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A systematic revision of the Hemilepidotinae, a subfamily of cottid fishes Peden, Alexander Edward 1964

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A SYSTEMATIC REVISION OF THE HEMILEPTJDOTINAE, A SUBFAMILY OF COTTTU FISHES. by Alexander Edward Peden B . S c . U n i v e r s i t y o f B r i t i s h Columbia , 19.60... A Thes i s Submit ted i n P a r t i a l F u l f i l m e n t o f the Requirements f o r t h e Degree o f Master o f Sc ience i n t he Department o f Zoology We accept t h i s t h e s i s as conforming t o t h e r e q u i r e d s tandard THE UNIVERSITY OF BRITISH COLUMBIA May, 1964 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of the requirements f o r an advanced degree at the U n i v e r s i t y of • B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study* I f u r t h e r agree that per-m i s s i o n f o r extensive copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s r e p r e s e n t a t i v e s . I t i s understood that;Copying or p u b l i -c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission^ Department of Z o o l o g y The U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, Canada Date J U l y 3 ' 1 9 6 4 i i ABSTRACT An e v a l u a t i o n i s made o f t h e v a l i d i t y , phylogeny, zoogeography, eco logy and taxonomy o f each named spec ies o f t he Hemi lep idot inaeo Hemi lep idotus  sp inosus represen t s t h e most g e n e r a l i z e d l i n e o f d i f f e r e n t i a t i o n w i t h i n t he H e m i l e p i d o t i n a e and be longs t o the monotypic subgenus G a l y c i l e p i d o t u s . H a p a p i l i o i s the most m o d i f i e d . l i n e wh ich belongs t o t h e monotypic subgenus He l l e t eSo The remain ing four spec ie s a re i n t he subgenus Hemi lep idotus o f wh ich H . hemi l ep ido tus i s most g e n e r a l i z e d . A l l s pec i e s a re r e a d i l y i d e n t i f i a b l e down t o s i z e s o f 16 o r 17 mm. The g e n e r a l i z e d spec ies o f t h e H e m i l e p i d o t i n a e are found towards t h e eas te rn p a r t o f t h e range i n t he Nor th P a c i f i c Ocean, w h i l e t h e most mod i f i ed spec ies a re found towards t h e w e s t . The p e l a g i c l a r v a e and p o s t l a r v a e d r i f t w i t h ocean c u r r e n t s wh ich a re p r o b a b l y t he most s i g n i f i c a n t means o f d i s p e r s a l f o r each s p e c i e s . The o r i g i n o f t h e genus Hemi lep ido tus was p r o b a b l y i n t he eas te rn Nor th P a c i f i c Ocean. The m o d i f i e d spec ies p robab ly arose from westward i n v a s i o n s o f t he a n c e s t r a l s tock i n t o t h e wes te rn Nor th P a c i f i c Ocean. The westward ocean cu r ren t across t he A l e u t i a n I s l ands p r o v i d e s a oneway f low o f l a r v a e and p o s t l a r v a e . The g e n e r a l i z e d spec ie s i n t he east have tended t o spread west ac ross t h i s b r i d g e but t he most m o d i f i e d spec ies i n t h e west do not appear t o have d i s p e r s e d back aga ins t t h i s cu r r en t sys tem. A l l s pec i e s o f t he H e m i l e p i d o t i n a e are g e o g r a p h i c a l l y sympat r ic w i t h t h e i r most a l l i e d form and appear t o have e c o l o g i c a l and b i o l o g i c a l d i f f e r e n c e s as w e l l as m o r p h o l o g i c a l d i f f e r e n c e s . X ACKNOWLEDGEMENTS The au thor g r a t e f u l l y acknowledges t he f o l l o w i n g p a r t i e s ; - The I n s t i t u t e o f F i s h e r i e s which made a l l t h e i r specimens and f a c i l i t i e s a v a i l a b l e . - The U n i t e d S ta tes N a t i o n a l Museum which loaned s e v e r a l specimens. - The N a t i o n a l Research C o u n c i l and A r c t i c I n s t i t u t e o f N o r t h America wh ich c o n t r i b u t e d f i n a n c i a l a i d f o r i c h t h y o l o g i c a l exped i t ions o f t he I n s t i t u t e o f F i s h e r i e s t o A l a s k a . - Mr . P . J . E l l i c k s o n , Mr . J . S.; N e l s o n , and D r . J . D . M a c P h a i l , who a s s i s t e d i n t r a n s l a t i o n o f some p e r t i n e n t l i t e r a t u r e . - D r . H . D . F i s h e r , D r . C. C . L i n d s e y , and D r . G . G . Scudder who have c r i t i c a l l y read t h e manuscr ip t . - D r . N . J . Wi l imovsky who o r i g i n a l l y suggested the problem and p r o v i d e d many v a l u a b l e recommendations d u r i n g t he course o f s t udy . 0. i i i TABLE OF CONTENTS page ABSTRACT - i i LIST OF FIGURES • •' v LIST OF TABLES i x ACKNOWLEDGEMENTS • • • • • x INTRODUCTION • ••• 1 MATERIALS AND METHODS ••• 2 Counts and Measurements • 2 D e f i n i t i o n s , ••••• 3 Graphs •• 4 DESCRIPTIONS OF THE HFJ-ITXEPIDOTINAE - 5 Genus Hemilepidotus . C u v i e r • • ••••• ••• 6 Subgenus C a l y c i l e p i d o t u s . Ayres — 9 Hemi lep ido tus ( C a l y s i l e p i d o t u s ) spinosus (Ayres ) 10 Subgenus Hemi lep idotus C u v i e r , 19 Hemilep ido tus (Hemilep i d o t u s ) hemi lep ido tus ( T i l e s i u s ) 24 Hemilep idotus (Hemi lep ido tus ) zapus . G i l b e r t and Burke.. 34 Hemi lep ido tus (Hemi lep ido tus ) .Jordani B e a n — 45 Hemilep i d o t us (Hemilep i do t us) g i l b e r t i Jordan and S t a r k s 56 Subgenus M e l l e t e s Bean 63 Hemilep ido tus ( M e l l e t es) p a p i l i o (Bean) 65 IDENTIFICATION OF THE SPECIES 71 ! Post l a r v a e • 71 Key t o t h e Post l a r v a e •• 72 Key t o t he J u v e n i l e s and A d u l t s 74 Key Charac te r s •• • • 76 Growth C h a r a c t e r i s t i c s and P r o p o r t i o n a l Measurements 76 PHYLOGENETIC RELATIONSHIPS 101 P o s i t i o n o f t he Hemilep i d o t inae w i t h i n the Fami ly C o t t l d a e — • 101 P h y l o g e n e t i c a l l y Important Charac te rs 102 M e r i s t i c Charac te r s ; 102 S t r u c t u r a l Charac te r s ; 102 A . Sca l e s 102 B . Tubu la r L a t e r a l L i n e Elements 107 C P a p i l l a e . on P e l v i c Fins , o f Males I l l 1. S k e l e t a l elements o f p e l v i c p a p i l l a e I l l 2„ E x t e r n a l appearance o f the p a p i l l a e •••• 112 3„ Terminology o f ^ p a p i l l a e " and " c t e n i " . , . 112 4» S i g n i f i c a n c e o f p a p i l l a e and c t e n i . . ^ 112 i v pa^e D. Spines on the Head 113 E . G i l l Membranes Fused t o t h e Isthmus 113 F . C i r r i , 114 P r o p o r t i o n a l Measurements 114 A , F i r i Measurements 114 B . Body Depth 114 Phy logene t i c R e l a t i o n s h i p s o f the Spec ies 115 TAXONOMT AND NOMENCLATURE '. 123 GENERAL LIFE HISTORY OF THE HEMILEPIDOTINAE 124 GEOGRAPHIC DISTRIBUTION • 126 Zoogeography , ••• 126 A . Fac to r s A s s o c i a t e d w i t h Temperature 126 1. Isotherms • 126 2. V e r t i c a l Temperature S t r a t i f i c a t i o n 127 3 . Ice 127 B . Depth •. 127 Co Oceanic C u r r e n t s . . . . . . . . . . . • 127 D i s t r i b u t i o n o f t h e Species- . . • 129 A 0 D i s t r i b u t i o n o f H , sp inosus 130 B . D i s t r i b u t i o n o f H . zapus 130 Co D i s t r i b u t i o n o f H . hemi lep ido tus 132 Do D i s t r i b u t i o n o f H . j o r d a n i - 134 E . D i s t r i b u t i o n o f H . g i l b e r t i 134 Fo D i s t r i b u t i o n o f H« p a p i l i o 137 POSSIBLE GEOGRAPHIC ORIGIN OF THE HEMILEPIDOTINAE — 137 BIOLOGIC AI, DIFFERENCES BETWEEN THE SPECIES.. . 143 Sympatry Between t h e Spec ies o f t h e Hemi l ep ido t inae 143 Depth and H a b i t a t P r e f e r e n c e . . . 143 S e x u a l Dimorphism in" the H e m i l e p i d o t i n a e 146 D i f f e r e n c e s o f Growth i n the Hemi l ep ido t i nae 149 CONCLUSIONS • 151 SUMMARY 153 LITERATURE CITED 158 V LIST OF FIGURES page F igu re 1 The d iagramat ic method o f g r a p h i c a l a n a l y s i s o f Hubbs and Hubbs (19 53) • 5 " 2 P o s t l a r v a e o f H . spinosus . 11 " 3 Hemi lep idotus spinosus • 12 tt 4 P o s t l a r v a e o f H 0 hemi l ep ido tus - .21 n 5 ? P o s t l a r v a e o f Hemilepidotus s p . 22 " 6 Hemi lep idotus hemi lep ido tus • 23 " 7 P o s t l a r v a e o f H . zapus . 35 " 8 Hemi lep ido tus z a p u s . 36 " 9 P o s t l a r v a e o f H . . jordani . 43 n 10 Hemi lep ido tus j o r d a n i - 44 n 11 Hemi lep ido tus g i l b e r t ! . 55 " 12 Sketch o f Hemi lep idotus g i l b e r t i t aken from F igu re 2 o f Gorbunova (1964)- 56 " 13 Hemi lep ido tus p a p i l i o . 64 n 14 Suggested stages o f l i f e h i s t o r y i n the Hemi lep ido t inae- 73 " 15 Approximate range and means o f number o f h o r i z o n t a l s c a l e rows i n d o r s a l s c a l e band o f the Hemi lep ido t inae . 77 " 16 D i s t r i b u t i o n o f l a t e r a l l i n e counts. 77 n 17 D i s t r i b u t i o n o f number o f a n a l f i n r a y s - 78 " 18 D i s t r i b u t i o n o f number o f so f t d o r s a l f i n r a y s . 78 " 19 D i s t r i b u t i o n o f number o f p e c t o r a l f i n rays- 80 M 20 Number o f f r i n g e s on n a s a l c i r r i o f H . hemi lep ido tus - 80 " 21 Ex ten t o f s c a l e s between l e v e l o f lowest row on d o r s a l s c a l e band and l a t e r a l l i n e o f cauda l pedunc le . 81 " 22 P o s i t i o n o f s c a l e s on c a u d a l peduncle i n the H e m i l e p i d o t i n a e . 81 v i page Figu re 23 D i s t r i b u t i o n o f p r o p o r t i o n o f m a x i l l a r y c i r r i w i d t h t o m a x i l l a r y c i r r i l e n g t h . 82 " 24 D i s t r i b u t i o n o f , -p ropor t ion o f he igh t o f f i n membrane on a n t e r i o r s i d e o f f ou r th d o r s a l sp ine compared t o he igh t o f fou r th d o r s a l spine". 82 M 25 Diagram showing r e l a t i v e l eng ths o f f i r s t and second spines o f d o r s a l f i n . 83 " 26 D i s t r i b u t i o n o f r a t i o : o f f i r s t d o r s a l sp ine l e n g t h d i v i d e d by second sp ine l e n g t h . 83 " 27 D i s t r i b u t i o n o f number o f t r a n s v e r s e rows i n d o r s a l s c a l e band . 84 " 28 D i s t r i b u t i o n o f number o f s c a l e s above l a t e r a l l i n e . 84 " 29 Range and approximate means o f ve r t eb rae counts . 85 " 30 D i s t r i b u t i o n o f r e l a t i v e head l eng th . . 85 " 31 D i s t r i b u t i o n o f cauda l peduncle l e n g t h . 86 n 32 D i s t r i b u t i o n o f cauda l peduncle depth . 86 w 33 D i s t r i b u t i o n o f body depth- 87 " 34 D i s t r i b u t i o n o f p e c t o r a l f i n l e n g t h . 87 " 35 Range and mean o f longes t sp ine on d o r s a l f i n . 88 M 36 Range and approximate mean o f number o f p y l o r i c caeca. 88 " 37 R e l a t i v e growth o f eye d iameter i n H . j o r d a n i and H . hemi lep ido tus . 90 1 1 38 R e l a t i v e growth of . i n t e r o r b i t a l w i d t h i n . p o s t l a r v a e o f H . spinosus . 91 » 39 R e l a t i v e growth o f i n t e r o r b i t a l w i d t h i n p o s t l a r v a e o f H . h e m i l e p i d o t u s . 92 " 40 R e l a t i v e growth of . i n t e r o r b i t a l w i d t h i n p o s t l a r v a e o f H . zapus . 93 " 41 R e l a t i v e growth o f i n t e r o r b i t a l w i d t h i n p o s t l a r v a e o f H . .j o r d a n i . 94 *» 42 Range o f head l e n g t h from data o f Gorbunova (1964): 95 v i i page F igure 43 Range o f head depth from da ta o f Gorbunova (1964) • 96 " 44 Range o f eye d iameter from da ta o f Gorbunova , (1964)• 97 " 45 Range o f body depth from da ta o f Gorbunova (1964) • 98 " 46 Range o f l e a s t body depth from da ta o f ' Gorbunova (1964) • 99 " 47 Range o f p r e - a n a l d i s t a n c e from da ta o f Gorbunova (1964) • 100 n 48 Development o f s c a l e s .. 104 " 49 P a t t e r n o f d o r s a l s c a l e band • 106 " 50 Bony t u b u l a r elements o f l a t e r a l l i n e • 108 » 51 P e l v i c c t e n i . • : 109 » 52 P e l v i c p a p i l l a e • 110 " 53 Phylogeny o f t h e H e m i l e p i d o t i n a e . 118 " 54 T h e o r e t i c a l p h y l o g e n e t i c diagram s i m i l a r t o t h a t o f B o l i n (1947) • 124 " 55 Ocean c u r r e n t s i n t he Nor th P a c i f i c Ocean.. 128 n 56 D i s t r i b u t i o n o f H . sp inosus • 131 »» 57 D i s t r i b u t i o n o f H a zapus . 131 " 58 D i s t r i b u t i o n o f H . hemi lep ido tus • 133 » 59 D i s t r i b u t i o n o f H . .Jordani . 135 » 60 D i s t r i b u t i o n o f Hemi lep idotus g i l b e r t i • 136 " 61 Geographic d i s t r i b u t i o n o f Hemi lep ido tus p a p l l i o • 138 " 62 H y p o t h e t i c a l zoogeographic h i s t o r y o f the H e m i l e p i d o t i n a e . * 140 " 63 S e x u a l dimorphism i n H . hemi lep ido tus . 147 " 64 V e n t r a l su r face o f male H . zapus . 147 " 65 S e x u a l dimorphism i n H» . jordani . 148 « 66 S e x u a l dimorphism i n H» g i l b e r t i • 148 v i i i page Figure 67 Sexual dimorphism in H. papilio. 149 n 68 Length frequency of H. hemilepidotus and H. .jordani from collection number BC 63-1010. 150 i x LIST OF TABLES Tab le I D i s t r i b u t i o n Records o f H . s p i n o s u s . " I I D i s t r i b u t i o n Records o f H . h e m i l e p i d o t u s . " n I I I D i s t r i b u t i o n Records o f H . zapus . " IV D i s t r i b u t i o n Records o f H . . j o r d a n i . " V D i s t r i b u t i o n Records o f H . g i l b e r t i . " V I D i s t r i b u t i o n Records o f H . p a p i l i o . " V I I Cha rac t e r index f o r H . s p i n o s u s , H„ zapus 9 H . g i l b e r t i and H . p a p i l i o a p p l i e d t o da ta o f S c h u l t s and Welander (1934). 1 1 V I I I Comparason o f m e r i s t i c c h a r a c t e r s . " IK Comparason,of cha rac t e r s i n the subgenus Hemi lep ido tus w i t h t hose possessed by H . p a p i l i o , " X A.few i n d i v i d u a l s p e c i a l i z a t i o n o f t he spec ies i n the subgenus H e m i l e p i d o t u s . «• " X I Phenet ic d i f f e r e n c e s a c c o r d i n g t o method o f C a i n and H a r r i s o n ( i 9 6 0 ) . " X I I P h y l e t l c d i f f e r e n c e s o f the H e m i l e p i d o t i n a e a c c o r d i n g t o t he method o f C a i n and H a r r i s o n ( i 9 6 0 ) . " X I I I P h y l e t i c d i f f e r e n c e s o f t he Hemilep i d o t inae when each p h y l e t i c c h a r a c t e r i s g i v e n equa l w e i g h t i n g i n s c a l e from n 0 n t o "10". " XIV Summary o f D i s t r i b u t i o n o f the Hemi lep ido t i n a e . " XV Summary on depth o f c o l l e c t i o n s o f t h e H e m i l e p i d o t i n a e . INTRODUCTION W i t h i n t e l e o s t . f i s h e s , t h e S c l e r o p a r e i , an order c h a r a c t e r i z e d by a s u b o r b i t a l s t a y , has r a d i a t e d i n t o many d ivergen t forms. One o f the l a r g e s t groups i s e i t h e r a f a m i l y o r c o l l e c t i o n o f f a m i l i e s making up the C o t t i d a e or C o t t o i d a e . Fur ther s tudy i s needed t o p r o p e r l y de f ine the l i m i t s o f the group 0 The c r i t e r i a u s u a l l y a p p l i e d t o these f i s h e s ares 1 - o p i s t h o t i c s m a l l and f a r removed from p r o o t i c ; 2 - ba s i spheno id absent ; 3 - p e l v i c f i n s w i t h f i v e or l e s s so f t rays and one s p i n e ; 4 - t r u e s c a l e s absent ; 5 - four p r e o p e r c u l a r sp ines u s u a l l y present i n t he younger s t ages . The subfami ly Hemi l ep ido t i nae represen ts t he most g e n e r a l i z e d contempo-r a r y l i n e w i t h i n the C o t t i d a e , The known spec ies o f t h i s subfami ly are c h a r a c t e r i z e d by a cont inuous d o r s a l f i n ; a s i n g l e sp ine and four sof t rays i n t h e p e l v i c f i n ; vomerine and p a l a t i n e t e e t h ; a pore beh ind the l a s t g i l l a r c h ; and four s i m p l e , s t r a i g h t , s t rong p r e o p e r c u l a r s p i n e s . The most d i s -t i n c t i v e fea tu re o f the subfami ly i s the presence o f t h r e e bands o f mod i f i ed s c a l e s . The d o r s a l band o f s c a l e s extends under the base o f the d o r s a l f i n and i s cont inuous ac ross t he nape a n t e r i o r l y . A second band o f s c a l e s extends a long the l a t e r a l l i n e . The v e n t r a l band occurs below the l a t e r a l l i n e . There are no known f o s s i l r e p r e s e n t a t i v e s o f t h i s sub fami ly . W h i l e u s i n g o n l y t h e c r i t e r i a o f d o r s a l l y p l a c e d s c a l e s , Watanabe ( i960) i n c l u d e s many forms i n t he Hemi l ep ido t inae o ther than those i n c l u d e d i n t h i s t h e s i s . Assuming t ha t s c a l e s are common t o a n c e s t r a l c o t t i d s , i t i s reasonable t o expect a p a r a l l e l l o s s o f these s c a l e s i n s e v e r a l p h y l o g e n e t i c l i n e s . Those forms l i s t e d by Watanabe do not possess most o f t he cha rac te r s c i t e d above, even though they have r e t a i n e d s c a l e s . Contemporary l i t e r a t u r e r ecogn izes two t o four generas- C a l y c i l e p i d o t u s wh ich I s synonymized w i t h Hemi lep ido tus ( i . e , B o l i n 1944) and Neohemilepidotus wh ich i s of ten 3ynonymized w i t h M e l l e t e s ( i . e . Andr iashev 1937)« Four t o seven 2 s p e c i e s are a l s o r e c o g n i z e d ; - H , hemi lep ido tus and H , j o r d a n i which a re subspecies a c c o r d i n g t o Schmidt (1929); H . zapus and H . g i l b e r t i wh ich are subspecies a c c o r d i n g t o Gorbunova (1964); H . p a p i l i o wh ich i s monotypic t o M e l l e t e s ; and p a c i f i c u s wh ich i s monotypic t o Neohemilepidotus and synonymized w i t h Ho p a p i l i o ( i . e . Andr io shev , 1937). The purpose o f t h i s r e v i s i o n i s t o e s t a b l i s h t he v a l i d i t y o f each named spec ies and c o n s i d e r t he p h y l o g e n e t i c , zoogeographic and c e r t a i n e c o l o g i c a l r e l a t i o n s between these s p e c i e s . MATERIALS AND METHODS Many i n d i v i d u a l c o l l e c t i o n s were a v a i l a b l e at ihe I n s t i t u t e o f F i s h e r i e s . The m a j o r i t y o f specimens from t h e A l e u t i a n and P r i b i l o f I s l ands were t aken by D r . N . J . Wi l imovsky and t h e author d u r i n g t he summers o f I96I t o 1963° A d d i t i o n a l specimens housed i n t h e I n s t i t u t e o f F i s h e r i e s were ob ta ined from surveys i n t h e G u l f o f A l a s k a conducted by t he I n t e r n a t i o n a l P a c i f i c H a l i b u t Commission. L i m i t e d A s i a t i c m a t e r i a l was a l s o a v a i l a b l e . A d d i t i o n a l specimens were ob ta ined on l o a n from t h e U n i t e d S ta t e s N a t i o n a l Museum. The above m a t e r i a l Inc luded 13 H e s p i n o s u s , approx imate ly 400 H . h e m i l e p i d o t u s , 21 H . zapus , approx imate ly 340 H . j o r d a n i , 9 H . g i l b e r t i » and 5 H . p a p i l i o . The procedure f o r coun t ing and measuring f o l l o w s Hubbs and L a g l e r (1958) except as g i v e n below. Counts and Measurements -Counts o f s c a l e s i n t h e row above l a t e r a l l i n e i n c l u d e a l l s c a l e s i n t h i s row but not any s c a l e s wh ich may be present above t h i s l e v e l , -Counts o f h o r i z o n t a l s c a l e rows i n d o r s a l s c a l e band a re made below t h e s p i n y d o r s a l f i n . Of ten an upper row i s represented by v e r y reduced s c a l e s 3 which t end t o l i e behind r a t h e r than above s c a l e s o f t he adjacent row. T h i s row was cons ide red t o be present o n l y when t h e r e were more t h a n s i x o f these s c a l e s . -Counts o f t r a n s v e r s e s c a l e rows i n d o r s a l band s t a r t from t h e f i r s t a n t e r o -d o r s a l l y d i r e c t e d row and extend t o t he l a s t s c a l e on c a u d a l pedunc le . The count does not i n c l u d e any s c a l e s on cauda l peduncle when they are below the l e v e l o f t he main h o r i z o n t a l rows. -The o r b i t a l d iameter i s t h e l o n g i t u d i n a l d iameter between t h e bony elements o f t he o r b i t . - I n t e r o r b i t a l w i d t h i s t he narrowest w i d t h between t he bony d o r s a l r ims o f each o r b i t . -Body w i d t h i s measured immedia te ly behind t h e c l e i t h r a l s p i n e s . -The body depth i s measured from a n t e r i o r base o f s p i n y d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i n . -He igh t o f f i n membrane a n t e r i o r t o f o u r t h d o r s a l sp ine i s from the a n t e r i o r base o f t h i s sp ine d i r e c t l y t o t he lowest pa r t o f t he margin o f t he f i n . T h i s measurement i s always expressed as a p r o p o r t i o n o f t h e he igh t o f t h e f o u r t h s p i n e . -The g rea te s t w i d t h o f t he m a x i l l a r y c i r r u s was always expressed as a p r o p o r t i o n o f t he g rea te s t l e n g t h o f t h i s c i r r u s . -Snout l e n g t h i s measured from m i d l i n e o f s u b o r b i t a l r i d g e t o o r b i t . P r e m a x i l l a e are not i n c l u d e d . - L a c r i m a l depth i s narrowest depth ac ross s u b o r b i t a l s below o r b i t . -S tandard l e n g t h i s always used i n s t e a d o f t o t a l or fork l e n g t h . D e f i n i t i o n s N a s a l c i r r i ; t h e c i r r i near t h e t i p o f t he n a s a l s p i n e s . M a x i l l a r y c i r r i ; t he c i r r i at t he p o s t e r b - l a t e r a l su r face o f the m a x i l l a e . P o s t o c u l a r c i r r i ; t h e c i r r i i n t he p o s t o c u l a r p o s i t i o n . 4 Frontal c i r r i : the pair of c i r r i between and slightly behind the postocular pair. Occipital c i r r i s the large pair of c i r r i above the posterior limit of the occiput, or over the occipital prominences. Opercular c i r r i s the large c i r r i ahead of the opercular s l i t and on the dorsal aspect of the operculum. Postocular spine; refers to the prominence above the posterodorsal aspect of the orbi t a l rim. Occipital spines refers to the paired prominences above the posterior portion of the occiput. Anal scale rows refers to the single rudimentary scale row which may be situated between the ventral scale band and the anal f i n . . Dorsal scale bands refers to the large scale band at the base of the dorsal f i n . Ventral scale band: refers to the large scale band below the lat e r a l l i n e . Lateral scale bands refers to the scale-like elements of the l a t e r a l l i n e and the scales immediately above these elements. Either d i a l calipers or dividers and rule were used for measurements. Postlarvae were measured with d i a l calipers accurate to 0.1 mm. Unless otherwise stated, measurements represent the proportion of the standard length. The terms "generalized" and "modified" are always used in terms of the degree of change from the supposed ancentral form. The terms "primitive" and "specialized" are rafeher anthropomorphic in outlook. More information is needed to ascertain how a structure is more advantageous than another i f these latte r terms are to be applied. . Graphs Numerous figures are presented in order to f a c i l i t a t e comparisons. This method (see Hubbs and Hubbs 1953) is summarized in Figure 1. 5 S A M P L E I V A L U E O F C H A R A C T E R S T A N D A R D V " " " " 1 1 / " " M E A N D E V I A T I O N J P ' S A M P L E H • •__ ^ i ^ ^ S i j • i j _ — —j , R A N G E — ' ' / p f S T A N D A R D E R R O R S [ O F T H E M E A N Figure 1. The diagrammatic method of graphical analysis of Hubbs and Hubbs (1953) As the dark portion of the bar on either side of the mean Increases in length compared to the light portion, the sample is statistically less adequate. Separation between the ranges of the dark bars of different samples indicates a high degree of significance between the differences of the means of,the samples. Occassionally only the means and ranges are given.. When more extreme values were found in the literature, I have extended the range by a dotted line. The samples represented by each graph are from the specimens used in the species descriptions, except where stated. DESCRIPTIONS OF THE HEMILEPIDOTINAE The genus, subgenera, and species which are recognized in this thesis are described fi r s t . The statistical parameters of the quantitative characters as well as important phylogenetic characters w i l l be discussed after the descriptions. Fin ray counts are recorded as the normal range of counts followed by the 6 extremes o f t he range o f each sample used . T h i s l a t t e r range i s i n b r a c k e t s . Measurements and s c a l e counts are g i v e n as t he mean f o l l o w e d by the extremes o f the range i n b r a c k e t s . Extreme counts and measurements found i n the l i t e r a t u r e a re o f t en i n c l u d e d i n square b r a c k e t s ; the accompanying r a i s e d f i g u r e r e f e r s t o t he a p p r o p r i a t e r e fe rence i n t h e b i b l i o g r a p h y . A l l measurements represent thousandths o f t h e s tandard l e n g t h (except where s t a t e d d i f f e r e n t l y ) . A l l d e s c r i p t i o n s are based on specimens w i t h extended f i n s , depressed operculum, and c l o s e d mouth (except where n o t e d ) . Co lours t end t o be a l t e r e d due t o p r e s e r v a t i o n . Consequent ly , c o l o u r d e s c r i p t i o n s a re based mos t ly on d i s t r i b u t i o n o f dark pigment on a l i g h t background. G e n e r a l d e s c r i p t i o n s are based on specimens above 100 mm i n s tandard l e n g t h , o r t he l a r g e s t specimen a v a i l a b l e (except where s p e c i f i e d ) . I n g e n e r a l , counts and measurements are based on specimens over f o r t y m i l l i m e t e r s , however p o s t l a r v a e were used i n t h e case o f H . zapus and H . s p i n o s u s . A t a b u l a t i o n o f t he l o c a l i t y records f o r each spec ies f o l l o w s t he d e s c r i p t i o n o f t he se s p e c i e s . Specimens from t h e I n s t i t u t e o f F i s h e r i e s have " B C " p r e f i x e d t o t he ca ta logue Mo. An a s t e r i s k denotes those c o l l e c t i o n s which were i d e n t i f i e d by t he au thor . Specimens w i t h d o u b t f u l l o c a l i t y da ta are not i n c l u d e d . S i n c e t h i s r e v i s i o n r ecogn izes one genus i n t he H e m i l e p i d o t i n a e , t he d e s c r i p t i o n o f Hemi lep ido tus serves f o r t ha t o f t he s u b f a m i l y . Genus Hemi lep ido tus C u v i e r Hemi lep ido tus C u v i e r , 1829, p» 165 (Type o f genus Cot tus hemi lep ido tus T i l e s i u s ) . D i a g n o s i s s - The form o f t he t h r e e s c a l e bands, t h e cont inuous d o r s a l f i n s and 7 t he I , 4 p e l v i c f i n s w i l l d i s t i n g u i s h t h i s genus. D e s c r i p t i o n s - S c a l e - l i k e p l a t e s i n t h r e e bands; f i r s t band under base o f d o r s a l f i n and cont inuous across nape w i t h t h e equ iva l en t band on t h e o ther s i d e ; t he second band o f s p i n u l e s on t he p o s t e r o - d o r s a l s i d e o f t he bony t u b u l a r elements o f the l a t e r a l l i n e and a row immediate ly above t h i s ; t he t h i r d band between l a t e r a l l i n e and a n a l f i n and on t a i l . The s c a l e - l i k e p l a t e s c o n s i s t o f many s p i n u l e s r a i s e d p o s t e r o - d o r s a l l y t o produce a p l a t e above t he o r i g i n a l base . . A d d i t i o n a l s c a l e s sometimes present between l a t e r a l and d o r s a l s c a l e bands* A l l p o s t l a r v a e possess an a n a l s c a l e row above base o f a n a l f i n ; t h i s row converges i n f ront o f anus w i t h t he equ iva l en t row on o ther s i d e and then cont inues a long mid l i n e t o b r e a s t . When present i n a d u l t s , s c a l e s o f a n a l row always reduced . Pa tch o f s c a l e s on s i d e o f body near a x i l o f p e c t o r a l f i n . L a t e r a l l i n e s t r a i g h t except fo r weak curve i n middle p o r t i o n . S e v e r a l s m a l l c i r r i o f t e n s c a t t e r e d a long l a t e r a l l i n e and another one beh ind l a s t l a t e r a l l i n e p o r e . Approx imate ly t h r e e l a t e r a l l i n e pores on base o f t a i l . P a i r o f c i r r i on t h e f o l l o w i n g l o c a t i o n s ? p o s t o c u l a r p o s i t i o n , o c c i p u t , p o s t e r o l a t e r a l angle o f m a x i l l a r y , near p o s t e r i o r end and above s u b o r b i t a l s t a y , operculum near upper end o f ope rcu l a r s l i t , l a t e r a l t o median p a i r o f c h i n p o r e s , and p o s t e r i o r t h i r d o f lower l i p . F l e shy t u b e r c l e - l i k e c i r r i on v e n t r a l border o f s u b o r b i t a l r i d g e * Many o f these c i r r i represented as v e s t i g e s i n H . p a p i l i o . Prominent n a s a l , sp ines at apex o f snout and reach l e v e l o f top o f p u p i l * Four p r e o p e r c u l a r s p i n e s , s imple (except f o r lowest sp ine i n some s p e c i e s ) , s t r o n g ; upper p a i r l o n g e r ; upper t h r e e p o i n t p o s t e r i o r l y o r p o s t e r o - d o r s a l l y and s i t u a t e d nearer t o upper h a l f o f p r e o p e r c l e , t he lower sp ine p o i n t e d v e n t r a l l y and s h o r t e r . S m a l l weak sp ines on p o s t e r o - d o r s a l , p o s t e r o - m e d i a l and 8 postere—ventra l edge o f fused ope rcu l a r and subopercu la r bones. These l a t t e r sp ines t end t o be h idden under integument. Short sp ine at t i p o f c l e i t h r u m , d o r s a l t o p e c t o r a l f i n base . Body e longa te , head robust o r s l i g h t l y depressed , t a i l compressed. ; Body deepest under sp iny d o r s a l f i n . Anus near o r i g i n o f a n a l f i n . 'Body wides t at l e v e l o f p r e o p e r c l e . Head s l i g h t l y more than one t h i r d o f s tandard l e n g t h . Eyes o f adu l t p l a c e d h i g h and face l a t e r a l l y . O r b i t s t end t o be r a i s e d above g e n e r a l p r o f i l e o f head. I n t e r o r b i t a l concave i n a d u l t s . Mouth t e r m i n a l , h o r i z o n t a l , p l a c e d v e n t r a l l y ; l a r g e m a x i l l a e extends t o under p o s t e r i o r p o r t i o n o f o r b i t . P r e m a x i l l a e p r o t r a c t i l e , p r o j e c t s beyond lower jaw, p a r t i a l l y f i t s i n t o groove under s u b o r b i t a l r i d g e . L i p s f l e s h y , t h i c k e r towards m i d l i n e . V i l l i f o r m t e e t h i n l a r g e bands on p r e m a x i l l a e , lower jaw, p a l a t i n e s , and vomer. W e l l developed pore beh ind l a s t g i l l a r c h . B r a n c h i o s t e g a l s 6. P o s t e r i o r end o f operculum a l i t t l e over and behind p e c t o r a l f i n base , s h a r p l y a n g u l a r , p o r t i o n o f o p e r c u l a r s l i t above t h i s angle about one h a l f o r b i t diameter o f a d u l t s . Upper p o r t i o n o f operculum at l e v e l o f upper p o r t i o n o f o r b i t . P e l v i c f i n s I , kl membrane i n c i s e d . Sp iny and so f t d o r s a l f i n s con t inuous , separa ted o n l y by n o t c h . P e c t o r a l f i n rounded, upper r ays t end t o be l o n g e r , lower rays t h i c k e r and membrane between them much more i n c i s e d . . P e c t o r a l f i n base broad and con f ined t o lower h a l f o f body, base immedia te ly beh ind operculum, base w i t h v e n t r a l p o r t i o n a n t e r i o r t o d o r s a l f i n . Caudal f i n w i t h up t o n i n e o r t e n branched r a y s . T y p i c a l l y f o u r dark bars ac ross back i n many specimens o f a l l s p e c i e s . Ranges- From Santa Barbara I s . , C a l i f , t o B e r i n g S t . , t o Hakodate, Japan. H e m i l e p i d o t u s ; - f romTM? rtmSwroS meaning " h a l f " " s c a l e d " . 9 Synops i s o f the Subgenera A Sca l e s o f d o r s a l band i n s i x or more h o r i z o n t a l rows. G i l l membranes j o i n e d t o isthmus w i t h no f ree f o l d . C a jyc i l e p i do tus p . 9 AA Sca l e s o f d o r s a l band i n f i v e or l e s s rows. G i l l membrane j o i n e d t o isthmus w i t h f ree f o l d p o s t e r i o r l y . B S c a l e s o f v e n t r a l band equa l i n s i z e t o those o f d o r s a l band. T h i r d sp ine o f d o r s a l f i n s h o r t e r than second and f o u r t h sp ine thus forming an apparent no tch i n spinous d o r s a l f i n . Hemi lep idotus . . . , . . o . , p . 19 BB Sca l e s o f v e n t r a l band l e s s than o n e - h a l f the s i z e o f s c a l e s i n d o r s a l band. T h i r d sp ine in t e rmed ia t e i n s i z e between second and f o u r t h sp ine thus p roduc ing no apparent n o t c h . M e l l e t e s . . o o . . . . . . . . p . 63 Subgenus C a l y c i l e p i d o t us Ayres C a l y c i l e p I d o t u s A y r e s , 1855, p« 76 (Type o f genus by subsequent d e s i g n a t i o n o f Jordan and Evermann, 1896, C a l v e i l e p i d o t u s sp inosus A y r e s ) ; Jordan and Evermann, 1898, p . .1936; Schmidt , 1929, p . 369. D i a g n o s i s ; - The s i x o r more rows o f s c a l e s i n t he d o r s a l band and t he fused isthmus comple te ly d i s t i n g u i s h e s t h i s form from o thers i n t he genus. D e s c r i p t i o n : - A l l s c a l e bands w i t h s i m i l a r s i z e d s c a l e s . S i x t o e igh t h o r i z o n t a l rows o f s c a l e s i n p o r t i o n o f d o r s a l band under s p i n y d o r s a l f i n ; s c a l e s i n upper rows s m a l l e r . V e n t r a l band o f s c a l e s w i t h o n l y four or f i v e h o r i z o n t a l rows at w ides t p o i n t . A n a l s c a l e row cont inuous behind l a s t a n a l f i n r ay i n p o s t l a r v a e ; absent i n j u v e n i l e s and a d u l t s . 10 Rounded and e longate prominence on a n t e r i o r p o r t i o n o f f r o n t o p a r i e t a l r i d g e . A s i m i l a r p a i r o f prominences m e d i a l t o t he g r o n t o p a r i e t a l prominence. G i l l membranes u n i t e d , j o i n e d t o isthmus at t h e i r p o s t e r i o r margin so as not t o produce a f ree f o l d . T h i r d spinous ray o f d o r s a l f i n tends t o be s h o r t e r than second o r f o u r t h spinous r a y s . Marg in o f f i n membrane g r e a t l y I n c i s e d between t h i r d and f o u r t h spinous r a y s , thus forming an obvious n o t c h . T h i s subgenus i s monotypic . R a n g e l - Santa Barbara I s . , C a l i f o r n i a t o Queen C h a r l o t t e Sound, B r i t i s h Columbia . C a l y c i l e p i d o t u s ; - from WAV^ ACn?6WT6j meaning "cup" " s c a l e d " . Hemilepidotus ( C a l y c i l e p i d o t u s ) spinosus (Ayres ) ( F i g . 2 and 3) C a l y c i l e p i d o t u s 3pinosus A y r e s , 1855, p . 76 (San F r a n c i s c o B a y ) ; Jordan and Evermann, 1898, p . 1937; S t a r k s and M o r r i s , 1907, p . 219; Hubbs, 1928, p . 14. Hemilepidotus sp inosus G i r a r d , 1856, p . 134* 1857, p . 13 ; 1858, p . 68; Gunther 1860, p . 174; Jordan and G i l b e r t , 1881, p . 454; 1882, p . 714; Schmidt , 1929, p . 369; Ba rnha r t , 1936, p . 66; B o l i n , 1944, p . 15 , f i g . 4; Clemens and W i l b y , 1949, p . 243, f i g ; 1961, p . 283 , f i g . D i a g n o s i s ; - t ha t o f subgenus. M a t e r i a l s ; - Counts are based on f i f t e e n specimens from c o l l e c t i o n No. BC 62-495» BC 53-92; BC 60-170. Measurements are based on t h r e e specimens from c o l l e c t i o n No. BC 62-495; BC 53-92. D e s c r i p t i o n ; - D o r s a l X I , 19 (18) [18-20J.1 A n a l ; 15 ( I 4 ) [ l 4 - l 6 p . P e c t o r a l s 15 \ (16) [ 1 4 - 1 6 ] n . V e r t e b r a e ; 3 5 . L a t e r a l l i n e po re s ; 63 (62-67X(57-66)+(3-6)fL, Transverse s c a l e rows i n d o r s a l band ;75 ( 6 6 -81 ) [75 -85 j^» Number o f s c a l e s i n row POSTLARVA OF H.SPINOSUS Figure 2 . F E M A L E : - C O L L . N O . B C 6 2 4 9 5 . , S Q M M . , Figure 3 : HEMILEPIDOTUS SPINOSUS. 13 above l a t e r a l l i n e s 29 (18-36) [ 2 2 - 3 7 ] 1 1 . Number o f h o r i z o n t a l s c a l e rows i n d o r s a l bands 6-8. P y l o r i c caeca ( l specimen) s 4« Standard l e n g t h o f specimenss 59«0-l64<>0 mm. Width immediate ly behind p e c t o r a l f i n s s (183-200). Depth from o r i g i n o f d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i n s s (225-261) [ 238 (215-273)] 1 1 . Depth from o r i g i n o f so f t d o r s a l f i n t o o r i g i n o f a n a l f i n s (190-222) [205 (185-233)] 1 1 . Head lengths (373-383) [390 ( 3 6 8 - 4 2 0 ) ] 1 1 . O r b i t diameters over 100 mm - (87-98) [106 (94-1 2 8 ) ] 1 1 ; under 100 mm - (109). O r b i t t o end o f operc les (189-213). I n t e r -o r b i t a l wid ths (40-48).- Snout l e n g t h ( e x c l u d i n g p r e m a x i l l a e ) s (75 -80) . Narrowest l a c r i m a l widths (33-38). Caudal peduncle lengths (83 -98) . Caudal peduncle depths (65-71) [73 ( 63-90)] 1 1 . Snout t o o r i g i n o f d o r s a l f i n s (312-321) [332 (305-370)1 n . Snout t o o r i g i n o f a n a l f i n s (591-623) [599 ( 566-11 627)] . Length o f d o r s a l f i n bases (630-644)« Height o f longes t sp ine on f i r s t d o r s a l f i n s (92-105) [110 (101-128)] 2 1 . Length o f a n a l f i n bases (296-309) [307 ( 273-347J}1. Length o f longes t sof t r a y on d o r s a l f i n s (132-165) [165 (150-183)] 1 1 . Length o f longes t r ay on a n a l f i n s (107-121) [132 (121-140)Ji Length o f longes t r ay on p e c t o r a l f i n s (233-253) [267 (242-3051 1 1 . Length o f longes t r a y on p e l v i c f i n s (174-193) [198 (176-233J] 1! Length o f longes t ray on c a u d a l f i n s (205-243) [227 (204-279 J 1 1 . Head depressed . I n l a t e r a l v i e w , p r o f i l e o f o c c i p u t not s l o p i n g g r e a t l y , and s t r a i g h t . Pa th o f t h i s p r o f i l e i n l i n e w i t h upper borde r o f p u p i l . The v a r i o u s sp ines on top o f s k u l l break up t h i s p r o f i l e . O r b i t s e l e v a t e d , but due t o n a r r o w i n t e r o r b i t upper border o f eye obscures d o r s a l p o r t i o n o f o r b i t a l r i m . Snout s t r a i g h t t o t i p o f n a s a l s p i n e . In p r o f i l e from d o r s a l v i e w , p r e m a x i l l a e rounded. Head s l i g h t l y p o i n t e d . The p r o f i l e o f m a x i l l a e and cheek o n l y s l i g h t l y rounded. Body deepest below about t h e f i f t h d o r s a l f i n s p i n e . Nape f l a t o r s l i g h t l y 14 convex when viewed from l a t e r a l s i d e . D o r s a l f i n base s l i g h t l y convex. D o r s a l p r o f i l e and v e n t r a l p r o f i l e o f cauda l peduncle s h a l l o w l y concave. A n a l f i n base s t r a i g h t . Anus In f ron t o f a n a l f i n o r i g i n . P r o f i l e o f b e l l y s t r a i g h t o r convex. P r o f i l e n e a r l y s t r a i g h t from p e l v i c f i n s t o c h i n . In p r o f i l e from d o r s a l v i e w , body wide and l a t e r a l p r o f i l e s t r a i g h t t o narrow cauda l pedunc l e . A n t e r i o r n o s t r i l q u i t e s h o r t , t u b u l a r ; r i m f l a r e d l i t t l e . P o s t e r i o r r i m o f n o s t r i l g r e a t l y e l eva ted i n t o f l a p more than t w i c e t he he ight o f a n t e r i o r s i d e o f n o s t r i l . From d o r s a l v i e w , a n t e r i o r n o s t r i l l a t e r a l t o n a s a l sp ines and d i r e c t l y a n t e r i o r t o upper edge o f i r i s o f eye. P o s t e r i o r n o s t r i l s h o r t , t u b u l a r , s l i g h t l y c o n s t r i c t e d at t i p . N o s t r i l d i r e c t l y i n l i n e from n a s a l s p i n e t o l a t e r a l border o f I n t e r o r b i t , and m e d i a l t o an te r io rmos t edge o f o r b i t . D i r e c t l y i n l i n e between n o s t r i l s and near base o f p o s t e r i o r n o s t r i l i s l a r g e r head p o r e . Pores abundant on s u b o r b i t a l , p r e o p e r c u l a r , p o s t o r b i t a l and o c c i p i t a l a r ea s . O r b i t l a r g e , becomes r e l a t i v e l y s m a l l e r at l a r g e r s i z e s . I n t e r o r b i t narrower and s h a r p l y concave. L o n g i t u d i n a l p a i r o f r i d g e s near m i d l i n e not p rominen t . N a s a l sp ines s t r ong and sharp , r a i s e d t o near l e v e l o f upper s i d e o f p u p i l . Spines d i r e c t l y a n t e r i o r t o l a t e r a l edge o f i n t e r o r b i t . P r e o p e r c u l a r sp ines s t rong and sharp . Second p r e o p e r c u l a r sp ine s l i g h t l y l o n g e r than uppermost one, about l e n g t h o f p u p i l d iamete r , and p o i n t s p o s t e r i o r l y . Upper p r e o p e r c u l a r sp ine p o i n t s p o s t e r o d o r s a l l y . T h i r d sp ine q u i t e s h o r t , sharp , and b road . Lowest or f o u r t h p r e o p e r c u l a r sp ine s h o r t , sharp , broad and l a r g e l y hidden beneath s k i n . Short sp ine on p o s t e r o d o r s a l edge o f o p e r c l e . Short sp ines on p o s t e r i o r and p o s t e r o v e n t r a l s i d e o f suboperc le wh ich i s fused t o o p e r c l e . These sp ines o f t en h idden under s k i n . C l e i t h r a l sp ine sharp but h idden under s k i n . Knobs o r rough r i d g e s on upper p o s t o c u l a r a r e a . O c c i p i t a l sp ines absen t . Rough t r a n s v e r s e r i d g e ac ross p o s t e r i o r end o f i n t e r o r b i t . 15 P t e r o t i c r i d g e prominent and con t inues past p o s t - t e m p o r a l a rea t o upper end o f ope rcu l a r s l i t . Pos t - t empora l r i d g e p r e s e n t , r i d g e on upper end o f o p e r c l e prominent . P a i r o f l o n g i t u d i n a l r i d g e s near m i d l i n e o f o r b i t not p rominent . Areas between r i d g e s ( i n c l u d i n g f r o n t o - p a r i e t a l r i d g e ) g r e a t l y depressed . These r i d g e s become more prominent and roughened i n l a r g e r specimens. C i r r i more numerous and t e n d t o be broad and l a r g e . P a i r below n a s a l sp ines v e r y s m a l l and t u b e r c l e - l i k e . N a s a l c i r r i s h o r t , t h i n , nar row, on a n t e r i o r s i d e o f n a s a l s p i n e . P o s t o c u l a r c i r r i weakly m u l t i f i d t o e n t i r e , b road . F r o n t a l c i r r i s m a l l e r , h i g h l y m u l t i f i d , c l o s e t o m i d l i n e , d i r e c t l y med ia l t o p o s t o r b i t a l p a i r . O c c i p i t a l c i r r i , b r o a d , t h i n , weakly m u l t i f i d . Reduced c i r r i on a n t e r i o r prominences o f f r o n t o - p a r i e t a l r i d g e . C i r r i o f t en present on prominence m e d i a l t o t h a t o f f r o n t o - p a r i e t a l r i d g e . Three t u b e r c l e -l i k e c i r r i below l a c r i m a l o f s u b o r b i t a l r i d g e . About two t u b e r c l e - l i k e c i r r i near lower borde r o f s u b o r b i t a l s t a y . S i m p l e , t h i n c i r r i near p o s t e r i o r l i m i t o f s u b o r b i t a l s t a y . L a r g e , t h i n , b road , weak ly m u l t i f i d c i r r i on upper operculum. C i r r i under lower jaw l o n g e r , s l i g h t l y f l a t t e n e d . C i r r i on a n t e r i o r face o f lower jaw s m a l l , t u b e r c l e - l i k e . C i r r i on m a x i l l a r y l a r g e r , f a i r l y b r o a d , e n t i r e . Many c i r r i s c a t t e r e d a long l a t e r a l l i n e ; many inconsp i cuous , s m a l l . Sometimes some, i n c l u d i n g one hear l a s t pore on cauda l f i n , a re l a r g e r , conspicuous and w h i t e . S c a l e s w i t h up t o twenty s p i n u l e s . A n t e r i o r l y t h e d o r s a l band o f s c a l e s occupies t he upper t h r e e - q u a r t e r s o f t h e space between d o r s a l f i n and l a t e r a l l i n e . The i n t e r v e n i n g naked space widens q u i c k l y above cu rva tu re o f l a t e r a l l i n e and then narrows toward cauda l pedunc le . D o r s a l s c a l e band separa te from s p i n y d o r s a l f i n by about one-quar te r o f the w i d t h o f t he band. The band then converges t o almost touch t h e p o s t e r i o r r ays o f t h e sof t d o r s a l f i n . H o r i z o n t a l rows o f s c a l e s on d o r s a l band s t r a i g h t except f o r t he bottom two rows which t u r n toward l a t e r a l l i n e near base o f cauda l f i n . S c a l e s v e r y crowded, l a r g e , 16 t end t o ove r l ap o the r s c a l e s . o n same and adjacent rows. The upper and s i x t h rows from bottom become reduced under s p i n y d o r s a l f i n w h i l e t h e f o u r t h and f i f t h rows are reduced and absent toward the end o f so f t d o r s a l f i n . T h i r d row i s reduced on c a u d a l pedunc le . F i r s t , second, and t h i r d rows cont inuous ac ross cauda l pedunc le . F i r s t row o f s c a l e s on d o r s a l band w i t h s c a l e s r a i s e d on d o r s a l s i d e o n l y w h i l e the o ther rows w i t h s c a l e s r a i s e d on a l l s i d e s . V e n t r a l band about two rows o f c l o s e l y spaced s c a l e s p o s t e r i o r t o t i p o f c l e i t h r a l s p i n e and upper base o f p e c t o r a l f i n . Band widens d i r e c t l y over anus and d i m i n i s h e s t o two d i s t i n c t rows o f s c a l e s on cauda l pedunc le . Transverse and h o r i z o n t a l s c a l e rows d i s t i n c t but anastamose over anus. Band d i s t i n c t l y separa ted from l a t e r a l l i n e by diameter o f two s c a l e s a n t e r i o r l y and h a l f a s c a l e p o s t e r i o r l y . V e n t r a l band o f s c a l e s separa ted from a n a l f i n by about two band w i d t h . A few v e r y l a r g e s c a l e s i n a x i l o f p e c t o r a l f i n . L a t e r a l l i n e curves upwards from above anus and o r i g i n o f a n a l f i n ; meets angle o f s u p r a c l e i t h r u m a n t e r i o r l y , and b i s e c t s t a i l i n t o equa l ha lve s p o s t e r i o r l y . O r i g i n o f s p i n y d o r s a l f i n d i r e c t l y over most a n t e r i o r p e c t o r a l f i n r a y . F i f t h sp ine o f d o r s a l f i n over p o s t e r i o r end o f depressed operculum. F i r s t t h r ee sp ines o f almost equa l he igh t but t w o - t h i r d s he igh t o f next fou r s p i n e s . F i r s t s p i n e does not extend beyond second d o r s a l s p i n e . Spines behind t he seventh become p r o g r e s s i v e l y s h o r t e r . Marg in o f f i n membrane modera te ly i n c i s e d between f i r s t s i x s p i n e s , deeply i n c i s e d between t h i r d and fou r th s p i n e . Most o f t h e so f t d o r s a l rays almost t h e same h e i g h t , t h e f i f t h t o n i n t h rays be ing t a l l e s t . Rays on e i t h e r s i d e become p r o g r e s s i v e l y s h o r t e r . Marg in o f membrane s l i g h t l y i n c i s e d between f i r s t four teen r a y s . Depressed d o r s a l f i n rays reach l e v e l o f base o f cauda l f i n r a y l e t s . O r i g i n o f a n a l under f i f t h so f t d o r s a l f i n r a y . L a s t a n a l f i n r ay under penu l t ima te d o r s a l f i n r a y . Margin o f f i n membrane i n c r e a s i n g l y i n c i s e d between r a y s , a n t e r i o r t o four t een th f i n 17 r a y . A n a l f i n rays appear o f s i m i l a r s i z e but the t w e l f t h ray i s about the l o n g e s t . D o r s a l and a n a l f i n rays become branched i n specimens above 90 mm. P e c t o r a l f i n rounded; e i g h t h o f n i n t h r ay from bottom l o n g e s t . P e l v i c s s h o r t , second ray l o n g e s t ; margin o r f i n membrane i n c i s e d . C o l o u r p a t t e r n dark above and l i g h t below; o therwise v a r i a b l e . Many b lo t ches and spots on d o r s a l s i d e . F ine s p e c k l i n g s p roduc ing dusky b lo t ches on c h i n , p e l v i c s , and behind p e c t o r a l f i n on body. Back w i t h four bars approx imate ly between base o f f i f t h and e i g h t h d o r s a l f i n s p i n e s , between t h i r d and seventh so f t d o r s a l f i n r a y s , between e leven th and four t een th sof t d o r s a l f i n r a y s , and between s i x t e e n t h and l a s t so f t d o r s a l f i n r a y s . I r -r e g u l a r ba r across cauda l f i n base and o thers across rays p o s t e r i o r t o t h i s . A n a l f i n w i t h s e v e r a l dark and l i g h t s t r e aks at r i g h t angles t o a n a l f i n r a y s . D o r s a l and p e c t o r a l f i n s w i t h i r r e g u l a r dark mark ings . L a t e r a l l i n e c i r r i may be consp icuous ly w h i t e . Leng ths - t o at l e a s t 203 mm. S p i n o s u s s - meaning " s p i n y " . Ranges- t ha t o f t he subgenus. 18 Tab le I . D i s t r i b u t i o n Records o f H . spinosus Reference o r L o c a t i o n c o l l e c t i o n No. Post l a r v a e 38°G3«50"N. 123°33 T 30"W. 51°N. 130°W. . (22-26 mm) (Northernmost record) F o l l e t , 1952 BC 60-170 J u v e n i l e s and A d u l t s C a l i f o r n i a ; 36°31»07"N. 36°37»35"N. 36°38»05"N. 37°48»23"N. 37°50«30»N. 37°50*40"N. 39°36»17"N. 40°01»05"N. 41°44» 57"N. 121 057 ?14"W. 121°53V50"W. 121°53 ,50»W. 122025T34"W. 122°51»00"W. 122°51»00"W. 123047 f17"W. 124°05'00"W. ( P a c i f i c Grove) (Monterey Bay) (Monterey Bay) (San F ranc i sco ) ( B o l i n a s P t . ) . ( B o l i n a s P t . ) ( S . o f B u s h e l P t . ) ( P t . Delgada) (Crescent C i t y ) B o l i n , 1944 124°12 t28wW, Santa Barbara I s . (Southernmost record) Sah F r a n c i s c o Bay Humboldt Bay Monterey and San F r a n c i s c o Oregon ; -43°18»N. 44°41 f N. 44°47 t N. 45°46»N. 45°55»N. Wash ing ton ; -Cape Johnson 124°24*W. 124°05»W. 124°05*W. 123°58«W. 123°58*W. (Cape Arago) (Yaqulna Head) (Cape Foulweather) (Cape Falcon) ( E c o l a Rocks) B r i t i s h C o l u m b i a ; -*Sooke, Vancouver I s . *13 m i l e s o f f S idney I n l e t *Ba ja P t . Nootka I s . *Ovouklnsk I n l e t near Burnsby I s . « « tt n n n « S t a r k s & M o r r i s , 1907 G i r a r d , 1856 tt Jordan & G i l b e r t , 1881 S c h u l t z & D e l a c y , I936 tt tt tt tt Hubbs, 1928 BC 62-495 ( i d e n t i f i e d by author) Clemens & W i l b y , 1949 BC 53-92 (uncata logued a t 1 . 0 . F . ) 19 Subgenus Hemilepidotus C u v i e r Hemi lep ido tus C u v i e r , 1829, p . 165 (Type o f genus Cot tus hemi lep ido tus T U e s i u s ) ; 1834> p . 141; Jo rdan and Evermann, 1898, p . 1934; Jo rdan and S t a r k s , 1904, p . 254; Schmidt , 1929, p . 360; So lda tov and L i n d b e r g , 1930, p . 199; Rendah l , 1931, p . 36; B o l i n , 1944* p« 13; Watanabe, 1958, p . 246; I960, p . 48. T e m n i s t i a R i c h a r d s o n , 1836, p . 59 (genotype B l e p s i a s v e n t r l c o s u s E s c h s c h o l t z = Cot tus hemi lep ido tus T i l e s i u s ) . D i a g n o s i s ? - The notched s p i n y d o r s a l f i n , four or f i v e h o r i z o n t a l rows i n d o r s a l s c a l e band, and s i x o r more h o r i z o n t a l rows i n v e n t r a l s c a l e band (a t w ides t p o i n t ) d i s t i n g u i s h e s t h i s subgenus from the r e l a t e d subgenera. D e s c r i p t i o n s - A l l bands o f s c a l e s w i t h s i m i l a r s i z e d s c a l e s . Four or f i v e h o r i z o n t a l rows o f s c a l e s i n p o r t i o n o f d o r s a l s c a l e band under s p i n y d o r s a l f i n . V e n t r a l band o f s c a l e s w i t h seven t o n i n e h o r i z o n t a l rows o f s c a l e s at w ides t p o i n t . A n a l s c a l e row d i scon t inuous i n area above l a s t depressed a n a l f i n ray o f p o s t - l a r v a e ; may or may not be present i n a d u l t s . No obvious prominences on top o f head a l though f i n e r i d g e s , g r a n u l a t i o n s , or s t r i a e r a d i a t e from low e l e v a t i o n s o f p o s t o c u l a r and o c c i p i t a l a reas . G i l l membranes u n i t e d , fused t o isthmus but w i t h a w e l l developed f ree f o l d behind at tachment . T h i r d spinous r ay on d o r s a l f i n u s u a l l y s h o r t e r than second or f ou r th spinous r a y s . Marg in o f f i n membrane g r e a t l y i n c i s e d between t h i r d and f o u r t h spinous r a y s , thus forming an obvious no tch i n the p r o f i l e o f the spinous d o r s a l f i n . Four spec i e s a re known i n t h i s subgenus. Range ; - Monterey Bay , C a l i f o r n i a ; t o B e r i n g Sea ; t o Hakodate, Japan. Hemilep i d o t u s ; - d e r i v a t i o n o f name i s t ha t o f genus. 20 Synopsis of the species of the subgenus Hemilepidotus A Dorsal and anal rays branched. Nasal c i r r i well developed with many branches, 36 or less vertebrae. H. hemilepidotus p. 24 AA Unbranched dorsal and anal fin rays. Nasal c i r r i simple or, at most, two branches, 36 or more vertebrae. B First spine on dorsal fin shorter than second spine. Lowest preopercular spine usually simple, sharp, round in cross-section. C Fifty-eight or fewer lateral line pores. At least one scale, usually two to six scales, between dorsal scale band and lateral line on caudal peduncle. Pelvic fin of male may extend past tip of pectoral fin. CC Fifty-nine or more lateral line pores. Practically never any scales between dorsal scale band and lateral line on caudal peduncle. . Pelvic fin of male never with large cteni, and never extend past tip of pectoral fin. H. .jordani p.45 BB First spine longer than second spine, even when spines are depressed. Lowest preopercular spine flattened and develops secondary barbs. Frontal c i r r i vestigial or absent. H. gilberti . . . . . . . . . . p . 56 10 M M (COLL. NO. BC. 63-ioiq) • :— 1 POSTLARVA OF H. HEMILEPI DOTUS Figure 4.. ? POSTLARVA OF HEMILEPIDOTUS SP Figure 5.. FEMALE:- COLL. NO. BC 63-882. " , SO_MM. H E M I L E P I D O T U S H E M I L E P I D O T U S . Figure 6. 24 Hemilep idotus (Hemi lep ido tus ) hemi lep ido tus ( T i l e s i s u ) ( T i g , 4 and 6, maybe 5) Cottus hemi lep ido tus ( T i l e s i u s , 1810, p . 262, f i g , ( " P e t r o p a v l o v s k , Kamchatka" - quote from B o l i n 19,44). Cottus t r a chu rus P a l l a s , 1814, p . 138 ( i n p a r t ) , Hemi lep idotus t i l e s i i C u v i e r and V a l e n c i e n n e s , 1829, p . 276, p i . 85 ( a f t e r T i l e s i u s and P a l l a s ) , B l e p s i a s v e n t r i c o s u s E s c h s c h o l t z , 1829, p<> 4 , p l « 13 ( N o r f o l k Sound and S i t k a ) . T e m n i s t i a v e n t r i c o s u s R i c h a r d s o n , 1836, p . 59» Hemilep idotus t r achurus Gunther , i 8 6 0 , p . 173 (probably i n p a r t ) ; Jordan and G i l b e r t , 1882, p . 7 1 5 . Hemi lep ido tus g i b b s i i G i l l , 1863, p , 13, Hemi lep ido tus hemi lep ido tus Jordan and Evermann, 1896, p . 439; 1898, p . 1935; 1900. f i g . 705; Jordan and G i l b e r t , 1899, p . 457; Schmidt , 1904, p . 113 ( i n p a r t ) ; 1929, p . 36O ( i n p a r t ) ; ; Evermann and Goldsborough, 1907, p . 304, f i g . 62 ; G i l b e r t and Burke , 1912, p . 53; Evermann, 1930, p . 573; So lda tov and L l n d g e r g , 1930, p . 202j Rendah l , 1931* p . 32; S c h u l t z and Welander, 1934s p« 5; B o l i n , 1944# p . 17, f i g . 5; Clemens and W i l b y , 1949, p . 244, f i g . 1961, p . 284, f i g ; Gorbunova, 1964, p . 241. The specimen drawn i n F igu re 5 i s t e n t a t i v e l y i d e n t i f i e d as H . h e m i l e p i d o t u s , P a l l a s , 1814, r e f e r s t o specimen from America spo t ted on under sur face - see Sherborn , 1934, and B u l l . Z o o l . Nomen., 1950, p , 403 , f o r p u b l i c a t i o n date o f P a l l a s • D i a g n o s i s ; - The branched d o r s a l and a n a l f i n r a y s , the 36 o r fewer v e r t e b r a e , t h e many f r i n g e d n a s a l c i r r i , t he 6 o r 7 p y l o r i c caecae and t he frequent p e r -s i s t ance o f a f i f t h s c a l e row i n t he d o r s a l s c a l e band are d i s t i n c t i v e fea tures w i t h i n t he subgenus. M a t e r i a l s ; - The d e s c r i p t i o n i s based, on 30 specimens rang ing from 41 t o 235 mm i n s tandard l e n g t h . These are from c o l l e c t i o n Nos, BC 63-1068, IO69, 1070, 1073 and IO76. D e s c r i p t i o n s on s e x u a l d i f f e r e n c e s a l s o i n c l u d e c o l l e c t i o n Nos. BC 54-450A; BC 61-649; BC 62-898; 547? BC 63-121, 782, 910, 1035 and 1077. Samples i n the I n s t i t u t e o f F i s h e r i e s 25 from throughout t he range were checked aga ins t t he d e s c r i p t i o n s . D e s c r i p t i o n ; - D o r s a l ; X I , 19 (18-20) [ 1 7 ] 5 9 . A n a l ; 15 (13-16). P e c t o r a l ; 16 ( 1 5 - 1 7 ) . V e r t e b r a e ; 3 5 ( 3 4 - 3 6 ) . L a t e r a l l i n e po re s ; 64 .7(60-68) [ 5 9 ] 1 1 . Transverse s c a l e rows i n d o r s a l band; 68.9 (64-75). Number o f s c a l e s i n row above l a t e r a l l i n e ; 18 .2 (9-27) [ 7] « Number o f h o r i z o n t a l s c a l e rows i n d o r s a l band; 4-5, P y l o r i c caeca ; 6-7. Width immedia te ly behind p e c t o r a l f i n s i n specimens under 100 mm; 180 (149-198) (16 specimens) . Width immediate ly beh ind p e c t o r a l f i n s i n specimens over 100 mm; 234 (194-271) (14 specimens) . Depth from o r i g i n o f d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i n s ; 270 (240-308)»•« Depth from o r i g i n ' o f sof t d o r s a l f i n t o o r i g i n o f a n a l f i n ; 231 (204-256). Head l e n g t h ; 369 (334-400), O r b i t d iameter i n specimens over 100 mm; 82 (70-90) (14 specimens) . O r b i t -d iameter i n specimens under 100 mm; 107 ,(97rl25) (16 specimens) . O r b i t t o end o f o p e r c l e ; 187 (164-234). I n t e r o r b i t a l widths 61 (53-72). Snout l e n g t h ( e x c l u d i n g p r e m a x i l l a r y ) s 76 (68-90). Narrowest l a c r i m a l w i d t h i n specimens above 100 mms 41 (37-45) (14 spec imens) . Narrowest l a c r i m a l w i d t h i n specimens below 100 mm; 33 (29-37) (16 specimens) . Cauda l peduncle l e n g t h ; 83 (62-105), Cauda l peduncle depth ; 72 (68 -82) . Snout t o o r i g i n o f d o r s a l f i n ; 327 (314-360). Snout t o o r i g i n o f a n a l f i n i n specimens above 100 mm: 600 (553-641) (14 spec imens) . Snout t o o r i g i n o f a n a l f i n i n specimens under 100 mm: 572 (551-584) (16 spec imens) . Height o f longes t sp ine on f i r s t d o r s a l f i n : 119 ( 9 3 - 1 3 8 ) . Length o f longes t sof t d o r s a l f i n r a y : 168 (143-196). Length o f a n a l f i n base : 323 (297-363) (28 specimens) . Length o f longes t ray on a n a l f i n : 135 (124-189). Length o f base o f p e c t o r a l f i n s 152 (132-180) . Length o f longes t r a y on p e c t o r a l f i n s 278 (243-319). Length o f longes t r ay on p e l v i c f i n i n specimens below 100 mms 212 (195-225) (16 spec imens) . Length o f l onges t r ay on p e l v i c f i n i n 13 females above 100 mm: 26 209 (176-22?) ( f rom c o l l e c t i o n Nos. BC 63-1068, 1069, 1070, 1073). Length o f longes t r ay on p e l v i c f i n In 12 females above 100 mms 222 (200-240) (From the c o l l e c t i o n s o ther t han the above) . Length o f longes t r a y on p e l v i c f i n i n 10 males above 100 mm; 247 (234-271). Length o f longes t r ay on cauda l f i n s (212-216), ' Head depressed . I n l a t e r a l v i e w , p r o f i l e o f o c c i p u t s t r a i g h t and s t e e p l y s l o p i n g toward snout . T h i s p r o f i l e i f extended would f o l l o w pa th through upper p o r t i o n o f p u p i l . O r b i t g r e a t l y elevatedo Snout s t r a i g h t a n t e r i o r l y , convex d o r s a l l y ; s l i g h t depress ion near a n t e r i o r i n t e r o r b i t a l a r e a . Snout . s l o p i n g s c h i n s l o p i n g toward i s thmus . In p r o f i l e from d o r s a l v i e w , m a x i l l a r y and p r e m a x i l l a r y s l i g h t l y rounded. P r o f i l e o f p r e o p e r c l e convex, Operc le ove r l aps p e c t o r a l f i n base . Head broado Body deepest under f i r s t sp ine o f d o r s a l f i n . Nape r a i s e d and convex. D o r s a l base s t r a i g h t from i t s o r i g i n t o cauda l pedunc le . D o r s a l and v e n t r a l p r o f i l e o f c a u d a l peduncle s i m i l a r l y concave. A n a l f i n base convex. P r o f i l e from a n a l f i n t o base o f p e l v i c f i n s e i t h e r s t r a i g h t o r convex. P r o f i l e from base o f p e l v i c f i n s t o isthmus s t r a i g h t o In d o r s a l v i e w , l a t e r a l p r o f i l e s t r a i g h t t o . na r row cauda l peduncle (except when stomach d i s tended w i t h food o r eggs ) . A n t e r i o r n o s t r i l s h o r t , t u b u l a r , d i r e c t l y l a t e r a l t o a n t e r i o r base o f n a s a l sp ine and d i r e c t l y a n t e r i o r t o d o r s a l margin o f o r b i t . D i s t a l p o r t i o n f l a r e d and w i d e r t h a n base . P o s t e r i o r margin o f n o s t r i l r a i s e d s l i g h t l y h ighe r t h a n a n t e r i o r s i d e . P o s t e r i o r n o s t r i l i n l i n e w i t h n a s a l sp ine and d o r s a l border o f o r b i t . D i s t a l margin s l i g h t l y c o n s t r i c t e d . Conspicuous head p o r e , same diameter as n o s t r i l , at a n t e r o l a t e r a l base o f p o s t e r i o r n o s t r i l . Many o ther s m a l l pores on o c c i p u t , i n t e r o r b i t , l a c r i m a l and cheek. O r b i t d iameter becomes r e l a t i v e l y s m a l l e r w i t h con t inued growth . Eye incomple t e ly f i l l s o r b i t 27 i n l a r g e r specimens.. I h t e r o r b i t h i g h l y concave., L o n g i t u d i n a l r i d g e s i n i n t e r o r b i t p r e s e n t . G i l l membranes u n i t e d , fused t o isthmus t o form a r e l a t i v e l y narrower f ree f o l d . Head sp ines appear h e a v i e r and more b l u n t , e s p e c i a l l y i n l a r g e r specimens. N a s a l sp ines s t r o n g , prominent . Upper p r e o p e r c u l a r sp ine same s i z e o r maybe longe r than second p r e o p e r c u l a r s p i n e , a p p r o x i m a t e l y , a p u p i l d iameter i n l e n g t h . Upper sp ine d i r e c t e d p o s t e r o - d o r s a l l y , second sp ine d i r e c t e d p o s t e r i o r l y . E s p e c i a l l y i n l a r g e r specimens, t h i r d sp ine q u i t e shor t and s t o u t ; may b a r e l y p r o j e c t th rough t h i c k integument. Rugose s t r i a t i o n s become s t r o n g e r w i t h age and r a d i a t e from p o s t o r b i t a l and o c c i p i t a l areas o f s k u l l . Low r i d g e s on d o r s a l margin o f o p e r c u l a r bone, l a t e r a l f r o n t o - p a r i e t a l , p t e r o t i c and p o s t - t e m p o r a l , and s u p r a c l e i t h r a l a reas . A l l r i d g e s , except o p e r c u l a r , become h e a v i l y rugose i n l a r g e specimens. C i r r i r e l a t i v e l y b road and more m u l t i f i d . The number o f f r i nges on c i r r i i nc rease w i t h s i z e . N a s a l c i r r i l a r g e w i t h up t o about a dozen f r i n g e s . P o s t o c u l a r c i r r i i n a n t e r o - l a t e r a l p lane and w i t h up t o about e igh teen f r i n g e s . F r o n t a l c i r r i m u l t i f i d and immediate ly p o s t e r o - m e d i a l t o p o s t o c u l a r c i r r i . C i r r i on occ ipu t broad and h i g h l y m u l t i f i d . L a c r i m a l p o r t i o n o f s u b o r b i t a l w i t h approx imate ly four f l e s h y t u b e r c l e - l i k e c i r r i , two s i m i l a r c i r r i near lower border o f s u b o r b i t a l s t a y . C i r r i near p o s t e r i o r l i m i t o f s u b o r b i t a l s t a y . C i r r i at upper p o r t i o n o f operculum, m u l t i f i d . C i r r i near c h i n pores s m a l l and b a r b e l - l i k e . C i r r i on lower jaw s m a l l . C i r r i on m a x i l l a r y broad a n d . l a r g e , i t s w i d t h .67 ( « 5 0 - . 8 3 ) i o f i t s l e n g t h . S c a l e s i n d o r s a l band t end t o be more rounded o r s e m i - c i r c u l a r on t h e i r d o r s a l s i d e . D o r s a l s c a l e band conf ined w i t h i n t he d o r s a l h a l f o f t he i - • - ' i d i s t a n c e between l a t e r a l l i n e and d o r s a l f i n . T h i s band separated from s p i n y d o r s a l f i n by about a h a l f o f t he band ' s w i d t h and then converges t o meet p o s t e r i o r p o r t i o n o f so f t d o r s a l f i n . : H o r i z o n t a l s c a l e rows s t r a i g h t fo r t he 28 most p a r t . However, p o s t e r i o r l y t he l a s t one o r two s c a l e s on lowest row of ten p l a c e d a l i t t l e lower than t he pa th f o l l o w e d by the r e s t o f t he s c a l e row. Often one o r two s c a l e s on cauda l pedunc le , midway between l a t e r a l l i n e and d o r s a l s c a l e band. S c a l e s crowded and of ten over lap o t h e r s , e s p e c i a l l y those on same row. A n t e r i o r l y , s c a l e s i n bottom four rows o f d o r s a l band o f equa l s i z e . Sca les on f i f t h row u s u a l l y l e s s than o n e - h a l f s i z e o f s c a l e s i n lower rows o f t h i s band. P e r s i s t e n c e o f dorsalmost s c a l e row v a r i a b l e , but more than a h a l f dozen s c a l e s are commonly present under s p i n y d o r s a l f i n . The next row i s reduced i n s i z e p o s t e r i o r l y and p e r s i s t s as f a r as the p o s t e r i o r t h i r d o f so f t d o r s a l f i n . The vent ra lmost row disappears before r each ing c a u d a l pedunc le , w h i l e t he remaining rows extend pas t d o r s a l f i n on cauda l pedunc le . Sca le s on lowest row r a i s e d on upper s i d e on ly w h i l e t h e upper rows are r a i s e d on a l l s i d e s . Tubu la r l a t e r a l l i n e bony elements w i t h s p i n u l e s on up t o p o s t e r i o r t h r e e - q u a r t e r s o f d o r s a l - s u r f a c e . V e n t r a l band approx imate ly two o r t h r ee h o r i z o n t a l s c a l e rows wide oppos i t e d o r s a l base o f p e c t o r a l f i n . Above anus and o r i g i n o f a n a l f i n t h i s band expands q u i c k l y t o s i x o r seven more r e g u l a r rows. Band d imin i she s t o one or two i r r e g u l a r rows on cauda l pedunc le . Rows tend t o anastamose w i t h t h e t a p e r i n g o f t h e band. The a n a l s c a l e row v e r y r a r e l y p e r s i s t i n a d u l t s as ex t remely reduced s c a l e s . Band separa te from l a t e r a l l i n e by two s c a l e diameters a n t e r i o r l y , and o n e - h a l f a s c a l e p o s t e r i o r l y . Band separate from a n a l f i n by approximate ly t he w i d t h o f t he band. L a t e r a l l i n e cu rva tu r e approx imate ly above anus and a n a l f i n . S e v e r a l c i r r i s c a t t e r e d on l a t e r a l l i n e . O r i g i n o f s p i n y d o r s a l f i n over base o f lowest p e c t o r a l r a y . F i f t h o r s i x t h d o r s a l f i n sp ine t a l l e s t . F i r s t sp ine seldom extends beyond t i p o f second sp ine o f d o r s a l f i n . T h i r d sp ine s h o r t e r than second s p i n e . F i f t h 29 sp ine over p o s t e r i o r l i m i t o f operculum. Spines become p r o g r e s s i v e l y s h o r t e r beh ind s i x t h d o r s a l f i n s p i n e . Marg in o f membrane on d o r s a l f i n i n c i s e d between f i r s t seven s p i n e s . Height o f membrane on a n t e r i o r s i d e o f f ou r th sp ine i s .67 (»55-°90) i n he igh t o f t h i r d s p i n e . S i x t h t o seventh so f t d o r s a l f i n rays t a l l e s t . F i n margin between t h i r d and four t een th rays almost s t r a i g h t . Other rays become p r o g r e s s i v e l y s h o r t e r on e i t h e r s i d e . D o r s a l f i n rounded p o s t e r i o r l y . Depressed d o r s a l f i n rays extend t o base o f upper -most cauda l f i n r a y l e t s or cauda l f i n base . P o s t e r i o r rays o f d o r s a l and a n a l f i n branched i n specimens above 140 mm. O r i g i n o f a n a l f i n under about t h e f i f t h s o f t - d o r s a l f i n r a y . Las t a n a l f i n ray under about t he penu l t ima te d o r s a l f i n r a y . Marg in o f f i n membrane deeply i n c i s e d between f i r s t t h i r t e e n a n a l f i n r a y s . Approx imate ly seventh o r e igh th a n a l f i n ray l o n g e s t . F i n rays p r o g r e s s i v e l y s h o r t e r on e i t h e r s i d e a l though t i p s o f t he m a j o r i t y o f f i n rays appear i n s t r a i g h t l i n e . Cauda l f i n s l i g h t l y rounded t o t r u n c a t e . P e c t o r a l f i n rounded; margin o f f i n membrane i n c i s e d most deeply between t e n lowest r a y s . E i g h t h t o e l even th r ay from lowest ray o f p e c t o r a l f i n l o n g e s t . The lower rays s h o r t e r and t h i c k e r than upper p e c t o r a l r a y s . P e l v i c f i n w i t h i n c i s e d margin o f membrane between r a y s . P e l v i c f i n rays t end t o be l o n g e r i n males . Rudimentary e l e v a t i o n s , p robab ly r e p r e s e n t i n g v e s t i g i a l c t e n i , p resent i n some males . A r i d g e o f p a p i l l a t e s k i n present on the lower m e d i a l su r face o f p e l v i c rays i n mature males . I n o ther specimens p e l v i c f i n rays and t i s s u e smooth. C o l o u r v a r i a b l e w i t h r e d , brown, w h i t e and b l a c k , however red p r e -domina t ing . Darker on d o r s a l su r face and almost w h i t e v e n t r a l l y . Body mot t l ed or spo t t ed a l l ove r , s p o t t i n g conspicuous over l i g h t v e n t r a l s u r f a c e . Spots always evident on c h i n and g i l l membranes i n specimens above 60 mm. Spots i r r e g u l a r and d i f f i c u l t t o see i n s m a l l e r specimens. Four i r r e g u l a r bars across back u s u a l l y p r e s e n t . F i r s t between approx imate ly t he t i p o f 30 t h e fou r th and seventh d o r s a l f i n sp ines t o the l a t e r a l l i n e , t he second between t he t h i r d and s i x t h sof t d o r s a l rays t o the l a t e r a l l i n e and then i r r e g u l a r l y t o a shor t d i s t a n c e above the o r i g i n o f a n a l f i n , t he t h i r d ba r between n i n t h t o t w e l f t h d o r s a l r ay i n i r r e g u l a r course t o n i n t h or t e n t h a n a l r a y , and t he f o u r t h ba r between s i x t e e n t h and l a s t sof t d o r s a l r ay t o l a t e r a l l i n e where i t fo rks i r r e g u l a r l y t o between t h i r t e e n t h and l a s t a n a l f i n r ay and t he o ther branch passes t o middle o f v e n t r a l border o f cauda l pedunc le . The bars t end t o reach margin o f d o r s a l f i n i n a p o s t e r i o r l y d i r e c t e d s l a n t . A dark a rea may be observed below f i r s t t h ree sp ines o f d o r s a l f i n . Cauda l f i n rays da rke r t han membrane, may have t h i n , v e r t i c a l , evenly spaced, l i g h t b a r s . In da rke r specimens the above p a t t e r n i s obscured. A n a l f i n l i g h t or da rk , m o t t l e d . Both s ides o f p e c o t r a l f i n s p o t t e d , w i t h i n d i c a t i o n s o f t h r e e or four dark v e r t i c a l b a r s . P e l v i c s u s u a l l y w h i t e or maybe s p o t t e d . In some l a r g e males t he p e l v i c f i n s become d a r k l y s p o t t e d , o r j e t b l a c k , e s p e c i a l l y a long the p a p i l l a t e d sur face o f the p e l v i c r a y s ; b l a c k patches and p a t t e r n s occur between the r a y s . The med ia l or p o s t e r i o r sur face o f the p e c t o r a l f i n s a l s o w i t h v e r y dark t o j e t b l a c k bars and pa tches . The p a t t e r n can spread across the a x i l o f f i n where l i g h t patches p r o v i d e s t r i k i n g con t r a s t t o the b l a c k . L e n g t h ; - t o about 300 mm. H e m i l e p i d o t u s s - D e r i v a t i o n o f name i s same as t ha t f o r genus. Ranges- Monterey B a y , C a l i f o r n i a t o P r i b i l o f I s l ands and t o Kamchatka. Geographic v a r i a t i o n i s not g r e a t . F i g . 27 and 28 i l l u s t r a t e minor d i f f e r e n c e s i n s c a l e counts between samples from the P r i b i l o f I s l a n d s , Umnak I s l a n d and Vancouver I s l a n d . More specimens from the coast o f the U n i t e d 31 Sta tes are needed t o complete a v a r i a t i o n a l s tudy . Tab l e I I . D i s t r i b u t i o n Records o f H . hemi lep ido tus L o c a t i o n Reference or c o l l e c t i o n No. Post l a r v a e P ioneer Seamount, C a l i f o r n i a * 52°32»N. 176°34 ?W. ( l e n g t h 13 mm doubt i d e n t . ) * 55°39*N. 170°00»W. ( l e n g t h 13 mm doubt i d e n t . ) * Amchitka I s . , A l a s k a ( l e n g t h 15-22 mm) * S t . P a u l I s . , A l a s k a ( l e n g t h 21 mm approx . ) * L i t t l e Konius I s . , Shumagin I s . , A l a s k a ( l e n g t h 20-31 mm) * Caton I s . near Sonak I s . , A l a s k a ( l e n g t h 22 mm) * 55°53?N. 165°33 ? W . ( l e n g t h 20-22 mm) J u v e n i l e s and A d u l t s C a l i f o r n i a San F r a n c i s c o H a l f Moon Bay 36°37»20»N. 121°54 T15"W. S. end o f Monterey Bay (most southern pa r t o f known range) Hanna, 1951 BC 62-847 BC 62-908 BC 63-1010 BC 63-1296 BC 63-1303 BC 63-1306 BC 63-1331 Jordan & G i l b e r t , 1881 Evermann, 1930 B o l i n , 1944 B o l i n , 1944 Oregon i-43°18'N. 43°22»N. 44°41 f N. 44°47fN. 45°28*N. 45°46»N. 45°55yN. 124°24fW. 127°17yW. 124°05tW. 1 2 4 ° 0 5 t w . 123°58*W. 123°58*W. 123°58»W. (Cape Arago) (Coos Bay) ( Y a q u i n a Head (Cape Foulweather) (Oceanside) (Cape Falcon) ( E c o l a Rocks) S c h u l t z & D e l a c y , 1936 n « it tt it tt Washingtons-47°52»N. 124°35 'W. 47°21 'N . 124°18»W. 47°18»N. 124°15»W. Cape Johnson Por t A n g e l i s Puget Sound S e a t t l e San Juan I s l a n d Neah Bay *Peak Bay ( L a Push) (Cape. E l i z a b e t h ) ( P t . G r e n v i l l e ) -tt tt it tt Meek, 1899 Jordon & G i l b e r t , 1881 K i n c a i d , 1919 Wismer & Swanson, 1935 Jordon & Jouy , 1882 BC 60-238 32 L o c a t i o n B r i t i s h Columbia ( southern) * Sooke, (Agate Beach) * " " » (depth 50 f t . ) * " " " (depth 50 f t . ) * San Juan P t . near P t . Renfew ( i n t e r t i d a l ) * Brooks Penn insu la ( i n t e r t i d a l ) * Fa l s e Narrows, Vancouver * S t a n l e y Pa rk , Vancouver * tt tt tt * Boyer I s . Howe Sound (depth 30-45 f t . ) * tt tt tt tt tt tt * J e r v i s I n l e t * Mouth o f F rase r R . * Saturna I s . 3JC tt tt * James I s . * W h y t e c l i f f * Depar ture Bay * Seymour I n l e t ( S . E . Arm) B r i t i s h Columbia (nor thern)%-* Goose I s . *" " * tt tt *Bute I n l e t *Work Channe l , Head o f T r a i l Bay *Hope I s . Ashby P t . *Go le t a s Channe l , N i g e i I s . T i p o f Boxer P t . *W. o f N a h w i t t i B a r , Queen C h a r l o t t e I s . (depth 72-90 f t . ) *Tasu Harbour , Queen C h a r l o t t e I s . Duncan Bay near P t . Simpson A l a s k a P a n h a n d l e i -N o r f o l k Sound and S i t k a Por t A l t h o r p Por t Etches (depth 72-96, f t . ) S i t k a H u n t e r ' s Bay L o r i n g *Auke Bay #it " * P o r t Armstrong *Port Herber t (depth 14-45 I t . ) * L i t t l e Por t Walker -* » « « (depth 3 I t . ) * I c y S t r a i t s , Cross ..Sound * N o r t h S h e l t e r I s . * C r a i g (depth 8 f t . ) *Monas i t ka Bay Reference o r c o l l e c t i o n No. BC 62-495 BC 62-880 BC 62-898 BC 62-899 BC 63-506 BC 60-228 BC 53- 20 BC 56-212 BC 61-649 BC 63-910 BC 63-340 BC 63-936 BC 59-540 BC 57-214 BC 54-450A BC 63-1465 BC 62-902 BC 55-232 BC 61-591 BC 53-215 BC 53-224 BC 53-228 BC 59-471 BC 61-257 BC 63-783 BC 63-785 BC 63-782 BC 61-684 Bean, I884 E s c h s c h o l t z , 1829 Bean, 1881 Evermann & Goldsborough, 1930 tt BC 61-505 BC 62-966 BC 62-591 BC 63-1254 BC 63-256 BC 63-1263 BC 63-234 BC 62-803 BC 6 3 - 82 BC 63-1028 33 Location Gulf of Alaska to Alaskan Penninsula;-Karluk and Uyak Bay * Chignik Bay * 58°3'0»N. 153°30»W. * Cohen Is. Kachemck Bay (intertidal) * 58°55*N. 152°00»3O"'i. (Amatuli Is.) % tt ti tt Reference or. collection No. Evermann & Goldsborough, 1930 BC 62-646 BC 62-666 BC 62-989 BC 62-456 BC 62-457 Aleutian Islandss-Urialaska Is. Jordan & Gilbert, 1899 Kiska Is. it Adak Is. n Atka'Is. it Amchitka Is. it Attu Is. it Attu Is. and Agattu Is." Gilbert & Burke, 1912 * Sarichef Unimak Is. (tide pools) BC 63-1080 * tt tt it tt BC 63-1081 * Tigalda Bay, Tigalda Is. (depth 12 in.) BC 63-1311 * Cape Point, Unimak Is. (depth 100-125 ft.) BC 63-347 * Umnak, Umnak Is. (tide pools) BC 63-1076 * Ashishik Pt. Umnak Is. BC 63-1078 * Nikolski tt n tt BC 63-1073, 1071, 1070, 1069, 1068 * Otter Pt. » " - BC 63-1077 * Igitkin Is. BC 63-919 * Kulak Bay, Adak Is. (tide pools) BC 63-906 * Sweeper Cove,, Adak Is. BC 63-1003 * " . n it tt (depth 0-20 ft.) BC 63-1429 * Constant ine Harbour, Amchitka Is. BC 63-309 , 63-1035, 62-938 * Makarius Bay " " BC 63-1019 * Lorah Stn. " n BC 64-1014 *Buldir Is. (depth 4 ft.) BC 63-1410 * » » (depth 2 ft.) BC 63-1412 *•» " (depth 4 ft.) BC 63-1414 * " " (depth 4 ft.) : BC 63-1421 * South side of Shemya Is. (tide pools) BC 63-897 * East " " " " " " BC 63-898 * S.W. « 11 " » t» tt BC 63-900 * Casco Pt., Attu Is. "• » BC 63-S80, 883 * Massacre Bay, Attu Is. " " BC 63-882, 888, 1009, 1013 * Murder Pt. » " " " BC 63-885, 886, 1011 * Alexal Pt. " " BC 63-1004, 893 * Eckman Creek " " BC 63-904 Bering Sea?-* Amak Is, (tide" pools) * Izembek Bay (depth 0-3 ft.) *St. Paul Is. (northern most record-tide pools) BC 63-1313 BC 63-1433 BC 63-1458, 1461 34 Location U . S . S . R . ; -* Medney I s . , ( t i de pool) Medni I s . ( t i de pools) Kamchatka (Petropavlovsk) (•westernmost record) . Reference or c o l l e c t i o n No. BC 63-1043 Gi lbe r t & Burke, 1912 T i l e s i u s , 1810 Hemilepidotus (Hemilep idotus) zapus Gi lbe r t and Burke ( F i g s . 7 and 8) Hemilepidotus zapus G i lbe r t and Burke, 1912, p . 54> fig» 10 (At tu Is land, Alaska) ; Schmidt, 1929, p . 366; Soldatov and Lindberg, 1930, p . 205; Rendahl, 1931. Hemilepidotus g i l b e r t i zapus Gorbunova, 1964, p . 241 (maybe i n p a r t ) . The synonymy of H. zapus i s unwarranted. Gorbunova provides no evidence other than the s i m i l a r i t y of the l a rvae . Species of te leos ts are most d i f fe ren t ia ted at mature s i ze s , consequently data should be presented on the known characters of mature specimens before synonymy can be considered. The larvae are characterized as being more darkly pigmented. The smallest postlarvae (17 mm) used i n t h i s study are l i g h t (except for some dark pigment on back, close to dorsa l f i n ) . These specimens have the d i s t i n c t i v e l a t e r a l l i n e pore counts and scale characters of the adul ts . Gorbunova presents no extrapolat ion of why her larvae are the same form as the adul t s . In view of these discrepancies, the i d e n t i f i c a t i o n of her larvae can not be t rusted. H. g i l b e r t i zapus and H. g i l b e r t i g i l b e r t i are d i f fe ren t ia ted by Gorbunova wi th propor t ional measurements. Her data which i s p lo t ted i n F i g . 42, 43, 44, 45, 46, and 47 could hardly be used to iden t i fy these forms, because of the overlap of ranges for each character. Types:- Type specimen, a male, 127 mm, Albatross s tn . 4782, i n U . S . N . M . ; cotypes from Albatross s tns. 4778 and 4779, U.S.N.M. No. 70866. 35 36 37 D i a g n o s i s ; - The f i r s t sp ine s h o r t e r than second sp ine o f d o r s a l f i n , the prominence o f s c a l e s , o n the cauda l pedunc le , and 58 o r fewer l a t e r a l l i n e pores a r e ' t h e bes t d i s t i n g u i s h i n g f ea tu r e s . M a t e r i a l s : - Counts are based on p o s t - l a r v a e from c o l l e c t i o n Nos. BC 63-843, 848, 904, 907, 914; cotypes (44-58 mm i n s tandard l eng th ) from U . S . N . M . 70866$ and s m a l l adu l t (84 mm) from BC 63-1426. Remaining d e s c r i p t i o n i s based on a l l but the p o s t - l a r v a e . Specimen from BC 63-1426 i s judged t o be a male because o f t h e l o n g e r p e l v i c f i n s , however no attempt has been made t o make a m i c r o s c o p i c d i s s e c t i o n o f the gonads. Counts o f f i n rays and l a t e r a l l i n e pores were t aken from U . B . C . c o l l e c t i o n s and those from G i l b e r t and Burke (1912). The U . S . N . M . specimens are presumed t o be de-s c r i b e d i n G i l b e r t and B u r k e . D e s c r i p t i o n : - Dorsa l s X I ( X I I ) , 20-21 (22) . Anals 17 ( l 6 ) . P e c t o r a l s 16 (15-17). -Ver tebrae : 36 o r 37. L a t e r a l l i n e poress 52.5 (47-58). Transverse s c a l e rows i n d o r s a l bands 66.7 (62-71). Number o f s c a l e s i n row above l a t e r a l l i n e s 22.5 (16-27)o Number o f h o r i z o n t a l s c a l e rows i n d o r s a l bands 4 ( 5 ) o F y l o r i c caecas 4 (3 specimens) . Width immediate ly beh ind p e c t o r a l f i n s 176 (168-189). Depth from o r i g i n o f d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i n s s 254 (239-278). Depth from o r i g i n o f so f t d o r s a l f i n t o o r i g i n o f a n a l f i n s 213 (202-227). Head l e n g t h : 368 ( 358-386). O r b i t diameters 121 (114-134). O r b i t t o end o f o p e r c l e ; 172 (166-181), I n t e r o r b i t a l w i d t h : 50 (41-55). Snout l e n g t h : 70 (64-75). Narrowest l a c r i m a l w i d t h : 32 (29-36). Caudal peduncle l e n g t h : 84 (70-95). Cauda l peduncle dep th : 67 (63-72). Snout t o o r i g i n o f d o r s a l f i n : 321 (307-339). Snout t o o r i g i n o f a n a l f i n : 546 (520-579). Length o f base o f d o r s a l f i n : 659 (620-688). Height o f longes t sp ine on d o r s a l f i n s 126 (108-145) (6 spec imens) . Height o f longes t so f t r a y on d o r s a l f i n s 3 8 ( 1 7 0 - 1 8 6 ) ( 2 spec imens) . Length o f a n a l f i n bases 3 5 9 ( 3 4 1 - 3 8 1 ) . Length o f l onges t r ay on, a n a l f i n s 1 2 6 ( 1 1 3 - 1 3 8 ) . Length o f base o f p e c t o r a l f i n s 1 4 1 ( 1 3 0 - 1 5 8 ) . Length o f longes t r ay on p e c t o r a l f i n s 3 1 3 ( 2 9 2 r - 3 3 6 ) . Length o f longes t c auda l f i n rays 2 6 5 ( 2 5 2 - 2 8 6 ) . Length o f longes t p e l v i c f i n r ay on s m a l l e r specimenss 2 3 3 - 2 6 3 . Length o f longes t r ay on p e l v i c f i n o f l a r g e s t specimens 2 9 0 . Head r o b u s t , depth a l i t t l e g r ea t e r than w i d t h . I n l a t e r a l v i e w , occ ipu t s t r a i g h t and f o l l o w s pa th t ha t would pass through d o r s a l s i d e o f o r b i t . O r b i t s j u s t s l i g h t l y e l eva t ed above d o r s a l p r o f i l e . Snout s t r a i g h t and s lopes from n a s a l s p i n e s . Upper p o r t i o n o f snout convex and then s l o p i n g t o o r b i t . Gh in s t r a i g h t and s l o p i n g t o i s thmus . In v iew from d o r s a l s i d e p r e m a x i l l a e rounded but head appears s l i g h t l y p o i n t e d . P r o f i l e o f m a x i l l a and p r e o p e r c l e s t r a i g h t . Operculum almost over laps base o f p e c t o r a l f i n r a y s . Body deepest under t h i r d o r f o u r t h sp ine o f d o r s a l f i n . In l a t e r a l v i e w , nape s t r a i g h t , or ve ry s l i g h t l y convex. Base o f d o r s a l f i n s t r a i g h t p o s t e r i o r t o f o u r t h or f i f t h s p i n e . D o r s a l and v e n t r a l p r o f i l e o f cauda l peduncle s i m i l a r l y s l i g h t l y concave. Base o f a n a l f i n s t r a i g h t or s l i g h t l y convex. P r o f i l e from a n a l f i n t o isthmus s t r a i g h t o r convex. I n v iew from d o r s a l s i d e , l a t e r a l p r o f i l e s t r a i g h t t o narrow cauda l pedunc le . A n t e r i o r n o s t r i l t u b u l a r , s h o r t , s t r a i g h t , s l i g h t l y r a i s e d on p o s t e r i o r s i d e . N o s t r i l d i r e c t l y l a t e r a l t o n a s a l spines, from l a t e r a l v iew n o s t r i l d i r e c t l y a n t e r i o r t o v e n t r a l bo rde r o f p u p i l . P o s t e r i o r n o s t r i l s m a l l e r , t u b u l a r , d i r e c t l y p o s t e r i o r t o n a s a l s p i n e , s l i g h t l y p o s t e r o - m e d i a l t o a n t e r i o r end o f o r b i t . Pore between n o s t r i l s a l i t t l e c l o s e r t o p o s t e r i o r t han a n t e r i o r n o s t r i l . Eye l a r g e . I n t e r o r b i t a l narrow, s h a l l o w l y concave. G i l l membranes u n i t e d , fused t o isthmus and forms a f a i r l y b road f o l d . Spines on head r e l a t i v e l y s t rong and sharp . I n d o r s a l v iew n a s a l sp ines 39 d i r e c t l y a n t e r i o r t o l a t e r a l m o s t edge o f i n t e r o r b i t when viewed d o r s a l l y . Uppermost p r e o p e r c l e s p i n e p o i n t e d p o s t e r o - d o r s a l l y , the next one p o i n t i n g p o s t e r i o r l y , t h e t h i r d one much s m a l l e r but evident under integument, t he lowest one sharp and p o i n t e d v e n t r a l l y . Spines on o p e r c l e and p o s t e r o -v e n t r a l border o f suboperc le e v i d e n t . C l e i t h r a l sp ine s h o r t , sharp . Specimens at hand t o o s m a l l t o observe development o f r i d g e s and s t r i a t i o n s . Marked depress ions on e i t h e r s i d e o f p t e r o t i c and p o s t - t e m p o r a l r i d g e s . A few c i r r i evident on l a t e r a l l i n e . C i r r i b road but t h i n and weak. V e r y s m a l l , weak, n a s a l c i r r i may be p r e s e n t . P o s t o c u l a r c i r r i q u i t e t h i n , b r o a d , weak ly m u l t i f i d . C o r o n a l c i r r i s m a l l , s i m p l e , p o s t e r o - m e d i a l t o p o s t o r b i t a l p a i r . C i r r i on occ ipu t t h i n , b r o a d , weak ly m u l t i f i d . L a c r i m a l p o r t i o n o f s u b o r b i t a l r i d g e w i t h about t h r e e b a r b e l - l i k e o r f l e s h y t u b e r c l e -l i k e c i r r i , a t l e a s t one on lower s i d e o f s u b o r b i t a l s t a y . Broad , t h i n , s i m p l e c i r r i near end o f s u b o r b i t a l s t a y . C i r r i near upper end o f o p e r c u l a r s l i t b r o a d , t h i n , weakly m u l t i f i d . C i r r i near c h i n pores b a r b e l - l i k e , s m a l l . C i r r i on m a x i l l a r y b r o a d , t h i n , s i m p l e . D o r s a l band o f s c a l e s conf ined t o at l e a s t t h e d o r s a l h a l f o f t he d i s t a n c e between base o f d o r s a l f i n and l a t e r a l l i n e . T h i s s c a l e band separa te from spinous d o r s a l f i n by about the w i d t h o f one or two s c a l e s . I t then converges t o meet base o f so f t d o r s a l f i n p o s t e r i o r l y . Always some s c a l e s misp laced on cauda l peduncle below l e v e l o f the s t r a i g h t pa th o f lowermost s c a l e row i n t he d o r s a l band. At l e a s t one, u s u a l l y two t o s i x s c a l e s , on c a u d a l pedunc le , and represent row between l a t e r a l l i n e and d o r s a l s c a l e band . Sca l e s not o b v i o u s l y crowded but edges may meet edges o f o ther s c a l e s i n d o r s a l band. A n t e r i o r l y , a l l rows o f equa l s i z e d s c a l e s . P o s t e r i o r l y the upper row d i m i n i s h e s i n s i z e and d i sappears under midd le o f sof t d o r s a l f i n . Ventra lmost row disappears beh ind p o s t e r i o r p o r t i o n o f so f t d o r s a l f i n . Midd l e two rows p e r s i s t on t o cauda l peduncle a l though the 40 s c a l e s o f the upper row on cauda l peduncle are reduced i n s i z e . Often a few s c a l e s above p r i n c i p l e two rows o f l a t e r a l s c a l e band. Sca l e s o f a x i l o f p e c t o r a l f i n s m a l l and s c a t t e r e d . Over o r i g i n o f a n a l f i n t h e v e n t r a l s c a l e band expands t o e ight o r n i n e more d e f i n i t e rows. Band then t a p e r s t o about two rows on cauda l pedunc le . Transverse and h o r i z o n t a l rows evident but anastamose w i t h t a p e r i n g o f band. A n a l s c a l e row always p r e s e n t , much l e s s than o n e - h a l f the s i z e o f s c a l e s on o the r bands. T h i s row of t en j o i n s same row from other s i d e o f a n a l f i n and con t inues up m i d l i n e o f b e l l y . P o s t e r i o r l y , v e n t r a l s c a l e band separa te from l a t e r a l l i n e by about w i d t h o f s c a l e . A n t e r i o r l y i t i s more separate but s c a l e s above and beh ind a x i l o f p e c t o r a l f i n t end t o f i l l i n i n t e r v e n i n g space . L a t e r a l l i n e curves s l i g h t l y upward above o r i g i n o f a n a l f i n and may recurve ve ry s l i g h t l y toward s u p r a c l e i t h r u m under spinous d o r s a l f i n . O r i g i n o f spinous d o r s a l f i n approx imate ly over the middle o f t he base o f t he p e c t o r a l f i n . F i f t h or s i x t h sp ine o f d o r s a l f i n t a l l e s t . T h i r d sp ine s h o r t e r t han second d o r s a l s p i n e . F i r s t sp ine always s h o r t e r than second sp ine o f d o r s a l f i n . Four th o r f i f t h sp ine over p o s t e r i o r l i m i t o f depressed operculum. Spines o f d o r s a l f i n become p r o g r e s s i v e l y s h o r t e r a f t e r t h e s i x t h s p i n e . Margin o f membrane between sp ines o f d o r s a l f i n b a r e l y i n c i s e d , i f at a l l . However, margin w e l l i n c i s e d between t h i r d and f o u r t h s p i n e . Sof t rays o f d o r s a l f i n appear weak. Approx imate ly t he t e n t h r ay i s l o n g e s t ; t he o ther rays p r o g r e s s i v e l y s l i g h t l y s h o r t e r on each s i d e . Marg in o f so f t d o r s a l f i n appears s l i g h t l y rounded, e s p e c i a l l y a n t e r i o r l y and p o s t e r i o r l y . Marg in o f f i n membrane may be ve ry s l i g h t l y i n c i s e d . Specimens known are t o o s m a l l t o determine i f t he d o r s a l o r a n a l rays b ranch . Depressed rays o f d o r s a l f i n b a r e l y , i f at a l l , meet base o f r a y l e t s on c a u d a l f i n . O r i g i n o f a n a l f i n under about the f o u r t h so f t ray o f d o r s a l f i n . 41 Las t a n a l f i n ray under about t h e penu l t ima te sof t d o r s a l f i n r a y . Rays o f a n a l f i n o f approx imate ly s i m i l a r he igh t except f o r f i r s t and l a s t few r a y s . Marg in o f membrane on a n a l f i n w e l l i n c i s e d , p a r t i c u l a r l y a n t e r i o r l y . Caudal f i n t r u n c a t e . P e c t o r a l f i n I n c i s e d between r a y s , p a r t i c u l a r l y between lower t e n r a y s . N i n t h o r t e n t h p e c t o r a l r ay from v e n t r a l s i d e l o n g e s t . Lower rays t h i c k e r and s h o r t e r compared t o upper ones. P e l v i c f i n s w i t h i n c i s e d membrane. I n males , at l e a s t , t he p e l v i c f i n s extend p o s t e r i o r l y pas t t i p o f p e c t o r a l f i n . A c c o r d i n g t o G i l b e r t and Burke (1912) t h e p e l v i c s o f the male have deeply i n c i s e d membranes, and a l s o have a dense s e r i e s o f s t a l k e d , c lub-shaped p a p i l l a e . I t i s assumed t ha t they have a bony support from the su ture between each a c t i n o t r i c h i a . The p e l v i c f i n s on o ther specimens appear t o be no rma l . G e n e r a l c o l o u r o f d o r s a l su r face l i g h t r e d , p i n k i s h , o r p a l e brown. V e n t r a l su r face p robab ly w h i t e . The four bars ac ross back between approx imate ly t he f i f t h and e i g h t h sp ine o f d o r s a l f i n , the second ba r between t h i r d and f i f t h so f t d o r s a l r a y , t h e t h i r d ba r between base o f n i n t h and t w e l f t h so f t d o r s a l r a y , and the f o u r t h bar between base o f s i x -t e e n t h and l a s t so f t d o r s a l r a y . The f i r s t bar extends toward d o r s a l p o r t i o n o f base o f p e c t o r a l f i n , the second extends i r r e g u l a r l y t o t he l a t e r a l l i n e , t h e t h i r d extends p o s t e r i o r l y toward l a t e r a l l i n e , and t he f o u r t h ba r extends a v a r i a b l e d i s t a n c e toward a n a l f i n . T h i s p a t t e r n can be broken up by l i g h t and dark pa t ches . These bars can extend a n t e r i o r l y on d o r s a l f i n . Dark b l o t c h e s appear near t i p s o f f i r s t t h r ee spines o f d o r s a l f i n and near t i p s o f f i f t h t o seventh sp ines o f d o r s a l f i n . Dark b l o t c h e s appear near t i p s o f f i r s t t h r e e sp ines o f d o r s a l f i n and near t i p s o f f i f t h t o seventh sp ines o f d o r s a l f i n . Many dark bars appear on a n a l f i n , p e r p e n d i c u l a r t o the r a y s . P e c t o r a l f i n w i t h s e v e r a l p e r p e n d i c u l a r i r r e g u l a r b a r s . Caudal f i n w i t h ba r ac ross base and w i t h another one o r two more d i s t a l l y on f i n . 42 P e l v i c f i n s w h i t e . In males , t h e a x i l , o f p e c t o r a l f i n appears t o be w h i t e w i t h a t h i n dark bar ac ross the base o f the r a y s . Maybe a s t r i k i n g dark b l o t c h or two across t h i s l i n e . P e l v i c f i n s may have a s e r i e s o f dark spots or bars on membrane between rays or on s i d e o f ray i t s e l f . L e n g t h ; - To at l e a s t 127 mm. Z a p u s : - R e f e r i n g f o o t . P robab ly r e f e r l n g t o l o n g p e l v i c f i n o f male . The genus Zapus r e f e r s t o t h e l o n g - l e g g e d "Jumping mouse", a rodent . Range: :- Chagulak I s . t o A t t u I s . A l a s k a . P robab ly throughout A l e u t i a n I s l a n d s . Tab le I I I . D i s t r i b u t i o n Records o f H . zapus Reference or L o c a t i o n c o l l e c t i o n No. J u v e n i l e s and A d u l t s 52°12»N. 179°52»E. (depth 198-258 f t . ) G i l b e r t & Burke 52°11*N. 179°57*W. (depth 324-336 f t . ) " 52°55»N. 173°27»E. (depth 342-354 f t . ) " * Semisopochnoi I s . (specimens from above c o l l e c t i o n s . ) U . S . N . M . 70866 * Chagulak I s . ( r e t r e i v e d from d i v i n g b i r d ) BC 63-1426 Post l a r v a e * 52°95 ? N. 175°00*W. BC 62-904 * 51°00»N. 1 6 9 ° 5 9 , W . BC 62-914 * 52°59*N. 179G00»W. ( l e n g t h 18-21 mm) BC 62^848 * Unknown l o c a l i t y . BC 62-843 * Unknown l o c a l i t y ( l e n g t h 16-19 mm) BC 62-907 | 0 M M . COLL. NO. BC. 63-1005 * • POSTLARVA OF H.JORDAN I Figure 9. FEMALE-- COLL. NO. BC. 63-404 HEMILEPIDOTUS JORDANI. Figure 1 O. 45 Hemi lep ido tus (Hemi lep ido tus ) . jordani Bean ( F i g s . 9 and 10) Hemilep idotus . jo rdani Bean, 1881, p . 153 (Unalaska I s . , A l a s k a ) ; G i l b e r t , 1895, p . 431; Jordan and Evermann, 1898, p . 1934; 1900, F i g . 704; Jordan and G i l b e r t , 1899, p . 457; Evermann and Goldsborough, 1907, p . 303 , F i g . 61; G i l b e r t and Burke , 1912, p . 53; S o l d a t o v and L i n d b e r g , 1930, p . 201; Rendah l , 1931, p . 34; S c h u l t z and Welander , 1934, p« 5» Gorbunova, 1964, p . 243• Hemilep ido tus hemi lep ido tus Schmidt , 1904, p . 113 ( i n pa r t non T i l e s i u s ) . Hemi lep ido tus hemi lep ido tus . jordani Schmidt , 1929, p . 362; 1950, p . 140. T y p e s ; - Type specimen, U . S . N . M . No. 27598. D i a g n o s i s ; - The narrow m a x i l l a r y c i r r i , unbranched d o r s a l and a n a l f i n r a y s , absence o f s c a l e s between d o r s a l band and l a t e r a l l i n e on cauda l pedunc le , the s imple lower p r e o p e r c u l a r s p i n e , s imple n a s a l c i r r i , f i r s t sp ine s h o r t e r t han second sp ine o f d o r s a l f i n , 59 o r more l a t e r a l l i n e pores and 17 t o 19 p e c t o r a l f i n rays a l l serve t o d i s t i n g u i s h t h i s s p e c i e s . M a t e r i a l s : - D e s c r i p t i o n based on specimens from c o l l e c t i o n no . BC 63-1026 (30 specimens between 72 and 238 mm were used) . Specimens a v a i l a b l e from o ther p a r t s o f t h e range were checked aga ins t the s i g n i f i c a n t c h a r a c t e r s . Specimens from c o l l e c t i o n nos . BC 62-470, 500, 503, 504, 644, 645, 661, 696; BC 63-404 and 1040 were used i n a d d i t i o n f o r s e x u a l d i f f e r e n c e s and p y l o r i c caeca coun t s . D e s c r i p t i o n : - D o r s a l : X I ( X I I ) , 20-21 ( 1 8 ) , [ 23 I 5 9 . A n a l : 16-17 [ l 8 ] 5 9 . P e c t o r a l : 18 (17-19). V e r t e b r a e : 36-37 (38). L a t e r a l l i n e p o r e s : 63.9 (59-68, "74") . Transverse s c a l e rows i n d o r s a l band: 71.9 (67-78). Number o f scales i n row above l a t e r a l l i n e : 27.0 (18-34) . Number o f h o r i z o n t a l rows o f s c a l e s i n d o r s a l band: 4 ( 5 ) . P y l o r i c caecae: 4-5. Width immediate ly behind p e c t o r a l f i n s i n specimens above 100 mm: 181 46 (166-239) (17 spec imens) , : Width immediately behind p e c t o r a l f i n s i n specimens under 10G mms 162 (151-181) (23 specimens) . Depth from o r i g i n o f d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i n s s 253 (234-275). Depth from o r i g i n o f so f t d o r s a l f i n t o o r i g i n o f a n a l f i n s 193 (175-223). Head lengths 360 (341-396). O r b i t diameter i n specimens over 100 mms 100 (88-115) (17 specimens) . O r b i t diameter i n specimens under 100 mms 107 (93-120) (23 specimens) . I n t e r o r b i t a l wid ths 58 (54-73). Snout l e n g t h ( e x c l u d i n g p r e m a x i l l a ) s 75 (68 -85) . Narrowest l a c r i m a l widths 34 (27-44), Caudal Peduncle lengths 98 (83-LL4). Cauda l peduncle depths 59 (50-71). Snout t o o r i g i n o f d o r s a l f i n s 319 (300-337). Snout t o o r i g i n o f a n a l f i n s 565 (537-589). Length o f base o f d o r s a l f i n s 627 (580-707). Height o f longes t sp ine on d o r s a l f i n s 122 (108-148) (29 specimens) . Length o f longes t so f t r ay on d o r s a l f i n s . 165 (145-183). Length o f base o f a n a l f i n s 321 (297-346). Length o f longes t r ay on a n a l f i n s 116 (105-128) (28 spec imens) . Length o f base o f p e c t o r a l f i n s 128 (111-159). Length o f longes t r ay on p e c t o r a l f i n s 268 (247-283) (29 specimens) . Length o f longes t c a u d a l rays 243 (209-283). Length o f longes t r ay o f p e l v i c f i n i n specimens under 100 mms 210 (180-223) (23 specimens) . Length o f longes t r ay o f p e l v i c f i n i n females over 100 mms 209 (169-231) (35 specimens) . Length o f longes t ray o f p e l v i c f i n i n males over 100 mms 235 (207-264) (14 spec imens) . Head robust t o s l i g h t l y depressed . I n l a t e r a l v i e w , p r o f i l e o f occ ipu t s l o p i n g , t he pa th i f extended passes th rough upper t h i r d o f eye. Snout s l o p i n g s t e e p l y and convex d o r s a l l y . P r o f i l e h o r i z o n t a l from n a s a l spines t o o r b i t . P r o f i l e from lower jaw t o isthmus more o r l e s s s t r a i g h t . In d o r s a l v i e w , p r e m a x i l l a e a n d ' m a x i l l a e rounded but head appear ing s l i g h t l y p o i n t e d ; p r e o p e r c u l a r p r o f i l e s t r a i g h t . Operculum over laps base o f p e c t o r a l f i n . Head not p a r t i c u l a r l y b road . 47 Body deepest under, anter iormost , few sp ines o f d o r s a l f i n . Nape s h o r t , maybe s l i g h t l y convex. Base o f d o r s a l f i n s l i g h t l y convex under s p i n y p o r t i o n , o therwise s t r a i g h t . In l a t e r a l v i e w , d o r s a l and v e n t r a l p r o f i l e o f cauda l peduncle both s i m i l a r l y s l i g h t l y concave. Base o f a n a l f i n ve ry l i g h t l y convex. Base o f a n a l f i n t o base o f p e l v i c f i n s t r a i g h t t o convex i n p r o f i l e . Base o f p e l v i c f i n t o isthmus s t r a i g h t . From d o r s a l v i e w , l a t e r a l p r o f i l e o f body s t r a i g h t t o concave. T a i l s l e n d e r . A n t e r i o r n o s t r i l s h o r t , t u b u l a r , d i r e c t l y l a t e r a l t o n a s a l s p i n e , i n l a t e r a l v iew d i r e c t l y a n t e r i o r t o v e n t r a l margin o f o r b i t . N o s t r i l not f l a r e d but o f un i fo rm d iameter . The p o s t e r i o r margin o f n o s t r i l r a i s e d t o he igh t o f d iameter o f n o s t r i l . In d o r s a l v i e w , p o s t e r i o r n o s t r i l i n l i n e between n a s a l sp ine and upper border o f p u p i l . N o s t r i l c o n s t r i c t e d d i s t a l l y . Conspicuous head pore o n e - t h i r d t o one-quar te r o f the way down i n d i s t a n c e from p o s t e r i o r t o a n t e r i o r n o s t r i l . Many s m a l l head pores on d o r s a l su r face o f head, p r e o p e r c l e , s u b o r b i t a l s , and snout . O r b i t s r e l a -t i v e l y s m a l l e r i n a d u l t s t han j u v e n i l e s but d i f f e r e n c e l e s s than i n H . hemilep i d o t u s . I n t e r o r b i t a l h i g h l y concave, l o n g i t u d i n a l r i d g e s on each s i d e o f m i d l i n e i n i n t e r o r b i t . G i l l membranes u n i t e d and fused t o isthmus and forms a moderately wide f ree f o l d . Head sp ines s t rong and sharp . N a s a l sp ines s t rong and sharp , on top o f snout at l e v e l midway between o r b i t and m i d l i n e . Upper p r e o p e r c u l a r s p i n e s t r o n g , sharp , p o i n t i n g p o s t e r o - d o r s a l l y , l e n g t h about t ha t o f p u p i l d iamete r . Next p r e o p e r c u l a r s p i n e s t rong and sharp , o f t en s l i g h t l y l onge r t han upper s p i n e . T h i r d sp ine much s h o r t e r , s t r o n g , sharp . Four th p r e -o p e r c u l a r s p i n e , sharp , s i m p l e , h idden beneath s k i n , p o i n t e d v e n t r a l l y . One l a r g e r specimen from the Commander I s l ands had some f l a t t e n i n g o f t h i s s p i n e . Spines on operculum and suboperculum weak. C l e i t h r a l sp ine s m a l l , sha rp . Reduced s t r i a t i o n s r a d i a t e from p o s t o c u l a r and o c c i p i t a l areas o f IS l a r g e r specimens. Ridges on o p e r c l e , f r o n t o - p a r i e t a l , p t e r o t i c and sup ra -c l e i t h r a l , and p o s t - t e m p o r a l a reas . A l l except ope rcu l a r r i d g e become h i g h l y rugose i n l a r g e r specimens. C i r r i t end t o be t h i n and weak. C i r r i below n a s a l sp ines s imple and p a p i l l a - l i k e . N a s a l c i r r i ve ry s m a l l , s i m p l e , narrow. P o s t o c u l a r c i r r i t h i n , w i d t h equa l t o h e i g h t . T h i s c i r r i weakly m u l t i f i d i n t o a maximum o f 10 f r i n g e s i n l a r g e s t specimens; c i r r i d i r e c t e d s l i g h t l y p o s t e r i o r t o t r a n s -v e r s e a x i s . F r o n t a l c i r r i immediate ly p o s t e r o - m e d i a l t o p o s t o c u l a r c i r r i , but c l o s e r t o m i d l i n e . C i r r i on occ ipu t s i m i l a r t o p o s t o c u l a r p a i r . Three or four f l e s h y t u b e r c l e - l i k e c i r r i on l a c r i m a l p o r t i o n o f s u b o r b i t a l r i d g e ; o thers may be present near s u b o r b i t a l s tay p o r t i o n . C i r r i near end o f s u b o r b i t a l s t a y , s m a l l , t h i n . C i r r i near upper end o f ope rcu l a r s l i t l a r g e r , t h i n , m u l t i f i d . C i r r i near c h i n pores s m a l l , b a r b e l - l i k e . C i r r i on lower jaw q u i t e s m a l l . C i r r i on m a x i l l a r y t h i n , i t s w i d t h .33 ( . 2 6 - . 4 9 ) o f i t s l e n g t h . Sca l e s w i t h up t o about twenty s p i n u l e s . D o r s a l s c a l e band con f ined w i t h i n the d o r s a l h a l f o f t he d i s t a n c e between l a t e r a l l i n e and d o r s a l f i n . Band separa te from most o f s p i n y d o r s a l f i n by approx imate ly o n e - h a l f or t w o - t h i r d s o f the w i d t h o f the band. P o s t e r i o r l y , d o r s a l s c a l e band converges v t o meet base o f so f t d o r s a l f i n . S c a l e rows on d o r s a l band s t r a i g h t on t." cauda l pedunc le , v e r y r a r e l y extend as f a r as base o f r a y l e t s on cauda l f i n . Ex t remely r a r e l y s c a l e s present between d o r s a l band and l a t e r a l l i n e on cauda l pedunc le . Sca l e s may meet edges o f adjacent s c a l e s on same row i n d o r s a l band. S c a l e s may meet edge o f s c a l e s on o ther rows. A few reduced s c a l e s r e p r e s e n t i n g an uppermost f i f t h row o c c a s s i o n a l l y present under spinous d o r s a l f i n , however, extremely r a r e l y are t h e r e over a h a l f dozen o f these s c a l e s . A n t e r i o r l y s c a l e s o f o ther four rows o f equa l s i z e . S c a l e s o f vent ra lmost row o f d o r s a l band and s c a l e s o f bo th lower bands r a i s e d on 49 d o r s a l s i d e o n l y . Other s c a l e s r a i s e d on a l l s i d e s . Uppermost row o f d o r s a l band reduced i n s i z e under sof t d o r s a l f i n and d isappears under p o s t e r i o r p o r t i o n o f sof t d o r s a l f i n . Lowest s c a l e row o f d o r s a l band d isappears under p o s t e r i o r p o r t i o n o f d o r s a l f i n . Two rows o f s m a l l e r s c a l e s p e r s i s t onto cauda l pedunc le . S p i n u l e s on bony elements o f l a t e r a l l i n e present on p o s t e r i o r one-h a l f t o t w o - t h i r d s o f d o r s a l s i d e o f the more developed s c a l e s . V e n t r a l band o f s c a l e s w i t h approximate ly two or t h r e e i r r e g u l a r rows o f w i d e l y spaced s c a l e s beh ind c l e i t h r a l s p i n e . Band expands r a p i d l y t o about e igh t more crowded and d i s t i n c t rows above anus. Band t ape r s t o about two rows on cauda l pedunc le . The s c a l e rows tend t o anastomose w i t h the t a p e r i n g o f t h e v e n t r a l band. Band separate from l a t e r a l band by about w i d t h o f t h r e e s c a l e s a n t e r i o r l y and about h a l f a l a r g e s c a l e p o s t e r i o r l y . V e n t r a l s c a l e band separa te from a n a l f i n by about the w i d t h o f the band. Sometimes a n a l s c a l e row p e r s i s t s i n a d u l t s as f e e b l e o s s i f i c a t i o n s i n f l a t t e d rudimentary p a p i l l a e . Les s f r e q u e n t l y i t p e r s i s t s i n m i d l i n e o f b e l l y . Curva tu re o f l a t e r a l l i n e above o r i g i n o f a n a l f i n . S e v e r a l weak c i r r i on l a t e r a l l i n e . O r i g i n o f d o r s a l f i n over lowest p e c t o r a l r a y . F i f t h o r s i x t h sp ine o f d o r s a l f i n t a l l e s t . F i r s t s p i n e never l o n g e r t han second sp ine o f d o r s a l f i n . F i f t h d o r s a l sp ine over p o s t e r i o r l i m i t o f depressed operculum. Spines o f d o r s a l f i n become p r o g r e s s i v e l y s h o r t e r a f t e r s i x t h s p i n e . Las t d o r s a l f i n sp ine may be t a l l e r t han penu l t ima te s p i n e . Margin o f f i n membrane moderate ly i n c i s e d between t h i r d and f o u r t h s p i n e s . Membrane between t h i r d and f o u r t h sp ine g r e a t l y i n c i s e d . Height o f membrane on a n t e r i o r s i d e o f f ou r th sp ine i s . 4 5 ( . 3 2 - . 6 6 ) i n he igh t o f t h i r d s p i n e . j F i f t h t o seventh sof t d o r s a l f i n ray t a l l e s t . Marg in appears almost s t r a i g h t 50 from t h i r d t o f i f t e e n t h sof t d o r s a l f i n r a y . Rays on e i t h e r s i d e p r o g r e s s i v e l y s h o r t e r and f i n rounded. Marg in o f membrane f a i n t l y i n c i s e d between rays o f d o r s a l f i n . No branched d o r s a l o r a n a l f i n rays* Depressed rays o f d o r s a l f i n w e l l shor t o f c auda l base or base o f r a y l e t s on cauda l f i n . O r i g i n o f a n a l f i n under about t he f o u r t h sof t d o r s a l r a y . Las t r ay o f a n a l f i n about under penu l t ima te r ay o f d o r s a l f i n . Marg in o f f i n membrane deeply i n c i s e d between f i r s t t h i r t e e n rays o f a n a l f i n . E i g h t h t o t h i r t e e n t h ray o f a n a l f i n l o n g e s t . Rays become s l i g h t l y s h o r t e r on e i t h e r s i d e . Caudal f i n t r u n c a t e . P e c t o r a l f i n rounded. Lower e l even rays w i t h margin o f membrane w e l l i n c i s e d between them. Lower r ays t h i c k e r . Marg in o f membrane between upper rays almost e n t i r e . Tenth o r e l even th r a y from v e n t r a l s i d e o f p e c t o r a l f i n l o n g e s t . Marg in o f membrane between p e l v i c f i n s s h a l l o w l y i n c i s e d . I n males t h e rays o f p e l v i c f i n average a l i t t l e l o n g e r than i n females . L o n g i -t u d i n a l r i d g e o f p a p i l l a t e s k i n on lower s i d e o f p e l v i c rays i n some mature males . No rudimentary bony c t e n i observed on p e l v i c r a y s . I n most specimens t i s s u e over r ays smooth. C o l o u r p a t t e r n v a r i a b l e w i t h l i g h t y e l l o w i s h brown or dark brown p redomina t ing . Some specimens from sha l low water appear d a r k e r . Dark spots sometimes present on upper s i d e s o f body, lower s i d e s o f t a i l and b e l l y . 'Dark pigment never present on v e n t r a l su r face from lower jaw t o a rea around base o f p e l v i c f i n s i n specimens over 100 mm. Smudges may appear here i n s m a l l e r specimens. Dark bars present between approx imate ly t he base o f f i f t h t o eighth d o r s a l s p i n e s , t h i r d and s i x t h sof t d o r s a l r a y s , t e n t h t o t h i r t e e n t h so f t d o r s a l r ays and seventeenth t o l a s t so f t d o r s a l r a y s . The f i r s t b a r extends i n i r r e g u l a r course t o l a t e r a l l i n e and upper base o f p e c t o r a l f i n base , second ba r a v a r i a b l e d i s t a n c e towards l a t e r a l l i n e , t h i r d ba r a v a r i a b l e d i s t a n c e towards or pas t l a t e r a l l i n e , w h i l e t he 51 f o u r t h ba r may fork towards, a n a l f i n and cauda l pedunc le . Often a dark a rea appears below and i n f ront o f f i r s t t h r e e d o r s a l f i n s p i n e s . These bands may cont inue t o margin o f d o r s a l f i n . L i g h t and dark m o t t l i n g on d o r s a l and l a t e r a l s i d e s . May obscure t h i s p a t t e r n . I n p a r t i c u l a r , p a t t e r n below l a t e r a l l i n e q u i t e v a r i a b l e . D o r s a l su r face always a l i t t l e darker than v e n t r a l s u r f a c e . Cauda l f i n may show t h r e e v e r t i c a l dark b a r s . A n a l f i n v a r i e s from l i g h t t o dark pigment . V a r i a b l e amounts o f s p o t t i n g may a l s o o c c u r . P e c t o r a l f i n s w i t h i r r e g u l a r bars and m i t t l i n g , o r l a r g e l y darkened. P e l v i c f i n s w h i t e . In many males the p e l v i c , p e c t o r a l , and a n a l f i n s become b l a c k . Often many f i n e spots appear on the lower s i d e s o f the body. A female from the Commander I s l a n d s had darker and s l i g h t l y p a p i l l a t e p e l v i c f i n s , a c o n d i t i o n wh ich i s no rma l ly found i n males . L e n g t h ; - To 410 mm (Gorbunova, 1964 - whether s tandard or t o t a l l e n g t h not known). j o r d a n i : - Named f o r D. S. Jo rdan . Range ; - S i t k a I s . , A l a s k a t o G u l f o f Anadyr , U . S . S . R . t o Southwest Kamchatka. 52 Table I V . D i s t r i b u t i o n Records o f H . . jordani L o c a t i o n Reference o r , c o l l e c t i o n no. Post l a r v a e * ( l e n g t h 26 mm) * ( l e n g t h 25-28 „mm) * Cons tan t ine H b r . , Amchi tka , A l a s k a (21-27 mm) * (22 mm) * 55°39 'N. 170°00»W. * Kiska"Harbour (22 mm) * Massacre Bay, A t t u I s . (depth" 30 f t . ) ( l e n g t h 23 mm) t? # tt # « * K i s k a Harbour * N i k o l s k i , Umnak I s . * 54°38»N. 159°01«W. J u v e n i l e s and A d u l t s ( su r f ace ) ( l e n g t h 23 mm) ( su r f ace ) ( l e n g t h 23 mm) (depth s o f t ) ( l e n g t h 21 mm) ( l e n g t h 23 mm) ( l e n g t h 25 mm) ( l e n g t h 24 mm) ( su r f ace ) ( l e n g t h 24 mm) ( l e n g t h 29 mm) BC 63-1303 BC 63-1293 BC 63-1010 BC 62-847 BC 62-908 BC 63-915 BC 63-887 BC 63-878 BC 63-9O3 BC 63-1013 BC 63-1005 BC 63-916 BC 63-IO7O BC 63-1292 BC 62-702 A l a s k a n Panhandles -S i t k a ( f rom A l b a t r o s s ) * P a t t e r s o n Bay , Baranof I s . / Po r t A l t h o r p / G u l f o f A l a s k a t o A la skan P e n k i n s u l a ; Uhga K a r l u k Por t Chatham, "Cook ' s I n l e t S t . "Pau l , Kodiak Humboldt Harbour , Shumagin I s . Afognak Bay C h i g n i k Bay 57°48»00"N. 5 4 ° 4 8 ? 0 0 M N . 54°49 ?00*.»N. 58°00"00"N. 54?17 700"N. 55°08' .20«N. 155°00»00"W. 160°00T00»¥. 162°59W»W. 152°00!CXW. 164°31T00"W. l60°19'00»'Wo Evermann & Goldsborough, 1930 BC 63-1270 Bean, 1881 (b) Jordan & Evermann, 1898 Jordan & Evermann, 1898 Bean 1881 (b) n ft Evermann & Goldsborough, 1930 tt BC 62-650 BC 62-657 BC 62-658 BC 62-660 BC 62-665 BC 62-424 53 Locat ion Reference or collection no. 54°00»00"F. 165°00»00"W. BC 62-427 54°33»30"N. l62o29*30»»W. BC 62-440 54°12»15'TN. 164°31 ?30«W. • BC 62-444 54°28»00"N. 163°31VG0"W. BC 62-487 55°24?00»N. 160°33?00«W. BC 62-488 54°54'00"N. 163°00»00"W. BC 62-498 55°33?30"N. 159°43?00"W. BC 62-499 55°36?00«N. 158°30»oo»W. BC 62-500 54°17T20"N. 163°30»00'«W. BC 62-501 * 54°05?50»N. 164°28*00«W. BC 62-502 * 57°36'00"N. 154°30»00«W. BC 62-503 54°I3t30"W. 163°30»30"W. BC 62-504 * 55°16»30"N. 159°40»00"W. BC 62-505 54°48'00"N. 160°58»30"W. BC 62-507 •A. 54°54,00"N. l6l°00»00«W. BC 62-454 54°39fOO"N. 159°O0»OO'W. BC 62-468 56°42»00"N. 152°30»00"W. BC 62-508 * 55°18»00"N. 153°00»00"W. BC 62-509 * 54°32»40"No 159°47»00«W. BC 62-458 * 53°54*00?pN, 163° 58»00"W. BC 62-471 * 57°10vOO"N. 155°02*00«W. BC 62-510 # 55°18»O0"N. 157°06?00"W. BC 62-511 * 54°12«oo"N. 161°30»00«'W. BC 62-512 •fi 54°28»00»N. 159°13950MW. BC 62-513 54°20»20?'N. 161°44?00»W. BC 62-420 * 54°18» N. 161°02» W. BC 62-670 57°06» N. 152°30» W. BC 62-645 * 57°06» N. 152°30» W. BC 62-677A 58°33 f N. 153°45 T W. BC 62-497 # 56°48» N. 154°30» W. BC 62-680 * 54°27» N. 159°45 ? W. 155500» . W. . BC 62-682 * 56°54* N. BC 62-526 * 56°42» N. 57°24 ? N. 154°30» W. BC 62-683 152°30» W. BC 62-684 58°36» N. 153°00» W. BC 62-697 * 56°48» N. 155°00» W. BC 62-651 * UyakBay, Kodiak Is. BC 59-486 # Kit 6 1 Bay, Afognak Is. BC 58-209 Kachemak Bay BC 61-518 * tt tt BC 61-520 * « tt (0-10 ft.) BC 62-991 * it it Near Homer (84 ft.) BC 62-995 * t t tt Aurora Lagoon (210-270 ft.) BC 62-998 * it tt Tutka Bay BC 62-999 * tt it Halibut Cove (150-210 ft.) BC 63-119 * it tt Kasitsna Bay (Intertidal) BC 63-118 * • t ! ' . " ' tl Nubble" Pt. (240-360 ft.) BC 63-120 -T* Kalsin Is. near Kodiak Is. BC 63-1032 Region Is. Woody. I s . hear Kodiak Is. BC 63-1026 Kelp Pt.,' Cold Bay BC 63-1438 (About 20 additional collections used with Incomplete data) 54 L o c a t i o n Reference or c o l l e c t i o n no . B e r i n g Seas -S t . P a u l I s . S t . George I s . Amak I s . * 75 m i l e s o f f St , * S t . P a u l I s . * S t . P a u l I s . * Izembek Bay-George I s . (144 f t . ) (6-24 f t . ) ( t i d e p o o l ) 52°11»N. 52°11»N. 52°11»K 0 A l e u t i a n I s . s -Unalaska I s . , I l i u l u k * Chernoffsky 179°49»E. I79052VE, I79°57*W. * Cons tan t ine H b r . , Amchitka I s . * Sweeper"Cove, Adak I s . * Cape P o i n t , Unimak I s . * N i k o l s k I , Umnak I s . * Chernabura I s . S . E . * tt tt * tt t? * 25 m i l e s o f f Akutan I s . * F inger Bay , Adak, I s . * Sweeper Cove, Adak, I s . U . S . S . R t — N a t a l s k y Bayr S.W. pa r t o f Anadyr G u l f (most n o r t h e r n -most record) * Cape U s h i n , U . S . S R . (50 f t . ) * 52 b 30»N.(53°10»N?) 155°10»E. * 49°30»N. 156°40»E. (most southern and wes te rn record) BC 63-404 (250-310 f t . ) tt G i l b e r t & ] (250-260 f t . ) tt (225-245 f t . ) tt (60 f t . ) BC 63-1015 BC 63-337 (114-144 f t . ) BC 63-347 (3 f t . ) BC 63-1068 BC 63-327 ( t i d e p o o l s ) BC 63-334 BC 63-331 BC 62-564 ( 5 - 2 0 f t . ) BC 63-1422 ( 0 - 2 0 f t . ) BC 63-1429 Jordan & Evermann, 1898 « G i l b e r t , 1895 BC 62-565 BC 63-1296 BC 63-1458 BC 63-1433 Bean, 1881 (a) (b) Andr i a shev , 1937 BC 63-1040 BC 63-402 F E M A L E : - COLL . NO. BC. 63-396. I S O M M - • 1 H E M I L E P I D O T U S G1LBERTI. Figure 11 56 17.5 MM.  Figure 12. Sketch of Hemilepidotus gilberti taken from Figure 2 of Gorbunova I964. Hemilep idotus (Hemilep idotus) g ilbert i Jordan and Starks . (Figs. 11 and 12) Cottus Trachurus Pallas, 1914j p» 138 (3n part - Kuriles). Hemilepidotus gilberti Jordan and Starks, 1904, p. 254, fig. 10 (Hakodate, Japan); Schmidt, 1929, p° 364; 1950/ p. 140; Soldatov and Lindberg, 1930, p. 203; Rendahl, 1931. Watanabe, i 9 6 0 , p. 48, pi. 9 - fig. 1. Hemilepidotus hemilepidotus Schmidt, 1904, p. 113 (in part). Hemilepidotus gilbrti Watanabe, 1958, p. 246. Hemilepidotus gilberti gilberti Gorbunova, I964, p. 238. North American references synonymized Cottus Trachurus of Pallas with H. hemilep idotus. The specimen he describes from North America is un-doubtedly the latter species. However, the reference of Pallas (1814) is concerned mainly with the Asiatic fauna. The locality (from Kuriles), fin ray counts (soft dorsal 21, anal 17, and pectoral 15-17) as well as colouration (unspotted uhdersurface) suggest that some specimens at least are H. gilberti. Some of the spotting on the undersurface may also be that of male H. gilberti. Consequently, there is a possibility that Trachurus refers mostly to H. gilberti and then Trachurus would have priority. 57 Ho g i l b e r t i has-been, used f o r over 50 yea r s and thus i t i s p r e f e r a b l e t o r ecogn ize t h i s name s i n c e t he r e has never been confus ion over i t . Types? - Type specimen i s number 7446, I c h t h y © l o g i c a l C o l l e c t i o n s , L e l a n d S t an fo rd J u n i o r U n i v e r s i t y Museum. Cotype i s No. 50916 U . S . N . M . D i a g n o s i s ; - The l o n g e r f i r s t d o r s a l f i n s p i n e , f l a t t e n e d lower p r e o p e r c l e s p i n e , p e r s i s t a n c e o f a n a l s c a l e row i n a d u l t s , reduced o r absent f r o n t a l c i r r i , and 37-38 ve r t eb rae he lp t o d i s t i n g u i s h t h i s s p e c i e s . M a t e r i a l s ; - Nine specimens from c o l l e c t i o n numbers BC 63-396, 675, 1040 and from U . S . N . M . 105146, 119857, 119865 were used i n t h i s d e s c r i p t i o n . They range i n s i z e from 40 t o 267 mm. Only one mature,male was a v a i l a b l e . D e s c r i p t i o n ; - D o r s a l ; X I [ X I I } 6 7 , 20-21 [ 2 2 ] 4 5 & 6 ? . A n a l : '17-18 (14) [ 1 9 ] 6 7 . P e c t o r a l : 1 6 - 1 7 ( 1 5 ) . V e r t e b r a e ; 37-38 [ 36] 6 ? . L a t e r a l l i n e p o r e s : 60 .8 (56-66) [ 5 5 - 6 7 ] T r a n s v e r s e s c a l e rows i n d o r s a l band: 73.3 ( 6 9 - 7 8 ) . Number o f s c a l e s i n row above l a t e r a l l i n e : 21 .8 (14 -35) . Number o f h o r i z o n t a l s c a l e rows i n d o r s a l band: 4 ( r a r e l y 5 ) . P y l o r i c caeca : 4 [ 5] Body w i d t h immediately beh ind p e c t o r a l f i n s : 179 (160-202) . Depth from o r i g i n o f d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i n : 259 (237-274). Depth from o r i g i n o f so f t d o r s a l f i n t o o r i g i n o f a n a l f i n : 217 (205-235). Head l e n g t h : 374-(356-399). O r b i t diameter In specimens over 100 mm: 88 (83-96) (7 spec imens) . O r b i t d iameter i n specimens below 100 mm: (108-136) . O r b i t t o end o f o p e r c l e : 2Q3 (179-236). I n t e r o r b i t a l w i d t h : -see H . zapus f o r comment on synonymy o f H . zapus and H . g i l b e r t i . 58 66 (58-72). . Snout l e n g t h , ( e x c l u d i n g p r e m a x i l l a e ) : 72 (66-78). Narrowest l a c r i m a l depths 34 (30-38). Caudal peduncle lengths 108 .(98-117). Caudal peduncle depths 65 (62-74)• Snout t o o r i g i n o f d o r s a l f i n : 311 (284-348). Snout t o o r i g i n o f a n a l f i n s 581 (557-600). Length o f base o f d o r s a l f i n s 658 (646-676). Height o f longes t d o r s a l f i n spines 131 (120-137). Height o f longes t so f t r ay o f d o r s a l f i n s 167 (151-178). Length o f base o f a n a l f i n s 340 (317-350). Height o f longes t r ay o f a n a l f i n s 130 (116-141). Length o f base o f p e c t o r a l f i n s 143 (132-160). Length o f longes t r ay on p e c t o r a l f i n s 300 (272-330). Length o f longes t rar on cauda l f i n s 217(192-285). Length o f p e l v i c f i n on j u v e n i l e s and females : 220 (204-243)o Length o f l onges t r ay on p e l v i c f i n o f males 304» Head a l i t t l e depressed , b r o a d . In l a t e r a l v i e w , p r o f i l e o f occ ipu t s l o p i n g , s l i g h t l y convex, i f extended pa th would pass through upper edge o f o r b i t . . O r b i t s s l i g h t l y e l e v a t e d above p r o f i l e . Snout s t r a i g h t s l o p i n g . P r o f i l e h o r i z o n t a l between n a s a l sp ines and o r b i t . C h i n s l o p i n g , s t r a i g h t t o convex towards i s thmus . In d o r s a l v i e w , m a x i l l a e and p r e m a x i l l a e rounded t o p r e o p e r c l e . P r e m a x i l l a e s l i g h t l y p o i n t e d . Operculum over l aps base o f p e c t o r a l f i n o r base o f p e c t o r a l f i n r a y s . Body deepest near t h i r d o r f o u r t h s p i n e . Nape s t r a i g h t when viewed l a t e r a l l y . Base o f d o r s a l f i n s t r a i g h t between f o u r t h d o r s a l sp ine and c a u d a l pedunc le . D o r s a l and v e n t r a l p r o f i l e o f cauda l peduncle s i m i l a r l y concave. Base o f a n a l f i n s t r a i g h t . P r o f i l e from o r i g i n o f a n a l f i n t o isthmus s t r a i g h t t o convex. In d o r s a l v iew l a t e r a l p r o f i l e s t r a i g h t t o c a u d a l pedunc le . A n t e r i o r n o s t r i l s h o r t , t u b u l a r , maybe w i t h a f l a r e d r i m . N o s t r i l d i r e c t l y l a t e r a l t o n a s a l sp ine and, i n d o r s a l v i e w , d i r e c t l y a n t e r i o r t o d o r s a l p o r t i o n o f o r b i t . P o s t e r i o r n o s t r i l ve ry shor t w i t h a broad base? appears almost as a s m a l l pore on apex o f a s m a l l s w e l l i n g . Head pore one-59 t h i r d d i s t a n c e toward a n t e r i o r n o s t r i l from p o s t e r i o r n o s t r i l . A few head pores s c a t t e r e d on s u b o r b i t a l s and d o r s a l aspect o f head. O r b i t diameter becomes r e l a t i v e l y s h o r t e r w i t h growth. I n t e r o r b i t s h a l l o w l y and f l a t l y concave. G i l l membranes u n i t e d , fused t o isthmus t o form a broad f o l d p o s t e r i o r l y . Head sp ines l o n g , s t r o n g , sharp . N a s a l sp ines s t r o n g , at l e v e l s l i g h t l y m e d i a l t o edge o f i n t e r o r b i t . Upper p a i r o f p r e o p e r c u l a r 3pines l o n g , l eng th may be up t o t w i c e t he diameter o f t he p u p i l . Upper sp ine d i r e c t e d s l i g h t l y p o s t e r i o r t o d o r s a l d i r e c t i o n . Second sp ine s l i g h t l y l onge r t han upper one, d i r e c t e d s l i g h t l y d o r s a l t o p o s t e r i o r d i r e c t i o n . T h i r d one much s m a l l e r but p rominent , sha rp , p o i n t e d , and d i r e c t e d p o s t e r i o r l y . Fou r th , o r l o w e s t , sp ine p o i n t e d v e n t r a l l y , f l a t b road , w i t h remnants o f secondary p o i n t s deve lop ing on l a t e r a l edge. These secondary p o i n t s more developed i n l a r g e r specimens. Spines on o p e r c l e and suboperc l e , shor t and s t r o n g . C l e i t h r a l sp ine s t r o n g , prominent . Rugose s t r i a t i o n s r a d i a t e from p o s t o c u l a r and, t o l e s s e r degree , on o c c i p i t a l areas i n l a r g e r specimens. Modera te ly r a i s e d , l i g h t l y rugose r i d g e s develop i n f r o n t o - p a r l e t a l , p t e r o t i c and p o s t - t e m p o r a l , and s u p r a c l e i t h r a l a r ea s . Operc le r i d g e s t rong and w e l l deve loped . C i r r i weak. S e v e r a l reduced c i r r i on l a t e r a l l i n e . C i r r i below n a s a l sp ines f l e s h y , t u b e r c l e - l i k e . N a s a l c i r r i absent o r extremely reduced . P o s t o c u l a r c i r r i b road , t h i n , i r r e g u l a r l y and weakly m u l t i f i d . F r o n t a l c i r r i ve ry reduced o r absent . T h i n weakly m u l t i f i d c i r r i on o c c i p u t . Three f l e s h y t u b e r c l e - l i k e c i r r i on l a c r i m a l p o r t i o n o f s u b o r b i t a l r i d g e . C i r r i near end o f s u b o r b i t a l s t ay s i m p l e , t h i n . C i r r i on upper operculum s m a l l t h i n , s imple o r weak ly m u l t i f i d . C i r r i near c h i n p o r e s , b a r b e l - l i k e . C i r r i on lower jaw v e r y s m a l l . C i r r i on m a x i l l a e l a r g e , t h i n , b road . Sca le s i n d o r s a l band conf ined t o at l e a s t upper h a l f o f d i s t a n c e between l a t e r a l l i n e and d o r s a l f i n . A n t e r i o r l y s c a l e s o f d o r s a l band separate from 6 0 d o r s a l f i n by o n e - h a l f w i d t h o f band. P o s t e r i o r l y t he band converges t o meet d o r s a l f i n . No s c a l e s misp laced below l e v e l o f s t r a i g h t o f lowest s c a l e row o f d o r s a l s c a l e band. Sca le s evenly spaced between those on same o r d i f f e r e n t rows. Sca l e s may or may not meet edge o f adjacent s c a l e s i n d o r s a l band. Sca l e s i n uppermost row d i m i n i s h i n s i z e p o s t e r i o r l y and d i sappear before end o f so f t d o r s a l f i n . Lowest row a l s o d isappears under p o s t e r i o r qua r t e r o f d o r s a l f i n . M i d d l e two rows p e r s i s t onto cauda l pedunc le . Upper rows w i t h s c a l e s r a i s e d on a l l s i d e s . Lowest row o f s c a l e s r a i s e d on d o r s a l s i d e o n l y . Bony elements o f l a t e r a l l i n e w i t h s p i n u l e s on the p o s t e r i o r h a l f o f t h e d o r s a l s u r f a c e . Many s m a l l s c a l e s i n a x i l o f p e c t o r a l f i n grade i n t o a few l a r g e , w i d e l y spaced , s c a l e s on upper s i d e o f b e l l y . These q u i c k l y expand i n t o e igh t or n ine more dense rows above anus. T h i s v e n t r a l band o f s c a l e s t ape r s t o two o r t h r ee rows on c a u d a l pedunc le . Transverse and h o r i z o n t a l rows t end t o anastamose w i t h t he t a p e r i n g o f band. A n a l s c a l e row c o n s i s t i n g o f h i g h l y reduced s c a l e s always p r e s e n t . T h i s row may p e r s i s t on m i d l i n e o f b e l l y , l e s s f r e q u e n t l y on b r e a s t . V e n t r a l band more separa te from l a t e r a l band a n t e r i o r l y than p o s t e r i o r l y . L a t e r a l l i n e curves s l i g h t l y upward at l e v e l above a n a l f i n , t h e n may recurve s l i g h t l y toward head. O r i g i n o f d o r s a l f i n over o r s l i g h t l y i n advance o f lowest ray o f p e c t o r a l f i n . Four th o r f i f t h sp ine o f d o r s a l f i n t a l l e s t . Second and t h i r d s p i n e s h o r t e r than f o u r t h sp ine o f d o r s a l f i n . F i r s t sp ine always t a l l e r than second s p i n e and extends f u r t h e r even when sp ines are depressed . Fourth or f i f t h sp ine d i r e c t l y over p o s t e r i o r l i m i t o f operculum. Spines become p r o g r e s s i v e l y s h o r t e r p o s t e r i o r t o f i f t h sp ine o f d o r s a l f i n . Las t sp ine o c c a s i o n a l l y t a l l e r than penu l t ima te s p i n e . Marg in o f f i n membrane i n c i s e d 61 between f i r s t f i v e s p i n e s , e s p e c i a l l y deeply I n c i s e d between t h i r d and f o u r t h spines*. Seventh t o t e n t h ray o f so f t d o r s a l f i n t a l l e s t . Rays on e i t h e r s i d e almost same he igh t but p r o g r e s s i v e l y s h o r t e r . F i n p r o f i l e rounded i a n t e r i o r l y and p o s t e r i o r l y . Depressed d o r s a l rays do not reach base o f c a u d a l rays o r r a y l e t s . No branched rays i n a n a l o r d o r s a l f i n . O r i g i n o f a n a l f i n approx imate ly under base o f f o u r t h or f i f t h so f t r ay o f d o r s a l f i n . L a s t ray o f a n a l f i n approximate ly under l a s t ray o f d o r s a l f i n . S i x t h t o f i f t e e n t h rays o f a n a l f i n t a l l e s t ; o ther p r o g r e s s i v e l y s h o r t e r on e i t h e r s i d e . Marg in o f f i n membrane deeply i n c i s e d e s p e c i a l l y a n t e r i o r l y . Caudal f i n t r u n c a t e . P e c t o r a l f i n rounded; margin o f f i n membrane s h a l l o w l y i n c i s e d , more i n c i s e d v e n t r a l l y than d o r s a l l y . Lower rays o f p e c t o r a l f i n t h i c k e r than upper ones. E l e v e n t h ray from v e n t r a l s i d e approximate ly l o n g e s t . P e l v i c f i n s s m a l l ; margin o f membrane i n c i s e d . I n l a r g e r males , c t e n i and p a p i l l a e present on p e l v i c f i n r a y s . Rays e longa te , extend beyond t i p o f p e c t o r a l f i n ; membrane between rays deeply i n c i s e d . C o l o u r v a r i a b l e . Bars o r bands on back between base o f f i f t h t o e i gh th sp ines o f d o r s a l f i n , between t h i r d and f i f t h rays o f sof t d o r s a l f i n , between t e n t h and t h i r t e e n t h ray o f so f t d o r s a l f i n , and between seventeenth and l a s t so f t r ay o f d o r s a l f i n . These bands extend i n an i r r e g u l a r course toward l a t e r a l l i n e . The bands a re v e r y i r r e g u l a r below l a t e r a l l i n e . Areas o f dark o r l i g h t pigment may d i s r u p t t h i s p a t t e r n . V e n t r a l s i d e l i g h t o r w h i t e , back d a r k e r , s c a t t e r e d spots o r b l o t c h e s p e r s i s t on d o r s a l s i d e . Base o f cauda l f i n w i t h v e r t i c a l dark b a r . Another t h i c k ba r may p e r s i s t ac ross t h i r d qua r t e r o f cauda l f i n . Bars on back may expand on d o r s a l f i n i n t o l a r g e dark a r ea s . A n a l f i n l i g h t o r dark w i t h i r r e g u l a r h o r i z o n t a l b a r s . P e c t o r a l f i n mo t t l ed or b l o t c h e d w i t h i n d i c a t i o n s o f t h r e e v e r t i c a l b a r s . C h i n , b e l l y , p e l v i c f i n s , and v e n t r a l su r face o f t a i l w h i t e . Sometimes p e l v i c 62 f i n s o f females have weak spots or bars a long t he rays. . In males , many dark spots may occur on b e l l y and p e l v i c f i n s . A x i l o f p e c t o r a l f i n w h i t e w i t h c o n t r a s t i n g b l ack bars and b l o t c h e s . Leng ths - t o 36O mm. g i l b e r t i ; - Named f o r C . H . G i l b e r t . Ranges- From Hakodate, Japan, th rough the Sea o f Okhotsk, t o at l e a s t B e r i n g I s . , U . S . S . R . , and p r o b a b l y f u r t h e r n o r t h than t h i s on the S i b e r i a n c o a s t . As shown by F i g s . 9 and 10, H . g i l b e r t i tends t o show more, v a r i a t i o n than a l l t he o ther s p e c i e s . More s tudy i s needed w i t h many more specimens. P o s s i b l y t h i s v a r i a t i o n persuaded Gorbunova t o i d e n t i f y some H , g i l b e r t i as H . zapus . Tab le V . D i s t r i b u t i o n Records o f Ho g i l b e r t i L o c a t i o n Hakodata , Japan Nemurd, Hokkaido , Japan (210 f t . ) A k k e s h i Bay , Hokkaido , Japan (180 f t . ) K u s h i r o " " (240 f t . ) Muroran » » ( 2 1 0 f t . ) . Funka Bay T» »» (210 f t . ) Westhlk Bay, east Kamchatka (not t o b e . t r u s t e d ) Aniwa Bay, south S a h k a l i n " Mauka, o f f Rebunto I s . , o f f '» Moneron I s . . Nor th Japan Sea *52°43*:N. 155 42*E. . *52°20»N. 155°30«.E. *53 '°IG»N."( 52°30»N?) 155°10»E. * B e r i n g I s . , U . S . S . R *42 G I6*30"ir. 142°04*E. *Sovgavan, T a r t a r S t . n tt tt tt tt tt Reference o r c o l l e c t i o n n o . Jordan & S t a r k s , 1904 Watenabe, I960 tt tt tt tt Schmidt , 1929 it tt BC 63-675 BC 63-396 BC 63-402 U . S . N . M . 119865 U . S . N . M . 119857 U . S . N . M . 105146 63 Subgenus M e l l e t e s Bean M e l l e t e s Bean,. 1 8 8 0 , p . 3 5 4 (Type o f genus M e l l e t e s p a p i l i o Bean) ; Jordan and Evermann, 1 8 9 6 , p . 438; 1 8 9 8 , p . ' 1 9 3 2 j Schmidt , 1 9 2 9 , p . 367? So lda tov and L l n d b e r g , 1930, p . 1 9 8 ; Andr i a shev , 1 9 5 4 , p.. 365. Neohemilepidotus Sakamoto. 1 9 3 2 , p . 4 ( T y p e ' o f gehu3 Neohemilepidotus  . japonicus Sakamoto); Watanabe, 1 9 5 8 , p . 242; I 9 6 0 , p . 44. D i a g n o s i s ; - V e n t r a l s c a l e band w i t h s c a l e s much s m a l l e r than those o f o ther bands. V e n t r a l s c a l e band w i t h h o r i z o n t a l rows w i d e l y spaced between l a t e r a l l i n e and a n a l f i n . D e s c r i p t i o n s - Sca l e s o f upper band r a t h e r spaced; s m a l l but more than t w i c e t he s i z e o f those o f v e n t r a l band. U s u a l l y t h r e e but maybe t w o t o four h o r i z o n t a l rows o f s c a l e s i n upper band . V e n t r a l band o f about four h o r i -z o n t a l s c a l e rows at w ides t p o i n t . These rows and t h e a n a l s c a l e row evenly spaced between a n a l f i n and l a t e r a l l i n e . P o s t o c u l a r and o c c i p i t a l prominences or sp ines en la rged . G i l l membranes u n i t e d , w i t h wide f ree f o l d , a t tached w e l l forward on i s thmus . F i r s t t h r e e d o r s a l sp ines i nc r ea se i n h e i g h t . Marg in o f f i n membrane between f i r s t four sp ines s i m i l a r l y i n c i s e d , thus t he r e i s no apparent n o t c h . T h i s subgenus i s monotypic . Ranges- Nosappo, Hokka ido , Japan t o eas te rn B e r i n g sea and B e r i n g S t . M e l l e t e s s - Refers to//fttAMVTYir meaning " a l o i t e r e r , remaining i n sha l low p o o l s as t i d e recedes" . O N HEMILEPIDOTUS  PAPILIO. ( SOMM. | M A L E : - C O L L . NO. BC. 63-396. Figure 13 65 Hemi lep idotus ( M e l l e t e s ) p a p i l i o (Bean) ( F i g . 13) M e l l e t e s p a p i l i o " Bean, 1880, p . 354* f i g . ( S t . P a u l I s . , P r i b i l o f I s . Alaska) . ; Jordan and Evermann, 1896, p . 438; 1898, p . 1932; 1900, f i g . 703; Schmidt , 1904, p . 116; 1929, p . 367; 1950, p . 141; So lda tov and L i n d b e r g , 1930, p . 199; Andr i a shev , 1937* P» 19; 1954* p . 365, f i g . 2Q3; Ta rene t z , 1937, p . H O ; Okada and Matsubara , 1938, p . 322; Gorbunova, I964, p . 246 ( i n p a r t ) . Hemi lep ido tus g i l b e r t i Schmidt , 1929, p . 366 ( i n pa r t non Jordan and S t a r k s ) . Neohemilepidotus j apon icus Sakamoto, 1932, p . 4, f i g . 2 (Nor the rn .Japan, l o c a l i t y unknown); Watanabe, 1953, p . 242. Neohemilepidotus p a c i f i s u s Watanabe.,1958. p . 15, f i g . 9 (Neohemilepidotus  pac i f l eas Watanabe « Meohemilep i d o t us .japonicus Sakamoto); I960, p . 45, p i . 21 - f i g . 2 . Gorbunova v s specimen has 11 d o r s a l sp ines a l though most H . p a p i l i o have 12 . The specimen shown i n F i g . 22 (BC 62-847) has a double o c c i p i t a l s p i n e , a l though i t does not have a pigmented u r o s t y l e ; bo th cha rac t e r s are used by Gorbunova t o i d e n t i f y H . p a p i l i o . S i m i l a r l y , even the f i g u r e o f Gorbunova does not show a pigmented u r o s t y l e . A l s o , the unnotched f i r s t d o r s a l f i n i s present i n a l l specimens o f t he Hemi lep idot i r . ae which are under about 17 mm. The i d e n t i f i c a t i o n s and records o f H . p a p i l i o by Gorbunova, I964, must t h e r e f o r e be r e j e c t e d . Because o f t he absence o f c i r r i , p a p i l l a e on p e l v i c f i n s and shape o f t h e d o r s a l f i n , H . j apon icus might be a v a l i d s p e c i e s . However, more than t h e o r i g i n a l two specimens must be examined f u r t h e r . Watanabe has c e r t a i n l y confused t he s i t u a t i o n . Types- Type specimen ca ta logued as U . S . N . M . No. 23751. D i a g n o s i s ; - That o f the subgenus. M a t e r i a l s , ; - F i v e specimens were a v a i l a b l e from ca ta logue No. BC 63-400, BC 63-396 and two c o l l e c t i o n s not ye t ca ta logued (W63-55 and Hau l 25 A r t h u r H . ) . 66 These c o n s i s t o f four males o f 128 mm t o 273 mm i n s tandard l e n g t h and one female 216 mm. D e s c r i p t i o n s - Dorsa l s X I I [ X I ] 2 , 20 (19) [ 19-22] 2 . Anals 16-17 [ 1 8 ] 2 . P e c t o r a l s 17-18 [ 16] 2 . Ver tebraes 39. L a t e r a l l i n e poress 56.6 (49-65). Transverse s c a l e rows i n d o r s a l bands 65.2 (63-69). Number o f s c a l e s i n row above l a t e r a l l i n e s 20 .8 (18-26). Number o f h o r i z o n t a l rows i n d o r s a l s c a l e bands 3-4. P y l o r i c caecas 6 (5) [ 7 ] ^ . Width immediate ly beh ind p e c t o r a l f i n s s 190 (172-224). Depth from o r i g i n o f d o r s a l f i n t o a n t e r i o r base o f p e l v i c f i a s 237 (219-251). Depth from o r i g i n o f so f t d o r s a l f i n t o o r i g i n o f a n a l f i n s 220 (209-240). Head lengths 369 (359-387). O r b i t diameters 88 (76-99). O r b i t t o end o f operc les 189(173-205). I n t e r o r b i t a l widths 7 2 ( 6 6 - 8 2 ) . Snout l e n g t h ( e x c l u d i n g p r e m a x i l l a e ) s 74 (67 -82) . Narrowest l a c r i m a l widths 29 (26-33). Cauda l peduncle lengths 117 (114-121). Caudal peduncle depths 63 (59-68). Snout t o o r i g i n o f d o r s a l f i n s 306 (289-320). Snout t o o r i g i n o f a n a l f i n s 604 (596-624). Length o f d o r s a l f i n bases 634 (61B-658). Height o f longes t sp ine on d o r s a l f i n o f females 160. Height o f longes t sp ine on d o r s a l f i n o f maless 221-276. Length o f longes t so f t r ay on d o r s a l f i n s 166 (151-179). Length o f a n a l f i n bases 299 (275-316). Length o f longes t r ay on a n a l f i n s 127 (113-151). Length o f base o f p e c t o r a l f i n s 156 (141-171). Length o f longes t r ay on p e c t o r a l f i n s 335 (311-352). Length o f longes t r ay on p e l v i c f i n s o f four maless 418 (330-482). Length o f longes t r ay on p e l v i c f i n s o f one females 219. Length o f longes t r ay on cauda l f i n s 204 (197-213). Head depressed. I n l a t e r a l v i e w , p r o f i l e o f o c c i p u t s l i g h t l y concave and s l o p i n g i n pa th wh ich would p r o j e c t th rough upper r i m o f o r b i t . O r b i t s o n l y s l i g h t l y e l e v a t e d . P r o f i l e o f snout s t r a i g h t , not s l o p i n g s t e e p l y ; 67 convex d o r s a l l y , at l e v e l o f n a s a l s p i n e s . P r o f i l e between o r b i t and n a s a l sp ines s l o p i n g o n l y s l i g h t l y . P r o f i l e from t i p o f lower jaw t o isthmus s t r a i g h t . Body deepest under t he fou r th t o s i x t h sp ine o f d o r s a l f i n . Nape s h o r t , s t r a i g h t , concave at edge o f o c c i p u t . Base o f d o r s a l f i n s t r a i g h t between s i x t h sp ine and l a s t so f t r a y ; convex a n t e r i o r l y . P r o f i l e o f d o r s a l arid v e n t r a l s i d e o f c a u d a l peduncle s l i g h t l y concave. Base o f a n a l f i n s l i g h t l y convex.'" P r o f i l e from a n a l f i n t o Isthmus s t r a i g h t or convex but l a r g e l y h idden by p e l v i c f i n s i n males . From d o r s a l v i e w , t he l a t e r a l p r o f i l e i s s t r a i g h t t o narrow cauda l pedunc le . A n t e r i o r n o s t r i l s h o r t , t u b u l a r , r i m on p o s t e r i o r s i d e r a i s e d about o n e - h a l f t o one t imes t h e he igh t o f a n t e r i o r s i d e o f n o s t r i l . N o s t r i l d i r e c t l y l a t e r a l t o n a s a l sp ines and a n t e r i o r t o l a t e r a l edge o f i n t e r o r b i t . P o s t e r i o r n o s t r i l b road , l a r g e r than a n t e r i o r one, n a s a l opening c o n s t r i c t e d . T h i s n o s t r i l d i r e c t l y m e d i a l t o a n t e r i o r edge o f o r b i t and p o s t e r i o r t o l a t e r a l edge o f n a s a l s p i n e . Head pore about o n e - t h i r d t o o n e - h a l f d i s t a n c e between p o s t e r i o r t o a n t e r i o r n o s t r i l ; pore at t i p o f s m a l l p a p i l l a e . I n t e r -o r b i t ve ry s h a l l o w l y concave. L o n g i t u d i n a l p a i r o f weak r i d g e s i n i n t e r -o r b i t . Spines on head prominent . N a s a l sp ines s h o r t e r , sharp . N a s a l sp ines a t l e v e l midway between o r b i t and m i d l i n e and o n e - h a l f d i s t a n c e from o r b i t t o snout . P r e o p e r c u l a r sp ines s t r o n g , sharp; upper sp ine d i r e c t e d p o s t e r o -d o r s a l l y ; next, one d i r e c t e d p o s t e r i o r l y t o s l i g h t l y d o r s a l l y , and i s s l i g h t l y l o n g e r t han upper one; t h i r d sp ine s h o r t , sharp , and d i r e c t e d p o s t e r i o r l y ; f o u r t h s p i n e at base o f p r e o p e r c l e and p o i n t e d v e n t r a l l y . , Short sharp sp ine on upper p o s t e r i o r edge o f o p e r c l e . P o s t e r o - m e d i a l and po , s t e ro -ven t r a l sp ines on .subopercle which i s fused t o o p e r c l e . S m a l l weak sp ine on p o s t e r i o r end o f i n t e r o p e r c l e . P o s t o c u l a r processes prominent and rounded. O c c i p i t a l 68 processes prominent , l o w , l o n g , and rounded, F r o n t o - p a r i e t a l , pos t t empora l and p t e r o t i c , and s u p r a c l e i t h r a l r i d g e s p resen t ; i n t e r v e n i n g areas markedly depressed . Ope rcu la r r i d g e w e l l deve loped , S t r i a t i o n s v i s i b l e o n l y where they r a d i a t e from p o s t o c u l a r p r o c e s s e s . C i r r i t end t o be much more reduced or absent . C i r r i below n a s a l sp ines v e r y s m a l l and t u b e r c l e - l i k e . N a s a l c i r r i appear t o be absent . C i r r i on p o s t e r i o r s i d e o f p o s t o r b i t a l p rocess h i g h l y reduced i n s i z e , t h i n , b road , weakly m u l t i f i d . F r o n t a l c i r r i absent o r ext remely reduced and s i m p l e . Three t o f i v e v e r y s m a l l t u b e r c l e - l i k e c i r r i under l a c r i m a l p o r t i o n o f s u b o r b i t a l r i d g e ; may be another one below s u b o r b i t a l s t ay p o r t i o n o f r i d g e . C i r r i near p o s t e r i o r end o f s u b o r b i t a l s tay reduced and almost t u b e r c l e - l i k e . S m a l l , f l a t , weakly m u l t i f i d c i r r i near upper end o f g i l l s l i t , on operculum. P a i r o f s m a l l , s t o u t , b a r b e l - l i k e c i r r i immediately l a t e r a l t o median p a i r o f c h i n p o r e s , ^ a i r o f v e r y s m a l l t u b e r c l e - l i k e c i r r i at p o s t e r i o r t h i r d o f a n t e r i o r face on lower jaw. Ve ry s m a l l , narrow, s i m p l e , c i r r i on m a x i l l a r y . Sca le s w i t h up t o one dozen s p i n u l e s . S c a l e band conf ined t o upper t h i r d o f d i s t a n c e between l a t e r a l l i n e and d o r s a l f i n . D o r s a l band c l o s e t o base o f d o r s a l f i n but d i s t a n c e between f i n and s c a l e band may be v a r i a b l e a n t e r i o r l y . S c a l e rows on d o r s a l band s t r a i g h t p o s t e r i o r l y and may extend onto c a u d a l pedunc le . S c a l e s w e l l spaced and separate from o thers on the same o r d i f f e r e n t rows. P e r s i s t e n c e o f s c a l e s i n each row h i g h l y v a r i a b l e so t h a t two, t h r e e , o r f o u r , rows o f s c a l e s may be seen under spinous d o r s a l f i n . Only one row p e r s i s t s onto cauda l pedunc le . , . ; Tubu la r bony elements o f l a t e r a l l i n e w i t h on ly a few c t e n i on the p o s t e r o - d o r s a l s i d e . Row o f s c a l e s above l a t e r a l l i n e separate from l a t e r a l l i n e by s e v e r a l s c a l e d i ame te r s . Often a few s c a l e s above t h i s row. S m a l l s c a l e s i n s m a l l pa t ch at a x i l o f p e c t o r a l f i n . S c a l e s o f v e n t r a l band much s m a l l e r and reduced , appear ing p a p i l l a t e . V e n t r a l band about two 69 rows wide on s i d e o f b e l l y , expanding t o four more d e f i n i t e rows above o r i g i n o f a n a l f i n , d i m i n i s h i n g t o one or two rows on cauda l pedunc le . Sca les on v e n t r a l band w i d e l y spaced . A n a l s c a l e row converges ahead o f anus and proceeds up m i d l i n e onto b r e a s t . A l l s c a l e rows between a n a l f i n and l a t e r a l l i n e evenly spaced. L a t e r a l l i n e w i t h a s l i g h t upward curve somewhere over a n t e r i o r h a l f o f a n a l f i n . Caudal peduncle b i s e c t e d i n t o equal p o r t i o n s by l a t e r a l l i n e . O r i g i n o f s p i n y d o r s a l f i n over p o s t e r i o r h a l f o f p e c t o r a l f i n . T h i r d or f o u r t h sp ine t a l l e s t . Fourth sp ine over p o s t e r i o r l i m i t o f operculum. A l l sp ines become p r o g r e s s i v e l y s h o r t e r on e i t h e r s i d e o f t a l l e s t s p i n e . Marg in o f d o r s a l f i n membrane deeply i n c i s e d between f i r s t four s p i n e s ; s h a l l o w l y i n c i s e d between r e s t o f s p i n e s . Las t d o r s a l f i n sp ine q u i t e s h o r t . Four th and f i f t h sof t d o r s a l rays t a l l e s t , rays on each s i d e become n o t i c e a b l y s h o r t e r . Depressed d o r s a l f i n rays extend t o p o i n t w e l l short, o f c auda l f i n base . Marg in o f so f t d o r s a l f i n membrane weakly i n c i s e d t o e n t i r e . O r i g i n o f a n a l f i n under about the t h i r d o r f o u r t h so f t d o r s a l r a y ; l a s t ray under l a s t so f t d o r s a l f i n r a y . Second t o e i g h t h a n a l f i n ray t a l l e s t ; those p o s t e r i o r t o t h i s become p r o g r e s s i v e l y s h o r t e r . Marg in o f a n a l f i n membrane g r e a t l y i n c i s e d , e s p e c i a l l y a n t e r i o r l y . No branched d o r s a l and a n a l f i n r a y s . Cauda l f i n t r u n c a t e . P e c t o r a l f i n rounded; e l even th t o t h i r t e e n t h ray from bot tom l o n g e s t . Marg in o f membrane on p e c t o r a l f i n s l e s s i n c i s e d . V e n t r a l l y membrane near margin o f p e c t o r a l f i n s l i g h t l y overgrown so tha t i t i s not f u l l y s t r e t c h e d when rays are spread a p a r t a P e l v i c s normal and i n c i s e d . I n m a l e s , t he d o r s a l and p e l v i c f i n s become g r e a t l y m o d i f i e d . The second t o seventh ( e s p e c i a l l y second t o f i f t h ) d o r s a l f i n sp ines become much l o n g e r (as much as t w i c e t he he igh t o f so f t d o r s a l f i n ) . Membrane between second and f o u r t h sp ines enormously i n c i s e d f o r t w o - t h i r d s t he l e n g t h o f t he 70 spineo F i n membrane i s overgrown near margin t o produce a l oose f ree f l a p behind the s p i n e . P e l v i c f i n s ve ry l o n g and extend p o s t e r i o r l y past t i p s o f p e c t o r a l f i n s . Membrane between rays i n c i s e d f o r more than o n e - t h i r d o f l e n g t h o f p e l v i c f i n r a y s . On m e d i a l - v e n t r a l su r face o f p e l v i c f i n r a y s , many l o n g , bony, c t e n i develop from su tu re between each a c t i n o t r i c h . C t e n i narrow; may be l o n g e r t h a n w i d t h o f t h i c k f i n r a y . Base o f p e l v i c f i n broader than t ha t o f female. C o l o u r p a t t e r n o f dark pigment v a r i e s . C h a r a c t e r i s t i c four bars ac ross back o f t en p r e s e n t . These bars between f i f t h and e i g h t h d o r s a l f i n s p i n e , between t h i r d and s i x t h so f t d o r s a l f i n r a y , between t e n t h and t h i r t e e n t h sof t d o r s a l f i n r a y , and between seventeenth t o l a s t sof t d o r s a l r a y . Other b l a c k markings p r e s e n t , e s p e c i a l l y on f i n s , v a r i a b l e . Body may be u n i f o r m l y c o l o u r e d . Spots and b l o t c h e s most ly absent . P e l v i c f i n s and under p a r t s w h i t e . In males , m o d i f i e d p o r t i o n o f s p i n y d o r s a l f i n b l a c k . P e c t o r a l f i n dark w i t h two o r t h r e e w h i t e v e r t i c a l s t r e aks or bars which are s h a r p l y s i l h o u e t t e d w i t h b l a c k . P e l v i c f i n s w i t h compl i ca t ed p a t t e r n s o f w h i t e and b l a c k bars or s t r e a k s . L e n g t h ; - To about 300 mm. p a p i l i o : - Meaning " b u t t e r f l y " i n re fe rence t o the en la rged p e l v i c f i n s o f t h e male . There i s l i k e l y some v a r i a t i o n which i s not shown by these f i v e specimens. Andr iashev (1954) r ecords d i f f e r e n c e s i n v e r t e b r a l counts o f H . p a p i l i o i n t he B e r i n g Sea and Sea o f Okhotsk. The specimens o f Sakamoto (1932) i l l u s t r a t e t h i s v a r i a b i l i t y . O b v i o u s l y , much more study o f t h i s s p e c i e s i s needed. 71 Tab le V I . D i s t r i b u t i o n Records o f H . p a p i l i o L o c a t i o n ( t i d e p o o l s ) (73-310 f t . ) S t . P a u l I s . , A l a s k a B e r i n g S t . K o r i a k Land U . S . S . R . Anadyr G u l f ' U . S . S . R . K r u s t e r n I s . U . S . S . R . Tkachen Bay (?67°25 tN„ 172°48*E.) = H . g i l b e r t i o f Schmidt . 1929 Bay o f Providence = H . . jordani o f Bean Tauyskaya Guba E r i n e y s k a y a Guba Perishinskaya Guba " Turpehya Bay Nossappim, Hokkaido , *52°20»N. 155°30 ? E. *60°41 ' 'N. 174°25 9 E. *east B e r i n g Sea? #east B e r i n g Sea Reference or c o l l e c t i o n no . Bean, 1880 Andr i a shev , 1937 it n Barsukov , 1958 Shantar I s . (not t o be t r u s t e d ) Schmidt , 1929 t» tt tt tt tt it t» tt tt tt tt tt tt Japan Watanabe, I960 BC 63-396 BC 63-400 A r t h u r H . H a u l 125 W63-55 IDENTIFICATION OF THE SPECIES C o n s i d e r i n g t h a t Schmidt synonymized H . hemi lep ido tus and H . . jo rdan i . and Gorbunova synonymized H . zapus and H . g i l b e r t i , t he d i f f e r e n c e s between these spec ies shou ld be s t r e s s e d . P o s t l a r v a e Larvae and p o s t l a r v a e change g r e a t l y i n body form d u r i n g ontogeny. M e r i s t i c s and s c a l e p a t t e r n s are s t a b l e a f t e r t h e i r i n i t i a l fo rmat ion and a re consequent ly p r e f e r a b l e f o r t he i d e n t i f i c a t i o n o f s p e c i e s . Few specimens wi thou t s c a l e s were a v a i l a b l e f o r s tudy so t ha t t he key presented here i s in tended f o r l a r g e r p e l a g i c p o s t l a r v a e ( i . e . 16 o r 20 mm and o v e r ) . There were no p o s t l a r v a e o f H . g i l b e r t i o r H . p a p i l i o a v a i l a b l e f o r t h i s s tudy . For reasons mentioned p r e v i o u s l y , Gorbunova 's (1964) specimens are not adequate ly i d e n t i f i e d . However her i l l u s t r a t i o n o f a 17.5 mm H . g i l b e r t i i s 72 d i f f e r e n t t o any specimens examined i n t h i s s tudy . Consequently t h i s i d e n t i f i c a t i o n i s p robab ly c o r r e c t (see f i g . 12). Without a complete range o f specimens from egg t o a d u l t , i t i s u n -p r o f i t a b l e t o d e s c r i b e the l a r v a e and p o s t l a r v a e i n great d e t a i l , s i n c e a l l o m e t r i c growth i 3 so g r e a t . Xn g e n e r a l the l a r v a e and p o s t l a r v a e are more compressed, t h e eyes are l a r g e r and l a t e r a l l y p l a c e d , sharp sp ines appear on top o f t he head, c i r r i a re absent , the c a u d a l f i n becomes f o r k e d , a t s m a l l e r s i z e s t he spinous d o r s a l f i n i s unnotched, and the i n t e r - o r b i t a l i s much w i d e r . The i n t e r o r b i t a l becomes r e l a t i v e l y narrower when t h e p o s t -l a r v a e i s about 30 mm. At the same t i m e , major changes appear ( i . e . the eyes are p l a c e d h i g h e r and the body becomes more depressed) and t he p o s t -l a r v a e i s then p robab ly more adapted t o a bottom h a b i t a t . T h i s change p r o v i d e s a d e f i n i t i o n between p o s t l a r v a e and j u v e n i l e s (see f i g . 14). Accord ing t o K o b a y a s h i , p o s t l a r v a e may reach 38 mm (Gorbunova, I964). Consequently the t ime o f metamorphosis i s p robab ly more v a r i a b l e than t ha t i n d i c a t e d by t he da ta i n t h i s t h e s i s . The a n a l s c a l e row i s present i n the p o s t l a r v a e o f a l l s p e c i e s , a l though i t i s l o s t i n the a d u l t s o f some. Key t o t h e P o s t l a r v a e : -The minimum s i z e s o f the p o s t l a r v a e at which each cha rac t e r i s known t o be present a re i n d i c a t e d i n t he b r a c k e t s . A , Isthmus fused w i t h g i l l membranes not p roduc ing a f ree f o l d (down t o at l e a s t 20 mm); 6 or more h o r i z o n t a l rows i n d o r s a l s c a l e band (down t o at l e a s t 20 mm) see f i g . 15 . . . . . . . . . . . . . H . spinosus AA. Isthmus fused t o g i l l membranes but p roduc ing a f ree f o l d p o s t e r i o r l y ; 5 .o r l e s s s c a l e rows i n d o r s a l s c a l e band. 73 A P P X . 30 M M . S T A N D A R D L E N G T H Figure 14. Suggest ed.;st ages of l i f e history, in the Hemilepidotinae. j . B. Unnotched spinous dorsal fin; about 4 rows.,,.of scales in ventral band, these scale rows evenly spaced between lateral line and anal scale row,; 12 spines in first dorsal fin (rarely 11). (Time at which characters appear unknown, dorsal spines probably at 12 to 14 mm) . . . . . H. papilio BB. Notched spinous dorsal fin (16-20 mm); 6 or more rows of scales in ventral band; these scale rows l i e closer to lateral line than anal scale row; 11 spines in spinous dorsal fin (very rarely 12). C. First spine of dorsal fin longer than second spine (probably at larger sizes only); (specimen wil l not have scales on caudal peduncle, between dorsal scale band and lateral line; it will not have IS or more pectoral fin rays. The illustration by Gorbunova (1964) shows, exceptionally long occipital spines which were not nearly as long on similar sized specimens of H. zapus. H. hemilepidotus and H. jordani), (Time of formation of these characters unknown) see fig. 19, 21, 22, 25 and 26 . . . . . . . . . . . H. gilberti 74 CC. First dorsal spine shorter than second spine. D. 58 or fewer lateral line pores; (specimen wil l have scales be-tween lateral line and dorsal scale band on caudal peduncle) (down to at least 17.5 nni) see fig. 16, 21 and 22 DD. 59 or more lateral line pores. E. 16 or fewer pectoral rays or i f 17 then total of soft dorsal, anal, and both pectorals is less than 70 rays (down to at least 14 mm); vertebrae 36 or fewer (down to 13 mm) see fig. 17, 18, 19, 29 and Table V I I . . . . . . . H. hemilepidotus EE. 18 or more pectoral rays or i f 17 then total number of soft dorsal, anal, and both pectorals is 70 or more rays.(down to at least 17 mm); vertebrae 36 or more Key to the Juveniles and Adultsi-The key to the postlarvae applies equally well here. However, other characters are more useful in larger specimens. A. G i l l membranes fused to isthmus without a free fold posteriorly; 6 or more scale rows in the dorsal scale band see fig. 15 H. spinosus AA. G i l l membranes form free fold posterior to their attachment to the isthmus; 5 or fewer scale rows in the dorsal scale band. B. Scales of ventral band less than half size of those of dorsal band (at least in specimens of 127 mm, probably down to 40 or 50 mm) . . . . . . . . . . H a papilxo BB. Scales of dorsal band and ventral band of about the same size. C. First dorsal fin spine always longer than the second spine 75 (down t o at l e a s t 40 mm); lower p r e o p e r c u l a r sp ine s p a t u l a t e and w i t h - s m a l l barbs on the, edge (down t o at l e a s t 190 mm); f r o n t a l c i r r i reduced o r absent (see f i g s . 19, 21, 22, 25 and 26) F i r s t d o r s a l f i n sp ine u s u a l l y s h o r t e r than second d o r s a l s p i n e ; lower p r e o p e r c u l a r sp ine s imple (except i n a few l a r g e H . hemi lep ido tus and r a r e l y i n H . . j o rdan i ) ; f r o n t a l c i r r i developed and m u l t i f i d i n l a r g e r specimens (except some H« zapus ) . Do L a t e r a l l i n e w i t h 58 o r fewer p o r e s ; s ca l e s always between d o r s a l s c a l e band and l a t e r a l l i n e on cauda l pedunc le ; a n a l s c a l e row always present (see f i g s . 16, 21 and 22) e e o e . o o . . . . . Ho ZapUS DD. L a t e r a l l i n e w i t h 59 o r more po re s ; s c a l e s absent between d o r s a l s c a l e band and l a t e r a l l i n e on cauda l peduncle (sometimes present on H . h e m i l e p i d o t u s . extremely r a r e l y on H . . j o rdan i ) ; a n a l s c a l e row absent (sometimes present i n H . j o r d a n i , ext remely r a r e l y i n H . hemilep i d o t u s ) . E . Width o f m a x i l l a r y c i r r i l e s s than h a l f i t s l e n g t h (above 50 mm); n a s a l c i r r i s i m p l e ; s c a l e s never between d o r s a l s c a l e band and l a t e r a l l i n e on cauda l pedunc le ; d o r s a l and a n a l f i n rays s i m p l e ; p y l o r i c caeca 4-5. (see f i g s . 16-36) . . . . . . . . . . . H . .j o r d a n i E E . Width o f m a x i l l a r y c i r r i more than h a l f i t s l e n g t h ; n a s a l c i r r i s t r o n g l y m u l t i f i d (above 60 o r 70 mm); s c a l e s some-t imes present between d o r s a l s c a l e band and l a t e r a l l i n e on c auda l pedunc le ; d o r s a l and a n a l f i n rays branched (above 140 mm); p y l o r i c caeca 6-7. (see f i g . 20) . . . . . . . . . H . hemi lep ido tus 76 Key C h a r a c t e r s ; - . The graphs on the f o l l o w i n g pages represent da ta on which many o f the cha rac te r s used i n t he keys were based . Growth C h a r a c t e r i s t i c s and P r o p o r t i o n a l Measurements:-In order t o d i s t i n g u i s h t h e spec ies o f the Hemilep ido t i n a e , i t i s important t ha t t he cha rac te r s used app ly t o as many i n d i v i d u a l s and t h r o u g h -out as much o f ontogeny as p o s s i b l e . Many workers have a t tached importance t o the s p e c i a l i z e d p e l v i c f i n o f males , c o l o u r p a t t e r n , and p r o p o r t i o n a l measurements. These cha rac t e r s maybe v a r i a b l e acco rd ing t o h a b i t a t , sex , o r age. The m e r i s t i c or s t r u c t u r a l cha rac t e r s p r e v i o u s l y d i s c u s s e d are much more u s a b l e . In p a r t i c u l a r , p r o p o r t i o n a l measurements have been commonly used . Eye d iamete r , i n t e r o r b i t a l w i d t h and snout l e n g t h were used by Rendahl (1931) and contemporary workers t o i d e n t i f y H . h e m i l e p i d o t u s , H . zapus , H . . jo rdani and H, g i l b e r t i . I f such measurements are used , t he degree o f a l l o m e t r i c growth must be de te rmined . These cha rac t e r s demonstrate s i g n i f i c a n t changes o f r e l a t i v e growth i n t he m a j o r i t y o f t he H e m i l e p i d o t i n a e . As shown i n F i g . 37, eye diameter o f H . hemi lep ido tus ove r l aps o r converges w i t h t ha t o f H. , jordani at standard, l eng th s under 110 mm. At much l a r g e r s i z e s H . h e m i l e p i -dotus has an i n f l e c t i o n which i l l u s t r a t e s reduced growth o f the eye . The i n t e r o r b i t a l w i d t h shows a l l o m e t r i c growth , ( i . e . H . h e m i l e p i d o t u s , H . zapus , H , . jo rdan i and H . sp inosus - f i g . 38, 39, 40 and 41). An. i n f l e c t i o n i s most n o t i c e a b l e i n H . sp inosus and H . zapus wh ich possess t he narrowest w i d t h d u r i n g t h e adu l t s t age . The data from the p l o t s on the l e f t o f these graphs represent p o s t l a r v a e i n which the eye i s l a t e r a l l y p l a c e d . Meta-morphosis i n body form f o r adap t ion t o a bot tom h a b i t a t occurs at a p p r o x i -mate ly 30 mm s tandard l e n g t h (see f i g . 14). The eyes are then h ighe r on the 77 H O R I Z O N T A L S C A L E R O W S . <> ^ c > 6 • — - - - f -7 - - * '> S P I N O S U S . .. J A • 1 J H E M I L E P I D O T U S , A • • — Z A P U S J O R D A N I ^ G I L B E R T I — -— -P A ' P I L I O , i •i • i I • — \ F i g u r e 15<> Approximate range and means o f number o f h o r i z o n t a l s c a l e rows i n d o r s a l s c a l e band o f the Hemi l ep idd t inae . LATERAL LINE PORES 55 .60 ' 65 70 • • • i • * : i' ' ' S P I N O S U S , .50 i l H E M I L E P I D O T U S U M N A K P R I B I L O F . V A N C O U V E R IS. f ZAPUS Figu re 16. D i s t r i b u t i o n o f l a t e r a l l i n e coun t s . 78 S P I N Q S U S H E M I L E P I D . Z A P U S  J O R D A N I  G I L B E R T I  P A P I L I O ANAL FIN 13 14 15 -j 'JJ^kl RAYS 16 1 17 uM L F T 18 19 Figure 17* D i s t r i b u t i o n o f number o f a n a l f i n r a y s » S O F T D O R S A L FIN RAYS i 1,8 19 2 0 2 1 2 2 S P I N O S U S • j i wkm i - j i i H E M I L E P I D . T * ! Z A P U S ! i : J O R D A N I i \- A • G I L B E R T ] 1 " f 1 1 B P A P I L I O ! 1 A 1 ! ' • 1 Figure 18. D i s t r i b u t i o n o f number o f so f t d o r s a l f i n r a / s , Table VII: Character index for H.spinosus, H.zapus, H.gilberti, and H.papilio applied to data for H.hemilepidotus and H.jordani from Schultz arid Welander (1934). Cumulative count of anal, soft dorsal and both pectoral fins 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 Total H.spinosus 2 8 3 13 H.hemilepidotus 3 17 15 45 8 1 89 H.zapus 1 3 4 3 1 1 13 H.jordani 9 32 28 43 16 3 1 1 133 H . gilberti 1 1 1 1 1 4 9 H.papilio 1 1 3 2 7 80 PECTORAL FIN RAYS 14 SPlNOSUS  HFM1I FP inOTUS Z A P U S JORDANI G ILBERT! PAPILIO 15 16 17 18 I 9 'F igure 19« D i s t r i b u t i o n o f number o f p e c t o r a l f i n r a y s . 20 • LT) i j . Z ^ O O o: cc 15 NUMBER FRINGES NASAL Ci 10 5 • — • • • • • • • — • • • • • • •• • t i | i i i i i i 1 i i i l i i i I i 1 i i i i i i i i \ 10 20 30 STANDARD LENGTH IN CM. F i g u r e 20. Number o f f r i n g e s on n a s a l c i r r i o f H . hemi lep ido tus 81 S P I N O S U S H E M I L E - P f D O T U S S C A L E S B E T W E E N L A T L I N E A N D DORS. B A N D S C A L E S B E L O W D O R S A L B A N D N O S C A L E B E T W E E N L A T LINE A N D D O R S . B A N D Z A P U S * 1 » J O R D A N I  G I L B E R T I  P A P I L I O Figure 21. Extent of scales between level of lowest row on dorsal scale band and lateral line of caudal peduncle. Figure 22. Position of scales on caudal peduncle in the Hemilepidotinae "A" - Posterior limit of scales in H. .jordani. H. gilberti and H. papilio. "B" - Lower limit of scales .in many H. hemilepidotus. "Cn - Scales extend to here in H. spinosus, H. zapus and many H. hemilepidotus. "D" - Anal scale row extends to here only in H. spinosus. M A X I L L A R Y C I R R U S W I D T H / C - I R R U S L E N G T H . 3 0 .40 . 5 0 .60 7 0 . 8 0 H E M I L E P I D O T U S J O R D A N I  G I L B E R T I 1 , f ft 1 A Figure 23: Distribution of proportion of maxillary c irr i width to maxillary c irr i length M E M B R A N E - H E I G H T S P I N Q S U S  H E M I L E P I D O T U S Z A P U S J O R D A N I G I L B E R T I A N T E R I O R TO 4TH S P I N E / 4 T H D O R S A L F I N S P I N E L E N G T H -40 J 5 0 _ £ 0 _ 7 f 0 £ p 3Q ! A - ' Figure 24 : Distribution of proportion of height of fia membrane on anterior side of fourth dorsal spine compared to height of fourth dorsal spine. \ 83 hi. G I L B E R T I O T H E R HEMILEPIDOTINAE Figure 25. Diagram showing r e l a t i v e l eng ths o f f i r s t and second sp ines o f d o r s a l f i n . 1ST D O R S A L S P I N E L E N G T H y ^ N D S P I N E L E N G T H .75 .85 .95 1.05 1.15 H E M I L E P I D O T U S Z A P U S J O R D A N I G I L B E R T I A. F i g u r e 26. D i s t r i b u t i o n o f r a t i o o f f i r s t d o r s a l spine l e n g t h d i v i d e d by second sp ine l e n g t h . 84 6.0 S P I N O S U S c H E M I L E P I D O T U S Z A P U S  J O R D A N I  G I L B E R T I  P A P I L I O Figure 2"\ D i s t r i b u t i o n o f number o f t r a n s v e r s e rows i n d o r s a l s c a l e band. . . . " A . , S P I N O S U S Z A P U S  J O R D A N I  G I L B E R T ]  P A P i t 10 Figure 28 . D i s t r i b u t i o n o f number o f s c a l e s above l a t e r a l l i n e . 85 H . S P I N Q S U S H . H E M I L E P I D O T U S H . Z A P U S H . J O R D A N I B G I L B E R T I ]H P A P I L I O N O . O F V E R T E B R A E 3f(4 3i5 3 6 3.7 3,8 3 9 I X A 4 0 Figure 29 . Range and approximate means o f ve r t eb rae coun t s . F igure 30. D i s t r i b u t i o n o f r e l a t i v e head, l e n g t h . 86 CAUDAL PEDUNCLE LENGTH .06 • .07 .08 .0.9 .10 .1,1 .12 S P I N O S U S ! 1 HEMILER : , Z A P U S 1 , 1 JORDANI i • m i G ILBERTI ! A P A P I L I O 1 ' ! , ' A i ! Figure 31° D i s t r i b u t i o n o f c a u d a l peduncle l e n g t h . CAUDAL PEDUNCLE DEPTH .0,5 .06 .0 7 .08 SP INOSUS HEM I LEP IDOTUS Z A P U S r JORDANI GILBERTI PA PILIO i milk i i Figure 32. D i s t r i b u t i o n o f c a u d a l peduncle dep th . 87 • 2 ? S P I N O S U S H E M I L E P I D O T U S  Z A P U S , J O R D A N I  G I L B E R T I P A P I L I O B O D Y .25 D E P T H • 2 / 2? .3J H-t-F i g u r e 33. D i s t r i b u t i o n o f body dep th . L E N G T H O F P E C T O R A L .24 .26 .28 . 3 0 .32 SPINQSUS] k ,_].. H E M I L E R Z A P U S  J O R D A N I Cf J U V ^ G I L B E R T I P A P I L I O F I N .34 .36 rai 1 i i I i ! • t " A i Figure 34. D i s t r i b u t i o n o f p e c t o r a l f i n l e n g t h . 88 H E I G H T O F S P I N Y D O R S A L F I N S P I N O S U S H E M I L E P ID. Z A P U S J O R D A N I  G I L B E R T I P A P I L I O ' F igure 35» Range and mean o f longes t sp ine on d o r s a l f i n . P Y L O R i C C A E C A 3 I •t c ) i ( S P I N O S U S j H F M I I E P I D O T U S A Z A P U S i L _ _ . J O R D A N I G I L B E R T I A ! j i _ ... P A P I L I O i i A Figu re 36. Range and approximate mean o f number o f p y l o r i c caecae. 89 t h e head and t he i n t e r o r b i t a l w i d t h i s co r r e spond ing ly nar rower . Such changes i n p r o p o r t i o n a l measurements make them unsafe f o r the i d e n t i f i c a t i o n o f specimens. The que r i ed p o i n t i n F igures 38, 39> 40 and 41 represents a ve ry s m a l l specimen (see f i g . 5) which i s l i k e l y H . hemi lep ido tus but has been i n c l u d e d on a l l the graphs . Gorbunova (1964) p resen ts i n Tabu la r form the " p l a s t i c " cha rac te r s o f t h e l a r v a e o f 5 s pec i e s o f Hemi lep ido tus . H . g i l b e r t i and H . zapus which she des igna tes as subspecies are d i f f e r e n t i a t e d by these cha rac t e r s ( i t i s p o s s i b l e she d i d not have H . zapus ) . For each s i z e ca t ego ry , on ly range and mean a re g i v e n . I have p l o t t e d these ranges o f hers ( f i g s . 42, 43» 44, 45, 46 and 47) which comple te ly merge between each s p e c i e s . S i m i l a r l y , H . ,j or dan i and H . p a p i l i o merge w i t h the range o f the above s p e c i e s . I f these da ta were s i g n i f i c a n t t h e n fu r t he r s t a t i s t i c a l parameters must be g i v e n t o show t h i s . Gorbunova 7 s da ta does not segregate t he spec ie s and can not be used t o i d e n t i f y specimens o r d i s t r i b u t i o n a l r e c o r d s . Moreover , any i n d i c a t i o n o f a r i t h m e t i c s i g n i f i c a n c e p robab ly r e f l e c t s t he d i f f e r e n t i a l s i z e range between specimens o f each s p e c i e s . A l l o m e t r i c growth i s p robab ly one c a u s a t i v e f a c t o r f o r some o f these d i f f e r e n c e s . 2 5 r 2 0 -r -Ld 15 < Q 10 LU LU • = K J O R D A N I © = H . H E M I L E P I D O T U S © 9 ®© 5 - §?>© © © ,0 © u J I I I L i I I I |_ © © © © 5 0 J I I L 1 0 0 150 S T A N D A R D L E N G T H IN M M . 2 0 0 © _l L Figure 37 : Relative growth of eye diameter in H. jordani and H. hemilepidotus I N T E R O R B I T A L W I D T H IN M M . 16 Figure 39: Relative growth of interorbital width in postlarvae of H. hemilepidotus . r N T E R O R B I T A L W I D T H IN M M , o •pa CD < CD era o o CD O M w rt-(U I—1 rt-o cn rt-P <! P CD N c cn £6 1 NTE RORBITAL WIDTH ro co 1N M M . C o O o 1 1 I 1 I I f 1 1 1 G o O cn Z o n o o > XI o 1 — m O z o @§ ' —\ I — C D z o « o O * • • C n O • 76 £ A D Q L E N G T H / B O D Y L E N G T H • i — I — I — I , I —I—'—I—r -i—i—t—i—i—r-ro O O T CD O 5 or m z CD —1 Cn ro o r o Cn • © II n « o N (— > OD TJ m C X) —i . in 9 9 9 • 9 • © © 9 9 0 0 9 90 0 99 © © 0 9 0 © ©9 0 9 © 0 0 0 0 & 0 © £6 CD o o -< m z o H Cn O Cn ro O ro Cn H E A D D E P T H B O D Y L E N G T H o ro O O t " ® © © • •© © • © • © • • © © © © © © • © • © © © 0 3 © © -J> o • © II ii I i I I « • N > p— CO m c JO H 96 OQ CD o cn E Y E - D I A M E T E R / B O D Y L E N G T H O o oo r o p> OQ CD O H+i CD' CD CD •1 O CD o o -< m z o • ® © • • © © • • © ® © O o o •1 cr c 0 o < h- © ro O ro • I© I i IX CD m H N > c © © « © • © © © © © © L6 B O D Y D E P T H A T A N U S B O D Y L E N G T H CD-O '5 m z o H X z o Cn ro o ro cn _ ro . P_ O O i i i i i i i i i i i i i i i i i i i i • • i i i i i i i T m • ® • ® © © ©S © • © • ® ©0 0 0 0 • © Ix n N r~ > CD T) n C XI cn —i 86 CD O O m z \ X Cn O Cn r o o r o Cn L E A S T B O D Y D E P T H B O D Y L E N G T H b b b b o o b p: OJ Cn CO ~ N | C P C O O © • © © © • © | I I I o f— CD m H N > Tl c in © © © © © © © © © © © • © © © © 0 © i 66 P R E - A N A L L E N G T H B O D Y L E N G T H OJ O O CD o o m z o H Cn r o o r o Cn T Cn O I ' ' ' ' O T © • ® © 0 ® • 0 © © © 0* 0 © 0 x ix ru m J3 N > T) C U) 0 ® 0 ® G 001 1G1 PHYLOGENETIC RELATIONSHIPS Position of the Hemilepidotinae within the Family Cottidae Most workers probably accept the hypothesis that the family Cottidae originated from an ancestor similar to the Scorpaenidae. Bolin ( 1 9 4 7 ) suggests ancestral cottids possessed the following characters: - a slightly compressed body (similar to Hemilepidctinae) - a large head (similar to Hemilepidotinae) - spines on the head of which there are four on the preopercle (present Hemilepidotinae) - teeth on vomer and palatines (present in Hemilepidotinae) - four complete g i l l arches and the s l i t behind the last g i l l arch complete (large pore s t i l l present behing last g i l l in Hemilepidotinae) - branched f i n rays (caudal and median fins remain branched in at least the generalized Hemilepidotinae) - g i l l membranes united but free from the isthmus (tends to be present in a l l species but H. spinosus) - soft dorsal and anal f i n short (relatively short in the generalized Hemilepidotinae) - both dorsals continuous (present in Hemilepidotinae) - at least five soft pelvic f i n rays (reduced to only four soft rays in . the Hemilepidotinae) - body covered with strong ctenoid scales (prominent in the Hemilepidotinae) - anus immediately in front of anal f i n (present in Hemilepidotinae) - large, well developed anal papilla or penis absent (absent in the Hemilepidotinae) - few c i r r i (many in Hemilepidotus, however, the author would argue as to what condition is more primitive) 102 The cottids have probably radiated from this basic form and within this framework the Hemilepidotinae are conspicuously generalized. Consequently, the differentiating characters and specializations wi l l tend to be conservative in the Hemilepidotinae. Phylogenetically Important Characters The preceeding account of generalized cottids provides a basis on which to evaluate the various structures in each species. Each character should be considered in terms of how it diverged from the ancestral plan. It will be shown that the species grade in the following order from generalized to modified, when the' characters are considered as a wholes H. spinosus, H. hemilepidotus, H. zapus. H. Jordani. H. gilberti and H. papilio. Meristic Characters Table VIII shows that H. spinosus has fewer fin rays, fewer vertebrae, more scale rows in the dorsal band, and more lateral line pores. H. hemilepidotus also has fewer meristic counts than the remaining species. These characters represent the generalized condition while on the other hand H« papilio is much more m o d i f i e d . Rt.nirjt^r-al Characters A. Scales In ontogeny, postlarvae o r i g i n a l l y develop a simple spinous or spinule-like scale (Fig. 4 8 ) . It grows by the addition of more spinules on each side. These subsequently form a fan-shaped plate above the original base of the scale. Advances in Hemilep idotus were accompanied simultaneously by the neotenic reduction of scales. Thus, in the more specialized condition, growth of scales is slowed or stopped before the scales have reached the Table VIII: Comparason of meristic characters. H.spinosus H. hemilepidotus H. zapus H. jordani H. gilberti H. papilio Total of soft fin .,ray counts(using Schultz and Welander character index) 63-65 63-68 67-72 71-78 65-71 69-73 Dorsal fin spines 11 11 11(12) 11 11 12 (11) Vertebrae 35 35(34-36) 36-37 36-37(38) 37-38 39 Lateral line pores 62-67 59-71 47-58 59-74 55-67 49-65 Number of horizontal rows in dorsal scale band 6-8 5(4) 4(5) 4(5) 4(5) 3(2-4) Number, of horizontal rows in ventral scale band 4-5 7-9 7-9 7-9 7-9 4-5 Pyloric caeca 4 6-7 4 4-5 4(3-5) 5-6(7) 104 D E V E L O P E M E N T OF S C A L E S . F O R L E G E N D F i g u r e 48. 105 size found, in the generalized species. The anal scale row is lost or develops l i t t l e beyond the post larval condition. It is sometimes represented only by minute centers of ossification buried deep in the skin. The scales of the ventral band in H. papilio acquire a few spinules and then develop-ment is reduced. The scales of the dorsal band in H. papilio appear to represent moderately arrested development compared to the well-formed scales of H. spinosus. The scale patterns in the dorsal band of mature specimens, are drawn in Fig. 49. This represents the pattern on a strip of skin below the spinous dorsal fin, and was drawn with a camera lucida. H. zapus is not shown because the dissection of such a rare specimen would not provide valuable enough information. H. spinosus has relatively larger and more crowded scales. H» papilio has specialized through the development of smaller scales, widely spaced scales, and fewer scale rows. H. hemilepidotus is more generalized (except for H. spinosus) because of the greater per-sistence of the fifth scale row and a greater tendency for overlapping of the edges between adjacent scales. This pattern changes somewhat towards the posterior direction and between individuals. The scales shown for H. Legend for Fig. 48 Simple spinule or scale found in postlarvae. Degenerate scale of anal scale row as in some H. jordani. Further development of scale in juvenile. Underdeveloped scale as found in ventral band of H. papilio. Farther development of normal scale (as in reduced dorsal row of H. hemilepidotus). Large scale in dorsal band of H. hemilepidotus. " » " " " » H. spinosus. " " " » " • « H. jordani. . " " " " " M H. papilio. (a) appearance of scale from peripheral view. (b) mid-sagital section of each scale in dorsal band. (c) mid-sagital section of scale in lowest row of dorsal band or from ventral band. 1: 2,3s 4: 5: 6: . 7 : 8: 9: 10: 106 PATTERN OF DORSAL SCALE BAND. ^ ^ ^ ^ ^ ^ ^ H. SPINOSUS —^ A ^5° ^ ^ ^ ^ ^ H J O R D A N I ^  ^ L * - -I^ GILJE H. PAPILIO F i g u r e 49• 107 hemilepidotus and H. .jordani are probably more widely spaced than is normal for this position on the dorsal scale band, A greater surface area covered by scales can be considered generalized as for example, the many scales on the caudal peduncle of H. zapus and the larger area on the back of H. spinosus. It appears that the subgsnus Hemilepidotus has retained a larger number and larger area of scales in the ventral scale band. Anteriorly the scales of the ventral band are larger and more regularly arranged into rows in H. spinosus. H. hemilep idotus also approximates this which is probably a generalized condition. The anal scale row is always present in the adults of those species which possess pelvic cteni (i.e. H. zapus, H. gilberti and H. papilio). This supports those phylogenies which stress the importance of these characters. This scale row is present in the postlarvae of a l l species and persists in some adults of H. .jordani as well as in the above species. This scale row is thus a weak character and its los3 in several species may be parallel occurrences. The scales in the more modified species appear to represent varying degrees of neoteny. B. Tubular Lateral Line Elements (fig. 50) In cottids the ossified tubular elements of the lateral line are probably derived from scales. The upper and posterior surface of these elements possess spinules similar to those on Hemilepidotus scales. Compared to the remaining species H. hemilepidotus approximates the condition of H. spinosus. There is a trend towards reduction of the number of spinules and lengthening of the lateral line element. H. papilio has progressed most in this feature. Spinules are most developed on those elements halfway along the lateral line. Consequently, a similar area on similar sized specimens must be compared. 108 H . S P I N O S U S i n n H . H E M 1 L E P I D O T U S m 1.. H . J O R D A N I in H . P A P I L I O F i g u r e 50. Bony t u b u l a r elements of l a t e r a l l i n e . 1 0 9 P E L V I C C T E N I Figure 5 1 « 110 PELVIC PAPILLAE H . H E M I L E P I D O T U S H . J O R D A N I H . Z A P U S H . P A P I L I O Figure 52. I l l C. Papillae on Pelvic Fins of Male3 Bolin (1944)» Rendahl (1931) and Schmidt (1929) were impressed by the larger pelvic papillae on the males of various Hemilepidotinae. The net result has been to consider H, zapus and H. gilberti as a distinct evo-lutionary line from H. hemilepidotus and H. .jordani. The author has not seen large papillae on H. zapus or H. gilberti, however this character is well documented by Gilbert and Burke (1912), Jordan and Starks (1904), and Watanabe ( i 9 6 0 ) . These so-called papillae are on the medial non-axillary surface of each soft pelvic ray. No males of H. spinosus were available for this study. Females have completely smooth short pelvic fins (except one specimen of H. .jordani). 1. Skeletal elements of pelvic.papillae;- A dissection of the pelvic fin on a mature male H. papilio reveals an ossified core in each papilla (fig. 51)• These skeletal elements arise near the proximal end of each actihotrich. A smaller barb-like structure usually branches at the same suture and near the adjacent actinptrich. These elements can become quite elongate. It is assumed that H, zapus and H, gilberti acquire similar but shorter skeletal elements. The single male H. gilberti available for this study had only rudimentary swellings on each suture between the actino-trichia (fig. 51). H. .jordani and H. hemilepidotus are generally considered to not have papillae. A l l pelvic fins of H. .jordani which the author had at his disposal were simple. These shorter, weaker and simpler soft pelvic rays are also found in the females of a l l species. In general, males of H. hemilepidotus possess the simple condition shown for H. .jordani or the rudimentary swelling shown for H. gilberti. However, one specimen of H. hemilepidotus (catalogue no. BC 62-686) possess well formed skeletal elements (fig. 51). These elements are short and not conspicuous externally, but 112 can be easily felt when touched with a finger. The origin of these skeletal elements are unknown. They are usually fused to the actinotrichia but there is often an apparent suture separating them from the actinotrichia or the adjacent small element. These ossifi-cations stained red with alizarin dye. In the specimen of H. hemilepidotus cited above, a few elements were observed anchored to the sides of another element or «ven lying free in the tissue. This suggests the possibility that these elements are modifications of some other structures such as tubercles or scales. 2 . External appearance of the papillae;- The material of H. spinosus, H. zapus or H. gilberti is insufficient for a complete description of papillae. The long pelvic fins of the large H. zapus did not have papillae but were smooth. Males of H. hemilepidotus and H. .jordani possess an undulating papillate ridge of fleshy tissue on the surface where the large papillae of the other species would occur (fife. 52). The undulations do not appear to correspond to the actinotrichia as do the ossified elements of the other species. At the base of this small ridge, a fleshy groove may often be seen. 3 . Terminology of "Papillae" and "Cteni";- When there is a central skeletal element this structure does not f i t the usual definition of "papillae". Because of their regular nature, the term "cteni" will be used. The term "papillae" wi l l apply to the low humps of the fleshy papillate ridge. The name "papilio" refers to the large "butterfly-like" fins of H. papilio and not the cteni or papillate-like ventral scale band. h> S i g n i f i c a n c e of pap-niaa and, r.t.P.niThe function of cteni is unkr. wn. They must have some selective value since the cteni and long pelvic fins appear rather cumbersome to be dragged over the bottom i f they 113 have no benefit. Because they are found only on males they must have some sexual Importance. The presence of cteni on some males of H. hemilepidotus suggest that this species, H. zapus, and H. gilberti, probably had a common ancestor with a potential for cteni development. Similarly, these species probably had an ancestor common with H. papilio which possessed cteni. Because cteni are described from more species than were originally known, they can not be as readily applied to classifications as previous workers have done. Whether the'papillae" represent the vestigial remnants of cteni and whether they have a sencory or other function is not known. D . Spines on the Head H. spinosus possesses additional heavier ridges or prominences on the occiput in comparison to other species. Since H. spinosus is to be considered relatively generalized,, these structures could represent a more generalized condition. H. papilio possesses postocular and occipital prominences which are represented in the o the r 1 species as raised areas with fine granulations and ridges. There is a trend towards reduction of spines and ridges on the occiput. H. gilberti is advanced in this respect. The modified lower preopercular spine o f H. gilberti is also probably a specialized feature. E. G-ill Membranes Fused to the Isthmus Bolin (1947) considers this character to be a specialized feature. Whether it is unique in the H. sp.inosus line or whether it is found in the ancestors of a l l Hemilep idot inae is not known. H. hemilep idotus appears to have a narrower fold than the other 3pecies which suggests its affinity to H. spinosus, however, no adequate method for quantitative measurement of this character was employed. 114 F. C i r r i Bolin (1947) considers many c i r r i to be an advanced character. Although c i r r i tned to be superficial, variable, and weak characters, their prevalence in some scorpaenids and the generalized Hemilepidotinae suggests that the opposite of Bolin?s statement might be true. H. hemilepidotus which has broad thick c i r r i , is a shallow water species. Another cottid genus, Sigmistess which was studied by the author, possessed the bushiest c i r r i in the shallow water species. C i r r i might reflect habitat rather than phylogeny. Proport ional Measurements A. Fin Measurements Figures 34 and 35 illustrate shorter fins in the generalized H. spinosus» and point towards more elaborate fins in the specialized forms. The pectoral fin length is variable but the average size is greater in the specialized species. The possible use of the pectoral fins for display purposes in behavior is discussed in a later section of the thesis. H. papilio possesses the longest pectoral fins in the male. Males of H. jordani tend to have longer pectorals than females. These larger fins could possibly promote greater exposure and flashing of conspicuous markings for communi-cation. Males of H. papilio have larger pelvic and spinous dorsal fins which might suggest an elaborate behavioral pattern. The modified pelvic fins are on the most specialized species. B. Body Depth H. hemilepidotus is the deepest bodie^/ form of the Hemilepidotinae. The caudal peduncle depth (Fig. 3 2 ) , caudal peduncle length (Fig. 3 1 )j and head length (Fig. 3 0 ) , also indicate that H. hemilepidotus is the deeper or robust bodied, and large headed species. H. jordani falls towards the other 115 extreme. In some groups of. cottids (i.e. Stgmistes)t the shallow water species is the robust form. Therefore the more turbulent conditions of shallow depths and the Intertidal zone possibly selects for this body form. Shorter fins may similarly be correlated with this. The use of proportional measurements should therefore be used with caution in this phylogenetic analysis. Phylogenetic Relationships of the Species The Hemilepidotinae can be grouped naturally into three major lines which have at one time or another been considered distinct genera. H. spinosus was described as Calycilepidotus (Ayres, 1855; Jordan and Ever-mann, I896) and H. papilio as Melletes (Bean, 1880; Jordan and Evermann, 1896). The remaining species were placed in Hemilepidotus (Ciivier, 1829; Jordan and Evermann, 1898). The characters which are listed in the sub-generic descriptions, represent major changes in the differentiation of the Hemilepidotinae. The three subgenera show similar degrees of divergence from each other although Hemilepidotus tends to be midway between the other two subgenera. Table IX lists the relative degree of divergence in each species for each character. No large specimens of H. zapus were available for study so that certain characters (i.e. branched soft dorsal fin rays) could not be evaluated and were designated by a question mark. In a l l cases spinosus represents the most generalized condition of known contemporary forms while H. papilio represents the most modified condition. Those species with the highest score are most modified. The evaluation of these characters is unavoidably j^ubjective. However it is apparent that H. hemilepidotus is highly generalized within the sub-genus, Hemilepidotus. The other species are relatively similar to H, gilberti. Tables IX and X demonstrate that this latter species is a l i t t l e more. 116 Table-IX. Comparison of characters in the subgenus Hemilepidotus with those possessed by H. papilio. Character possessed by H. papilio / species * Hemilepidotus zapus jordani gilberti reduced size and number of c i r r i 0 1 or 1 | 1 few horizontal rows of scales in dorsal scale band 0 1 1 1 fewer spinules over length of lateral line element 0 ? 1 1 unbranched dorsal and anal fin rays 0 ? 1 1 higher fin ray counts 0 • 1 \ more vertebrae 0 h h 1 fold of g i l l membranes across isthmus, deeper i. 1 1 1 development of large, functionally distinct, cteni on pelvic fins • 0 1 0 1 fewer scales on caudal peduncle between dorsal scale and lateral line g 0 1 1 lateral line elements elongate 0 ? 1 1 persistance of anal scale . row in adults 0 1 ^ 1 Total score 1 5i or 9 8j 10| 117 Table X. A few Individual specializations of the species in the subgenus, Hemilepidotus. Character/ species Hemilepidotus zapus jordani gilberti longer first dorsal spine than second dorsal spine 0 0 G 1 broad lower preopercular spine 1 2 G G 1 low no. of lateral line pores 0 1 0 l 2 narrow maxillary c i r r i o 1 G Total 1 2 1 1 2* modified. H. zapus and H. jordani are quite similar and their position could be easily interchanged. Ih considering H. zapus to be slightly more generalized, stress has been laid on the scale pattern of the caudal peduncle. Reasons for not stressing the cteni of the pelvic fins or the presence of the anal scale row were considered previously. Examination of larger specimens is necessary in order to place this species. Figure 53 illustrates the new concept of the phylogeny of the Hemilepidotinae compared to that of Bolin (1947) who typified a l l previous concepts. The phylogeny expressed here is partially supported by the method of phyletic weighting of Cain and Harrison (i960). H. spinosus, H. hemilepidotus and H. papilio can be appropriately differentiated from the other species. However, there are many objections to the method. The range of a certain character compared to the maximum value of that character may be much greater than that of another character and thus unjust weighting is given 118 I S P I N O S U S ' i i i i i LxJ K ft' — - > M O R P H O L O G I C A L D I V E R G E N C E - ^ P H Y L O G E N Y R E C O G N I Z E D IN T H I S S T U D Y Figure 53° Phylogeny of the HemilepidotInae. 119 to i t . For example, the height of the dorsal fin has a range of 1G (Table XI and XII). The figures might be misleading since vertebrae might be phylogenetically a more important character. Subjectivity is employed during the i n i t i a l selection of pertinent characters, consequently, appli-cation of numbers provides a greater sense of objectivity than is the actual case. Many characters such as branched fin rays, presence or absence of spines, or simple and multifid c i r r i are a l l or none characters and without intermediate values they would provide unnecessary weighting. Similarly, some characters change continuously during ontogeny and even though important can not be used quantitatively. The mean of a character can only be used, however a more variable character ought to be treated differently. Table XI and XII demonstrate overall affinity and phyletic affinity of Cain and Harrison (1958; I 9 6 0 ) . Table XIII gives equal value to each character by designating."©" and " 1 0 " as the minimum and maximum value of the mean for each character. As a result we are then dealing with the number of uncorrelated characters instead. In essence, the characters become tabulated similarly to those described in Table IX. Phylogeny can be considered to be a product of the rate of divergence and the time over which this has occurred. Because there is no fossil record, morphological characters must be weighted in order to construct such a phylogeny. Figure 53(b) represents Time (-subjective evaluation of morphology) plotted against the overall divergence or modification of each species. This subjective evaluation of morphology is derived from the previous discussion of characters in respect to convergence, divergence, homology, stability against selective pressure, and a knowledge of the probable origin of the Hemilepidotinae. Tablg XT: Phenetic differences according to method of Cain and Harrison (I960). sh _sz sj sg sp hz hg hp zg zp jg JP gP fin rays 3 6 13 6 10 3 10 3 7 7 0 4 7 3 4 pyloric caecae 29 0 0 0 29 29 29 29 0 0 0 29 0 29 29 vertebrae 0 5 5 5 10 5 5 5 10 0 0 5 0 5 5 body depth 10 5 5 6 0 5 5 4 10 0 1 5 1 5 4 dorsal fin height 3 6 4 8 51 3 1 5 48 2 2 45 4 46 43 caudal peduncle length 6 5 6 14 22 1 12 20 28 11 19 27 8 16 8 caud. peduncle depth 1 7 17 10 12 6 16 9 11 10 3 5 7 5 2 scales above lateral line 25 19 5 19 22 6 20 6 3 14 0 3 14 17 3 1st dorsal spine/2nd dorsal spine 1 7 7 9 14 8 8 10 13 0 16 7 16 7 23 horizontal rows in dorsal band 25 37 37 37 50 12 12 12 25 0 0 13 0 13 13 lateral line pores 2 14 0 4 9 16 2 6 11 14 10 5 4 9 5 transverse rows in dorsal band 7 9 3 0 12 2 4 7 5 6 9 3 3 9 12 interorbital width 22 8 1 $ 29 36 14 4 7 15 10 21 29 11 19 . 8 pelvic fin length 12 22 9 25 53 10 3 23 41 13 3 31 16 44 28 eye diameter 14 16 0 9 9 30 7 5 5 16 25 25 9 9 0 maxillary c i rr i 9 2 32 2 60 11 41 11 69 30 0 58 30 28 58 height of fin membrane 22 23 3 6 9 1 25 16 31 26 17 32 9 6 15 head length 1 4 5 1 2 5 6 2 3 1 3 2 1 2 1 Mean Character Difference 10.7 10.8 9.4 10.6 22.8 9.3 11.7 10.0 18.6 8.9 7.2 18.2 7.8 15.1 1.4.5 s=H. spinosus h=H.hemilepidotus z=H.zapus j=H. jordani ,_g=H.-gilberti p=H. papilio Table XII: Phyletic differences of the Hemilepidotinae according to the method of Cain and Harrison (I960) . sh sz sj sg sp hz hj . hg hp zj zg zp jg JP gP ' Fin rays 3 6 13 6 10 3 1.0 3 7 7 0 4 7 3 4 vertebrae 0 5 5 5 10 5 5 5 10 0 0 5 0 5 5 dorsal fin height 3 6 4 8 51 3 1 5 48 2 2 45 4 46 43 horizontal rows in dorsal band 25 37 37 37 50 12 12 12 25 0 0 13 0 13 13 Phyletic difference 7.7 13.5 14.7 14.0 3.2 5.7 7.0 6.2 22.5 2.2 0.5 16:7 2.7 2677 16..2 s=H. spinosus h= H.hemilepidotus z=H.zapus j=H.jordani g= H. gilberti p=H. papilio Table XIII: Phyletic differences of the Hemilepidotinae when each phyletic character is given equal weighting in "scale from "0" to '10". sh sz sj sg sp hz hj hg hp Z J zg zp zg JP gP fin rays 2 5 10 5 8 3 8 3 6 5 0 3 5 2 3 vertebrae 0 5 5 5 10 5 5 5 10 0 0 5 0 5 5 dorsal fin height 0.1 0.1 0.9 1.6 10 0.0 0.8 1.5 9.9 0.8 1.5 9.9 0.5 9.1 8.4 horizontal rows in dorsal band 5 7.5 7.5 7.5 10 2.5 2.5 2.5 5 0 0 2.5 0 2.5 2.5 Phyletic Difference 1.8 4.4 5.8 4.8 9.5 2.6 4.1 3.0 7.7 1.4 0.4 5.1 1.4 4.6 4.7 s=H. spinosus h hfeH. he milep idotu s z=H. zapus j=H.jordani g=H. gilberti p=H.papilio 122 On the basis of the generalized nature of the Calve ilep idotus line, it is reasonable to assume that this line diverged early from the postulated main stem of the Hemilepidotinae but changed less in form since then. In contrast Melletes diverged greatly in form in apparently the same time. The subgenus Hemilepidotus is midway between these extremes. The trend towards reduction of scales, simplification of fin rays, persistence of anal scale row in most adults, as well as specialization of pelvic papillae and cteni suggest greater genetic affinity and more recent divergence of Melletes from Hemilepidotus than from Calve ilep idotus. Except for possibly the pelvic cteni and absence of a fold of g i l l membrane across the isthmus, C aly c ilep idot us probably has characters which are ancestral to a l l the Hemilep idot inae. H. hemilepidotus shows a similar relationship to the subgenus Hemilepidotus as does Calyc ilep idotus to the rest of the Hemilep idot inae. Fewer fin rays and vertebrae, branched dorsal and anal fin rays, and certain scale characters suggest that this species is most generalized and earliest derived in the subgenus. Compared to H. .jordani, the greater number of scales on the caudal peduncle may designate H. zapus as the generalized form, while the elongate first dorsal spine and modified pre-opercular spine of H. gilberti may designate this latter species as the modified form. However, these species are quite similar and probably should be considered to have a triphyletic origin. The greater variability in H.. papilio and H. gilberti suggest that these species are most flexible and have the greatest potential for future change. 123 TAXONOMY" AND NOMENCLATURE Since one objective of taxonomy is to reflect phylogeny, the relation-ships in the preceeding section must be considered. Bolin (1947) shows a between the specific and family levels. It is then stated that a l l phylogenetic lines which diverge before this level are genera. Bolin complains that many workers recognise many twigs towards the right hand side as genera. On this basis alone the probable more recent divergence of Melletes should cause i t to be synonymized with Hemilepidotus i f Calycilepidotus is also synonymized with Hemilepidotus. Although the relationships between organisms are most naturally shown in a framework similar to Fig. 54, such a phylogenetic representation is highly idealistic. Great subjectivity on deciding where the phylogenetic lines diverge must exist, without a more objective approach it is futile to be involved with whether a marginal genus has really diverged before or after the generic level (i.e. line "CC" in fig. 54). However when a line is established as a certain taxon, a l l distal divergent lines should be included and subordinate in rank. With the present day Linnean concepts of taxonomy, the phylogenetic objectives of taxonomy cannot be met. Often equivalent and more recent phylogenetic lines are given a higher taxonomic level than more basic portions of the phylogenetic stem (i.e. the classes Mammalia and Aves are recent lines compared to the ancestral Reptilia and Amphibia). Although phylogenetic concepts in taxonomy have less use, until new practices are generally followed, these previously mentioned ideals should be employed only where contempory systems of taxonomy permit. In view of the fact that Bolin (1944) synonymizes Calycilepidotus (a practice fallowed by other recent workers), then the synonymy of Melletes demonstrates detailed diagram of the phylogeny of many cottids. A line is drawn halfway Xi 124 Figure 54. Theoretical Phylogenetic diagram similar to that of Bolin (1947). the above principles. If these taxa were given generic status then H. hemilepidotus should be placed in one subgenus while H. zapus, H. .jordani and H. gilberti should be placed in another. GENERAL LIFE HISTORY OF THE HEMTLEPIDOTINAE Little is known about the l i f e history of the Hemilepidotinae. Barsukov (1958) refers to breeding condition of H. papilio while Clemens and Wilby (1949) mention that eggs of H. hemilepidotus were observed near the intertidal zone in March. The spawning season was not established with the specimens studied in this thesis, however, Gorbunova (1964) provides new 125 Information. Eggs (presumably of H. gilberti) were observed at Kamchatka In depths of 10-20 meters In August at temperatures of 5-10°C. Some eggs were hatched experimentally at 2-3°C. The yellow eggs are adhesive and dlmersal. Divers have observed clusters of these eggs on rocks. It is suggested that males probably guard the eggs. Some females of H, gilberti and H. .jordani were observed to be spawned out at this season. Specimens of H. gilberti between 260-360 mm long possessed between 25,455 to 88,084 eggs. A specimen of H. .1ordanit 410 mm length, possessed 139,633 eggs. The eggs, reported by Gorbunova, were about 1.3 mm in diameter and took 5 weeks to hatch. The young were 4 to 4*3 mm long at hatching and reached 10 to 14 mm in the spring. By June and July they were 25 to 38 mm. These postlarvae were taken at great distances from shore and must have been drifting in the open sea for almost 12 months. The specimens available for this study support the preceeding data of Gorbunova, even though many of her specific identifications are doubted. The smallest postlarvae examined by the author was about 12 mm. Mary-spec imens of H. spinosus, H. hemilepidotus, H. zapus and H. .jordani between 17 and 25 mm were examined. These were taken by night light and mid-water trawl far out to sea. It appears that these larvae are carried great distances by ocean currents before metamorphosis into the juvenile and adult stages. Fig. 14 summaries Fig. 38, 39, 40 and 41 and provides a definition between postlarvae and juveniles or adults. The line nAB n designates the size at which the interorbital resumes normal growth of the adult stage. Metamorphosis occurs at approximately 30 mm and specimens above this length are not known from pelagic waters (except the 38 mm specimens reported by Gorbunova). There/is) probably specific differences in time at which 126 metamorphosis occurs. The postlarvae probably settle to the bottom randomly and subsequently there is either selection for the proper habitat or elimination of those over unfavorable habitat. No information beyond what has already been discussed, is known on age or growth. Food appears to consist of any animal material available in the habitat. H. .jordani and H. hemilepidotus have been captured in crab and shrimp traps as well as hook and line, when animal material was used as bait. Shrimps and crabs were found to be abundant in the stomachs of a l l species. H. hemilepidotus frequently devoured marine annelids, while in addition to these items, H. .jordani had devoured Ammodytes, ophiuroids, young octopus, and clypeastroid echinoderms. Hemilepidotus appears to move deliberately and rather awkwardly so that they are probably restricted to slow moving prey. No obvious differences in feeding habits were noted between the species. GEOGRAPHIC DISTRIBUTION Zoogeography The recorded range of the Hemilepidotinae is from the Santa Barbara Islands of California to Bering Strait and then to northern Japan. In order to understand the factors controlling the distribution of the Hemilepidotinae, pertinent oceanographic factors must be considered. A. Factors Associated with Temperature 1. Is ot herms i-/There is obviously a latitudinal cline of temperature and associated ecological conditions. In the north Pacific Ocean this will 127 undoubtedly be a factor i n determining the d i s t r i b u t i o n of each species. Because of the prevalent oceanic current systems, the isotherms are much further north i n the high la t i tudes of the eastern North P a c i f i c than on the western s ide . 2 . V e r t i c a l Temperature S t r a t i f i c a t i o n s - Because of warmer climate towards the south, waters of equivalent temperature are deeper here than i n the nor th . This must be s ign i f i can t i n shallow water species such as H . hemilepidotus which are common i n the t i d e pools of Alaska but are not found i n them so r ead i ly towards the south. The occurrence of H. p a p i l i o i n a t i d e pool of the P r i b i l o f Island (Bean 1580) and i n the Bay of Providence (Barsukov 1958) compared to deeper records further south may be of s i m i l a r s ign i f i cance . 3 . Ices- Much of the Bering Sea and the Sea of Okhotsk freeze over i n win te r . The grinding of ice i n shallow water destroys t i d e pool habitat and a lso a narrow zone immediately below lowest t i d e l e v e l . This must reduce the ava i lab le habitat of shallow water species such as H. hemilepidotus. B . Depth The juveni les and adults of the Hemilepidotinae are en t i r e ly bottom dwel l ing . Therefore movement and migration of the older stages w i l l be near the shoreline i n the case of shallow water forms, and on the cont inenta l she l f i n the case of predominantly deep-water forms. Very deep water w i l l prevent movement across of the adult stages. C. Oceanic Currents (Figure 55) Because postlarvae of the Hemilepidotinae are pe lag ic , current systems are probably the greatest s ingle factor i n the d i s t r i b u t i o n of each species. Their geographic range i s therefore l a rge ly determined by the speed of the OCEAN CURRENTS IN THE NORTH PACIFIC OCEAN Figure 55. 129 current and the length of time before metamorphosis of the postlarvae. The North Pacific Drift moves eastward In the North Pacific. From this current, water is directed northwards along the coast of British Columbia. Water converges into the anti-clockwise Alaskan Gyrol in the Gulf of Alaska and then some is directed westward across the south side of the Aleutian Island chain. Some water filters from here through the Aleutian Islands to the Bering Sea. In general, the Bering Sea has an anti-clock-wise circulation which sends water southward along the coast of Kamchatka and towards northern Japan (Dodlmead et.al. 1962). A oneway flow exists across the Aleutian Islands so that species formed in the east (i.e. H. hemilep idotus and H. .jordani) can easily spread westward. H. zapus is too rare for consideration. Species such as H. gilberti and H. papilio do not appear to have moved eastwards from Kamchatka or the Commander Islands and are probably restricted by this westward flow across the Aleutian Islands. Occurrances of H. papilio in the eastern Bering Sea have probably spread across the continental shelf to the north. Distribution of the Species Many of the published records are included in the descriptions along with the l i s t of specimens which were available for this study. Where possible these records are plotted on the distribution maps. Postlarvae and adult records are indicated separately. On the maps the numbered records refer, to the authority possessing the equivalent number in the bibliography. The shading represents an estimated range for the postlarvae. Off the continental shelf some weighting of this range should be made according to the dominant direction of ocean currents. Wilimovsky (1964) has recently recorded many new records from the Aleutian Islands. 130 Gorbunova*s (1964) records are not recorded here because of the doubtful method of identification of many of her specimens. , A. Distribution of H. spinosus (Fig. 56) H. spinosus has been recorded from the Santa Barbara Island, California (Starks and Morris 1907) to Nootka Island, British Columbia (Clemens and Wilby 1949)* Two collections of adults and one of postlarvae from British Columbia were available for this study. Five additional specimens from near the Brooks Peninsula are also known from British Columbia. Follet (1952) records postlarvae about 46 miles out to sea from an area slightly north of San Francisco. The northern limit of the known range of this species is represented as postlarvae caught by G. McT. Cowan in May, 1959, at 51°N. and 130°W. This record suggests that adults of H, spinosus occur further north than already known, or that there is a con-tinual flow of postlarvae northward. Presumably these young would perish from an unfavourable environment in the north. There are no other obvious geographical barriers preventing their dispersal northwards. In British Columbia H. spinosus probably occurs mostly on the outer coast. Compared to the many ichthyological collections from the Gulf of Georgia, specimens have been taken only from the more inaccessible and poorly collected west coast of Vancouver Island. B. Distribution of H. zapus (Fig. 57) Only Gilbert and Burke (1912) have recorded this species previously. A specimen collected by the author from Chagulak Island extended the known range from Attu and Semisopochnoi Island. These few known specimens suggest that H. zapus occurs throughout the Aleutian Island Chain. There is most likely a large adult population since the postlarvae can be captured so readily in the few midwater trawl hauls made in the region. The rarity \2A% I22°w. 32°N. F i g u r e 56. F i g u r e 57. 132 of the adults is probably a product of the few collections made in their appropriate habitat. The deep records of Gilbert and Burke suggest that H. zapus inhabits areas throughout the continental shelf. C. Distribution of H. hemilepidotus (Fig. 58) H. hemilepidotus is the most wide-ranging species of the Hemilepidotinae. In North America it is recorded from Mussel Point, California (Bolin 1944) to the Alaskan Peninsula and Aleutian Islands (Gilbert and Burke 1912). Contrary to the theory of Schmidt (1929) this species was found by the author at St. Paul Island in the Bering Sea. The Institute of Fisheries has a specimen from the Commander Islands in the U.S.S.R. Schmidt (1929), who considered H. hemilepidotus to consist of a subspecies, H. h. hemil-epidotus from the south and H. h. .jordani from the Bering Sea, appears to have misidentified some of his specimens. Therefore, his records of this species are untrustworthy. The type specimen of Tilesius (1810) from Petropavlosk, U.S.S.R. compares favourably with that form generally described to this species and represents the western limit of the known range (figure of Tilesius is comparable). The specimens of Pallas from the Kuriles (Cottus Trachurus 1814) are not likely to be this species because of some differences in fin ray counts and colour descriptions. H. hemilepidotus occurs in shallower water than most of the other species of the Hemilepidotinae. The more variable conditions of the shallower water and intertidal zones, and the reduced competition from other Hemilepidotinae species in the shallower water zone may account for the wider geographic range of H. hemilepidotus. Along Kamchatka most of the specimens of Hemilepidotus sp. known to science are possibly taken during exploration of commercial species (using trawls etc.). Conseuqently the 133 DISTRIBUTION O F H. HEMILEPIDOTUS. ( F O R K E Y S E E M A P O F H . J O R D A N I . ) Figure 58. 134 habitat near the lew tide level is not fully sampled. H. hemilepidotus will probably be found In this habitat of Kamchatka. As suggested previously the formation of ice in the Okhotsk and Bering Seas may help to limit distribution. Records from St. Paul and Amak Islands are near the southern limit of winter ice. D. Distribution of H. .jordani (Fig. 59) H. .jordani was previously known from near Baranof Island and throughout the Aleutian Islands In Alaska. The records of Schmidt are discounted as they were for H. hemilepidotus. The specimens examined in this study from southwest Kamchatka represent the south-western limit of the known range. Previous records from this area are not verifiable. The records of Andriashev (1937) from the Gulf of Anadyr are accepted because a l l his specimens f i t into the character index of Schultz and Welander (1934) and samples probably would not have f i t i f they were of any other species. The records from the Gulf of Alaska (as shown in Fig. 24) makes it apparent that H. .jordani occurs throughout the continental shelf. For that reason it is supposed that the bottom stages of H. .jordani occur along- the edge of the continental shelf between the Pribilof Islands and the Gulf of Anadyr. North of the Pribilof Islands, however, there are no large specimens from the eastern Bering Sea. E. Distribution of H. gilberti (Fig. 60) Previous distribution records from northern Japan (Watanabe I960) and Kamchatka ( i f Schmidt's records (1929) are to be trusted) are known. Specimens from the Tarter Strait, Commander Islands, northern Japan and southwest Kamchatka were used in this 3tudy. Schmidt (1929) records specimens from further north on the Siberian coast. Since some of his identifications have been challenged by other workers, it seems desirable Figure 59. 137 to consider the northern limit of the known range as unknown. This species has records from deeper waters on the continental shelf. Consequently it probably inhabits at least the edge of the continental shelf around the Sea of Okhotsk. Much of the shallower inshore area might be too dilute for the occurrence of this species. F. Distribution of H. papilio (Fig. 61) H. papilio was described from St. Paul Island (Bean 1880). The known range extends from here to the Amur River estuary and Bering Strait. The form described by Sakamoto (1932) is synonymized and thus represents the southern limit of the species. Andriashev (1937) reidentifies a specimen of Schmidt (1929) as this species. This record is indicated on the distri-bution map, however, since the latitude and longitude given by Schmidt (1929) does not seem reasonable (he mistakingly records an east longitude for west longitude, when he quoted the H. zapus records of Gilbert and Burke, 1912), the record should be considered with caution. In addition to the records on the distribution map, there are two other specimens (one labelled only east Bering Sea) which represent the second and third records from the Alaskan side of the Bering Sea. Since there are no specimens from the Aleutian Islands, it is possible that the species has invaded the east Bering Sea from Siberia via the northern continental shelf. POSSIBLE GEOGRAPHIC ORIGIN OF THE HEMILEPIDOTINAE Considering that the two most generalized species in the Hemilepidotinae (H, spinosus and H. hemilepidotus) are found on the west coast of North America it is most likely that the ancestral forms of this subfamily will 139 have or iginated here a l so . The southern range of H. spinosus would suggest that t h i s species was separated from the main Hemilepidotus stem as shown i n F i g , 62A, I t may be conjectured that an early Pleistocene g l a c i e r had destroyed a large s t r i p of coast l i n e of B r i t i s h Columbia. Neave (1958) postulates the separation of Oncorhynchus from Salmo since the s tar t of t h i s per iod. Enough time has probably elapsed, therefore, so that c lose ly related genera or subgenera could have diverged af ter the s ta r t of the Pleis tocene. H. p a p i l i o most l i k e l y arose from a western off-shoot of the main eastern Hemilep idotus st em ( F i g . 62B). Fluctuat ing water l eve l s of the Sea of Japan and the Sea of Okhotsk could provide a means for i s o l a t i n g H . p a p i l i o i n these waters. Lindberg has observed that the inland seas so produced, remained sa l ine enough for marine l i f e during some periods of . lowered water l eve l s (Neave 1958). Subsequently there was probably a d i s -pe r sa l of the species t o Bering S t r a i t . As suggested prev ious ly , the one way current flow probably prevented d i spe r sa l back across the Aleut ian Island Chain. After the i s o l a t i n g mechanisms i n eastern Asia broke down, there was probably reinvasion of Hemilepidotus stock which could r ead i ly d r i f t across the Aleut ian Islands. In t h i s fashion another stock was i so la ted and probably gave r i s e to H. g i l b e r t i . Lindberg postulates at leas t two Pleistocene water l e v e l changes i n the region (Neave 1958). Geographically, the subgenus Hemilepidotus represents an intermediary connection between Calyc i l e p idotus and Mellet es. The der iva t ion of the remaining species i s more obscure. Figure 62 demonstrates the r e l a t i v e geographic areas from which the species arose, H . hemilepidotus was most l i k e l y derived from the eastern part of the range of the subgenus Hemilepidotus. I t probably merged wi th the geographic range of H. spinosus and might have even been s p l i t from the main stem by a sub-Figure 62A :j^ $o««\Xv. = Area of differentiation of ; v y;.'.;.'i,\; •;• Hemilep idot us-line _ Area of development of Melletes-line 141 Figure 62C Figure 62D u. x ft. Range of H. ; hemilepidotus - Range of H., papilio Area of development of H., zapus _ . Area of development of ~ .i ordani-gilbert i-line 142 Figure 62E Area of development of H. .jordani Area of development of H. gilberti sequent glacial period. The stock that gave rise to H. zapus, H. .jordani and H. gilberti probably arose in the Aleutian Islands and Bering Sea area. H. .jordani likely arose in the Gulf of Alaska or Bering Sea, H. zapus in the Aleutian Island chain and H. gilberti in the Okhotsk sea. Whether H, .jordani or or H . zapus split off first or whether there was a triphyletic origin is debatable. The far western occurrences of H. .jordani and H. hemilep idotus are no doubt subsequent invasions across the one-way flow of the Aleutian Island bridge. The greatest evolutionary change and speciation as well as the greatest potential for future change appears to be in the Bering Sea or Sea of Okhotsk. It is interesting that the two most advanced species occur at the western-143 most part of the range and each have tended to follow similar evolutionary-trends (i.e. specialization of pelvic cteni). It would be most interesting to isolate those causal factors for this great differentiation and contrast them to the factors maintaining the generalized forms on the North American coast. BIOLOGICAL DIFFERENCES BETWEEN THE SPECIES Sympatrv Between the Species of the Hemilepidotinae The distribution of the species of Hemilepidotus is summarized in Table XIV. The most significant point is that, on a geographic bases, a l l the species are sympatric with the form with which they are most related. In addition, a l l the species which are underlined In the table have been taken in the same collection as the other species underlined in the same column. Sympatry should indicate the likelihood of biological differences between species when they are also morphologically distinct. Depth and Habitat Preference Information on depth is absent for many of the collections used In this study. Table XV summarizes the known depth of some of the collections of each species of Hemilep idot us. As noted previously, preferred depth probably changes with latitude. H. hemilepidotus is found in shallower depths than the other species. Depth preference of H. spinosus is not known by the author. The specimens from the Halibut Commission were taken mostly by trawl and are presumed to be from greater depths. Depth preference is a clue to the habitat of the species concerned. Although more observations are needed, habitats of H. hemilepidotus and H. ,1 ordani were observed in Alaska. H. hemilepidotus is generally taken in Table XIV: Summary of Distribution of the Hemilepidotinae. Southern Calif. North Calif. British Alaskan Columbia Panhandle Aleutian Islands Bering ; Sea Kamch-atka Sea of Okhotsk Northern Japan H. spinosus XX XX XX H. h emil epidotu s XX XX XX XX XX XX H. zapus XX H. jordani XX XX XX XX XX w H. gilberti XX XX XX XX H. papilio XX XX XX XX 145 Table XV. Summary on depth of collections of the H emilep idot inae H. hemilepi-dotus H. zapus H. jor-dani H. gilberti H. papilio Tide pools 49 collections 6 collec-tions 1 collection (3) Deeper water 10 collections (0-90 ft.) 200-350 feet (1) 15 collec-tions (10-360-feet) . 2 collections (2) 3 - -collections 4 collections (4) (5) Halibut Commission collections 3 collections 65 " collec-tions (1) Gilbert and Burke (1912) (2) Watanabe (I960) (3) Bean (1880) (4) Andriashev (1937) (5) Barsukov (1958) rocky tide pools or immediately below this where the water and bottom was clean, often where surge from waves predominates, and often where crusty and coral-like algae occur. The 3 l i g h t l y deeper habitat which is probably preferred to the south, will differ slightly. Specimens of H. .jordani, with the usual brownish colour of the species, were observed during skin-diving operations. They were in a bay where the water was undisturbed. A vertical ten foot strip overgrown with brown algae occurred along the shore. Beyond this the bottom was composed of brown sediment or muck that was easily disturbed. This dirtied the normally clear water. Fishermen have described specimens (presumably of this species) in shrimp traps on areas of similar bottom. A few juvenile specimens, which are supposedly strays from their normal habitat, were observed with collections of H^  hemilepidotus in the 146 typical habitat of the latter. These specimens of H. .jordani were coloured reddish similar to the H. hemilepidotus. The general colour differences could be an indicator of the preferred habitat. Sexual Dimorphism in the Hemilepidotinae The males of a l l species have distinctive features in morphology and colour pattern. Females of a l l species tend to have shorter pelvic fins and pale coloured ventral surfaces. Details of the structure of papillae and cteni were discussed previously. Colour differences are described in the species descriptions. H. zapus, H. gilberti and H. papilio probably have long enough pelvic fins and large enough cteni to be functionally important, however, large cteni were observed only on H. papilio by the author. The under surface of the males is illustrated in fig. 63, 64, 65, 66 and 67. No males of H. spinosus were available for this study. Breder (1963) describes defensive behaviour in Scorpaena, a group which may possess much in common with ancestral cottids. Apparently some species can flip over the pectoral fins and expose conspicuous colour patterns on the axillary surface. An examination of several species of Scorpaena at the Institute of Fisheries indicates that conspicuous patterns are common in the axil of the pectoral fin. It is reasonable that the Hemilepidotinae could have inherited similar mechanisms of communication. Since only the males adopt these patterns then this behavioral mechanism should be of reproductive or sexual,significance. If these colour patterns produce distinct but different responses between sympatric species during breeding activity, then we have an important mechanism by which the species of the Hemilepidotinae can remain distinct. Many cottids such as Myoxocephalus have been observed with distinct patterns on the belly by the author, and consequently factors associated with this dimorphism may be highly important 147 Figure 63 . Sexual dimorphism in H. hemilepidotus; male is about 200 mm in standard length. (Note pattern in axil of pectoral fin, short pelvic fins and darker pelvic fins of male) Figure 64. Ventral surface of male H. zapus; standard length is 87 mm. (Note light colour of ventral surface, suggestion of a dark streak at base of pectoral fin and long pelvic fin) 148 Figure 65 . Sexual dimorphism In H. .jordani: male is approximately 200 mm in standard length. ^Note absence of pattern in axil of pectoral fin; dark colour of anal, pectoral and pelvic fins; and relatively shorter pelvic fin of male) and longer pelvic fin in male) 149 Figure 67. Sexual dimorphism in H. papilio; female is 216 mm in standard length. (Note pattern on pectoral and pelvic fins, the very long pelvic fins, cteni (papillae) on pelvic fin in male) in many cottids. The large pectoral fins of cottids would certainly facilitate flashing of these signals. Differences of Growth in the Hemilepidotinae In the present study, no aging or analysis of growth rates were per-formed. The specimens used in Fig. 68 are from a sample of postlarvae taken with a night light on July 31, 1961 at Amchitka Island. The differences in length would suggest that these species have different growth rates, are born at different sizes or breed during different seasons. At this point the differences in allometric growth of the eye and interorbital should be noted (fig. 37, 38, 39, 40 and 41). The preceeding evidence shows that the species of the Hemilepidotinae are biologically, as well as morphological, distinct. Since H. hemilepidotus and H. .jordani were considered to be subspecies by Schmidt, the differences S T A N D A R D L E N G T H I N . M M . Figure 68 : Length frequency of H. hemilepidotus and H. jmrdaini from collection number BC .63-4010. 151 between these species should be stressed. The basis for subspecies designation of H. gilberti and H. zapus by Gorbunova (1964) are not'known by the author. The distinct morphology and apparent differences of the males as well as differences in the known maximum size of these species, suggest that they are distinct. CONCLUSIONS 1. The Hemilepidotinae are a distinct group of six species within the Cottidae. 2. Meristic and structural characters are better for the identification of species than proportional measurements. 3 . Postlarvae are characterised by major changes of relative growth of Interorbital width and other body parts at standard lengths between 20 aid 30 mm. 4. Juveniles and adults are.characterized by the stabilization of the proportional width of the interorbital width after metamorphosis of the postlarvae. 5. A l l species can be identified after fin rays, lateral line pores, and scale bands have formed in the postlarvae. 6. The Hemilepidotinae are generalized cottids. . 7 . There are three major phylogenic lines in the Hemilepidotinae which have been given subgeneric status. 8 . Calyc ilep idotus (type spinosus) is* the most generalized subgenus of the Hemilepidotinae. 9. Melletes (type papilio) is the most modified subgenus of the Hemilepidotinae. , 152 10. The subgenus Hemilepidotus (type hemilep idot us) forms the third phylo-genetic line and is composed of four species. 11. H. hemilepidotus is most generalized in this line. 12. H. zapus, H. .jordani and H. gilberti might have a triphyletic base although the former tends to show more generalized characters while the latter has a few more modifications. 13* The generalized H. spinosus and H. hemilepidotus are found and probably originated in the eastern Pacific Ocean, and this suggests that the Hemilepidotinae originated here also. 14. The more modified H. gilberti and H. papilio probably originated in Asian waters. 1 5 . H. zapus probably originated in the Aleutian Island chain. 16. H. .jordani probably originated in either the Gulf of Alaska or Bering Sea. 17. Occurrences of H. hemilepidotus and H. .jordani in the Aleutian Islands and Siberia probably arrived by means of the westward water current across the Aleutian Islands. 18. Different morphological and colour patterns in the males suggest distinct patterns of sexual recognition and communication in each species of the Hemilepidotinae. 19. Differences in depth preferences, differences of characters providing possible means of sexual recognition, different patterns of allometric growth, and sympatry between closely related species on a broad geo-graphic basis suggest that a l l six species of the Hemilepidotinae are also biologically and ecologically distinct. 153 SUMMARY The Hemilepidotinae are restricted to those species of cottids with three scale bands of which the upper one is continuous across the midline anteriorly, continuous dorsal fin* and 4 soft rays of the pelvic fins. Some definitions of counts and measurementsj as well as certain c i r r i , spines and scale bands are given, Allometric growth is prominent, especially in eye and interorbital proportions of young specimens. Interorbital width shows reduced growth during metamorphosis from post-larvae to juveniles. This occurs at about 20-30 mm. Meristic and structural characters are better means of identification of the Hemilepidotinae than proportional measurements. H. spinosus can be identified at sizes down to at least 20 mm, by the 6 or more rows in the dorsal scale band as well as by nb free fold of g i l l membrane across the isthmus. H. hemilepidotus can be identified by the four or five scale rows of dorsal band,. branched anal and dorsal rays, and the 68 or less rays in both pectoral, anal and soft dorsal fin. This species was identified at sizes down to 16 mm. H. zapus has scales on caudal peduncle above lateral line,, as well as 58 or less lateral line pores. Specimens were: identified down, to 17 mm. H. jordani possesses 17 to 19 pectoral raysJ or 72 or more rays in the soft dorsal, anal, and both pectoral fin rays. There are also no scales immediately above lateral line on caudal peduncle. This species was identified down to sizes of 17 mm. H. gilberti is distinguished at sizes above 40 mm by the longer first spine compared to the second dorsal spine, 17 or fewer pectoral rays, 154 reduced or absent frontal cirri., a flattened lower preopercular spine as well as no scales on caudal peduncle above lateral line. ~ M.» papilio has 12 dorsal fin spines as well as reduced scales in the ventral scale band, - Ar t i f i c i a l keys to the postlarvae and adults as well as phylogenetic keys are presented, - All species, subgenera and a single genus are described, - In cottids, many compact scales, many spines, fewer counts of fin rays, teeth on palatine and vomer, g i l l s l i t behind last arch, branched rays, and high pelvic fin counts are the ancestral condition of the Cottidae. - The Hemilepidotinae are generalized cottids. - H. spinosus is most generalized*of the Hemilepidotinae. - H. papilio is most modified of the Hemilepidotinae. -The remaining species form a phylogenetic line midway in development between the line of H. spinosus and the line of H. papilio. - H. hemilep idot us is most generalized in this line. - H. zapus might be more generalized than H. .jordani or H. gilberti but the three species are closely related compared to H. hemilep idotus. - S» gilberti is the most modified of the species in the middle phylogenetic line. - The vestigial remains of cteni on some male H. hemilepidotus breaks down the phylogenetic concept that groups H. hemilepidotus with H. .jordani and H. zapus with H. gilberti. - Branched fin rays, number of vertebrae, number of scale rows, scale size, number of spinules on lateral line elements, number of fin rays, and abundance of c i r r i indicate the following sequence of generalized to modified species. H. spinosus, H. hemilepidotus, H. zapus, H. jordani, H. gilberti and Ho papilio. 155 For the most part H., papilio shows modifications already suggested in other species of Hemilepidotus. ' H. papilio probably has more recent affinity to the H. hemilepidotus line than H. spinosus and therefore should be synonimized within the genus Hemilepidotus i f H. spinosus is also. Gajycilepidotus (type spinosus), Hemilepidotus (type hemilepidotus) and Melletes (type papilio) are retained as subgenera within genus Hemilepidotus. Factors associated with temperature probably restricts the northern and southern distribution of the Hemilepidotinae. Oceanic currents are probably the most significant factor in the distri-bution of the pelagic young. The westward flow of water across the Aleutian Islands has probably transported species westward but not eastward. The juveniles and adults are probably restricted to the bottom of the continental shelf and consequently do not disperse across deeper water. H. spinosus is known from the Santa Barbara Islands, California to Queen Charlotte Sound. H. hemilepidotus is known from San Francisco and Monterey to Kamchatka and the Pribilof Islands. H. zapus is probably distributed throughout the Aleutian Island chain. H. jordani is known from Baranof Island, Alaska to the Gulf of Anadyr and southwest Kamchatka. H. gilberti is,known from northern Japan to the Commander Islands, U.S.S.R. and probably further north. H. papilio is known from northern Japan to Bering Strait and East Bering Sea. 156 - All.species are sympatric in some part of their range with their most allied or closely related species. - The two most generalized species are in the eastern Pacific Ocean while the two most modified species are in the western Pacific Ocean. - The Hemilepidotinae probably originated in the eastern Pacific Ocean. - Pleistocene glaciation might have played a part in the speciation of H. hemilepidotus and H. spinosus. - Isolation in the Sea of Japan and Sea of Okhotsk might have permitted the speciation of H. gilberti and H. papilio. - H. zapus and H. .jordani probably arose in the Gulf of Alaska, Aleutian Islands or Bering Sea. - Occurrences of H. hemilepidotus and H . .jordani in Asian waters reflect subsequent invasion across the Aleutian Islands. - Postlarvae are known only for H. hemilepidotus,. H. spinosus, H. .jordani and H. zapus. - Little is known of the l i f e history of the Hemilepidotinae. - Most of the species of Hemilepidotus are deeper water forms but H. hemilepidotus occurs in the tidal and subtidal zone. - The males of a l l species of Hemilepidotus have different and distinct colour patterns and morphology. - The function of these characters are unknown. - It is suggested that these markings have behavioral significance and it is suggested that the markings in the axil of the pectoral fin might have had a homologous origin as a means of communication similar to that described for Scorpaena. - Differences of growth are indicated in the young of H. hemilepidotus and H<, .jordani. 157 - A l l six species in the Hemilepidotinae are biologically as well as morphologically distinct. 158 LITERATURE CITED References marked with an asterisk were not seen by the author. Okada and Matsubara,and Taranets were taken from Schmidt (1950). Except for Cuvier, Cuvier and Valenciennes, Ayres, and Girard, these references were quoted by Schultz and Delacy (1936) as distributional records. 1. Andriashev, A. P. 1937* A contribution to the knowledge of the fishes from the Bering and Chukchi Seas. Liza Lanz and N. J. Wilimovsky, (trahsl.). U. S. Dept. Interior, Fish and Wildlife Service. Spec. Sci. Rept. Fisheries No. 145, 1955. 80 pp. 2. Andriashev, A. P. 1954. Ryby Severnykh Morel SSSR. Akademii Nauk SSSR. Moskva. 3 . *Ayres, W. 0. 1854. The Pacific. 3(42): 166. 4. Ayres, W. 0. 1855. Proc. California Acad. Nat. Sci., 1:75-77. 5. Barnhart, P. S. 1936. Marine Fishes of Southern California. Univ. California, Berkely. 209 pp. 6. Barsukov,T. V. 1958. Rybr bukhty Lrovideniia i sopredelnykh vod Chukotskogo poluostova. Trudy Zool. in-ta, t. 25. 7. Bean, T. H. 1880. Descriptions of some genera and species of Alaskan fishes. Proc. U. S. Nat. Museum (1879). 2:353-359. 8. Bean, T. H. 1882a." Descriptions of new fishes from Alaska and Siberia. Proc. U. S. Nat. Museum {1881). 4:144-159. 9. Bean, T, H. 1882b. A preliminary catalogue of the fishes of Alaska and adjacent waters. Proc. U. S. Nat. Museum (1881). 4:239-272. 10. Bean, T. H. 1884. Notes on a collection of fishes made in 1882 and 1883 by Capt. Henrv e. Nichols, U.S.'N., in Alaska and British Columbia, with a description of "a new genus and species, Pripnistius macellus. Proc. U. S. Nat. Museum (1883). 6:353-361. 11. Bolin, R. R. 1944. A review of"the marine cottid fishes of California. Stanford Ichthyological Bull. 3(l):1^135 .12., Bolin, R. R. 1947. The evolution of the marine Cottidae of California with a discussion of the genus as a Systematic Category. Stanford Ichthyological Bull. 3(3):153-l68. 13. Breder, CM. 1963. Defensive behavior and venom in Scorpaena and Dactylopterus. Copeia. (4):698-700. 159 14a. B u l l . Z o o l . N o m e n . 1950. 4:403. 14b. C a i n , A . J . and H a r r i s o n G. A . 1958. An a n a l y s i s o f the t a x o n o m i s t ' s judgement o f A f f i n i t y . P r o c . Z o o l . Soc . London. 131:85. 1 4 c C a i n , A . J . and H a r r i s o n G. A . I 9 6 0 . P h y l e t i c W e i g h t i n g . P r o c . Z o o l . Soc . London. 135:1-31. 15. Clemens, W. A . and W i l b y , G. V . 1949. P i shes o f the P a c i f i c Coast o f Canada, B u l l . F i s h e r i e s Research Board o f Canada. ( 6 8 ) : l - 3 6 8 . 16. Clemens, W. A . and W i l b y , G . V . 1961. P i shes o f the P a c i f i c Coast o f Canada, B u l l . F i s h e r i e s Research Board o f Canada. ( 6 8 ) : l - 4 4 3 . 17. C u v i e r , G . 1829. Le regne an imal d i s t r i b u e M d*apres son o r g a n i s a t i o n , pour . s e rv i c de base a l ' h i s t o i r e n a t u r e l l e des animaux et d T i n t r o d u c t i o n a l T a h a t o m i e comparee, P o i s s o n s . (second e d i t i o n ) P a r i s . V o l . 2 , 5 3 2 p p . 18. C u v i e r , G. 1834. The c l a s s P i s c e s . 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