n u l i to correspond t o the known time of f r e e -dom. Van Oosten (I92-3) found agreement between t h e known ages of whit.ef.ish i n t h e i r e i g h t h and n i n t h y e a r s and the number of a n n u l i . I n t h e case of the h e r r i n g , p r o o f of the s c a l e t h e o r y i s a l l by i n d i r e c t evidence, but i s , n e v e r t h e l e s s , c o n c l u -s i v e . The dominance of one year c l a s s over a p e r i o d of " y e a r s has served a s the means of p r o o f . Lea (as r e p o r t e d by Dr. H j o r t f o r t h e Canadian F i s h e r i e s E x p e d i t i o n , 1 9 1 9 ) bad t r a c e d t h e 1 9 0 4 year c l a s s of Norwegian h e r r i n g f o r e l e v e n y e a r s . I n 1 9 0 8 he found t h a t the f o u r - r i n g e d f i s h were p r e -dominant, i n 1 9 0 9 f i v e - r i n g e d f i s h , and so on. Thus i n each year one annulus was added t o t h e s c a l e s of the dominant age-group. Over a.period'of e l e v e n years t h i s dominant age-group p e r s i s t e d . i n t h e f i s h e r y , grew l a r g e r , and added one r i n g t o I t s s c a l e s each.year. An abnormal arrangement of the r i n g s . appeared i n these f i s h - the second and t h i r d r i n g s were more c l o s e l y approximated than i n a normal s c a l e . T h i s p e c u l i a r -i t y s e r v e d t o "mark" the f i s h and i n 1 9 1 0 s i x - r i n g e d f i s h were p a r t i c u l a r l y marked, the f o l l o w i n g year c h i e f l y seven-r i n g e d f i s h showed the f o r m a t i o n . T h i s n a t u r a l marking served v e r y w e l l as proof where e x p e r i m e n t a l .work was l a c k -i n g . Other i n v e s t i g a t o r s have found s i m i l a r p r o o f f o r d i f f e r -ent r a c e s of h e r r i n g . I n a p o p u l a t i o n of h e r r i n g i n a g i v e n l o c a l i t y t h e r e i s found t o be a predominant age-group which o f t e n d i s t i n g u i s h e s t h a t p o p u l a t i o n from those of nearby l o c a l i t i e s . Dahl ( 1 9 0 7 ) was able t o d i s t i n g u i s h between h e r r i n g from f o u r d i f f e r e n t l o c a l i t i e s a c c o r d i n g t o the dom-i n a n t age-group i n each l o c a l i t y . Lea (1910) gave f u r t h e r d e t a i l s on the ra c e s of Norwegian h e r r i n g i n a r e p o r t i n which he compared the samples from the same l o c a l i t y f o r d i f f e r e n t y e a r s , and i n a l l c a s e s i f h e r r i n g w i t h f o u r r i n g s preponderate in"one y e a r the f o l l o w i n g year f i v e - r i n g f i s h are i n t h e m a j o r i t y . 'In Dr. H j o r t * s ' r e p o r t f o r the Canadian F i s h e r i e s E x p e d i t i o n f o r 1914-15, Lea has t a b l e s w h i c h show the d i v i s i o n of t h e A t l a n t i c h e r r i n g i n the v i c i n i t y of the G u l f of S t . Lawrence and Newfoundland i n t o f o u r , s eparate p o p u l a t i o n s . "The d i f f e r e n c e between f i s h from t h e d i f f e r -ent l o c a l i t i e s made i t s e l f apparent p a r t l y i n the more or l e s s d i s t i n c t markings of the annual r i n g s on t h e s c a l e s r e n d e r i n g number and d i s t a n c e more or l e s s easy t o re a d ; p a r t l y i n t h e f a c t t h a t the dominant age-groups (year c l a s s e s ) d i f f e r e d i n samples "from d i f f e rent l o c a l i t i e s ; and f i n a l l y , i n t h e d i f f e r e n t c h a r a c t e r of the growth, as i n d i c a t e d in. the s c a l e s . " T e s t e r (19J7) showed dominant y e a r c l a s s e s i n the P a c i f i c h e r r i n g o f d i f f e r e n t l o c a l i t i e s . The time a t whi c h the w i n t e r r i n g i s formed has been determined f o r s e v e r a l s p e c i e s o f f i s h , and s e r v e s as a d d i -t i o n a l evidence t o s u b s t a n t i a t e t h e w i n t e r r i n g t h e o r y . Dahl (190?) found t h a t i n the h e r r i n g t a k e n i n March t h e r e was a c l e a r zone around the margin of t h e s c a l e , whereas i n the summer the s t r i a e were present on the ve r y margin of the s c a l e . Lea (1910)- confirmed t h i s work of Da h l , and found t h a t the growth of the new summer zone began i n A p r i l w i t h s t r i a e b e i n g formed o u t s i d e t h e c l e a r zone, whereas s c a l e s taken i n November showed a c l e a r zone f o r m i n g at the edge. I n a l a t e r r e p o r t Lea ( 1 9 1 9 ) f o l l o w e d the development of t h e summer band month b y month. As above, s t r i a e began t o f o r m o u t s i d e the c l e a r zone i n A p r i l and as the s c a l e widened d u r i n g the summer t h e s t r i a e c o n t i n u e d t o be approx-imated t o t h e margin u n t i l 'about September. The c l e a r w i n t e r band was formed from October t o March. Hodgson ( 1 9 2 5 ) from an e x t e n s i v e s t u d y o f the E n g l i s h h e r r i n g , has d a t a which shows t h a t "the s c a l e s of the, E n g l i s h h e r r i n g b e g i n t o grow i n A p r i l and cease t o grow i n . September, i r r e s p e c t i v e of t h e ge o g r a p h i c a l p o s i t i o n . " Eor the P a c i f i c h e r r i n g , Thompson ( 1 9 1 7 ) c o n s i d e r e d t h a t the w i n t e r checks of t h e h e r r i n g s c a l e were formed I n t h e l a t t e r p a r t of the w i n t e r , perhaps i n con-n e c t i o n w i t h spawning t i m e . T e s t e r ( 1 9 3 7 ) c a r r i e d out a study on the P a c i f i c h e r r i n g s i m i l a r t o t h a t of Hodgson on the A t l a n t i c h e r r i n g and found complete agreement. In.some cases s c a l e r e a d i n g i s found t o be com p l i c a t e d by the presence of s o - c a l l e d " f a l s e " checks. S c a l e s t a k e n from f i s h of southern B r i t i s h Columbia waters are p a r t i c u -l a r l y d i f f i c u l t to i n t e r p r e t . However, t h e e r r o r i n t r o d u c e d i s not h i g h enough t o r a d i c a l l y a l t e r the r e s u l t s ( T e s t e r , . 19357). The Norwegian workers, H j o r t , D a h l , and Lea, found t h a t they were a b l e to d i f f e r e n t i a t e a c c u r a t e l y between f a l s e checks and t r u e w i n t e r r i n g s . Savage (1919) i n t r o d u c e d the use of p o l a r i z e d l i g h t as a means of d i s t i n g u i s h i n g f a l s e checks. -D'Arcy Thompson (.Sherriff, 1922) b e l i e v e s t h a t the r i n g s on t h e h e r r i n g s c a l e s are s u b j e c t to wide v a r i a t i o n i n a s i n g l e age-group, and, t h e r e f o r e , cannot be used f o r age de-t e r m i n a t i o n s . H i s argument has been r e p l i e d t o and r e f u t e d by t h e Norwegian w o r k e r s . Growth C a l c u l a t i o n s Perhaps the most important development from t h e s c a l e r e a d i n g , and the one w i t h which the present paper is~most concerned, i s , the b o d y / s c a l e growth r e l a t i o n s h i p . W a l t e r (1901) suggested t h a t i f each r i n g on t h e s c a l e r e p r e s e n t e d one year's growth, then i t would seem reasonable t h a t the growth of the f i s h f o r each y e a r of i t s l i f e c o u l d be c a l -c u l a t e d from t h e r e l a t i o n of each zone on the s c a l e t o t h e e n t i r e l e n g t h o f t h e s c a l e i n r e s p e c t to the t o t a l l e n g t h of t h e f i s h . Lea (1910), i n the Report on Norwegian F i s h e r y and.Marine I n v e s t i g a t i o n by Dr.. H j o r t , g i v e s t a b l e s of. measurements of a l a r g e number of f i s h showing t h a t he can c a l c u l a t e t o ' h i s s a t i s f a c t i o n the y e a r l y growth of t h e f i s h . He f i n d s t h a t the measurement (T) ( t h a t i s , the l e n g t h of the s c a l e d p o r t i o n of the body) g i v e s a more exact and con-s t a n t r a t i o to" t h e s c a l e growth than does the t o t a l l e n g t l f of t h e f i s h ( L ) . H e - i n t r o d u c e s a c o r r e c t i o n f a c t o r t o be a p p l i e d t o (L) as i t i s t h e more used l e n g t h , but a c t u a l l y the e r r o r i s so s m a l l t h a t he f e e l s t h a t I t i s not n e c e s s a r y t o use the c o r r e c t i o n f a c t o r . H i s e q u a t i o n i s s i m p l y L/7 = 1W 1/ vl» where y i , V2, e t c . r e p r e s e n t the measurements f o r each y e a r zone. The v a l i d i t y of h i s f o r m u l a i s q u e s t i o n e d by M i s s Lee .(1912) who f i n d s i n t h e c a l c u l a t e d increments f o r each year's growth what she c a l l s t h e "phenomenon of apparent change i n growth r a t e . " As c a l c u l a t i o n s a r e made f r o m the s c a l e s of f i s h of I n c r e a s i n g age t h e r e i s a decrease i n the c a l c u l a t e d l e n g t h of the f i s h i n i t s e a r l y y e a r s ; t h a t i s , t h e r e i s an apparent s h r i n k i n g i n the s i z e of t h e s c a l e as the f i s h gets o l d e r . Lee o f f e r s s e v e r a l e x p l a n a t i o n s f o r t h i s phenomenon, a l l of them m e c h a n i c a l c o n s i d e r a t i o n s . Lea (1913.) e x p l a i n s t h i s phenomenon as t h e n a t u r a l r e s u l t of the d i f f e r e n t i a l a d d i t i o n of slow- and f a s t - g r o w i n g f i s h t o mature s c h o o l s combined, w i t h the d i f f e r e n t i a l m o r t a l i t y of slow- .and f a s t -growing f i s h of the s c h o o l s of mature f i s h . Meek (1916) - 10 -f i n d s the e x p l a n a t i o n of Lea does not cover a l l the f a c t s . Lea has omitted t o take, i n t o c o n s i d e r a t i o n the f a c t t h a t the s c a l e s are not formed u n t i l the f i s h has reached some l e n g t h and must, t h e r e f o r e , grow r a p i d l y f o r a time to become l a r g e enough t o o v e r l a p . Meek i s a b l e t o show t o h i s s a t i s f a c t i o n , from samples of autumn spawned h e r r i n g , t h a t i f a c o r r e c t i o n f a c t o r i s i n t r o d u c e d t o maka up f o r the f a c t t h a t t h e s c a l e s are not formed u n t i l the f i s h i s 4 cm. l o n g the c a l c u l a t e d s i z e s a r e v e r y c l o s e t o the a c t u a l . E r a s e r (1916-17) a l s o .suggests t h a t Lee's phenomenon may be accounted f o r by t a k i n g i n t o c o n s i d e r a t i o n the f a c t t h a t the s c a l e s are not formed u n t i l the f i s h has a t t a i n e d some s i z e , and i n c a l c u l a t i o n s f o r t h e growth of the salmon shows t h a t Lee's phenomenon i s taken c a r e of by i n t r o d u c i n g t h i s c o r r e c t i o n . Molander (1918) shows t h a t i n h e r r i n g t h e r e was an u n d u l a t i n g e f f e c t between the growth of the f i s h and s c a l e over a p e r i o d of y e a r s . I n o t h e r words, the growth of t h e f i s h i s r e l a t i v e l y more r a p i d d u r i n g the e a r l y y e a r s , and that o f the s c a l e s i s more r a p i d d u r i n g t h e l a t e r y e a r s . He does not f i n d t h a t the c o r r e c t i o n f a c t o r a l t o g e t h e r e l i m i n a t e s the d i s c r e p a n c i e s encountered from th e c a l c u l a t i o n s suggested by the Norwegian w o r k e r s . Huntsman (1918) d e s c r i b e s graphs and measurements f o r f o u r s p e c i e s o f f i s h , among them Clupea harengus, i n w h i c h he f i n d s t h a t L/V ( t r a n s v e r s e diameter) decreases c o n t i n u o u s l y w i t h i n c r e a s i n g l e n g t h of f i s h ; L/W ( l o n g i t u d i n a l diameter) de-- 11 -c r e a s e s w i t h g r o w t h , a t f i r s t r a p i d l y h u t l a t e r e x t r e m e l y s l o w l y ; w h i l e L / X ( l o n g i t u d i n a l d i a m e t e r o f a n t e r i o r f i e l d ) d e c r e a s e s r a p i d l y a t f i r s t t h e n m o r e s l o w l y u n t i l i t b e c o m e s a p p r o x i m a t e l y $0 a t a b o d y l e n g t h o f a b o u t 22 c m . w h e n i t i n c r e a s e s s l i g h t l y u n t i l d e a t h . L e e (192 0) , i n a f u r t h e r s t u d y o f t h e a g e a n d g r o w t h o f s e v e r a l s p e c i e s o f f i s h , f i n d s t h a t , t h e s c a l e i n c r e m e n t i s , o n t h e Whole, i n d i r e c t p r o p o r -t i o n , t o . t h e i n c r e m e n t o f t h e f i s h . S h e b e l i e v e s t h a t t h e p h e n o m e n o n i s p a r t l y t a k e n c a r e o f b y t h e c o r r e c t i o n f o r t h e t i m e a t w h i c h t h e ' s c a l e s a r e f o r m e d . U s i n g E r a s e r ' s c o r r e c -t i o n f a c t o r , s h e s u g g e s t s t h e f o r m u l a L x = C + ( v l / V ( L - C ) i n w h i c h C i s t h e l e n g t h o f t h e f i s h a t t h e t i m e t h e s c a l e i s f o r m e d , L i s t h e l e n g t h o f t h e f i s h a t d e a t h , Y i s t h e s c a l e d i m e n s i on a t d e a t h , i s t h e c o m p u t e d l e n g t h ' a t t h e e n d o f t h e f i r s t y e a r , a n d v n i s t h e s c a l e l e n g t h a t t h e e n d . of t h e f i r s t a n n u l u s . M i s s S h e r r i f f (1928), f r o m m a t h e m a t i c a l d e -d u c t i o n s , o f f e r s t h e f o r m u l a L = A V 2 + B V + C w h i c h g i v e s a p a r a b o l i c c u r v e . M o n a s t u i r s k y (1926), a f t e r r e v i e w i n g t h e p r e v i o u s w o r k d o n e i n a n a t t e m p t t o c o r r e l a t e t h e s c a l e a n d b o d y g r o w t h , s u g g e s t s t h a t t h e o n l y w a y t o g e t - 1 2 -a s t r a i g h t l i n e r e l a t i o n s h i p i s t o t a k e t h e l o g a r i t h m i c v a l u e s and i n t h a t way t o e l i m i n a t e t h e d i f f e r e n c e b e t w e e n m a l e s a n d f e m a l e s . H i s s u g g e s t e d f o r m u l a I s L o g L = L o g K + n L o g x where x i s t h e measure o f t h e s c a l e s , n i s t h e s l o p e o f t h e l i n e , and t h e t e r m K I s f o u n d b y e x t r a p o l a t i o n . Van O o s t e n ( 1 9 2 8 ) s t u d i e d t h e s c a l e / b o d y r e l a t i o n i n t h e L a k e h e r r i n g t o a c c o u n t f o r L e e ' s phenomenon and t o d e t e r m i n e w h e t h e r o r n o t h e r f o r m u l a was a p p l i c a b l e . He d e c i d e d t h a t t h e l a t e a p p e a r -ance o f t h e s c a l e s was compensated f o r by t h e i r s u b s e q u e n t r a p i d r a t e o f g r o w t h i m m e d i a t e l y a f t e r f o r m a t i o n . He i s , t h e r e f o r e , of t h e o p i n i o n t h a t i n t h e L a k e h e r r i n g y e a r l y g r o w t h can b e more a c c u r a t e l y c a l c u l a t e d b y t h e s i m p l e s t r a i g h t l i n e r e l a t i o n s h i p L/V. 1 3 -THE,PROBLEM Prom a study o f ' t h e l i t e r a t u r e we f i n d t h a t t h e r e i s ample p r o o f , e i t h e r d i r e c t or I n d i r e c t , f o r the t h e o r y of age d e t e r m i n a t i o n based on the a n n u l i of s c a l e s of many s p e c i e s of f i s h i n g e n e r a l , and Of t h e h e r r i n g i n p a r t i c u l a r . The r e -l a t i o n of s c a l e growth t o body growth has been t h e s u b j e c t of i n v e s t i g a t i o n b y many a u t h o r s . I n some s p e c i e s of f i s h t h e r e l a t i o n i s found to be a simple one; i n othe r s i t i s complex. I n e i t h e r case there i s l i t t l e agreement between i n v e s t i g a -t o r s of d i f f e r e n t s p e c i e s , o r even between i n v e s t i g a t o r s of the same s p e c i e s . Much of t h i s work has been done on the A t l a n t i c h e r r i n g (Clupea harengus), a s p e c i e s c l o s e l y r e l a t e d to t he P a c i f i c h e r r i n g . The l a c k o f agreement between authors i n . t h i s s i n g l e s p e c i e s i n d i c a t e s the n e c e s s i t y Of f u r t h e r r e -s e a r c h . I t i s of p r i m a r y importance t o r a c i a l work t o have t h i s r e l a t i o n s h i p e s t a b l i s h e d , because v/hen i t i s p o s s i b l e t o c a l -c u l a t e a c c u r a t e l y the annual growth of the f i s h much of i t s l i f e h i s t o r y w i l l be r e v e a l e d . I n t h i s way t h e n e c e s s i t y f o r much of t h e sampling and t a g g i n g of h e r r i n g vdaich i s now b e i n g c a r r i e d on w i l l be e l i m i n a t e d . At t h e s u g g e s t i o n of Dr. A.L. T e s t e r , P a c i f i c B i o l o g i c a l S t a t i o n , Nanalmo, a d e t a i l e d study of the r e l a t i o n between the s c a l e and body growth of t h e h e r r i n g d u r i n g i t s f i r s t y e a r was - 14 -u n d e r t a k e n and t h e r e s u l t s o f - t h i s i n v e s t i g a t i o n w e r e com-b i n e d w i t h d a t a of a s i m i l a r n a t u r e f o r t h e a d u l t f i s h as c o m p i l e d by D r . T e s t e r . A s t u d y was a l s o made o f t h e v a r i -a t i o n i n s i z e and shape of t h e s c a l e s of t h e h e r r i n g and t h e t i m e o f f o r m a t i o n and o r d e r o f a p p e a r a n c e of t h e s c a l e s . T h i s a s p e c t o f t h e work w i l l b e c o n s i d e r e d f i r s t . - I M -MATERIAL The m a t e r i a l used f o r t h e p r e s e n t s t u d y c o n s i s t e d of twenty-one samples of h e r r i n g i n t h e i r f i r s t y e a r , c o n t a i n i n g i n a l l some f i f t e e n hundred f i s h r a n g i n g i n s i z e from 25 t o 120 mm., t o g e t h e r w i t h d a t a on s c a l e and body measurements of f i s h from I I t o 71 y e a r s of age p r o v i d e d by Dr. A.L* Tester. The c o l l e c t i o n s were made over a p e r i o d of n i n e y e a r s (l,9jl-1939) from v a r i o u s p o i n t s on t he south e a s t c o a s t of Vancouver i s l a n d . P r e v i o u s t o 1939, each y e a r was r e p r e s e n t e d by o n l y two or three'samples . Eor 1939 , however, i t was pos- / s i b l e t o g e t a s e r i e s of samples r e p r e s e n t i n g f i s h i n a l l the stages of t h e i r f i r s t y e a r . By combining the d a t a f o r s e v e r a l y e ars v a r i a t i o n which a r i s e s because of s m a l l numbers i n v o l v e d i n i n d i v i d u a l samples and v a r i a t i o n a r i s i n g from l o c a l and an-n u a l d i f f e r e n c e i n e n v i r o n m e n t a l f a c t o r s are both e l i m i n a t e d . The above samples were p r e s e r v e d i n e i t h e r f o r m a l i n o r a l c o -h o l . The a d u l t f i s h were from t h e commercial s c h o o l s t a k e n w i t h purse s e i n e from v a r i o u s f i s h i n g grounds i n s o u t h e r n B r i t i s h Columbia. Most samples were t a k e n during, the w i n t e r s 1931-32 t o 1934-35, and the d a t a f o r t h e separate age-groups were combined. I n f o r m a t i o n r e g a r d i n g t h e t i m e , p l a c e , and method of c a p t u r e , and the number and average l e n g t h of each o f t h e -.16 -samples of f i r s t year f i s h , and f o r t h e age groups of the o l d e r f i s h , i s g i v e n i n Table 1. - 17 -, -METHODS. For t h e s t u d y of t h e s i z e and shape of the s c a l e s of the h e r r i n g the m a t e r i a l c o n s i s t e d of a f i s h of.150 mm. i n l e n g t h i n i t s second year t a k e n from Pender harbour, J u l y , 1939' The s c a l e s were removed from t h e r i g h t s i d e of the body and mounted i n g l y c e r i n e . They were arranged on s l i d e s a c c o r d i n g t o l o n g i t u d i n a l rows s t a r t i n g from t h e V e n t r a l s u r f a c e . Mea-surements were made f o r t h e l e n g t h ( l ) of the l o n g i t u d i n a l a x i s of t h e a n t e r i o r f i e l d of the s c a l e , f o r the t o t a l l e n g t h ( t ) t a k e n a l o n g the a n t e r i o r - p o s t e r i o r a x i s , and f o r the w i d t h (w) . The s c a l e s on t h e l e f t s i d e were s i m i l a r l y r e -moved and mounted I n g l y c e r i n e on,a l a r g e p i e c e of paper to be used f or, r e f e r e n c e * From j u s t p o s t e r i o r t o the a n a l f i n the rows are r a t h e r d i f f i c u l t to f o l l o w o u t , and i n some- cases complete rows c o u l d not be o b t a i n e d . To study the time of f o r m a t i o n of the s c a l e s h e r r i n g from 25-4-0 mm. i n l e n g t h were used. These were taken f r o m the Departure bay samples f o r May, 193&, 'and 1939* The s k i n was removed from t h e l e f t s i d e of t h e body i n each case and c a r e -f u l l y c l e a n e d t o remove the pigment and adhe r i n g muscle t i s s u e . I n many cases the s k i n s were s t a i n e d i n Haematoxylin t o show up the p a p i l l a e more c l e a r l y . The p r e p a r a t i o n s were t h e n p l a c e d on g l a s s s l i d e s as w a t e r mounts, and, a f t e r examina-t i o n , a l l o w e d t o d r y . - 18 -To study the scale/body growth r e l a t i o n s h i p f o r t h e f i s h i n t h e i r f i r s t y e a r , measurements f o r each of t h e samples were c o n s i d e r e d s e p a r a t e l y , t h e n t h e complete s e r i e s c o n s i d e r e d t o -ge t h e r . I n most c a s e s f i v e s c a l e s were t a k e n f r o m each f i s h from t h e area at t h e end o f t h e f l e x e d p e c t o r a l f i n . When t a k i n g s c a l e s f rom t h e v e r y young f i s h c a r e must be t a k e n t o get r e p r e s e n t a t i v e s c a l e s f rom t h e rows below t h e l a t e r a l l i n e , s i n c e i t i s these s c a l e s which are used i n . t h e a d u l t f i s h a l -though the development i s l a t e r here t h a n p o s t e r i o r l y . The s c a l e measurements f o r each f i s h were averaged, t h e f i s h - w e r e arranged I n mm. groups, and the averages were t a k e n f o r the number of f i s h i n t h e group. S c a l e measurements c o n s i s t e d of the l e n g t h - o f t h e a n t e r i o r f i e l d ( l ) , and t o t a l l e n g t h of t h e s c a l e s ( t ) . I n a l l cases t h e s t a n d a r d l e n g t h .of t h e body (L) was used, t h a t i s , t h e l e n g t h from the t i p of t h e snout t o t h e end of t h e s i l v e r y a r e a on t he caudal peduncle. Data f o r t h e s t a n d a r d l e n g t h of t h e s c a l e f o r each mm. group, t o -gether w i t h t h e number of f i s h averaged f o r each mm. group, i s g i v e n i n Table 2 ( a ) . Table 2 (b); g i v e s s i m i l a r d ata f o r t h e t o t a l l e n g t h of t h e s c a l e . The s t a n d a r d l e n g t h measurements of the a d u l t h e r r i n g were t a k e n i n t h e f i e l d from samples of-the-.commercial catches. S c a l e s were t a k e n from i n d i v i d u a l s and p l a c e d i n books t o be examined l a t e r i n t h e l a b o r a t o r y . To f a c i l i t a t e r e a d i n g of the s c a l e s they were f i r s t soaked i n w a t e r then cleaned and - 19 -mounted between two g l a s s s l i d e s , . Only the • l e n g t h of the a n t e r i o r f i e l d ( 1 ) of t h e s e s c a l e s was measured. I n t h e 1 9 3 1 - 3 2 samples, s c a l e measurements were based on two to f o u r s c a l e s from each f i s h ; f o r t h e 1 9 3 2 - 3 3 and 1 9 3 4 - 3 5 m a t e r i a l , measurements were based on a s i n g l e s c a l e o n l y from each f i s h . The s c a l e s were examined w i t h the a i d of a Promar p r o j e c t o r and measured at a m a g n i f i c a t i o n of 1 8 . 8 f o r t h e young f i s h and 1 9 . 6 f o r t h e a d u l t f i s h . A l l o f th e measurements were converted t o a c t u a l s i z e . The young f i s h were p r e s e r v e d In e i t h e r a l c o h o l or f o r -m a l i n . I n some cases, the samples had been i n t h e p r e s e r v a t i v e f o r a number of ye a r s b e f o r e the measurements were made. I t was c o n s i d e r e d a d v i s a b l e to determine whether or not s h r i n k a g e had o c c u r r e d and, I f so, t o c a l c u l a t e a c o r r e c t i o n f a c t o r . Table 3 g i v e s the, l i s t of t h e m a t e r i a l i n which both f r e s h and p r e s e r v e d measurements were a v a i l a b l e , the p r e s e r v a t i v e i n each c a s e , the l e n g t h of time which t h e y were i n the p r e -s e r v a t i v e , and the amount of s h r i n k i n g . I t w i l l be n o t i c e d t h a t the r e s u l t s are v e r y e r r a t i c , p a r t i c u l a r l y i n a l c o h o l . The f i s h were I n c l i n e d t o s o f t e n i n t h e l a t t e r and. i n some cases have been s t r e t c h e d I n measuring. S i n c e t h e shri n k a g e i s so s l i g h t and w i l l be more or l e s s constant f o r the young f i s h , a c o r r e c t i o n f a c t o r was not used In t h e young f i s h . However, i n the g e n e r a l curve i t was c o n s i d e r e d a d v i s a b l e t o - 20 -use a c o r r e c t i o n f a c t o r o f + 1 mm. f o r the preserved m a t e r i a l . Van Oosten ( 1 9 2 8 ) c a l c u l a t e d a c o r r e c t i o n f a c t o r of 1 . 0 1 6 f o r the Lake h e r r i n g which he had preserved i n A% f o r m a l i n f o r a month. T h i s agrees v e r y c l o s e l y w i t h the above c a l c u l a -t i o n . W i l l i a m s o n ( 1 9 1 4 ) found no shri n k a g e i n the marine her-r i n g . - 21 -THE STRUCTURE, SHAPE, AND SIZE OE THE SCALES S t r u c t u r e The s c a l e s of the h e r r i n g are f l a t p l a t e s v a r y i n g i n s i z e and shape i n d i f f e r e n t p a r t s of the body. The m a j o r i t y of them are f a i r l y c l o s e to the shape 'shown i n F i g u r e 1. The a n t e r i o r p a r t of the s c a l e i s embedded i n the s k i n and i s d i s t i n g u i s h e d from the p o s t e r i o r (exposed) f i e l d by t h e r e g u -l a r p a t t e r n oh t h e s u r f a c e . The two f i e l d s are d i v i d e d by a t r a n s v e r s e b a s e - l i n e . The p o s t e r i o r f i e l d has, as a r u l e , a r a t h e r uneven edge, and the s u r f a c e markings are r e s t r i c t e d t o uneven l i n e s r u n n i n g i n from t h e margin. The markings on the a n t e r i o r f i e l d are f i n e t r a n s v e r s e s t r i a e l a i d v e r y c l o s e t o g e t h e r . Running aeross th e s t r i a e at a s l i g h t angle are f h e a v i e r l i n e s , the " r a d i i " , c o r r e s p o n d i n g t o l i n e s w h i c h ' r a d i -ate from t h e c e n t r e of the s c a l e i n many s p e c i e s of f i s h . I n the o l d e r f i s h another type of marking i s p r e s e n t . T h i s con-s i s t s of narrow c l e a r zones conforming t o the shape of the s c a l e . These zones are the " a n n u l i " or w i n t e r checks used i n age d e t e r m i n a t i o n s and'.growth c a l c u l a t i o n s . I n c r o s s - s e c t i o n , , t h e h e r r i n g , s c a l e i s f l a t l y t r i a n g u l a r i n shape, w i t h t h e base of the t r i a n g l e l y i n g i n c o n t a c t w i t h t h e body. The s c a l e i s composed of an i n n e r f i b r o u s l a y e r , and an outer c a l c i f i e d l a y e r c o v e r i n g the s i d e of the s c a l e away from th e body. The i n n e r l a y e r i n c r e a s e s i n s i z e by the f o r m a t i o n ' o f s u c c e s s i v e l y l a r g e r l a y e r s of f i b r e s which r u n both t a n g e n t i a l l y and r a d i a l l y (Lea, 1919)• Thus t h e h e i g h t of t h e t r i a n g l e i s formed by t h e l a r g e r number of l a m e l l a e . The outer l a y e r i s of e q u a l t h i c k n e s s over the e n t i r e s c a l e and i n c r e a s e s i n s i z e by the a d d i t i o n o f new m a t e r i a l t o the o u t s i d e edge. The s u p e r f i c i a l s c a l e markings are r e s t r i c t e d to t h i s l a y e r . . The macroscopic appearance of the h e r r i n g s c a l e d i f f e r s q u i t e e x t e n s i v e l y from the t y p i c a l c y c l o i d s c a l e . T h i s d i f -f e r e n c e i s apparent o n l y i n the s u p e r f i c i a l markings. I n c y -c l o i d s c a l e s of s p e c i e s of salmon and t r o u t the markings on the s c a l e a r e c i r c u l a r i n n a t u r e , conforming c l o s e l y t o the Shape of t h e s c a l e and c o v e r i n g the e n t i r e s c a l e i n younger f i s h . - Olde r s c a l e s have the p a t t e r n l e s s d i s t i n g u i s h a b l e p o s t e r i o r l y . W i n t e r checks a re formed'by zones of c i r c u l l formed c l o s e t o g e t h e r , as d i f f e r e n t i a t e d f r o m the summer zones of wide a p a r t markings. Shape and S i z e H e r r i n g s c a l e s a r e so ar r a n g e d t h a t b oth l o n g i t u d i n a l . and d i a g o n a l rows maybe r e c o g n i z e d . I n t h e pre s e n t d i s c u s s i o n the l o n g i t u d i n a l rows are numbered from t h e v e n t r a l t o . t h e d o r s a l s u r f a c e . A t t h e a n t e r i o r end of the f i s h t h e r e are t h i r t e e n l o n g i t u d i n a l rows oh e i t h e r s i d e of the body and a median d o r s a l row. To compensate f o r t h e .narrowing of the - 23 -body p o s t e r i o r l y , n o t o n l y do t h e s c a l e s become v e r y much s m a l l e r , b u t a l s o some o f the rows t a p e r o f f . The two v e n t r a l rows s t o p a t t h e a n a l f i n and t h e f i r s t d o r s a l row t a p e r s o f f j u s t p o s t e r i o r t o t h e d o r s a l f i n , l e a v i n g o n l y n i n e r o w s a t t h e p o s t e r i o r end o f t h e f i s h . L o n g i t u d i n a l l y t h e c o m p l e t e r o w s , i . e . , t h e f o u r t h t o t w e l f t h r o w s i n c l u s i v e , make up f i f t y - t w o (_+) v e r t i c a l r o w s . These rows r e a c h t o j u s t p a s t t h e o r i g i n o f t h e t a i l f i n . T h e r e a r e s m a l l s c a l e s on t h e c a u d a l p e d u n c l e , b u t t h e y d o n o t c o n f o r m t o t h e e x a c t r o w s . To c l a r i f y t h e f o l l o w i n g d i s c u s s i o n a f e w p o i n t s may be n o t e d . The shape of t h e s c a l e s w i l l be d i s c u s s e d f i r s t i n t h e f o u r t h t o t e n t h l o n g i t u d i n a l rows s i n c e t h e s e a r e t h e most r e g u l a r l y s h a p e d , t h e n i n t h e more d o r s a l rows and, f i n a l l y , i n t h e v e n t r a l r o w s . U n l e s s o t h e r w i s e s t a t e d , t h e "rows" r e f e r r e d t o a r e the l o n g i t u d i n a l r o w s . The shape o f t h e s c a l e s shows c o n s i d e r a b l e v a r i a t i o n . Over t h e body t h e most r e g u l a r l y shaped a r e t h e " s i d e " s c a l e s f o u n d i n t h e m e d i a n a n t e r i o r p a r t o f t h e b o d y . These s c a l e s a r e r o u g h l y c i r c u l a r , a s shown f o r t h e t y p i c a l s c a l e I n F i g u r e . 1 . P r o c e e d i n g p o s t e r i o r l y , t h e shape becomes more e l o n g a t e d , t h a t I s , t h e t r a n s v e r s e diameter- d e c r e a s e s more r a p i d l y t h a n d o e s t h e a n t e r o - p o s t e r i o r d i a m e t e r , u n t i l t h e s c a l e s a r e r o u g h l y e l l i p t i c a l i n t h e t a i l r e g i o n . T h i s change I n shape i s shown i n P l a t e s I I , I I I , and I V . O t h e r changes a r e a l s o n o t i c e a b l e . The p r o p o r t i o n o f a n t e r i o r t o p o s t e r i o r - 24 -f i e l d becomes l e s s and the contour i s more i r r e g u l a r . I n most ca s e s , as s c a l e s nearer t h e head are examined,(from about t h e e i g h t h v e r t i c a l row) -they become s m a l l e r w i t h l e s s decrease i n t r a n s v e r s e diameter. Those i n immediate c o n t a c t w i t h the g i l l c a v i t y are d i s t i n c t l y m o d i f i e d . I n g e n e r a l , these s c a l e s have the a n t e r i o r edge f l a t t e n e d to conform t o the shape of the edge of the c a v i t y . The f i r s t s c a l e of the f o u r t h row has the p o s t e r i o r f i e l d s l i g h t l y m o d i f i e d and the second s c a l e d e e p l y notched t o f I t around the p e c t o r a l f i n ( P l a t e IV; Row IV, 1, 2 ) . The f i r s t s c a l e of the f i f t h row. i s known as the b r a n c h i a l s c a l e and has a very c h a r a c t e r i s t i c shape ( P l a t e I I I ; Roy/ V, l ) . The second s c a l e of t h i s r o w . i s oblong, w i t h the p o s t e r i o r f i e l d t a p e r e d . The f i r s t s c a l e of each of t h e rows above a r e , i n g e n e r a l , w i d e r than l o n g , w i t h the a n t e r i o r edge f l a t t e n e d ( P l a t e IT; Row:XI, 1 ) . I n t h e two rows d o r s a l t o t h e t e n t h row. t h e s c a l e s , on t h e whole are of a comparative shape t o t h e m e d i a l rows. However, t h o s e c o r r e s -ponding t o the s i d e s c a l e s are n o t i c e a b l y s m a l l e r and l e s s e v e n l y shaped ( P l a t e I I ; Rows X I and X I I ) . I n the t h i r t e e n t h row the s c a l e s are v a r i e d i n shape; t h e y a r e s m a l l w i t h a l -most c i r c u l a r s t r i a e and t h e r e i s c o n s i d e r a b l e v a r i a t i o n i n the form.of t h e p o s t e r i o r f i e l d . The l a c k o f u n i f o r m i t y of the s c a l e s here i s due t o t h e extreme v a r i a t i o n i n t h e median d o r s a l s c a l e s . I n t h i s row the s c a l e s are r a d i c a l l y d i f f e r -ent i n appearance to; anywhere e l s e on the body. They a r e , 25 -f o r t h e most p a r t , b i l o b e d , and t h e a n t e r i o r f i e l d Is round and forms more th a n a s e m i - c i r c l e . Thus the s c a l e i s pinched i n at t h e b a s e - l i n e and spreads out a g a i n i n the p o s t e r i o r f i e l d , g i v i n g r a t h e r t h e appearance of a c l o s e d f a n . Towards the t a i l the p o s t e r i o r f i e l d becomes v e r y s m a l l and i s j u s t a bulbous p r o j e c t i o n from t h e a n t e r i o r f i e l d ( P l a t e I I ; Row XIV) . V e n t r a l l y from the f o u r t h row the shape of the s c a l e s becomes more r e c t a n g u l a r . The f i r s t v e n t r a l row i s m o d i f i e d i n t h a t i t has a f l a t edge l y i n g a l o n g the v e n t r a l l i n e . The s t r i a e change d i r e c t i o n somewhat t o converge t o t h e b a s e - l i n e on the f l a t s i d e of the s c a l e . The two v e n t r a l rows extend a n t e r i o r l y beyond t h e l i m i t s of t h e m a j o r i t y o f the v e r t i c a l rows to f i l l the v-shaped c a v i t y below t h e p e c t o r a l f i n . - I n t h i s r e g i o n t h e s c a l e s are v e r y s m a l l and i n t he second row the ones next t o the p e c t o r a l f i n have the p o s t e r i o r f i e l d grooved and notched t o f o r m a r e c e s s which e x a c t l y c o r r e s -ponds w i t h the contour of the a n t e r i o r edge of t h e p e c t o r a l f i n . As the two v e n t r a l rows proceed p o s t e r i o r l y they a r e i n t e r r u p t e d by the v e n t r a l f i n . I n t h e f i r s t row the a d j a c e n t s c a l e s are s m a l l , but i n the second row a v e r y much m o d i f i e d s c a l e d e v e l o p s . The a n t e r i o r f i e l d i s about the same s i z e as i n a d j o i n i n g s c a l e s , but t h e p o s t e r i o r f i e l d i s v e r y much elon g a t e d , so t h a t the edge of the f i n f i t s a l o n g the s i d e of - 26 -the s c a l e ( P l a t e IV; Row I I , 24). Between t h e v e n t r a l f i n and the a n a l f i n t h e f i r s t row i s m o d i f i e d to f o r m t h e "keeled"' s c a l e s . The lower edge of these s c a l e s i s notched t o f o r m a p r o j e c t i n g s p i n e i n t h e extreme cases. T h i s can he f e l t by r u n n i n g the f i n g e r a l o n g the v e n t r a l s u r f a c e . The second row i s m o d i f i e d s l i g h t l y i n t h a t the p o s t e r i o r f i e l d forms a r a t h e r sharp p r o j e c t i o n t o f i t over th e f i r s t row P l a t e IV; Row I I , 3 D . The l e n g t h of the a n t e r i o r f i e l d and the w i d t h of the s c a l e s f o r each l o n g i t u d i n a l , row have been f i g u r e d g r a p h i c a l l y by p l o t t i n g a g a i n s t t h e number of the s c a l e i n ^ r M - c ^ r l row ( F i g u r e 3)« Measurements and graphs show t h a t the r e g i o n of l e a s t v a r i a t i o n i n the s i z e of the " s i d e " s c a l e s l i e s be-tween t h e f o u r t h and t e n t h l o n g i t u d i n a l rows, and between.the e i g h t h and f i f t e e n t h o r t w e n t i e t h v e r t i c a l rows. From a con-s i d e r a t i o n of the. curves f o r the f o u r t h to t e n t h l o n g i t u d i n a l rows i t i s noted t h a t I n g e n e r a l t h e shape i s t h a t of an i n -v e r t e d "U", w i t h the maximum h e i g h t i n t h e r e g i o n of t h e s i d e s c a l e s . I t i s f u r t h e r noted t h a t the curve f o r the s i x t h row i s the f l a t t e s t , t h a t i s , shows l e a s t change' i n l e n g t h of t h e a n t e r i o r s c a l e from the head to t h e t a i l , o f t h e f i s h . On e i t h e r s i d e of t h i s row the curves become.progressively deeper, t h a t i s , the s i d e s c a l e s r e a c h a g r e a t e r - l e n g t h , but the pos-t e r i o r s c a l e s f o r a l l t h e rows'are comparative i n s i z e . Curves f o r t h e w i d t h g i v e a much g r e a t e r maximum yet both ends a p p r o x i m a t e t h e c u r v e s f o r t h e l e n g t h . T h i s i l l u s t r a t e s t h e change i n s h a p e m e n t i o n e d p r e v i o u s l y - a g r e a t e r . d e c r e a s e i n w i d t h t h a n i n l e n g t h a s s c a l e s a r e c o n s i d e r e d p r o g r e s s i v e l y p o s t e r i o r l y . I t has been m e n t i o n e d i n c e r t a i n c a s e s , e.g., a t t h e • v e n t r a l f i n , t h a t s p e c i a l i z e d s c a l e s a r e f o r m e d . To compen-s a t e f o r t h i s d i f f e r e n c e i n shape,.the a d j a c e n t s c a l e s a r e a l s o s l i g h t l y m o d i f i e d i n a l l cases.- The rows o f s c a l e s l y i n ^ a l o n g t h e s i d e o f t h e d o r s a l and a n a l f i n s a r e v e r y s m a l l and t h e e f f e c t i s f e l t i n t h e n e x t row. I f an a b n o r m a l l y s m a l l s c a l e i s f o u n d on t h e b o d y t h o s e s u r r o u n d i n g i t h ave become c o r r e s p o n d i n g l y l a r g e t o c o m p e n s a t e . I n t h e N o r w e g i a n h e r r i n g E i n a r L e a (1911) went t o some l e n g t h t o d e t e r m i n e w h i c h s c a l e d i m e n s i o n s show t h e b e s t c o r -r e l a t i o n , and a l s o ' w h i c h s c a l e s show t h e g r e a t e s t r e g u l a r i t y i n s h a p e . He p r e s e n t s g r a p h s t o show t h e v a r i a t i o n i n t h e s i z e (as measured f o r t h e a n t e r i o r f i e l d ) of t h e s c a l e s f r o m t h r e e rows of t h e same f i s h . V a r i a t i o n s I n t h e r ows a r e i n -d e p e n d e n t o f e a c h o t h e r , w h e r e a s e a c h o f t h e w i n t e r z o n e s o f s c a l e s f o r t h e same r o w shows t h e same t y p e o f c u r v e . He d e t e r m i n e d t h a t t h e l e a s t e r r o r i s t o be f o u n d i n s c a l e s t a k e n f r o m t h e a n t e r i o r p a r t o f t h e b o d y , and more p a r t i c u l a r -l y f r o m t h e 2^-29 myotome. Huntsman (1.918 )• c o r r o b o r a t e s L e a ' s f i n d i n g s . He t o o k v e r t i c a l r o w s i n . t h e A t l a n t i c h e r r i n g , , i n w h i c h t h r e e t o f i v e s c a l e s were t a k e n f r o m e a c h o f t h e l o n g i -- 28 -t u d i n a l rows, and t h e measurements a v e r a g e d . He c o n c l u d e d t h a t t h e s c a l e s f r o m t h e l 8 - 2 6 t h myomeres and t h e 4-10th . l o n g i t u d i n a l rows show t h e most u n i f o r m i t y . These r e s u l t s c o n f o r m q u i t e c l o s e l y w i t h t h o s e f o r t h e P a c i f i c h e r r i n g . The d i f f e r e n c e i n s i z e and shape o f t h e two f o r m s a l l o w s f o r some v a r i a t i o n i n r e s u l t s . M i s s E s d a i l e (1912) s t u d i e d t h e s c a l e s o f t h r e e s p e c i m e n s o f Salmo s a l a r , h u t a p a r t f r o m f i n d i n g a d i f f e r e n c e i n number o f a n n u l ! , h e r c o n c l u s i o n s a r e n o t d e f i n i t e . - 2 9 -TIME AND ORDER OF APPEARANCE OE SCALES Although an e x t e n s i v e 'amount of work has been done on the h i s t o l o g y and development of t h e t e l e o s t s c a l e , these i n v e s -t i g a t i o n s have not i n c l u d e d work on the h e r r i n g s c a l e . Time o n l y p e r m i t t e d the study of a s m a l l number of s e c t i o n s of the s k i n of t h e h e r r i n g , but by comparison w i t h work of such i n -v e s t i g a t o r s a s P a r r o t t ( 1 9 J 4 ) , Setna (1934) , and Neave (1936.),. on s p e c i e s of Salmo, f o r m a t i o n of the p a p i l l a e and subsequent s e c r e t i o n of s c a l e substance appears to be e s s e n t i a l l y s i m i l a r . The p a p i l l a e develop as a g g r e g a t i o n s of c e l l s i n the dermal l a y e r of t h e s k i n . Almost im m e d i a t e l y the c e l l s b e g i n t o se-c r e t e the s c a l e substance of both the f i b r o u s and the c a l c i -f i e d l a y e r s . P a p i l l a e f i r s t appear on the l a t e r a l l i n e i n the r e g i o n of and p o s t e r i o r to t h e a n a l f i n . A row of p a p i l l a e then develop above and below those f i r s t formed. At t h e time t h a t the t h r e e rows of p a p i l l a e are formed p o s t e r i o r l y a s ec-ond c e n t r e of growth appears a t t h e a n t e r i o r end j u s t below the l a t e r a l l i n e ( F i g u r e 4 ) . By the time the p a p i l l a e are formed most of t h e way along t h i s l i n e , v e r y s m a l l s c a l e s are apparent on the l a t e r a l l i n e p o s t e r i o r l y . Development p r o -ceeds more r a p i d l y d o r s a l l y and v e n t r a l l y than l o n g i t u d i n a l l y . A t h i r d c e n t r e of growth makes i t s appearance on the a n t e r i o r v e n t r a l s u r f a c e . At about t h e time t h a t t h e r e are t h r e e rows of p a p i l l a e formed i n the median, r e g i o n , t h e r e are two rows of s c a l e s present on the v e n t r a l s u r f a c e . D i f f e r e n c e i n the time of appearance of the s c a l e s i s i l l u s t r a t e d i n F i g u r e 5. P o s t e r i o r l y f i v e rows of s c a l e s and a row of p a p i l l a e are p r e s e n t , w h i l e a n t e r i o r l y t h e r e are t h r e e rows of s c a l e s w i t h a row of p a p i l l a e above and below, and m e d i a l l y t h r e e rows of s c a l e s occur w i t h two rows of s c a l e s below; t h e r e are two rows of s c a l e s v e n t r a l l y . From t h i s i t may be conclu d e d t h a t t h e development i s more r a p i d f r o m p o s t e r i o r t o a n t e r i o r t h a n from a n t e r i o r t o p o s t e r i o r . I n the next stage ( F i g u r e 6) a row of p a p i l l a e has formed above the v e n t r a l ro¥*rs of s c a l e s , and a l a s t row below the a n t e r i o r median growth c e n t r e . There are now i n d i c a t i o n s of t e n complete rows of s c a l e s a n t e r i o r t o t h e a n a l f i n , and seven rows p o s t e r i o r t o t h i s p o i n t . S c a l e s are pre s e n t on -a l l the p o s t e r i o r , roves'-.with the e x c e p t i o n of t h e most d o r s a l and the most v e n t r a l . The l a t t e r I s c o n t i n u o u s - w i t h the t h i r d row of s c a l e s a t the a n t e r i o r end. The t h r e e rows of d o r s a l s c a l e s , and the median d o r s a l row are developed l a t e r . K l a a t s c h ( I 8 9 O ) was of the o p i n i o n t h a t t h e s c a l e s of the t r o u t were formed f i r s t a t t h e a n t e r i o r of t h e body. Huntsman (1918} found t h a t i n t h e A t l a n t i c h e r r i n g t h e r e are th r e e growth c e n t r e s ; one p o s t e r i o r , a second a n t e r i o r , and a t h i r d v e n t r a l , which agrees w i t h those found i n the P a c i f i c h e r r i n g . Paget ( 1 9 2 0 ) confirms t h e s e f i n d i n g s . I n t h e t r o u t the f i r s t s c a l e s formed are those of the l a t e r a l l i n e w i t h the - 3 1 -p a p i l l a e f o r m i n g above a n d b e l o w b e f o r e t h e c o m p l e t i o n of t h e l i n e a n t e r i o r l y and p o s t e r i o r l y . D e velopment i s more r a p i d p o s t e r i o r t o t h e d o r s a l f i n t h a n a n t e r i o r l y . I n s p e c i e s o f S almo, P a r r o t t ( 1 9 3 4 ) , S e t n a ( 1 9 3 4 ) , and Weave (1936) have r e c o r d e d s i m i l a r o b s e r v a t i o n s . E l s o n (1939) n o t o n l y c o n -f i r m s t h e above, but a l s o g i v e s i n much more d e t a i l a d e s c r i p -t i o n o f t h e way i n w h i c h t h e s p r e a d o f t h e s c a l e s t a k e s p l a c e . He c o n c l u d e s t h a t " f r o m t h e p l a c e o f o r i g i n on t h e l a t e r a l l i n e d e v e l o p m e n t p r o c e e d s by o u t g r o w t h s f r o m t h e p r i m a r y p a p i l l a e , w h i c h grow o b l i q u e l y f o r w a r d n e a r l y a t r i g h t a n g l e s • t o t h e d i r e c t i o n o f t h e myocommata. S c a l e p a p i l l a e a r e f o r m e d a l o n g t h e s e o u t g r o w t h s . Development i s most r a p i d i n t h e a r e a b e l o w t h e a d i p o s e f i n , and s l o w e s t b e l o w t h e d o r s a l f i n . " F i r s t p a p i l l a e a r e f o u n d i n t h e h e r r i n g a t a b o u t 2 9 mm. w i t h s l i g h t v a r i a t i o n i n i n d i v i d u a l f i s h . A t 3 .2-34 mm. s c a l a -. t i o n i s c o m p l e t e d t o t h e p o i n t where e i t h e r t h e s c a l e s - o r t h e p a p i l l a e g i v e a . p i c t u r e o f a l l b u t t h e t h r e e d o r s a l rows o f s c a l e s . A t - t h i s t i m e , i n most c a s e s , s m a l l s c a l e s a r e p r e s e n t i n t h e a r e a u s e d I n measurements. From 35-37 mm. s c a l e s a r e f o r m e d on a l l b u t t h e d o r s a l r o w s . The t i m e o f f o r m a t i o n of t h e s c a l e s v a r i e s i n d i f f e r e n t s p e c i e s . Huntsman ( 1 9 1 8 ) f o u n d t h a t I n t h e A t l a n t i c h e r r i n g t h e s c a l e s b e g i n t o f o r m a t a b o u t 4 8 mm. The f i r s t a p p e a r i n g s c a l e s a r e r o u n d t o o v o i d r e g u l a r -shaped b o d i e s f r e e f r o m s t r i a t i o n . W i t h i n a v e r y s h o r t t i m e - 32 s t r i a e b e g i n to f o r m a t the a n t e r i o r end. The s c a l e s develop very r a p i d l y , f o r by the time t h r e e rows are formed a n t e r i o r l y the p o s t e r i o r s c a l e s show some s t r i a t i o n , and s c a l e s of the f i r s t formed row have two or t h r e e s t r i a e by the time the p a p i l l a e show i n t h e t h i r d row down. On the same f i s h , s c a l e s showing no s t r i a e and s c a l e s w i t h 10-15 rows of s t r i a e a re t o be found ( F i g u r e 2). The b a s e - l i n e which s e p a r a t e s t h e an-t e r i o r and p o s t e r i o r f i e l d s i s not apparent when t h e s t r i a t i o n s are f i r s t formed. When t h e s c a l e s a r e f i r s t formed t h e y a re co m p l e t e l y covered by e p i d e r m i s , t h a t i s , they l i e r i g h t i n -s i d e the s c a l e pocket,. I t i s the o p i n i o n of t h e author t h a t the b a s e - l i n e i s formed a t t h e time t h a t t h e p o s t e r i o r f i e l d comes .to p r o j e c t o u t s i d e the pocket t o t h e p o i n t at which the s t r i a e are formed. The b a s a l l i n e ' i s p r esent i n s c a l e s of \ 37'mm. f i s h i n the r e g i o n s where the s c a l e s are f i r s t formed, and at the time t h a t the b a s e - l i n e i s p r e s e n t t h e r e a r e t e n t o f i f t e e n s t r i a e formed. The r e l a t i v e growth of the a n t e r i o r and p o s t e r i o r f i e l d s i s d i s c u s s e d l a t e r i n c o n n e c t i o n w i t h s c a l e growth d u r i n g the f i r s t y e a r . The p o s t e r i o r s c a l e s de-ve l o p more r a p i d l y at f i r s t and are l a r g e r t h a n -elsewhere on the body. T h i s i s due i n p a r t t o the f a c t t h a t they a r e the f i r s t t o develop. Before v e r y l o n g , however, the a n t e r i o r s c a l e s i n c r e a s e i n s i z e t o exceed those i n the p o s t e r i o r , which c o n d i t i o n shows v e r y markedly i n the a d u l t f i s h . P a p i l l a e are formed one t o each myomere u n t i l a p p r o x i -- 33 -mately the r e g i o n of the a n a l f i n , when they take up a p o s i -t i o n i n r e l a t i o n t o the intermyotomic r e g i o n s . Thus one s c a l e l e s s i s formed than t h e r e are myotomes. Towards the p o s t e r i o r end t h i s r e l a t i o n becomes m o d i f i e d , and t a k i n g i n t o c o n s i d e r a t i o n the s c a l e s on the c a u d a l peduncle, the number of s c a l e s i s s l i g h t l y i n excess of t h e number of v e r t e b r a e . T h i s i s i n s u b s t a n t i a l agreement w i t h the f i n d i n g s of Huntsman f o r the A t l a n t i c h e r r i n g . The r e l a t i o n of the s c a l e s to the myo-meres has been a s u b j e c t of some d i s c u s s i o n and t h e r e i s a ve r y great d i f f e r e n c e i n v a r i o u s s p e c i e s of f i s h . I vlaatsch found no r e l a t i o n i n the. t r o u t between the s c a l e s and myomeres. Hase (1907) s t a t e s t h a t i n the normal t e l e o s t s d e f i n i t e seg-mental r e l a t i o n e x i s t s between s c a l e s and somites. Regan (1910) found t h a t of two s p e c i e s o f E l o p s , E. machnata has 63-64 v e r t e b r a e and 94-98 s c a l e s ; E. l a c e r t a has 74 v e r t e b r a e and 74-83 s c a l e s ; o t h e r s 78-79 v e r t e b r a e and 102-118 s c a l e s . Paget (1,920) s t a t e s t h a t f o r t h e A t l a n t i c h e r r i n g there a r e 56- 59 v e r t e b r a e and 53-59 scales'; the p i l c h a r d has 53 v e r t e -brae and 32 s c a l e s ; w h i l e I n Salmonldae the number of s c a l e s i s r o u g h l y double the number of v e r t e b r a e , S. f a r i o h aving 57- 58 v e r t e b r a e and 120 s c a l e s . There a r e , however, v a r i a -t i o n s even w i t h i n a group. - 34 -RELATIVE GROWTH OF BODY AND SCALE R e l a t i v e Growth of Body and S c a l e of F i s h i n T h e i r F i r s t Year. I n t h i s i n v e s t i g a t i o n the data f o r each sample were worked out s e p a r a t e l y . I n each case the l e n g t h of the s c a l e (1) was p l o t t e d a g a i n s t the l e n g t h of t h e f i s h (L) , and the best free-hand curve drawn through the p o i n t s . A l l the cur v e s were then put on a s i n g l e graph and the b e s t free-hand curve drawn through them. The curves f o r t h e s e p a r a t e samples p l o t t e d on the same c o o r d i n a t e s o f f e r some i n t e r e s t i n g comparisons ( F i g u r e 7)-C o n s i d e r a t i o n of some of the i n d i v i d u a l curves shows a v a r i -a t i o n i n growth r a t e , not o n l y between samples from d i f f e r e n t l o c a l i t i e s i n the same y e a r , but a l s o between samples from the same l o c a l i t i e s f o r d i f f e r e n t y e a r s . I n F i g u r e ^y^JEj^aiiA-^ H r e p r e s e n t curves f o r samples taken i n t h r e e d i f f e r e n t y e a r s from Departure bay. Sample C was taken a t t h e end of May, 1931, and sample H at t h e end of June, 1939- The average s i z e o f these samples i s t h e same, y e t i t yrould appear t h a t t h e r e was a month's d i f f e r e n c e i n age. Sample E, taken i n -1934, l i e s w e l l above t h e g e n e r a l t r e n d of t h e c u r v e , which means t h a t t h e s c a l e s were l a r g e r than average I n p r o p o r t i o n t o t he s i z e of t h e f i s h . Samples from A c t i v e pass and P o r l i e r pass taken on the same day have a d i f f e r e n c e i n average l e n g t h of 10 mm. A sample from Deep bay, June, 1939 ( F i g u r e 7, K) - 35 -and a sample f r o m D e p a r t u r e bay t a k e n ' t w o weeks l a t e r ( F i g u r e 7, H) show o n l y a s l i g h t d i f f e r e n c e i n a v e r a g e l e n g t h , b u t t h e s c a l e s o f t h e Deep b a y f i s h a r e n o t i c e a b l y s m a l l e r t h a n a r e t h o s e o f t h e D e p a r t u r e b a y sam p l e . The Deep bay sa m p l e i s a p p a r e n t l y t h e f a s t e r - g r o w i n g s a m p l e , and has s m a l l e r s c a l e s . T h i s shows a p p a r e n t c o n f i r m a t i o n o f t h e t h e o r y t h a t f a s t -g r o w i n g f i s h have s m a l l e r s c a l e s t h a n s l o w - g r o w i n g f i s h o f t h e . .same l e n g t h . On t h e o t h e r h a n d , s a m p l e s f r o m T r i n c o m a l i c h a n -n e l and L a d y s m i t h h a r b o u r show j u s t t h e o p p o s i t e r e l a t i o n . ' The T r i n c o m a l i c h a n n e l f i s h were t a k e n I n November, 1936, and t h e L a d y s m i t h harbour, f i s h w e r e t a k e n i n M a r c h , 1939. I t w o u l d b e ' - r e a s o n a b l e t o assume t h a t t h e f o r m e r w e r e a t t h e end o f t h e i r f i r s t y e a r ' s g r o w t h and t h e l a t t e r had n o t y e t s t a r t e d t h e i r s e c o n d y e a r ' s g r o w t h , s i n c e L e a (1919), F r a s e r (1917), and Hodgson ( 1 9 2 5 ) , have f o u n d f o r t h e h e r r i n g , s a l m o n , and E n g l i s h h e r r i n g , r e s p e c t i v e l y , t h a t g r o w t h i s c h e c k e d f r o m November t o M a r c h . The w i n t e r c h e c k s on t h e s c a l e s a r e ap-p a r e n t d u r i n g t h i s t i m e . The two s a m p l e s , t h e r e f o r e , a r e com-p a r a b l e i n age a s b e i n g a t t h e end o f t h e i r f i r s t y e a r ' s g r o w t h . The a v e r a g e l e n g t h f o r t h e T r i n c o m a l i sample i s 95.1 mm., and f o r t h e L a d y s m i t h sample 89.0 mm.; t h u s we w o u l d a s -sume t h a t t h e f o r m e r a r e t h e f a s t e r - g r o w i n g f i s h . However, I n d i r e c t c o n t r a d i c t i o n o f t h e t h e o r y , t h e s c a l e s o f t h e f o r m e r a r e t h e l a r g e r . Thus f o r t h e f i r s t y e a r f i s h we f i n d v a r i a t i o n i n t h e - 356 -growth r a t e v e r y marked. The d i f f e r e n c e w i t h i n the same year samples may he accounted f o r i n p a r t by d i f f e r e n t spawning t i m e s , and i n p a r t by environmental f a c t o r s such as tempera-t u r e and food . V a r i a t i o n s i n these f a c t o r s w i l l have a'much more n o t i c e a b l e e f f e c t on young than on mature f i s h . A g a i n , the e r r a t i c shape of some o f t h e curves can be accounted f o r by the s m a l l s i z e o f the samples. The T o r l i e r pass sample was l a r g e enough t o get t e n f i s h f o r each mm. group, and thus have a r e p r e s e n t a t i v e average. The r e s u l t was a v e r y smooth curve which was i n c o n t r a s t t o the shape of the curve f o r such a s m a l l sample as t h a t of Departure bay, 19351 ( F i g u r e 7, C ) . We w i l l f i r s t c o n s i d e r the curve f o r growth o f the an-t e r i o r f i e l d ( F i g u r e 8,.(1))-. The o r i g i n of the curve i s . at about 32 mm. body l e n g t h . There i s a s h a r p r i s e i n the curve up t o a p p r o x i m a t e l y 40-30 mm-., which i n d i c a t e s t h a t t h e s c a l e i s growing f a s t e r than the body. The sl o p e g r a d u a l l y drops o f f t o n e a r l y a s t r a i g h t l i n e . However, i f a s t r a i g h t l i n e i s drawn t o u c h i n g t h e h e i g h t s . o f t h e c u r v e , i t i s observed t h a t the. g r e a t e r p a r t of t h e p l o t t e d graph l i e s below the l i n e . T h i s d i p i n the curve shows t h a t the growth r a t e of the s c a l e p r o g r e s s i v e l y decreases i n r e s p e c t t o t h a t of the body f o r a t i m e , then shows a s l i g h t I n c r e a s e i n r a t e . The r a t e , however, remains, below t h a t ' of t h e body. At a p p r o x i -m a t e l y 40 mm. the a n t e r i o r f i e l d of t h e s c a l e i s a c t u a l l y - 37 -l a r g e r i n r e l a t i o n t o t h e b o d y t h a n a t any o t h e r t i m e . The r a t i o o f l / L i s g i v e n b e l o w f o r e a c h cm. i n c r e a s e o f bo d y l e n g t h ( t a k e n f r o m t h e c o m p l e t e c u r v e , F i g u r e 11). When l / L i's g r e a t e r t h a n one t h e s c a l e has t h e f a s t e r r a t e o f g r o w t h . W i t h d e c r e a s e i n t h e r a t i o t h e s c a l e r a t e i s d e c r e a s i n g b e l o w t h a t o f t h e body , e.g., f r o m 50-110 mm., w i t h a s l i g h t i n -c r e a s e i n s c a l e r a t e a t a b o u t 60-70 mm. L i n mm. l / L 35 1.27 40 1.00 50 .77 60 .65 70 .69 80 .67 90 .67 100 .67 110 t .67 120 .67 130 .76 140 -73 150 .76 160 .79 170 .85 180 .85 190 .89 200 1.00 205 .92 210 .67 The g r o w t h r a t e o f t o t a l s c a l e ( t ) i n r e l a t i o n t o t h e g r o w t h r a t e o f body l e n g t h (L) g i v e s a much more c o n s t a n t c u r v e . The r a p i d g r o w t h r a t e o f t h e s c a l e c o n t i n u e s u n t i l t h e f i s h i s a b o u t 50 mm. l o n g , t h e n t h e g r o w t h o f t h e s c a l e and t h e g r o w t h - 38 -of the body con t i n u e a t a u n i f o r m r a t e f o r some time. At about 8-0 mm. the s c a l e rate,becomes v e r y s l i g h t l y l e s s than t h a t of t h e body. The r a t i o t / L f o r each cm. group i s g i v e n below: L i n mm. t/ L 50 40 = 1.10 50 1.00 6o 1.00 70 .1.00 8o .92-90 .92 100 . 92 110 •92 115 .92 120 • 92 I t i s apparent t h a t a f t e r t h e s c a l e s have reached a c e r t a i n s i z e the . growth r a t e of body and ..scale i s almost c o n s t a n t . The n a t u r e of the c u r v e Is shown i n F i g u r e 8. The c u r v e , wnen p r o j e c t e d to t h e b a s e - l i n e , g i v e s too g r a d u a l a slope at t h e b e g i n n i n g . The a c t u a l curve i s r e p r e s e n t e d by the d o t t e d l i n e . T h i s v e r y sharp r i s e a t the bottom of the curve i s caused.by the i n s t a n t a n e o u s f o r m a t i o n of a -scale of n o t i c e a b l e s i z e by the p a p i l l a e , r a t h e r than the expected g r a d u a l growth from a v e r y s m a l l c e n t r e . The p r o j e c t e d curve g i v e s the approximate time of f i r s t f o r m a t i o n of the p a p i l l a e . : I n t r a c i n g the growth of the a n t e r i o r f i e l d of the s c a l e w i t h growth of the body t h e r e a r e two d i f f e r e n t i a l growth r a t e s t o be c o n s i d e r e d . At f i r s t the a n t e r i o r f i e l d i s I n -c r e a s i n g i n p r o p o r t i o n t o t h e e n t i r e s c a l e , w h i c h i s i n t u r n i n c r e a s i n g i n r e l a t i o n , t o t h e body. The change i n p r o p o r t i o n of t h e a n t e r i o r f i e l d t o t h e e n t i r e s c a l e i s shown g r a p h i c a l l y i"n F i g u r e 9, and c a l c u l a t i o n s o f (1) as a p e r c e n t o f ( t ) a r e g i v e n i n T a b l e 4. The l / t r a t i o f o r e a c h era. g r o u p i s shown b e l o w : L i n mm. l / t 35 =26 40 .43 50 .56 60 .57 70 .58 80 .60 90 .62 100 .65 110 .65 120 .67 A t t h e end o f t h e f i r s t y e a r t h e a n t e r i o r f i e l d i s a p p r o x i -m a t e l y s i x t y - f o u r p e r c e n t o f t h e s c a l e . I t i n c r e a s e s f r o m z e r o p e r c e n t t o f i f t y p e r c e n t b y a body l e n g t h o f a b o u t 45 mm., t h e r e m a i n i n g f o u r t e e n p e r c e n t i s n o t a t t a i n e d u n t i l t h e end o f t h e y e a r ' s g r o w t h (120 mm.). The c u r v e f o r t h e e n t i r e s c a l e ( F i g u r e 8), as d i s c u s s e d e a r l i e r , shows a r a p i d r a t e o f g r o w t h up t o a b o d y l e n g t h o f a b o u t 40-50 mm., t h e n a d e c r e a s e I n r a t e t o b e c o n s t a n t w i t h t h a t o f t h e body; whereas t h e c u r v e f o r t h e a n t e r i o r f i e l d shows a s i m i l a r r a p i d r a t e o f g r o w t h up t o a b o d y l e n g t h o f a b o u t 40 mm., t h e n a d e c r e a s e i n r a t e u n t i l i t becomes e v e n l e s s t h a n t h a t o f t h e body. - 4 0 -T h i s c o n t i n u e d s l o w i n g up o f t h e g r o w t h r a t e o f t h e a n t e r i o r f i e l d i s a c c o u n t e d f o r by t h e f a c t t h a t b o t h t h e r a t e o f g r o w t h o f t h e e n t i r e s c a l e and t h a t of t h e a n t e r i o r f i e l d show a d e c r e a s e i n r e l a t i o n t o t h e r a t e of g r o w t h o f t h e body. G r o w t h i n L e n g t h o f t h e A n t e r i o r F i e l d ( l ) i n R e l a t i o n t o t h e  Body L e n g t h (L) i n F i s h More t h a n One Y e a r o f Age. F o r t h e I I y e a r a g e - g r o u p t h e combined r e s u l t s o f s e v e r a l y e a r s were c o n s i d e r e d t o g e t h e r and g r o u p e d i n t o t h r e e a v e r a g e s w i t h an e q u a l number of f i s h i n e a c h g r o u p . The a v e r a g e s were t a k e n f o r b o t h (1) and (L) and t h e p o i n t s p l o t t e d . F o r t h e I I I , I V , V, and V I y e a r a g e - g r o u p s i n w h i c h t h e r e w e r e more f i s h t a k e n o v e r l e s s number o f y e a r s , t h e same t h i n g was done b u t o n l y f o r t h e s e p a r a t e y e a r s 1931-32, 1932-33, and 1934-35. Then i n ea c h c a s e t h e t h r e e s e t s o f a v e r a g e s were t h e m s e l v e s a v e r a g e d u s i n g t h e u n w e i g h t e d a v e r a g e . The d a t a i s g i v e n i n T a b l e 5 . The s e p a r a t e a g e - g r o u p s w e r e p l o t t e d a s i n F i g u r e 10, and t h e g e n e r a l c u r v e drawn t h r o u g h t h e m i d - p o i n t o f e a c h c u r v e . The r e s u l t i n g g r a p h s show a p a r t i c u l a r l y i n t e r e s t i n g f e a t u r e i n t h a t e ach o f t h e a g e - g r o u p s l i e s on a d i s t i n c t c u r v e w h i c h i s a t a s l i g h t a n g l e t o t h e g e n e r a l c u r v e . T h i s may be e x p l a i n e d on t h e b a s i s o f d i f f e r e n t i a l s c a l e g r o w t h b e t w e e n f a s t - and s l o w - g r o w i n g f i s h . The s l o w - g r o w i n g f i s h h a s a l a r g e r s c a l e ( on t h e a v e r a g e ) t h a n a f a s t - g r o w i n g f i s h - 41 -of* the same s i z e . F o r example, a f i s h of 190 mm. i n i t s t h i r d year has an average s c a l e l e n g t h of 4 . 6 mm., whereas a f i s h of the same l e n g t h i n i t s f o u r t h year has an average s c a l e l e n g t h o"f 4 . 7 mm. F u r t h e r , t h e r e i s a tendency f o r each of the curves to be convex. T h i s I n d i c a t e s a r e t a r d i n g of the d i f f e r e n t i a l growth of the s c a l e and t h e body i n the f a s t - g r o w i n g f i s h . The g e n e r a l curve drawn through the c e n t r e of t h e averages of the i n d i v i d u a l c u r v e s f o r t h e d i f f e r e n t ages shows a d e f i n i t e r e t a r d i n g of the r a t e of s c a l e growth between t h e ages of V and V I . I t i s u n f o r t u n a t e t h a t t h e r e are not da t a a v a i l a b l e i n the o l d e r age-groups t o show the c o n t i n u a t i o n of t h e curve f o r the ( t ) t o (L) r e l a t i o n as i t has been worked out f o r t h e f i r s t year f i s h The combined d a t a f o r a l l the specimens examined are shown g r a p h i c a l l y i n F i g u r e 11. T h i s curve r e p r e s e n t s the growth curve f o r t h e b o d y / s c a l e r e l a t i o n f o r t h e f i r s t s i x yea r s of the l i f e of the P a c i f i c h e r r i n g . I t i s obvious t h a t t h e r e i s c o n s i d e r a b l e d i v e r s i o n from t h e s t r a i g h t l i n e r e l a -t i o n such as t h a t suggested by Lea (1910') or even as g i v e n i n the f o r m u l a i n v o l v i n g t h e c o r r e c t i o n f a c t o r of F r a s e r . The v a r y i n g growth r a t e d u r i n g the f i r s t year has a l r e a d y been g i v e n i n some d e t a i l (p. " 5 6 ) . : A t t h e end of the f i r s t year the s c a l e i s s t i l l growing more s l o w l y than t h e body. S h o r t l y a f t e r t h e b e g i n n i n g of the second y e a r , between 120 mm. and - 42 -130 mm., the s c a l e growth begins to i n c r e a s e s l i g h t l y over t h a t of the body and c o n t i n u e s to i n c r e a s e up t o t h e end of the t h i r d year (190-200 mm.). D u r i n g the f o u r t h and f i f t h y ears the curve f l a t t e n s out somewhat and from the c a l c u l a -t i o n s on page 3 7 . f o r the l / L r e l a t i o n s h i p i t i s seen t h a t the s c a l e and body are.growing at the same-rate f o r a t i m e . The end of t h e curve drops o f f a g a i n , I n d i c a t i n g that t h e s c a l e growth has slowed i n r e l a t i o n to t h e body. Due t o t h i s con-t i n u a l change i n r e l a t i v e growth r a t e , i t would be d i f f i c u l t t o d e r i v e a s i n g l e f o r m u l a to f i t the e n t i r e c u r v e . Huntsman (1918) worked out t h e growth r e l a t i o n of f o u r s p e c i e s of f i s h f o r s e v e r a l d i f f e r e n t s c a l e dimensions. I n -cluded i n t h i s m a t e r i a l i s t h e growth r e l a t i o n of t h e a n t e r i o r f i e l d of the s c a l e t o the body l e n g t h ( i n c l u d i n g the t a i l f i n ) of t h e f i s h i n the A t l a n t i c h e r r i n g . T h i s curve I s much more n e a r l y i n the n a t u r e o f the curve o b t a i n e d of the t o t a l l e n g t h of t h e s c a l e i n t h e present case. He f i n d s t h a t a f t e r the f i r s t r a p i d .growth of the s c a l e the r a t e decreases to s l i g h t l y l e s s t h a n t h a t of t h e body growth of t h e f i s h , i s c o n s t a n t f o r some time, then i n t h e o l d e r f i s h i n c r e a s e s . Van Oosten (1928) has a curve f o r t h e s c a l e / b o d y r e l a t i o n i n the young Lake h e r r i n g , which shows a steady i n c r e a s e i n s c a l e growth over t h a t of body growth up t o a p p r o x i m a t e l y 60 mm., then a g r a d u a l dropping o f f . T h i s g i v e s a p a r b o l i c t ype of curve. For t h e o l d e r f i s h he f i n d s t h a t , i f t h e t o t a l l e n g t h of the - 4 3 -s c a l e i s used, the f o r m u l a of Lea h o l d s more n e a r l y than does . the f o r m u l a i n v o l v i n g the c o r r e c t i o n f a c t o r . Van Oosten con-s i d e r s t h a t the r a p i d growth j u s t a f t e r the time of f o r m a t i o n of the s c a l e makes up f o r the l a t e r development. However, Greaser . (1926), working w i t h the s u n f i s h Eupomo.tis gibbosus, f i n d s growth r e l a t i o n s t h a t g i v e a curve almost i d e n t i c a l w i t h the present curve. "The l i n e s t a r t s at 15 mm. from the z e r o -zero p o i n t and q u i c k l y r i s e s t o 30 mm., due to the i n c l u s i o n of one-half of t h e f o c u s i n the measurements. At f i r s t the s c a l e g r a d u a l l y grows more r a p i d l y i n p r o p o r t i o n than the f i s h l e n g t h and the curve, bends upward. T h i s continues and i s i n -creased more at a f i s h l e n g t h of about 80 mm. As the f i s h reaches 120 mm. i n l e n g t h , the s c a l e grows p r o p o r t i o n a t e l y l e s s than the f i s h , r e s u l t i n g i n / a sharp t u r n of t h e curve f o l l o w e d by a g r a d u a l downward t r e n d . I n t h i s manner a char-a c t e r i s t i c sigmoid curve i s formed, showing t h a t the r e l a t i o n of the a n t e r i o r l e n g t h of the s c a l e t o t h e l e n g t h o f the f i s h i s a changing one. The form of the curve has important r e l a -t i o n s t o the form of the growth curve f o r t h i s f i s h .." A curve s i m i l a r t o t h a t i n t h e p r e s e n t case f o r the t o t a l l e n g t h of the s c a l e i s a l s o shown. He f i n d s a s t r a i g h t l i n e r e l a t i o n a f t e r the f i r s t sharp r i s e up to 120 mm., i n which F r a s e r ' s f o r m u l a i s a p p l i c a b l e . D i f f e r e n t s p e c i e s of f i s h w i l l , without doubt, show d i f -f e r e n t growth c u r v e s , and t h e r e w i l l be v a r i a t i o n even w i t h i n _ 44 -t h e same s p e c i e s . F o r t h e P a c i f i c h e r r i n g , g r a p h s were made o f t h e f i r s t y e a r m a t e r i a l p r e v i o u s t o 193?> and f o r t h e 1939 m a t e r i a l , b e f o r e t h e d a t a were combined. I n a l l t h e s e c u r v e s t h e same g e n e r a l f o r m was f o u n d t o e z i s t . We, t h e r e f o r e , f e e l j u s t i f i e d i n r e g a r d i n g F i g u r e l j l as t h e c u r v e w h i c h d e -p i c t s t h e t r u e g r o w t h o f t h e a n t e r i o r f i e l d o f t h e s c a l e r e -l a t i v e t o t h e g r o w t h i n l e n g t h of t h e bod y . B e c a u s e o f t h e c h a n g i n g g r o w t h r a t e o f s c a l e i n r e l a t i o n t o body i t may be c o n c l u d e d t h a t n e i t h e r a s t r a i g h t l i n e r e l a t i o n o n l y , n o r a s t r a i g h t l i n e r e l a t i o n w i t h a c o r r e c t i o n f a c t o r , can be a p p l i e d i n c a l c u l a t i n g t h e g r o w t h o f t h e P a c i f i c h e r r i n g f r o m t h e s c a l e s . I t s h o u l d b e p o s s i b l e , h owever, h a v i n g t h e t r u e s c a l e l e n g t h / b o d y l e n g t h r e l a t i o n s h i p , t o c o n s t r u c t a movable g r a p h t o make su c h c a l c u l a t i o n s p o s s i b l e . - 45 SUMMARY AND CONCLUSIONS The i n v e s t i g a t i o n ' of-.the s c a l e g r o w t h / b o d y - g r o w t h r e l a -t i o n e n t a i l e d b o d y and s c a l e measurements o f some f i f t e e n h u n d r e d h e r r i n g i n t h e i r f i r s t y e a r , combined w i t h s i m i l a r d a t a f o r an a l m o s t e q u a l number o f f i s h I n t h e i r s e c o n d t o s i x t h y e a r s . F u r t h e r , t h e s c a l e s f r o m a l l o v e r t h e b ody o f an a d u l t h e r r i n g were examined f o r v a r i a t i o n i n s i z e , and s h a p e , and s k i n s f r o m a l a r g e number o f h e r r i n g f r o m .25-40 mm. were examined t o ' f i n d t h e t i m e and o r d e r o f a p p e a r a n c e o f t h e s c a l e s . • '. ' The r e s u l t s o f t h e v a r i o u s a s p e c t s o f t h e work a r e sum-m a r i z e d b e l o w . 1. T h e r e i s v e r y e x t e n s i v e v a r i a t i o n i n - s i z e a n d shape of t h e s c a l e s I n d i f f e r e n t p a r t s of t h e body. 2. The most r e g u l a r s h a p e d s c a l e s a r e f o u n d i n t h e a n -t e r i o r m e d i a n r e g i o n , t h a t i s , f r o m t h e f o u r t h t o t h e n i n t h l o n g i t u d i n a l r o w s , and a b o u t t h e e i g h t h t o t h e f i f t e e n t h o r t w e n t i e t h v e r t i c a l r o w s . .3. T h e r e a r e s e v e r a l e x a m p l e s o f s p e c i a l i z a t i o n i n shape o f t h e s c a l e w h i c h a r e c o n s t a n t i n a l l h e r r i n g . These a r e (a) b r a n c h i a l s c a l e , (b) v e n t r a l s c a l e , and ( c j k e e l e d s c a l e s . 4. The s i z e o f t h e s c a l e s I s g r e a t e s t j u s t p o s t e r i o r t o t h e h e a d , w i t h d e c r e a s e i n l e n g t h and w i d t h p o s t e r i o r l y and d o r s a l l y . - 4 6 -35.. The c u r v e f o r the (w) dimension shows a g r e a t e r maxi-mum than the curve f o r the ( l ) dimension, but both c u r v e s con-verge v e r y c l o s e l y at each end. 6 . S c a l e p a p i l l a e f i r s t f o r m i n the young h e r r i n g a t about 29 mm. 7. S c a l e s develop from t h r e e c e n t r e s of growth. The f i r s t i s on the l a t e r a l l i n e i n t h e t a i l r e g i o n , the second i s i n the a n t e r i o r median r e g i o n , and t h e t h i r d i s i n the a n t e r i o r v e n t r a l r e g i o n . 8 . Development spreads from t h e s e c e n t r e s more r a p i d l y d o r s a l l y and v e n t r a l l y than l o n g i t u d i n a l l y . 9. Squamation i s completed except f o r the d o r s a l rows i n f i s h of 356-358 -mm... 10. F i r s t formed s c a l e s have no a n t e r i o r f i e l d , but w i t h i n a'short time s t r i a e are formed and the a n t e r i o r f i e l d develops at a much f a s t e r r a t e than the e n t i r e s c a l e . -11. F o r the g r e a t e r p a r t of t h e body s c a l e s are formed one to each myotome. P o s t e r i o r l y the s c a l e s l i e intermyotom-i c a l l y , so t h a t i n g e n e r a l .there i s one l e s s s c a l e than myo-tome. 12. I n the body g r o w t h / s c a l e growth r e l a t i o n s h i p t h e growth r a t e of the a n t e r i o r f i e l d o f the s c a l e i s g r e a t e r than the body growth r a t e up to about 40-350 mm. Then i t decreases t o become n o t i c e a b l y l e s s than body growth r a t e . - 4 7 -13V The growth r a t e f o r the e n t i r e s c a l e i s more r a p i d than t h e body up t o .40-50 mm., then f o r t h e remainder, of the f i r s t year i s v e r y c l o s e to that of the body. 14. The growth r a t e of the a n t e r i o r f i e l d i n r e s p e c t t o the e n t i r e s c a l e w i t h i n c r e a s e i n body l e n g t h shows an i n c r e a s e from zero t o f i f t y percent b y a body l e n g t h of 40 mm. Then the r a t e i s v e r y l i t t l e g r e a t e r than t h a t of the e n t i r e s c a l e and the curve i s v e r y c l o s e to a s t r a i g h t l i n e . 15. The two d i f f e r e n t i a l s of growth r a t e f o r the a n t e r i o r f i e l d account f o r the g r e a t e r d i v e r g e n c e of the growth curve than i n the t o t a l s c a l e curve. 16. I n t h e a d u l t f i s h age-groups a r e found to form i n d i v -i d u a l curves l y i n g at an angle t o t h e g e n e r a l c u r v e . T h i s i s e x p l a i n e d on the b a s i s of f a s t - and slow-growing f i s h . 17. The shape, of the g e n e r a l curve f o r h e r r i n g up t o s i x years of age i s found to be " s i g m o i d " . With t h e s c a l e growing at a s l i g h t l y s lower r a t e t h a n t h e body up to the f o u r t h t o f i f t h year when i t i s . found t o have i n c r e a s e d up t o t h a t of the body. The curve drops -off a g a i n a t t h e end. 18. Because of the changing growth r a t e of growth of s c a l e i n r e s p e c t t o body a s t r a i g h t l i n e r e l a t i o n i s not ac-c e p t a b l e f o r the P a c i f i c h e r r i n g . The most convenient means of c a l c u l a t i n g the annual growth of the body from t h e growth of t h e s c a l e s w i l l be t o c o n s t r u c t a movable graph r a t h e r than t o attempt t o work out a f o r m u l a . - 48 -BIBLIOGRAPHY C r e a s e r , C.W. The S t r u c t u r e and Growth of the S c a l e s of F i s h e s i n R e l a t i o n t o the I n t e r p r e t a t i o n of T h e i r L i f e H i s t o r y , w i t h S p e c i a l Reference t o the S u n f i s h (Eupo-motis g i b b o s u s ) . M i s c . Pub. U n i v . Mich. Mus. Z o o l . No. 17, pp. 1-82, 1926. Da h l , Knut. The S c a l e s of t h e H e r r i n g as a Means o f Determin-i n g Age, Growth, and M i g r a t i o n . Rep. Nor. F i s h , and Mar. I n v e s t . V o l . I I , No. 6, P t . 2, 36 pp., 1907. The Assessment of Age and Growth i n F i s h . I n t . Rev. H y d r o b i o l . Hydrog. V o l . I I , Nos. 4 & 5, pp. 758-769, 1909. The Age and Growth of Salmon and Trout i n Norway as Shown by T h e i r S c a l e s . ( E n g l . Trans. ) Salmon and Trout Assoc., London. 141 pp., 1911. E l s o n , P.F. Order of Appearance of S c a l e s i n Speckled Trout. J our 11. F i s h . Res. Bd. Can. V o l . I V , No. 4, pp. 302-308, 1939-E s d a i l e , P.O. I n t e n s i v e Study of the S c a l e s of Three S p e c i -mens of Salmo s a l a r . Mem. and P r o c . Man. L i t . and P h i l . Soc. V o l . L V I , No. 3, 1912. F r a s e r , C. McLean. Growth of the S p r i n g Salmon. Trans. Pac. F i s h . Soc. pp. 29-39, 1915 (1916). On the S c a l e s of" the S p r i n g Salmon. Cont. Canad. B i o l . pp. 21-38, (1915-16) 1917. On t h e - L i f e H i s t o r y of the Coho or S i l v e r Salmon. Cont. Canad. B i o l . pp. 39-52, (1915-16) 1917. R e a r i n g Sockeye Salmon i n F r e s h Water. 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Frequency Curves i n H e r r i n g I n v e s t i g a t i o n . Rep. Nor. F i s h , and Mar. I n v e s t . V o l . I l l , No. 4 , pp. 1-27, 1924. Lee, Rosa M. An I n v e s t i g a t i o n i n t o the Methods of Growth De-t e r m i n a t i o n i n F i s h e s . Pub. de C i r c . No. 63, pp. 1-35, 1912. A Review o f the Methods of Age and Growth Determina-t i o n i n F i s h e s by Means of S c a l e s . M i n i s t r y of A g r i c . and F i s h . F i s h . . I n v e s t . S e r i e s I I , V o l . IV, No. 2, pp. 1-32, 1920. M a r e t t Tims, H.W.. The Development of S t r u c t u r e and Morphology of the S c a l e s i n Some T e l e o s t e a n F i s h . Quart. Journ. M i c r . S c i . V o l . XLIX, pp. 39-68, 1906. Meek, A. The S c a l e s of the H e r r i n g and T h e i r Value as an A i d t o I n v e s t i g a t i o n s . N o r t h . N..S.V. p. 11, Newcastle-upon-Tyne, 1916. *Molander, A.R. S t u d i e s i n the Growth of the H e r r i n g , Espec-i a l l y w i t h Regard t o t h e Examination of the S c a l e s f o r Determining i t s Growth. Ur Svenska H y d r o g r a f i s k -h i o l o g l ska Kommissionens S k r i f t e r . V I , Stockholm, 1918. *Monastuirsky, G.N. On the Technique of Determining the Rate of Growth of E i s h by Measurements of the S c a l e s . S i b e r -i a n I c h t h y . Lab. K rasnoyarsk. 1926. Neave, F e r r i s . The Development' of the S c a l e s of the Salmo. Trans. Roy. Soc. Can. 30 ( 5 ) , 1936.. Nusbaum, "T. M a t e r i a l i e n zur v e r g l e i c h e n d e n H i s t o l o g i e der Hautdeoke der ¥virbeltiere, i i i . Zur Kis t o g e n e s e der Lederhaut und der C y c l o i d - s c h u p p e n d i e r K n o c k e n f i s c h e . Anat. Anz. "50, 1907-. Paget, G.W.. Report on the S c a l e s of,Some T e l e o s t e a n F i s h w i t h S p e c i a l Reference t o T h e i r Method, o f Growth. M i n i -s t r y of A g r i e . and F i s h . , F i s h . I n v e s t . S e r i e s I I , ' Vol.'. IV, No. 3, pp. 1-24, 1920. P a r r o t t , A.W. The V a r i a b i l i t y and Growth, of the S c a l e s of the Brown Trout (Salmo. t r u t t a ) i n New Zealand. Trans, and Pr o c . N.Z. I n s t . No. 63, pp. 497-516, 1934. Regan, T. Caudal F i n of E l o p i d a e and Some Other T e l e o s t e a n F i s h e s . Ann. and Mag. Nat. H i s t . S e r i e s B, 1910. Savage, R.E. Report on Age D e t e r m i n a t i o n from S c a l e s of Young H e r r i n g , w i t h S p e c i a l Reference to the Use of P o l a r i z e d L i g h t . M i n i s t r y of A g r i c . and F i s h . , F i s h . I n v e s t . S e r i e s I I , V o l . IV, No. 1, 191.9. Setna, S.B.. Development of the Trout S c a l e . J o u r n . U n i v . Bombay ( B i o l . S c i e n c e s ) . No.' 2, P t . 5, 1934. S h e r r i f f , C a t h e r i n e V\T.M. H e r r i n g I n v e s t i g a t i o n s . Report on the M a t h e m a t i c a l A n a l y s i s of Random Samples of H e r r i n g s . W ith a n I n t r o d u c t o r y Note by P r o f . D'Arcy Thompson* F i s h . Bd. f o r S c o t l a n d , S c i e n t i f i c I n v e s t . No". 1, 1922. T e s t e r , A.L. The Length and Age Composition of the H e r r i n g (Clupea p a l l a s i - i T . i n the C o a s t a l Waters-of B r i t i s h Colum-b i a . J o u m . B i o l . Bd. Can. V o l . I ' l l , No. 2, 1937. Thompson, W.F. A C o n t r i b u t i o n t o t h e L i f e H i s t o r y of t h e P a c i -f i c H e r r i n g : I t s B e a r i n g on the C o n d i t i o n and Future o f the F i s h e r y . Rep. B.C. Comm. F i s h . 19l6 (1917), pp. 539-587, 1917.. -51 -Thomson, J". S t u a r t . The P e r i o d i c Growth of S c a l e s i n Gadidae and P l e u r o n e c t l d a e as an Index of Age. Jo u r n . Mar. B i o l . Assoc.. V o l . V I , No. 3, pp. 373-375, 1902. 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London. V o l . CXLI, pp. 643-702, I 8 5 I . WInge, O t t o . On the Value of the Rings i n t h e S c a l e s of the Cod as a Means of Age D e t e r m i n a t i o n . M e d d e l e l s e r f r a Kommissionen f o r Havunderso'geiser, s e r i e E i s k e r i . V o l . T V , No. .8',. 1915 • * I n f o r m a t i o n from Graham, 1929. 52 -fH 0 -P : f t o O ' o -P , 0 © -P • H 0 0 d a • H - H 0 © 0 0 © © 0 d . p - p o i - p to p -p -p cd © -rH - H - H - H «H ftdcddcdddd.H •H pi © P> © p-> P> P a 0 ^  . 0 d • H 0 ra © ra fn-0 c-© d 0 m-•p • H rH -P • H i •p ra • H © o- © a d .d*— d .d cd o o d ft cd 60 ft cd P s - H 0 >H - H © © -P • H © A r t |3 ft p5 P i P s - H r H -P 01 o © © © © d d d d • H • r l • H 0 © © © K) 03 ra ' 0 © © © ra ra ra ra fH u fn Fn 0 0 ?f PH PH PH PH © d © © m PH © • tiD^ d CM r-r\ON-<H/ K\ CM NO ON O ccS - p • • • • • • • • • fn W C— r H K\ ON C— ON ^- ^ © d KNLTW" "vh LT\ U\ K\-3" NO > 0 H ON C O H O O L f \ r H U \ O J " ^ N O LTN-vi- CM NO K\ONC—-<H/ ITWOVONG ON N\ f- K\ LT\OO K\LT\ C- LT\ TO\ c— H CO o • « « 9 O <=* c— NO K\ u\ CO ON o H H H H OJ CM O OJ CO rH C— ON ON K\ LT\ O CM C—-NO NO UN. 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A Y u r e b a y h a r b o u r u r e bay ' h a r b o u r L. 1931 1953 1935 1934 19J4 51 52 - - - -55 - - _ . . 54 ( 1) 0.08 -55 ( 7 ) 0.04 56 ( 9) 0.10 -57 (12) 0.09 -• 58 (11) 0.15 - - ( 1) 0.56 59 ( 9) 0.22 - ( 4 ) 0.51 40 ( 2) 0.54 - ( 4) 0.51 ( 1) 0.59 41 ( 1) 0.27 - ( 4) O.58 ( 5) O.56 42 - ( 7) 0.59 - ( 1) 0 .46 45 - ( 4) 0.48 ( 5) 0.61 44 - ( 1) O.70 ( 8) 0.46 ( 6) 0.68 45 - ( 1) 0.65 ( 6) 0.52 ( 7) 0.75 ( 1) 0.49 46 - ( 2) 0.74 ( 6), 0.46 ( 7) O.76 47 - ( 5) 0.66 - (12) 0.76 48 - ( 9) 0.68 ( 2) 0.70 ( 5) 0.78 49 ( 8) 0.69 ( 5) 0.68 (15) O.76 50 - ( 7) 0.70 ( 2) 0.72 (16) 0.79 51 - ( 6) 0.72 - {17) 0.81 52 - ( 2) 0.74 C l ) 0.75 ( 1 2 ) 0 . 8 5 5J - ( 5) O.75 - ( 9) O.89 54 - ( 2) 0.74 - ( 4) 0.86 ( 1) 0.80 55 ( D O.92 - ( 2) 0.88 56 • - ( 1) 0.88 - ( 5) 0.90 ( 1) 0.84 57- - ( 1) 0.75 - ( 1) O.87 ( 1) 1.02 58 - ( 4) O.87 - ( 1) 0.97 59 - ( 5) 1-02 -60 - - - ( 1) 1.00 61 - - - - ( 1) 0.95 62 - - - ( 1) 0.88 65 -64 -65 - ( 2) 1.1? - ( 1) 1.08 66 - - ( 1) 1.18 67 - -68 - - - - -69 - -70 71 - . _ - -72 73 - - -74 - - - - -75 - - _ 76 77 78 79 80 - - -8 1 ' - -82 - - -85 - . -84 85 - - -86 87 ' - - -88 -89 -90 - - -91 92 - - -93 - - _ 94 - - -95 -96 - - -97 -( D 1.47 100 - - I 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 T e l e g r a p h Hanoose H e i d tt„<„- m • h a r b o u r b a y i s l ^ d U ^ ° n ^ i n o o m a l i 1 9 3 4 «55 1955 193T ^ f 1 31 32 33 -54 _ | I ' " < ?) 0.18 57 ' 58 39 40 41 42' 6) 0.25 (• 7) 0.25 ( 1) 0.27 ? 0-29 ( 2.) 0.54 ( 4) 0.29 ( 4) 0.5? ••( i ) o.4o 3 0-39 ( 2) 0.46 ( 1) o.44 0.41 ( 1) 0.49 ( 4) 0*42 44 , _, v.-rj \ <±i  45 - ( 1) 0.44 ( 2) 0 . . 46 - ( 2) 0.39 -47 - ( 6') 0;56 48 ( 1) 0.72 ( 1) 0.54 49 ( 2) 0.54 •- -50 ( 1) 0.75 - -51 -52 ( 1) 0.67 ( 1) 0.81 53 ( 3) 0.78 - ( 1) 0.76 54 ( 2) 0.86 - •( 1) 0.78 55 ( 1) 0.85 -56 ( 2 ) 0.80 - ( 1) 1.01 57 ( 2) 0.84 { 1) 1.00 58 ( 2) 0.87 ( 1) O.89 -59 ( 2) 0.82 - - ( 1) 0.98 60 - - - ( 1) 1.0? 61 - - - -62 65 -64 - - - ( 1) 1.15 65 - - - ( 2) 1.15 66 - - - ( 2) 1.08 67 - ( 1) 1.22 68 -6? - - - -70 71 - - - - -72 - ' - - ( 1) 1.27 73 - - - - , . -74 - - - :' ' -75 -76 _ _ _ _ _ 77 78 - - -79 - -80 - • - - ( 1 ) 1.60 81 - -82 - - : - • - ( 2) 1.6? 83 - - - - ( 4) 1.63 84 - - - - ! 2) 1.55 85 - - - - ( 5) 1-97 86 - • - - , - ( 4} 1.95 87 - - - -88 - - - - ( 5) 1.72" 8? - - - - ( 3) 1.88-90 - - - - ( 4) 1.77 ?1 - - - - ( 2) 1.86. 92 - - - - ( 6) 2.57 ?5 - - - - ( 3) 1-99 ?4 - - - - ( 4) 2.25 95 - - - - ( 3) 2-33 ?6 - - - - ( 4) 1.97 97 - - - - ( 5) 1-94 98 - - - - ( 4) 1.93 99 - - - - ( 3) 1.82 100  2 2 27 101 _ - ( 7) ? OS 102 _ - ( fa) 00 103 _ - ( 1! 2 11 104 _ - ( •'.) 2 22 l o j - ( 2) 106 - - ( 1) 2 107 _ - i 2) 2 lot; - ( 3) 0 27 10? _ - ( 1) n JO 110 - ( 1) 2 i n -- - -112 113 114 115 116 117 118 119 120 121 122 Tabl© 2 (a) 57 -L. D e p a r t -u r e bay 1938 h a r b o u r 1938 M i l l b a y 1938 D e p a r t -u r e bay 1939 F u l f o r d h a r b o u r 1939 31 32 33 34 35 36 37, 38 . 39 40 41 42 43 44 4I 46 47 48 4? 50 51 52 53 54 55 56 57 58 ^ 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 7 ^ 76 77 78 79 80 81 82 83 84 85 86 87 ( 9) 0 . 0 6 (12) 0 . 0 9 ( 6) 0 . 1 3 ( 3) 0 . 9 5 1) 1.14 1.12 1.17 1 . 1 5 1.14 1) 3) 5) 5) 6) 1 .21 2) 1 .16 3) 1 .15 1) 1 . 3 4 1 ) 1 .47 1 ) 1 .38 1) 1 .49 1 .43 1 .45 1-51 3) 1-51 2) 1 . 6 1 1) 1-57 1) 1-53 3) 1 .68 2) 1 .60 3) I . 6 3 ( 2) 0 . 1 7 ( 4) 0 . 1 6 ( 6) 0 . 2 1 (10) Q.26 ( 5) 0 . 3 2 ( 8) 0 . 3 4 ( 5) 0 . 3 8 ( 2) 0 . 4 6 ( 2) 0 . 4 3 ( 1 ) 0 . 4 4 1 ) 0 . 5 4 ( 5) 0 .67 ( 4) 0 . 6 6 ( 8) 0 . 6 7 ( 8) 0 . 7 3 ( 3) O .69 ( 2) 0 . 7 4 ( 2) 0 . 8 6 ( 1) 0 . ? 4 ( 1 ) O .85 ( 1 ) O .99 ( 2) 0 . 0 0 ( 9) 0 . 0 9 ( 6) 0 . 1 2 ( 7) 0 . 1 6 ( 9) 0.24 ( 7) 0 . 2 3 ( 5) 0 . 3 0 ( 5) O .36 (14) 0 . 3 9 (16) 0 . 4 5 ( 9) 0 . 4 8 ( 7) 0 . 4 8 ( 3) O .52 ( 1) O .56 ( 1) 0 . 2 8 ( 1) 0.14 ( 2) O .36 ( 1) O .33 ( 2) 0.44 ( 1) O .58 ( 4) ( -3) ( 4) ( 3) ( 1 ) ( 1) ( 1) ( 1) ( 1) O .65 0 . 6 6 0 . 5 7 0 . 8 0 0 . 8 0 0 . 7 0 0.79 0 . 8 2 0 . 8 3 0 . 7 6 O/85 0.?4 1 . 0 0 1.14 0 . 9 7 0 . 9 5 1 .05 ( 1) 1 .08 ( 1 ) 1 .46 ( 1 ) 1 .74 91 92 93 94 95 96 97 ( 1 ) 1. 99 100 101 102 1 0 J 104 103 10 6-107 108 10? 110 an 112 113 114 115 116 117 118 119 •120 121 122 Table 2 (a) 5? -8? 90 91 92 93 94 93 96 97 B i d w e l l A c t i v e P o r l i e r L a d y s m i t h bay p a s s p a s s h a r b o u r Deep -o, i«>ba p a s s h a r b o u r bay L. 1939 1939 1939 1939 1939 31 - - - - -32 33 - - - - (13) 0.04 34 - - - - (12) 0 . 1 0 33 - (16) 0.14 36 - ( 2) 0 . 3 7 - (21) 0 . 1 8 37 - (27) 0 . 2 3 38 - - ( 3) 0 . 3 4 - (20) 0.24 39 (10) 0.41 - (11) 0 . 3 3 40 - - ( 9) 0.42 - ( 7) O . 3 4 41 - (11) 0 . 5 0 - ( 1 ) O .39 42 - - ( 9) 0 . 5 3 - ( 2) 0-41 43 - - (11) 0 . 5 2 - ( 1 ) 0.48 44 ( 1) 0 . 5 7 (10) 0 . 5 5 45 - - (11) 0 . 3 7 46 (10) 0 . 5 9 47 ( 3) 0 . 6 3 (10) 0 . 6 4 48 - ( 1) 0 . 6 8 (10) 0 . 6 8 49 ( 3 ) 0 . 6 8 (10) O .69 50 - - (10) 0 . 7 3 51 - ( 1) 0 . 7 1 ( 9) 0 .77 32 - ( 6) 0 . 7 9 ( 7) O .76 53 - ( 2) O .78 ( 7) 0.82 54 - ( 7) 0 . 8 6 ( 6) 0 . 8 5 55 - (11) 0 . 8 7 ( 6) 0 . 8 7 56 ( 8) 0 . 9 0 -57 - ( 1) 0 . 9 8 -58 ( 5) 1-06 ( 8) 0 . 9 4 ( 1) 1.04 59 - ( 5) 1 . 0 1 60 ( 2) 1 . 0 8 -61 ( 1 ) 1 . 0 1 ( 6) 1 . 0 1 ( 2) 1 . 12 62 ( 1) 0 . 8 7 ( 5) 1-12 - - -65 - ( 5) 1 .13 . . . 64 ( 2) 1 .07 ( 2) 1 .08 -65 ( 3 ) 1 .09 66 ( 2) 1.07 -67 ( 1 ) 1 . 1 3 68 ( 2) 1 .17 -69 ( 3 ) 1 . 13 - - - -70 ( 3) 1-21 -71 ( 3 ) 1 - 1 9 - ( 1) 1 .38 72 ( 2) 1 .22 - -73 ( 2) 1 .11 74 ( 1) 1.40 2) 1 .52 In ol - - ( 6) 1 . 5 2 Vf\ ? - - ( 5) 1 .58 81 ( 1) 1 . 5 4 - . { I] 84 1.45 1.41 ( 6) 1 .45 ( 4) 1 .60 85 86 ( 1) 1 .55 . - ( 4) 87 ( 1) 1 .72 • ' ( 5) 1-57 7) 1 .56 100 ( i ) 2 1.53 ( 7) 1.68 1) 1.75 - - ( 4) 1 . 7 6 1 1 _ - - ( 6) 1 .6? - ( 6) 1.70 - ( 7) 1 .69 - I 3) l . g o - ( 5) 1 .88 ( 4) 1 . 8 6 - ( 4) l . ? l - ( 3) 1-93 - ( 7) 2-00 ( 1) 2.14 - - ( 6) 1 . 8 8 K 1 1 ( 8) 1 .95 Table 2 (a) - 60 -101 102 10? 104 105 106 107 108 109 110 111 -112 11? 114 115 ( 1) 2 .48 116 117 118 11? ( 1) 2 . 5 4 120 121 122 ( 1 ) 2 .62 ( 4) ( 1) ( 1 ) ( 5) ( 2) ( 1) 2) 2 . 16 2 . 0 9 1 .99 1 .97 2 . 1 2 2 . 1 0 2.49 ( 5) 2 . 1 9 .( 1) 2.65 ( 1) 3-17 Table 2 (b) - 61 -D e p a r t -u r e bay 1951 D e p a r t -u r e b a y 1953 F u l f o r d h a r b o u r 1935 D e p a r t -u r e b a y 1 9 M F u l f o r d h a r b o u r 1954 31 - _ 32 -53 - _ 34 ( 1) 0.43 _ ?5 ( 7) 0.-38 _ 36.. ( 9) 0.47 _ 57 (12) 0.50 _ 38 (11) 0.56 39 ( 9) 0.64 _ 40 ( 2) 0.81 _ 41 { 1) O.67 _ _ 42 _ _ 45 - _ 44 - < 1) 45 - 1 1 ) 1.16 46 - < 2) 1.37 47 - ( 5) 1.25 48 - ( 9) 1.28 . 49 - ( 8) 1.32 50 - ( 7) 1.35 31 - ( 6) 1.37 52 - ( 2) 1.37 53 - ( 3) 1.34 54 - ( 2) 1.42 55 ( 1) I . 6 5 56 - ( 1) 1.31 57 - ( 1) 1.34 58 - ( 4) 1.33 I9 - ( 3) 1.77 6o _ 61 _ 62 _ 65 _ _ 64 _ & - ( 2) 1.87 66 _ 67 - -DO 69 _ 70 - _ 71 - _ 72 - _ 75 - . -74 -75 _ _ 76 - _ 77 _ _ 78 -79 80 _ 81 _ 82 _ _ 85 _ 84 _• _ 85 fi£ - -oo 87 8a..... -89 90 - -91 - _ 92 - _ 95 - _ 94 -95 - _ 96 _ „ - -7° 99 _ _ 100 - _ 4) 0.83 4) O.76 ( 4) O.89 ( 7) O.95 ( I . 0 5 1.01 1.06 6) 1.00 ( 2) 2) 1.50 3) 1.28 1.50 1) 1.36 ( 1) 1.08 { 1) 0.81 ( 5) 1.12 3) 1.21 " 1-31 1-35 1.42 1.43 6) 7) 7) (12) ( 5) 1.38 (15) 1.36 (16) 1.57 (17) (12) ( 9) 1.40 1.44 1.49 ( 4) 1.45 1.49 1.55 1.52 1) 1.71 ( 1) 1.47 1) 1. ( 1) 0. ( 1) 0.82 1) 1.46 ( 1) ( 1). 1.48 1.69 1) 1.71 1) 1.68 1) 1. 1) 2.51 Table 2 (b) 101 102 1 0 J 104 105 106 107 108 109 110 111 112 115 114 115 116 117 118 119 120 121 122 L- 1 9 2 4 l? 3 5 1935 193& 193T 51 52 - _ 33 • - i ~ -54 - . " - • -| | - ( 2) 0.36 - " f£ ' - ( 6) 0.62 _ - " : 12 " ! 7! ( 1) 0.77 - • 39 - ( 4) 0.79 -j ? - - 4*) o-9o ( i ) o.8o : 41 42 fA - < 1) i.ol • ( 3) s: ^ - •• i.) i - o i ( 4) -3) 0.E 2) 0.93 ( 1) 0. 0.95 :i - \ -LI u-oy I 2) 0.97 In " - ( 2) 1.04 4 ( 1 ) 1 . 1 5 " ( D l . l i ((>)1-1l 49 ( 2) 1.21 _ 50 ( 1) 1.34 51 -52 ( 1) 1.32 - ( 1) 1.52 - • 53 • ( 3) 1-42 - C l ) 1.36 - • 54 ( 2) 1.57 - ( D 1.49 55 ( D I . 3 8 - - - -56 ( 2) 1.46 - ( D 1.73 57 ( 2) 1.40 ( 1) 1.77 - - -58 ( 2) 1.45 ( 1) 1 .59 - -59 ( 2) 1.50 - - ( 1) 1.89 60 - - C D 2.05 61 - - -62 - - -63 - - - - -64 - - - ( 1) 2.10 65 - - - ( 2) 1.96 66 - - - { 2) 1.94 67 - - - ( 1) 2.69 68 • - - - -69 -70 - - - - -71 - - - - . -72 - . - - - C D 2.05 • • 73 - - -74 - - - - -75 -76 - - - - • -77 - - - - ' -78 . - - -80 - - - - ( D 2-52' 81 - - - -82 - - - ( 2) 2-80 83 - - - - ( 4)' 2.62 84 - - - - ( 2) 2.49 85 - - - - ( 5) 3-05 86 - - - 4) 3.-27 87 - - - - -88 - - - i 3) 2-69 89 - - - - ( 3) 2.91 90 - ' - - ( 4) 2.70 91 _ - - - ( 2) 2.79, 92 - - - - ( 6.) 3.65 93 - - - - ( 3) 3.02 94 - - - - ( 4) 3-53' 95 - - - - ( 3 ) 3-84 96 - - - - ( 4) 2.98 97 - - - - • • ( • ? ) 3-10 98 - - ( 4). 3.04 99 - - - - ( 3) 2-95 100 - - - ( 2) 3.44 101 102 - - ( 7) 3-18 1 0 J _ - - { 6) - x ! 3 . 2 6 104 3 - 3 1 103 • - 4 3 . 4 3 i o 6 - 2 2.91 107 3 . 3 2 108 - - 2 4 . 6 3 10? - " : 3 3.41 110 - ( 1) .3.67 i n - •• - (• 1 ) 3 . 8 2 112 -113 - - -114 -115 -116 117 -118 -119 - • I -120 -121 -D e p a r t - Ganges. M i l l D e p a r t - F u l f o r d u r e b a y h a r b o u r b a y u r e b a y h a r b o u r L . 1958 1938 1938 ,1959 1939 3 1 - - ( 2) O .33 32 - ( 2) 0 . 6 0 - f 9) 0 . 4 6 33 ( 4) 0 . 5 2 - ( 6) 0 . 5 2 -3 4 ( 9) 0.42 ( 6) O .57 - ( 7 ) O .56 35 (12) 0 . 4 5 (10) O .72 - ( 9 ) O .67 36 ( 6) 0 . 5 7 (' 5) 0 . 7 4 , - ( 7) O .67 37 - ( 8) 0 . 8 1 - ( 5) 0 . 7 8 38 - ( 5) 0.'82 - ( 5) 0 . 8 4 39 - ( 2) 0 . 9 9 - (14) 0 . 9 0 ( 1) 0 . 7 0 40 . ' - ( 2) O .92 - (16) O .93 41 - :"( 1 ) 0.92 - ( 9) 1-02 ( 1) 0 . 4 8 42 - - - ( 7) 1-02 ( 2) 0 . 8 4 43 - - - ( 3) 1.04 ( 1 ) 0 . 8 2 . 44 - - ( 1) 1.17 ( 2) 0 . 9 9 ' 43 ( 1 ) O .54 - -46 - - - - ( 1 ) 1.17 47 - - -48 , - - - ( 1) 1 .21 49 - - - - ( 2) 1 .29 50 - . - ( 5) 1-24 - ( 1 ) O .93 51 - - ( 4) 1 .26 - ( l ) 1 .45 52 - - ( 8) 1 .26 - ( 1 ) 1-.51 53 - - ( 8) 1 .38 - ( 4) 1 .33 3 4 - - ( 3) I . 3 0 - ( 2) 1 .45 35 - - ( 2) 1 .38 - ( 8) 1 . 5 2 56 - - ( 2) 1 . 5 5 - ( 4) 1 .55 57 - - ( 1 ) 1 .59 - (, 5) 1-41 58 - - - ( 4) I . 5 6 59 - - ( 1) 1 .58 - ( 5) 1 .67 60 ( 5) 1 .89 - ( 1 ) 1 .68 - ( 1) 1 .77 61 - - - - ( 1) 1 . 8 5 62 ( 1 ) 1 .19 - - ( 1) 1 .69 65 ( 1 ) 2 . 0 2 - - - ( 1) 1 .78 64 ( 3) 2 . 0 9 - - - ( 1 ) 1 . 9 8 65 ( 5) 2 . 0 7 - - -66 ( 5) 2-10 - - - -67 ( 6) 2 . 1 0 - - - -68 ( 2) 2 . 0 2 - - - ( 1) 1 . 9 3 69 ( 5) 2 .12 . -70 ( 1). 2.18 - -71 - - - - ( 1 ) - 2 - 4 8 72 ( 1 ) 2 . 3 6 . - - . - ' -73 ( 1 ) 2 . 3 8 -74 - - - - -75 ( 1) 2-54 - - - -76 ( 2) 2 . 5 8 - • - - -77 ( 2) 2.42 - - - . -78 ( 2) 2 . 5 0 - - - - . 79 ( 5) 2-50 - - - -80 - . _ 81 ( 2) 2 . 5 1 - -82 ( 1 ) 2 .55 - - -85 ( 1 ) 2 .47 . - - -84 ( 5) 2 . 7 1 - - - -85 ( 2) 2 .57 - - -86 ( 3) 2 . 5 1 -87 -88 — — — _ _ 89 i n 2 .78 - - - -90 - - -91 - - . - -92 - - -95 - - - - -94 - -95 - - - -96 ( 1) 2 . 9 7 - - -98 - - - -99 - - -100 THUTi 101 102 103 104 105 106 107 108 109 110 111 112 11? 114 115 116 117 118 119 120 121 122 B i d n e l l A c t i v e P o r l i e r L a d y s m i t h Deep bay p a s s p a s s h a r b o u r boy L- 193? 1939 1939 1939 1939 31 -32 - - - - -33 - (13) 0.45 34 - (12) 0 . 5 3 35 - (16) 0.57 36 - ( 2) 0 . 6 2 - (21) 0 . 6 3 37 - - - (27) O .69 38 - ( 3) O .82 - (20) 0 . 7 0 39 (10) 0 . 9 2 - ( I D 0.82 40 - - ( 9) O .96 - ( 7) 0.82 41 - - (11) 1.04 - ( 1) 1 . 0 3 42 - ( 9) 1 . 1 0 - ( 2) 0.95 43 - - (11) 1 . 0 9 - ( 1) 0.97 44 - ( 1 ) 1 .18 (10) 1 .13 45 - - (11) 1 . 1 7 46 (10) 1.19 47 - ( 3) 1 - 2 8 (10) 1.27 48 - ( 1) 1 . 2 5 (10) 1 . 3 0 49 - ( 3) 1-31 (10) 1.34 50 (10) 1.37 51 ( 1) 1-45 ( 9) 1-43 52 - ( 6) 1.45 ( 7) 1.46 53 - t 2) 1.51 ( 7) 1 - 5 1 54 - ( 7) 1-59 ( 6) 1.59 55 - ( I D 1-61 ( 6) 1 .56 56 - ( 8) 1 .63 -57 - ( 1) 1.68 58 ( 3) 1 . 8 3 ( 8) I . 6 9 ( 1) 1-75 59 - ( 5) 1-75 -60 - ( 2) 1.83 -61 ( 1) 1 . 7 0 ( 6) 1 . 8 0 ( 2) 1 . 8 1 62 ( 1) 1 . 6 3 ( 3) 1 - 9 6 -63 - ( 5) 1 - 9 2 -64 ( 2) 1 . 8 5 ( 2)- 1.88 -65 ( 3) 1 . 8 0 66 ( 2) 1 . 8 1 -67 ( 1) 1 .91 - - - _ 68 ( 2) 1 .95 -69 ( 3) 1 -90 -70 ( 3) 1 .97 -71 ( 3) 1.98 72 ( 2) 2 . 0 5 75 ( 2) 1 . 9 1 74 75 ( 1) 2.15 76 ( 1) 2 . 2 5 77 ( 2) 2.24 78 ( 1) 2 .32 79 ( 2) 2.33 80 ( 2) 2 . 5 0 81 ( 1) 2 . 3 1 82 -63 64 -85 - - -86 ( 1) 2 .45 87 ( 1 ) 2 . 8 0 B9 90 91 ( 1) 2 . 7 6 - _ f 93 94 9 ? 96 97 96 ( 1) 3 . 3 4 99 100 ( 1) 5 . 0 7 1) 2 . 4 3 1) 2 .36 2) 2 . 6 0 5) 2 .27 6) 2 .37 5) 2-34 7) 2 .37 5) 2 .46 6) 2 . 5 2 4) 2.40 6) 2-51 4) 2.66 5) 2 .61 7) 2 . 5 9 4) 2.72 7) 2.78 4) 2.87 6) 2 . 6 9 6) 2 . 8 3 7) 2 .61 5) 2 . 8 7 5) 3 . 0 1 4) 2 .97 4) 3 . 0 5 5) 3 . 0 2 7) 3-14 6) 2 . 9 6 8) 3 . 0 9 101 102 10? 104 105 106 i b 7 108 10° no i n _ 112 115 114 115 ( 1 ) 5-67 l l o 117 118 1 1 ° ' •(•• 1) 5.8° 120 121 122 ( l j 4.15 2) 4) 1) 1) 5) 2) 1) 3-29 5-54 5-55 5.1? 5.26 •36 • 78 5) 5. 97 59 - 69 -d o •H -P Crf i> u © co © Pi Pi © cd •«d a . . d to CD U <H I d © u 0 m erf 0 to © H PH 00 CD , & U -H O -P -d <H -P O PI tt! CD , d H +3 tiD © d tiD © Cd H Pi © tO Crf O •H Pi cd E> crf •P Pi crf o P q-i d o © H cd EH H I I rH" H i © •H EH © > •H +3 cd t> Pi © CO © Pi PH o 15 - p •H H cd o o H i © - p cd p o o CM -=4-^O rH co CM « 9 « e e © O rH o r-i H H 1 ! -f l • 1 S o tr- o o K \ NO rH v O « « © • e e ON HN NO v O c— CA CM K \ K \ KN K \ CQ CO CO CO CO CO ">> !>. l » !>> !>» cd Crf crf cd cd crf - d - d •d - d - d •d CM CM CM NO ON ON Pi •H rH Crf Pi O o crf © Pi d •P Pi cd P i © ON K\ ON rH CO H d o • d o o rH < o crf rP © -P Pi crf PH © P O s K\ ON H CM © Pi d CO cd - d CM 0 NO CO CO Pi cd © d •H rH cd o PR H O xi o o H O O ON CO ON CM O Pi d crf o Pi © © © cd Pj > . d d o O -P o O CO Pi © crf -P Pi o O d © o o cd P f p pq & ON ON ON ON K"\ K \ K \ K \ ON ON ON ON H H r-i H «% CO CO ON CM CM c\J CM © © © © Pi Pi Pi d d d CO ON CO Pi cd © Pi d •H H rH •H rH H O o rH O cd , d Crf 1^ j3 o o o H o o H o O rH H o rH PR < <4 FR < o O KN O NO C - H ON CM C— CO H ON CO H 0 ® • e « o t-C"\ CO CO o NO c-- CO K N KN KN CO ON o o CM r-i © crf si Crf o to CO •H cd © rH P i a cd d © o i> Pi o •H Crf d +3 P i •H O © Pi < P EH ON uc\ NO K \ K \ K \ ON ON ON H rH H », ON rH rH CM H rH © -P e d P i s> P © o Ho CQ 12! Table 4 - 7 0 -Data on the d i f f e r e n t i a l growth of the l e n p t h of t b e a n t e r i o r f i e l d ( 1 ) r e l a t i v e t o t h e t o t a l l S g t S ( t ) of the s c a l e w i t h ' i n c r e a s e i n l e n g t h ( L ) . 3 1 3 2 3 3 3 4 3 5 36 3 7 3 8 3 9 40 41 42 4 3 44 4 5 4 6 4 7 4 8 4 ? 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 ' 5 * 5 9 6 o 6 l 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 6 No. 1 t 2 0 . 0 0 0 . 3 3 1 1 0 . 1 0 0 . 4 8 2 3 o . o 8 . 0 . 4 7 3 5 0 . 1 2 0 . 5 1 5 6 0 . 1 6 O . 5 6 5 6 0 . 1 9 0 . 6 2 6 o 0 . 2 2 O . 6 7 5 6 0 . 2 6 0 . 7 1 5 5 0 . 3 4 O . 8 3 4 6 0.40 O . 8 9 3 4 0 . 4 5 O . 9 8 3 1 0 . 4 6 1 . 0 0 2 7 0 . 5 0 1 . 0 5 3 4 0 . 5 3 1 . 1 1 3 0 0 . 5 8 1 . 1 3 2 8 0 . 6 0 1 . 2 1 3 6 0 . 6 7 1 . 3 0 3 0 0 . 6 9 1 . 2 9 4 3 0 . 7 0 1 . 3 3 42 0 . 7 4 1 . 3 4 3 8 0 . 7 7 1 . 3 9 3 9 0 . 7 7 1.40 3 7 0 . 7 9 1 . 4 3 2 8 0 . 8 2 1 . 4 6 3 1 0 . 8 5 1 . 5 5 2 2 0 . 8 8 1 . 5 7 1 1 0 . 8 7 1 . 5 1 24 0 . 9 2 1 . 6 4 1 5 0 . 9 7 1 . 7 0 9 1 , 0 1 1 . 8 4 1 1 1.04 1 . 7 9 7 1 . 0 3 I . 6 9 7 1 . 0 3 1 . 9 1 9 1 . 1 1 I . 9 8 1 3 1.14 1 . 9 5 1 0 1 . 1 2 1 . 9 9 8 1 . 2 0 2 . 1 5 1 . 1 5 1 . 9 7 6 1.14 2 . 0 1 4 1.24 2 . 0 2 5 1 . 2 8 2 . 1 7 4 1 . 2 9 2 . 1 3 4 1 . 2 5 2.14 2 1 . 5 2 2 . 6 0 7 1 . 3 2 2 . 2 9 1 0 1 . 3 7 2 . 3 5 1 0 0 l / t 0 0 . 0 2 0 . 8 1 7 . 0 2 3 . 5 2 8 , 6 3 0 . 7 3 2 . 8 3 6 . 6 41.0 4 5 . 0 4 5 . 9 4 6 . 0 4 7 . 6 4 7 . 7 5 1 . 3 4 9 . 6 5 1 . 5 5 3 . 5 5 2 . 6 5 5 . 2 5 5 . 4 5 5 . 0 5 5 . 2 5 6 . 2 5 4 . 8 5 6 . 0 5 7 . 6 5 6 . 1 5 7 . 1 5 4 . 9 5 8 . 1 6 1 . 0 5 4 . 0 5 6 . 1 5 8 . 5 5 6 . 3 5 5 . 8 5 8 . 4 5 6 . 7 6 1 . 7 5 9 . 0 6 0 . 6 5 8 . 4 5 8 . 5 5 7 . 7 5 8 . 3 L No. 1 7 7 9 1.41 7 8 1 0 1 . 4 8 7 9 1 0 1 . 4 6 8 0 9 1 . 5 0 8 1 7 1 . 5 0 8 2 9 1 . 5 2 8 3 9 1 . 6 1 8 4 1 0 1 . 6 0 8 5 14 1 . 7 1 8 6 1 2 1 . 7 0 8 7 8 I . 6 9 8 8 8 1 . 7 4 8 9 1 0 1 . 7 5 9 0 1 0 1 . 7 3 9 1 9 1 . 7 3 9 2 1 1 2 . 1 1 9 3 8 . 1 . 9 2 9 4 8 2 . 0 5 9 5 7 2 . 0 9 9 6 8 1 . 9 2 9 7 1 2 I . 9 8 9 8 1 1 1 . 9 2 9 9 1 1 1 . 9 1 1 0 0 3 2 . 1 8 1 0 1 9 2 . 1 0 1 0 2 1 0 2.04 1 0 3 2 2 . 0 5 104 5 2 . 1 7 1 0 5 5 2 . 1 6 1 0 6 3 2.14 1 0 7 3 2 . 8 2 1 0 8 3 2 . 2 7 1 0 9 1 2 . 3 0 1 1 0 4 2 . 2 7 i l l - _ 1 1 2 — „ 1 1 3 — 114 _ — 1 1 5 1 2 . 4 8 1 1 6 — _ 1 1 7 1 2 . 6 5 1 1 8 1 3 . 1 7 1 1 9 1 2 . 3 4 1 2 0 _ _ 1 2 1 — 1 2 2 . 1 2 . 6 2 t 1 0 0 l / t 2 . 3 4 2 . 3 9 2 . 4 5 2 . 5 2 2.42 2 . 5 8 2 . 6 2 2 . 6 2 2 . 7 5 2 . 8 3 2 . 7 8 2 . 7 9 2 . 7 6 2 . 7 8 2 . 8 0 3 . 2 9 3 . 0 1 3 . 2 5 3 . 3 9 2 . 9 9 3 . 1 2 3 . 0 3 3 . 0 5 3 . 2 5 3 . 2 0 3 . 2 9 3-42 3 . 3 8 3.14 3 . 3 5 4 . 3 5 3.41 3 . 6 7 3 . 5 5 6 0 . 3 6 2 . 0 5 9 . 6 5 9 . 6 6 2 . 0 5 8 . 9 6 1 . 5 6 1 . 1 6 2 . 2 6 0 . I 6 0 . 8 6 2 . 4 6 3 . 4 6 2 . 2 6 1 . 8 6 4 . 2 6 3 . 4 6 3 . I 6 1 . 7 6 4 . 2 6 3 . 5 6 3 . 4 6 2 . 6 6 7 . I 6 5 . 6 6 2 . 0 6 0 . 0 6 4 . 2 6 8 . 8 6 3 . 8 6 4 . 8 6 6 . 6 6 2 . 7 6 4 . 0 3 . 6 7 6 7 . 6 3 - 9 7 3 . 5 9 3 . 8 9 6 6 . 8 8 8 . 2 6 0 . 2 4 . 1 5 6 3 . 2 - 71 -H CD cd fn • H cd o f n cd CD -p a cd o O 0 . d 0 EH •p U CO O fn cd EJO 0 d f>> • H fH U d © cd CO fH © o cd > — © fH © 0 H <tH CO <H O - r i CO f t f <iH fH O O A U - P cd CJD-H © a H P H <d O O 9 © — • C O © r l H f n — -p xi • H - p A fH co p q <JH • H . o <M d fH U 0 o . d -p +3 O t)D taD ra d 0 <H H O CO P i p o fH P i • H O cd o -p J=I co © cd fH © cd o i> co cd © cd fH © • 0 ft ft 0 tiD cd fH © © H -d • H © cd fH © < J fH © 8 H " o 5 3 o o 121 K N UN « O N O r H K N K N UN • O U N rH N O N O H • K N U N * O H K N ^ - N O K N C — C — C O • © # ^ ^ OO N O CM CM H CM K N K N CM O K N K N UN. N O CM CM O N O N O N H H H O O O N U N * * N O N O O O • * « t K N C - T H C O K N a 9 c— K N CO O N CM K N U"\ © H K N K N K N r H 1 I 1 ft o H O J ^ -• P K N K N K N w CO O N O N O N H H H H K N ! O K N O N H KN ON r H O N U N H C " - N O CM C O U N * * C— tf\ CM N O * * U N LT\ C O C O C O C O H H H H O O N O O N U N K N C — U \ H K N C - O N O CM CM K N K N ** « U N N O K N N O c-~c—c— H H H H O O O N O N U \ « * \ J 3 LT \ © W cd fn © cd © - P • H © PI u \ u \ u \ ir\ N O N O H • B 0 U\ LT\ L f \ o o o CM CM CM K N * * O N U N U N * * U N C — C M O N O O e » © ** i i M f \ ** ** ** CM ONt U N O CM N O U N r H H O • U N U \ O N * * K N • • • • C T - O - N O c—• O N O N O N O N H H H H CM U N C O ur\ U \ L T \ * * u\ H N O C O CM O N C — C — O O C — N O O N C O C — C O C O O O C O C O H H H H K N ' * O N N O U \ U \ * * i r \ H © cd fH © CM K N UN, J> K N K N K N Cd I I I H CM " * >d K N K N K N © O N O N O N +3 H H H ,d •QD • H 2 s: d 1=3 O H H * * U N U \ • • • UN U~\ Lf\ C O K N U \ • « • * * K N C O H H H CM CM CM O N CM CM CM K N H C - H O CM * * U N • • • • U N U N U N O H O O • • » o o o CM CM CM O K N O J K N K N H H H N O C~~NO N O • • # • U N U N ^ U N U N N O • * o « • e U N C ~ C O K N H H CM CM CM CM CM CM K N O O O CM H K N O N H N O K N U N U \ U N • » 9 • U N U N U N U N K N CM O H • e • 9 H C— O H H H CM CM CM CM U N . H H CM CM H K N O N C - U N H N O O N I—1 C M H K N K N K N • * « • • • « • "Nt U N ^ U N U N U \ U N Hco N O O O K N O N H • • « e « e © O N C O U N c-- K N N O U N O N O N O N O N O O O H H H H CM CM CM O N C M CM K N O O O CM K N H C — CM H K N CM K N UN, K N K N K N I I I H CM *=d-K N K N K N O N O N O N H H H © W cd fn © cd © , d w • H © K N K N I CM K N O N © cd fH © cd © •a • H © 1= © H H H H H i> H H - 7 2 -EXPLANATION OF PLATES AND FIGURES P l a t e I F i g u r e 1. T y p i c a l s c a l e o f a h e r r i n g i n the f i r s t y e a r . F i g u r e 2 . R e l a t i v e s i z e o f t h e a n t e r i o r f i e l d o f s c a l e s f r o m a f i s h o f 3 7 mm. P l a t e s I I , I I I , and I V Drawings o f s c a l e s t a k e n i n l o n g i t u d i n a l s e r i e s t o show the v a r i a t i o n i n shape and s i z e . P l a t e V F i g u r e 3 . Graphs i l l u s t r a t i n g t h e change o f s i z e i n t h e t h i r d , f i f t h , s i x t h , s e v e n t h , e i g h t h and t e n t h rows o f s c a l e s . P l a t e V I F i g u r e s 4, 3 , and- 6 . Diagrams o f t h e s k i n o f young h e r r i n g t o show t h e d e v e l o p m e n t a l a r r a n g e -ment o f the s c a l e s . P l a t e V I I F i g u r e 7 . Smoothed i n d i v i d u a l c u r v e s showing the r e l a -t i o n s h i p between l e n g t h of s c a l e ( l ) and l e n g t h of f i s h (L) f o r a l l samples o f f i s h i n t h e i r f i r s t y e a r . - 7 3 -P l a t e V I I I F i g u r e 8 . Smoothed curve showing t h e g e n e r a l r e l a -t i o n s h i p between l e n g t h s of s c a l e (1) and ( t ) and l e n g t h of f i s h (L) f o r f i s h i n t h e i r f i r s t y e a r . P l a t e I X F i g u r e 9. Curve i l l u s t r a t i n g t h e d i f f e r e n t i a l growth. of (1) i n r e s p e c t t o ( t ) w i t h i n c r e a s e i n l e n g t h of f i s h ( L ) . P l a t e X F i g u r e 10. Curves showing the r e l a t i o n s h i p between l e n g t h o f s c a l e (1) and l e n g t h of f i s h (L) a t ages I I to V I . P l a t e X I F i g u r e 11. G e n e r a l c u r v e showing the average r e l a t i o n -s h i p between l e n g t h of s c a l e (1) and l e n g t h o f f i s h ( L ) . P L A T E I Fig 2 P L A T E II P L A T E I I I P L A T E I V r P L A T E V I P L A T E V I I 1 1111 ^' ^ S% Q 5 N X? S i ! >0 r - HJ.ONJ7 . 3 7 ¥3? 00 CD 13 HlDN.il IV/DS P L A T E I X 1 5 CD i_ n CP ? JO 7~ ••3D\/J_NJJtiJcJ P L A T E X I 

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