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The anatomy of the North American Pilchard and its bearing on the genetic relationships and nomenclature Whittaker, William R. 1932

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The Anatomy o f t h e K o r t h Amer i can . P i l c h a r d and i t s B e a r i n g on the G e n e r i c • R e l a t i o n s h i p s a n t nomenclature- • . W i l l i a m R. Whlttafcer A T h e s i s s u b m i t t e d f o r the degree o f Mas te r o f A r t s i n the Department o f Z o o l o g y U n i v e r s i t y o f B r i t i s h Co lumbia September - 1932 THE ANATOMY OF THE HORTK AMERICAS PILCHARD AHD ITS BEARISH? OH THE GENERIC RELATIONSHIPS 1ID NOMENCLATURE INTRODUCTION The p i l c h a r d or sardine of the P a c i f i c coast of North America i s known commerically under two names which are gener-a l l y interchangeable„ In California, where both mature and immature f i s h are caught both terms, " C a l i f o r n i a sardine" and "pilchard" are used f o r the species„ the l a t t e r terra generally r e f e r r i n g to the adult„ On the North P a c i f i c coast where the adults alone are talc en i n commercial hauls , the name "pilchard" i s u n i v e r s a l l y used. To avoid any confusion and i n order to di s t i n g u i s h t h i s species from the European species» Sardina pilchardus, the term "North American p i l c h a r d " i s used i n t h i s paper.o A few years ago the question was raised as to whether the pi l c h a r d of North America could be r i g h t l y considered as occupying the same genus as that occupied by the pilchards of the North A t l a n t i c . In f a c t Hubbs (1929) S i n separating these two species g e n e r i c a l l y expresses doubt as to the i r having an immediate common o r i g i n . Accordingly the present work was undertaken with the i i p u r p o s e i n view o f comparing the two t y p e s o f p i l c h a r d s anatom-i c a l l y I n some d e t a i l , n o t i n g s i m i l a r i t i e s and differences» and c o n s i d e r i n g the h e a r i n g o f these on the q u e s t i o n of g e n e r i c nomenc lature* For the g r e a t e r p a r t o f the work the v a r i o u s p o i n t s o f anatomy were determined on f o u r p i l c h a r d s t a k e n i n B a r k l e y sound on the west c o a s t o f Vancouver i s l a n d s She body ( s t a n d a r d ) lengths o f these r a n g e d f rom 218 to ?.6o mil l imetres« These specimens were compared with four European pilchards from P lymouth which had standard lengths o f about 210 m i l l i -metres* 1 h e r e t a k e t h e opportunity o f t h a n k i n g D r , 0, M c l e a n Fraser o f the U n i v e r s i t y o f B r i t i s h Columbia, and D r . W*A. Clemens and D r . J . L . H a r t o f 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 t o whom I am indebted f o r a s s i s t a n c e and r e f e r e n c e s . HISSORT OF SHE HOMENCLATUHE She g e n e r i c and. s p e c i f i c nomenc la ture o f the H o r t h .Ameri-can s a r d i n e o r p i l c h a r d has been l o n g disputed*, She h i s t o r y o f the nomenc la ture o f the s p e c i e s i s b r i e f l y as f o l l o w s : -She p i l c h a r d was f i r s t r e c o r d e d f rom the C a l i f o r n i a ! ! c o a s t by G l r a r d i n 18^ 4 and d e s c r i b e d under the name o f H e l e t t a c a e r u l e a 0 L a t e r 9 l862a G i l l r e d e s e r i b e d the same s p e c i e s as A l a n s a c a l i f o r n l c a and n o t e d i t s p o s s i b l e p o s i t i o n i n t h e ' g e n u s Clupea*.. Jo rdan and G i l b e r t 1881* l i s t the p i l c h a r d as C l u p e a s a j a x 5 ( J e n y n s 9 1842), e v i d e n t l y c o n s i d e r i n g i t to be the same as the C h i l e a n s a r d i n e , y e t a few y e a r s l a t e r J o r d a n and E v e r -mann changed the s p e c i f i c name to c a e r u l e a ( G i r a r d ) . In 1896 the same a u t h o r s changed the g e n e r i c name to Clupanodon LaCepede 9 Jo rdan and G i l b e r t h a v i n g p r e v i o u s l y (l88;5) r e f e r r e d the European p i l c h a r d to t h i s g e n u s 0 I t may be ment ioned here t h a t none o f these g e n e r i c names are i n order-—Clupanodon i s r e s e r v e d f o r a Ch inese genus o f f i s h e s , w h i l e H e l e t t a and A l a n s a are synonyms f o r S p r a t e l l a and A l o s a r e s p e c t i v e l y . Regan's l?l6 c o n t r i b u t i o n has more b e a r i n g on the p r e s e n t nomenclature of the Hor th Amer ican p i l c h a r d . In h i s paper on the C l u p e i n a e he r e f e r s the p i l c h a r d s o f the P a c i f i c to the European genus S a r d i n a A n t i p a , 1906* S h o r t l y a f t e r Regan had - 2 -done so Jo rdan synonym!se& S a r d i n a A n t i p a w i t h S a r d i n i a P o e y t l8j?8« T h i s synonomy was n o t g e n e r a l l y r e c o g n i s e d . F i n a l l y 9 Hubbs (1929) p roposes t o d i v o r c e the two types o f p i l c h a r d s g e n e r i e a l l y a Thus he r e t a i n s S a r d i n a f o r the European p i l c h a r d and s u p p l i e s a new genus s S a r d i n o p s , f o r the H o r t h Amer i can and r e l a t e d s p e c i e s * At p r e s e n t the v a l i d i t y o f S a r d i n o p s as a s e p a r a t e genus i s q u e s t i o n a b l e . The d i a g n o s i s o f S a r d i n a as g i v e n by Regan to forms o f p i l c h a r d s i s as f o l l o w s : -i n c l u d e both S a r d i n a A n t i p a s 1^06 "Elongates, m o d e r a t e l y compressed; abdomen no t s h a r p - e d g e d , Houth moderates the m a x i l l a r y n o t e x t e n d i n g beyond the midd le of the e y e ; l ower jaw no t prominent? u p p e r w i t h o u t or w i t h s l i g h t median n o t c h ; t e e t h minute o r a b s e n t 9 none on vomer. Eye w i t h w e l l - d e v e l o p e d a d i p o s e l i d s * Operculum w i t h grooves r a d i a t i n g towards subopereu lum. D o r s a l f i n o f 16 to 20 r a y s s h i g h e s t a n t e r i o r l y , i n the midd le o f the l e n g t h o f the f i s h , i t s o r i g i n n e a r e r to end o f snout t h a n to base o f c a u d a l ; a s c a l y s h e a t h a t base e x t e n d i n g to t i p o f l a s t r a y . A n a l o f 16 to 20 r a y s r l ow, d e p r e s s i b l e i n a s c a l y s h e a t h ; two l a s t r ays enlarged, , Cauda l f o r k e d ; on each s i d e e n l a r g e d s c a l e s on the i n n e r edge o f the s c a l y p a r t o f each l obe* P e c t o r a l s s c a l y a t base ; p e r v i e s 8~rayed* i n s e r t e d below midd le or p o s t e r i o r p a r t o f d o r s a l . About j H s c a l e s i n a l o n g i t u d i n a l and 10 to 14 i n a t r a n s v e r s e s e r i e s ; v e n t r a l s c u t e s k e e l e d but no t p r o j e c t -i n g beyond the edges o f the groove i n which they l i e . V e r t e -brae j>0 to j j j . " - 4 -The d i a g n o s i s o f S a r d i n o p s which i n c l u d e s the P a c i f i c type o f p i l c h a r d o n l y , I s g i v e n "by Hubbs as f o l l o w s : -S a r d i n o p s Hubbs , 1 9 2 9 T y p e - s p e c i e s , M e l e t t a c a e r u l e a G- irard, 1 8 5 4 . "C lupe i&ae w i t h the u p p e r jaw not n o t i c e a b l y n o t c h e d on the m i d - l i n e ; the g i l l r a k e r s o f the upper l i m b f o l d e d over those o f the lower l i m b , wh ich become marked ly and p r o g r e s s i v e l y s h o r t e n e d towards the a n g l e ; c a r i n a o f g l o s s o h y a l n o t d e n t i c u l a t e ; no b i l o b e d dermal f l a p on the s h o u l d e r - g i r d l e ; o p e r c l e w i t h s t r o n g and marked ly o b l i q u e r i d g e s ; p r e o p e r c u l a r edge s t r o n g l y s l o p i n g ; i n t e r o p e r c l e w i d e l y exposed b e h i n d p r e o p e r c l e ; s c a l e - r o w s r e g u l a r l y s p a c e d , the l a t e r a l s c a l e s a l l w i t h subequa l exposed a r e a s ; r a d i i on the s c a l e s n e a r l y v e r t i c a l ^ and p a i r e d on each s i d e o f median l i n e ; k e e l s on v e n t r a l s c u t e s weak; l a s t two r a y s o f d o r s a l and a n a l f i n s somewhat e n l a r g e d ; a row of dark spo ts t y p i c a l l y deve loped on upper s i d e s beh i n d head*" - 5 -E X T E M A I . • FS/LSHJESS—JTortli Amer ican p i l c h a r d HEAD; The head i s g r e e n - b l u e d o r s a l l y , brown and s i l v e r on the s i d e s and s i l v e r below and compr i ses about t w e n t y - f o u r p e r cent o f the body l e n g t h o f the a d u l t f i sh„ The d o r s a l s u r f a c e and s i d e s a re c o v e r e d by a tough dermal l a y e r which i s marked by a r b o r e s c e n t mucous canals.,. The eyes a re m o d e r a t e l y l a r g e and are p r o v i d e d w i t h a w e l l - d e v e l o p e d adop ise e y e - l i d . The e x t e r n a l na res a re s u p e r o -l a t e r a l , n e a r e r the snout t h a n the e y e s and doub le on each s i d e . The a n t e r i o r s e t i s s m a l l e r and p r o v i d e d w i t h a f l a p - l i l c e v a l v e * The o p e r c l e has f o u r to s i x s t r o n g l y marked r i d g e s e x t e n d -i n g o b l i q u e l y f rom the p r e o p e r c l e to the s u b o p e r c l e be low. The p o s t e r i o r marg in o f the o p e r c l e i s n o t i c e a b l y s l o p i n g ; the p o s t e r i o r edge o f the p r e o p e r e l e i s s t r o n g l y s o . The i n t e r -o p e r c l e i s c o n s i d e r a b l y exposed below the p r e o p e r c l e . The u p p e r jaw i s not n o t i c e a b l y n o t c h e d on the m i d - l i n e ; the lower jaw i s dark i n c o l o u r and the t i p p r o j e c t s but s l i g h t l y beyond the upper jaw. The p o s t e r i o r edge o f the max-i l l a i s below the m i d d l e o r a n t e r i o r p a r t o f the e y e . BODY: She body I s m o d e r a t e l y compressed, rounded d o r s a l l y and weakly k e e l e d on the m i d - v e n t r a l l i n e c She c o l o u r i s dark b l u e on the u p p e r p a r t s and s i l v e r y w i t h m e t a l l i c l u s t r e be low. L a t e r a l l y , e x t e n d i n g backwards f rom the head t h e r e I s g e n e r a l l y one s e r i e s o f l a r g e dark s p o t s i n the same l e v e l as the eye and sometimes a l e s s e r , s e r i e s below t h i s l e v e l . S h i s f e a t u r e i s no t c h a r a c t e r i s t i c o f the i n d i v i d u a l however f o r i n a s m a l l p o r t i o n o f the' s p e c i e s i t I s l a c k i n g a l t o g e t h e r . In a l l the specimens so f a r examined there i s a s m a l l dark spot a t the base o f each s c a l e on the d o r s a l s u r f a c e , making i n a l l about f i v e s e r i e s o f dots on each s i d e o f the m i d - d o r s a l line<> She s c a l e s are t h i n 9 . t ransparent ; , c y c l o i d , and dec iduouso Shey are a r r a n g e d i n r e g u l a r rows and become markedly s m a l l e r towards the base o f the c a u d a l f i n . Shere a re f i f t y - t h r e e to f i f t y - f o u r rows i n a l o n g i t u d i n a l s e r i e s and t e n t o f o u r t e e n i n a t r a n s v e r s e . S c a l e s on the m i d - l i n e below the d o r s a l f i n have about o n e - f i f t h o f t h e i r t o t a l a r e a exposed . She marg in o f the esrposed a r e a i s f i m b r i a t e , t h a t o f the c o v e r e d p o r t i o n i s r e g u l a r o She c i r c u l i on these s c a l e s are p a r a l l e l but not c o n c e n t r i c ; the r a d i i a re s l i g h t l y bowed, p a i r e d on each s i d e o f the median l i n e and n e a r l y v e r t i c a l when the s c a l e i s i n p o s i t i o n , , She s c a l e s on the d o r s a l s u r f a c e a re s m a l l e r than the l a t e r a l s c a l e s and more or l e s s c i r c u l a r ; the c i r c u l i are c o n -c e n t r i c o r n e a r l y s o ; the r a d i i ex tend out r a d i a l l y f rom the c e n t r e . Otherwise the s c a l e s are s i m i l a r to those o f the s i d e s . Around the s h o u l d e r s t h e r e i s a f r i l l o f s c a l e s wh ich i s covered by a dermal f l a p marked by the a r b o r e s c e n t mucous c a n a l s « The m i d - v e n t r a l l i n e i s p r o v i d e d w i t h a s e r i e s o f k e e l e d s c a l e s e x t e n d i n g from the I n s e r t i o n o f the p e c t o r a l s to the i n s e r t i o n o f the v e n t r a l f i n s and f rom b e h i n d the v e n t r a l s to the vent* There a r e about e i g h t e e n b e f o r e and f o u r t e e n b e h i n d the v e n t r a l s ; those b e h i n d the v e n t r a l s become marked ly s m a l l e r towards the v e n t . The k e e l s c a l e s a re composed o f t h r e e p a r t s 0 The two o u t e r p a r t s a re s i m i l a r b e i n g wide a t the base and e x t e n d i n g w e l l up the s i d e o f the f i s h by a s t r o n g s p i n e - l i k e p r o j e c t i o n . The median p o r t i o n i s v e r y s l i g h t l y kee led^ b l u n t l y p o i n t e d a n t e r i o r l y but more expanded p o s t e r i o r l y <> From the p o s t e r i o r end t h e r e are p r o j e c t i n g one median and two o u t e r , minute s p i n e so The d o r s a l f i n i s i n s e r t e d n e a r e r to the snout than to the base o f the c a u d a l f i n . I t i s h i g h e s t a n t e r i o r l y and has seventeen o r e i g h t e e n f i n r a y s . The f i r s t two or three r a y s are s o f t y e t unbranched ; the r e s t o f the r a y s a re branched* The l a s t two r a y s a re n o t n o t i c e a b l y e n l a r g e d but there i s a s c a l y s h e a t h a t the base of the f i n w h i c h extends to the t i p s o f these r a y s . The a n a l f i n has s i x t e e n to e i g h t e e n r a y s : the l a s t two rays are d e c i d e d l y enlarged and l o n g e r * The "/hole f i n has a sealy 'sheath at the base* The caudal f i n i s deeply forked and has twenty-four to twenty-six r a y s . On each side on the inner ed^e of the s e a l y part of each lobe o f the caudal f i n there is an enlarged modi-f i e d s c a l e 0 The pectoral f i n s are inserted bc-low the angle of the subopercle; they are scaly at the base and have sixteen to eighteen dermal rays» The v e n t r a l f i n s are eight-rayed and are inserted below the middle or posterior part of the dorsal» EXTERNAL FEA5?D*EES—European p i l c h a r d HEAD: ' The head compr i ses twenty-two to t w e n t y - t h r e e p e r cent o f the body l e n g t h and resembles t h a t o f the N o r t h Amer ican p i l -c h a r d i n c o l o u r i n g , a r b o r e s c e n t m a r k i n g s , and eye w i t h a d i p o s e l i d o The o p e r c l e h a s s l i k e t h a t o f the o t h e r s p e c i e s , f o u r to s i x r i d g e s , but these d i f f e r i n b e i n g more n e a r l y v e r t i c a l * L i k e w i s e the p o s t e r i o r marg ins o f the p r e o p e r e l e and o p e r c l e are more n e a r l y v e r t i c a l , , The i n t e r o p e r c l e i s exposed below the p r e o p e r e l e to l e s s than h a l f the e x t e n t i t i s I n the N o r t h Amer ican s p e c i e s * The u p p e r jaw i s not n o t i c e a b l y n o t c h e d on the m i d - l i n e : and the t i p o f the lower jaw p r o j e c t s s l i g h t l y beyond the s n o u t . The m a x i l l a r y extends back to a p o i n t below the a n t e r i o r h a l f o f the eye wh ich i s s l i g h t l y a n t e r i o r to the p o i n t reached by the m a x i l l a r y o f the N o r t h Amer ican p i l c h a r d . BODY: The body i s l i k e t h a t o f the N o r t h Amer ican p i l c h a r d i n g e n e r a l c o l o u r and s h a p e . The l a r g e dark spo ts on the s i d e s common to the N o r t h Amer ican s p e c i e s a re g e n e r a l l y l a c k i n g but a c c o r d i n g to Couch (18^5) , and Regan ( l ? l 6 ) , these may be p r e s e n t but not to any g r e a t e x t e n t . The dark s p o t s a t the - 10 -base of the scales on the back are lacking on the specimens examined.,, ' . The scale arrangement i s markedly d i f f e r e n t to that of the Uorth American p i l c h a r d , Here the scales are a l t e r n a t e l y larger and smaller so that the" l a r g e r scales almost completely cover the smaller scales immediately p o s t e r i o r . This arrange-ment gives an apparent number of t h i r t y large scales i n a long-i t u d i n a l series whereas the true number of scales i s about the same as that of the other p i l c h a r d * — f i f t y - t h r e e to f i f t y - f o u r . There are ten to fourteen scales i n a transverse s e r i e s . I t may be pointed out here that t h i s peculiar scale arrange-ment 5 although c h a r a c t e r i s t i c o f the species, i s not t y p i c a l f o r every i n d i v i d u a l of the species 0 According to Day (l88'7) 9 and Bateson (1890, l 8 ? 4 ) s a small proportion of the European pilchards are i n part scaled l i k e the pilchards o f the P a c i f i c » The scales below the dor-sal f i n on the mid-line are approximately the same as those of the North American p i l c h a r d as to the nature of the c i r c u l i and the shape*' The l a r g e r scales are considerably, and the smaller s l i g h t l y l a r g e r than the scales of the North American species from the same body region* The r a d i i are s l i g h t l y more bowed than i n the scales of the other species but are paired on eaeh side of the median l i n e and are nearly v e r t i c a l when the scale Is i n p o s i t i o n . The scales of the back are al t e r n a t e l y larger and smaller. Those anterior to the dorsal f i n are of the same shape as the scales of the sides; those posterior to the dorsal are ovulateo I n a l l o f these as i n s c a l e s frora the same p o s i t i o n on the H o r t h Amer i can s a r d i n e 9 the c i r c u l i a re c o n c e n t r i c ana the r a d i i r a d i a t e f rom the c e n t r e s She f r i l l o f s c a l e s and the dermal f l a p about the s h o u l d e r s a re the same i n bo th spec ies . . The k e e l s c a l e s a r e s i m i l a r i n number to those o f the N o r t h Amer ican p i l c h a r d * The o u t e r p a r t s a re s i m i l a r i n d e s i g n i n both s a r d i n e s exeept t h a t the s p i n e - l i k e p r o j e c t i o n s a re not so s t r o n g i n the European species'* The median p a r t i s s i m i l a r i n b o t h when v iewed f rom above 9 but on the under- s u r f a c e the European s a r d i n e has a pronounced k e e l on the p o s t e r i o r h a l f of the s c u t e . The d o r s a l and p e c t o r a l f i n s a re i n the came r e l a t i v e p o s i t i o n s i n the two p i l c h a r d s ; the d o r s a l has seventeen o r e i g h t e e n f i n r a y s and the p e c t o r a l s i x t e e n to e i g h t e e n r a y s . The a n a l and v e n t r a l f i n s are f a r t h e r fo rward i n the European s a r d i n e but the number o f r a y s f o r each f i n i s the same i n the two s p e c i e s . The European p i l c h a r d has the l a s t two r a y s o f the a n a l f i n e n l a r g e d as i n the N o r t h Amer ican s p e c i e s . The s c a l y sheaths a t t h e base o f a l l the f i n s e x c e p t the v e n t r a l s are p r e s e n t i n bo th s p e c i e s . The c a u d a l f i n Is s i m i l a r i n b o t h , even to the two e n l a r g e d s c a l e s on each a ide o f the f i n . - 12 SKELETAL SYSTEM—North Amer i can p i l c h a r d • •CRANIUM: The o c c i p i t a l r e g i o n i s composed o f f o u r bones: the s u p r a -o c c i p i t a l , , the p a i r e d e x o c c i p i t a l s , , and the b a s i o e c i p i t a l . The s u p r a o c c i p i t a l i s exposed d o r s a l l y i n an e l e v a t e d , t r i a n g u l a r p o r t i o n wh ich s e p a r a t e s the p a r i e t a l s and extends a s h o r t d i s -tance between the frontals» I t i s marked by a pronounced median r i d g e and two t r a n s v e r s e r i d g e s p a i r e d on each s i d e o f the median r i d g e . The median r i d g e c o n t i n u e s down the p o s t e r -i o r s u r f a c e o f the s u p r a o c c i p i t a l t o the magnum fo ramen . The e x o c c i p i t a l s a re j o i n e d below the s u p r a o c c i p i t a l to fo rm the d o r s a l and l a t e r a l b o u n d a r i e s o f the magnum foramen. The b a s i -o c c i p i t a l bounding the foramen v e n t r a l l y i s a l o n g centrum-shaped bone.. The p o s t e r i o r s u r f a c e o f the b a s i o c e i p i t a l i s e n l a r g e d and concave and a r t i c u l a t e s w i t h the f i r s t v e r t e b r a . The v e n t r a l s u r f a c e Is p r o v i d e d w i t h two t h i n p l a t e s wh ich a re f i t t e d to s i m i l a r p l a t e s o f the p a r a s p h e n o i d to form the e y e -muscle cana l . . I n the r e g i o n o f each a u d i t o r y c a p s u l e t h e r e a re f i v e bones. The s p h e n o t i c i s a n t e r i o r and d o r s a l ; i t has a s t o u t l a t e r a l p r o c e s s which marks the p o s t e r i o r boundary o f the o r b i t o f the e y e s and i s p a r t i a l l y c o v e r e d by a l a t e r a l e x t e n -s i o n o f the f r o n t a l . The p r o o t i c i s v e n t r a l to the s p h e n o t i c and the p r o o t i e s o f each s i d e a re j o i n e d v e n t r a l l y to form the base o f the c r a n i u m c The p t e r o t i c marks the p o s t e r i o r l i m i t s o f the c a p s u l e e I t i s c h a r a c t e r i s e d by a l o n g s p i n e - l i k e p r o c e s s w i t h which one b r a n c h o f the p o s t - t e m p o r a l bone a r t i c u -l a t e s * The o p i s t h o t i e i s v e n t r a l to the p t e r o t i c and e x t e r n a l to the e x o c c i p i t a l o I t i s a s m a l l bone and has a s h o r t p o s -t e r i o r p r o j e c t i o n . . The e p i o t i c i s between the s u p r a o e c i p i t a l and the p t e r o t i c and extends backwards to a b l u n t p o i n t , The a n t e r i o r p a r t o f the b r a i n case Is completed by the b a s i s p h e n o l d and the p a i r e d a l i s p h e n o l & s . The b a s i s p h e n c i d Is a s m a l l Y - s h a p e d bone f o r m i n g p a r t o f the boundary o f the o p t i c fo ramen 0 I t ex tends v e n t r a l l y by the base o f the "Y" and I s jo ined, by c a r t i l a g e to the p a r a s p h e h o i d * The a l i s p h e n o i d s are l a r g e r bones d o r s a l t o the foramen; and a re j o i n e d s u t u r a l l y t o the f u s e d o r b i t o s p h e n o i d s * The o r b i t o s p h e n o i d c o n s i s t s o f two f u s e d bones ; a wide p o r t i o n bounds the o p t i c foramen a n t e r i o r l y and a t h i n n e r p o r t i o n extends f o r w a r d to the p r e f r o n t a l to s e p a r a t e the o r b i t s o f the e y e a . The parasphenoid" i s a l o n g u n p a i r e d bone e x t e n d i n g f rom the vomer to b e h i n d the b a s i o c e i p i t a l . The p o s t e r i o r p o r t i o n i s s p l i t i n t o two t h i n n o n - d i v e r g e n t p r o c e s s e s * Below the b a s i o c c i p i t a l the p a r a s p h e n o i d i s t rough-shaped and w i t h t h a t bone i t forms the eye-musc le c a n a l which i s open i n f r o n t and b e h i n d . The p a i r e d f r o n t a l s and p a r i e t a l s compr ise the d o r s a l s u r f a c e o f the skul l„ The f r o n t a l s a r e l a r g e bones e x t e n d i n g f rom the mesethmold i n f r o n t to the s u p r a o c c i p i t a l b e h i n d . They a re narrow and p o i n t e d a n t e r i o r l y but a re expanded p o s t e r -i o r l y . The p o s t e r i o r p o r t i o n i s f o r k e d to a l l o w f o r the s u p e r i o r - t e m p o r a l f o s s a e * The s u r f a c e o f the f r o n t a l s i s mark-ed by d e l i c a t e l o n g i t u d i n a l r i d g e s . The p a r i e t a l s a re s m a l l and a re s i t u a t e d b e h i n d the median e x t e n s i o n s o f the f r o n t a l s . They are s e p a r a t e d f rom each o t h e r by the s u p r a o c c i p i t a l , p o s s e s s s h o r t r i d g e s on the s u r f a c e , and mark the a n t e r i o r l i m i t s o f the e p i o t i e f o s s a e . The bones o f the ethmoid r e g i o n are the p a i r e d p r e -f r o n t a l s „ the menethmoid and vomer* The p r e f r o n t a l s mark the a n t e r i o r b o u n d a r i e s o f the o r b i t s . They are each p r o v i d e d 'with two l a t e r a l p r o c e s s e s ; the l a r g e r s l o p e s downwards and back-wards around the o r b i t a l space ; the other-, s m a l l e r and more v e n t r a l , s l o p e s downwards and fo rwards around the n a s a l s p a c e . The menethmoid i s med ian , a n t e r i o r t o the f r o n t a l s ; i t g i v e s o f f two t h i n p l a t e s p o s t e r i o r l y v/hich ex tend f o r a c o n s i d e r -a b l e d i s t a n c e below the f r o n t a l s . A n t e r i o r l y i t j o i n s the r o s t r a l c a r t i l a g e . The vomer i s a narrow bone f o r m i n g the r o o f o f the mouth below the mesethmoid. I t c u r l s over the r o s t r a l c a r t i l a g e a n t e r i o r l y and extends backwards i n a groove i n the p a r a s p h e n o i d to b e h i n d the p r e f r o n t a l s . The a n t e r i o r p a r t of the mesethmoid g i v e s o f f two bony p r o c e s s e s ; one to each s i d e , w i t h wh i ch the p re raax i l l ae a r t i c u l a t e . The n a s a l s are j o i n e d to the a n t e r i o r p a r t o f the mesethmoid b e h i n d the p r e r a a x i l l a e ; they are s m a l l f l a t bones s l i g h t l y expanded. a n t e r i o r l y , . Around the o r b i t o f the eye are s e v e r a l s c a l e - l i k e bones; the l a r g e p r e o r b i t a l , the s u p r a o r b i t a l and f o u r o r more s u b -o r b i t a l s * SUSPEHSORIUM: She suspensor ium o f each s i d e c o n t a i n s two bones , the hyomand ihu lar and the syraplcct iCt. The hyomand ibu lar i s a l a r g e bone a r t i c u l a t i n g w i t h the s p h e n o t i c and the p t e r o t i c * On the p o s t e r i o r marg in some d i s t a n c e from the p o i n t o f a r t i c u -l a t i o n w i t h the bones o f the a u d i t o r y c a p s u l e the hyomand ibu lar sends o f f a s h o r t rounded p r o c e s s which i s the p o i n t o f a r t i c u -l a t i o n o f the o p e r c l e . Y e n t r a l l y the hyomand ihu lar extends by a narrow p r o c e s s between the p r e o p e r c l e and the quadrate* The s y m p l e c t i e i s a s h o r t r o d - l i k e bone connec ted to the v e n t r a l e x t e n s i o n o f the hyomand ibu lar by c a r t i l a g e ; i t s v e n t r a l e x t r e m i t y f i t s i n t o a s l i g h t n o t c h i n the quadrates JAWS t The bones o f the upper jaw are pa i red , , and t o o t h l e s s . The p r e m a x i l l a e a re most a n t e r i o r ; they a re s m a l l f l a t bones meet ing i n the median l i n e to form the t i p o f the snout„ At the p o i n t o f a r t i c u l a t i o n w i t h the mesethmoid the p r e m a x i l l a e are somewhat t h i c k e n e d . The m a x i l l a e a re l a r g e r bones; they a r t i c u l a t e w i t h the l a t e r a l p r o c e s s e s o f the mesethmoid and extend back to below the o r b i t o f the e y e . Above the m a x i l l a anil a t t a c h e d to I t are the f i r s t and second a d m a x i l l a r i e s . The second a d m a x i l l a r y p a r t i a l l y o v e r l a p s the p o s t e r i o r p o r t i o n o f the m a x i l l a * F i v e p a i r s of r e p l a c i n g bones f o r m the roof and s i d e s of the b u c c a l c a v i t y w i t h i n * These are f o r each s i d e : the p a l a t i n e , a t h i n bone f o r m i n g the r o o f a n t e r i o r l y , and e x t e n d -i n g f rom the a n t e r i o r end o f the vomer to the m e s o p t e r y g o i d and p t e r y g o i d ; the p t e r y g o i d s e x t e n d i n g a long the s i d e f rom the p a l a t i n e to the quadra te ; the mesopte rygo id f o r m i n g the r o o f p o s t e r i o r to the p a l a t i n e and bounded b e h i n d by a h o r i -z o n t a l p r o c e s s from the rne tapterygo id ; the m e t a p t e r y g o i d , fo rming the upper p o r t i o n o f the s i d e p o s t e r i o r l y and t o u c h i n g the hyomand ibu la r ; the q u a d r a t e , below the m e t a p t e r y g o i d and p a r t l y s e p a r a t e d f rom t h a t bone by an o v a l f e n e s t r a . The p r e o p e r e l e I s a t t a c h e d to t h e p o s t e r i o r marg in o f the quadrate where t h e r e i s a s l i g h t t h i c k e n i n g o f the bone; the v e n t r a l end o f t h i s t h i c k e n e d p o r t i o n o f the quadra te a r t i c u l a t e s w i t h the a r t i c u l a r o f the lower jaw* The bones o f the l ower jaw a r e p a i r e d and t o o t h l e s s . The dentary i s a n t e r i o r E l a r g e , and d e e p l y no tched to r e c e i v e a t h i n p r o c e s s f rom the a r t i c u l a r * The a r t i c u l a r i s s m a l l e r , t h i n a n t e r i o r l y but t h i c k e n e d p o s t e r i o r l y so t h a t the t h i c k e n -ed p o r t i o n resembles a "Y" . The p o s t e r i o r p r o j e c t i o n a r t i c u -l a t e s w i t h the q u a d r a t e . The a n g u l a r i s a s m a l l i n c o n s p i c u o u s bone below the r e g i o n o f a r t i c u l a t i o n o f the a r t i c u l a r * OPERCULUM: The opercu lum c o n s i s t s o f f o u r bones: the p r e o p e r c l e , the i n t e r o p e r e l e , o p e r c l e and s u b o p e r c l e 0 The p r e o p e r c l e i s r o u g h l y e r e s c e n t i c i n shape , s l i g h t l y t h i c k e n e d a l o n g the c o n -cave m a r g i n and a t t a c h e d to the quadrate a t the lower end o f t h i s t h i c k e n e d p o r t i o n 0 The i n t e r o p e r e l e i s below the p r e -o p e r c l e and more t h a n h a l f c o v e r e d by i t and i s a t t a c h e d a t i t s a n t e r i o r end to - the a r t i c u l a r o f the mandib le* The o p e r c l e i s the l a r g e s t o f the f o u r bones; i t i s w i d e s t d o r s a l l y and a r t i c u l a t e s w i t h the hyomand ibu lar by a concave d i s c s i t u a t e d about a t h i r d o f the d i s t a n c e down the a n t e r i o r margin* On the e x t e r n a l s u r f a c e o f the o p e r c l e are f o u r to s i x s t r o n g , s l i g h t l y c u r v e d r i d g e s which r a d i a t e f rom the r e g i o n o f a r t i c u -l a t i o n to the whole e x t e n t o f the v e n t r a l margin* The sub-o p e r c l e i s a s m a l l f l a t bone below the o p e r c l e and i s a t t a c h e d to t h a t bone a t i t s v e n t r a l margin* A s h o r t p r o c e s s f r o m the s u b o p e r c l e ex tends up the a n t e r i o r marg in o f the o p e r c l e * HYOTD ARCH AHD ASSOCIATED BOIES: The hyoid. a r c h i s j o i n e d to the c a r t i l a g e between the hyomandibular and s y m p l e c t i c by a s m a l l p a d - l i k e bone, the i n t e r h y a l . The a r c h c o n t a i n s t h r e e bones: the e p i h y a l , c e r a t o -h y a l , and double basihyal» The e p i h y a l i s f l a t , t r i a n g u l a r , and j o i n s the i n t e r h y a l a t the apex and the e e r a t o h y a l a t the b a s e The c e r a t o h y a l i s l o n g and f l a t and bears near i t s - 18 -v e n t r a l m a r g i n , t h r e e or f o u r p e r f o r a t i o n s i n t o which the b r a n c h i o s t e g a l r a y s are hooked . The b a s i h y a l i s s m a l l r -double 9 and i s a n t e r i o r to the c e r a t o h y a l . A t t a c h e d to the h y o i d a r c h are c e r t a i n i n v e s t i n g bones« The g l o s s o h y a l i s s u p p o r t e d by the b a s i h y a l s ; i t I s a s m a l l s k e e l e d , t o o t h l e s s bone s u p p o r t i n g the tongue . The b r a n c h i o -s t e g a l s are a t t a c h e d to the c e r a t o h y a l and the e p i h y a l . They are sword-shaped bones i n c r e a s i n g 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 i n the s e r i e s * There a re s i x . s e v e n , or e i g h t , as the case may b e , i n the s e r i e s ; the f i r s t t h r e e o r f o u r hook i n t o the h o l e s I n the c e r a t o h y a l ; the r e s t o f the r a y s are a t t a c h e d to the f l a t s u r f a c e o f the c e r a t o h y a l and e p i h y a l , and to the i n t e r o p e r c l e by the b r a n c h i o s t e g a l membrane. The u r o h y a l forms the is thmus o f the t h r o a t ; i t i s a sabre shaped bone, a t t a c h e d to the b a s i h y a l a n t e r i o r l y and by c a r t i l a g e to the p e c t o r a l g i r d l e p o s t e r i o r l y . BRANCHIAL ARCHES: There are f i v e b r a n c h i a l a r ches d i m i n i s h i n g i n s i z e from i n f r o n t backwards . The f i r s t t h r e e a r ches a re comple te , c o n -s i s t i n g o f f o u r segments; the h y p o b r a n c h i a l and c e r a t o b r a n c h i a l form the l ower l i m b . o f the a r c h ; the e p i b r a n c h i a l and s u p e r i o r p h a r y n g e a l form the upper l i m b . I n the f o u r t h a r c h the hypo-b r a n c h i a l i s s m a l l and the s u p e r i o r p h a r y n g e a l l a u n o s o i f i e d . The e p i b r a n e h i a l has s t r o n g t e e t h a long p o s t e r i o r marg in and bears d o r s a l l y a l a r g e p l a t e - l i k e bone, the s u p e r i o r e p i b r a n -c h i a l * The f i f t h arch contains a single bone, the i n f e r i o r pharyngeal (ceratobranchial) which bears teeth along i t s pos- . t e r i o r margin. The epibranchials of the f i r s t three arches bear single posterior projections vrhieh extend over the super-i o r pharyngeals of the arches immediately behind,, The hypobranehials of the arches of the two sides are connected by an unpaired basibranchial which i s In f i v e seg-ments , one segment to each areh* The f i r s t segment i s joined . to the glossohyal; the l a s t two segments are not o s s i f i e d . The l a s t segment extends beyond the f i f t h arch i n a c a r t i l a g -inous p l a t e . The superior pharyngeals of the f i r s t arch are connected to the parasphenoid by a small bone s the suspensory pharyngeal* Each branchial arch bears a single row of long g i l l rakers on i t s d o r s a l surface and a double row of g i l l filaments on the vent r a l surface. In the adult f i s h there are about ninety rakers on the lower limb of the f i r s t arch and about f i f t y on the upper limb* In the f i r s t arch the rakers of the upper limb f o l d over the rakers of.the lower of which a number (about ten) become markedly shortened at the angle * Each g i l l raker bears a row of minute processes on each side; these are about h a l f as long as the raker i s wide, and are spaced about three to the distance of t h e i r length. Each process consists of a flask-shaped s t a l k with a fimbriated l e a f - l i k e terminal element. - 20 -VERTEBRAL COLUMN: G e n e r a l l y t h e r e a re twenty abdominal and t h i r t y - o n e c a u d a l v e r t e b r a e e x c l u s i v e o f the h y p u r a l b o n e a . Each abdomin-a l v e r t e b r a , w i t h the e x c e p t i o n o f the f i r s t , c o n s i s t s o f a centrum, shaped l i k e an h o u r - g l a s s 9 concave a t the a n t e r i o r find p o s t e r i o r s u r f a c e s ; parapophyses on each s i d e o f the c e n -trum v e n t r a l l y ; l o n g p l e u r a l s o r r i b s s e x t e n d i n g v e n t r a l l y to l i e under the p r o c e s s e s f rom the k e e l s c a l e s and a t t a c h e d to the parapophyses ; s h o r t e r e p i p l e u r a l s a t t a c h e d to the p a r a p o -physes above the p l e u r a l s ; a d p l e u r a l s o r e x t r a e o s t a l bones j o i n e d h o r i z o n t a l l y to the s i d e s o f the centrum; neurapophyses which a re j o i n e d d o r s a l l y t o form the n e u r a l a r e h and c o n t i n u e i n the n e u r a l s p i n e ; zygapophyses p r o j e c t i n g fo rwards from the base o f the neurapophysea and c o n n e c t i n g then w i t h the neurapo-physes of the v e r t e b r a i n f r o n t ; e p i n e u r a l s a t the base o f the neurapophyses, , > • ' The f i r s t abdominal v e r t e b r a o r a t l a s p o s s e s s e s o n l y the neurapophyses and e p i p l e u r a l s ; the f i r s t few n e u r a l s p i n e s a re s p l i t a t the ends„ The e p i p l e u r a l s , a d p l e u r a l s and e p i n e u r a l s a re absent In the p o s t e r i o r c a u d a l v e r t e b r a e : the p l e u r a l s become m o d i f i e d to form the haemapophyses<> The haemapophyses o f the f i r s t few e a u d a l v e r t e b r a e a re connec ted v e n t r a l l y by a t r a n s v e r s e bar which i s g r a d u a l l y r e p l a c e d by the haemal spine„ The parapophyses remain,, but i n the p o s t e r i o r v e r t e b r a e they be-come s i m i l a r i n shape to the zygapophyses and touch the haemapophyses o f the v e r t e b r a e i n f r o n t * I n a d d i t i o n to these bones connected to the v e r t e b r a e , t h e r e are c e r t a i n numerous s m a l l bones s i t u a t e d i n the rayo-s e p t a ; a s h e a t h o f f i n e bones i s a t t a c h e d to the e p i o t i e and extends backwards i n a f o l d o f the i n t e r m u s c u l a r 5 septum; below and above the a d p l e u r a l s i s a s e r i e s o f t h i n i n t e r m u s c u -l a r bones wh ich are f o r k e d a t the a n t e r i o r end,, These bones become widened to form a s h e a t h over the c a u d a l p l a t e , APPENDICULAR SKELETON: The p e c t o r a l g i r d l e i s suspended f rom the s k u l l by the p o s t - t e m p o r a l s The g i r d l e has the u s u a l number o f p a r t s ; the p o s t - t e m p o r a l i s f l a t and f o r k e d a t one end to a r t i c u l a t e w i t h the e p i o t i e by one f o r k and the p t e r o t i c p r o c e s s by the o t h e r ; the s u p r a c l a v i e l e i s a s c a l e - l i k e bone j o i n i n g the p o s t -tempora l w i t h the c l a v i c l e below by e x t e n d i n g a l o n g the o u t e r s u r f a c e o f eacho The p o s t c l a v i c l e i s i n two p a r t s , the upper p a r t i s a t t a c h e d t o the o u t e r s u r f a c e o f the c l a v i c l e and the lower p a r t extends to a p o i n t beh ind the I n s e r t i o n o f the p e c t o r a l f i n s * The d o r s a l p a r t o f the c l a v i c l e i s narrow and sharp-edged p o s t e r i o r l y ; the v e n t r a l p a r t i s bent to the h o r i -z o n t a l p lane and the edge i s r o l l e d inwards ; a t the angle o f the bend there i s a s h o r t p r o c e s s w i t h an expanded end* The c l a v i c l e s o f eaeh s i d e l o c k a n t e r i o r l y by means o f two o r th ree shor t p r o c e s s e s . The v e n t r a l and i n n e r p a r t o f the g i r d l e i s formed by - 22 -three bones; the hypocoracoid i s f l a t s with a rounded margin a n t e r i o r l y and pointed behind and joins i t s fellow of the other side to form a keel v e n t r a l l y ; the raesaeoracoid i s above the hypocoracoid and anterior to the glenoid facet; the hyper-eoracoid i s small, p o s t e r i o r to the mesacoracoid and dorsal to the glenoid facet• The pectoral rays are segmented and s p l i t lengthwise; they are connected to the g i r d l e by four r o d - l i k e aetinosts and a number of small pad-like aetinosts over which the bent ends of the two halves of each ray f i t * The p e l v i c plate consists of a single f l a t bone,, pointed a n t e r i o r l y , thickened and expanded p o s t e r i o r l y : the p e l v i c rays are s i m i l a r i n form to the pectoral rays and are attached to the plate by two or three small aetinosts. The rays of the dorsal f i n are i n d i v i d u a l l y supported by the interneural spines and two aetinosts. Smaller interneural spines extend from the head to behind the dorsal f i n . The rays of the anal f i n are s i m i l a r l y supported by i n t e r -haemal spines and a c t i n o s t s s the f i r s t spine being the largest of the s e r i e s . The hypural plate supports the caudal f i n . I t i s formed by the haemal and neural spines of the l a s t two vertebrae and the spines associated with the hypural bone. The spines from the l a s t vertebra support the outer rays of the f i n but they are not greatly modified and do not enter into the composition of the plate proper. The neural spine of the l a s t vertebra - 23 -may bear a s m a l l a c c e s s o r y s p i n e i n f r o n t * The h y p u r a l bone and the s p i n e s a t t a c h e d to I t form a s o l i d p l a t e i n the f o l l o w i n g manner: the h y p u r a l i s a c t u a l l y a m o d i f i e d v e r t e b r a e p r e s e n t i n g an u p t u r n e d end p o s t e r i o r l y which r e p r e s e n t s the o s s i f i e d t i p o f the n o t o c h o r d ; the n e u r a l s p i n e f rom the h y p u r a l l i e s over the upturned, t i p and bears an a c c e s s o r y s p i n e a n t e r i o r l y ; the haemal s p i n e i s d o u b l e , p l a t e -l i k e , and forms a g r e a t p o r t i o n o f the c a u d a l p l a t e . The lower p o r t i o n o f the haemal sp ine f rom the h y p u r a l bears a t h i n h o r i z o n t a l r i d g e which s e r v e s to suppor t the b a s a l s e g -ment o f the two median c a u d a l r a y s . A l o n g the t i p o f the h y p u r a l , p o s t e r i o r to the n e u r a l s p i n e , are two s m a l l bones wh ich e v i d e n t l y r e p r e s e n t the n e u r a l s p i n e s o f unformed v e r t e -b r a e . Below one o f these a p i n e s t h e r e i s a double haemal s p i n e , s m a l l e r than t h a t o f the h y p u r a l but s i m i l a r l y m o d i f i e d ; the haemal sp ine below the l a s t n e u r a l s p i n e i s s m a l l and not so w e l l developed*. The r a y s o f the c a u d a l f i n w i t h the e x c e p t i o n o f the two median ones a re s i m i l a r to those o f the d o r s a l o r p e c t o r a l f i n * In the two median r a y s the b a s a l segments a re e n l a r g e d on each s i d e i n t o p a d d l e - s h a p e d p i e c e s which, extend over the c a u d a l p l a t e . SKELETAL SYSTEM—European p i l c h a r d The s k e l e t o n o f the European p i l c h a r d i s m a i n l y s i m i l a r to t h a t of the N o r t h Amer ican p i l c h a r d : i n the r e g i o n o f the s k u l l the c a r i n a o f the g l o s s o h y a l , the vomer, and the p a l a -t i n e s b e a r no t e e t h ; the o e r a t o b r a n c h i a l s o f the f o u r t h b r a n -c h i a l a r e h , and the i n f e r i o r p h a r y n g e a l s bear s t r o n g s p i n e -l i k e t e e t h e The c h i e f d i f f e r e n c e s a re f o u n d i n the shape o f the f o u r o p e r c u l a r bones ; the p r e o p e r e l e i s not e r e s c e n t - s h a p e but more L -shaped a l o n g i t s p o s t e r i o r marg in thus c a u s i n g the i n t e r o p e r c l e to be more t h a n t w o - t h i r d s c o v e r e d ; the o p e r c l e i s as wide a t i t s v e n t r a l edge as i t I s a t the a r t i c u l a r f a c e t , the r i d g e s a r e v e r y s l i g h t l y c u r v e d but o t h e r w i s e the same as those o f the o t h e r p i l c h a r d ; the s u b o p e r e l e i s s l i g h t l y w i d e r Mian t h a t o f the N o r t h Amer i can s p e c i e s because o f the expanded v e n t r a l marg in o f the o p e r c l e ; the i n t e r o p e r c l e i s s l i g h t l y l e s s expanded p o s t e r i o r l y and the marg in i s more c u r v e d so t h a t the p o s t e r i o r e x t r e m i t y i s v e n t r a l i n s t e a d o f d o r s a l as i n the N o r t h Amer i can p i l c h a r d o Another d i f f e r e n c e i s found i n the n a t u r e o f the g i l l r a k e r s : h e r e the r a k e r s are s l i g h t l y s h o r t e r and l e s s numerous— t h e r e b e i n g about s e v e n t y - f i v e on the lower l imb o f the f i r s t arche Only one or two r a k e r s are s h o r t e n e d a t the ang le and the r a k e r s o f the upper l i m b l i e a l o n g those o f the l ower l i m b but do no t o v e r l a p them. T h i s f e a t u r e i s perhaps due to the s h o r t e r r a k e r s and a s l i g h t l y deeper m o u t h . f o r i f the two l i m b s are p r e s s e d c l o s e r t o g e t h e r a r t i f i c a l l y the g i l l r a k e r s o v e r l a p as i n the N o r t h Amer ican p i l c h a r d . The minute p r o c e s s e s on the g i l l r a k e r s a r e , i n the s p e c i -mens examined, s i m i l a r i n shape to those o f the Nor th Amer ican p i l c h a r d , and o f ahout the same l e n g t h and s p a c i n g . T h i s i s c o n t r a r y to the f i n d i n g s o f B ige low as r e p o r t e d by Hubbs (1929) who s t a t e s t h a t the p r o c e s s e s are s h o r t , s p i n e - l i k e , and s l i g h t l y b e n t . T h i s type o f p r o c e s s i s found on the g i l l r a k e r s of the h e r r i n g , C lupea p a l l a s i i , but so f a r has no t been d i s c o v e r e d by an e x a m i n a t i o n o f s e v e r a l specimens o f each type o f s a r d i n e . • The v e r t e b r a l co lumn, the g i r d l e s and the h y p u r a l p l a t e s a re s i m i l a r i n both s p e c i e s , There are g e n e r a l l y f i f t y - o n e . v e r t e b r a e i n each s p e c i e s , but specimens wi th one v e r t e b r a more or l e s s than t h i s number a re f o u n d a - 26 MUSCULATURE The muscle system may be c o n v e n i e n t l y s e p a r a t e d i n t o the f o l l o w i n g : -1. M u s c l e s o f the t r u n k , the g r e a t l o n g i t u d i n a l muscles« 2 . Musc les o f the head r e g i o n * 3« M u s c l e s o f the d o r s a l f i n . 4. M u s c l e s o f the a n a l f i n . J?„ M u s c l e s o f the p e c t o r a l f i n . 6 . Musc les o f the p e l v i c f i n . 7« M u s c l e s o f the c a u d a l f i n . MUSCLES OF THE TRUNK, THE LONGITUDINAL MUSCLES: T h i s group o f musc les forms the major mass o f muscle subs tance on each s i d e o f the body o f the s a r d i n e . There a re t h r e e s e t s o f musc les i n t h i s group; d o r s a l l y there a re the s u p r a e a r i n a l e s ; l a t e r a l l y are the g rea t l a t e r a l musc les p r o p e r , and v e n t r a l l y t h e r e are the i n f r a c a r i n a l e s . The g r e a t l a t e r a l musc les p r o p e r These are the l a r g e s t muscles i n the f i s h and extend from the base o f the s k u l l t o a tendon a t the base o f the c a u d a l f i n on each s i d e o f the f i s h . The e n t i r e muscle l a y e r i s d i v i d e d In to segments o r myomeres 9 each myomere c o r r e s p o n d i n g w i t h a v e r t e b r a be low. The myomeres are s e p a r a t e d from one another by t h i n s e p t a , the rayoeommata. The shape o f the myomeres i s complex b e i n g somewhat o f the form of the l e t t e r "W" but so a r r a n g e d t h a t one myomere o v e r l a p s i n the c e n t r a l p o r t i o n the myomere d i r e c t l y beh ind i t , but i s i n t u r n o v e r l a p p e d a t the a n g l e s by the same myomere. S h i s o v e r l a p p i n g becomes more n o t i c e a b l e i n the c a u d a l myomeres where the c e n t r a l p o r t i o n s become s e r a i c o n i c a l . She g r e a t l a t e r a l muscle c o n s i s t s o f a s u p e r f i c i a l d i v i -s i o n and a deep divis ion*,, She s u p e r f i c i a l d i v i s i o n or musculus l a t e r a l i s 3 u p e r f i c i a l i s , i s s e p a r a t e d f rom the deep d i v i s i o n by a t h i n septum, and i s i t s e l f d i v i d e d by a much l e s s n o t i c e a b l e septum i n t o an e p a x i a l , and a h y p a x i a l d i v i s i o n . , S h i a muscle extends over o n l y p a r t o f the deeper musc le , i s dark brown i n c o l o u r , and i s s e p a r a t e d from the dermis by a l a y e r o f f a t . She deep d i v i s i o n or musculus l a t e r a l i s p r o f u n d i s , has an e p a x i a l and h y p a x i a l d i v i s i o n , extends over the whole s i d e and i s salmon p i n k to whi te i n c o l o u r . These musc les are the c h i e f swimming m u s c l e s , t h e i r purpose b e i n g to bend the body l a t e r a l l y . She d o r s a l l o n g i t u d i n a l m u s c l e s , the s u p r a c a r i n a l e s Shese musc les a re p a i r e d on eaeh s i d e o f the m i d - d o r s a l l i n e , w e l l d e v e l o p e d , and e n c l o s e d i n heavy c o n n e c t i v e s h e a t h s . She myomeres are s p i r a l l y a r r a n g e d . There are two d i v i s i o n s to t h i s g roup ; the a n t e r i o r one i s the p r o t r a c t o r d o r s a l i s , the p o s t e r i o r s the r e t r a c t o r d o r s a l i s . She p r o t r a c t o r d o r s a l i s i s a t t a c h e d by a s h e e t - l i k e tendon or f a s c i a to the s h o u l d e r g i r d l e and o c c i p i t a l bones . I t i s - 28 -i n s e r t e d a t the base o f the f i r s t r a y o f the d o r s a l f i n , and a l s o a l o n g the f u l l l e n g t h o f the median septum hy s l i p s f rom the myomeres. S h i s musc le draws the d o r s a l f i n f o r w a r d and f l e x e s the body dorsally« She r e t r a c t o r d o r s a l i s i s no t so w e l l deve loped as the protractor» I t i s i n s e r t e d a t the base of the l a s t r a y o f the d o r s a l f i n and i s a t t a c h e d p o s t e r i o r l y , by a b r o a d tendon , to the c o n n e c t i v e t i s s u e around the n e u r a l s p i n e s o f the c a u d a l reg ion , , S h i s musc le s e r v e s to move the d o r s a l f i n backwards and c o - o p e r a t e s w i t h the p r o t r a c t o r to f l e x the body. Both o f these musc les are c y l i n d r i c a l i n f orm„ She v e n t r a l l o n g i t u d i n a l m u s c l e s , the i n f r a c a r i n a l e s Shese musc les are p a i r e d on each s i d e o f the m i d v e n t r a l l i n e . • Shey are w e l l deve loped , c y l i n d r i c a l , and e n c l o s e d f o r the most p a r t i n a d i s t i n c t s h e a t h . Shere are th ree d i v i s i o n s to t h i s musc le ; the a n t e r i o r d i v i s i o n i s the p r o t r a c t o r i s e h i i , the mid d i v i s i o n i s the r e t r a c t o r i s e h i i o r p r o t r a c t o r a n a l i s , the p o s t e r i o r d i v i s i o n i s the r e t r a c t o r a n a l i s . She p r o t r a c t o r i s e h i i o r i g i n a t e s i n a f l a t tendon mass which i s a t t a c h e d to the v e n t r a l edge o f the h y p o c o r a c o i d and to the s e p t a o f the l a t e r a l musc les o f t h i s r e g i o n . I t i s i n s e r t e d p o s t e r i o r l y i n t o the a n t e r o - v e n t r a l border o f the i s c h i a l or p e l v i c p l a t e and i n t o a tendon connected w i t h the r e t r a c t o r i s - c h i i * S h i s musc le a c t i n g a l o n e se rves to draw fo rward the p e l v i c g i rd le„ The r e t r a c t o r i s c h i i arises from the posterior border of the ischium as well as from the tendon from the protractor i s c h i i * I t passes around the anal opening and has i t s i n s e r -t i o n at the base of the f i r s t interhaemal spine. Acting alone or with the protractor i s e h i i I t protracts the anal f i n . Act-ing with the r e t r a c t o r a n a l i s , i t re t r a c t s the p e l v i c f i n . The r e t r a c t o r analis i s quite slender. I t i s inserted at the base of the l a s t interhaemal spine and originates from the ends of the haemal spines of the caudal plate. This muscle eo-operates with the other muscles to f l e x the body v e n t r a l l y or acts alone to draw the anal f i n backward. HU5CLES OF TEE HEAD: The dorsal head muscles may be divided into a s u p e r f i c i a l set and a deep set. Of the s u p e r f i c i a l set the adductor manibulae i s by f a r the most noticeable. This muscle forms the "cheek" of the p i l c h a r d and has a cephalic and a mandibular d i v i s i o n . The 'cephalic d i v i s i o n originates p a r t l y from each of the follow-ing: the anterior border of the preoperele, the surface of the quadrate, the raetapterygoid. the hyomandebular, and the con-nective tissues around the levator arcus p a l a t i n i . At•the place of convergence of t h i s fan-shaped muscle a tendon arises which goes out to the angle of the jaw.. The mandibular por-t i o n originates from the tendon over the quadrate bone and i s Inserted Into the inner surface of the mandible. Both of these - 30 -musc les s e r v e to c l o s e the mouth. The l e v a t o r a r c u s p a l a t i n i f i l l s the space i m m e d i a t e l y p o s t e r i o r to the e y e b a l l * I t o r i g i n a t e s f r o m the s p h e n o t i e and p a s s e s undernea th the c e p h a l i c d i v i s i o n o f the adductor man&i-bu lae to i t s p o i n t o f i n s e r t i o n on the s u p e r i o r p o r t i o n o f the hyomandibu lar and the s u p e r i o r marg in o f the m e t a p t e r y g o i d . The d i l a t o r o p e r e u l i i s s m a l l and i s s i t u a t e d immed ia te l y beh ind the l e v a t o r a r c u s p a l a t i n i . I t o r i g i n a t e s f rom the p t e r o t i c and s p h e n o t i e and p a s s e s backward and downward to a l i t t l e knob where the o p e r c l e a r t i c u l a t e s w i t h the hyo-mandi b u l a r * The muscle i s a t t a c h e d by a s t r o n g tendon to the u p p e r marg in of the o p e r c l e 9 thus c a u s i n g the o p e r c l e to s p r e a d outwards when the muscle c o n t r a c t s . The l e v a t o r o p e r e u l i o r i g i n a t e s from the p o s t e r i o r b o r d e r o f the p t e r o t i c and spreads f o r w a r d to be i n s e r t e d i n the u p p e r p o s t e r i o r marg in o f the o p e r c l e . C o n t r a c t i o n o f t h i s muscle e l e v a t e s the opercu lum. There are two deep d o r s a l head m u s c l e s , one concerned w i t h the o p e r c l e and the o t h e r w i t h the p a l a t i n e a r c h . The f i r s t o f t h e s e , the adduc tor o p e r e u l i , o r i g i n a t e s on the v e n t r a l s u r f a c e o f the p t e r o t i c u n d e r the p o i n t o f o r i g i n o f the l e v a t o r oper -e u l i . I t i s i n s e r t e d i n t o the i n n e r s u r f a c e o f the o p e r c l e and opposes the d i l a t o r o p e r e u l i i n a c t i o n . The seeond o f t h e s e , the adduc to r a r c u s p a l a t i n i , o r i g i n -a tes f rom the o u t e r s u r f a c e o f the o r b i t o s p h e n o i d and i s i n s e r t e d by a broad attachment i n t o the m e t a p t e r y g o i d and mesopterygo id* The musc les o f the eye The s i x c h a r a c t e r i s t i c eye musc les appear i n the s a r d i n e , w e l l d e v e l o p e d . The s u p e r i o r and I n f e r i o r o b l i q u e musc les a re a t t a c h e d to the marg in o f the c o r n e a o f the e y e b a l l and pass f o r w a r d to be a t t a c h e d near the a n t e r i o r end o f the i n t e r -o r b i t a l septum* The s u p e r i o r o b l i q u e i s d i r e c t l y above the I n f e r i o r o b l i q u e . The s u p e r i o r r e c t u s and the i n f e r i o r r e c t u s musc les are a t t a c h e d to the eye near the p l a c e o f attachment o f the s u p e r i o r and I n f e r i o r o b l i q u e s r e s p e c t i v e l y . The e x t e r n a l o r a n t e r i o r r e c t u s and the i n t e r n a l or p o s t e r i o r r e c t u s a re a t t a c h e d a t the o t h e r two quadrants o f the e y e b a l l . These f o u r musc les pass backwards i n c l o s e p r o x i m i t y to each o t h e r i n an "eye muscle c a n a l " to t h e i r o r i g i n a t the base o f the skulls There a re i n a d d i t i o n to these musc les many which are much l e s s n o t i c e a b l e . These are the b r a n c h i a l a r c h , and m a n d i b u l a r and h y o i d a r c h musc les wh ich a i d i n r e s p i r a t i o n . MUSCLES O F THE D O R S A L F I H : The musc les o f the d o r s a l f i n a re a l l s l i p - l i k e I n fo rm, one s l i p b e i n g p r e s e n t f o r each r a y . The s u p e r f i c i a l musc le Is the i n e l i n a t o r d b r s a l i s which i s p a i r e d to each s i d e o f the f i n . T h i s muscle i s composed o f n i n e t e e n s m a l l s l i p s each o f which takes i t s o r i g i n i n a f a s c i a which i s a t t a c h e d to the s k i n . T h i s muscle bends the f i n to the s i d e . Below the i n c l i n a t o r a re the s l i p s o f the e r e c t o r d o r s a l i s and the d e p r e s s o r d o r s a l i s , a l t e r n a t i n g w i t h each o t h e r and one o f each l y i n g between two i n t e r n e u r a l s p i n e s . The e r e c t o r i s the more a n t e r i o r o f the s e t o f two * The e r e c t o r d o r s a l i s s l i p s o r i g i n a t e from the i n t e r n e u r a l s p i n e s and the f a s c i a e between the s l i p s o f the two types o f m u s c l e s . I t i s i n s e r t e d on the " t o e " o f the f o o t - s h a p e d end ing o f the dermal f i n r a y . The d e p r e s s o r o r i g i n a t e s s i m i l a r l y and i s i n s e r t e d on the " h e e l " o f the b a s a l p o r t i o n o f the r a y . THIS iUSCLES OF THE ANAL F I N : The musc les o f t h i s f i n a re on the same p l a n as those o f the d o r s a l f i n and a r e ; the i n c l i n a t o r a n a l i s , the e r e c t o r a n a l i s , and the d e p r e s s o r a n a l i s . . There are a l s o s l i g h t muscle f i b r e s between the dermal f i n r a y s c o n s t i t u t i n g tho i n t e r f i l -amenti a n a l i s . THE 1 U S C L E S OF THE PECTORAL F I N : The musc les o f the p e c t o r a l g i r d l e a r e : the abduc tor p e c t o r a l i s s u p e r f i c i a l i s , the a b d u c t o r p e c t o r a l i s p r o f u n d u s , ' the a d d u c t o r p e c t o r a l i s s u p e r f i c i a l i s , the adduc tor p e c t o r a l i s p r o f u n d u s , the e x t e n s o r p e c t o r a l i s . , and the i n t e r f i l a m e n t i p e c t o r a l i s . Qf these the i n t e r f i l a m e n t i , e x t e n s o r , and abduc tor s u p e r f i c i a l i s are f i r s t exposed . The i n f e r i ' i l a m e n t i p e c t o r a l i s c o n s i s t s o f a group o f muscle f i b r e e x t e n d i n g f rom r a y to r a y near t h e i r b a s e s . The a b d u c t o r s u p e r - f i e i a l i s has i t s o r i g i n a l o n g the a n t e r i o r v e n t r a l b o r d e r o f the c o r a c o i d as f a r back as the base o f the f i n c A n t e r i o r l y some o f the tendons from t h i s muscle j o i n w i t h those o f the p r o t r a c t o r I s e h i i * The muscle i s i n -s e r t e d by a s e r i e s o f t e n d i n o u s s l i p s to the t i p s o f the p r o -c e s s e s o f the v e n t r a l 5 or e x t e r n a l h a l f r a y s o f the f i n 5 i n Eueh a way t h a t c o n t r a c t i o n o f the musc le bends the f i n s down-ward and forward* The e x t e n s o r takes i t s o r i g i n f rom the v e n t r a l p o r t i o n o f the c l a v i c l e , under the a n t e r i o r marg in o f the abduc tor s u p e r -f i c i a l i s and I s i n s e r t e d i n t o the base o f the f i r s t f i n r a y e T h i s muscle s e r v e s to spread the f i n I n the h o r i z o n t a l p lane* The abduc to r p e e t o r a l i s p ro fundus l i e s beneath the s u p e r -f a c i a l i s c I t o r i g i n a t e s f rom the v e n t r a l p o r t i o n o f the c o r a c o i d and i s i n s e r t e d by a s e r i e s o f tendon s l i p s , i n t o the i n n e r margins o f the b a s a l p r o c e s s e s o f the v e n t r a l h a l v e s o f the f i n r a y s . T h i s muscle p u l l s the f i n downwards «, The adduc to r p e c t o r a l i s s u p e r f i c i a l i s and p ro fundus both l i e i n t h e - a n g l e between the c l a v i c l e and the c o r a c o i d and are bo th I n s e r t e d i n t o the d o r s a l o r i n t e r n a l h a l f r a y s . They both s e r v e to p r e s s the f i n a g a i n s t the s i d e s o f the body* The s u p e r f i c i a l i s a r i s e s from the p o s t e r i o r v e n t r a l s u r f a c e o f the c o r a c o i d and f rom the mesacoraco id* The pro fundus a r i s e s from the most v e n t r a l p o r t i o n o f the c l a v i c l e and from the d o r s a l marg in o f the c o r a c o i d * - 34 -MUSCLES OF THE PELVIC FIN: The abductor v e n t r a l i s s u p e r f i c i a l i s i s the most median of the muscles of the p e l v i c f i n and has i t s o r i g i n on the vent r a l border of the anterior portion of the p e l v i c plate <, I t i s also connected v/ith the tendon between the protractor i s c h i i and the r e t r a c t o r i s e h i i - I t i s inserted into the tips' of the v e n t r a l halves of the rays of the fin„ The abductor profundus l i e s next to t h i s * I t takes i t s o r i g i n from the v e n t r a l surface of the p e l v i c plate and i s inserted into the v e n t r a l halves of the rays. Both these muscles tend to move the f i n downward, away from the body; and to proximate the f i n rays. The adductor profundus i s a large muscle l a t e r a l to the abductors«, I t arises from the dorsal surface of the pelvic plate and i s inserted into the median borders of the dorsal halves of the rays. The adductor s u p e r f i e i a l i s i s dorsal to a l l of these muscles, and has i t s o r i g i n on the dorsal surface of the p e l v i c plate and i t s insertions into the dorsal halves of the rays of the fin« The adductor group accomplish the spreading of the rays of the pe l v i c f i n , as well as ro t a t i n g the f i n i n towards the body. MUSCLES OF THE CAUDAL FIN: The only s u p e r i f i e i a l muscles here are the i n t e r f i l a m e n t i eaudalis which are muscle f i b r e s running obliquely between the ~ 3>5 ~ c e n t r a l f i n r a y s . A f t e r the tendons o f the l a t e r a l musc les have been removed the deep c a u d a l musc les a r e exposed,, The a d d u c t o r c a u d a l i s v e n t r a l i s i s r o u g h l y t r i a n g u l a r - i n shape„ I t a r i s e s from the d o r s a l marg in and the s u r f a c e o f the l owes t h y p u r a l bones and i s i n s e r t e d i n t o the more d o r s a l c a u d a l f i n rays„ T h i s muscle f l e x e s the d o r s a l l o b e o f the f i n . The f l e x o r c a u d a l i s d o r s a l i s s u p e r i o r , and i n f e r i o r both f l e x the t a i l f o r sv/imming and s t e e r i n g . The s u p e r i o r o r i g i n -a tes i n the median septum over the n e u r a l s p i n e s near the -c a u d a l end, and i s I n s e r t e d on the s i d e s o f the most d o r s a l f i n r a y s . The o r i g i n o f the i n f e r i o r i s f rom the marg in o f the h y p u r a l p l a t e and the i n s e r t i o n i s on the s l i g h t l y more median r a y s o f the d o r s a l l o b e 0 The f l e x o r c a u d a l i s v e n t r a l i s a r i s e s from the bases o f the haemal s p i n e s and extends backward to I t s i n s e r t i o n on the s i d e s o f the r a y s o f the v e n t r a l l o b e I t has an i n f e r i o r and s u p e r i o r d i v i s i o n . The above account a p p l i e s i n g e n e r a l to the muscu la ture o f the European p i l c h a r d . I t i s q u i t e p o s s i b l e t h a t a s i m i l a r type o f muscu la ture a p p l i e s i n g e n e r a l to a l l the members o f the s u b - f a m i l y C l u p e i n a e . DIGESTIVE SYSTEM The pharynx has two pouches or p o c k e t s d o r s a l l y and i s connected, w i t h the s o - c a l l e d stomach by a s h o r t oesophagus; . the stomach i s Y - s h a p e d , the lower branch b e i n g connec ted w i t h the duodenum; the median caecum, wh i ch forms the base o f the "Y" , i s connected w i t h the swim b l a d d e r by the pneumatic duct which passes forwards and upvirards f rom the end o f the caecum to the swim b l a d d e r . The duodenum i s o f s l i g h t l y l a r g e r d i a m e t e r than the r e s t o f the i n t e s t i n e , i s s e p a r a t e d f rom the l ower l i m b o f the stomach by a s l i g h t c o n s t r i c t i o n , and bears the numerous p y l o r i c c a e c a . The duodenum i s f o l d e d upon i t -s e l f and then passes backwards to the vent by the s t r a i g h t t u b u l a r i n t e s t i n e * I n the v i c i n i t y o f the duodenum are the g a l l b l a d d e r , l i v e r , s p l e e n and p a n c r e a s ; the l i v e r i s r i g h t and l e f t l o b e d , the l e f t l o b e b e i n g l a r g e r , and c o v e r i n g the duodenum and p y l o r i c c a e c a ; the g a l l b l a d d e r i s i n the r i g h t l o b e o f the l i v e r , i s f a i r l y l a r g e , g l o b u l a r , and empt ies I n t o the duo-denum by means o f a s h o r t c y s t i c d u c t ; the panceas i s found i n the mesentery -of the f o l d o f the duodenum; the s p l e e n i s brown, l o n g , narrow and three r i d g e d , and i s s i t u a t e d i m m e d i a t e l y d o r s a l to the i n t e s t i n e below the median caecum. The swim b l a d d e r i s w e l l deve loped and b e s i d e s p o s s e s s i n g a pneumatic duct i t opens upon the v e n t by a p o s t e r i o r d u c t , / u l t e r i o r l y the a i r b l a d d e r passes i n t o the neck r e g i o n where - 37 -i t f i r s t narrows and then d i l a t e s to forra a bu lb from which the d i v e r t i c u l a pass forward to rest against the auditory fenes-t r a s The digestive system of the European pilc h a r d i s In general s i m i l a r to that of the North Amer i can p i l c h a r d as described a P y l o r i c eaeea were not counted but should the number be s l i g h t l y d i f f e r e n t f o r the two species any such dif-ference i s not of generic Importance, TTRINO -GElvTTAL SYSTEM The k i d n e y o f bo th s a r d i n e s i s o f the u s u a l C l u p e o i d t y p e ; i t I s an u n p a i r e d , brown, g l a n d u l a r mass e x t e n d i n g the l e n g t h o f the body c a v i t y immed ia te l y v e n t r a l to the v e r t e b r a l column,, and empt ies i n t o the v e n t by a mesonephr ic d u e t . The gonads a re p a i r e d and. are s u p p o r t e d i n the body c a v -i t y by the d o r s a l m e s e n t e r y ; - t h e gonoduct f rom each gonad p a s s e s backwards to a common duet w h i c h opens upon the v e n t . The o v a r i e s are t o r p e d o - s h a p e d and are o f a s e m i - g r a n u l a r c o n -s i s t e n c y i n an immature f e m a l e . I n the na tu re female the ova can be seen I n the o v a r y . The t e s t e s are k n i f e - s h a p e d and are o f a more even texture« The p e l a g i c ova, may be ment ioned h e r e . The ova of the European p i l c h a r d as d e s c r i b e d by Cunningham ( 1 8 % ) , have a r e t i c u l a t e d y o l k , a s i n g l e o i l g l o h u l e and a g rea t space-between the y o l k and the egg membrane. Regan ( 1 9 1 6 ) , and B a r n a r d (1^2.5), d e s c r i b e a s i m i l a r type o f egg f o r the p i l -chards o f the s o u t h e r n hemisphere , £ n e o p i l c h a r d u s and S a r d i n a  s a j a x , wh ich are c l o s e l y r e l a t e d t o the ITorth Amer ican p i l c h a r d . • ~ 39 . BLOOD- VASCULAR SYS T E H Dae t o the d i f f i c u l t i e s presented, by n o n - i n j e c t e d , mater-i a l t h e v a s c u l a r s ystem i s h e r e m e r e l y ou t l i ned , i n g r o s s a s p e c t . I t i s n o t t o be expec ted t h a t any g r e a t d i f f e r e n c e s s h o u l d o c c u r i n t h i s f o r the system i s s i m i l a r f o r a l l t h e ' C l u p e o i d f i s h * HEART: The h e a r t i s s i t u a t e d i n t h e p e r i c a r d i a l c a v i t y between the two h a l v e s of t h e p e c t o r a l a r c h and v e n t r a l t o the oesophagus . I t i s e n c l o s e d w i t h i n the pe rca rd ium and c o n s i s t s o f t h r e e chambers; t h e s i n u s venosus i s d o r s a l and p o s t e r i o r , and r e c e i v e s the two d u c t i C u v i e r i ; the a u r i c l e i s t h i n - w a l l e d a n t e r i o r and d o r s a l , and i s connec ted w i t h the s i n u s venosus by the s i n u - a u r i c u l a r open ing ; the v e n t r i c l e i s v e n t r a l .and p o s t e r i o r , t h i c k - w a l l e d and muscu lar and connec ted w i t h the ; a u r i c l e . A n t e r i o r l y the v e n t r i c l e opens i n t o the bulbous a o r t a which i s a t h i c k e n e d p o r t i o n o f the v e n t r a l a o r t a . ARTERIES: , , From the bulbous a o r t a the v e n t r a l a o r t a extends a n t e r -i o r l y to g i v e o f f the p a i r s o f a f f e r e n t b r a n c h i a l a r t e r i e s . There a re f o u r such p a i r e d a r t e r i e s . The f o u r t h and the t h i r d branches a r i s e f rom a common stem; the second b r a n c h i a l a r i s e s s e p a r a t e l y ; and the f i r s t s e t i s formed by the b i f u r c a t i o n of the v e n t r a l a o r t a . The a f f e r e n t a r t e r i e s are j o i n e d by c a p i l l a r i e s to the e f f e r e n t b r a n c h i a l b r a n c h e s . The e f f e r e n t b r a n c h i a l a rches come t o g e t h e r d o r s a l l y to form a r i n g from which the e x t e r n a l c a r o t i d s are g i v e n o f f a n t e r i o r l y and the d o r s a l a o r t a p o s t e r -i o r l y . The d o r s a l a o r t a passes back to the c a u d a l r e g i o n c l o s e under the v e r t e b r a e f f e n c l o s e d by the haemal a r c h e s f o r p a r t o f the d i s t a n c e . I n I t s a n t e r i o r p o r t i o n the d o r s a l a o r t a g i v e s o f f a r t e r i e s which s u p p l y the v i s c e r a o f the body, YEIHS: Three s e t s o f v e i n s empty i n t o the s i n u s v e n o s u s . The f i r s t of these I s the s e t w h i c h s u p p l i e s the p e c t o r a l g i r d l e and the v e n t r a l abdominal w a l l . The second o f these a re the h e p a t i c v e i n s which connect by s i n u s o i d s w i t h the h e p a t i c p o r t a l system which g a t h e r s the b l o o d from the v i s c e r a . The t h i r d s e t a re the two s t o u t C u v i e r i a n ducts which r e c e i v e the j u g u l a r v e i n s f rom each s i d e o f the head r e g i o n , the g e n i t a l v e i n s and the p o s t e r i o r c a r d i n a l v e i n s . The j u g u l a r v e i n s a re e n l a r g e d and bulbous b e f o r e they j o i n the d u c t i C u v i e r i ; the g e n i t a l v e i n s e n t e r v e n t r a l to the j u g u l a r s . The r i g h t p o s t e r i o r c a r d i n a l o n l y i s x r/ell - 41 -deve loped and e x t e n d s t h e whole l e n g t h o f t h e k i d n e y , imbedded there in , , I t r e c e i v e s * a t i t s p o s t e r i o r end the c a u d a l v e i n w h i c h l i e s below t h e d o r s a l a o r t a e n c l o s e d w i t h i n the haemal arches*. The nervous system i s p r e s e n t e d o n l y i n g r o s s a s p e c t : i t i s s i m i l a r f o r both, s p e c i e s * BRA EH AMD CRANIAL KERVES: The p r o s e n c e p h a l o n i s d i v i d e d i n t o the u s u a l d i v i s i o n s o f t e l e n c e p h a l o n and d ieneepha lon* The t e l e n c e p h a l o n i s d i v i d e d i n t o t w o ' f a i r l y s m a l l c e r e b r a l hemispheres o r corpora, s t r i a t a p o s t e r i o r l y and i n t o two s m a l l e r o l f a c t o r y l o b e s a n t e r i o r l y . The o l f a c t o r y l o b e s each g i v e o f f an o l f a c t o r y nerve which i s about two c e n t i m e t r e s l o n g and ends on the i n n e r s u r f a c e o f the n a s a l capsules» The d i e n e e p h a l o n i s not w e l l d e v e l o p e d and i s c o v e r e d fi.orsa.Ily by the o p t i c l o b e s . I t s p o s i t i o n i s marked however by the p i n e a l body wh ich r i s e s on i t s d o r s a l s u r f a c e and p r o -j e c t s between the o p t i c l o b e s and over the t e l e n c e p h a l o n to a s l i g h t e x t e n t . T e n t r a l l y the d i e n e e p h a l o n g i v e s r i s e to two k i d n e y - s h a p e d b o d i e s , the l o b i i n f e r l o r e s and between these i o b i a re s i t u a t e d the hypophys i s a n t e r i o r l y and the saecus v a s c u l o s u s p o s t e r i o r l y , , These bod ies are both r o u g h l y s p h e r o i d and are e a s i l y detached,, From the a n t e r i o - v e n t r a l margin o f the d i e n e e p h a l o n .the seeond c r a n i a l o r o p t i c n e r v e s are g i v e n o f f . They are v e r y l a r g e and t h i c k and do not merge to form 45 -an o p t i c c h i a s m a , h u t pass out d i r e c t l y , the r i g h t nerve c r o s -s i n g o v e r the l e f t t o go to the l e f t eye and the l e f t nerve to the r i g h t e y e . The mesencephalon i s w e l l deve loped i n t o two l a r g e o p t i c l o b e s a and i a c u r i o u s l y bent upon i t s e l f , due to the e x t e n s i v e growth f o r w a r d . The m i d - b r a i n g i v e s r i s e to the t h i r d c r a n i a l or ocu lomotor nerve on I t s extreme p o s t e r i o r d o r s a l s u r f a c e . The rhombencephalon has the u s u a l d i v i s i o n s i n t o meten-cehha lon and n y e l e n e e p h a l o n . The metencephalon or- c e r e b e l l u m Is pronounced but s m a l l . I t I l e a above the m e d u l l a o b l o n g a t a and i a s l i g h t l y wedge-shaped*, The n y e l e n e e p h a l o n o r m e d u l l a o b l o n g a t a extends f rom the p o s t e r i o r m a r g i n o f the m i d - b r a i n to merge p o s t e r i o r l y i n t o the s p i n a l c o r d . I t g i v e s r i s e to the r e s t o f the c r a n i a l n e r v e s . The s i x t h c r a n i a l or abducens a r i s e s from the v e n t r a l s u r f a c e of the m e d u l l a ; a l l the o t h e r s have l a t e r a l o r i g i n s . AXIAL SERVOUS SYSTEM: The s p i n a l nerve c o r d Is e n c l o s e d by the n e u r a l s p i n e s . I t i s exposed between the s p i n e s and i n these r e g i o n s I t g i v e s o f f the d o r s a l and v e n t r a l r o o t s o f the body n e r v e s . 3ESSE ORGAHS i T h e i n n e r e a r c o n s i s t s o f a r a t h e r l a r g e s y s t e m o f s e m i -c i r c u l a r c a n a l s , u t r i c u i u s , a n d s a c c u l u s , w h i c h c o n t a i n t h e en& o l y r a p h , . T h i s w h o l e s y s t e m i s s u r r o u n d e d b y the p e r i l y m p h c o n t a i n e d i n t h e b o n y a u d i t o r y c a p s u l e , , The p e r i l y m p h i s s e p a r a t e d b y a m e m b r a n e f r o m t h e b r a i n a n d h a s a f e n e s t r a I n t h e p o s t e r i o r o t i c b o n e s * T h e s a c c u l u s c o n t a i n s t h e l a r g e o t o l i t h , t h e s a g ! t t a p the u t r i c u i u s c o n t a i n s two s m a l l e r o t o l i t h s . T h e s a g i t t a e o f t h e t w o s a r d i n e s a r e s i m i l a r a n d c a n h a r d l y be d i s t i n g u i s h e d , t h e o n e f r o m t h e o t h e r * ' -T h e a n t e r i o r c h a m b e r o f t h e eye i s v e r y s h a l l o w s o t h a t t h e c o r n e a l i e s c l o s e " a g a i n s t t h e l e n s * T h e l e n s i s s p h e r i c a l , v e r y h a r d , a n d c a n n o t b e a l t e r e d , i n s h a p e . T h e r e i s f o u n d i n t h e p o s t e r i o r c h a m b e r o f t h e e y e t h e f a l c i f o r n p r o c e s s o f t h e c h o r o i d w h i c h j o i n s w i t h t h e s l e n d e r c a m p a n u l i H a l l e r i a n d s e r v e s t o a d j u s t t h e f o c a l l e n g t h f o r t h e s i g h t o f o b j e c t s * mZSAEX AND CONCLUSIONS The iiuropeari and North. American pilchards when compared anatomically present the following differences: -(1) The size and arrangement of the scales I s d i f f e r e n t f o r the type of each species. ( 2 ) The series of dark body spots I s usually present i n the North American and generally lacking i n the European species; the se r i e s of smaller spots at the base of the dorsal scales i s present only i n the North American p i l c h a r d . (3) The keel scales d i f f e r In shape and i n the prominence of the v e n t r a l k e e l . • (4) The opercular bones are d i f f e r e n t i n shape. ( 3 ) The g i l l rakers of the f i r s t branchial arch d i f f e r i n each type as to numbers and to length; i n the North American p i l c h a r d the shortening of the rakers at the angle i s more noticeable and the rakers of the upper limb f o l d over those of Ibhe lower.*. The two types of pilchards present the following s i m i l -a r i t i e s ; -(1) Each has approximately the same number of vertebrae. (2) The same type of ridges on the opercle. ( 3 ) The same type of caudal scales. (4) The same type of g i l l raker, processes. (j?) The absence of vomerine and palatine teeth. - 46 -The remaining s i m i l a r i t i e s a re mainly/ c h a r a c t e r i s t i c o f the s u b - f a m i l y . The genera o f the s u b - f a m i l y C l u p e l n a e a r e c l o s e l y r e -l a t e d , p a r t i c u l a r l y those t h a t were f o r m e r l y c o n s i d e r e d as occupy ing the genus GTupea, and i n o r d e r to separa te the v a r i o u s groups g e n e r i c a l l y i t " is n e c e s s a r y t o r e s o r t m a i n l y to s u p e r f i c i a l d i f f e r e n c e s 0 However, w i t h r e s p e c t to t h i s f a c t , and I n v iew of the d i f f e r e n c e s and s i m i l a r i t i e s p r e s e n t e d by the two s p e c i e s i n q u e s t i o n , i t Is d i f f i c u l t to c o n s i d e r the two types o f p i l c h a r d s as occupy ing separa te g e n e r a . Two o f the d i f f e r e n c e s n o t e d , the s p o t t i n g and tho s c a l e arrangement , are i n t e r g r a & i n g ; the s l i g h t d i f f e r e n c e i n shape o f the k e e l s c a l e s and o p e r c u l a r bones does not exc lude the p o s s i b i l i t y tha t the two s p e c i e s were d e r i v e d f rom the same a n c e s t r a l type* The o n l y d i f f e r e n c e t h a t c o u l d p o s s i b l y be c o n s i d e r e d as o f g e n e r i c impor tance i s the p e c u l i a r o v e r l a p p i n g of the g i l l r a k e r s found i n .One type but not i n the other* T h i s f e a t u r e taken a lone would s e r v e to s e p a r a t e the IJorth Amer i can p i l c h a r d from the European p i l c h a r d but would not s e p a r a t e the l a t t e r f rom the members o f the genus C l u p e a . However s i n c e b o t h types o f p i l c h a r d s have s i m i l a r g i l l r a k e r p r o c e s s e s i t i s q u i t e p r o b a b l e tha t the o v e r l a p p i n g arrangement o f the r a k e r s as found I n the Hor th American, p i l c h a r d , i s a m o d i f i e d form o f the arrangement f o u n d i n the European p i l c h a r d s Two o t h e r p o i n t s show the c l o s e r e l a t i o n s h i p between the two p i l c h a r d s . I t i s d i f f i c u l t to c o n c e i v e how two p e c u l i a r -i t i e s auch as the o p e r c u l a r s t r i a t i o n s and the c a u d a l s c a l e s a rose i n bo th types o f p i l c h a r d s i f the types d i d not have an immediate common o r i g i n . So t h i s arguement i s added the f a c t t h a t the r e s p e c t i v e p i l c h a r d s have s i m i l a r h a b i t s and the same type o f p e l a g i c o v a . The r e l a t i o n s h i p o f the N o r t h Amer i can p i l c h a r d , ( S a r d i n a c a e r u l e a ) to the p i l c h a r d s o f A u s t r a l i a , Japan , C h i l e , and South A f r i c a , i s as y e t no t f u l l y known. F u r t h e r i n v e s t i g a t i o n i a n e c e s s a r y In o r d e r to dec ide whether these are s e p a r a t e s p e c i e s j s u b - s p e c i e s , or g e o g r a p h i c a l p o p u l a t i o n s . I f the l a t t e r p o s s i b i l i t y s h o u l d be the ease the N o r t h Amer ican p i l c h a r d becomes S a r d i n a s a j a x , ( Jenyns)* A l i s t o f the p o s s i b l e s p e c i e s o f the genus S a r d i n a f r e q u e n t i n g the P a c i f i c and South A t l a n t i c oceans i s g i v e n b e l o w : -S a r d i n a s a jape ( J e n y n s ) , 1842-. C h i l e a n . S a r d i n a c a e r u l e a (G- i ra rd) , 1&54. N o r t h Amer i can . S a r d i n a m e l a i i o s t i c t a (Temminck and S e h l e g e l ) , 1846* Japanese . S a r d i n a o c c l l a t a (Pappe) , 18^3° South A f r i c a n . S a r d i n a neopIIchardus ( 3 t e i n d a c h n e r ) , l8j?3* A u s t r a l i a n . - 48 -T/1TEPATURE C I T E D Barnard;. E.H» 1925« A monograph o f the mar ine f i s h e s o f South A f r i c a . , 3 c A f r i c a n I k s , , A r m . , 21, p t . 1, p . 112... B a t e s o n , W i l l i a m * l890» P i l c h a r d s (C lupea p l l c h a r d u s ) w i t h the number o f s c a l e s .abnormal ly I n c r e a s e d * Eool> S o c . b o n d . , P r c e , p 6 586 - 588. l894.. (On spec imens o f the common p i l c h a r d , C l u p e a p l l c h a r d u s r showing v a r i a t i o n i n the number and s i z e o f the scale's*.) 2 o o l E S o c . L.pnd*, P r o e . , p* 164* Couch, Jonathan* 1835° E s s a y on the p i l c h a r d ( C l u p e a p l l c h a r d u s ) . R* C o r n w a l l P o l y tech.. S o c , Ann . R e p t . y 3, p* $3 101» Cunningham,. J . T . 18.96. The n a t u r a l h i s t o r y o f the marke tab le marine f i s h e s o f t h e B r i t i s h I s l a n d s . London, p . 168 - 177. Day, F r a n c i s * 1887. On a supposed h y b r i d between the p i l c h a r d , (C lupea p l l c h a r d u s ) and the h e r r i n g , ( C . h a r e n g u s ) , and on a specimen o f Salmo purpura tus* Z o o l . S o c L o n d . , P r o e , p* 129 - 130, p i * X P V G i l l , ' T . N . l863« N o t i c e o f a c o l l e c t i o n o f the f i s h e s o f C a l i f o r n i a p r e s e n t e d to the S m i t h s o n i a n I n s t i t u t i o n by I.h% Samuel Hubbard, A c a d . H a t . S c i . P h i l . , P r o c , ( f o r 1862) 14, p . 28l. G i r a r d , C h a r l e s . 1854. D e s c r i p t i o n o f new f i s h e s , c o l l e c t e d by D r . A . L . Heermann, n a t u r a l i s t a t t a c h e d to the survey of the P a c i f i c r a i l r o a d r o u t e , under L i e u t . R .S . V r i l l i a m -s o n 5 C 3 . A . A c a d . H a t . S c i . P h i l . , P r o c , 7 s p.> 138 ~ 199* Hubbs, C a r l L* 1929<• 5?he g e n e r i c r e l a t i o n s h i p s and nomenclature o f the C a l i f o r n i a s a r d i n e . C a l i f . A c a d . S o l . . P r o c * , 4 . s e r . , 1 8 , n o . 11, p . 261 - 263. J o r d a n , D a v i d S t a r r . 19l8. Notes on s i l v e r s i d e s and . s a r d i n e s . G o p e i a , n o . 36 » p, 45- - 46. J o r d a n , D .S . and B.W. Evermann. 18?6« A cheek l i s t o f the f i s h e s and f i s h - l i i c e v e r t e -b r a t e s o f n o r t h and midd le A m e r i c a . U . S . Comm. F i s h . , Rept . ( f o r l8? 3 ) P- 28l. I896. The f i s h e s o f n o r t h and midd le A m e r i c a . U . S . N a t . Huso, B u l l 3 , n o . 47 p t* 1, p . 423 - 424. J o r d a n ; D .S . and C H . G i l b e r t . l88l Notes on a c o l l e c t i o n o f f i s h e s f rom San D iego , C a l -i f o r n i a . U . S . E a t . l i u s . , P r o c . ( f o r i860) 3 , p. 30. 1883. On c e r t a i n n e g l e c t e d names o f L a c e p e d e . U . S . N a t . 3Ius«, P r o c . ( f o r 1882) 3 , p . 374. - 50 Began, Charles T a i t . 193.6, The B r i t i s h , f i s h e s o f the sub - fami l y C lupe inae and r e l a t e d s p e c i e s In o t h e r seas. Ann . Hag. H a t . Hist».jr ser.« 8, 18, n o , lO^V p* 14, pis, 1 » 3 , 1 9 1 6 * l a r v a l and p o s t - l a r v a l f i s h e s . B r i t i s h A n t a r t i c ( T e r r a l lova) E x p e d i t i o n , 1 9 1 0 , 2ool« ? 1-,- n o . 4ti • p* 13 6* V a r r e l l , W i l l i a m . 1836. (Quoting--M.S. o f C o u c h ) . A h i s t o r y o f B r i t i s h f i s h e s , , Londonj 2 , p.,. 1 0 8 » BIBLIOGRAPHIES Wheeler, Genevieve Oorwin. 1 9 3 1 * A" bibliography o f the sardines. Calif., Pi v . 'Fish * Game,, Fish Bull* , ho. 3^ -< Plate 1 "Figure 1 ... ms%h Amer ican P i l c h a r d •+ S ^ r j l n a c a ^ r u l e a P l a t e I Plate II Figures .1, 2, and 2 - Dorsal, lateral, and ventral views of the skull a. mesethmoid b. prefrontal c. frontal d. parietal e. sphenotie f. epiotie g. . pterotic h. prootic i . opisthotic j« supraoocipitai k» exoccipitals 1. basioccipital m, vomer n. parasphenoid o. allsphenoid p. basisphenoid q. orbitosphenoid Figure 4 - The operculum, suspensorium, jaws, and associated bones a. premaxilla b. maxilla c. admaxillaries d. palatine e. pterygoid f # mesopterygoid g» metapterygoid h. quadrate i . hyomandibular j . symplectic k. preopercle 1. interoperele m« opercle n« subopercle o. articular p. angular q. dentary r. glossohyal s, basihyal t. ceratohyal U i epihyal v. urohyal w, branchiostegals P l a t e I I Plate III Figures 1 and 2 - External and internal views of l e f t arch of pectoral girdle a. post-temporal to, supraclavicle c. postclavicles d. clavicle e. hypercoraeoid f. hypocoracoid g. mesocoracoid h. •aetinosts Figure J> - Pelvic plate - internal aspect Figure 4 - Anterior and lateral view of dorsal ray and radial Figures 5 and 6 - Anterior and lateral views of abdom-inal and caudal vertebra© a, centrum b* parapophyses c. neurapophyses d. ; neural spines e. neural arch f• epineurals g. adpleurals h. epipleurals i . pleurals j . haemal arch zygapophyses Figure 7 - Branchial arches - from the l e f t side a f i r s t , second, and third basi-branchials b. hypobranchials c. ceratobranchials &* epibranchials e. superior pharyngeals f. suspensory pharyngeal g. inferior pharyngeal h. superior epibranchial P l a t e I I I F i g . 7 Plate IV Figure 1 -Figure 2 -Figure 3 -Opercular bones of S3, caerulea Op ercular,bones of S. pilchardus Figure 4 -G i l l rakers and g i l l raker process of caerulea a. Arrangement of g i l l rakers at the angle of the f i r s t branchial arch b. G i l l raker process (magnified about 200 X) G i l l rakers and g i l l raker process of S_. pilchardus a. Arrangement of g i l l rakers at the angle of the f i r s t branchial arch b. G i l l raker process (magnified about 200 X) Figure 3 - Keel scale of S. caerulea Figure 6 - Keel scale of S. pilchardus P l a t e I V P l a t e ¥ F i g u r e 1 «• M u s c l e s o f the head and trunk* a* adduc to r mand ibu la r - mandibu lar p o r t i o n b„ a d d u c t o r mandibu lar -» c e p h a l i c p o r t i o n c* g i l l f i l a m e n t s ' el* -musculus l a t e r a l i s profundus -i h j p a x i a l d i v i s i o n ' v~--' e* musculus l a t e r a l i s s u p e r f i c i a l i s f e myomere of the g r e a t l a t e r a l muscle g. l e v a t o r a r cus p a l a t i n e h* d i l a t o r opercu lae I e l e v a t o r o p e r c u l a e k<, musculus l a t e r a l i s p r o f u n d i s <=• e p a x i a l d i v i s i o n 1* m a x i l l a V f l e x e d out o f p l a c e P l a t e V P l a t e VI f i le v e n t r a l l o n g i t u d i n a l muscles a« . p r o t r a c t o r i s c h i i b o r e t r a c t o r i s c h i i Co r e t r a c t o r a n a l i s Musc les o f the p e c t o r a l g i r d l e a, abductor p e c t o r a l i s s u p e r f i c i a l i s b t p r o t r a c t o r i s c h i i C o e x t e n s o r p e c t o r a l i s Musc les o f the p e l v i c g i r d l e a„ p r o t r a c t o r i s c h i i bo adductor v e n t r a l i s profundus c 0 abductor v e n t r a l i s profundus d 0 abductor v e n t r a l i s s u p e r f i c i a l i s e, r e t r a c t o r i s c h i i P l a t e VI P l a t e ¥11 S u p e r f i c i a l musc les o f d o r s a l ana a n a l .fins.g and the d o r s a l • l o n g i t u d i n a l musc les a B p r o t r a c t o r d o r s a l i s h* I n c l i n a t o r d o r s a l i s C e r e t r a c t o r d o r s a l i s d c I n c l i n a t o r a n a l i s Deep musc les o f a n a l f i n s &o d e p r e s s o r a n a l i s ho. e r e c t o r a n a l i s Deep musc les o f the d o r s a l f i n a.„. e r e c t o r d o r s a l i s h e depressor d o r s a l i s Plate VII P l a t e 7IXI F i g u r e 1 ~ S u p e r f i c i a l c a u d a l musc les a<> d o r s a l s l i p o f l a t e r a l i s s u p e r f i c i a l i s b 0 d o r s a l tendon o f <! " e e v e n t r a l , s l i p o f « • . « d« v e n t r a l tendon o f " " e. i n t e r f i l a m e n t i c a u d a l i s F i g u r e 2 - C ross s e c t i o n o f body s l i g h t l y a n t e r i o r to a n a l opening* F i g u r e 3 - Deep c a u d a l musc les a . f l e x o r c a u d a l i s d o r s a l i s s u p e r i o r b* " (} i n f e r i o r C o s r " v e n t r a l i s s u u e r f i c i a l i s d„ » '» " i n f e r i o r e.» " " s u p e r i o r f 0 adduc to r c a u d a l i s v e n t r a l i s g« i n t e r f i l a m e n t i c a u d a l i s Plate V I I I Plate IX f i g u r e 1 " L a t e r a l aspec t o f b r a i n F i g u r e 2 - Ventral a spec t o f b r a i n a. 1st C r a n i a l « O l f a c t o r y nerve b o olfactory l o b e s c* c o r p o r a s t r i a t a d. 2nd Cranial ~ O p t i c nerve e« o p t i c l o b e s f . 3rd. C r a n i a l oculomotor nerve g. 4th C r a n i a l - t r o c h l e a r nerve h. l o b i I n f e r i o r e s hypophys i s k. saccus vasculosus 1« 6th Cranial - abducens nerve m. medulla oblongata n« 5th Cranial « trigeminal nerve 0« 7th " « f a s c i a l nerve p. oth " - auditory nerve q. 9th " - glossopharyngeal nerve r« 10th " vagus nerve a. s p i n a l a c c e s s o r y nerve Figure 3 - Dorsal view of brain Figure 4 - Longitudinal section through the eye a*, cornea b o lens G« s c l e r o t i c d o choroid e* r e t i n a f o campanuli H a l l e r i g o choroid glands Figure 5 - Ear system a» 3aeculus b. u t r i c u i u s C o semi-circular canals d 0 o t o l i t h s Figure 6 « Muscles of th© eye a„ i n f e r i o r oblique muscle b « superior oblique muscle Co i n f e r i o r rectus d o posterior rectus e c i n t e r n a l rectus f« superior rectus P l a t e IX 

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